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Zeitschrift für Zoologie

Herpetofauna Caboverdiana

Von Hans Hermann Schleich

SPIXIANA Supplement 12

München, 30. Juni 1987

ISSN 0177-7424

SPIAIANA

ZEITSCHRIFT FÜR ZOOLOGIE

herausgegeben von der

ZOOLOGISCHEN STAATSSAMMLUNG MÜNCHEN

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Schwerpunkten in Morphologie, Phylogenie, Tiergeographie und Ökologie. Manuskripte werden

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Umfangreiche Beiträge können in Supplementbänden herausgegeben werden.

SPIXIANA publishes original papers on Zoological Systematics, with emphasis on Morphology,

Phylogeny, Zoogeography and Ecology. Manuscripts will be accepted in German, English or

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edited in supplement volumes.

Redaktion — Editor-in-chief Schriftleitung — Managing Editor

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ZOOLOGISCHE STAATSSAMMLUNG MÜNCHEN

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SPIXIANA - Journal of Zoology

published by

The State Zoological Collections München

Herpetofauna Caboverdiana

Von Hans Hermann Schleich

Gewidmet dem jungen Staate Cabo Verde -

für eine Kontinuität der Regenzeit

München, 30. Juni 1987 ISSN 0177-7424

SPIXIANA Supplement 12

[N —— —n

Dr. Hans Hermann Schleich,

c/o Zoologische Staatssammlung, Münchhausenstraße 21, D-8000 München 60

Gesamtherstellung: Gebr. Geiselberger, Altötting

Herpetofauna Caboverdiana

Von Hans Hermann Schleich

Inhaltsverzeichnis

MoLworte. 22 2 OTTO SATIODDEE SITHETTRORERITTEN AAN AAN,

AN Samamentassungs hun Ar re EN ee a ER ee ee Are:

Emleitunga me N ee SORT Lew Stck erEtt,

INIlgEmeimersKeill nr ANTRIEBE ERTENIEN SSER RN,

WiesKapyerdischenälnselnsn®NTakaronesienen en ee

Bemerkungen zur Geographie, Geologie und Klimatologie ..............

Narerialsund N ethocik rer NL N N N NND

Artenliste mit Merkmalscharakteristik und Verbreitungstabelle der

kapyerdischen, Echsen A. mer Insassen name een Mask angel

SystematischerTeil >... Merle bestnelal. yobnu. en IRAR

Seineidae — Genuse@Mabuyalaoieid ort IEREBINIRLIEN IA ER.

Mabuyardelalandu (DUNErm Se BERONMISSS)E2 2

WIGbuyaaogoenSsıs(O)SEAUGEINESSK LSA). en:

Mabuya fogoensis fogoensis (O’SHAUGHNESSY, 1874) .. 2.2.2222 nenennn

NIab2ayaBo 20ensısim.colauensisnovasspa

INabeyalosoensissantaoensis noy..sspa nr. 2

Pdrbuyasstangen. (GRAS 1SAD) 2 200 0 cn ne ee:

NiEboyalstangen stanzen.(Gras 182) ee:

Mabeyalstanzenmaioensisa MERTENSIOSSE. TE Senn

ab ayastanzen salensısu N Gen, 19550 ee:

WIabuyastangenuspinahs BOULENGER, 19006, 2. 2 20 0:

NiAbuyakvanllantıBoulENeerylBSZ

Grekkondae &enuspljarentolaWe

LlanentolaanudısaB OUBENGERSL OO

Manentolamedismudis HB OUNEN GER IO

DanentolammudısunaioenSsishS CENEICEI SA

Manerttolamudıspnotogigas|OGERSTOSAE De reg:

Manentolandarwınıs]ocER, 1984 zo. en oe

NckentolarcabovendianauS eEmE CE 98T

Tarentola caboverdiana caboverdiana SCHLEICH, 1984 . 2 2... 2 once.

Tarentola caboverdiana nicolauensis SCHLEICH, IIBA . . . oo... ....n nn.

ManentolarcabovendianamazianaS emrrıern, 198 ee

Manentolalcaboverdianasubstitutaloc 19842. anne

Manentolaaasası Bockern 189) a a

Üanentolasıgasigas BoctcHMls96)E a ee

Manentolassasasibrancoensis,Semueiem, [984 Mr Ware

4.2 Gekkonidae Genus Hemidachyjlus 2... 2...2. . Denken ee 51

4.2.5 Hemidactylus brooki angulatus TIALLOWELL, 1852 „. .n ureenocrocnonens 51

4.2.6 Hemidactylas bondier (BOCouRT, 1870). een 52

4.2.6.1 Hemidactylus:bouvieri boavien (Boca; 1870) 2.2 An nn nun: 52

4.2.6.2 Hemidactylus bouvieri boavistensis BOULENGER, 1906 . .... 2.2... cn eenen 53

4.2.6.3 Hemidactylus bouvieri razoensis GRUBER & SCHLEICH, 1982 ........ cr... 54

4.3 Testmehmesun Ste, ee ee 55

4.4 Amphibia Butonidaer 2 era ee el lt ee er en ee 56

5. Die einzelnen Inseln. - Zu ihrer Geographie und

Biologie - Okologie der einzelnen Arten... u. a oa en 56

51 SAL, Artenspektrum, Fundorte, Biologsie-OÖkologie... .. 2... Meran 56

52 BOA VISTA, Artenspektrum, Fundorte, Biologie-Ökologie ............. 59

53 MAIO, Artenspektrum, Fundorte, Biologie-Ökologie ........ 2.2.22. 60

5.4 SAO THIAGO, ILHEU ST. MARIA, Artenspektrum, Fundorte,

Biologie-Okolosie.. 2: sus4 20 2008208 20er. BNEE 60

55 FOGO, Artenspektrum, Fundorte, Biologie-Ökologie ................ 63

5.6 BRAVA, Artenspektrum, Fundorte, Biologie-Ökologie ............... 64

De ET EISDORHOMBOL,. 37 Klara aan waren kr 66

5.8 SANTO ANTÄO, Artenspektrum, Fundorte, Biologie-Ökologie ......... 66

5.9 SAO VICENTE, Artenspektrum, Fundorte, Biologie-Ökologie .......... 67

5.10 SANTA LUZIA, Artenspektrum, Fundorte, Biologie-Ökologie .......... 68

Se BRANCO, Artenspektrum, Fundorte, Biologie-Ökologie .............. 69

5412 RAZO, Artenspektrum, Fundorte, Biologie-Ökologie ................ 70

5313 SAN NICOLAU, Artenspektrum, Fundorte, Biologie-Ökologie .......... 72

6. Biotop-und’Artenschutze. 3.24 Sen... 020er N 73

2: Schrifttum. 2... en Se ee ee Se 74

Abkürzungen für Sammlungen und Museen:

ZSM Zoologische Staatssammlung München

BMNH British Museum (Natural History) London

ZFMK Zoologisches Forschungsinstitut und Museum A. Koenig Bonn

MCNG Museo Civico di Storia Naturale Genova

Vorwort

Eine ungezählte Namensliste kapverdischer Bürger, seien es Bauern, Fischer oder Kinder, die mit Rat und Tat,

mit Fanghilfe und Hinweisen meine Arbeiten unterstützten, aber auch viele entgegenkommende Entwicklungshel-

fer, die sich um mein und meiner Begleiter Wohlergehen kümmerten, wären hier anzufügen. Allen sei hier aufrich-

tigst gedankt.

Einen wohl kaum in Dankesworte ausdrückbaren Beitrag zum Gelingen des Projektes leisteten meine Reise-

begleiter D. BENDER (Basel), H.-J. GRUBER (Schliersee) und M. WUTTKE (Frankfurt).

Mein allerherzlichster Dank für ihr persönliches Engagement und zuweilen auch tatkräftige Unterstützung bei

Recherchen, aber auch im Gelände, gilt besonders E. STEIN, weiter I. FARIA, M. DUARTE-ALMEIDA, H. SOARES,

A. PırEs, X. BARBOZA, M. ALMEIDA (alle genannten Cabo Verde) sowie H. & K. ZILLER.

Dr. U. GRUBER und Dr. W. KASTLE korrigierten freundlicherweise das Manuskript und trugen durch ihre

freundschaftlich kollegiale Kooperationsbereitschaft sehr zum Gelingen dieser Arbeit bei.

Für die Exkrement-Rückstandsuntersuchung zur Analyse der Nahrungszusammensetzung bin ich Dr. E.-

G. BURMEISTER zu verbindlichstem Dank verpflichtet.

Leihmaterial stand mir dankenswerterweise aus den Museen Wien, Turin und London zur Verfügung. Eine Aus-

leihe des MERTENS’schen Materials aus dem Senckenberg-Museum Frankfurt war leider auch nach mehrmaligen

Versuchen über Jahre hinweg nicht möglich. Der Deutsche Akademische Austauschdienst (DAAD) förderte durch

seine Finanzierungshilfe der zweiten Reise im Frühjahr 1981 den Fortgang der Arbeit.

Besonderer Dank gebührt auch Dr. W. LoBIn (Frankfurt) für die Zusammenarbeit und Korrektur dieser und

vorhergehender Kapverdenarbeiten.

Sowohl einige Exemplare der beiden Riesengeckoarten der Inseln Branco und Razo als auch Mabuya vaillanti

von $. Thiago befinden sich zu Beobachtungs- und Nachzuchtzwecken in Terrarienhaltung bei Herrn H.-J. GRU-

BER und mir. Betreffs letztgenannter Art Mabuya vaillantı danke ich herzlichst Herrn R. STEFFENS für die Fund-

orthinweise, aber auch für das Überlassen von drei Zuchttieren.

Aus Prioritätsgründen mußte leider ein Teil zur Systematik der Geckos (Gattung Tarentola) vorab (SCHLEICH

1984) publiziert werden.

Zusammenfassung

Die Herpetofauna der Kapverdischen Inseln wird beschrieben und revidiert. Die der Beschreibung zugrunde lie-

genden Informationen stammen aus der bisher bekannten Literatur sowie aus dem in den Jahren 1977-1981 neu auf-

gesammelten Material.

Sieben Formen werden neu beschrieben (incl. SCHLEICH 1984), zwei revalidisiert sowie zwei aus dem Unterart- in

den Art-Status erhoben. Aufgrund der Ausbeute von Fischern, der auf den Märkten angebotenen Ware und den In-

formationen durch die SCAPA-Dienststellen waren Angaben über das Vorkommen von Seeschildkröten möglich.

Neben den systematisch-taxonomischen Beschreibungen werden Angaben über die einzelnen Inseln sowie zur

Ökologie und Biologie ihrer Herpetofaunen, basierend auf Geländenotizen, Terrarienhaltung, Zuchterfolgen und

Kotuntersuchungen gegeben. Ein dem Autor dringend notwendig erscheinender Arten- bzw. Biotopschutz wird

diskutiert.

Summary

The herpetofauna of the Cape Verde Islands is described and partially revised. The paper is based on the literature

and own collections and descriptions of the material collected in the years 1977-1981. Several new forms are descri-

bed or their taxonomic status is discussed.

Informations about the seaturtles are due to the public merchandise or informations from SCAPA. Beside the sy-

stematic and taxonomic work informations about the islands, their biological-herpetological facts based on fieldno-

tes or terraristic observations are given. The necessity of a general protection of species and habitats is discussed.

Resume

L’herpetofaune des Iles du Cap Vert est descrite et revisee (en partie). Les informations qui font la base de cette

description, sont tirees de la litterature d&ja existante A ce sujet, ainsi que du materielnouvellement collectionne dans

les annees 1977-1981. Il s’agit d’une nouvelle description pour sept formes (incl. SCHLEICH 1984), ainsi que d’une

discussion de leur statut taxonomique.

Les informations sur les tortues de mer sont dues ä la marchandise publique et aux renseignements de laSCAPA.

En plus des descriptions systematiques et taxonomiques, ce rapport contient des informations sur les differentes is-

les, concernant l’Ecologie et la biologie de leur composition herpetofaunistique, bas&es sur des notes sur le terrain,

des observations en captivite, des succes d’elevage et des analyses des excrements.

L’auteur y ajoute une discussion sur la necessite absolue d’une protection des especes et des habitats.

Resumo

A herpetofauna das Ilhas de Cabo Verde & descrita e revisada. As descrigöes baseam-se na literatura e no material

das colecgöes novas dos anos 1977-1981.

Sete formas säo descritas de novo (incl. SCHLEICH 1984), duas säo revälidadas e duas subespecies säo elevadas ao

nivel de especies.

O estudo da pesca e dos mercados de peixe assım como as informagöes da SCAPA facilitaram indicagöes da pre-

senga das tartarugas do mar. Junto das descrigöes sistemäticas e taxonömicas esta publicagäo contem informagöes da

cada Ilha por sı e da ecologia e biologia da herpetofauna. Estas indicagöes baseam-se nas noticias do terreno, nas

estudos do terrario, nas criagdes e nos anälises dos excrementos.

O autor discute a necessidade e a maneira prätica duma protecgäo das especies e dos biotopos.

1. Einleitung

Die Untersuchungen und Bearbeitungen der Kriechtiere der Kapverden wurden durch die erneute

Aufnahme von Nachforschungen zur Überlebensfrage von Macroscincus coctei (s. SCHLEICH 1979a

und 1982) eingeleitet. Eine erste Sammelreise im Sommer 1979 führte dabei auf die Inseln Boa Vista,

Sao Thiago, Fogo, Branco, Brava und S. Nicolau. Während eines Aufenthaltes auf den Inseln Branco

und Razo, dem einstigen Lebensraum von Macroscincus coctei, konnten leider keinerlei Spuren und

Anzeichen für ein Überleben des kapverdischen Riesenskinks gefunden werden. Zwei weitere Reisen

im Frühjahr und Spätsommer 1981, mit jeweils mehrwöchigen Aufenthalten auf den Inseln Santa Lu-

zia, Branco und Razo, erbrachten ebenfalls nur negative Ergebnisse bezüglich der Nachforschungen

zur Überlebenschance dieser Großechse. Der Hinweis von Einheimischen über das Vorkommen eines

großen Skinkes (kreol.: ‚‚Chinel‘; s. SCHLEICH 1982) auf Fogo und Sao Thiago lenkte erneut die Auf-

merksamkeit des Autors auf die Suche nach Macroscincus coctei. Nach Abschluß der letzten Sammel-

reise im September/Oktober 1981 konnten nun von allen Inseln (außer den unbewohnten Felsinsel-

chen der Rhombosgruppe) Reptilien und Amphibien aufgesammelt werden, so daß vorliegende Zu-

sammenfassung ermöglicht wurde.

Die Widersprüche der einzelnen Bearbeiter zur Herpetofauna der Kapverden kamen in der Tabelle

über die Verbreitung der Reptilien auf den Kapverden (SCHLEICH 1982: 246) deutlich zur Geltung.

Nicht nur aus diesen Unklarheiten bisheriger Bearbeitungen, sondern auch im Rahmen eines inter-

nationalen Gesamtforschungsprojektes ‚‚Makaronesische Inseln“, war nun endlich die Neubearbei-

tung der kapverdischen Herpetofauna erwünscht. Bewußst wurde dabei versucht, alle Inseln selbst zu

bereisen, um über die wirkliche Herkunft der Aufsammlungen Klarheit zu haben. So basiert auch, bis

auf wenige Ausnahmen, der Hauptteil dieser Arbeit auf eigenen Aufsammlungen.

Historischer Überblick

Mit der Erstbeschreibung von Macroscincus coctei als Euprepes Coctei lieferten DUMERIL & BIBRON (1839) den

ersten herpetologischen Beitrag über die Kapverden. Weitere Grundlagen zur Kenntnis der Kapverden-Reptilien

sind durch BOCAGE (1896), ANGEL (1935, 1937) und MERTENS (1955) geschaffen worden. BANNERMAN & BAN-

NERMAN (1968) geben einen zusammenfassenden Überblick über die Herpetofaunenverteilung auf Gesamt-Maka-

ronesien wieder. GREER (1976) berichtete in einer ersten Detailstudie ausführlich über Macroscincus coctei, worauf

SCHLEICH (1979, 1980, 1982) sowie GRUBER & SCHLEICH (1982) mit weiteren Nachforschungen und Bearbeitungen

zu den Riesenechsen der Inseln Branco und Razo als letzte Bearbeiter folgten. Weitere systematische Beschreibun-

gen lieferten GRUBER & SCHLEICH (1982) und SCHLEICH (1984).

2. Allgemeiner Teil

2.1 Die Kapverdischen Inseln in Makaronesien

Nachdem in letzter Zeit von verschiedensten Autoren verstärkt Augenmerk auf die wissenschaftli-

che Bearbeitung der Kanaren gelegt wurde, stehen nun die Azoren, Madeira, Selvagens und die Kap-

verden in vergleichender Betrachtung zu ihrer Faunen- und Florenentwicklung. Während das Problem

von Migrationen und Besiedlung dieser atlantischen Inselgruppen zum Hauptdiskussionspunkt wur-

de, drängt sich natürlich die Frage nach bestandenen Festlandsverbindungen, aber auch jene der inter-

und intrainsularen Zusammenhänge auf. Besteht auf den Azoren die Kriechtierbesiedlung aus nur einer

Form (Lacerta dugesi), so weist das südlichere, festlandsnähere Madeira bereits ein Faunenspektrum

von acht Arten auf. Auf den Kanaren steigt die Artenfülle bereits auf 19 Arten an, die sich letztlich bei

den Kapverden auf nun 23 Formen beläuft.

Bemerkenswert ist ebenfalls das Verhältnis von ‚‚Kriechtier-Endemiten‘ auf den jeweiligen Insel-

gruppen. Sie stehen im Verhältnis (nach BANNERMAN & BANNERMAN, 1968, BISCHOFF, 1985 und vorlie-

gender Bearbeitung)

MARK: A RMOFNDERSCOIMERN

+50 Fer ar ar Ar 0+

50 40 30 20 10 0

+0 = ; Azoren 40

= >

Madeira.

a :

Salvages

+30 a: 30+

Ga Kanarische Inseln ’.®, 4

+20 »

R Kr)

Kapverdische 8

Inseln *

Azoren (0) : Madeira (4) : Kanaren (11) : Kapverden (21),

woraus sich für die Kapverden der höchste Endemitenanteil ergibt.

Zu berücksichtigen sind bei solchen Zahlen jedoch noch vielerlei Umstände bei der Diskussion um

die Speziationen, wie beispielsweise Landbrücken, Klima, Driftmöglichkeiten, Entfernung u. a. m.

2.2 Bemerkungen zur Geographie, Geologie und Klimatologie

Die Kapverden, seit 1975 unabhängiger, selbständiger Inselstaat, bestehend aus 10 größeren und 8

nennenswerten kleineren Inseln, liegen ca. 400-600 km westlich vor Senegal zwischen 14°48 ’ und

17°12 30’ ’ nördlicher Breite und zwischen 22°44 ’ und 25°22’ westlicher Länge zwischen dem

Wendekreis des Krebses und dem Äquator. Orographisch betrachtet lassen sich die Inseln in flache,

wüstenhafte Inseln (Sal, Boa Vista, Maio, Säo Vicente, Ilhas do Rhombo, Branco, Razo und Santa Lu-

zia) sowie in die vegetationsreicheren gebirgigeren Inseln Sao Thiago, Brava, Fogo, St. Antäo und

S. Nicolau gliedern.

In nachstehender Tabelle sind die höchsten Erhebungen (H, in m) der Inseln sowie ihre ungefähre

Flächenerstreckung (F, in qkm) in ‚,‚ca.““-Werten angegeben.

Sal Boa Vista Maıio S. Vicente Rhombos Branco Razo

lol 406 390 436 774 96 327 164

F 200 620 250 220 3 7

St. Luzia S. Thiago Fogo Brava St. Antäo S. Nicolau

H 395 1392 2829 976 1979 1304

F 16 1000 500 65 730 350

Ein allgemeiner Abrif3 zum Vegetationsbild der Inseln wurde von Login und GroH (1979, 1980) und

von SCHLEICH & WUTTKE (1983) für die kleinen Eilande St. Luzia, Branco und Razo gegeben. Die geo-

logischen Fakten zur Entstehung des Archipels bzw. der einzelnen Inseln sind noch relativ undurch-

sichtig. Verallgemeinert kann gesagt werden, daß der gesamte Archipel atlantischen Ursprungs ıst. Die

vulkanische Entstehung der Inseln im Tertiär, wahrscheinlich Miozän, kann angenommen werden;

eine Festlandsverbindung zum afrıkanischen Kontinent, auch in jüngeren geologischen Epochen, er-

scheint aber aufgrund der bis zu 3000 m tiefen Beckenregionen zwischen dem Archipel und dem Fest-

land nıe bestanden zu haben. Daß viele Inseln noch bis in die Quartärzeit (Fogo bis rezent) vulkanisch

aktıv waren, ist durch die unverwitterten vulkanischen Formationen zu erwarten.

Einen Eindruck zur Klimatographie der Kapverden sollen folgende Angaben liefern:

Klimatologisch können die Kapverden als gemäßigt ozeanisch, aber mit einem sehr trockenen Klima

betrachtet werden. Die Regenzeit fällt dabei in die heißeste Jahreszeit während der Monate Juli bis Ok-

tober, während die kälteste Jahreszeit auf die Monate November bis Dezember kommt.

Aus den Klimatabellen des Meteorological Office sind für Mindelho ($. Vicente) und für Praia

(S. Thiago) folgende Werte zu entnehmen:

Mindelho für die Zeitperiode von 1892 bis 1925:

tägl. Max. Temp.: 25 °C; monatl. © Max. Temp.:30 °C

tägl. Min. Temp.: 20,6°C; monatl. © Min. Temp.: 14,4°C

bei einer durchschnittlichen monatlichen Niederschlagsmenge von 99,1 mm.

Praia für die Zeitperiode von 1904 bis 1930:

tägl. Max. Temp.: 27,2°C; monatl. @ Max. Temp.: 32,2°C

tägl. Min. Temp.: 22,2°C; monatl. @ Min. Temp.: 17,2°C

bei einer durchschnittlichen monatlichen Niederschlagsmenge von 259,1 mm.

nl DER

Ss „Grande. a

l S SANTO ANTAO

2 Pt. Novo \\ BR

Mindelho SAO VICENTE = %gr

Io, OS

Blava r

"ISANTA zRS BRANCı

| SAO NIcoLAU

_ ee

VN.Cintra fg S.Filipe .. Re a ur 2

25° BRAVA .. nn... er ei 23°

Nach dem Annuario Climatolögico de Portugal (1973, Vol. 27) können folgende jährliche Gesamt-

niederschlagsmengen für 1973 wiedergegeben werden:

$. Vicente (Mindelho) —

jährlicher Niederschlag: 1012,8 mm

mittl. Lufttemp. (°C): 21,4 (2 Min.); 30,2 (Max.) -6. Sept.

17.82 Vin.) 20. eb.

Sal -

jährlicher Niederschlag: 1008,6 mm

mittl. Lufttemp. (°C): 20,1 (® Min.); 31,4 (Max.)- 8. Sept.

26,2 (© Max.)

15,4 (Min.).— 16. Feb.

S. Thiago (S. Jorge dos Orgaos) —

mittl. Lufttemp. (°C): 18,3 (® Min.); 12,5 (Min.) - 21., 25. Jan.

Fogo (Monte Velha) —

mittl. Lufttemp. (°C): 11,0 (Min.);— 23. Jan.

Fogo (S. Felipe) —

mittl. Lufttemp. (°C): 25,8 (® Min.); 29,0 (Max.)

33,5 (Max.) - 8. Okt.

18,0 (Min.) — 6. Feb.

Nachstehende Tabelle (aus Sasına, A. A., 1977**) gibt die „‚Krisenjahre‘“ der Inseln mit dem Aus-

bleiben der Regenzeit an:

mit ljähriger Dauer mit 2-3jähriger Dauer Jahre mit Regen Jahre ohne Regen

(partiell) (generell)

1719 1748-1750 1775 1718

1754 1773-1775 1778 1743

1764 1789-1791 1855 1753

1814 1811 1864 1763

1825 1831-1833 1884 1772

1845 1864-1866 1886 1788

1850 1885 1887 1810

1875 1901-1903 1906 1813

1889 1921 1918 1824

1896-1900 1941 1919 1831

1946-1948 1927 1845

1951-1952 1928 1850

1959-1960 1938 1854

1962-1965 1943 1858

1966-1967 1950 1863

1968-1976 1951 1874

1952 1883

1953 1896

1954 1902

1955 1920

1956 1932

1957 1940

1958 1941

1946

1947

1949

2.3 Material und Methodik zZ

Das gesammelte Material verteilt sich auf die jeweiligen Inseln wie folgt:

Insel Mabuya Tarentola Hemidactylus

Exemplare Exemplare Exemplare

Sal 9 — 7

Boa Vista 4 - 23

Maio 30 9 -

S. Thiago 47 9 2

Ilheu St. Maria 11 - -

Fogo 6 11 4

Brava 4 — =

St. Antäo 9 17 =

S. Vicente 3 11 4 (2)*

St. Luzia 9 10 _

Branco 5 18 =

Razo 14 27, 7,

S. Nicolau 7. 11 5

> 158 123 47

* (BMNH, Hemidactylus bouvieri bouvieri)

** Eingesehen am Landwirtschaftsministerium Praia/CABOVERDE.

Auf verschiedenen Inseln oder insbesondere von Lokalitäten mit schwach oder aber schützenswert erscheinen-

den Populationen wurden aus Gründen des Artenschutzes,keine Tiere oder, falls unumgänglich, nur eine geringe

Anzahl gefangen.

Die Bearbeitung beinhaltet neben der rein deskriptiven Darstellung Abbildungen zu den Zeichnungsmustern

bzw. Färbungsvarianten jeweiliger Formen bzw. sind diese in den Tafeln dargestellt. Für die umfangreichen Unter-

suchungen der Gattungen Mabuya und Tarentola wurden bei ersterer 8 und bei letzterer 12 Parameter zur allome-

trischen Streckenerfassung gemessen. Soweit Tiere zur Beobachtung im Terrarium gehalten wurden, entstammten

die Maßangaben von diesen lebenden Exemplaren, die nun von den konservierten Tieren verschieden sein können.

Die Allometrieangaben beinhalten folgende Meßstrecken:

GL: Gesamtlänge = Schnauze-Schwanzspitze (KR + SL)

SL: Schwanzlänge = Strecke Kloakenspalt-Schwanzspitze

KR: Kopfrumpflänge = Strecke Schnauzenspitze-Kloakenspalt

KL: Kopflänge = Strecke Schnauzenspitze-post. Nuchaliagrenze

KB: Kopfbreite = Strecke über den Augen

A: Augdurchmesser

O: _ Ohrdurchmesser

OA: Strecke anteriorer Ohr-posteriorer Augrand

AS: Strecke Auge-Schnauzenspitze

MSP: Maulspaltlänge von Schnauzenspitze bis Mundwinkel

ML: Mentale-Länge

MB: Mentale-Breite (mittlere)

Anhand der gemessenen und auf Lochkarten gespeicherten Parameter wurden am Leibniz-Rechenzentrum

(LRZ) München nach dem angebotenen Statistik-Programm-System SPSS 8 (NIE, N. H. & Hurt, L. K., 1980)

„„Scattergrams“ erstellt und daraus sowie über die Statistikprozedur ‚‚Pearson Correlation“ die notwendigen stati-

stischen Berechnungen!) gewonnen. Die dazugehörigen Reinzeichnungen der Diagramme!) wurden ebenfalls am

selbigen Institut gefahren. Für die kollegiale Unterstützung bei der Durchführung dieser Arbeiten möchte ich

Herrn W. SPIEGEL (München) herzlichst danken.

Die ın den Diagrammen und variationsstatistischen Angaben verwendeten Abkürzungen lauten:

Gliederung zu den Diagrammen allometrischer Meßwerte an kapverdischen Skinken der Gattung Mabuya -

Dargestellt wurden je 9 Diagramme mit folgenden Parametern:

GL/KR = Gesamtlänge : Kopfrumpflänge

SL/KR = Schwanzlänge : Kopfrumpflänge

KL/KR = Kopflänge : Kopfrumpflänge

A/KL = Augdurchmesser : Kopflänge

OA/KL = Strecke anteriorer Ohrrand-posteriorer Augrand : Kopflänge

A/O = Augdurchmesser : Ohrdurchmesser

KB/KL = Kopfbreite : Kopflänge

O/KL = Ohrdurchmesser : Kopflänge

A/OA = Augdurchmesser : Strecke OA

Gliederung zu den Diagrammen allometrischer Meßwerte an kapverdischen Geckos der Gattung Tarentola —

Dargestellt werden je 12 Diagramme mit folgenden Parametern:

GL/KR = Gesamtlänge : Kopfrumpflänge

SL/KR = Schwanzlänge : Kopfrumpflänge

KL/KR = Kopflänge : Kopfrumpflänge

A/KL = Augdurchmesser : Kopflänge

OA/KL = Strecke Ohr-Auge : Kopflänge

AS/KL = Strecke Auge-Schnauzenspitze : Kopflänge

") Aus finanziellen und drucktechnischen Gründen war eine Übernahme in vorliegender Arbeit nicht mehr mög-

lich. Die Diagramme können zusammen mit den Berechnungen in der Bibliothek der ZSM eingesehen werden.

KB/KL = Kopfbreite : Kopflänge

O/KL = OÖhrdurchmesser : Kopflänge

A/O = Augdurchmesser : Ohrdurchmesser

MSP/KL = Maulspaltlänge : Kopflänge

MSP/OA = Maulspaltlänge : Strecke OA

A/OA = Augdurchmesser : Strecke OA

Diagramme zur Allometrie und variationsstatistischen Analysen.

Die variationsstatistischen Angaben sind im Anschluß an die jeweiligen Diagramme eines Verbreitungsgebietes

bzw. einer Art aufgeführt.

Ihre Abkürzungen lauten:

n = Anzahl der ausgewerteten Exemplare

R = Pearson Correlation; Grad der Beziehung zwischen 2 Variablengruppen

Corr = Korrelationskoeffizient

Intcp A = Schnittpunkt mit der y-Achse

Slope B = Steigung der Regressionsgeraden

Std. Err = geschätzter Standardfehler (= Std. Err. Est.)

Std. Err (A) = geschätzte Standardabweichung der y-Werte

Std. Err (B) = geschätzte Standardabweichung der x-Werte

MW = Mittelwert

Std.Dev. = Standardabweichung

Neben diesen statistisch verwerteten Messungen kommen noch folgende weitere Informationen zur Indikation

und Beschreibung der jeweiligen Formen hinzu:

Anzahl der Schuppen um die Rumpfmitte

Anzahl der Supra/Sublabialia (OL/UL)

Anzahl der Aurikulartuberkel

Anzahl der Subdigitallamellen

Anzahl der Streifen- bzw. Bänder des Zeichnungsmusters.

Die dritte, hinter den Inventarnummern der ZSM erfolgte Durchnumerierung diente ebenfalls den statistischen

Berechnungen bei der EDV.

3. Artenliste mit Merkmalscharakteristik und Verbreitungstabelle der kapverdischen Echsen

Scincidae:

Glatte bis schwach gekielte Schuppen, keine verbreiterten Finger mit Haftlamellen:

Nur 1 Gattung Genus Mabuya

KR bis ca. 68 mm, 42-50 Schuppenreihen, dunkle Achselocellen, rostbrauner breiter, hell gesäumter

Rückenstreifen M. delalandii

KR bis ca. 80 mm, 58-60 Schuppenreihen um die Rumpfmitte, dunkel, fast melanotisch, Dorsalıa un-

gekielt bis tricarinat; 3-6 anteriore Aurikulartuberkel M. fogoensis fogoensis

KR bis ca. 87 mm, 52-60 Schuppenreihen um die Rumpfmitte, Dorsalia ungekielt bis doppelkielig, 0°

ventrolateral deutlich rot gefärbt; 2-3 anteriore Aurikulartuberkel M. fogoensis nicolauensis

KR bis ca. 67 mm, 50-54 Rumpfschuppen, Dorsalia doppelkielig, sehr dunkel gefärbt; 3 anteriore Au-

rikulartuberkel M. fogoensis antaoensıs

KR bis max. 80 mm, meist 4246 Schuppenreihen, je einen hellen Dorsolateralstreifen

M. stangeri stangeri

KR bis max. 80 mm, 3640 Schuppenreihen, schwach tricarinat, helle Oberschenkelocellen

M. stangerı maioensis

KR bis max. 87 mm, 36-44 Rumpfschuppen, zwei- bis dreikielig. Schmaler dunkler Dorsomedian-

streifen, der von Dorsolateralstreifen gesäumt wird M. stangeri salensıs

KR bis max. 70 mm, 34-38 Schuppenreihen, Dorsalıa tricarinat, helle Schenkelocellen, Ohröffnung

oft nur halber Augendurchmesser, M. stangeri spinalıs

KR bis 122 mm, Schnauzenspitze orange, 50-56 Schuppenreihen, 3 helle Rückenstreifen

M. vaillantı

Gekkonidae:

Finger und Zehen verbreitert, mit Subdigitallamellen

Subdigitallamellen ungeteilt Genus Tarentola

Subdigitallamellen geteilt Genus Hemidactylus

Genus Tarentola

KR-Länge adulter Tiere zwischen 55 bis 72 mm, 12-18 Tuberkelreihen, Mentalialänge entspricht etwa

doppelter Mentaliabreite, 4-5 Rückenbänder Tarentola rudis

Verhältnis von Supra-/Sublabialia 9/11 bis 7/11; 16-18 Tuberkelreihen, ca. 72 mm max. KR-Länge,

4-5 Rückenbänder T. rudıs rudis

KR-Länge bis ca. 68 mm, 12-18 Tuberkelreihen, meist jedoch 14, Verhältnis von Supra-/Sublabialia

8/9 bis 7/8; 5 Rückenbänder T. rudis maioensis

KR-Länge max. 98,5 mm, 144-181 Schuppen um die Körpermitte, Verhältnis Supra-/Sublabialia

11(-9)/11(-7), 16 Tuberkelreihen, 4-5 Rückenbänder T. rudis protogigas

Beschuppung ‚‚samtig“, perlgrau, 14-18, meist 16 Tuberkelreihen; bis ca. 54 mm KR-Länge, Supra-/

Sublabialia 8/13 bis 7/10; 4, meist 5 Transversalbänder Tarentola darwini

KR-Länge max. bis 60 mm, 14-20 relativ kräftige Tuberkelreihen, 3-6 Rückenbänder, Mentale länger

als seine halbe Breite Tarentola caboverdiana

KR-Länge max. bis 48 mm, 14-16 Tuberkelreihen (meist 16!), 5(4)-6 Bänder, meist 5.

T. caboverdiana caboverdiana

KR-Länge max. bis ca. 54 mm, 14 bis meist 18 Tuberkelreihen, 4-6, meist 5 Transversalbänder

T. caboverdiana nicolauensis

KR-Länge max. bis ca. 53 mm, 16(-18) Tuberkelreihen, 3—4 Transversalbänder

T. caboverdiana razıana

KR-Länge max. 60 mm, 146-167 Schuppen um die Körpermitte, Vorderbeine kürzer als Kopflänge,

Verhältnis Supra-/Sublabialia 11(-8)/9(-7), 14-20 Tuberkelreihen, 4-5 Rückenbänder

T. caboverdiana substituta

Kräftig gackernde Stimme, massige große Tiere, KR bis ca. 135 mm, Mentalelänge mehr als doppelte

Breite Tarentola gıgas

KR-Länge bis ca. 135 mm T. gigas gigas

KR-Länge bis ca. 114 mm T. gigas brancoensis

Genus Hemidactylus

Rosa bis graubraun mit warzigen Tuberkeln Hemidactylus brooki angulatus

Einfarbig bis bunt gezeichnet, glatte Schuppen Hemidactylus bouvieri

Vorkommen

REPTILIA - SCINCIDAE EISSTLLLLE

Mabuya delalandii

| dogoensAÄs

fogoensÄis $ogoensis

gogoensis antaovensÄd

gogoensis nicolauensid

stangeni

stangeni sLangeni

stangeri masoensis

stangeni salensÄs

stangeni spinalis

vailkanti Man IE RN EI EEE

S.Vicente

Branco

9, Nicola

Boa Vista

9, Mage

Saar

Rhombos

REPTILIA - GEKKONIDAE

Tarnentola caboverdiana

Tarentola cabovendiana caboverdiana

Tarentola caboverdiana nicolauensis

Tarentola caboverdiana raziana

Tarentola caboverdiana Aubstituta

Tarnentola darnwini

Tarnentola

Tarentola gÄ gigas

Tanentola gÄ brancoensis

Tarentola

Tanentola (3 nudis

Tanentola (34 maLoendis

Tarentola (4 protogigas

HemidactyLus bouvieni

Hemidacty£Lus bouvierni bouvieri

HemidactyLus bouvieri boavistensis

HemidactyLus bouvieri nazoensis

@ - in vorliegender Arbeit behandelt u. nachgewiesen (s.a.SCHLEICH, 1982)

Vorkommen wahrscheinlich oder von vorhergehenden Autoren angeführt;

als Neunachweis bislang noch nicht bestätigt

il]

KR-Länge bis ca. 38 mm H. bouvieri bouvieri

KR-Länge bis ca. 50 mm H. bouvieri boavistensis

KR-Länge bis ca. 29 mm H. bouvieri razoensis

4. Systematischer Teil

4.1 Scincidae - Genus Mabunya

Nach intensiven Recherchen und Nachforschungen zur Überlebensfrage von Macroscincus coctei,

dem kapverdischen Riesenskink, wird dieser von mir (s. SCHLEICH 1982 b) als ausgestorben betrachtet.

Die einzig verbleibende Skinkgattung Mabuya kommt nach meinen Untersuchungen mit4 Arten und

9 Unterarten/Arten vor.

4.1.1 Mabuya delalandıi (Dumeril & Bibron, 1839). (Taf. I, Fig. 1)

Material:

4 Ex. ZSM 367/1978; Brava. 7 Ex. ZSM 5/1982; S. Thiago-Praia, Flughafen. 12 Ex. ZSM 7/1982; S. Thiago-Praia,

Stadt. 4 Ex. ZSM 369/1978; S. Thiago-C. Velha. 1 Ex. ZSM 154/1981;S. Thiago-Tarrafal.2 Ex. ZSM 157/1981; S.

Thiago-St. Cruz. 2 Ex. ZSM 8/1982; Ilheu St. Maria (S. Thiago). 4 Ex. ZSM 11/1982; Fogo-S. Felipe. 3 Ex. ZSM

373/1978; Boa Vista. 5 Ex. ZSM 155/1981; S. Thiago — 5 km S.-Tarrafal.

1839 Euprepis delalandii DUMERIL & BIBRON

1845 Euprepis delalandii. — GRAY

1845 Euprepis belcheri. — GRAY

1857 Euprepis venustus. — GIRARD

1867 Euprepis delalandü. — BOCAGE

1869 Euprepis delalandii. — PETERS

1875 Euprepis delalandi. — BOCAGE

18837 Mabuya vaillanti. - BOULENGER (non del.)

1935 Mabuya delalandei. — ANGEL

1937 Mabuya vaillanti. — ANGEL (non del.)

1951 Mabuya delalandei. — DEKEYSER & VILLIERS

1955 Mabuya delalandıi. - MERTENS

1976 Mabuya delalandı. — GREER

1982b Mabuya delalandii. — SCHLEICH

Terra typica: Cap de Bonne Esperance

Bisherige Verbreitung: S. Thiago, Fogo, Brava, Rhombos

Nachgewiesen von: S. Thiago, Ilheu St. Maria, Fogo, Brava, Boa Vista

Artcharakteristika nach:

MERTENS (1955) —

14 Exemplare, von verschiedenen Inseln; (er bezeichnet seine 6 Stücke von Praia als Topotypen).

Frontoparietalia sowie Parietalia mit Interparietalia zu einem Schild verschmolzen. 5. (6.) Supralabiale ist eigent-

lich Suboculare. Zwischen 48 und 52 Rückenschuppen, dorsal scharf dreikielig, heller Dorsolateralstreifen gezackt

abgegrenzt.

GREER (1976) —

KR: 80 mm. Schuppenreihen um Körpermitte: 46-52

DEKEYSER & VILLIERS (1951) —

18 Exemplare; min. 48, max. 80 mm (74 mm; reg. Schwanz). MW: 67,7 mm KR-Länge. 46 Schuppenreihen um

den Körper, Verbreitung: Brava, Fogo, Rhombos, S. Thiago

ANGEL (1935) —

Frontoparietalia verwachsen, 46-50 Schuppen um die Körpermitte, 5. Labiale viel größer als 6., bildet Subokula-

re. Augdurchmesser gleich Ohrdurchmesser

Diagnose:

KR-Länge bis ca. 80 mm, SL ca. 1,6% KR. 42-50 (?52) Schuppenreihen um die Rumpfmitte.

Ohrdurchmesser gleich oder nur geringfügig kleiner als Orbit. 7-8 Supra- bei 6-7 Sublabialia. 5. Su-

pralabiale vergrößert unter dem Auge. Dorsalia 2-3 kielig, Ventralia glatt. Interparietale mit Parietale

verschmolzen; darauf Pinealforamen. Braune Rückenfärbung, mit einem Dorsalstreifen mit lateraler

Begrenzung durch helle Streifen. Ventral beige; dunkle Achselocellen.

Beschreibung:

Anmerkung

Von der vor Praia (Sao Thiago) liegenden kleinen Insel St. Maria, sowie von Brava und Boa Vista

konnte Mabuya delalandii nachgewiesen werden. Für Boa Vista ist dies ein Neunachweis.

Habitus

Der allgemeine Habitus von Mabuya delalandii ıst von kräftiger, mittelgroßer Gestalt. Während der

Kopf gut bis weniger stark vom Rumpf abgesetzt ist, ist dies beim Schwanz meist deutlich der Fall. Die

Schwanzlänge beträgt immer weniger als die anderthalbfache KR-Länge. Die Beine sind kurz, der

Schwanz rund bis dorsoventral abgeflacht. Die O'C’ sind von den PP durch ihre dickere Wangenpartie

und den so breiter erscheinenden Kopf unterscheidbar.

Pholidose

Zwischen 42 bis max. 50 Schuppen können um die Rumpfmitte gezählt werden, wenngleich sich

auch bei Populationen bestimmter Verbreitungsgebiete enger begrenzte Schuppenzahlen zu stabilisie-

ren scheinen. So konnten für Brava und Boa Vista wie auch für die kleine vor Praia liegende Insel

St. Maria bei Mabuya delalandii 44-50 Schuppen gezählt werden, wo hingegen die Tiere von

S. Thiago (5 km S-Tarrafal) 44 Schuppen aufweisen.

Generell können 8 Supra- und 6 Sublabialia ausgebildet sein, wo wiederum auch eine deutliche

Häufung von Exemplaren mit je 7 Supra- und 6 Sublabialıa auffällig ist.

Die Dorsalschuppen sind im Normalfall drei-, seltener zweikielig, die Ventralia glatt. Die Pileus-

pholidose ist neben der Färbung wohl das deutlichste Artmerkmal. Zwischen drei und fünf vordere

Ohrrandtuberkel sind ausgebildet. Von den meist sechs Supralabialia ist das fünfte oft deutlich vergrö-

ßert und liegt direkt unter dem Auge. Eine transparente, relativ klare Palpebralschuppe ist ausgebildet.

Die Supranasalia trennen das Rostrale vom Frontonasale, welches wieder durch die Präfrontalia vom

Frontale getrennt ist. Vier Supraocularıa und bis zu 6 Supraciliaria sind ausgebildet. Nur ein, meist tra-

pezförmiges, Frontoparietale ist typisch. Das Interparietale ist mit den Parietalia zu einer großen

halbmondförmigen Schuppe verschmolzen, worauf ein Pinealforamen ausgebildet ist. Zwei Nuchalıa

begrenzen caudal die Pileuspholidose.

Färbung

Die Tiere sind dorsal hell- bis kupferbraun, ventral cremefarben bis gelbbraun. Unter den Achseln

sind typische dunkle Achselocellen ausgebildet. Die Zeichnung besteht fast ausnahmslos aus dem brei-

ten unıfarbenen Rückenlängsstreifen, der dorsolateral von zwei hellen, weißlichen Streifen begrenzt

ist. Diese beginnen über den Augen und ziehen bis zur Schwanzbasis, meist von einer dunkelbraunen

Punktreihe gesäumt. Lateral schließt ein dunkler — meist dunkler als die Rückenfärbung -Flankenstrei-

fen an, der vom Rostrale bis zur Schwanzbasis zieht. Ventrolateral kann daran noch ein weißlicher

Streifen folgen, meist aber schließt direkt an den dunklen Flankenstreifen eine dunkelbraune Tüpfe-

lung an, die sich auch auf den Kehlbereich ausdehnen kann. Die Ventralseite ist bis auf eben den bei ei-

nigen Individuen getüpfelten Kehlbereich einfarbig hell.

Tafel I

Tafel 1: Fig. 1: Mabuya delalandiüi von S. Thiago. Fig. 2: Mabuya vaillanti von S. Thiago. Fig. 3: Mabuya stangeri

spinalis von S. Thiago. Fig. 4: Mabuya stangeri salensis von Sal. Fig. 5: Mabuya stangeri maioensis von Maio.

Fig. 6: Mabuya fogoensis fogoensis von Fogo. Fig. 7: Mabuya fogoensis nicolauensis von S. Nicolau. Fig. 8: Ma-

buya stangeri stangeri von Branco.

17.

Maßtabelle

Mabuya delalandii von S. THIAGO - Praia, Flughafen

Nr. ZSM GL KR SL KL KB [(@) A OA OL/UL Schuppen

3827. 114,0 40,0 74,0 8,0 540 1,6 DD. 4,0 7/7 43

5.82.6 146,0 56,0 90,0 1255 755 2,6 3,0 4,1 8/7 44

5.82.5 152,0 54,0 98,0 11-3) 6,1 1077 DT, 5,0 7/6 43

5.82.4 179,0 78,0 101,0 14,0 8,4 11,55) 353 6,5 8/7 42

5.82.3 178,0 73,0 105,0 1550 8,6 1,8 387, 6,0 7/6 42

5.82.2 192,0 78,0 114,0 1545 10,0 1,6 3,6 6,5 7/6 42

en 185,0 75,0 110,0 14,8 11,0 DR, 3,0 7,0 7/6 42

— Praia, Stadt

Has 108,0 40,0 68,0 9,8 58 0,0 0,0 0,0 _ _

7.82.11 116,0 BAU 63,0 11,4 6,3 5) 3,0 4,2 7/6 46

7.82.10 148,0 51,0 97,0 11,4 6,0 155 255 4,0 7/6 _

2) 151,0 55,0 _ 13 6,6 0,0 0,0 0,0 _ _

7.82.8 _ 76,0 _ 14,6 985 20 5 6,7 7/6 46

182.7: _ 80,0 - 17,0 11,0 159 BZ 6,5 7/6 46

7.82.6 _ 68,0 _ 127 725 2il DR2 555 7/6 46

7.82.5 165,0 60,0 105,0 12,8 72 20) 2,5 359 6/6 48

7.82.4 175,0 64,0 111,0 l.®, 8,0 2AO 255 53 7/7 47

7.82.3 _ 72,0 _ 14,4 10,2 2,4 353 6,0 ZT 47

TRS282 _ 72,0 = 14,7 10,3 2] 355 5,6 7/7 48

SZ - 80,0 _ 14,5 NZ 23 3,5 6,1 8/7 48

— Cidade Velha

369.78.3 130,0 52,0 78,0 10155 7,0 1,8 2,5 4,5 ZT. 42

369.78.2 99,0 44,0 55,0 10,0 555 152 253 Sl 7/6 44

369.78.1 135,0 54,0 81,0 12,0 6,8 185 23 4,0 7/7 42

— Tarrafal

154.81.1 TBeo re O5 11,5 3.0 ro 7/7 44

— 5 km S-Tarrafal

155.81.5 = 75,0 4 Baier 5,8 717 44

155.81.4 eos 118 25 30.68 7/7 44

155.81.3 iss 770 111.0, 155° or ae 7/6 44

155.81.2 2000 ao 125,0 70149 95. Bor De 7/7 44

155.81.1 = 68,0 E ee 5,2 7/6 44

— St. Cruz

157.81.2 = 78,0 % TEIOR, M1IkOs a; zi7 44

157.31@1 173,0 ZE\-{0) 102,0 1555 90 255 332 5,8 7/6 46

von ILHEU ST. MARIA

8.82.2 a 68,0 2 len oh oe 7/6 44

8.82.1 eo 720 O1 loan en 7/6 48

von FOGO

11.82.4 120,0 48,0 72,0 ılao) 6,0 1,8 v0) 4,2 8/7 46

11.82.3 120,0 50,0 70,0 11,8 6,0 1,6 220) 4,0 7/6 46

10822 116,0 5240 64,0 1055 6,3 2,0 28 4,1 7/6 50

11.82.1 135,0 51,0 84,0 11,0 6,3 1,8 2,0 4,5 8/7 46

von BRAVA

367.78.4 (114,0) 46,0 ( 68,0) 10,5 5,8 11577 20) 3,7 7/6 46

367.78.3 _ 72,0 - 13,5 110,0) 2,1l 2,3 6,2 8/7 46

367.78.2 _ 75,0 _ 14,6 82 2,0 Al 6,3 8/7 50

367.78.1 _ 80,0 _ 1555 12,8 3,0 3,0 7,0 7/6 48

von BOA VISTA

373.78.3 110,0 46,0 64,0 10,0 6,5 1,4 2,0 3,9 8/7 42

373.78.2 _ 43,0 - 10,2 6,0 1,5 AN 4,0 8/7 42

373.78.1 = 58,0 _ 13,0 8,0 1,8 — 4,9 8/7 44

4.1.2 Mabuya fogoensis (O’Shaugnessy, 1874)

1874 Euprepes fogoensis O’SHAUGNESSY

1887 Mabuya fogoensis. - BOULENGER

1935 Mabuya fogoensis. - ANGEL

1937 Mabuya fogoensis. — ANGEL

1951 Mabuya fogoensis. — DEKEYSER & VILLIERS

1955 Mabuya fogoensis. -— MERTENS

1976 Mabuya fogoensis. — GREER

1982 Mabuya fogoensis. - SCHLEICH

Terra typıca: Fogo, Sao Vicente

Bisherige Verbreitung: Fogo, St. Antäo, Sao Vicente, San Nicolau

Nachgewiesen von: Fogo, San Nicolau, St. Antao

Artcharakteristika nach:

MERTENS (1955) —

60-66 Schuppenreihen, Schuppen dorsal zweikielig bis undeutlich dreikielig; Neigung zum Melanısmus; Kopf-

unter-, Bauchseite dunkelgrau bis schwarz gefleckt. KR: 66 mm, SL: 95 mm

GREER (1976) —

KR: 78 mm, 60-66 Schuppenreihen

DEKEYSER & VILLIERS (1951) —

46 Exemplare; KR max: 81 mm, min. 54 mm; KR - MW: 67,2 mm, zwischen 53 und 68 Schuppenreihen

4.1.2.1 Mabuya fogoensis fogoensis (O’Shaugnessy, 1874) (Taf. I, Fig. 6)

Material: 4 Exemplare BMNH 1946 8.18.8-11.

Terra typıca: Fogo

Bisherige Verbreitung: Fogo

Nachgewiesen von: Fogo

Diagnose:

KR-Länge ca. 80 mm, Schwanzlänge ca. 1,1X KR-Länge. 58-60 Schuppenreihen um die Rumpf-

mitte; Dorsalia ungekielt bis schwach dreikielig; 7 Supra- und 6 Sublabialia;4 Supraocularıa. 3-6 ante-

rıore Aurikulartuberkel. Dunkelbraun mit Tüpfelung oder Flankenstreifen.

Beschreibung:

Habitus

Die Skinke sind von lacertiformem Habitus mit abgesetztem Kopf und rundem Schwanz. Die KR-

Länge beträgt max. ca. 8 cm bei 17 bis 18 cm Gesamtlänge. Die Ohröffnung ist etwa gleich groß wie

der Augdurchmesser.

Pholidose

58-60 Schuppen werden um die Rumpfmitte gezählt. Die Dorsalschuppen sind ungekielt bis

schwach dreikielig, die Ventralia glatt. 8-10 vergrößerte Präanalschuppen und je 7 Supra- und Subla-

bialia (1X 6), sowie 3-6 sehr kleine, vordere Ohrrandschuppen sind ausgebildet. Das5. Supralabiale ist

vergrößert und liegt direkt unter dem Auge. Die transparente Palpebralscheibe ist relativ groß. 4 Su-

praocularıa und 6 Supraciliaria können gezählt werden. Das Interparietale mit einem schwach sichtba-

ren Pinealforamen ist deutlich kleiner als die beiden Frontoparietalia. Die Parietalia sind sehr groß und

umgreifen fast das Interparietale. Posterocranial begrenzen zwei Nuchalia die Pileuspholidose.

Färbung

Typisch für die 4 Alkohol-Exemplare aus der Sammlung des BMNH ist eine dunkelbraune Tüpfe-

lung auf hellbraunem Grund. Dabei sind die Flecken dorsal in2 Doppeltüpfelreihen, die meist einen

hellen kleineren Mittel- oder Randfleck aufweisen, angeordnet. Dorsolateral tritt dieses gleiche Zeich-

nungsmuster — in einer intensivierten Dunkelfärbung zuweilen auf einem dunkleren Farbrand — das

vom Auge bis zum Schenkelansatz reicht, auf. Die Flanken sind gesprenkelt, der Kopf gelblich braun.

Maßtabelle

Mabuya fogoensis fogoensis von FOGO

BMNH-Inv. Nr. GL KR SL ISIE KB (6) A OA OL/UL Schuppen

1946.8.18.8 173,0 80,0 93,0 16,5 11,2 2,2/0 3 6,8 7/6 60

1946.8.18.9 173,0 76,0 97,0 15,5 11,0 2,6/4 3,0 6,6 7/6 58

1946.8.18.10 — 66,0 z 14,0 9,5 2,0/3 259 3,2 7/6 58

1946.8.18.11 108,0 58,0 50,0 13,0 8,0 2,0/2 2,6 >51 7/6 58

4.1.2.2 Mabuya fogoensis nicolauensis nov. ssp. (Taf. I, Fig. 7)

Material: 7 Exemplare; Holotypus — ZSM 1.82.1, S. Nicolau. Paratypen — ZSM 1.82.2-7.

Terra typica: $. Nicolau

Verbreitung: $. Nicolau

Derivatio nominis: nicolauensis, von S. Nicolau

Diagnose:

KR-Länge bis 87 mm; zwischen 52-60 Schuppen um die Körpermitte. In der Regel 7 Supralabialıa.

2-3 anteriore Aurikulartuberkel. Dorsalia ungekielt bis doppelkielig; Ventralia glatt. Bei O’O’ auffäl-

lige Rotfärbung der Bauchseite.

Beschreibung des Holotypus:

Der Holotypus ist das größte Individuum der gefangenen Serie von sieben Exemplaren. Seine Ge-

samtlänge beträgt 16,4 cm, wobei 8,7 cm auf die Kopf-Rumpflänge entfallen; 5,5 cm der Schwanz-

länge sind regeneriert. Die Kopflänge mißt 17,5 mm bei 13,0 mm Breite. Ohr- und Augdurchmesser

sind mit je 3,1 mm gleich groß. Sieben Supra- und Sublabialia sind ausgebildet, um die Rumpfmitte

zählt man 54 Schuppen. Das Exemplar ist dorsal braun gefärbt und trägt die zwei typischen hellen

Dorsolateralstreifen, die zu den Flanken hin mit einer dunklen Punktreihe gesäumt sind. Die hellen

Streifen reichen vom hinteren Augenwinkel bis auf den Schwanz. Die mittlere linke Zehe ist kupiert,

ebenfalls der linke Hemipenis. Ventral ist das Tier ab den Vorderextremitäten rötlich beige gefärbt mit

einer lateralen Rotintensivierung.

Beschreibung:

Habitus

Der Habitus der max. bis 87 mm (KR) lang werdenden Tiere ist von typisch lacertider Form mit vom

Rumpf abgesetzten Kopf und Schwanz. Insbesondere bei den adultenoO° ist deutlich die Wangen-

region verdickt. Der Schwanz ist dorsolateral rundlich und mit basaler Abflachung.

Abb. 1-5: Schematisierte Rückenzeichnungen kapverdischer Skinke der Gattung Mabuya. 1) Mabuya dela-

landii; 2) Mabuya fogoensis fogoensis; 3) Mabuya fogoensis nicolanensis; 4a, b,) Mabuya fogoensis antaoensıis;

5) Mabuya stangeri stangenı.

Pholidose

52-60 Schuppen können um die Rumpfmitte gezählt werden. Von den Lippenschildern sind mit je

einer Ausnahme (1X8 Supralabialia, 1x6 Sublabialia) 7 ausgebildet.

Das Augenlid bedeckt eine relativ große, klare Palpebralscheibe. Den vorderen Ohrrand zieren

meist nur 2-3 kleine Spinalschuppen. Zwei Supranasalia trennen das Rostrale vom Frontonasale. Die

Frontoparietalia sind nur wenig größer als das Interparietale. Vier Supraocularia sind vorhanden. Zwei

Nuchalia begrenzen caudal die Schädelpholidose. Die Dorsalschuppen weisen, falls überhaupt er-

kennbar, nur eine schwache Doppelkielung auf, die Ventralia sind glatt.

Färbung

Vier der konservierten Exemplare zeigen eine deutlich orange-rote Kehl- und Bauchfärbung. Mögli-

cherweise handelt es sich bei dieser Färbung um einen Geschlechtsdimorphismus, da alle vier Exem-

plare ausgestülpte Hemipenes zeigen, während die drei verbleibenden kleineren Exemplare ventral nur

gräulich gefärbt sind. Alle Exemplare zeigen zwei helle Lateralstreifen, die zur Rückenmitte von dun-

kelbraunen Tupfen gesäumt werden und an den Flanken von einer feinen, abwechselnd dunkelbrau-

nen, hellbeigen Tüpfelung begrenzt sind. Helle Tüpfel zieren die Caudalseiten der Oberarme. Die

Rückengrundfärbung ist mitteldunkelbraun, die Flecken sind graubraun.

Maßtabelle

Mabuya fogoensis nicolanensis nov. ssp. von $S. NICOLAU

Nr. ZSM GL KR SL KL KB (®) A OA OL/UL Schuppen

12822] - 87,0 _ 1743 1550 I! Sl Te NT. 54

1822 164,0 72V 92,0 14,0 11,4 2,6 2,8 6,2 7/7 54

823 15940 64,0 95,0 IB 8,7 2,4 2,6 5385 7/7 60

1.82.4 147,0 58,0 89,0 155 7,0) 1,8 2>2 4,7 7/6 52

1.82.5 121,0 49,0 72,0 10,8 6,5 220) 3,0 4,1 ZT, 52

1.82.6 94,0 42,0 52,0 10,0 5330 12,0 S)-0) Be 7/7 54

1982R72 —_ 5220 _ 14163 Tail 2,0 0,0 4,2 8/7 52

BMNH-Inv. Nr.

1906.3.30.36 160,0 72,0 80,0 15,0 11,0 2,5/3 3,0 6,0 THRTE 58

4.1.2.3 Mabuya fogoensis antaoensis nov. Ssp.

Material: 9 Exemplare; Holotypus — ZSM 23.1982.1. Paratypen — ZSM 23.1982.2-9.

Terra typica: St. Antäo

Verbreitung: St. Antao

Derivatio nominis: antaoensis, von St. Antao

Diagnose:

KR-Länge bis ca. 67 mm, SL meist ca. 1,3-1,5X KR; 50-54 Schuppen um die Rumpfmitte. In der

Regel 7 Supra- und 6 Sublabialia; 3 anteriore Aurikulartuberkel. Dorsalia doppelkielig; Ventralia

glatt.

Beschreibung des Holotypus:

Als Holotypus wurde das Exemplar mit der größten KR-Länge gewählt. Seine Gesamtlänge beträgt

13,6 cm bei 6,7 cm KR-Länge, wobei 4 cm auf das Schwanzregenerat entfallen.

Das Tier ist dunkelbraun und erscheint am Rücken nur sehr schwach gezeichnet. Lateroventral löst

sich die Färbung in eine unregelmäßige Fleckenreihe auf; dieselbe Fleckung erstreckt sich auch auf den

gesamten Kehlbereich. Die Bauchmitte sowie die Extremitätenunterseite sind hellbeige/grau gefärbt.

Auf dem heller gefärbten Schwanzregenerat wird die Streifung wieder deutlicher. Weitere allometri-

sche Werte sind nachfolgender Maßtabelle zu entnehmen.

Beschreibung:

Habitus

Das von der Ostseite von St. Antäo stammende Material umfaßt eine kleine Serie von 9 Skinken ju-

veniler bis adulter Tiere. Ihrer phänotypischen Erscheinung nach gleichen sie sehr denen von $. Vicen-

te. Mit 67 mm KR-Länge erscheint das größte ©’ vom Körperbau her bedeutend gedrungener als ein

vergleichbares Exemplar von S. Nicolau. Der Kopf erscheint kürzer als bei besagter Form und ist

ebenso wie der Schwanz nur undeutlich vom Rumpf abgesetzt. Der Körper ist ziemlich gleichförmig,

der Schwanz rund.

Pholidose

Zwischen 50 und 54 Schuppen können um die Rumpfmitte gezählt werden. 7 Supra- und 6 (eine

Ausnahme mit 7) Sublabialia sind ausgebildet. Das Augenlid besitzt eine klare Palpebralscheibe; am

vorderen Ohrrand sind 3 Spinalschuppen vorhanden. Die Dorsalschuppen sind doppelkielig, die Ven-

tralia glatt. Die Supranasalia trennen das Rostrale vom Frontonasale. Die Zahl der Supraocularschilder

beträgt 4. Die Frontoparietalia sind nur geringfügig größer als das Interparietale. Die Pileuspholidose

wird durch zwei Nuchalia begrenzt.

Färbung

Die Tiere sind von tief dunkelbrauner Grundfärbung am Rücken und den Flanken, die Bauchseite ist

dunkel graubraun.

Bis auf ein Exemplar ist der Mittelstreifen nur als dünner heller Strich, zuweilen ganz als Strichlinie

ausgebildet. Seitlich davon liegen ebenfalls in mehr oder wenig deutlicher Ausbildung noch je zwei

feine helle Streifen, die oft auch nur als Punktlinien angedeutet sein können. Die Flanken sind leicht

hell gesprenkelt. Der Übergang zur helleren Bauchfärbung erfolgt meist erst auf der Ventralseite und

nicht scharf begrenzt. Die Kehle erscheint ebenfalls deutlich marmoriert.

Maßtabelle

Mabuya fogoensis antaoensis nov. ssp. von ST. ANTAO

Nr. ZSM GL KR SL KL KB (6) A OA OL/UL Schuppen

23.82.9 122,0 47,0 75,0 10,5 6,5 1,6 2,0 353 7/6 50

23.82.8 - 42,0 — 8,8 5,5 — — — — —

23.82.7 102,0 38,0 64,0 9,0 553 _ — _ — —

23.82.6 100,0 43,0 57,0 10,0 552 — — = = —

23.82.5 _ 53,0 — 12,0 753 DEP. 2,6 5,0 7/6 50

23.82.4 104,0 42,0 62,0 10,2 6,5 1,9 2,2 4,0 7/6 54

23.82.3 100,0 43,0 57,0 10,0 6,5 2,0 2,0 3,8 7/6 52

23.82.2 — 63,0 _ 1255 8,2 23 2,5 4,0 7/6 50

23.82.1 — 67,0 _ 1855 2) 2,0 255 559 7/7 54

4.1.3 Mabuya stangeri

4.1.3.1 Mabuya stangeri stangeri (Gray, 1845) (Taf. I, Fig. 8)

Material: 3 Ex. ZSM 159/81; S. Vicente. 3 Ex. ZSM 15/82; Branco. 3 Ex. ZSM 14/82; Branco. 14 Ex. ZSM 3/82;

Razo. 1 Ex. ZSM 13/82; Razo. 2 Ex. ZSM 12/82; Razo. 11 Ex. ZSM 2/82; St. Luzia.

1845 Euprepis stangerı GRAY

1869 Euprepis polylepis. — PETERS

1875 Euprepis hopfferi. - BOCAGE

1887 Mabuya stangeri. — BOULENGER

1937 Mabuya stangeri. — ANGEL

1951 Mabuya stangeri. — DEKEYSER & VILLIERS

1955 Mabuya stangeri stangeri. — MERTENS

1976 Mabuya stangeri. — GREER

Terra typica: Westafrika

Bisherige Verbreitung: (?) Sal, (?) Boa Vista, (?) Brava, $. Vicente St. Luzia, Branco, Razo

Nachgewiesen von: $. Vicente, St. Luzia, Branco, Razo

228)

7 8 9

Abb. 6-9: Schematisierte Rückenzeichnungen kapverdischer Skinke der Gattung Mabuya. 6a-d) Mabuya stan-

geri maioensis; 7) Mabuya stangeri salensis; 8) Mabnya stangeri spinalis; 9) Mabuya vaıllantı.

Artcharakteristika nach:

MERTENS (1955) — von 54 Exemplaren:

42—46 Schuppenreihen, dreikielig; 5-7 Supraciliaria, 5-6 Labialia unter dem Auge, Praefrontale kann mit den

Frontoparietalia und das Interparietale mit dem Parietale verschmolzen sein.

GREER (1976) —

KR-Länge für M. stangeri: 90 mm. 36-46 Körperschuppen um die Rumpfmitte.

DEKEYSER & VILLIERS (1951) —

Verbreitung: $. Vicente, Branco, Razo, Sal, Boa Vista, Brava.

KR: max. 90 mm, min. 50 mm. 49-44 Schuppenreihen um den Körper, 5 Supraciliaria.

Diagnose:

KR-Länge bis max. 80 mm, SL wenig mehr als KR.

42 (1x38)46 Schuppenreihen um die Rumpfmitte, 6/5 bis max. 7/7 Supra- bzw. Sublabialia. 34 an-

teriore Aurikulartuberkel. Dorsalia tricarinat. Ventralia glatt. Ohrdurchmesser oft kleiner als Aug-

durchmesser. Dorsal dunkelbraun mit je einem hellen Dorsolateralstreifen, ventral hellgrau bis rötlich.

Beschreibung:

Habitus

Kopf und Schwanz der bis zu 160 mm lang werdenden Tiere sind deutlich vom Rumpf abgesetzt.

Der Augdurchmesser ıst meist größer oder gleich groß dem Ohrdurchmesser.

Pholidose

Zwischen 42 und 46 Schuppen können um die Rumpfmitte gezählt werden, wobei die Tiere mit

42 Schuppenreihen häufiger sind. Von den Labialıa sind meistens 7 Supra- sowie 7 Sublabialıa ausge-

bildet, wenngleich auch Exemplare mit je 6 oder 7 Supra- und 5 Sublabialia vorkommen. Der vordere

Ohrrand ist von 3 oder 4 kleinen Spinalschuppen bestanden. Die Dorsalschuppen sind schwach drei-

gekielt. Vier Supraocularıa und 5 Supraciliarıa sind ausgebildet.

Das Rostrale wird durch die Supranasalia deutlich vom Frontonasale getrennt. Das Interparietale ist

gleich groß oder nur geringfügig kleiner als die Frontoparietalia. Sowohl das 5. als auch 6. Sublabiale

liegen unter dem Auge. Zwei Nuchalia begrenzen caudal die Pileuspholidose.

Färbung

Die dorsal relativ dunkelbraun gefärbten Tiere zeigen alle beiderseits je einen helleren Dorsolateral-

streifen. Dazwischen kann eine dunkle Fleckung, teilweise mit hellen Punkten versehen, auftreten, die

auf den Flanken bei allen Exemplaren vorkommen. Die Bauchseite ist hellgrau gefärbt und bei einigen

Exemplaren (O'C") mit rötlichen Ventrolateralstreifen gezeichnet.

Maßtabelle

Mabuya stangeri stangeri von $S. VICENTE

Nr. ZSM GL KR SL KL KB OÖ A OA OL/UL Schuppen

159.81.5 _ 72,0 —_ 13,0 10,0 ZA 2,3 5,5 6/6 44

159.81.4 155,0 75,0 80,0 13,8 11,1 Dal 3,1 5,5 6/6 46

159.81.3 92,0 36,0 56,0 8,5 5,0 12 1,8 3,0 6/6 42

159.81.2 _ 56,0 _ 13,0 8,5 2,5 2,3 4,8 _ _

159.81.1 130,0 52,5 77,5 10,9 Ze 1,8 DD, 4,5 - _

von St. LUZIA

2.82.9 13820. 02 62/0) 760, 1212:5 So EL. MO 6/5 44

2.82.8 oo 3a a 0 52 hs 1,8 3,2 7/6 42

2.82.7 nano 50 Aa oo Ro 9 a0 7/5 44

2.82.6 117,0 44,0 73,0 9,1 5,5 1,3 2,5 Do 7/6 42

2.82.5 117,0 43,0 74,0 9,0 6,0 1,5 2,0 3,5 6/5 42

2.82.4 — 51,0 — 10,0 6,5 1,5 2,6 4,1 7/5 44

2.82.3 116,0 63,0 53,0 12,2 8,5 2,0 2,8 4,9 6/5 42

2.82.2 130,0 57,0 73,0 12,0 8,1 1,8 | 4,1 7/6 44

2.82.1 _ 63,0 — 12,9 9,4 1,8 za 5,0 7/6 42

von BRANCO

15.82.3 = 74,0 ® 14,5 OO 3 5,0 7/6 38

15.82.2 0 370. 220 00.0 0 = _

15.82.1 = 80,0 & DOW OO ERE oE 355 7/7 44

von ?BRANCO

14.82.1 Sole az er ae an a7 7/6 44

14.82.2 a 74,0 s 141) all 10:70, 3 6,0 7/6 44

3.82.14 133,0 65,0 68,0 1350 8,0 2,0 25H 5,0 7/6 42

3R8213 143,0 66,0 77,0 13,0 8,2 22 ZN 352 7/6 an

3282212. - 75,0 _ 14,0 9,8 11577 3,0 5,8 7/7 42

3.82.11 - 72,0 — 14,0 9,0 2,6 2,8 4,3 6/6 -

3.82.10 154,0 71,0 83,0 az 352 2,0 23 3 7/6 42

328259, _ 68,0 _ 13,0 953 2 2,8 5,3 7/7 42

3.82.8 _ 74,0 - 13,9 10,1 2 253 539 717 42

382.7, 151,0 68,0 83,0 12,8 10,2 2 29 555 7/7 42

3.82.6 — 65,0 — 13,0 9,0 2,8 25 >53 717 44

3.82.5 154,0 75,0 79,0 14,0 10,0 3,0 3,0 353 7/7 44

3.82.4 151,0 67,0 84,0 12,8 950 259 22) 553 7/7 42

3.82.3 156,0 71,0 85,0 13,0 10,0 2,0 2,8 558 717 44

BNS22 140,0 63,0 77,0 1252 8,2 2,0 253 5,0 717 46

382 _ 76,0 Z— 1554 10,0 2,0 257 5,8 717 44

4.1.3.2 Mabuya stangeri maioensis Mertens, 1955 (Taf. I, Fig. 5)

Material: 30 Ex. ZSM 160/1981; Maio. 1 Ex. ZSM 10/1982; Pt.-Cais-Maio.

1955 Mabuya stangeri maioensis MERTENS

1982 Mabuya stangeri maioensis. — SCHLEICH

31 semiadulte bis adulte Exemplare liegen von Maio vor. Während 30 Individuen aus dem Gebiet ca. 10-20 km

nördlich von Vila do Maio gefangen wurden, stammt ein juveniles Exemplar (ZSM 10/1982) von der Fischerstation

Pt. Cais im Norden der Insel.

Terra typica: Maio

Bisherige Verbreitung: Maio

Nachgewiesen von: Maio

Artcharakteristika nach:

MERTENS (1955) —

‚„‚Sehr nahestehend der spinalis-Rasse von Fogo, aber die Längsstreifen, auch der dunkle Vertebralstreifen, ver-

löschen bzw. werden durch dunkle, in Längsreihen stehende Fleckchen ersetzt.“

Anmerkung: Die Artbeschreibung von MERTENS geschah nur nach einem einzigen Exemplar. Eine Diagnose und

Beschreibung wird nachfolgend gegeben.

Diagnose:

KR-Länge bis 80 mm, SL ca. 1,2x KR. 3640 Schuppen um die Rumpfmitte. Dorsalia schwach tri-

carınat, Ventralia glatt. 4-5 vordere Ohrrandtuberkel. Dorsal graubraune Färbung dominierend; helle

Oberschenkelocellen.

Beschreibung:

Habitus

Der Gesamthabitus der Skinke ist von rundlichem Körper mit wenig bis nicht abgesetztem Kopf-

und Schwanzansatz. KR-Länge bis 80 mm, SL oftüber 1,5x KR-Länge. Augdurchmesser nur gering-

fügig kleiner oder größer als Ohrdurchmesser. Der Schwanz ist rundlich viereckig, der Kopf kurz und

gedrungen.

Pholidose

36-40 Schuppen können um die Rumpfmitte gezählt werden. Sechs bis acht, meist jedoch sieben Su-

pralabialia sowie sechs bis sieben, meist jedoch ebenfalls sieben Sublabialia sind ausgebildet. Zwischen

vier bis fünf, meist jedoch vier Spinalschuppen liegen am vorderen Ohrrand.

Eine große Palpebralscheibe bedeckt das Augenlid. Das Rostrale stößt mit den Supranasalia und dem

Frontonasale zusammen. Die beiden Frontoparietalia sind geringfügig größer als das Interparietale.

Zwei Nuchalia begrenzen caudal die Schädelpholidose. Die Dorsalia sind schwach dreigekielt, die

Ventralia glatt.

Färbung

Während ein Drittel der Maio- Aufsammlung aus dorsal mehr oder weniger einfarbig graubraun ge-

färbten Tieren besteht, scheint der Rest aus ‚‚Intergrades“ aller auf den Kapverden vorkommenden Ar-

ten zu bestehen. Die abgebildeten Zeichnungsmuster liegen innerhalb des Bereiches der auf Maio vor-

kommenden Typen. Wie bei Mabuya stangeri spinalis sind auch hier helle Oberschenkelocellen ausge-

bildet.

Eine typische Juvenilfärbung ist nicht zu erkennen, wenngleich auch die Mehrzahl der gefleckten

bzw. mit Streifen gemusterten Tiere kleinere Exemplare sind. Die Dunkelfärbung des Rückens reicht

auf Flankenhöhe ziemlich genau bis auf die Höhe der Beinoberseite bzw. des Beinansatzes. Alle Exem-

plare tragen auf der Caudalseite der Oberschenkel zwei bis drei, meist kreisrunde, helle Flecken. In

Abbildung 6.a-d ist die Variabilität der Zeichnungsvariationen von Mabuya stangeri maioensis darge-

stellt.

Anmerkung: Für Maio, bislang geologisch als älteste der kapverdischen Inseln betrachtet, läge nun

der Gedanke nahe, an eine Entwicklung oder Erstbesiedlung dieser Insel zu glauben, von wo aus radia-

tiv die verschiedenen Nachbarinseln besiedelt worden wären. Ein Vergleich aller typischen Zeich-

nungsmuster verwandter Arten könnte diese Ansicht stützen.

Maßtabelle

Mabuya stangeri maioensis von MAIO

Nr. ZSM GL KR SL KL KB (6) A OA OL/UL Schuppen

160.81.30 96,0 43,0 53,0 953 5,8 1,6 21 4,0 7/7 40

160.81.29 _ 37,0 — 9,6 7,1 1,8 ol 3,5 717 36

160.81.28 _ 45,0 85,0 10,3 6,8 2,0 253 4,0 8/7 36

160.81.27 _ 52,0 74,0 11,0 6,6 2,0 252 5,0 7/7 36

160.81.26 _ 43,0 84,0 12,0 5,4 25] 252 ) 7/7 38

160.81.25 - 51,0 79,0 12,0 6,5 2,0 253 4,5 7/7 38

160.81.24 - 59,0 — 15,0 76 BR 252. De) 717 38

160.81.23 - 56,0 - 12,0 8,1 2,3 29 5,0 8/7 36

160.81.22 154,0 56,0 98,0 12,0 763 2,0 2,4 4,7. 8/7 36

160.81.21 _ 74,0 — 13,2 9,8 255 Sal 6,0 7/7 36

160.81.20 153,0 56,0 97,0 1155 8,0 1,8 2,4 4,6 7/7 36

160.81.19 _ 71,0 — 1553 8,7 2,4 252 6,0 6/6 36

160.81.18 166,0 68,0 98,0 13,0 8,0 2,0 De) 6,0 7/7 40

160.81.17 _ 75,0 - 1357 9,0 N! 253 6,0 7/17 40

160.81.16 — 75,0 - 14,0 8,8 15%) 253 5,9 7/7 38

160.81.15 — 77,0 _ 14,8 9,0 22. 252. 6,5 ZUR 36

160.81.14 130,0 75,0 55,0 14,1 10,1 2) 2,8 6,0 7/6 38

160.81.13 — 80,0 — 15,0 9,8 349 3,0 6,0 7/6 38

160.81.12 _ 65,0 - 12,2 9,0 2,4 2,6 5,0 7/6 38

160.81.11 — 76,0 - 12,5 359 Da 2,6 552 ZUR 36

160.81.10 — 71,0 - 13,8 10,0 253 2,6 6,0 7/17 38

160.81.9 - 67,0 — 12,1 8,9 2,0 253 5,5 7/7 38

160.81.8 — 75,0 — 13,5 9,5 DRS 2,5 6,3 8/7 36

160.81.7 _ ZARO - 13,9 Do 2,6 ZN 6,0 7/7 38

160.81.6 _ 76,0 _ 14,5 10,0 2,6 2,8 6,0 7/7 38

160.81.5 175,0 80,0 95,0 1355 9,5 3,1 2,8 6,1 717 38

160.81.4 - 53,0 — 11,1 6,5 243 I, 5,0 7/7 38

160.81.3 — 75,0 _ 14,0 9,8 253 27 6,0 7/7 40

160.81.2 — 77,0 — 13,3 8,5 23 2,6 6,1 7/6 38

160.81.1 - 74,0 — 14,0 10,7 DAT 243 357. 8/7 38

4.1.3.3 Mabuya stangeri salensis Angel, 1935 (Taf. I, Fig. 4)

Material: 8 Exemplare, ZSM 4/1982; Sal. 1 Exemplar, ZSM 16/1982; Sal. 1 Exemplar, ZSM 374/1978; Boa Vista.

1935 Mabnya salensis ANGEL

1937 Mabuya salensis. - ANGEL

1951 Mabuya stangeri. — DEKEYSER & VILLIERS

1955 Mabuya stangeri salensis. - MERTENS

1982 Mabuya stangeri salensis. — SCHLEICH

Terra typica: Sal

Bisherige Verbreitung: Sal, (?) Boa Vista

Nachgewiesen von: Sal, Boa Vista

Artcharakteristika nach:

ANGEL (1935) —

Schnauze kurz, längengleich zu dem Abstand zwischen Augenhinterrand und Ohröffnung. Nasenlöcher auf der

Naht zwischen Rostrale und dem ersten oberen Labiale. Oberes Augenlid mit einer durchsichtigen Scheibe, unge-

teilt. Ein Postnasale. Vorderes ‚‚Frenale“ nicht in Kontakt mit erstem Labiale, Supranasalia posterior von Rostrale

begrenzt. Frontonasale viel breiter als lang, in Kontakt mit Frontale. Präfrontalia ohne Mediansutur. Frontale viel

länger als Frontoparietalia und Interparietale zusammen. 2. und 3. Supraokulare in Kontakt mit Frontale. 4 Su-

praokularıa nach Größen geordnet: 2.3.4. und 1.

DreiSupraciliaria, das erste gleich lang wie die beiden verbleibenden. Frontoparietalia größer als das Interparieta-

le. Parietalia mit einer kurzen Sutur an die nachfolgenden Schuppen. Ein Paar Nuchalia. Sieben obere Labialia, das

5. und 6. unter dem Auge, kaum verkleinert; Ohröffnung groß, oval, ihr Vertikaldurchmesser gleich lang wie der

Augendurchmesser, mit davorliegenden, etwas vergrößerten Schuppen.

Dorsal- und Nackenschuppen unterschiedlich, dreigekielt. 39-40 Schuppen um die Körpermitte. Glatte Finger-

lamellen. Schwanz ca. 1,2 mal der KR-Länge. 3 Supraciliarıa, 5. und 6. Labiale unter dem Auge.

Maße:

GE: 124 mm Vordergliedmaßen: 16 mm

KL: 12 mm Hintergliedmaßen: 22 mm

KB: 8 mm SL: 67 mm

KR: 45 mm

Färbung:

Dunkelgrau mit Spuren einer Rückenlinie, irregulär mit hellen, dorsolateralen Schuppen.

Reihen weißer Tupfen auf den Flanken, die ebenfalls Anzeichen zu zwei Längsreihen bilden.

Anmerkung:

Nach einem Vergleich mit ‚‚Mabuya spinalis“ gibt ANGEL (1937) für Mabnya stangeri salensis folgende Unter-

scheidungsmerkmale an: 32-40 Schuppen um die Körpermitte, 5-6 Labiale unter dem Auge, 3 Supraciliaria, vorde-

res „‚Frenale“ nicht in Kontaktmit 1. Labiale, Präfrontalia ohne Mediansutur, kürzerer Schwanz, unterschiedliche

Färbung.

MERTENS (1955) —

KR: 90 mm; SL: 119 mm; Sal.

MERTENS betrachtete seine Exemplare von Sal als Topotypen. Praefrontalia bilden untereinander eine Naht.

1. Supralabiale steht mit vorderem Loreale (1 Ausnahme) in Kontakt; ferner 4 (1 Ausnahme mit drei) Supraciliaria

ausgebildet. 39-(40)42 Schuppenreihen, Schuppen dreikielig (bei Juvenilen zweikielig!).

Diagnose:

KR-Länge bis ca. 90 mm, Gesamtlänge bis ca. 210 mm. 3644 Schuppenreihen um die Körpermitte;

zweigekielt, selten dreigekielt. 6-7 Supra-/Sublabialia. 3-6 anteriore Aurikulartuberkel. Dunkler Dor-

somedianstreifen, helle Oberschenkelocellen.

Beschreibung:

Habitus

Nach Mabuya vaillanti ist dies die zweitgrößte und auch kräftigste Mabuyenart der Kapverden. Bis

87 mm KR-Länge und 210 mm Gesamtlänge maß das größte, im Frühjahr 1981 gefangene Exemplar

von Sal. Der Schwanz mißt nie über anderthalbmal die KR-Länge. Der Kopf ist deutlicher als bei Ma-

buya vaillanti vom Rumpf abgesetzt. Der kreisrunde Schwanz verschmälert sich ab Rumpfbasis deut-

lich. Der Augdurchmesser ist deutlich größer als der Ohrdurchmesser. Meist 7 (6) Supra- und 7 (6)

Sublabialia sind ausgebildet.

Pholidose

Zwischen 36 und 42 (?44) Schuppen zählt man bei den Tieren von Sal um die Körpermitte, bei den

Exemplaren von Boa Vista 40. Die Rückenschuppen sind schwach zweigekielt, selten dreigekielt. Das

Exemplar von Boa Vista hat7 Supra- und 6 Sublabialia, ebenso ein Exemplar von Sal, wo auch ein wei-

teres mitje 6 Lippenschildern vorkam. Eine deutlich erkennbare, große Palpebralschuppe ist ausgebil-

det. Drei bis sechs (meist 5) gut sichtbare vordere Ohrrandtuberkel sind vorhanden. Sowohl das 5. als

auch 6. Supralabiale kommen unter dem Auge zu liegen.

Die Supranasalnaht zwischen Rostrale und Frontonasale ist sehr kurz. Das Nasale wird vom Ro-

strale und 1. Supralabiale begrenzt. 4 Supraokularia und 4-5 Supraciliaria sind ausgebildet. Die Fron-

toparietalia sind geringfügig größer als das Interparietale. Ein Foramen parietale ist nicht erkennbar.

Zwei Nuchalia begrenzen caudal die Schädelpholidose.

Färbung

Das typische Zeichnungsmuster der meist hell- bis gelbbraun gefärbten Tiere ist ein dorsomedianer,

schmaler, dunkler Streifen, der von hellen Längsstreifen begrenzt oder nur als Fleckenlinie auf einem

hellen Mittelstreifen liegt. Daran folgt beiderseits ein dunkelfleckiger Längsstreifen, der wieder von

zwei schmalen, hellen Dorso-Lateralstreifen gesäumt wird. Den Übergang zu den Flanken bilden

meist ein bis zwei hellgetüpfelte Punktreihen. Die Bauchfärbung der Tiere ist hell-weißlich. Auf der

Hinterseite der Oberschenkel sind, wie für spinalis und maioensis ebenfalls typisch, helle Ocellen aus-

gebildet.

Maßtabelle

Mabuya stangeri salensis von SAL

Nr. ZSM GL KR SL KL KB (6) A OA OL/UL Schuppen

4.82.8 _ 86,0 —_ 16,0 12,0 3,0 353 6,5 7/7 42

4.82.7 98,0 38,0 60,0 353 2 1,8 2 952 7/7 —

4.82.6 _ 72,0 — 13,9 8,5 249 29 5,2 7/7 38

4.82.5 153,0 65,0 88,0 13,0 9,0 2,3 3,0 4,5 7/7 36

4.82.4 166,0 70,0 96,0 13,0 10,5 3,0 357, 4,5 7/7 40

4.82.3 210,0 87,0 123,0 14,9 11,0 3,1 4,0 6,0 TUT, 42

4.82.2 — 64,0 _ 13,0 8,7 2,4 3,0 4,9 6/6 38

4.82.1 _ 86,0 — 15,0 10,0 243 3,2 559 7/6 40

16.82.1 88,0 31,0 57,0 8,5 4,8 — = = — =

16.82.2 90,0 — — _ — _ — = — —

16.82.3 95,0 — — — — _ - _ = —

von BOA VISTA

Mabuya stangeri salensis vel M. st. stangeri (= M. st. salensis)

ZSMS a 10 0 an Be 40

4.1.3.4 Mabuya stangeri spinalis Boulenger, 1906 (Taf. I, Fig. 3)

Material: 9 Ex. ZSM 30/82; (1X intermed. mit Mabuya vaillantı); Insel St. Maria. 1 Ex. ZSM 6/82; Praia —

S. Thiago. 3 Ex. ZSM 363/78; Tarrafal — S. Thiago. 1 Ex. ZSM 364/78; (intermed. mit M. vaillantı; Tarrafal -

S. Thiago).2 Ex. ZSM 156/81;5 kmS. Tarrafal-S. Thiago.2 Ex. ZSM 368/78; Fogo. 1 Ex. BMNH 1906.3.30.40;

Fogo.

1906 Mabuya spinalis BOULENGER

1935 Mabuya spinalıs. - ANGEL

1937 Mabuya spinalıs. -— ANGEL

1955 Mabuya stangeri spinalis. — MERTENS

1982 Mabuya stangeri spinalis. — SCHLEICH

Terra typica: Igreja und S. Filipe, Fogo

Bisherige Verbreitung: Fogo, Sal

Nachgewiesen von: Praia, Tarrafal, S. Thiago; Ilheu St. Maria; Fogo

Artcharakteristika nach:

BOULENGER (1906) — Übersetzung

Schwanz kurz, stumpf. Unteres Augenlid mit ungeteilter, transparenter Scheibe. Nasenlöcher hinter der Vertika-

len der Sutur zwischen Rostrale und 1. Labiale; 1 Postnasale; vorderes Loreale gewöhnlich in Kontakt mit 1. La-

biale. Supranasalia in Kontakt hinter dem Rostrale. Frontonasale breiter als lang. Präfrontalia in Kontakt mit ihrem

inneren Winkel, oder eine kurze Sutur bildend.

Frontalia so lang wie Frontoparietalia und Interparietaliazusammen; in Kontakt mitdem2. und 3. oder 1.,2. und

3. Supraoculare. Vier Supraocularia, wobei das 2. am größten ist. Vier oder fünf Supraciliaria; unterschiedliche

Frontoparietalia, größer als das Interparietale. Parietalia bilden eine Sutur hinter dem letzten Schild; ein Paar Nu-

chalia. 5. oder 6. oberes Labiale am größten und unter dem Auge gelegen.

Ohröffnung oval; beinahe so groß wie Orbit; mit wenigen, leicht vorspringenden Granulae an der Vordergrenze.

Nuchale und Nackenschuppen streng dreigekielt (tricarinat). 36 oder 38 ungleiche Schuppen um die Körpermitte.

Die Hinterextremität reicht bis zum Handgelenk oder dem Ellbogen der abgespreizten Vorderextremität. Glatte

Subdigitallamellen. Schwanz etwa 1,5 mal so lang wie KR. Dorsal oliv, mit oder ohne kleine schwarze Flecken; eine

mehr oder weniger differenzierbare schwarze Vertebrallinie und ein blasser Streifen auf jeder Rückenseite vom Su-

praciliarrand bis zur Schwanzbasis. Unter diesem hellen Streifen ein dunkler oder schwarzer Streifen, der wieder

von einem helleren unteren begrenzt wird; Nacken und manchmal Körperseiten mit kleinen weißen Flecken; weiße

Flecken auf der Hinterseite der Schenkel; unterer Teil gelblich-weiß.

GL 146 mm

Kopflänge 13 mm Vorderextremität 185 mm

Kopfbreite 9 mm Hinterextremität 26 mm

KR 46 mm Schwanz 86 mm

Verbreitung: Igreja und $. Filipe, Fogo

Artverwandt mit M. stangeri, aber leicht unterscheidbar durch das Vorhandensein eines einzigen echten Sub-

oculare, der geringeren Anzahl von Körperschuppen um die Rumpfmitte und dem Vorhandensein einer dunklen

Vertebrallinie.

MERTENS (1955) — 1 Exemplar von Fogo:

Interparietale vollständig entwickelt, 3 (statt 4-5) Supraciliaria. Unter dem Auge befindet sich 6. Supralabiale,

das größer aber niedriger als das 7. ist. Um den Körper stehen 36 (36-38 bei den Cotypen) Schuppenreihen, dreikie-

lig; dunkle, hellgesäumte Vertebrallinie ist vorhanden.

ANGEL (1937) —

Nur 1 Labiale (5. oder 6.) unter dem Auge. 4 oder 5 Supraciliaria; eine weiße Dorsolateral-Linie.

Diagnose:

KR-Länge bis ca. 70 mm bei ca. 180 mm Gesamtlänge. 34-38 Schuppenreihen um die Körpermitte;

8-6 Supra- und 5-7 Sublabialia; Dorsalia tricarinat, Ventralia glatt. 4-6 anteriore Aurikulartuberkel;

4 Supraciliaria, -ocularia. 2 Schenkelocellen auf der Rückseite der Oberschenkel, helle Tüpfelung zwi-

schen Ohr und Achsel.

Beschreibung:

Vorab ist zu erwähnen, daß aus zwei verschiedenen Kollektionen, nämlich einer Aufsammlung von

der Insel St. Maria vor Praia (ZSM 30/82) sowie aus einer Aufsammlung von Tarrafal (ZSM 363/78)

zwei phänotypisch intermediäre Formen mit M. vaıllanti auftraten. Beide Exemplare heben sich auf-

grund ihrer Rückenzeichnung (Längsstreifen mit hell/dunklen Sattelflecken) deutlich von den übrigen

Tieren ab, sind aber möglicherweise als intermediäre Formen abzugrenzen. Zudem besitzen sie die

größte KR-Länge dieser Serie. Das Exemplar von Tarrafal (ZSM 364/78) zeichnet sich zudem noch

durch die höhere Rumpfschuppenzahl (44) in seiner Serie ebenso wie das Tier von Ilheu St. Maria

(ZSM 30/82.9) mit 40 Rumpfschuppen aus. Die Pileuspholidose entspricht wieder der von Mabuya

stangeri stangerı.

Habitus

Die kräftig gebauten Tiere sind von relativ kurzer, gedrungener Gestalt. Der stumpfe Kopf hebt sich

ebenso wie der dorsoventral abgeflachte Schwanz kaum vom Rumpf ab. Die Ohröffnung ist kleiner als

bei den anderen Formen, oft nur halb so groß wie der Augdurchmesser.

Pholidose

34-38 Rumpfschuppen sind ausgebildet, zwischen 9-6 Supra- und 5-7 Sublabialia kommen vor.

Dorsal sind die Schuppen dreigekielt, ventral glatt. Vier bis sechs vordere Ohrrandschuppen, 4 Su-

praciliaria und 4 Supraocularia sind ausgebildet. Die übrige Schädelpholidose entspricht dem stange-

rı-Typ.

Färbung

Für alle phänotypisch reinen ‚‚stangeri spinalis“-Tiere ist ein einheitliches Zeichnungsmuster cha-

rakteristisch.

Am auffallendsten sind dabei die systematisch wertvollen Schenkelocellen auf der Rückseite der

Oberschenkel. Meist sind zwei runde, weiße Flecken auf dem Oberschenkel, beimanchen Exemplaren

ebenfalls auf dem Unterschenkel vorhanden. Ebenfalls ist eine unregelmäßige, helle Tüpfelung zwi-

schen Ohr und Achsel typisch.

Die Tiere sind dorsal von graubrauner Grundfärbung, ventral cremefarben. Dorsolateral flankieren

dünne helle Streifen, vom hinteren Augenrand ausgehend bis über die Schwanzbasis hinaus, den ein-

farbigen oder dunkel gesprenkelten Rücken. Ein deutlich hellerer oder gleich heller Fleckenstreifen

verläuft zwischen Achsel und Oberschenkelansatz, worauf lateroventral nochmals ein dunkler Streifen

folgt, der dann von der hellen Bauchfärbung abgelöst wird.

Maßtabelle

Mabuya stangeri spinalis von ILHEU ST. MARIA

Nr. ZSM GL KR SL KL KB oO A OA OL/UL Schuppen

30.82.9 _ 72,0 _ 13,4 9,0 2,8 3,6 4,9 6/6 40

30.82.8 104,0 36,0 68,0 8,9 5,2 1574 2,5 IH 7/6 -

30.82.7 95,0 33,5 61,5 9,0 Sol 1,5 Al 7, 7/6 _

30.82.6 85,0 37,5 48,5 9,8 Sa A 2,9 4,0 7/6 36

30.82.5 _ 40,0 _ 9,5 6,2 59 3,3 RD. 6/5 36

30.82.4 180,0 67,0 113,0 13,6 8,9 1,9 4,0 SR 7/5 36

30.82.3 _ 61,0 _ 195 7,5 2,0 3,9 4,5 8/5 37

30.82.2 168,0 69,0 99,0 18%5 8,0 2,1 4,0 5,0 6/5 36

30.82.1 7AO 62,5 114,5 1252 9,0 2,5 3,6 5,4 7/5 36

Praia Hafenstraße / S. THIAGO

6.82.1 0 20 2 ae 6/7 30

S - Tarrafal / S. THIAGO

363.78.3 - 60,0 - 13,5 8,5 255 4,0 5,0 6/5 36

363.78.2 157,0 64,0 83.0 12,6 8,2 255 3,0 4,8 7/6 34

363.78.1 188,0 72,0 116,0 125 353 2,4 255 5,0 6/6 38

364.78, _ 80,0 — 15,0 10,0 3,0 3 6,0 7/6 44

156.81.2 180,0 67,0 113,0 13,5 355 25 3 4,8 7/7 36

156.81.1 177,0 67,0 110,0 13,0 9,0 3,0 3,0 5,0 7/6 36

368.78.2 - 53,0 — lo) 6,0 153 3,0 4,0 7/6 40

368.78.1 _ 57,0 — 12,2 Zi 167 3,0 5,0 7/7 36

FOGO

BMNH

1906.3.30.40 120,0 57,0 63,0 12,0 PR. 2, Mo 258 353 7/6 36

4.1.4 Mabuya vaillanti Boulenger, 1887 (Taf. I, Fig. 2)

Material: 6 Ex. ZSM 152/1981, St. Cruz -S. Thiago. 1 Ex. ZSM 364/1978, Tarrafal - S. Thiago (intermed. mit

M. stangeri spinalıis; s. dort). 3 Ex. BMNH 1946/81825, 81826; BMNH 1906/33043, Fogo.

1887 Mabuya vaillanti BOULENGER

1937 Mabuya vaillanti. - ANGEL

1955 Mabuya delalandü. — MERTENS

1976 Mabuya vaillanti. — GREER

1982 Mabuya vaillanti. — SCHLEICH

Terra typica: $S. Thiago

Bisherige Verbreitung: S. Thiago, Fogo, Rhombos

Nachgewiesen von: $. Thiago, Fogo (Material: BMNH)

Artcharakteristika nach:

ANGEL (1935) —

54 Schuppen um die Körpermitte, 5. und 6. Supralabiale verschieden, bilden Subokularia. Ohröffnung viel grö-

ßer als Augöffnung.

GREER (1976) —

KR: 122 mm; 54 Schuppenreihen.

Diagnose:

KR-Länge ca. 13 cm bei ca. 29,5 cm Gesamtlänge. Kopf klein, stumpf, wenig abgesetzt.

50-56 Schuppen um die Rumpfmitte, 7 Supra- und 6 Sublabialia. 4-6 posteriore Aurikulartuberkel.

5. Supralabiale vergrößert unter Auge. Parietalia mit Interparietale verschmolzen; ein oder zwei Nu-

chalia; Dorsalia schwach tricarinat. Gelb-orange Schnauzenfärbung; Rückenzeichnung typisch drei-

streifig.

Beschreibung:

Diese größte kapverdische Art der Gattung Mabuya konnte aufgrund ihrer scheuen, relativ ver-

steckten Lebensweise erst spät, nach einem Hinweis von Herrn R. STEFFENS, entdeckt werden. Ihm

und Frau E. STE gebührt hier ein besonderer Dank beı der Auffindung dieser von MERTENs 1955 mit

Mabuya delalandii synonymisierten Art.

Sympatrisch mit Mabuya delalandii und Mabuya stangeri spinalis konnte Mabuya vaillanti bei

Tarrafal (S. Thiago) beobachtet werden.

Habitus

Bis zu 295 mm Gesamtlänge bei 122 mm KR-Länge maß ein Tier von St. Cruz. Der Schwanz ist nıe

länger als das 1,5fache der KR-Länge. Der Körper der Tiere ist relativ breit, rundlich und Kopf wie

Schwanz nur wenigvom Rumpf abgesetzt. Der Schwanz ist rundlich, relativ lang. Der Kopf ist in Rela-

tion zur Körpergröße klein und schmal. Die Ohröffnung ist größer oder nur geringfügig kleiner als der

Augdurchmesser.

Pholidose

Die sechseckigen Dorsalia sind zwei- bis meist dreikielig, auf den Ventralia ist die Dreikielung nur

noch angedeutet zu erkennen. Zwischen 50 und 56 Schuppenreihen zählt man um die Rumpfmitte.

Der Pileus zeigt eine deutliche, nur für vazllantı spezifische Pholidose. Das breite Rostrale reicht bis

unter die Nasalia. Zwei Supranasalia schließen sich an. Das folgende Frontonasale ist von gleicher

Breite wie die beiden Supranasalia zusammen. Zwei Präfrontalia, 1 Frontale sowie 4 Supraocular-

schilder begrenzen seitlich und von vorne das ungeteilte Frontoparietale. Ein großes ungeteiltes Parie-

tale reicht beidseits fast bis an die hinteren Ciliarıa. Ein großes Nuchale von fast der Breite des Fronto-

parietale grenzt die Pileusschuppen von den Dorsalia ab. Eine typische Palpebralscheibe ist ausgebil-

det.

Zwei Postocularia und zwei Postsubocularia sind erkennbar. Neun Temporalia liegen zwischen dem

Postoculare und den Aurikulartuberkeln. Meist sind 7 Supra- und 6 Sublabialia zu zählen, zwei Post-

labialia begrenzen Oberlippen und Ohr. Die ventrale Kopfbeschuppung besteht aus einem umfassen-

den, bis unter die Mitte des 1. Supralabiale reichenden Mentale, einem bis unter die Mitte des 2. Sub-

labiale reichenden Postmentale sowie je Seite 3 Submaxillaria, deren 1. und 2. ein Inframaxillare flan-

kieren.

Färbung

Von der graubraunen bis kräftig braunen Grundfärbung heben sich auf dem Rücken drei hellbraune

Streifen ab. Die Seitenstreifen sind etwa anderthalbmal bis doppelt so breit wie der Mittelstreifen und

beginnen kurz oberhalb und hinter dem Ohr, um dann nach etwas mehr als der Hälfte der Schwanz-

länge wieder zusammenzulaufen. Der dünnere Mittelstreifen dagegen beginnt erst ab Achselhöhe. Die

gesamte Schnauzenspitze bis etwa zum 2. Labiale inklusive des Mentalbereiches ist kräftig orange ge-

färbt. Die dazwischen liegenden dunkleren Streifen werden von noch dunkleren braunen Punkten im

Abstand von ca. 0,5 mm eingefaßt. Ab dem hinteren Ohrrand hebt sich ein schwarz/gelber Fleck ab,

dessen Farbgrenze ziemlich genau auf der Mitte des Ohres liegt. Entlang der Flanken zieht eine Punkt-

reihe weißer Flecken. Darunter wechselt die Flankenfärbung nach grau mit schwarzen Sprenkeln, die

sich über die Ventralseite fortsetzen können. Die Hand- bzw. Fußsohlen sind ebenso wie Schnauze

und Kehle orange gefärbt. Bei O’CO’ tritt diese Färbung auch auf dem Bauch auf. Bemerkenswert ist das

Zeichnungsmuster noch zwischen Kehl- und Achsel-Bereich. Ab dem posterioren Ende der orangen

Kehlfärbung verlaufen beidseits je drei Streifen zum Ohr, zwei etwa auf halber Länge Ohr/Achsel und

der dritte bis kurz vor der Achsel. Die Vorder- wie Hinterextremitäten können ebenfalls gesprenkelt

erscheinen. Schwach bis deutlich erkennbare schwarz/(gelbe) Achselocellen können auftreten.

Maßtabelle

Mabuya vaillanti von St. Cruz / S. THIAGO

Nr. ZSM GL KR SL KL KB (@) A OA OL/UL Schuppen

152.81 _ 130,0 _ 23,0 15,0 4,0 4,5 9,5 7/6 52

152.81 295,0 122,0 173,0 21,6 14,5 4,7 4,0 9,0 7/6 54

152.81 233,0 116,0 117,0 12 14,5 3,8 4,2 8,0 7/6 52

152.81 _ 80,0 _ 16,0 10,5 So) Sal 6,0 7/6 54

152.81 151,0 71,0 80,0 13,5 9,0 Dal 2) 5,8 7/6 50

152.81 281,0 110,0 171,0 21,6 15,9 4,1 4,2 9,0 7/6 54

Trockenexemplar 21,2 17,0 5,2 3,8 8,6 7/6 52

BMNH

1946.81825 2 zo ee a ze ie 50

1946.81826 25,0 100,0 123,0 180 a az 50

von FOGO

1906.33043 0 a a a az er 7/6 56

4.2 Gekkonidae - Genus Tarentola

Während Dumrrır & Bıgron (1836) Platydactylus (= Tarentola) delalandii als Form mit einfachen,

ovalen, nur sehr schwach gekielten Dorsaltuberkeln mit einem Ohrloch ohne zackige Randschuppen

und einer Schuppenreihenzahl von 6 auf dem Schwanz als Tiere von Tenerife darstellen, beschreibt

STEINDACHNER (1891) Tarentola delalandi boettgeri als eine Form mit den gleichen Merkmalen, die er

durch die höhere Anzahl von 16-18 Tuberkelreihen von Tieren von Gran Canaria und jenen mit

12-14 Reihen von Tenerife, Palma und Gomera unterscheidet. Hinzu kommt, daß Dumerı & BiBRON

keine Angaben über die Anzahl der Tuberkelreihen gemacht hatten, ihre Beschreibung aber gänzlich

der Art boettgeri sensu STEINDACHNER, entsprechen würde. Nach Materialvergleichen mit Aufsamm-

lungen sowohl von den Canaren als auch den Kapverden und all den verwirrenden Beschreibungen

nachfolgender Autoren wäre fast an eine Fundortverwechselung von Dumrrir & Bısron zu denken.

Verwirrender wird letztlich noch die Synonymisierung von LOVERIDGE (1947, S. 5, 6) mit folgender

Erklärung: ‚As a further consequence of these studies nearly fifty described forms or species appear to

be unrecognizable... The following, however, are believed to be referred to the synonymy for the first

time:

‚„‚Tarentola gigantea (lapsus) Scherer = T. delalandii gigas (Bocage)

Tarentola d. var. boettgeri Steindachner = T. d. delalandü Dumeril & Bibron

Tarentola d. var. boettgeri Boulenger = T. d. delalandii Dumeril & Bibron.“

MerTens (1955, $. 6) stellte sämtliche kapverdischen Tarentolas (außer den Tieren von Branco und

Razo) zu Tarentola delalandii rudis, ‚‚weil das vorliegende Material trotz seiner Variabilität in seiner

Gesamtheit einen anderen Eindruck macht, als die auf den Kanaren lebenden Angehörigen von Ta-

rentola delalandii“, und sieht sowohl Tarentola delalandii delalandii (von Tenerife) und delalandıi

boettgeri von Gran Canaria als lediglich auf den Kanaren verbreitete Rassen an. Bei den Tieren von

Fogo erwähnt er nur ein durch seine Größe von der übrigen Aufsammlung herausfallendes Exemplar,

sieht dieses jedoch wiederum innerhalb der Variabilität dieses Rassenkreises an:

T. delalandii: Schnauze ein wenig länger als die Entfernung zwischen Auge- und Ohröffnung. Das Mentale ist in

der Mitte nicht länger als seine dreifache Breite.

T. gigas: Schnauze nicht viel länger als der Aug/Ohr-Abstand. Die Kinnschuppenlänge entspricht ihrer dreifa-

chen Breite.

Während er für 7. delalandıi die ‚Varietäten‘ wie folgt abgrenzt:

T. del. delalandii: 12 Längsreihen vergrößerter Tuberkel, glatt oder gekielt.

T. del. rudis: 16-18 Tuberkelreihen, deutlich gekielt, 4 Transversalbänder mit hellen Mittelflecken, Schwanz-

schuppen spinös.

T. del boettgeri: 18-20 glatte Tuberkelreihen, gerade Transversalbänder.

JOGER (1984) trägt nun, nach meinem Versuch (SCHLEICH 1984) die Systematik der kapverdischen

Tarentolas zu er- und überarbeiten, erneut zu größerer, hauptsächlich nomenklatorisch taxonomischer

Verwirrung bei, indem er längst ‚‚verschollene“ nomina oblita zu revalidisieren versucht.

Eine Klärung dieser neu entstandenen Problematik ergibt sich jedoch zwangsläufig aus der Beach-

tung der Internationalen Regeln für die Zoologische Nomenklatur, insbesondere mit ihren 1970 erfolg-

ten Ergänzungen, so daß die alteingeführten nomina conservanda von Tarentola gigas und Ta-

rentolarudis selbstredend erhalten bleiben können. Hierfür einige Auszüge aus besagten Ergänzungen:

(Erklärung 43 n. Fassung v. 1.1.1970 zu $ 23b I-VIII)

$ 33bI: ‚Einschränkung. — Ein Name, der sich in fortlaufendem, allgemeinen Gebrauch befindet und minde-

stens50 Jahre verfügbar gewesen ist, darf nach 1960 nicht durch ein unbenutztes älteres Synonym ersetzt werden.“

„Ein Name ist als in fortlaufenden allgemeinem Gebrauch befindlich anzusehen, wenn er in den unmittelbar vor-

ausgehenden 50 Jahren von mindestens fünf verschiedenen Autoren und in mindestens zehn Veröffentlichungen als

mutmaßlich gültiger Name auf ein bestimmtes Taxon bezogen wurde.

(II) Ein älteres Synonym ist als unbenutzt anzusehen, wenn es im Laufe der unmittelbar vorausgehenden 50 Jahre

kein einziges Mal auf ein bestimmtes Taxon als dessen mutmaßlich gültiger Name bezogen wurde. Ein älteres unbe-

nutztes Synonym, das nach 1960 unter Verletzung der Vorschriften von Artikel 23 b verwendet wurde, sei es, um

das jüngere Synonym ausdrücklich zu ersetzen oder nicht, verliert hierdurch nicht seinen Status als unbenutzter

Name.

(III) Die Erwähnung eines Namens in der Synonymie oder lediglich dessen Aufnahme in eine Referier-Veröffent-

lichung, einen Nomenklator oder sonstigen Index oder eine Liste von Namen bedeutet keinen Gebrauch im Sinne

von Artikel 23b.

(IV) Jedes Zitat ist für sich zu werten, ohne Rücksicht auf Art oder Titel der Arbeit, in welcher der Name vor-

kommt.“

Zusammenfassen läßt sich so nach Absatz 4 der Normen der Verfügbarkeit:

Tarentola borneensis wurde 1965 von WERMUTH im Sinne von LOVERIDGE (1947: 33!) als synonym zu Tarentola

delalandıi delalandii betrachtet und wäre außerdem so nicht als für Tarentola gigas angewandt zu betrachten. Un-

abhängig davon wurde von beiden Autoren Tarentola delalandii gigas als eigenständig gültiges Taxon dargestellt.

Außerdem tritt für den von JOGER zitierten Fall eindeutig $ 23b Ill in Kraft. Eine gesonderte Synonymieliste zur

Demonstration der Erfüllung dieser Verfügbarkeitsnormen ($ 23b I) ist im Kapitel Tarentola gigas aufgeführt. Die

Forderung von $ 23b sind alle rechtmäßig erfüllt, die Zoologische Nomanklaturkommission wurde dazu von mir,

folgend $ 23b V, informiert.

Bestehen bleibt, daß sich sowohl nach morphologischen als auch nach Pholidose- und Zeichnungs-

merkmalen folgende 4 Artengruppen von Tarentola als eigenständige Formenkreise ausgliedern las-

sen:

Tarentola rudıs, eine durch besonderes Größenwachstum gezeichnete Art, wird von den Inseln

Fogo, S. Thiago mit St. Maria und Maio beschrieben. Sie kommt auf $S. Thiago neben einer ‚‚samt-

schuppigen‘“ Form, mittlerweile als Tarentola darwini beschrieben, vor. Das sympatrische Vorkom-

men dieser beiden phänologisch eindeutig unterscheidbaren Arten zieht taxonomische und nomenkla-

torische Schlußfolgerungen nach sich. Dementsprechend wird neben Tarentola darwini Tarentola ru-

dis als eigenständige Kapverden-Art betrachtet, deren verschieden phänotypisch-morphologische

Ausbildungen der unterschiedlichen Inselformen, Rassenkreise unter den Kapverden-Inseln abzu-

grenzen zwingt. Daneben konnte Tarentola caboverdiana Schleich 1984 mit diversen Unterarten aus-

gegliedert werden.

Die Riesengeckos der beiden Inseln Branco und Razo werden wiederum als eigene Formen der Art

gigas betrachtet.

Entgegen meiner Darstellung (SCHLEICH, 1982a) in der Tabelle 1 zu ‚‚Vorläufige Mitteilungen zur

Herpetofauna der Kapverden“ mit Tarentola delalandıi delalandii als ‚Hauptform“ der kapverdi-

schen ‚„‚Mauer‘geckos und rudis als Rassenkreis davon, bleibt die von Dumezrır & Bısron (1836) be-

schriebene ‚‚Platydactylus“ delalandıı für Tenerife, Madeira und Senegal bestehen.

Tarentola delalandii rudis Boulenger wurde 1906 mit der Terra typica-Angabe: S. Filipe auf Fogo

sowie Praia von $. Thiago bekannt und war so neben den Riesengeckos die einzige von Caboverde be-

schriebene Tarentola.

Daraus ergab sich für ‚‚rudis“ die Notwendigkeit, jene Form als kapverden-typisch anzusprechen,

während die Unterarten Tarentola delalandıi boettgeri und Tarentola delalandır nur im nordmakaro-

nesischen Raum mit den Unterarten boettgeri und delalandı vertreten sind.

Zu dem von mir für Tarentola gigas betrachteten eigenen Artstatus stimme ich in bezug auf Größe

und Morphologieaffinitäten zu Tarentola rudis protogigas mit JOGER’s Ansicht überein, jedoch sind

die Formen von Branco und Razo in ihrem Verhalten und in ihren ökologischen Ansprüchen so von al-

len übrigen Kapverdengeckos verschieden, daß eine Gleichheit auf Artniveau angezweifelt werden

muß. Als gravierende Merkmalsunterschiede wären nach umfangreichen Gelände- und Gefangen-

schaftsbeobachtungen zwischen den beiden erwähnten Formen zu nennen:

Tarentola (r.) protogigas

Lautäußerungen konnten nıe wahrgenom-

men werden.

Eiablage ungeklärt, erfolgt den Lebensge-

wohnheiten nach wahrscheinlich an Mauer oder

vergleichbarem Trockensubstrat.

Tiere bewegen sich speziell auf vertikalen

Wänden.

Tarentola gigas

Lautäußerungen spielen im sozialen Verhal-

tensmuster eine deutliche Rolle (spez. epiga-

misch).

Eiablagen erfolgen nur unter ausgesucht spe-

ziellen Bedingungen (s. SCHLEICH 1980).

Adulte Tiere sind kaum in der Lage, sich in

der Vertikalen zu halten (zu großes Körperge-

wicht), und konnten als über 95% bodenstän-

dig in ihren Lebensäußerungen beobachtet

werden. Eine positive Selektion zum Riesen-

wuchs war durch fehlende inter-/intraspezifi-

sche Competition möglich.

Eine besondere Fettspeicherung fehlt. Eine extreme Fettspeicherung ist typisch.

4.2.1 Tarentola rudis Boulenger, 1906

4.2.1.1 Tarentola rudis rudıs Boulenger 1906 (Taf. II, Fig. 3; Taf. IV, Fig. 2)

Material: 3 Ex. ZSM 139/1981 -S. Thiago / Praia-Flughafen. 1 Ex. ZSM 372/1978-S. Thiago / C. Velha. 2 Ex.

ZSM 135/1981 — S. Thiago / Praia. — Paralectotypus n. JOGER (1984: 101); MCNG 28149/2.

1906 Tarentola delalandii var. rudis BOULENGER

1935 Tarentola delalandii. -— ANGEL

1937 Tarentola delalandii. -— ANGEL

1947 Tarentola d. var. rudis BOULENGER

= Tarentola d. delalandii (DUMERIL & BIBRON). — LOVERIDGE

1955 Tarentola delalandıi del.. — DEKEYSER & VILLIERS

1955 Tarentola delalandii rudis. — MERTENS

1984 Tarentola rudis rudis. — SCHLEICH (part.)

1984 Tarentola b. (= borneensis) rudis. — JOGER

Terra typıca restricta: Praia — S. Thıago, ?S. Filipe-Fogo

Bisherige Verbreitung: Sal, Boa Vista, Maio, $. Thiago, Fogo, Brava, St. Luzia

Nachgewiesen von: $. Thiago (Praia), Ilheu St. Maria (von dem kleinen Eiland St. Maria vor Praia wurden aus

Schutzgründen keine Tiere gefangen)

Artcharakteristik nach:

BOULENGER (1906) —

Hervorstechende Tuberkel, mehr oder weniger unterschiedlich gekielt, auf dem Schwanz fast spinös; in 16-18

Transversalreihen auf der Körpermitte; Schläfe mit sehr unterschiedlich vergrößerten Tuberkeln; 4 verschiedene,

wellige oder w-förmige dunkle Querbänder, deren mittlerer Sinus an einen hellen Fleck grenzt. KR bis 88 mm.

Diagnose:

Eine Diagnose ergibt sich nach SCHLEICH (1984) und JOGER (1984) wie folgt:

Kopf-Rumpflänge max. 8,5 cm, PP kleiner (6,5 cm). 16 Tuberkelreihen; zwischen 10/11 Supra-

und 9/10 Sublabialia. 16-19 Interorbitalschuppen; 130-165 Schuppenreihen um die Körpermitte. Dor-

saltuberkel undeutlich gekielt. Fünf dunkle Transversalbänder. Orbit meist doppelt so groß wıe Ohr-

durchmesser. Auf dem ersten Finger 12, auf dem dritten 13 Subdigitallamellen. Verhältnis Mentalia-

länge : -breite ca. 2:1. Maulspaltlänge um mindest die Hälfte länger als die Strecke zwischen vorderem

Augenrand bis Schnauzenspitze. Entfernung Ohr-Auge geringer als Augvorderrand-Schnauzenspit-

ze.

Beschreibung des Lectotypus: s. JOGER (1984: 101)

Anmerkung: JOGER designierte aus BOULENGER’s Syntypen von ‚,S. Jago“ eines der Exemplare als

Paralectotypus zu ‚7. b. rudis Boul., 1906“ von S. Thiago mit der Terra typica restricta: „‚Dist. di

Praia, Calhetta de S. Martinho“. Da mir seinerzeit dieses Originalmaterial nicht zur Verfügung stand,

ich selbst aber umfangreichere Aufsammlungen von Fogo, St. Maria (S. Thiago) und von Praia selbst

tätigte, glaubte ich, daß die von genannten Lokalitäten gefangenen Exemplare alle einem sehr variablen

Rassenkreis angehören würden. Mittlerweile möchte ich mich aber der Ansicht JoGEr’s anschließen

und den südl. Bereich von S. Thiago als Vorkommen für Tarentola rudis rudis bestätigen.

Beschreibung:

Für die von mir gesammelten Exemplare von Praia ergibt sich: Kopf-Rumpflänge bis 58,5 mm, der

Schwanz ist meist länger, so daß sich max. eine Gesamtlänge von 120,5 mm ergibt. Der Schädel ist

deutlich länger als breit, der Augdurchmesser etwa doppelt so groß wie der Ohrdurchmesser. Der Ab-

stand Ohr/Auge ist wenig kleiner als der Abstand Auge/Schnauzenspitze. Der Maulspalt reicht bis

hinter das Auge. Das Mentale ist etwa doppelt so lang wie breit. An Lippenschildern sind 10-11 Supra-

und 9-10 Sublabialia ausgebildet. Die Rückentuberkel sind in 16 Längsreihen angeordnet. Die Tiere

sind grau bis graubraun gefärbt und tragen 5 Transversalbänder.

Maßtabelle

Tarentola rudis rudıs von S. THIAGO;

Mentale Tub. Lam.

Nr. ZSM GL KR SL KL KB © A OA AS MSP L BIOR/ULZERE Jr Bd

139.81.1 10 SD SD Az el zZ rl 70 0 Az 5 2,8 10/9 16 _ 5

139.81.2 1116,0556,0000,0818,0212,8 2 2312 AN E77 8122050235 10/9 16 12/13 5

139.81.3 2.059585568:0818:021055251585.770=26752:23:0527125 8,5: 0002 5 le 16 = 5

4.2.1.2 Tarentola rudis maioensis Schleich, 1984 (Taf. II, Fig. 5; Taf. IV, Fig. 1)

Material: Holotypus — ZSM 136/81.6. Paratypen — ZSM 136/81.1-5 u. 7-9.

1954 Tarentola delalandii rudıs. - MERTENS (part.)

1984 Tarentola rudis maioensis. — SCHLEICH

1984 T. b. (borneensis) maioensis. — JOGER

Terra typica: Maio

Verbreitung: Maio und n. JOGER (1984: 102) auch Boa Vista

Diagnose:

Gesamtlänge bis ca. 139 mm bei max. 68 mm KR-Länge. Die Kopflänge entspricht dem 1,3-1,6fa-

chen der Kopfbreite. Der Abstand zwischen vorderem Ohrrand und Auge ist meist gleich groß der

Entfernung vom Auge zur Schnauzenspitze. 12-18 Tuberkelreihen; eine Dorsalbänderung ist kaum

wahrnehmbar.

Beschreibung:

Die Tiere sind von relativ plumper, kräftiger Gestalt und bis ca. 139 mm lang, ihr Kopf erscheint

groß und breit. Der Maulspalt reicht deutlich hinter den Pupillenspalt, der Ohrschlitz ist auffallend

klein und etwa nur halb so groß wie der Augdurchmesser. Die Tuberkelschuppen sind konisch bis

leicht apikal.

Alle Tiere sind graubraun gefärbt, ohne deutliche Rückenbänderung. Falls erkennbar sind 5, relativ

breite, schwach dunkle Transversalbänder mit hellem Kernfleck oder Mittelstreifen typisch. Zwischen

Ohr und Auge zieht sich bis zu den Nasenlöchern ein heller Streifen.

97.

Maßtabelle

Tarentola rudis maioensis von MAIO

Mentale Tub. Lam.

Nr. ZSM GEW KR SE: U KL "KBim@s "AH 1OAU ASHMSPILTITM BI OPT ORTE

136.81.1 _ 0 TEE IE 9/7 27 AV

136.81.2 — fo —_ ae ee re I I re 8/7 14 87 -

136.81.3 = Ge are 2 er oe RD ee Fe 8/7 6105

136.81.4 - 47.0 — 25,0 710,0 21,87 020772050 5229 8/7 14 11/11 5

136.81.5 112875960500.6255, 2.0101 379223277 089 O PS ENTE 14 14/11 5

136.81.6 159208.68:0872130822, 0516. 0,2775 ES Sales Bis 9/7 18713715775

136.81.7 955084820. 45:0 75 SE 155 270 E40 E60 6500529 9/8 16021225

136.81.8 = ln ee 7 2 8 el 5 ee 9/8 14 11/12

136.81.9 102053593 5E175217 25 14 02H IA OEA IE 9/8 14 11/13

4.2.1.3 Tarentola rudis protogigas Joger, 1984 (Taf. II, Fig. 1, 2, 4; Taf. IV, Fig. 7)

Material: 11 Ex., ZSM 145/1981 - Fogo. Holotypus: ZSM 145/1981.1 — Fogo. Paratypus: ZSM 145/1981.2 -

Fogo und Exemplare des BMNH u. MCNG C. E. (s. JOGER, 1984: 101).

1906 Tarentola delalandiüi var. rudis BOULENGER (part.)

1947 Tarentola delalandii. — LOVERIDGE (part.)

1984 Tarentola rudis rudis. — SCHLEICH (part.)

1984 Tarentola b. (borneensis) protogigas. — JOGER

Terra typıca: Fogo

Anmerkung: Ein isoliertes Vorkommen von einer allerdings recht kleinen Population konnte auf dem Inselchen

St. Maria in der Hafenbucht vor Praia entdeckt werden. Die Tiere erreichten mindest die Körperlänge der Exem-

plare von Fogo. Aus Schutzgründen wurde hier jedoch selbst auf Belegexemplare verzichtet, da wir nur sehr wenige

große Individuen zu sehen bekamen.

Verbreitung: Fogo, Ilheu St. Maria (S. Thiago) und nach JOGER (1984: 101) Brava, Rhombos-Inseln

Diagnose:

Mit Tarentola gigas die größte kapverdische Tarentola mit morphologischer Merkmalsausbildung

der T. rudis-Gruppe. Rückentuberkel schwach gekielt, 144-181 Schuppen um die Körpermitte. Max.

KR-Länge 98,5 mm. Schwanz kürzer als Kopf-Rumpflänge. Die Anzahl der Subdigitallamellen be-

trägt bei der 1. Zehe 12-14 und an der 5. Zehe 22-26 bis zur Zehenbasis gezählt.

Beschreibung:

Die bis knapp 10 cm (KR) lang werdenden Tiere sind kräftig gebaut, jedoch nicht so massig und dick

wie die Riesengeckos (T. gigas) der Inseln Branco und Razo. Auf die Kopflänge entfallen dabei etwa

!/; bis '/ der KR-Länge.

Der Maulspalt reicht bis hinter das Auge. Der Ohrschlitz ist meist halb so groß wie der Augdurch-

messer. Die Tuberkelschuppen sind relativ flach, leicht apıkal und von entsprechend kleinen Schuppen

umgeben. Die leicht vorgezogene Spitze ist nur wenig erhaben. In meist 16 Längsreihen (1 Ausnahme

mit 18) sind die Rückentuberkel angeordnet. Die Mentalialänge entspricht etwa zweimal ihrer mittle-

ren Breite. Meist sind 10-11 Supra- und 8-9 (7, 11) Sublabialia ausgebildet.

Die grau bis graubraun gefärbten Tiere tragen 4-5 Transversalbänder.

Tafel II

Tafel II: Fig. 1: Tarentola rudis protogigas von Fogo; adultes Exemplar, in katasematischer Färbung. Fig. 2: Ta-

rentolarndis cf. protogigas von Fogo. Fig. 3: Tarentolarudisrudis von S. Thiago. Fig. 4: Tarentola rudıis cf. pro-

togigas von Ilheu St. Maria. Fig. 5: Tarentola rudis maioensis von Maio. Fig. 6: Tarentola darwini von Tarrafal-

S-Thiago. Fig. 7: Tarentola caboverdiana substituta von S. Vicente. Fig. 8: Tarentola caboverdiana raziana von

St. Luzıa.

Maßtabelle

Tarentola rudis protogigas von FOGO; n=11

Mentale Tub. Lam.

Nr. ZSM GE SR SIE IS 108 (© NEO REENSESNSPEEITE B OL/UL R. % Bd

145.81.1 _ SD 7925 3,8 6:0 Sole 020 16 4

145.81.2 - u 2er Is ee ed Br 10 16 4

145.81.3 = FE ZI Ir 755 10,0 15,0 30 3 OR 16 13/15 4

145.81.4 _ 2,0 = I Per 2 5 2 1ER 16 5

145.81.5 _ SU Teams 418 16:5, 725015705508 16 5

145.81.6 = SSR EE 09025 5507 10:0 7115021720767 la 18 5

145.81.7 - SO OeE r57255545, 650, 10FOE5 3703 16 4

145.81.8 - ee ae I 25 50 7 1 re 3 eo 16 5

145.81.9 - I = A I 30 7 Del 8 3 9/7 16 5

145.81.10 - DON =11910515555554,0,,478,,.06,80278012557276,0,2286 9/8 16 4

145.81.11 = 20 ee 2a er Te Ne rev 16 5

4.2.2 Tarentola darwini Joger, 1984 (Taf. II, Fig. 6; Taf. IV, Fig. 3)

Material: 22 Ex. Holotypus - ZFMK 37256 (coll. et don. Schleich, 1978), S. Thiago-Tarrafal, und 21 Ex. ZSM

365/78, 146/81, 147/81, 29/82, S. Thiago-Tarrafal. Paratypen (nach JOGER, 1984): 365/78, 146/1981, 147/1981.

1906 Tarentola delalandıi boettgeri BOULENGER

1947 Tarentola delalandii delalandii. — LOVERIDGE

1954 Tarentola delalandii rudıs. — MERTENS

1982 Tarentola sp. — SCHLEICH

1984 Tarentola sp. - SCHLEICH

1984 Tarentola darwini. — JOGER

Terra typica: Tarrafal, S. Thiago

Verbreitung: Tarrafal, S. Thiago; Igreja, Fogo

Die Vermutung JOGER’s (1984) für ein Vorkommen auf Nicolau und Sal erscheint mir spekulativ, für Fogo dage-

gen wahrscheinlich.

Diagnose:

Eine Diagnose ergibt sich nach JOGER (1984: 96) und SCHLEICH (1984: 102):

Kopf-Rumpflänge bis 62 mm. Flache, meist glatte Tuberkel mit nur schwachem Kiel. 18-20 Tuber-

kelreihen.

Deutlich abgesetzte, zu Ketten verwobene Härchen der Schuppenoberflächen, die sich dadurch

samtig anfühlen. Zahlreiche, meist glatte Rückentuberkel zwischen den Hinterseiten der Vorder- und

Hinterbeinansätze. Dorsal enge bis unregelmäßige Transversalbänder bis marmorierter Zeichnungs-

typ.

Die weiteren von JOGER (op. cit.) unter „Diagnose“ erwähnten Merkmale sind differentialdiagnostisch.

Beschreibung:

Für die Tiere von Tarrafal ergeben sich folgende Merkmale: Die bis 110 mm Gesamtlänge messen-

den Tiere sind von relativ kleinem Habitus mit auffallend kleinem Kopf. Die Kopflänge entspricht etwa

1,1- bis 1,4mal der Kopfbreite. Die Ohröffnung ist klein, meist kleiner als der halbe Augdurchmesser.

Der Abstand Ohr-Auge ist nur geringfügig kleiner als der vom Auge zur Schnauzenspitze. Zwischen

8-13 Supra- sowie 7-10 Sublabialia können ausgebildet sein. An einem Exemplar konnten an der ersten

Zehe 9 und an der dritten 13 Subdigitallamellen gezählt werden. Die Mentaliabreite entspricht etwa der

halben Mentalialänge. Die Tuberkel sind rundlich bis oval, schwach gekielt und kaum erhaben; sie sind

in 14-18 Längsreihen angeordnet. Das Mentale wird von 2 Submaxillarıa flankiert.

der Gattung Tarentola (s. SCHLEICH,

Abb. 10-13: Schematisierte Rückenzeichnungen kapverdischer Geckos

Tarentola caboverdiana nicolauensıs;

1984): 10a, b) Tarentola gigas brancoensis; 11) Tarentola gigas gigas; 12)

13) Tarentola caboverdiana caboverdiana.

In der Färbung und Zeichnung sind die Tiere von allen anderen kapverdischen Geckos gut zu unter-

scheiden. Die Grundfärbung ist gräulich bis graubraun, der Bauch hell. Eine dorsale Bänderzeichnung

ist kaum bis nur sehr schwer erkennbar; vier bis fünf Bänder können gezählt werden. Im allgemeinen

ist jedoch eine silbrig graue Grundfärbung mit dunkler Sprenkelung vorhanden. Die Supralabialia sind

dunkel gefleckt, die Sublabialia können hell bleiben. Ein schwacher heller Streifen verläuft von den Na-

senlöchern zum vorderen Ohrrand.

Anmerkung: JOGER (1984: 98) gibt an, daß von Sal ein Einzeltier in sehr schlechtem Erhaltungszu-

stand überliefert ist und begründet die mögliche Artzurechnung zu darwini lediglich mit der hohen

Anzahl der Rückentuberkel. Weitere Angaben über Herkunft und Aufsammlung fehlen. Für die

Zuordnung von Jocer’schen darwini-Typen von $. Nicolau bestehen ebenfalls Zweifel, und er selbst

schreibt (S. 97): „Die Zuordnung dieser beiden Exemplare erfolgt unter Vorbehalt.“

Maßtabelle

Tarentola darwini von Tarrafal / S. THIAGO

Mentale Tub. Lam.

Nr. ZSM EIMIKR/STL KL KBlO A,/ON AS MP L BNOLUE TRIER

29.82.1 _ 520) — ls, RA 22 8 70 75 US es 2 EyAlle) 16 _ =

29282.2 = ler 1A 22 7 ae ee NS 18 - 4

295823 = 56 oO l6 OR 238 A 6 97123 16 - (4)5

29.82.4 = a er rear 7 NN 25 OT 16 _ 5

298223 - elzeeearere 2e Taro — = 8/7 16 - 4

29.82.6 = Te nee er ee 7 re 2 le 16 = 4

DORSDT, - re Ze ae TO N aD en 255 9/7 16 = 4

29.82.8 - DNB 4565ER 9/9 16 _ 5

365.78.1 _ ee OS rs er = _ 10/7 16 - 5

147.81.1 - 56081611450, 2755410618 740 O5 E20 14 931375

147.81.2 - 560 SROSTTEZEE 25373,97 24794 O7 9/7 16 _ 5

146.82.1 10,0, GELERNT AT er la 52 16 = 5

146.82.2 - Do 8012720 530551701255 022 9/7 16 _ 5

4.2.3 Tarentola caboverdiana Schleich, 1984

4.2.3.1 Tarentola caboverdiana caboverdiana Schleich, 1984 (Taf. IV, Fig. 6)

Material: 30 Ex. Holotypus — ZSM 141/81.03, Paratypen — ZSM 141/81.01-02; 141/81.04 - 17.

1947 Tarentola delalandıi delalandii. — LOVERIDGE (part.)

1954 Tarentola delalandii rudis. — MERTENS (part.)

1984 Tarentola caboverdianus. — SCHLEICH

1984 Tarentola caboverdiana. — JOGER

Terra typica: St. Antao

Diagnose (s. SCHLEICH 1984: 98 ff.):

Gesamtlänge bis 103 mm, wobei die Schwanzlänge immer geringer als die Kopfrumpflänge ist. Die

Kopflänge ist 1,5-1,8mal die Kopfbreite (Imal 1,2X). Der Ohrschlitz ist meist länger als der halbe

Augdurchmesser. Die Entfernung Ohr-Auge ist gleich lang oder kürzer als Auge/Schnauzenspitze.

3 Submaxillaria. 14-16 Tuberkelreihen mit caudaler Farbintensivierung.

Beschreibung:

Die bis 103 mm lang werdenden Tiere besitzen einen relativ schlanken Kopf. Der Schwanz ist dor-

soventral abgeflacht, immer kürzer als die Kopfrumpflänge und erscheint dorsal hell/dunkel geringelt.

Die Dorsaltuberkel sind leicht gekielt und in 14 bis 16 Längsreihen angeordnet. Die Zahl der Oberlip-

penschilder schwankt zwischen 9 und 13, jene der Unterlippenschilder zwischen 7 und 10. Zwischen

Augen und Nasenlöchern verläuft ein heller, dunkel gesäumter Streifen, der sich zwischen Auge und

Ohr fortsetzt. Die Labialıa sind weiß gefärbt. Die Rückenzeichnung besteht aus 4-6, meist 5, Trans-

versalbändern.

Maßtabelle

Tarentola caboverdiana caboverdiana von St. ANTÄO

Mentale Tub. Lam.

Nr. ZSM GL KR SL KL KB © A OA AS MSP L B OL/UL R. Bel

141.81.1 _ > 1 120 28 7 7,0 1,5550 2,0 Ay 16 - (4)5

141.81.2 _ 6O,0BE E15, 095 022720519, 012 1519, 16 - (4)5

141.81.3 217528 59, 02.477,2016, 68.115,42 2,052, 222 6,1277,0,1 03,5 ET 5

141.81.4 _ 52 2 510555010,03702735 5,06: AO Orr rg 9/8 14 _ 6

141.81.5 _ A = 10 I AO 5 50 Zi 100 A522 OR 16 - 5

141.81.6 m re ae 2 2 a ae 23 9/9 14 _ 5

141.81.7 103,0 0 EI 1 2 12er 751 5 10 16 = 5

141.81.8 2,0 20 300 az NOS 1,8 8 ed aa a2 9/8 14 - 4

141.81.9 0 29 A re 1 2,070 7 ee 2‘ 9/8 14 = 5

141.81.10 _ 42 = aD NO IR ie rel 9/- 16 = 6

141.81.11 ID SD 40,0 15,0 12,0 227 BZ 80 74 953020 9/8 16 5

141.81.12 90,0 45,5 44,5 15,0 11,6 2,1 4,0 6,0 7,0 10,0 4,0 1,6 10/9 16 = 5

141.81.13 O1, 32,0 29,0 Als ZI a 2 re ee 7 30 9/8 16 _ 3

141.81.14 _ 790 = 15 105 23° 39 80 98 32 1.9 WY 16 = 5

141.81.15 _ 2 IR 2,0 re 7a 1 20 9/8 14 _ 5

141.81.16 770 32 ID 1,2 18,9 2.0 927500 ed ee TR 16 _ 5

141.81.17 31,0 Al 3 1er 1 2er ee ae 207 1 16 _ 5

4.2.3.2 Tarentola caboverdiana nicolauensis Schleich, 1984 (Taf. IV, Fig. 4)

Material: 11 Ex. Holotypus — ZSM 138/81.2, Paratypen — ZSM 138/81.1,3-11.

1984 Tarentola caboverdianus nicolauensis SCHLEICH

1984 Tarentola c. nicolanensis. — JOGER

Terra typıca: S. Nicolau

Verbreitung: S. Nicolau

Diagnose (s. ScHLEicH 1984: 100):

Kopf-Rumpflänge bis 61 mm, Schwanzlänge annähernd gleichlang KR-Länge. Kopflänge

1,2-1,5mal Kopfbreite. Der Abstand Ohr-Auge ist deutlich kleiner als vom Auge zur Schnauzenspit-

ze. 14-18 Tuberkelreihen; Tuberkel länglich, ‚‚kantig“. 9-11 Supra- und 8-9 Sublabialia. 4-6, meist

aber 5 Transversalbänder.

Beschreibung:

Die Art wird bis ca. 120 mm lang, ihr Kopf-Rumpf/Schwanz Längenindex entspricht 1,1. Der Schä-

del ist etwa 1,1- bis 1,5mal so lang wie breit. Zwischen 9-11 Supra- und 8-9 Sublabialia sind ausgebil-

det. Auf dem dorsalen Ohrrand liegen meist zwei kleinere Schuppengranulae. 14-18 dorsale Tuberkel-

längsreihen können ausgebildet sein. Zwischen Nasenloch und Augrand verläuft je ein heller, dunkel

gesäumter Streifen, die Lippenschilder sind einheitlich weiß. Die Rückenzeichnung besteht aus 4-6

meist dünnen Transversalbändern.

Maßtabelle

Tarentola caboverdiana nicolauensis von S. NICOLAU

Mentale Tub. Lam.

Nr. ZSM CIZERRZESEE NT FERB © Ar OA AS MS IE 1% KOILAUIE IR: Bel

138.81.1 115,08572055840216 951358722 SEA 0 6, 75 OO SER 9/8 18 12/14 5

138.81.2 0,20 32,0 30 ZINN ee 7 a ee 10/9 14 12/14 5

138.81.3 DR 0245, 024620212, 8511105922205 355,4787658,,,977.0535 80145 10/9 16 12/14 5

138.81.4 8420.43, 02414 0215572.11105222403, 272570, 60a 10/8 15° BABrE5

138.81.5 = 57.022 1550155858755, 27 0575er 10/8 18 12/14 5

138.81.6 - H2RO Et 108 203595527740 AR6 2RO 11/8 16. [BHBzE5

138.81.7 - (SL) EZ lee) MS 2 ee ee, eo ARE) 9/8 18 12/14 6

138.81.8 = Ikea 1 I ae aloe 11/9 Io aD >

138.81.9 1720500805572. 20 I EB SEP BESTE 9358469, 9/9 16 14/14 5

138.81.10 10553570259, 0, PAD TZD BT ED 5107727 3A A, 11/9 16 12/14 4

138.81.11 _ SS 0 OB SEO, 10/9 17 BMAZ5

4.2.3.3 Tarentola caboverdiana raziana Schleich, 1984 (Taf. II, Fig. 8; Taf. III, Fig. 1; Taf. IV,

Fig. 5)

Material: 20 Ex. Holotypus - ZSM 133/81.1 von St. Luzia, Paratypen — ZSM 133/81.2-10 v. St. Luzia,

134/81.1-10 v. Razo.

1984 Tarentola caboverdianus razianus SCHLEICH

1984 T. c. raziana. — JOGER

Terra typica: St. Luzia

Verbreitung: St. Luzia, Razo

Diagnose (s. SCHLEICH 1984: 101):

Kopf-Rumpflänge bis 60 mm; die Kopflänge beträgt 1,2-1,6mal der Kopfbreite. Die Entfernung

zwischen Auge-Schnauzenspitze ist deutlich länger als jene zwischen Ohr und Auge. 9-11 Supra- und

8-9 Sublabialia sind ausgebildet. 16 Tuberkelreihen und 3 bis max. 4 Transversalbänder zieren den

Rücken.

Beschreibung:

Lediglich ein Individuum der Serie von 20 Exemplaren hat einen unregenerierten Schwanz, der et-

was länger als die Kopf-Rumpflänge ist. Die KR-Länge beträgt max. 60 mm. Die Kopflänge mißt

1,2-1,6mal der Kopfbreite. Der kleine Ohrschlitz ist nur etwa halb so lang wie der Augdurchmesser.

Der Abstand vom vorderen Ohrrand zum Auge ist deutlich kürzer als jener zwischen Augvorderrand

und Schnauzenspitze. Zwischen 9-11 Supra- (meist 10) und 7-9 (meist 8 od. 9) Sublabialıa sind ausge-

bildet. Die Tuberkel sind in 16-18 Längsreihen angeordnet und von rundlich kegelförmiger, leicht ge-

kielter Gestalt. Die Tiere sind einheitlich braungrau gefärbt und tragen drei bis vier sich caudal verdun-

kelnde Transversalbänder, wobei entweder das 4. oder 5. auf der Schwanzbasis liegt. Das Schwanz-

regenerat ist bei lebenden Exemplaren rötlich. Zwischen Nasenloch und Ohröffnung verläuft je ein

heller, dunkel gesäumter Streifen. Zumindest die Sublabialıa sind kaum oder nicht gefleckt und weiß.

Maßtabelle

Tarentola caboverdiana raziana von St. LUZIA

Mentale Tub. Lam.

Nr. ZSM GIZEIRRTESISERITERBEE OZZENGEOATENSENSPERTE B; OL ORG ae

133.81.1 = rare ee ee 0 5 ll 3,0 2,8 10/9 16 10/13 4

133.81.2 = rd N I 2.5 2 6 7 Me DZ 1/9, 16 _ 4

IS = Sa re DE A Di 2 11/9 16 _ 3

Tafel III

Tafel III: Fig. 1: Tarentola caboverdiana raziana von Razo; adultes Exemplar. Fig. 2: Tarentola gigas gigas von

Razo; frisch geschlüpftes Jungtier. Fig. 3: Tarentola gigas brancoensis von Branco; adultes Exemplar. Fig. 4: He-

midactylus bouvieri boavistensis von Sal; Straße nach St. Maria; adultes Tier. Fig. 5: Hemidactylus bouvieri razo-

ensis von Razo; adultes Tier. Fig. 6: Hemidaciylus brooki angulatus von Sal; adultes Tier.

133.81.4 - Ries Are Deo So) 2 9/8 16 - 3

133.81.5 - ar A 3 NO Al 11/9 16 - 3

133.81.6 - Sl, = DO 20 are ra ee 10/8 16 = 4

133.81.7 _ u) = 17 NO 2 ee Te en 10/9 16 _ 4

133.81.8 = SB. 0m OS 358, 510072010585 47700226 10/8 16 - 3

133.81.9 - DO Fer 43076: 0,6, 925] 9/8 16 - 4+

133.81.10 - A902 116551052 40 50 AO 6:83.45, 2°0 10/9 16 - 4

von RAZO

134.81.1 - 60,022 28,52.114.02207 74,87 16,2. 7.07 Ne 10/9 16 - 3

134.81.2 - Ad 1 le ie 7a OD 7 250 10/9 16 - 4

134.81.3 _ 56:0 8315029407 76:25 50er 10/9 16 - 4

134.81.4 89, 05242,5240,595:1353331105955270, 23597 55: 077208117 2E4575229 9/8 18 - 4

134.81.5 = 48:02 215, 0ER E2XOF 4075,07 00 0270 10/9 18 - 3

134.81.6 - 42,0 A503 AO EE5S O7 11055402209 10/8 16 - 4

134.81.7 _ SB, 0,2 21650 ROSEN O5 20700122052 10/8 16 - 4

134.81.8 - ee TO re eb ler 10/7 18 _ 4

134.81.9 = 54,0, Zee 55725150 4,6, 255 10/8 18 - 4

134.81.10 = ee ie re ee 10/8 18 - 4

4.2.3.4 Tarentola caboverdiana substituta Joger, 1984 (Taf. II, Fig. 7)

Material: 17 Ex. ZSM 371/78; 140/81.1-10.

1984 Tarentola caboverdianus caboverdianus. — SCHLEICH (part.)

1984 Tarentola caboverdiana substituta. — JOGER

Terra typica: $. Vicente

Verbreitung: $S. Vicente

Diagnose (s. JOGER 1984: 103):

„Schuppen kleiner als bei den anderen Subspezies, daher mehr Schuppen um die Körpermitte:

146-167. Vorderbeine kürzer als die Kopflänge (bei allen anderen Kapverdengeckos mindestens

ebenso lang). 4-5 Rückensättel, caudal oft von weißen Tuberkeln umrahmt, vom Nacken bis in die

Sakralregion. Maximale Kopf-Rumpf-Länge 60 mm (J), 56 mm (P).“

Anmerkung: Eigenartigerweise erwähnte JOGER das ihm zur Verfügung gestandene Münchner Ma-

terial hier nicht. Es ist zu hoffen, daß das von ihm beschriebene Material dieser zum Großteil lang zu-

rückliegenden Aufsammlungen wirklich von $. Vicente stammt und dessen Herkunft - wie bei vielen

anderen alten Kapverdenkollekten - nicht nur gemutmaßt wurde.

An den von mir selbst gesammelten Tieren von $. Vicente und deren untersuchten Merkmalen erga-

ben sich für mich keine signifikant genug erscheinenden Differenzierungsmöglichkeiten für einen be-

rechtigten eigenen Unterartstatus, so daß ich nach den von JOGER untersuchten Merkmalen die Art hier

übernehme und die morphometrischen Werte der Münchner Exemplare hier mit anführe.

Beschreibung:

Für die von mir selbst gesammelten und in der ZSM hinterlegten Exemplare gilt:

Die Kopf-Rumpflänge des größten Exemplares beträgt 57,3 mm bei einer Schwanzlänge von

55,7 mm; hieraus ergibt sich eine max. Gesamtlänge für die gefangenen Exemplare von 113 mm.

Die Kopflänge mißt 1,2 bis 1,8(MW 1,6)mal der Kopfbreite. Der Ohrspalt ist etwa so lang oder grö-

ßer als der halbe Augdurchmesser.

Der Abstand vom vorderen Ohrrand zum Auge ist kürzer als der vom Augvorderrand zur Schnau-

zenspitze. Die Länge des Maulspaltes reicht deutlich hinter den hinteren Augenrand. Das Mentale ist

etwa doppelt so lang wie breit. An Lippenschildern sind 8-11 Supra-/, 7-9 Sublabialia ausgebildet.

Zwischen 14 u. 20 Tuberkelreihen können gezählt werden. Die Rückenzeichnung besteht aus

4-5 Bändern.

8 9

Tafel IV: Dorsolateraltuberkel kapverdischer Tarentola; REM-Aufnahmen. Fig. 1: Tarentola rudis maioensis von

Maio; 22x. Fig. 2: Tarentola vudis rudis von Praia-S. Thiago; 25x. Fig. 3: Tarentola darwini von Tarrafal-

S. Thiago; 24x. Fig. 4: Tarentola caboverdiana nicolanensis von S. Nicolau; 22X. Fig. 5: Tarentola caboverdiana

raziana von St. Luzia; 23x. Fig. 6: Tarentola caboverdiana caboverdiana von St. Antäo; 23x. Fig. 7: Tarentola

rudis cf. protogigas von Fogo; 24%. Fig. 8: Tarentola gigas brancoensis von Branco; 29x. Fig. 9: Tarentola gigas

gigas von Razo; 26x.

Maßtabelle

Tarentola caboverdiana substituta von S. VICENTE

Mentale Tub. Lam.

Nr. ZSM SL IR SL SC 18 (© A OA AS MSP L B OL/UL R. Bd

377,821 - 40,9 —- = 9202.2,05.4,05 4:0556: 02111070, = - 10/8 14 = 4

140.81.1 - 21,07 2 8,1 45 - 1,9, Pa 255 - 9/8 14 - 4

140.81.2 50=26:0224:082.9,0725,02 A035 6,0 - 10/7 20 - 4

140.81.3 - 2.0 = Ah Fe a Zee EZ az 10/8 18 = 4

140.81.4 _ 0.0 = I Tai 2 ee ae le 8/7 16 - 5

140.81.5 0 An ee ler re 90 or 2,0 9/8 14 _ 4

140.81.6 105410555205 527 0.218.012 52 24 24126387, 011240) 4585255 9/9 18 - 5

140.81.7 IE 056:0555, 0217, 0 EN 52,5 E24,93 76:10 7971270 O2 11/9 16 - 5

140.81.8 15208575355577..118,05 125527822, 556:555,9X 01140) 530238 10/9 16 - 5

140.81.9 100:0=55,0=25,0=15,05.19552 2757 2750561022770 2105 5022. 10/8 16 - 4

140.81.10 1072050585502 218702 330223 1 6 IE 752 11/9 14 - 4

4.2.4 Tarentola gigas (Bocage, 1875)

1875 Ascalabotes gigas BOCAGE

1884 Platydactylus gigas. -— ROCHEBRUNE

1885 Tarentola gigas. — BOULENGER

1896 Tarentola gigas. -— BOCAGE

1907 Tarentola giganten. — SCHERER

1937 Tarentola gigas. — ANGEL

1947 Tarentola delalandii gigas. — LOVERIDGE

1955 Tarentola delalandii gigas. — MERTENS

1979 Tarentola delalandii gigas. — SCHLEICH

1980 Tarentola delalandıi gigas. — SCHLEICH

1982 Tarentola ‚‚delalandii“ gigas. — SCHLEICH

1982 Tarentola delalandı gigas. - GRUBER & SCHLEICH

1983 Tarentola delalandü gigas. - BAEZ & SANCHEZ-PINTO

1983 Tarentola delalandii gigas. —- SCHLEICH & WUTTKE

1984 Tarentola gigas. — SCHLEICH

1984 Tarentola borneensis. — JOGER

4.2.4.1 Tarentola gigas gigas (Bocage, 1875) (Taf. III, Fig. 2; Taf. IV, Fig. 9)

Material: 17 Ex. ZSM 131/1981; Razo.

1875 Ascalabotes gigas BOCAGE

1884 Platydactylus gigas. — ROCHEBRUNE

1885 Tarentola gigas. — BOULENGER

1896 Tarentola gigas. — BOCAGE

1907 Tarentola giganten. — SCHERER

1937 Tarentola gigas. — ANGEL

1947 Tarentola delalandi gigas. —- LOVERIDGE

1955 Tarentola delalandii gigas. -— MERTENS

1984 Tarentola gigas gigas. — SCHLEICH

1984 Tarentola b. borneensis. — JOGER

Terra typica: Razo

Verbreitung: Razo; s. a. SCHLEICH & WUTTKE (1983). Die JOGER’sche Verbreitungsangabe (1984: 100) erscheint

unbelegt, stammt aber vermutlich aus SCHLEICH & WUTTKE (op. cit.).

Diagnose:

KR-Länge bis 13,5 cm; kräftig gackernde Stimme. Kopflänge 1,2- bis 1,8mal der Kopfbreite. Kleine

Rückentuberkel in 16 Längsreihen. Entfernung Ohr-Auge nur geringfügig kleiner als Entfernun

Auge-Schnauzenspitze. Zwischen 9 und 12 (meist 10) Supra- und 7-9 (meist 8 oder 9) Sublabialia.

Schwanz deutlich kürzer als KR-Länge. Mentale etwa doppelt so lang wie breit. 2-3 Submaxillaria sind

ausgebildet.

Beschreibung:

Der kräftige Habitus der bis 13,5 cm KR-Länge (freilebend) messenden plumpen Tiere unterschei-

det sie deutlich von allen anderen Arten der Gattung Tarentola. Das schwerste ©’ von über 80 auf Razo

gemessenen Exemplaren wog 81 g. Geschlechtsdimorph sind die O'C’ von den PP durch ihren deut-

lich breiteren Kopf zu unterscheiden. Der Maulspalt reicht bis hinter den Pupillenspalt. Die Tuberkel

der in einzelnen Querreihen angeordneten Schwanzsegmente sind alternierend unterschiedlich groß.

Die Rückenfärbung der Tiere ist meist hellgrau, die Bauchseite cremefarben. Die Zeichnung besteht

aus in der Dorsomedianen unterbrochenen, aber auch durchgehenden Transversalbändern, die poste-

rior dunkel gesäumt sind. Die Flanken sind gesprenkelt. Zwischen Nasenlöchern und Ohr verläuft ein

dunkel gesäumter, heller Streifen. An einem Exemplar wurden an der ersten Zehe 14 und an der dritten

15 Subdigitallamellen gezählt. Eine Auswertung der umfangreichen Freilanduntersuchungen befindet

sich in Bearbeitung.

Maßtabelle

Tarentola gigas gigas von RAZO

Mentale Tub. Lam.

Nr. ZSM GL KR SL KL KB © A OA AS MSP L B OL/UL R. 5 adl

131.81.1 1:80 02105,0775,05 0, 022155722438725. 97712 0713 a1 ES 10/9 16 - 5

131.81.2 184,0 98,0 86,0 27,0 22,0 4,0 6,0 12,5 13,0 20,0 7,8 3,3 10/7 16 - 4

131.81.3 — 1082006 253.0,37235,07 5.07 6:012,5213597 2240728722430 10/9 16 = 4

131.81.4 -— 120,0 - 36,3 26,0 5,0 6,4 15,0 15,0 25,0 8,9 4,5 10/9 16 - 4

131.81.5 165508288:.0275, 02220219, 0 54,85 25772 10 0 E57 527, A355 10/9 16 = 5

131.81.6 212507120,0792,0/ 34,5, 25,0 24,57 76,5. 13,42 16,.0724,379,7. 452 12/9 16 - 6

131.81.7 — 520,022 735072653: 74555 76,55 14,74 16,0035525,9587 2550 10/8 16 14/15 6

131.81.8 — O3 0 PZFSERR, 26:05 12585135 OB - 9/9 16 _ 5

131.81.9 — 527.072 235, 0925,0 24555 26,35. 15,0516,2292583710,07274,0 11/9 16 = 6

131.81.10 — 117,0 - 31,0 26,5 4,5 6,0 14,0 14,5 19,0 9,0 3,8 10/8 16 - 4

131.81.11 271610511116, 0/100,0729X0.724 567 74,07 25,42 13,52 14,3 21755, 78,173,9 10/9 16 - 4

131.81.12 29320, 171140924072 370224, 02385 25,52.,121e5135068118,22@787 2356 10/7 16 _ 4

131.81.13 — 105,0 - Ze a Ne 1256 II 7 10/8 16 - 4

131.81.14 1924051072.0585,0229,55.22,0723,27 25.521052 135 6,419,077:98 5354 10/8 16 _ _

131.81.15 1614 02285,027706:022558. 318,0, 54.05 55,52 10 DS, 0 E F7EE350 9/8 16 - 4

131.81.16 199105107,02831022970221,67 24,35 76,55 14, 51.552. 2055, 28,022440 9/8 16 _ 5

131.81.17 — E50 33, 0528357 74,55 26, 015,52 15,022556, 10,027555 9/8 16 _ 4

4.2.4.2 Tarentola gigas brancoensis Schleich, 1984 (Taf. III, Fig. 3; Taf. IV, Fig. 8)

Material: 18 Ex. Holotypus — ZSM 362/78.1, Paratypen - ZSM 362/78.2-6, 19/82.1-12.

1875 Ascalabotes gigas BOCAGE

1884 Platydactylus gigas. - ROCHEBRUNE

1885 Tarentola gigas. — BOULENGER

1907 Tarentola giganten. — SCHERER

1937 Tarentola gigas. — ANGEL

1947 Tarentola delalandıi gigas. — LOVERIDGE

1955 Tarentola delalandıi gigas. — MERTENS

1980 Tarentola delalandii gigas. — SCHLEICH

1982 Tarentola ‚‚delalandii“ gigas. — SCHLEICH

1984 Tarentola gigas brancoensis. — SCHLEICH

1984 Tarentola b. borneensis. — JOGER

Terra typıca: Branco

Verbreitung: Branco

Diagnose:

Unterscheidet sich von der Nominatform durch geringere Körpergröße. KR-Länge nur bis 11,4 cm.

Kräftig gackernde Stimme. KL 1,2- bis 2,2mal KB; kleine Rückentuberkel in 16 Längsreihen; Entfer-

nung OA nur wenig kleiner als AS; Ohröffnung ?/s bis mehr als die Hälfte des Augdurchmessers. Zwi-

schen 8-12 Supra- und 7-9 Sublabialia. Schwanz kürzer als KR-Länge. 2-3 Submaxillarıa.

Beschreibung:

Die Maximalwerte von Geländemessungen an 100 Tieren ım Frühjahr 1981 betrugen 11,4 cm KR-

Länge mit einem Gewicht von ca. 53 g. Durch ihren kräftigen, gedrungen wirkenden Körperbau sind

sie von den anderen Tarentola- Arten gut zu unterscheiden. Der Ohrspalt ist meist größer als die Hälfte

des Augdurchmessers. Die Entfernung zwischen Auge/Schnauze ist nur wenig größer als zwischen

Ohr/Auge. Falls eine Transversalbänderung ausgebildet ist, zählt man zwischen 4 und 6 Querbänder,

die jedoch selten durchgehend sind, sondern durch einen hellen Dorsomedianstreifen oder längliche,

miteinander verschmolzene Kernflecken getrennt sein können, die wiederum anterior dunkel gesäumt

sind. Weitere Angaben s. SCHLEICH, 1980.

Eine Auswertung der umfangreichen Freilanduntersuchungen befindet sich in Bearbeitung.

Maßtabelle

Mentale Tub. Lam.

Nr. ZSM GEEIKRESLIKE CKBLTO A 4 IOAN ZASZMSPL Bi: !OL/UE RZ YytaBa

| 75400.42.0.33.07440909.8 1537 4;0,5,3°'7,0710,07 451,3 9/8 16 - -

19.82.2 - 113,0 - 35.074025 13203, 12/9 16 - 5

19.82.3 - 98.54 —- 26020304537 555711,5°12,2° 208,0’ 7,0 3,6 11/9 16 - 5

19.82.4 — 10807 E7312724.0 4,1769 73, BIT 753 750 12/8 16 - 5

19.82.5 _ 98.077 781,972293,6 6,1 12,0 13,0720,37 8.1 4,0 12/9 16 - 6

19.82.6 u ER e e e e B Aolı IAZE EI EN et 1179 16 - 5

19.82.7 - 99,0 - - 2.548 60 2,0 30 195 30740 9/8 16 - 5

19.82.8 = 408.0 TB 5222 Bez 8/8 16 - 4

19.82.9 - 92.0°— "25,07 20,673, 5.374132, 3555 7.0 30 10/7 16 - 4

19.82.10 174:0° 94.080.0255722,0 3,2 5,2 10,1712;0 46,2 6,7 3,1 11/8 16 _ _

le Rs | - 22.0 25,049,53.0%5 57410.071,53. 3.077.132 9/7 16 - +

19.82.12 — #100 750.024 47357563037 9/8 16 _ 4

362.78.1 - 9137,85 2,2555, 3 11/9 16 13/14 5

362.78.2 - ar - 232 150 32 350739 102 165 ©&1 31 9/7 16 _ -

362.78.3 - 35.0 = 20 853 = - - - - - - - _ _ _

362.78.4 - 87.8 - 29.0 26,0 3,2. 5557 16.9, 11,0, 19,0. 753 30 10/9 16 - “

362.78.5 - 500 — ' 25,0721,07 2,8 4,8 1051107180 75 3,0 10/8 16 - -

362.78.6 = 920 = .e28,5 2255 12,6 - 5561100: 12,0) 19,0 77,3 738 9/8 16 _ -

4.2 Gekkonidae - Genus Hemidactylus

Anmerkung: Herr Dr. R. CromBıE, Washington, teilte mir freundlicherweise mit, daß Hemidacty-

lus cyanogaster von GirARD (1858) von den Kapverden beschrieben wurde, jedoch erscheint eine

Zuordnung bislang noch nicht möglich').

4.2.5 Hemidactylus brooki angulatus Hallowell, 1852 (Taf. III, Fig. 6)

Material: 2 Ex. ZSM 149/1981; Praia, S. Thiago. 7 Ex. ZSM 151/1981; Sal. 4 Ex. ZSM 150/1981; S. Vicente.

4 Ex. ZSM 148/1981; Fogo.

1852 Hemidactylus angulatus HALLOWELL

1858 Hemidactylus cyanogaster. — GIRARD

1937 Hemidactylus brookı. -— ANGEL

1947 Hemidactylus brookiı angulatus. — LOVERIDGE

1951 Hemidactylus brookii angulatus. — DEKEYSER & VILLIERS

1955 Hemidactylus brookiı angulatus. — MERTENS

1982 Hemidactylus brooküi angulatus. — SCHLEICH

Terra typica: ‚‚afrikanische Westküste“

Bisherige Verbreitung: Sal, Boa Vista, S. Thiago, Fogo, Brava, St. Antäo, $. Vicente, IIheu Grande

Nachgewiesen von: S. Thiago, Sal, S. Vicente, Fogo, ? Ilheu St. Maria

Artcharakteristik nach:

LOVERIDGE (1947) —

KR-Länge 51-69 mm; gewöhnlich 8 (4+4) Tuberkel auf4 Schwanzsegmenten. Tuberkel der Dorsolateralia drei-

flächig; weniger Supralabialia, durchschnittlich 8.

Diagnose:

KR-Länge bis 54 (69) mm, im Habitus ähnlich Hemidactylus brookı brooki mit Rückentuberkeln in

10-20 Längsreihen.JJ mit 22-25 Präanofemoralporen; 4-6 Subdigitallamellen am ersten und 7-8 am

dritten Finger.

Maßangaben für vom Autor gesammelte Exemplare:

Sal KR GL S. Vicente KR GL

ZSM 151/81 6,0 cm 12,4 cm Mindelho 5,4cm

5,4cm reg ZSM 150/81 6,1cm

5,0cm reg DANCE

4,6cm reg 1,9 cm 5,1cm

4,8cm reg

3,6cm 7,6 cm Fogo,S. Felipe 5,2cm

4,8 cm ZSM 148/81 5,4cm

4,6cm 6,1cm

5,9cm

S. Thiago 5,9cm

Praia 3,2 cm 5,1cm 10,7 cm

ZSM 149/81 3,6cm

") Während der Drucklegung der Arbeit teilte mir Dr. R. CROMBIE/Washington freundlicherweise die Erwähnung

von Hemidactylus cyanogaster durch GIRARD (1858) für die Kapverden mit, wofür ich ihm herzlich danken

möchte. GIRARD gab eine kurze Beschreibung und Charakteristik sowie eine relativ umfangreiche Synonymie

für die von ihm „‚ım Oktober 1838 nach dem Leben gezeichneten Tiere von Praia (‚Port Praja, San Jago‘)‘‘, wor-

aus ersichtlich wird, daß er sein Material seinerzeit (1858: 284-285, Taf. 25) Hemidactylus turcicus zuschrieb.

4.2.6 Hemidactylus bouvieri (Bocourt, 1870)

4.2.6.1 Hemidactylus bouvieri bouvieri (Bocourt, 1870)

Material: 2 Ex. BMNH. 66.4.12.3,4; S. Vicente.

1870 Emydactylus bouviere BOCOURT

1873 Hemidactylus cessacii. -— BOCAGE

1884 Hemidactylus bouvieri. - ROCHEBRUNE

1885 Hemidactylus bouvieri. -— BOULENGER

1935 Hemidactylus bouvieri. — ANGEL

1937 Hemidactylus bouvieri. -— ANGEL

1947 Hemidactylus bouvieri bouvieri. — LOVERIDGE

1955 Hemidactylus bouvieri bouvieri. -— MERTENS

1982 Hemidactylus bouvieri bouvieri. — SCHLEICH

1982 Hemidactylus bouvieri bouvieri. - GRUBER & SCHLEICH

Terra typica: $. Vicente

Bisherige Verbreitung: $. Vicente, St. Antäo, $. Thiago, Fogo, Brave

Nachgewiesen von: $. Vicente

Artcharakteristika nach:

MERTENS (1955) —

Von einem Exemplar von Praia; unter dem 1. Finger 4 (5, falls prox. dazugezählt wird) Haftlamellen. Oberseite

dunkelgrau, durch 4 schmale, helle Querbinden unterbrochen. 7 Querbänder auf der Schwanzoberseite, jene auf

der Schwanzwurzel bilden nach hinten gerichteten Winkel.

LOVERIDGE (1947) —

Schnauze wenig länger als Entfernung zwischen Aug- und Ohröffnung. Schnauzengranula größer als rückwärti-

ge. Nasenlöcher begrenzt vom Rostrale, 1 Labiale und 3—4 kleinen Nasalıa.

7-8 obere Labialia, 6-7 untere Labialia. Ein Paar große Postmentalia in Kontakt mit Mittellinie sowie ein äußeres,

aber kleines Paar Kinnschilder. Rücken mit ziemlich großen einheitlichen Granula. Ventralia glatt, gerundet,

schindelartig. O0" 2 Präanalporen; Finger frei, leicht verschmälert mit ziemlich kurzen Endgliedern, innerhalb

Haftlamellen, 3 unter 1. Zehe, 4-5 unter Mittelzehe. Schwanz zylindrisch, dorsal mit glatten einheitlichen Schup-

pen, ventral mit glatten Schindelschuppen, deren Mittelreihe transversal vergrößert ist. Schwanzlänge geringfügig

kürzer als KR-Länge.

Färbung: Dorsal schwach braun, mit dunklem Streifen von den Nasenlöchern über das Auge, der auf der Flanke

und der Schwanzbasis fortgesetzt sein kann. Oberlippe weiß. Rücken mit 5-6 dunkelbraunen Querbändern, ventral

hell, GL 67 (36+31) mm.

Diagnose:

Hemidactylus bouvieri bouvieri mißt bis ca. 38,5 mm KR-Länge. Unter der ersten Zehe besitzt

Hemidactylus bonvieri bonvieri 344 und unter der 3. Zehe 4-5 Subdigitallamellen. Eine dorsale Bän-

derzeichnung ist für die Art charakteristisch, muß aber nicht immer ausgebildet sein.

Beschreibung:

Hemidactylus bouvieri unterscheidet sich von Hemidactylus brooki durch das Fehlen der Rük-

kentuberkel. H. bouvieri konnte leider nie selbst von mir gefangen werden, so daß sich diese Angaben

auf Leihmaterial aus dem Britischen Museum sowie auf Literaturangaben beziehen (s. GRUBER &

SCHLEICH 1982). Die Größenvariabilität von 5 Exemplaren konnte in einem Diagramm (op. cit.: 305,

Abb. 2) dargestellt werden. Die maximale Kopf/Rumpflänge scheint ca. 38 mm zu betragen und liegt

damit zwischen razoensis und boavistensis. Die Anzahl der Subdigitallamellen beträgt 34 an der er-

sten und 4-5 an der dritten Zehe. Das Supralabiale ist mit der Nasenöffnung in Kontakt. Die beiden

vorliegenden Exemplare tragen 5 sich caudal verdunkelnde braune Transversalbänder, wobei das erste

ım Nacken und das 6. auf Höhe des Oberschenkelansatzes beginnt.

4.2.6.2 Hemidactylus bouvieri boavistensis Boulenger, 1906 (Taf. III, Fig. 4)

Material: 8 Ex. ZSM 366/1978; Boa Vista. 9 Ex. ZSM 20/82; Sal, Straße zur Saline. 3 Ex. ZSM 21/82; Sal, Straße

nach St. Maria.

1906 Hemidactylus boavistensis BOULENGER

1935 Hemidactylus chevalieri. - ANGEL

1937 Hemidactylus chevalieri. - ANGEL

1937 Hemidactylus boavistensis. — ANGEL

1947 Hemidactylus bouvieri boavistensis. — LOVERIDGE

1947 Hemidactylus bonvieri chevalieri. - LOVERIDGE

1951 Hemidactylus bouvieri chevalieri. - DEKEYSER & VILLIERS

1955 Hemidactylus bouvieri boavistensis. — MERTENS

1982 Hemidactylus bouvieri boavistensis. - GRUBER & SCHLEICH

Terra typıca: Boa Vista

Bisherige Verbreitung: Boa Vista, Sal

Nachgewiesen von: Boa Vista, Sal

Artcharakteristika nach:

BOULENGER (1905) —

In Pholidose und Morphologie Hemidactylus bouvieri ähnlich, aber ziemlich stark verlängerte Zehen und mit

mehr Lamellen. 4-5 davon unter der 1. und 6-8 unter der 3. und 4. Zehe. Das Paar Kinnschilder hinter der Sym-

physe ist ebenfalls länger als bei bouvieri. Dorsal gelb, rötlich oder blaß-graubraun, gewöhnlich dunkelbraun ge-

sprenkelt, mit 5 gebogenen und welligen braunen, dunkelgesäumten Querbändern, wovon das erste zwischen den

Augen über den Occiput und das zweite im Nacken läuft. Diese Bänder können von einer Reihe dunkelbrauner

Flecken entlang der Rückenmittellinie unterbrochen sein, oder sie sind alle zusammen kaum unterscheidbar. Ein

dunkler brauner Strich verläuft beidseits der Schnauze, mit einem gelblichen darüber liegenden. Schwanz mit mehr

oder weniger unterschiedlich halbmondförmigen dunklen Rückenbändern bei weißer Ventralseite. Bei Hemidacty-

lus bouvieri fehlen die hellen Streifen auf der Schnauze, und die dunklen Bänder seitlich des Kopfes reichen bis zum

dunklen Nackenband.

Boa Vista von 0-600 m

6) 2 6) 2

GL 81 83 Körper 28 29

SL 39 39 Vorderextr. 14 14

Kopflänge 14 15 Hinterextr. 18 19

Kopfbreite 10 11 (Angaben in mm)

LOVERIDGE (1947) —

Hemidactylus bouvieri boavistensis unterscheidet sich von Hemidactylus bouvieri bouvieri nur in leicht verlän-

gerten Fingern mit konsequent mehr Haftreihen, nämlich 4-5 unter der ersten Zehe, 6-8 unter der dritten und vier-

ten. Dorsal, gräulich, gelblich oder rötlichbraun, ein dunkelbrauner Streifen vom Rostrale ist von oben gelb be-

grenzt und reicht bis zum Auge. Von Auge zu Auge zieht ein welliges dunkelgesäumtes Band, ein weiteres im Ge-

nick, drei weitere auf dem Rücken, der auch dunkel gesprenkelt oder einfarbig sein kann. Die Bänder können mesial

von einer Serie dunkelbrauner Flecken unterbrochen oder beinahe weißlich sein. Dieser leichte helle canthal-Strei-

fen fehlt bei Hemidactylus bouvieri, wobei der dunkle sich niemals bis zum Nackenband oder den Flanken wie bei

bouvieri fortsetzt. - Größe: GesamtlängeQ’ 81 (42 + 39),Q 83 (44 + 39).

MERTENS (1955) —

Von 33 Exemplaren von Sal und Boa Vista

Hemidactylus chevalieri von Sal ist identisch mit boavistensis, da die Zahl der Lamellen unter dem 1. Finger (4-5)

sich auf 6 erhöhen kann. Zeichnung sehr variabel, neben Individuen mit dunklen Querbändern (1 Nacken-, 3 Rük-

ken-, 1 Schwanzwurzelband) haben einige eine helle, dorsale Mittellinie, andere dagegen sind bis auf eine feine

Tüpfelung zeichnungslos. Der regenerierte Schwanz ist meist hell mit 3 dunklen Längsstreifen.

Diagnose:

Von der Nominatform bouvieri bonvieri deutlich durch die etwa '/; größere Körperlänge unter-

scheidbar. Die Anzahl der Subdigitallamellen beträgt an der ersten Zehe 5-6 und an der dritten Zehe

7-8. Kopf/Rumpflänge bis ca. 50 mm.

Beschreibung:

Hemidactylus bouvieri boavistensis ähnelt im Habitus mehr einer kleinen Tarentola caboverdiana

als Hemidactylus brooki angulatus. Von letzterem ist er durch die gleichmäßig glattere Pholidose gut

zu unterscheiden. Das erste Supralabiale steht in Kontakt mit dem Nasale. Die Tiere erreichen eine Ge-

samtlänge von ca. 80 mm. Die Größenverteilung der gesammelten Exemplare ist in GRUBER &

SCHLEICH (1982: 305, Abb. 2) in einem Diagramm dargestellt.

Die Anzahl der Subdigitallamellen beträgt bei Hemidactylus bouvieri boavistensis 5-6 an der ersten

und 6-8 an der dritten Zehe.

Die Tiere können einfarbig hell gefärbt oder stark akzentuiert gebändert sein, wobei dann die meist

vier bis fünf Transversalbänder dunkel gesäumt sind.

4.2.6.3 Hemidactylus bouvieri razoensis Gruber & Schleich, 1982 (Taf. III, Fig. 5)

Material: Holotypus: ZSM 129/1981; Razo. Paratypen:5 Ex. ZSM 130/1981; Razo. 1 Ex. ZSM 22/1982; Razo.

1982 Hemidactylus bouvieri razoensis GRUBER & SCHLEICH

1983 Hemidactylus bouvieri razoensis. -— SCHLEICH & WUTTKE

Terra typica: Razo

Verbreitung: Razo

Diagnose (nach GRrUBER & SCHLEICH 1982):

„Die neue Form zeichnet sich durch ihre geringere Körpergröße (KR max. = 29 mm) sowie eine

verringerte Anzahl von Subdigitallamellen (1. Finger: 2-3; 3. Finger: 4, selten 5) aus. Die Nasenöff-

nungen stehen nicht in direktem Kontakt mit den ersten Labialıa.‘“

Beschreibung (nach GRUBER & SCHLEICH 1982):

Habitus

Die Schnauze der Tiere ist spitz, ihre Beine sind kurz; das nach vorne gelegte Hinterbein reicht nicht

bis zur Achselhöhle. Der runde Schwanz ist spitz auslaufend und an der Basis etwas dünner als in der

Mitte. Der Augdurchmesser beträgt durchschnittlich 1,9 mm.

Pholidose

Die Schnauze ist- vor allem zwischen Auge und Nasale — von gegenüber den Dorsalia vergrößerten,

oft hexagonalen Schuppen bedeckt. Von den beiden Postnasalia trennt das jeweils untere die Nasenöff-

nung vom ersten Supralabiale. In Ausnahmefällen kann das untere Postnasale — unter Verbleib einer

schwachen Naht - mit dem Labiale verwachsen sein. Je sieben bis acht Supralabialia sind ausgebildet,

wobei das letzte hinter der Augenmitte liegt. Sie sind durch mindestens eine Reihe kleiner Schuppen

vom Auge getrennt. Die Pileusschuppen sind nur geringfügig kleiner als die Dorsalıa.

Das nach hinten spitz zulaufende Mentale endet auf halber Länge zwischen dem ersten der beiden

aneinanderliegenden Paare der Postmentalia. Die je sechs bis sieben Sublabialia werden von vergrößer-

ten Gularıa gesäumt. Die Gularia sind von rundlicher Form, flach und kleiner als die Dorsalıa. Diese

sind glatt, rundlich und nach hinten zunehmend gewölbt. Alle Dorsalia sind annähernd gleich groß.

Die dorsalen Schwanzschuppen sind rechteckig, größer als die Dorsalia und in Querreihen angeordnet.

Die vergrößerten Ventralia haben eckige Gestalt, überlagern sich und sind in der Körpermitte ın ca. 25

versetzten Längsreihen angeordnet. Die stark vergrößerten Subcaudalia sind oft in nur einer Reihe vor-

zufinden. Alle Exemplare weisen beiderseits der Schwanzwurzel zwei, selten einen Postanaltuberkel

auf, die bei den &'C° deutlicher ausgeprägt sind. Die Anzahl der mit Haftborsten versehenen Subdigi-

tallamellen beträgt am 1. Finger drei und am 3. Finger vier (selten fünf oder eine zusätzliche, vergrö-

ßerte basale Schuppe). Weist der 1. Finger nur zwei Lamellen auf, ist meist ebenfalls eine vergrößerte

basale Schuppe zu erkennen. Das Rostrale ist hinten schmäler und von dort zu ”/z seiner Länge in der

Mitte gespalten. Im Anschluß daran befinden sich zwei, selten drei große Internasalia.

Färbung

Kopf und Körperseite sind sandfarben. Die Rückenzeichnung besteht aus fünf 2-3 mm breiten,

dunkelbraunen Transversalbändern, die nach caudal zunehmend dunkler werden. Das erste befindet

sich am Nacken, das letzte an der Schwanzbasis. Auf der Schwanzoberseite verwischt die Bänderstruk-

tur der Zeichnung, während die Grundfarbe des Schwanzes in Orange übergeht. Der vordere, obere

Augenrand ist gelblich, ebenso die Labialia. Über den Supralabialia zieht sich ein dunkler Streifen von

der Schnauzenspitze über das Auge bis zum lateralen Ende des ersten, selten des zweiten Transversal-

bandes. Ventral dieses Streifens erstreckt sich ein weißes Lateralband vom Mundwinkel bis zur

Schwanzwurzel. Die Bauchseite ist weißlich, etwas durchscheinend und mit feinen, schwarzen Spren-

keln überzogen.

Diskussion (siehe GRUBER & SCHLEICH 1982).

4.3 Testudines

Nur sehr wenige Literaturangaben gibt es zu Schildkrötenhinweisen auf den Kapverden.

BOULENGER (1905: 197) erwähnte ‚‚Chelone imbricata‘‘ von Fogo, und BANNERMAN & BANNERMAN

(1968: 40) berichten aus Bourne’s Expedition von 1951 über die Rhombos-Inseln: ... ,,as one comes

ashore on little beaches of white sand marked with the excavations of turtles“, und von Maio (op. cit.:

S. 14)...,,‚and ata certain Season ofthe Year, as May, June, July and August, a sort of small sea-tor-

toise came hithere to lay their eggs; but these turtles are not so sweet as those in the West Indies.“

SCHLEICH (1979) berichtete über eine ‚‚Sea Turtle Protection needed at the Cape Verde Islands“ und

erwähnt dabei ‚‚Hawksbill‘“ and ‚‚Loggerhead‘“ als die durch intensiven Fang gefährdeten Arten. In

den ‚vorläufigen Mitteilungen zur Herpetofauna der Kapverden“ SCHLEICH (1982: 247) werden vier

Arten von Seeschildkröten, die auf den Kapverden vorkommen und intensivst bejagt werden, erwähnt:

Caretta caretta caretta, Eretmochelys imbricata, Chelonia mydas mydas und Lepidochelys olivacea

olivacea.

Neben dem großen Schaden durch hohe jährliche Abfangraten und Sammeln sowohl der Tiere als

auch ıhrer Gelege durch Privatleute, Fischer und bislang der staatlichen Fischereiindustrie, wurden die

Bestände der Seeschildkröten stark dezimiert. Nicht unerwähnt soll trotzdem ihre Bedeutung auf den

Kapverden als Eiweißlieferant, aber auch die Verwendung des Rückenpanzers als Transportbehältnis

sein. Nach meinem Bericht (SCHLEICH 1979) und intensivem Drängen bei den Fischereibehörden wurde

mir von Direktor M. DUARTE-ALMEIDA seitens der SCAPA versichert, daß der industriell-gewerbliche

Schildkrötenfang verboten wird. Andererseits werden natürlich noch immer genug Tiere auf dem

Markt angeboten oder direkt von den Fischern gehandelt. Auch das Absuchen weiter Sandstrände nach

frischen Gelegen seitens der Einheimischen sowie streunender Hunde führt zu einer weiteren intensi-

ven Bedrohung der Tiere. Die Verfolgung eines Schutzprojektes wäre hier dringlichst notwendig!

Als bevorzugt ‚‚angelaufene“ Inseln gelten natürlich alle mit flachen Sandstränden und Buchten, so

besonders Sal, Boa Vista, Maio, S. Vicente, St. Lucia und Branco. Auf Maio wurde mir berichtet, daß

die Seeschildkröten zwischen Mai und Juni zur Eiablage an Land gehen und dort sofort von den Ein-

heimischen gefangen werden. Offiziell sei nur eine Rate von ca. 70 Tieren bekannt, Herr M. Rıvas

(frdl. mdl. Mittlg./20.1.81) nahm jedoch eine Dunkelziffer von mehr als 100 Tieren pro Jahr an.

Das von MErTENSs (1955) für Ilheu St. Maria (in der Hafenbucht vor Praia gelegene kleine Insel) er-

wähnte Vorkommen von Süßwasserschildkröten der Art Pelusios subniger derbianus wurde von mir

(SCHLEICH 1982) als nicht mehr existent und seinerzeit anthropogen eingeführt, abgehandelt.

4.4 Amphibia — Bufonidae

Bufo regularıs Reuss

Material: 11 Ex. ZSM 18/1982; S. Nicolau. 3 Ex. ZSM 17/1982; S. Thiago.

SCHLEICH (1982) erwähnte erstmals Froschlurche von den Kapverden. Herr M. Tanpy bestätigte die

Zugehörigkeit der Kröten zu Bufo regularıs. Von Angestellten des Landwirtschaftsministeriums der

Kapverden (MDR) wurde versichert, daß die Kröten, bislang von S. Thiago und $S. Nicolau bekannt,

bereits zur Zeit der portugiesischen Kolonialherrschaft zur Dezimierung der Mückenplage in den öf-

fentlichen Wasserreservoirs eingesetzt wurden.

5. Die einzelnen Inseln

Zu ihrer Geographie und Biologie-Ökologie der einzelnen Arten (s. a. Pkt. 2)

Verwendete Abkürzungen:

LT°C = Luft-Temperatur in °C

BT’C = Boden-(Substrat)-Temperatur in °C

KT°’C = Körper-(Kloaken)-Temperatur in °C

LF% = relative Luftfeuchte in %

5.1 Sea

Sal, die südöstlichste, als einzig autonome Insel der Republik Cabo Verde hat eine Oberfläche von

ca. 200 km? und eine maximale Höhe von 406 m. Die gestreckte, ca. 24 km lange und nur knapp 10 km

breite Insel zeigt durch ihre geringen orographischen Unterschiede ein einheitliches Bild - eine mono-

tone Steinwüste in typischer Serirausbildung geprägt. Gelegentliche Sanddünen, Bodenbildungen und

Basaltberge wechseln das sonst eintönige Landschaftsbild.

Nur wenige größere Ribeiras und ebensowenige Oasen zeichnen sich ab.

Palmeira aPedra Lume

Ilheu de Rabo ,

de Junco D Ilheus

‚fe do Chano

Sudpedih VA S’A MARIA

Abb. 14: Sal; Straße nach St. Maria; Biotop von Mabuya stangeri salensis und Hemidactylus bouvieri boa-

vistensis.— Abb. 15: Dünenlandschaft auf Boa Vista; Biotop von Hemidactylus bouvieri boavistensis und Mabuya

delalandü. — Abb. 16: Im Inselinnern von Maio; Biotop von Mabuya stangeri maioensis und Tarentola rudis

maioensis.

Artenspektrum

Nachgewiesen wurden: Mabuya stangeri salensis, Hemidactylus brooki, Hemidactylus bouvieri

boavistensis. In der Geschichte der kapverdischen Herpetofaunistik wurden für Sal bereits 8 verschie-

dene Reptilienarten nachgewiesen, unter anderem auch eine Schlange, Psammophis sibilans sibilans

durch DEkEYsER & VILLiers (1951), die selbst aber berichteten, daß dieser einmalige Nachweis sicher auf

eine anthropogene Verschleppung zurückzuführen ist (s. auch SCHLEICH 1982).

MERTENS konnte für Salnur Mabuya stangeri salensıs als einzigen Skink bestätigen. Das von AnGeı

(1935, 1937) gemutmaßte Vorkommen von Mabuya (stangerı) spinalis, konnte von mır bislang nicht

bestätigt werden. Die 9 Exemplare umfassende Kollektion an Skinken beinhaltet ebenfalls nur Ma-

buya stangeri salensis. Die Variationsbreiten ihrer Merkmalsausbildungen sind in den Diagrammen

und in der Beschreibung (4.1.3.3) dargestellt. Leider gelang es mir auf keinem meiner insgesamt sechs-

maligen Aufenthalte auf Sal eine Tarentola nachzuweisen.

Fundorte

1 ca. 3 km vor Palmeira: Mabuya stangeri salensıs

2 Oase auf der Strecke nach St. Maria: Mabuya stangeri salensis und Hemidactylus bouvieri bouvieri, unter Pal-

menresten und Lesesteinen im Dünensand

Lagerhaus beim Flughafenhotel: Hemidactylus brooki angulatus

4 Steinwüste ca. 5 km südlich Flughafen: Mabuya stangeri salensis

Straße zur Saline (Pedra Lume): Mabuya stangeri salensis, Hemidactylus bouvieri boavistensis

Straße nach St. Maria: Mabuya stangeri salensis, Hemidactylus bouvieri boavistensis

Zur Biologie - Ökologie

Geländenotizen, Körpertemperaturen, Biotopcharakteristika:

16. 1. 81 — windig, bedeckt

Datum Uhrzeit LT LF BT Taxon KT Bemerkungen

€ % ® ®

61% 14.00 24 - 30 Mabuya 29 unter Lavafels auf Sandboden

28 45 28,5 Mabuya (70% LF, 23,5°C unter Stein);

Boden leicht lehmig feucht

14.45 - - - Mabuya 26,5 (24°C unter Stein)

Mabuya 25,0

15.008522 — 29 Mabuya 30,0 (28°C unter Stein, semiadult)

ab 20.00 - — - Hemid. 24-26 Lagerhaus bei Hotel

ab 24.00 - - 22 brooki

17.2. Hemid. kühl und windig, 7 Tiere gesichtet,

b. boav. Richtung Pedra Lume unter Steinen

Anmerkungen:

Bei einem Gespräch am 19.9. 1977 auf Sal (St. Maria) mit Einheimischen bezüglich Reptilienvorkommen auf der

Insel wurde mir einstimmig entgegnet, daß es auf Sal eigentlich gar nichts gibt, weder Pflanzen noch Echsen, und

nur im Norden der Insel vereinzelt kleine Skinke vorkommen.

Zur Biologie der Echsen insbesondere von der zweitgrößten kapverdischen Mabuyenart Mabuya

stangeri salensis ıst bislang noch nichts bekannt.

Die Kotanalysen erbrachten folgendes Nahrungsspektrum für die im Januar gefangenen Mabuyen:

Acridiidae, Curculionidae, Heteroptera, Myrmeleonidae, Tenebrionidae.

Für Hemidactylus brooki angulatus von den Lagerhallen des Flughafenhotels setzten sich die Rück-

stände wie folgt zusammen: Coleoptera, Tenebrionidae, Arachnidae, zahlreiche Blattaria und Ortho-

ptera.

Für Hemidactylus bouvieri boavistensis ergaben die Rückstandsuntersuchungen folgende Zusam-

mensetzung: Aranea, Tettigoniidae, Aphidina, Hymenoptera, Formicidae, Diptera, Coleoptera.

5.2 BOA VISTA

Boa Vista ist neben Sal und Maio eine der wüstenhaftesten bewohnten Inseln des Archipels, von

rundlicher Form, ca. 620 qkm Flächenausdehnung und mit 390 m höchster Erhebung ebenfalls relativ

flach. Der N-S-Mittelstreifen der Insel ist etwas gebirgiger und trennt die Bereiche der großen Sanddü-

nen, die hauptsächlich auflandig der Westküste der Insel auftreten. Das typische Landschaftsbild der

Insel sind vulkanische Gesteins- bzw. Gebirgsmassive mit Sanddünen, teilweise Sedimentgesteinen

und Oasen- bzw. Ribeirabewirtschaftung.

BEOFA VS A

Iheu dos Pässaros

a !!heu do Cascalho

Ilheu Holandes

Ilheu do Sal Rei ee

VADE SAL REI \

x Se ae @ Ilheu do Baluarte

ER N _>fFundo de Figueras

Ya „iCabego de Tarafes

Ilheu do

Lagosteiro

Artenspektrum

Neben Mabuya delalandü (ZSM 373/78) wurde noch eine größere Mabuya gesichtet. Leider

konnte das Tier nicht gefangen werden. An Geckos kamen noch Tarentola (nach Aussagen der Bevöl-

kerung) und Hemidactylus bouvieri boavistensis (ZSM 366/78) vor. Leider gelang uns nur der Nach-

weis von letzterem!

Fundorte

1 Aufgrund verkehrstechnischer Verbindungsschwierigkeiten waren wir auf ein Sammelgebiet im

Umkreis der Stadt Sal Rei beschränkt.

Steinwüste, Talalluvionen und Sanddünen beherrschen diesen Landschaftsabschnitt. Einzelne Lese-

steine oder gestürzte Palmstämme bzw. kleine Pflanzeninseln auf den Sanddünen waren Unter-

schlupf und Zufluchtsraum für Skinke (Mabuya delalandı, M. sp. indet.) und Halbfingergeckos

(Hemidactylus bouvieri boavistensis). Die Mauern der Straßeneinfriedungen waren ebenfalls belieb-

ter Zufluchtsort der Skinke.

Zur Biologie - Ökologie

Geländenotizen:

Auf eine Fläche von ca. 1-2 qkm kamen etwa 1 Skink und ca. 3 Geckos. In einem ca. 4 qkm großen Areal wur-

den 15 Hemidactyli und 8 Mabuyen gefangen. Am 22.7.1977 entdeckte ich unter einem Palmstück 37 Hemidacty-

lus-Eier, wovon zu diesem Zeitpunkt bereits etwa '/3 geschlüpft waren. Die Fluchtdistanz junger Mabuyen betrug

ca. 15 m.

Sehr hell klingendes ‚‚Geckogekreische‘“ war in Gefangenschaftshaltung von 2 verbissenen Hemi-

dactyli (? Streit, Kopula) sowohl am 9.3. als auch am 10.3.79 nachmittags zu hören. Weder vorher

noch nach diesem Zeitraum war jemals wieder solch eine Lautäußerung vernommen worden. Am

9.3.79 wurde ein rundes, kleines hartschaliges Ei von etwa 2 g und 0,85 cm Durchmesser abgelegt.

Von Einheimischen wurde mir berichtet, daß eine größere Geckoart (? Tarentola) innerhalb der

Häuser anzutreffen sei, leider glückte mir selbst der Nachweis bislang nicht.

An ‚‚natürlichen“ Freßfeinden konnten wir hauptsächlich Katzen sehen.

5.3 MAIO

Maio gehört neben $. Thiago, Fogo, Brava und den Rhombos-Inseln ebenfalls noch zur Sotavento-

gruppe und hat mit die kleinste Einwohnerzahl aller kapverdischen Inseln. Maio ist von ovalem Umriß

mit einer Flächenausdehnung von ca. 250 qkm und mit dem Mt. Renose im Inselzentrum, bis auf

436 m aufragend. Das Inselinnere ist relativ stark zergliedert. Tief eingeschnittene Ribeiras, Hochpla-

teaus, Sandstrände und Kalkgesteinmassive wechseln mit Steinwüsten, Sanden, Talalluvionen und

Vulkaniten ım Landschaftsbild. Plattig absondernde Kalkschichten bieten mit ihren wie scherbenüber-

streuten weiten Bodenflächen hervorragende Kleinstbiotope und Unterschlupf für Reptilien. Selbst

auf den Salzklippen der Salınen waren Mabuyen zu finden.

X Aue da Horta

Falk

Artenspektrum

Nachgewiesen wurden Mabuya stangeri maioensis sowie Tarentola rudis maioensıs.

Fundorte

1 Ribeira zwischen Vila do Maio und Morro (19.1.81): Mabuya, Tarentola

2 Salınas: Mabuya

Zur Biologie-Ökologie

Die Populationsdichte in einem Biotopausschnitt von ca. 100 qm betrug etwa 30 Skinke und 10 Ta-

rentolas, wobei meist 1-2 Skinke auf 1 Gecko unter einer Gesteinsplatte kamen.

Die Kotrückstandsuntersuchungen ergaben folgende Bestandteile für die Mabuyen:

Coreidae-Lygaeidae, Formicidae, Tenebrionidae

und für die Tarentolas:

Tettigonüdae, Blattaria, Cydnidae, Tenebrionidae, Acridiidae.

5.4 SAO THIAGO

S. Thiago (Santiagio, S. Jago) ist mit der Hauptstadt Praia flächenmäßig und der Einwohnerzahl

nach die größte aller kapverdischen Inseln. Die Insel ist von tropfenförmigem Umriß mit einer Flä-

chenausdehnung von ca. 1000 qkm und reicht mit dem Pico da Antonia auf eine maximale Höhe von

1392 m. Landschaftlich, orographisch wie faunistisch und floristisch istS. Thiago wohl die vielgestal-

tigste aller Cabo-Verde-Inseln. Die gesamte Insel ist relativ stark zerklüftet, mit tief einschneidenden

Ribeiras, schroffen Vulkanschluchten und grünen Plantagen. Herpetologisch von Bedeutung war die

kleine ehemalige Strafgefangeneninsel Ilheu St. Maria in der Hafenbucht vor Praia mit ihren alten Rui-

nenmauern.

Artenspektrum

Bislang waren von S. Thiago an Skinken nur Mabuya delalandii (und Mabuya stangeri) bekannt,

d. h. vor MERTENs’ Synonymisierung von Mabuya vaillanti mit Mabuya delalandiüi auch jene Art.

Nach der Bearbeitung der Geckos (s. Pkt. 4.2) verbleiben für S. Thiago Tarentola darwini (Tarrafal),

Tarentola rudis rudıs und Hemidactylus brooki angulatus.

An Skinken verbleiben so der größte Kapverdische Skink (kreol.: Chinel) Mabuya vaillanti (ZSM

364/78), Mabuya delalandıi (ZSM 363/78) und Mabuya stangeri spinalıs, die meist sympatrisch einen

Biotop besetzen.

Fundorte und Belege

1 Praia-Hafenstraße : Mabnya delalandii, M. stangeri

Tarentola rudis rudıs

2 Praia-Flughafen : Mabuya delalandıı

Tarentola rudis rudıs

3 Cidade Velha : Tarentola rudis rudıs

Mabuya delalandıi

4 S. Jorge dos Orgaos : Bufo regularis

Su arcafal : Tarentola darwiniı

Mabuya delalandıi

6 5km von Tarrafal : Mabuya vaillantı

Mabuya delalandıi

Mabuya stangeri

7 er Cm : Mabuya vaillanti

8 Ilheu St. Maria : Mabuya spec.

Tarentola spec.

Zur Biologie - Ökologie

Geländenotizen, Körpertemperaturen:

Die Messungen erfolgten kurz nach einem Regenschauer, alle Skinke wurden unter Steinen gefan-

gen.

Datums Uhrzeit LT JeE: BT Taxon KT Bemerkungen

IC % RC IC

I 17.00 24,5 90 22 M. del. 24,5 teilweise noch aktıv

17.15 25 90 23,50 Midel: 23,5

17.20 24,5 90 DOSE Madel. 2353

17.25 24,5 90 22,5 M. sta. 25 leichter Regen

12502215 2 DASS M Adel. uWE251 22%) Regen

18.00 21 - 25, M. del. 2332559525

21-21.30 20 _ 20 Tarentola 23; 23; 22,5; 23,5 BT = Mauertemperatur

18 7.0 01.9 95 - — windig

11.20 26,5 71 — - windig, Mabuyen aktiv

SE el. [en RED EINE)

-643 /

RN DO TARRAFAL 23°30°

IS. jo

»Rib. da. Prata / \ |

SIUWRZDN

Figleira BER Naus’

_1021( „..| J020 Teves z BES

1392 PA

) ‚>. Domingos I; - _#°” 2

| ; N Milho Branco-”

ee "550 / ste Ana

| Da are

23°50' \ RS PN N

"Ile ei \l)) AL bs Nova

& PRAIA

Cidade Velhs IIh&u de S.f@ Maria

Auffallend ist die äußerst hohe Konzentration von M. delalandii ım Bereich der Poussada Praia-

Mar bis zur Hafenbucht. Sie erscheinen dort augenfällig als Kulturfolger und profitieren dabei aus den

Fäkalien- und Abfallansammlungen am Straßenrand.

Interessant ist das Artenverhältnis in einem etwa 1 qkm großen Beobachtungsgebiet ca. 5 km süd-

lich von Tarrafal, wo auf 2 adulte und 2 juvenile Mabuya vaıllantı ca. 30 bis50 Mabuya stangeri und

einige hundert Mabuya delalandıi kamen. In der Nähe von Praia ließe sich ein Verhältnis von etwa 100

Mabuya delalandii auf 10 Mabuya stangeri und 1 Mabuya vaillanti angeben. In Straßenunterführun-

gen konzentrieren sich Geckos der Gattung Tarentola, die an Hauswänden nur seltenst anzutreffen

sind, dafür diese aber öfters von einzelnen Hemidactylus brooki angulatus besetzt sind.

Interessante Beobachtungen glückten bei der erfolgreichen Nachzucht von Mabuya vaillantı. Als

epigamische Reaktionen können ein hochfrequentes Vertikalnicken des Kopfes angesprochen werden,

anschließend verbeißen die O'C’ sich kurz in die P, um diese an der Flucht zu hindern, und dann zur

Paarung einen Flankenbiß anzusetzen.

Bei Paarungsunwilligkeit des @ verbeißt dieses das ©’. Die Kopula erfolgt mit Flankenbiß. Sowohl

kurz nach der Kopula als auch während dieser, kann das intervallartige hochfrequente Nicken während

des Verbissenseins bei beiden Partnern anhalten. Mabuya vaıllanti ist lebendgebärend. Die Größe der

in Gefangenschaft mehrmals nachgezüchteten, neugeborenen Jungtiere betrug ca. 8 cm.

In Terrarienhaltung konnte beobachtet werden, daß Mabuya vaillanti meist omniphag, weniger

Mabuya delalandii, und Mabuya stangeri nur noch carnıvor war.

Die Kotanalyse getrockneter Exkrementrückstände von Mabuya vaillanti von St. Cruz ergab fol-

gende Bestandteile: Acridiidae, Formicidae, Coleoptera, viele junge Blätter und Knospen möglicher-

weise von Cistaceae.

Die Nahrungsrückstände von Mabuya delalandii und Mabuya stangeri setzten sich wie folgt zu-

sammen:

Heteroptera, Coreidae-Lygaeidae, Coleoptera, Tenebrionidae.

Bei den Tarentolas wurde getrennt nach den Formen von $. Thiago sowie nach jenen von Ilheu

St. Maria ausgewertet:

Praia: Aranea, Heteroptera, Cydnidae, Coreidae-Lygaeidae, Hymenoptera, Formicidae, Coleopte-

ra, Tenebrionidae.

St. Maria: Acridiidae, Heteroptera, Cydnidae, Pentatomidae, Coreidae-Lygaeidae, Formicidae,

Diptera, Tenebrionidae.

Freßfeinde

Sowohl aufS. Thiago wie auf Fogo konnten relativ viele Eisvögel (Halcyon leucocephala acteon) be-

obachtet werden. BANNERMAN & BANNERMAN (1968: 17) berichteten dazu: ‚,...the kingfisher which

tamely sits on the branches of the castor-oil plants and thence darts on grasshoppers and lizards.“

DOLEOCO

Fogo ist von rundlicher Gestalt und typischem Schichtvulkan-Habitus. Bis 2829 m ragt der Pico de

Fogo aus dem Meer, die Flächenausdehnung der Insel beträgt ca. 500 qkm, Hauptstadt ist S. Felipe.

Der Pico, die große Caldeira, weite Lavafelder und tiefeingeschnittene Ribeiras kennzeichnen die In-

sel.

Fajasinha

S. Lourengo ze

ee 2469 1790

FEE Dr, N IN ß

ı IIDN

1 N

CEDE SFELIPE

Artenspektrum

Obwohl Mabuya vaillanti aus der Aufsammlung des British Museum für Fogo (Igreja) bekannt

war, gelang es mir nicht, diese Art nachzuweisen. Der geographisch höchste Nachweis eines Skinkes

auf den Kapverden gelang auf dem Pico bei ca. 2800 m. Leider wurde das Tier nur gesichtet, so daß

seine Feldbestimmung als Mabuya cf. fogoensis unbestätigt bleibt. Mabuya stangeri kommt im offe-

nen Gelände zusammen mit Mabuya fogoensis vor. Hemidactylus brooki angulatus wurde vornehm-

lich an Gemäuern in S. Felipe gefunden. Tarentola rudis rudis dagegen hauptsächlich im offenen Ge-

lände, unter Wasserdurchlässen oder an freistehenden alten Häusern.

Fundorte

1 Pico de Fogo : Mabuya spec.

DIES Relipe : Mabuya, Tarentola, Hemidactylus

3 3km N-Mira Mira : Mabuya

4 S. Lourengo : Tarentola

5 Igreja : M. vaıllanti (BMNH)

Zur Biologie - Ökologie

Geländenotizen, Körpertemperaturen:

Datum Uhrzeit LT LF BT Taxon KT Bemerkungen

TC % _@ EC

dot. 7.00 #29. „., Sonne: > S. Felipe

21.45 24 24 Hemidact. 24 BT = Wand/Mauertemperatur

22.30 23 23 Hemidact. 26

23.45 23 23 Hemidact. 26 zusammen unter Brücke

Tarentola 26

24.00 225 >» Tarentola 23

2235 19 20 Tarentola 21 S. Lourengo

20 22 Tarentola 22

Die Kotanalysen ergaben folgende Nahrungskomponenten:

Tarentola: Aphidina, Coleoptera, Tenebrionidae, Scarabaeidae, Tettigoniidae, Mantidae, Formici-

dae.

Hemidactylus: Isoptera, Formicidae, Diptera, Coleoptera.

5.6 BRAVA

Brava ist die südlichste Insel des Archipels und gehört noch zur Sotavento-Gruppe. Die Insel ist von

rundlicher Gestalt bei einer Flächenausdehnung von ca. 65 qkm. Mit dem Monte Mato erreicht Brava

976 müber NN. Quellen, Wasserläufe, massives Kalkgestein und eine üppigere Vegetation geben der

Insel ihr eigenes charakteristisches Aussehen.

BRAVA

Ilheu da Areia oO

[>)

VAN SINTRA

Artenspektrum

Bislang waren von Brava (s. ScHLEıcH 1982: 246) Mabuya delalandi, Mabuya stangeri, Tarentola

delalandii (rudis) sowie Hemidactylus bouvieri bouvieri und Hemidactylus brooki angulatus bekannt.

Bei einem leider nur kurzen Aufenthalt konnten lediglich 8 Mabuya delalandii gefangen werden.

Fundorte

1 Forainhas bei Mato Grosso; vor dem Friedhof unter Lesesteinen: Mabuya delalandıii (ZSM 370/78).

Abb. 17: Tarrafal-S. Thiago; Biotop von Mabnya vaillantı, M. delalandii und M. stangeri. - Abb. 18: Brava;

Biotop von Mabuya delalandı. — Abb. 19: S. Vicente; Biotop von Tarentola caboverdiana substituta im Inselin-

nern.

Zur Biologie - Ökologie

Geländenotizen:

Am 29.7.1977 wurden auf einer Ackerbaufläche von ca. 250-300 qm unter Lesesteinen 8 Skinke gefangen.

5.7 ILHEUS DO RHOMBO

Die nahe vor Brava liegenden Ilheus do Rhombo sind unbewohnte Felseilande, deren drei größte In-

seln Luz Carneiro, Sapado Grande und Cima sind. Mit96 m erreichen sie auf Ilheu Grande die höchste

Erhebung.

Leider konnten die Inseln von mir selbst nie besucht werden. SCHLEICH (1982: 246) gibt nach MEr-

TENS und AnGer Mabuya delalandıi, Mabuya vaillanti, Tarentola delalandii = (T. rudis) und Hemi-

dactylus brooki angulatus an. JOGER (1984: 101) führt auch ‚‚Tarentola borneensis protogigas“

(= T. rudis protogigas) an.

2 o Q

ILHEU GRANDE o

{e] ß

9° ILHEU DE CIMA

Be ILHEU LUZ

CARNEIRO

5.3 SANTO ANTÄO

Santo Antäo, die nordwestlichste der kapverdischen Inseln, zählt neben $. Vicente, den Islas Deser-

tas, S. Nicolau, Sal und Boa Vista zur Gruppe der Barlaventos.

Die Insel besitzt annähernd rechteckige Grundform bei einer Fläche von 730 qkm. Santo Antäo ist

auch in extremsten Trockenperioden neben $. Thiago und Brava eine der Inseln mit ständig offenen

Fließwässern. St. Antäo ist die gebirgigste, zerklüftetste und landschaftlich beeindruckendste aller In-

seln. Die Südostseite ist bis zu dem längs über die Insel ziehenden Gebirgskamm trocken, steinig und

von wüstenartigem Gepräge. Die NW -Seite dagegen zeichnet sich durch üppig grüne Landschaften mit

steilen Felswänden, Tälern und überraschender Fruchtbarkeit aus.

Artenspektrum

Erst der zweite Besuch auf St. Antäo war bezüglich des Reptilienfanges erfolgreich. Wir suchten auf

der trockenen SE-Seite der Insel erfolgreich, wogegen auf der NW-Seite im Bereich von Ribeira-

Grande weder Geekos noch Skinke zu finden waren. Einheimische meinten, daß es hier keine Echsen,

insbesondere Geckos mehr gäbe, da sie von den erst vor kurzem durch Saatkartoffeln aus Kontinental-

afrika eingeschleppten Skolopendern ‚‚aufgefressen‘‘ worden wären. Leider glückte auch hier kein

Nachweis von Hemidactylus bouvieri bouvieri (s. SCHLEICH, 1982: 246), dagegen von Mabuya fogoen-

sis antaoensis und Tarentola caboverdiana caboverdiana ca. 4 bis 10 km nördlich der Straße von Porto

Novo nach Chä da Monte.

m LA

25020" S a A N f A 0) N FAR ae

e —uN

171° — —

ÄS

' N

\ Rib.da Cruz‘,

Ilheu Lombo

„Janela

z de Boi

1586

PORTO NOVO

—— 17°

25°

Tarrafal do z

Monte Trigo /F

Fundorte

1 Gebiet ca. 4-10 km nördl. der Straße von Porto Novo nach Chä da Monte: Mabnya fogoensis antaoensis, Ta-

rentola caboverdiana caboverdiana.

Zur Biologie — Ökologie

Geländenotizen:

Am 24.9. 81 konnten wir nachmittags im beschriebenen Areal bei einer LT von ca. 23—40°C aktive Mabuyen und

Tarentolas unter Lesesteinen fangen. Auffallend war dabei ein großer Anteil diesjähriger Jungtiere, der auch zu die-

ser Zeit auf allen anderen Inseln bemerkt wurde.

5.9 SAO VICENTE

Sao Vicente liegt im NW des Archipels zwischen St. Antäo und den Islas Desertas. Mit der Ilha do Sal

ist Sao Vicente als eine der wüstenhaftesten des Archipels zu bezeichnen. Die Insel ist von rundlich-

rautenförmiger Gestalt und hat eine Gesamtausdehnung von ca. 220 qkm bei einer maximalen Höhe

SIeRY ERBEN ITNE

von 774 m mit dem Monte Verde. Große Gebirgszüge, weite öde Wüstenstrecken und junge, von vul-

kanischer Aktivität gezeichnete weite Lavagebiete sowie Dünengürtel kennzeichnen die Insel.

Artenspektrum

In einer früheren Arbeit (SCHLEICH 1982: 246) erwähnte ich nach verschiedenen Autoren Mabuya fo-

goensis und Mabuya stangeri, die ich vorbehaltlich noch als Mabuya spp. indet. ansprach und nach der

systematischen Überarbeitung als zu Mabuya stangeri stangeri gehörig betrachte.

An Geckos wurde ‚‚Tarentola delalandu“ als Tarentola caboverdiana substituta (s. Pkt. 4.2.3.4)

berichtet, Hemidactylus brooki angulatus nachgewiesen und Hemidactylus bouvieri bouvieri durch

zwei Exemplare aus dem British Museum (GRUBER & SCHLEICH 1982) belegt.

Fundorte

1 Küstenabschnitt südl. von Madeiral:

Mabuya stangeri stangeri

2 ca. 3 km westl. von Madeiral:

Tarentola caboverdiana substituta

3 Häuserfront der Hafenstadt Mindelho:

Hemidactylus brooki angulatus

Anmerkung: Weite Bereiche der Insel wurden neben den genannten Fundpunkten abgesucht, jedoch ohne Er-

folg.

Zur Biologie - Ökologie

Geländenotizen:

Der Biotop (Fundpunkt ‚,2‘“) der am 4.8.77 gefangenen Tarentolas bestand in einer im Umkreis weniger qkm ve-

getationslosen Sand-Stein-Wüste, wo unter einzelnen, wenigen Lesesteinen meist 2 Tarentolas zu finden waren.

Skinke wurden hier keine gesehen. Etwa 2 km westlich von Punkt 2 konnten zwei vereinzelte Jungtiere von Ma-

buya stangeri stangeri gefangen werden. Im gesamten Bereich zwischen Mindelho und $S. Pedro wurde intensivst

5 8 ger 5 5 5

gesucht, jedoch ohne einen einzigen Nachweis erbringen zu können.

Für Tarentola caboverdiana substituta erbrachte die Analyse der Kotproben folgende Zusammen-

setzung:

Zahlreiche Cydnidae, viele Coreidae-Lygaeidae und Coleoptera sowie Pentatomidae, Tettigonii-

dae, Scarabaeidae, eine Spinne (Aranea) sowie eine Lepidopteren-Larve und drei Geckoschwanz-

enden.

510 STALUZIA

In SCHLEICH & WUTTkE (1983) wurden die Islas Desertas mit St. Luzia, Branco und Razo gesondert

abgehandelt, so daß eine detaillierte Wiederholung hier entbehrt werden kann. Weitere Angaben zu

den Islas Desertas finden sich bei SCHLEICH (1977, 1980 und 1982) und GRUBER & SCHLEICH (1982).

ShelaslnZz I» A

Artenspektrum

Nach einem längeren Aufenthalt und intensiven Begehungen und Suchaktionen konnte lediglich

Mabuya stangeri stangeri und Tarentola caboverdiana raziana nachgewiesen werden.

Fundorte

Gesammelt wurde auf der ganzen Insel, vornehmlich im Bereich des Mittelteiles.

Zur Biologie - Ökologie

Geländenotizen, Körpertemperaturen:

Datum Uhrzeit LT LIE BT Taxon KT Bemerkungen

@ % C SC

2,281 1 225 31,0 Tarentola 29,0 windig, ca. 300 m NN auf windexponiertem

ungeschütztem Felsgrat

T°C unter Stein: 21,5°C, auf Stein: 26,5°C

RS 29,0 Tarentola 25,5

11.40 225 29,0 Tarentola 27 T°C unter Stein: 1°C

DRS 29 Mabuya 28

22,5 29 Mabuya 26,5

13.00 22 32,5 Mabuya 30 adultes Tier

.20 22 33,5 Mabuya 30 adultes Tier, sehr windig, sonnig

14.25 22 33 Tarentola 28,5 26°C unter Stein

Tarentola 29,5 adultes Tier

14.45 22,5 33 Tarentola 31,5 °2 mit 1 Eı (Schatten - T°C)

26,5 33,5 Tarentola - (Sonne — T°C)

14.55 32 al Q mit 1Eı

RS d, T°C unter Steinplatte: 28°C

Hemipenes beim Messen ausgestülpt

15.00 2355 5255 Hl Q mit Ei

15.10 25 35 Mabuya 28,5 Adulti, unter Steinplatte eingegraben

(BT unter Platte: 26°C)

15.30 24 32 Mabuya 30,5 aktiv, semiadult

21.20 21 20 Tarentola 20

21.45 19 18 Tarentola 19 19°C unter Steinplatte

32. 13.30 Tarentola 27 25°C in ca. 20 cm ‚„‚Bautiefe“

24.00 19 82

4.2 6.30 17 Sonnenaufgang

Do 7.00 17 82

9.00 24 65

SIEBRLNNEO,

Branco, die mittlere der3 Islas Desertas, ist an einigen Sandbuchten von der SW-Seite aus anlandbar.

Detaillierte Informationen zur Inselbeschreibung finden sich bei SCHLEICH & WUTTKE (1983) sowie bei

SCHLEICH (1979, 1980).

Artenspektrum

Mehrere längere Aufenthalte auf Branco bestätigten lediglich 2 Echsenarten: Tarentola gigas bran-

coensis und Mabuya stangeri stangeri. Ein frisches Gelege einer Seeschildkröte konnte im Sommer

1977 entdeckt werden. Nach SCHLEICH (1982) gilt Macroscincus als ausgestorben.

Fundorte

Gesammelt wurde auf der ganzen Insel, jedoch konzentrieren sich die Riesengeckos auf den mittleren Bereich ım

BRANCO

SW der Insel. Detaillierte Angaben zur Struktur, Fauna und Flora der Insel s. SCHLEICH (1980) u. SCHLEICH &

WUTTKE (1983).

Zur Biologie - Ökologie

Geländenotizen, Körpertemperaturen:

Datum Uhrzeit LT EE BT Taxon KT Bemerkungen

ce % I® @

6.1.8, 212.20 22 86 32 Tarentola 24,5 ausgegraben, ca. 80 cm tief in Sandhöhle

(T°C: 25)

14.45 23 - 23 Mabuya 29 ca.20m NN, Steilhang N-Seite

17.05 22 - 25 Mabuya 28 (T°C unter Steinplatte: 28)

in ca. 40 cm Bautiefe: 86 % LF; 22°C

5.12 RAZO

Eine detaillierte Beschreibung zu Razo findet sich bei SCHLEICH & WUTTKE (1983).

24°37'

6 RAZO

Abb. 20: Blick auf Branco; deutlich sind die Sanddünen - das Hauptverbreitungsgebiet von Tarentola gigas bran-

coensis - zu erkennen. Im Hintergrund rechts: St. Luzia. Im Vordergrund Razo mit dem Biotop von Hemidactylus

bouvieri razoensis, Tarentola gigas gigas und Mabuya stangeri stangeri.

RN SIEERIIRAN x x 58 ben I

Abb. 21: Bereich im Innern von St. Luzia, der Biotop von Tarentola caboverdiana raziana und Mabuya stangeri

stangert.

Artenspektrum

Razo wurde von meinen Begleitern und mir in den letzten Jahren gründlich herpetologisch durch-

forscht, wobei folgende Taxa studiert wurden:

Tarentola gigas gigas

Tarentola caboverdiana razıana

Mabuya stangeri stangeri

Hemidactylus bouvieri razoensis

Fundorte

Gesammelt wurde auf der ganzen Insel, jedoch scheinen außer Mabuya die übrigen Arten auf bestimmte Inselab-

schnitte gebunden zu sein oder zumindest sich dort zu konzentrieren (s. SCHLEICH & WUTTKE 1983).

Zur Biologie - Ökologie

Geländenotizen, Körpertemperaturen:

Datum Uhrzeit LT BT Taxon KT Bemerkungen

@ °C ®

10,2 8 22) 2765 Mabuya LS) aktıv

19.00 il Tarentola 21

Zur Biologie von Hemidactylus bouvieri razoensis Gruber & Schleich wurde 1982 gesondert berich-

tet; eine detaillierte Sammeldarstellung zu Tarentola gigas ist in Vorbereitung. Weitere Angaben sind

aus SCHLEICH & WUTTKE (1983) zu entnehmen.

Die Kotanalyse erbrachte folgendes Nahrungsspektrum für

Mabuya:

Acridiidae, Tettigoniidae, Cydnidae, Pentatomidae, Coreidae-Lygaeidae, Formicidae, Coleoptera,

Tenebrionidae

Tarentola caboverdiana razıana:

Opiliones, Acridiidae, Cercopidae, Heteroptera, Pentatomidae, Diptera, Coleoptera, Tenebrionidae,

pflanzliche Reste

Tarentola gigas gigas:

Opiliones, Tettigoniidae, Mantidae, Formicidae, Coleoptera, Tenebrionidae, Carabidae.

5.13 35. NICOLAU

S. Nicolau, zu den Barlaventos gehörend, hat eine Flächenerstreckung von 350 qkm bei einer maxi-

malen Höhe von 1304 m. Ähnlich St. Antäo ist auch hier eine fast lineare landschaftsklimatologische

Zweigliederung der Insel in den sehr trockenen Südteil und den vegetationsreicheren Nordteil zu er-

kennen, wo in letzterem auch vorwiegend die Reptilien gefangen werden konnten.

Ss. NRErOTEA U

436

‚Praia Branca

„$Fiqueirae

deCoxe +619

Fontainhas

Tarrafal ,.- 790

...- " Prequiga'

WiS

R Castilhiano

2 km

Artenspektrum

Mabuya fogoensis und Tarentola caboverdiana nicolauensis scheinen hier als einzige Arten neben

Bufo regularis vorzukommen.

Fundorte

1 Ribeira Brava: Bufo regularıs

2 Etwa halbe Strecke zwischen Preguiga und Rib. Brava: Tarentola, Mabuya

Zur Biologie - Ökologie

Geländenotizen:

1977 konnten bei den offenen Wasserbecken der Quellfassung von Ribeira Brava ca. 100 Kröten gezählt werden,

wovon 1981 nach Verbauung der Quelle nur noch wenige einzelne Tiere vorhanden waren.

Die Kotanalyse erbrachte folgendes Nahrungsspektrum für

Mabuya: Cercopidae, Heteroptera, Coreidae-Lygaeidae, Formicidae, Coleoptera, Carabidae,

pflanzl. Reste

Tarentola: Aranea, Cydnidae, Heteroptera, Coreidae-Lygaeidae, Formicidae, Coleoptera, Curcu-

lionidae.

Die Tiere wurden am 20.7.81 gefangen.

6. Biotop- und Artenschutz

BANNERMAN & BANNERMAN (1968) wiesen wohl als erste auf einen dringend notwendigen Arten-

schutz der kapverdischen Fauna hin. Ihr Bericht (op. cit., S. 5) über die ‚laws for the protection of na-

ture“ basiert verständlicherweise noch auf den Angaben aus der portugiesischen Kolonialzeit mit Ge-

setzen von 1955. Aus BANNERMAN & BANNERMAN (Op. cit.) sei zitiert:

„Here is a list of the birds that cannot be killed in the Archipelago, it being true that the only other

animal whose life is safeguarded by law is the skink Macroscincus coctei: most of the species of swifts,

larks, flamingos (Phoenicopterus ruber) and the cattle egret (Ardeola ıbıs).

The following species can only be hunted during the game season from 1st December to 31st July:

rock-pigeon Columba livia, the quail Coturnix coturnix, and the guinea-fow| Numida galeata.“

Von weiteren Bedrohungen bzw. Ausrottungen berichten BANNERMAN & BANNERMAN (Op. cit.,

S. 42). |

Jedoch wiesen bereits viele Autoren schon kurz nach der Erstbeschreibung von Macroscincus coctei

auf dessen Schutzwürdigkeit bzw. bevorstehende Ausrottung (s. SCHLEICH 1979) hin.

In gleicher Weise sprach ich bereits von einer dringlichen Notwendigkeit eines Schildkrötenschutz-

projektes.

SCHLEICH & WUTTKE (1983: 41) müssen leider von intensivsten Nachstellungen der Avifauna auf den

kleinen unbewohnten Inseln Branco und Razo berichten, denen sicher auch die Ausrottung von Ma-

croscincus coctei anzulasten ist.

Als dringlich erachte ich in Zusammenarbeit mit der kapverdischen Regierung die Erstellung von

Arten- bzw. Biotopschutzprogrammen für folgende Reptilien:

Cheloniidae spp.

Mabuya vaillantı

Tarentola gigas ssp.

Hemidactylus bouvieri razoensıs

Ein umfassender zusätzlicher Biotopschutz, insbesondere der unbewohnten Inseln Branco und

Razo (möglicherweise auch der Rhombos-Inseln, jedoch liegen mir hier keine persönlichen Aufzeich-

nungen vor), würde nicht nur deren Herpetofauna, sondern auch die Avifauna schützen! Entscheidend

scheint mir dabei, die beiden Inseln überhaupt vor unkontrolliertem Betreten zu schützen, um nicht

Mäuse und Ratten einzuschleppen. Auf St. Luzia (s. SCHLEICH & WUTTkE 1983) können die Mäuse be-

reits als richtige Plage bezeichnet werden, die sicher auch für den niedrigen Reptilienbesatz auf dieser

unbewohnten Insel verantwortlich sind.

Ein detailliertes Aufklärungsprogramm über sinnvolle Nutzung und Schutz sowohl der Herpeto-

fauna als auch der Avifauna scheint mir unabdingbar.

Nachtrag: Nach Fertigstellung des Manuskripts erfolgte von IUCN/WWF erfreulicherweise die

Herausgabe von Ersttagsbriefen, -karten mit englischem und deutschem Begleittext zu den schutzbe-

dürftigsten Arten kapverdischer Reptilien. Hierzu sind Tarentola g. gigas, T. g. brancoensis, Hemı-

dactylus bouvieri razoensis und Mabuya vaillanti abgebildet auf 4 verschiedenen Briefmarken der Re-

publik Cabo Verde unter dem Titel ‚‚Protecäo de Especies em Via de Extingäo“ erschienen.

7. Schrifttum

ANGEL, F. 1935: Lezards des Iles du Cap Vert, rapportes par M. le Professeur Chevalier. Description de deux espe-

ces nouvelles. — Bull. Mus. Hist. nat. Paris (2)7: 165-169

— — 1937: Sur la faune herpetologique de !’Archipel du Cap-Vert. — XIIe Congres International Zool., Sec-

tion IX: 1693-1700, Lisbonne 1935

BAEZ, M. & SANCHEZ-PINTO, L. 1983: Islas de Fuego y Agua. - Edit. Regional Canaria, Las Palmas-Gran Canaria,

184 S.

BANNERMAN, D. A. & BANNERMAN, W. M. 1968: Birds of the Atlantic Islands. 4 Vol. IV. History of the Birds of

the Cape Verde Islands. — Oliver & Boyd, 459 S.

BERTIN, C. 1946: Le peuplement des Iles Atlantides en vertebres heterothermes. - Mem. Soc. Biogeogr. 7: 87-107

BISCHOFF, W. 1985: Die Herpetofauna der Kanarischen Inseln I. — Herpetofauna 7 (34): 11-22

BOCAGE, J. V. B. du 1873a: Notice sur !’habitat et les characteres du Macroscincus coctei (Euprepes coctei Dum. et

Bibr.). — J. Sci. math. phys. nat. 16: 12 S.; ebenfalls nachgedruckt in: J. Zool. 3: 1-15

— — 1873b: Note sur l’habitat de l’Euprepes coctei Dum. et Bibr. - Proc. Zool. Soc.: 703-704

— — 1875: Sur deux reptiles nouveaux de !’Archipel du Cap Vert. — J. Sci. math. phys. nat. 5: 108-112

— — 1896: Reptis des algumas possessöes portuguesas d’Africa que existem no museu de Lisboa. — J. Sci. math.

phys. nat. 14 (2): 65-73

BOULENGER, G. A. 1887: Catalogue of the lizards in the British Museum (Natural History), 3: 1-575

— — 1906: Report on the reptiles collected by the late L. Fea in West Africa. - Ann. Mus. civ. storia nat. Genova

3 (2): 196-213

CHABANAUD, P. 1924: Reptiles recueillis par M. Th. Monod en Mauritanie et aux Iles du Cap-Vert. — Bull. Mus.

Hist. Nat. 1: 54-56

DEKEYSER, P. L. & VILLIERS, A. 1951: Mission J. Cadenat aux Iles du Cap Vert. — Bull. Inst. frangais d’Afrique

noire 13(4): 1152-1158

Dumekrir, A.M.G. & BIBRON, G. 1839: Erpetologie generale ou histoire naturelle complete des reptiles. — 854 S.

GERVAıS, P. 1874: Plate of the head and skull of Macroscincus coctei the reprint of Bocage’s 1873a article. - J. Zool.

Bla

GIRARD, CH. 1858: United States Exploring Expedition. Herpetology. — Lippincott & Co., Philadelphia

GREER, A. E. (1976): On the evolution of the giant Cape Verde scincid lizard Macroscincus coctei. — J. nat. Hist. 10:

691-712

GRUBER, H. J. & SCHLEICH, H.-H. 1982: Hemidactylus bouvieri razoensis nov. ssp. von den Kapverdischen In-

seln. — Spixiana 5 (3): 303-310

HONEGGER, R. 1975: Red Data Book 3, IUCN 1975 9(1) F. Code: 24 10211

JOGER, U. 1984: Die Radiation der Gattung Tarentola in Makaronesien. - Cour. Forsch. Inst. Senckenberg 71:

91-111

Kraus, O. 1970: Internationale Regeln für die Zoologische Nomenklatur. - W. Kramer-Verlag; Frankfurt; 92 S.

LOBIN, W. & GROH, K. 1979: Die Kapverdischen Inseln - ein Reisebericht. — Natur u. Museum 109 (12): 394405

— — & — — 1980: Die Kapverdischen Inseln -— ein Reisebericht II. - Natur u. Museum 110 (10): 289-304

LOVERIDGE, A. 1947: Revision of the African lizards of the family Gekkonidae. - Bull. Mus. comp. Zool. 98: 1469

METEROLOGICAL OFFICE 1975: Tables of Temperature, Relative Humidity and Precipitation for the World.

Part. IV. Africa, the Atlantic Ocean south of 35 N and the Indian Ocean. Her Majesty’s Stationery Office,

London 208 S.

MERTENS, R. 1934: Die Inselreptilien, ihre Ausbreitung, Variation und Artbildung. —- Zoologica 84: 209 S.

— — 1955: Die Eidechsen der Kapverden. - Soc. Sci. fenn. Comment. biol. 15(5): 1-16

Nie, N. H. & Hutr, C. H. 1980: SPSS 8. Statistik-Programm-System für die Sozialwissenschaften. —3. Auflage,

300 S., Fischer-Verlag; Stuttgart

ORLANDI, $. 1894: Note anatomiche sul Macroscincus coctei (Bearb. d. Boc.). — Atti. Soc. Lingustica 5: 175-204

O’SHAUGNESSY, A. W. E. 1874: Descriptions of new species of Scincidae in the collection of the British Museum. —

Ann. Mag. nat. Hist. 13 (4): 298-301

PERACCA, M. G. 1891a: Sulla oviparitä del Macroscincus coctaei Dum. e Bibr. - Boll. Mus. Lab. Zool. Anat. comp.

R. Univ. Genova 6 (105): 1 S.

— — 1891b: Osservazioni sul Macroscincus coctaei D. B.- Boll. Mus. Lab. Zoo. Anat. comp. R. Univ. Genova

6 (107): 1-5

SCHLEICH, H.-H. 1979a: Der Kapverdische Riesenskink, Macroscincus coctei, eine ausgestorbene Echse? - Natur

und Museum 109 (5): 133-138

— — 1979b: Sea Turtle Protection needed at the Cape Verde Islands. - Marine Turtle Newsletter 12: 12

— — 1979c: Kapverden, Vergessene Inseln im Atlantik. - (Farbabbildungen von Echsen der Kapverden); Film

und Video 12: 56-65

— — 1980: Der kapverdische Riesengecko, Tarentola delalandı gigas (Bocage, 1896). — Spixiana 3 (2): 147-155

— — 1982a: Vorläufige Mitteilung zur Herpetofauna der Kapverden. - Cour. Forsch.-Inst. Senckenberg 52:

215-248

— — 1982b: Letzte Nachforschungen zum Kapverdischen Riesenskink Macroscincus coctei (D. & B.). — Sala-

mandra 18 (1/2): 78-85

— — 1984: Die Geckos der Gattung Tarentola der Kapverden. Reptilia: Sauria: Gekkonidae- Tarentola. - Cour.

Forsch.-Inst. Senckenberg, 68: 95-106

— — 1986: Geckos und Skinke (engl. Desert Island Lizards). - (Begleittext zu Ersttagsbriefen, Marken u. Karten

etc. von Cabo Verde) 6 S., 10 Farbabb. (Hrsg.: IUCN/WWF)

SCHLEICH; H.-H. & WUTTKE, M. 1983: Die kapverdischen Eilande, St. Luzia, Branco und Razo - ein Reisebericht.

Natur und Museum 113 (2): 3344

TROSCHEL, F. H. 1875: Über die Rieseneidechse der Inseln des Grünen Vorgebirges. - Arch. Naturgesch. 41 (1):

111-121

VAILLANT, L. 1882: Sur les Macroscincus cocteı, D. B., recemment arrıves a la menagerie du Museum d’Histoire na-

turelle. - C. r. hebd. Seanc. Acad. Sci. 94 (12): 811-812

WERMUTH, H. 1965: Gekkonidae, Pygopodidae, Xantusiidae. Das Tierreich 80: I-XXII, 1-246; (R. MERTENS &

W. HENNIG, Hrsg.); DE GRUYTER, Berlin

Manuskriptannahme: 1. März 1985

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Bisher erschienene Supplementbände der SPıxıana:

Supplementband 1: GUSTAV PETERS, 1978

Vergleichende Untersuchung zur Lautgebung einiger Feliden

(Mammalia, Felidae).

206 Seiten und 80 Seiten mit 324 Abbildungen und 20 Tabellen.

Supplementband 2: HERMANN ELLENBERG, 1978

Zur Populationsökologie des Rehes (Capreolus capreolus L., Cervidae)

in Mitteleuropa.

211 Seiten mit 47 Abbildungen und 42 + 6 Tabellen.

Supplementband 3: JENSLEHMANN, 1979

Chironomidae (Diptera) aus Fließgewässern Zentralafrikas.

(Systematik, Ökologie, Verbreitung und Produktionsbiologie).

Teil I: Kivu-Gebiet, Ostzaire.

144 Seiten mit 252 Abbildungen und 11 Tabellen.

Supplementband 4: KLAUS HORSTMANN, 1980

Revision der europäischen Tersilochinae Il

(Hymenoptera, Ichneumonidae).

76 Seiten mit 150 Abbildungen und 2 Tabellen.

G. VAN ROSSEM, 1980

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oPIAIMN

Zeitschrift für Zoologie

A Revision of the Holarctic

Species of Orthocladius

(Euorthocladius)

(Diptera: Chironomidae)

By Annelle R. Soponis

Herausgegeben

von

E. J. Fittkau

Zoologische Staatssammlung München

SPIXIANA Supplement 13 München, 31. Januar 1990 ISSN 0177—7424

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A Revision of the Holarctic

Species of Orthocladius

(Euorthocladius)

(Diptera: Chironomidae)

By Annelle R. Soponis

SPIXIANA Supplement 13 München, 31. Januar 1990 ISSN 0177— 7424

Gesamtherstellung: Gebr. Geiselberger, Altötting

A Revision of the Holarctic Species of Orthocladius

(Euorthocladius)

(Diptera: Chironomidae)

By Annelle R. Soponis

Contents

A N EEE N 3

EoELOdUCLONN ER. et ren a LER Bl ee a er N er Se 4

Er-knowledementst te. as ne 3 LEN Re ee ee RR 5

Berertalsand4Methodset..n..1. 0 et ee ER 6

fine Subgenus Orthocladıus (Euorthocladinus) Thienemann ... ........uceoeeoeeueeauen. 6

PB 20.nosisgand3Deseriptions:. ne ee EEE SEE HE 7

Keys to Holarctic Species of O. (Enorthocladius) and Subgenera of Orthocladins:

ErdultsNlalesgBupaeslarvae. 15.000000. N ee RE 9

abiskoensis Thienemann and Krüger rl

Erreilist (Ro back ERTL E EN N L TrER IR. ee ee Das uleredkt 16

SETS ae al A TE ENTER HE BEhe r5 2 Er REINER NEED 17

220908 naar ct ee ee oe NR OR EEE 19

SD na hi EN large yattekc Kits 20

Bzalısleundbeck)i zur. sen: 2u::ue eg Re re Eu ng al

3.2008 (OSTEN 22

Erzipes1@oetshebuers.ne.rn. Sr let 23

Br co/aATelen A N N ae eh 26

BarloynmsKuelternse ra u a re N la ar 5 30

EERSSEH ZEIT EST REN AEON IE, I ERTENN, 2 a e 34

5230929 (Molke) Re 3 Se 36

EZSpensus okumara)a a. det. ae 27 herckcr ea En Arsen era adag Kl" anne 38

Rlochaerus3Banstonsn Tec Se ee RE 40

BurenemannuKuelter.e 2.2.5 0 ce et en 40

Rieierencesppg ns RT ee a BE NE 44

MlaStraTon St PN u 49

Abstract:

The classification of the Holarctic species of the subgenus Orthocladius (Enorthocladius) is revised. Keys to spe-

cies for adult males, pupae, and larvae are given. Keys to subgenera of Orthocladius are included for adult males,

pupae, and larvae. Fifteen names of Orthocladius (Euorthocladius) are recognized as valıd. Redescriptions or notes

on previously-known species and descriptions of three new species are presented. The new species are: ashei, coff-

mani, and roussellae. Type Material was examined for the following species: abiskoensis, anteılis, calvus, difficalis,

kanii, luteipes, saxosus, suspensus, telochaetus, and thienemanni. Lectotypes are designated for abiskoensis Thiene-

mann and Krüger, rivulorum Kieffer, and for the type of the subgenus, thienemanni Kieffer. The type of rivicola

Kieffer could not be located.

Introduction

Adults of Orthocladius (Enorthocladins) are small to medium-sized, yellow, green, brown, or black

chironomids. In the temperate zone they emerge primarily in spring, fall and winter, although some

are taken at high altıtudes and latitudes in summer.

Live larvae have brown head capsules and green, brown, or yellow bodies, and live in gelatinous

tubes in cold springs or fast-flowing waters of inlets, creeks, streams, and rivers. These gelatinous

tubes are inıtially clear, and can be covered with sand or algae, appearing brown or green depending

on the habitat. In the rivicola-group, larvae live in ellipsoid tubes fastened along their margins to sto-

nes and rocks. In some species, e. g. saxosus and thienemanni, larvae are gregarious. In the rzvulorum-

group, larvae live in suspended tubes, attached by one end to stones and rocks. Larvae of certain spe-

cies have been collected from moss (ashei, abiskoensis, rivolorum), reeds (thienemanni), Ranunculus

(ashei), and algal mats (roussellae). Larvae collected from mud have probably occured there acciden-

tally.

Species of Orthocladius (Euorthocladius) can be sympatric ands synchronous, and it is not unusual

to collect exuviae of different species, e. g. rivicola with either ashei, Inteipes, or thienemanni, in the

same sample.

Based primarily on pupal characters, the species can be placed in one of two groups: the rivicola-

group (ashei, calvus, difficılis, kanıı, Iuteipes, rivicola, thienemanni); and the rivulorum-group (antei-

lis, coffmani, rivulorum, roussellae, suspensus). The species abiskoensis and saxosus share characteris-

tics of both groups.

Characters separating species can be very subtle, and several species remain undescribed. Intraspe-

cific variation is apparently large, and needs to be studied further. Three species found in Japan are

almost identical morphologically to certain species from Europe and/or North America, but because

of important morphological differences in the pupa or adult male, the incipient species in the sub-

genus, and the lack of series of reared specimens, these species are not synonymized here: suspensus

and rivulorum, kanii and Iuteipes, and saxosus and telochaetus.

Although names of Orthocladius (Euorthocladius) have appeared often in the literature, actual mu-

seum specimens are few, given the number of species. Whether this reflects low abundances or

inadequate collecting of natural populations remains unanswered.

The subgenus Orthocladius (Enorthocladius) is recorded only from the Holarctic Region. Because

the genus Orthocladius is distributed worldwide (Soponis 1977), it would not be surprising to find

species of Orthocladius (Euorthocladins) in other regions of the world in cold or fast-flowing waters.

Species of O. (Enorthocladius) are most easily identified in the pupal stage. Specific identification

of adult males is particularly difficult because of intraspecific variation and morphological similarity

of congeners. Specific identifications of larvae can be made with more confidence than of adult males,

but for accurate specific identification larvae should be associated with the pupal stage.

Prior to the present study, sıx species of Orthocladius (Enorthocladius) were recorded from the

Palearctic region (Sasa & Yamamoto 1977, Fittkau & Reiss 1978) and two species from the Nearctic

region (Sublette & Sublette 1965). Presently ten species are reported from the Nearctic region and nine

species from the Palearctic region, with six species occuring in both regions. There has been uncer-

tainty about whether or not to place frigidus and abiskoensis in the subgenus O. (Euorthocladins). Or-

thocladius frigidus belongs to Orthocladins (Orthocladins). (Soponis, 1987), and Orthocladins abisko-

ensis has been placed in O. (Euorthocladins) by Säwedal (1978), which is accepted here.

The primary objective of this study was to revise the classification of the Holarctic species of Ortho-

cladius (Euorthocladius), thereby gaining a better understanding of the genus Orthocladius. Adult

males of Orthocladius are difficult to place to subgenus, but pupae and larvae can be assigned with re-

lative ease. Adult males and larvae of Orthocladius are difficult and sometimes impossible to place to

species, but pupae can be determined to species with relative ease.

The following keys should be considered tentative; the most accurate specific identifications of Or-

thocladius (Euorthocladins) will be those based on specimens associated with the pupal stage. Caution

must be used in identifying Palearctic material since the species-rich Orthocladius (Orthocladius) of

the Palearctic still needs revision.

Acknowledgments

It is my pleasure to thank all those who contributed to this project over the years, and I apologize for any omis-

sions: W. P. Coffman for testing the pupal key; P. S. Cranston for helping with literature and types; J.H. Epler for

discussing systematic problems; E.]. Fittkau for providing the opportunity to publish; R.W. Flowers for transla-

ting Tokunaga 1964; P. Grootaert for facılitating loans of the Goetghebuer types; L.C.V. Pinder for providing an

early version of the Orthocladius calvus manuscript; F.Reiss for facilitating access to the ZSM collection and the

Thienemann types and notebooks; and, J.E. Sublette for supplying photographs of Orthocladins anteilıs.

I also thank W.M. Beck, Jr., L.C. Ferrington, Jr., D.R. Oliver, and M.E. Roussel Dillon for help with the lit-

erature; J.H. Epler, F. Reiss, A.E. Gordon, M.D. Hubbard and W.L. Peters for discussions on nomenclature and

taxonomy; P.S. Cranston, D.R. Oliver, M. Rodriguez, and K.W. Simpson for comments on the manuscript; and

USDA/CSRS, Florida A&M University, and the Alexander von Humboldt-Stiftung for supporting this research

project.

The following individuals and institutions kindly loaned specimens:

ANSP Academy of Natural Sciences of Philadelphia, S.S. Roback, D. Azuma

BMNH _ British Museum (Natural History), London, P. S. Cranston

BRUX Institut Royal des Sciences Naturelles de Belgique, Bruxelles, P. Grootaert; also R.1I.Sc. NB.

CALD B. A. Caldwell, Stone Mountain, Georgia

ENE Canadian National Collection, Ottawa, D.R. Oliver

CORE W.P. Coffman, Pittsburgh, Pennsylvanıa

DUB University College, Dublin, Ireland, P. Ashe, D. Murray

FBA Freshwater Biological Association, England, L. C. V. Pinder

FDER Florida Department of Environment, Punta Gorda, A.S. Walton, Jr.

FSCA Florida State Collection of Arthropods, Tallahassee

FWI Freshwater Institute, Winnipeg, Canada, B. Bilyj

GUTH P. Guthrie, Gainesville, Florida

HEY . M.W.Heyn, Columbus, Georgia

HUD P.L. Hudson, Ann Arbor, Michigan

INHS Illinois Natural History Survey, Urbana, D. W. Webb

KYU Kyushu University, Japan, Y. Hirashima

LUND University of Lund, Sweden, R. Danielsson

MAS P. Mason, Regina, Canada

MINN University of Minnesota, St. Paul, E.F. Cook, P.J. Clausen

NAT N. Natchev, Bistriza, Bulgarıa

NCNR North Carolina Department of Natural Resources, Raleigh, D. R. Lenat

NMI National Museum of Ireland, Dublin

NYSH New York State Department of Health, Albany, K. W. Simpson

OSU Oregon State University, Corvallis, J. Furnish

ROSS B.Rossaro, Milan, Italy

SBSK State Biological Survey of Kansas, Lawrence, L. C. Ferrington, Jr.

SUB J. E. Sublette, Pueblo, Colorado

UCOP University of Copenhagen, C. Lindegaard

USGS United States GeologicalSurvey, Atlanta and Tucson, B. Steiner, J. Doughman

USNM United States National Museum, Washington, D.C.,W. W. Wirth, R. V. Peterson

YAM M. Yamamoto, Fukuoka, Japan

ZMB Zoological Museum Bergen, Norway, O. A. Saether, G. Halvorsen

ZSM Zoologische Staatssammlung, Munich, West Germany, F. Reiss, E. J. Fittkau

Materials and Methods

This study was based on an examination of approximately 700 specimens, representing 15 species of Orthocladius

(Enorthocladius). Type material was remounted in Canada balsam after treating the specimen in sequential baths of

10% potassium hydroxide (KOH), distilled water, glacial acetic acid, 2-propanol, and 2-propanol-cedarwood oil

(Oliver & Roussel 1983). Specimens stored for long periods in alcohol dehydrate and fade. Maceration (“clearing“)

of the internal tissues is often impossible to accomplish without simultaneously making the exoskeleton invisible.

Using cold 2% KOH for a longer time rather than hot 10% KOH briefly macerates tissue while maintaining spe-

cimens in better condition. Euparal appears to cause more collapse of structures than balsam. Although balsam may

be considered the choice for mounting museum specimens, Hoyer’s may be a better alternative for teneral and pro-

blem specimens. Handling specimens through successive dehydrations and chemical treatments ıs avoided with the

use of Hoyer’s.

For the most part, terminology follows Saether (1980). Figures, counts, and measurements are my own, and fol-

low Soponis (1977). In adult males of Orthocladius, two kinds of eyes can be distinguished: those with typically,

slightly extended medial margıns, here called male-like (Fig. 12), and those with widely separated medial margıns,

here called female-like (Fig. 13). The entire length of the leg was measured, rather than the median axis (cf. Schlee

1968). Since Orthocladius adults do not have prominent tibial extensions as do adults of, e.g. Corynoneura, there

is no advantage in measuring along the median axis. In adult males, AW is the antepronotal width measured me-

dially.

Here, cast-off pupal skin(s) is/are referred to as exuviae (Ex) as recommended by Langton (1984). A cast off larval

skin is referred to as a larval skin (LS).

Only the fourth larval instar was studied, unless otherwise indicated. For the larvae the mental ratio (MR) (Sopo-

nis 1987) is used: width of the median tooth/width of fırst lateral tooth of the mentum.

Other standard ratios used are:

AR — (male, — antennal ratio (Edwards 1929): length of last flagellomere/total length of flagellomeres 1-12

AR — (larva — antennal ratio (Fittkau 1962, modified by Schlee 1966): length of basal segment/total length of ter-

minal segments

LR — legratio (Edwards 1929): length of first tarsal segment/length of tıbia

HR — hypopygium ratio (Saether 1968): length of gonocoxite/length of gonostylus

The Subgenus Orthocladius (Euorthocladius)

Euorthocladius Thienemann, 1935; 1944.

Spaniotoma (Orthocladius) group Euorthocladius. Johannsen, 1937.

Orthocladius (Orthocladius) [partim]. Andersen, 1937. Goetghebuer, 1942.

Hydrobaenus (Bryophaenocladius) [partim]. Kloet & Hincks, 1945.

Orthocladius van der Wulp [partim]. Chernovskii, 1949. Sublette & Sublette, 1965. Pankratova, 1970. Sasa& Yama-

moto, 1977. Oliver, 1981.

Orthocladius (Enorthocladius). Brundin, 1956. Fittkau et al., 1967. Hamilton etal., 1969. Kloet & Hincks, 1975. So-

ponis, 1977. Fittkau & Reiss, 1978. Pinder, 1978. Cranston, 1982. Ashe, 1983. Cranston et al., 1983. Oliver &

Roussel, 1983. Coffman & Ferrington, 1984.

Hydrobaenus group Euorthocladius. Roback, 1957a.

Hydrobaenus Fries [partim]. Wirth & Stone, 1968. Cole, 1969.

?Lapporthocladius Thienemann. Brundin, 1956.

Lapporthocladius Thienemann. Säwedal, 1978.

Type of Subgenus: Orthocladius thienemanni Kieffer, designated by Thienemann, 1935: 201.

Diagnosis

Adult males and immature stages of Orthocladius (Euorthocladius) resemble those of the other sub-

genera of Orthocladius. Most adult males of Orthocladius (Euorthocladius) can be distinguished by

the multiserial scutellars and the rounded anal point. If the scutellars are uniserial, then the female-like

eyes will distinguish most males of Orthocladius (Euorthocladins).

Pupae of O. (Eunorthocladius) can be distinguished by the lack of anal macrosetae.

Larvae of O. (Euorthocladius), except for rivulorum, can be distinguished by the brown head cap-

sule, 5-segmented antenna, robust Lauterborn organs, sparse chaetulae laterales, shortened teeth of

the mandible, and 13 to 15 teeth on the mentum.

Description

Adult Male

Small, medium or large chironomids. Yellow, green, brown, or black. Head. Eyes widely separated;

temporals uniserial except in roussellae, doubled or clumped near the coronal suture. Postorbitals

present except in suspensus and some rivulorum. AR usually 1.00-2.15 (0.80 recorded for alpine rivi-

cola). Palps normal to long, with segment 3>4, except in anteilıs, suspensus, and thienemanni, segment

3<4. Thorax. Antepronotal lobes weak to robust, wide or narrow medially. Lateral antepronotals

present; acrostichals present or absent within and between species; prealars present; dorsocentrals

uniserial except in some roussellae, biserial; scutellars usually biserial or multiserial, except in some ri-

vicola and some abiskoensis, uniserial. Mesonotal pit usually absent; if present, weakly developed.

Wings. Length 1.30—3.45 mm. Finely to moderately punctate with anal lobe right-angled, or slightly

to strongly produced. VR 1.00-1.22. Brachiolum with 1—2 setae, R with 2-12, squama with 8-40.

R4+5 bare except in single specimens of abiskoensis, calvus, roussellae, and telochaetus. Costa ending

above or distal to M3+4. R2+3 ending about '/; the distance between Ri and R4+5. Legs. Hind tibial

comb composed of 6-13 setae. Tarsal spines absent on p1; 2 spines usually present on tal and ta2 of

p2 and p3. Tarsal beard present on p3. LR1 0.57-0.77 (0.80 recorded for saxosus); LR2 0.43—0.56;

LR3 0.47—0.60. Sensilla chaetica present or absent between and within species. Sensilla chaetica pre-

sent on basal half of tal in p2 and sometimes in p3.

Hypopygium (Fig. 1). Virga present or absent between species and within species. Anal point me-

dıum to long, weak to robust, armed laterally with setae, and usually with a rounded tip, pointed tip

only in some abiskoensis and coffmani. Superior volsella collarlike except in abiskoensis, triangular. In-

ferior volsella well developed with dorsal part squared, rounded, nose-like, either covering most of

ventral part, or with ventral part extended weakly or prominently below. Gonostylus haired, robust,

with grooved spine between two strong setae. Crista dorsalis long, absent only in abiskoensis. HR

close to 2.00 or higher, except in some specimens of rivicola and thienemanni.

Variation. The tip of the anal point will apear more rounded if it is flattened down under the coverslip. Microtri-

chia appear on the anal point ın specimens from the high arctic (e. g. telochaetus, thienemannı).

The crista dorsalis can appear strongly to weakly developed, depending on the orientation of the gonostylus. The

effect of orientation on the gonostylus has been illustrated by, e.g. Oliver (1976, figs. 10-12) for Oliveria tricornis

(Oliver).

The spine on the gonostylus is grooved and appears bifid. This may not be noticed unless the gonostylus is exten-

ded, exposing the spine. Usually the spine appears to’be blunt at high magnification because of light shining between

the bifurcation. This condition is not as pronounced as in some other chironomids, e.g., Zalutschia briani Soponis

(1979).

The apodemes of the hypopygia are thickened in some species but this character is too variable within Zuortho-

cladius to be of diagnostic value, as Schlee (1968) found for the Corynoneura group. Schlein and Gratz (1972) used

daily growth in the skeletal apodemes of mosquitoes and in genitalic apodemes of muscoid flies in their studies, sug-

gesting that the genitalic apodemes of chironomids may also be subject to variations produced by daily growth.

The virga is not agood diagnostic character in the subgenus Euorthocladius. Most members of the rivicola-group

appear to have a virga present, but in some species of this group it may be present or absent. Whether the virga was

never present or subsequently lost (e.g. after mating) is unknown.

The length of the palps is positively correlated with the interocular distance in rivicola, but not in roussellae, the

only two species examined for this relationship.

The degree of the development of the antepronotal lobes corresponds to the orientation of the specimen, and is

not a useful species character.

Oliver (1970) was the first to point out that acrostichals could be present or absent within O. (Euorthocladius).

Pupa (Exuviae)

Light to dark brown; darker shading on cephalothorax; apophyses on tergites. Length 2.5—6.5 mm,

variation in all species. Largest individuals belong to roussellae, luteipes, thienemanni, and coffmani;

smallest individuals belong to rivicola, rivulorum, and saxosus.

Cephalothorax. Frontal setae absent. Frontal warts weak or absent, except in saxosus, robust; ce-

phalic tubercles usually absent, but may be weakly developed in some species; 3 precorneals, usually

clumped; 1—2 median antepronotals, 0-1 lateral antepronotals, 3 dorsocentrals in rivicola-group,

4 dorsocentrals in rivulorum-group and abiskoensis and saxosus. Development of cephalothoracic

setae variable, from weak to well developed, except in ashei, thick and robust. Thoracic horn absent

in abiskoensis; short (30— 110 um), ellipsoid, and stalked in rzvicola-group and saxosus; long (170—440

m), tubular, and bubbled or smooth in rivulorum-group. Cephalothorax can be dorsally smooth,

wrinkled, granular, or rugose, and is variable and inconsistent within species except in roussellae and

rugose.

Abdomen. Tergites with spine arrangements of 2 kinds: rows of straight spines on III-IV to VIII

(rivicola-group); or hooklets on II or II-V (rivulorum-group). Some species (abiskoensıis, rivulorum,

roussellae) with shagreen of the Orthocladius (Orthocladius) type. In all species except roussellae, seg-

ment I bare of spines. Sternites with spinules on II-VII in different patterns. Dorsal o-setae in coff-

mani, luteipes, rivulorum, roussellae, and saxosus; absent in abiskoensis, ashei, calvus, rivicola, and

thienemanni. Anal macrosetae absent. Anal lobe reduced laterally in rivzcola-group and saxosus;

strongly developed as large, circular lobes in coffmanı, suspensus, and rivnlorum-group; extended dis-

tally as long lobes ın abiskoensis and roussellae. Spines on lobes of roussellae. Seta on distal half of lobe

in ashei, abiskoensis, coffmani, kanıi, rivicola, and roussellae; at midpoint in abiskoensis, kanıı, rivi-

cola, roussellae, and thienemanni; absent ın calvus, Iuteipes, rivulorum, and saxosus. Lobe bare ven-

trally, covered with spinules dorsally, usually in a small anterior patch. Genital sheaths of males extend-

ed beyond tips of anal lobe in all species; extended beyond tips in females of /uteipes, rivicola, saxosus,

and thienemanni. Pedes spurii B absent in abiskoensis, and rivicola-group; present in saxosus and rivn-

lorum-group. Pedes spurii A present in calvus, luteipes, roussellae, and saxosus.

Variation. The hooklets and spines in the spine rows on tergites II-VIII may be directed anteriorly or poste-

riorly. The direction of these moveable spines has no value as a taxonomic character. Hooklets and spines are prob-

ably moved as the pupa leaves its site upon eclosion. Hooklets, or recurved spines, are similar to those in Orthocla-

dius (Orthocladius). In the rivicola-group, calvus has a patch of spines on tergite II; these are not recurved, but

straight and thorn-like (Fig. 37a).

The thoracic horn is stalked in the rivicola-group. The horn of saxosus is similar to that of /uteipes, with a weakly

developed stalk and a more easily collapsed horn. The thoracic horns of rivicola, thienemanni, and ashei are similar,

with amore prominent stalk and a less easily collapsed horn.

Five dorsocentrals occur in some specimens of rivulorum, and no dorsocentrals can be found in some specimens

of saxosus. The arrangement of dorsocentrals is inconsistent within species (see rivulorum, roussellae, saxosus).

Dorsocentrals can be branched or forked (ashei, rivicola).

Larva (Fourth Instar)

Live larvae with body yellow, brown, or green, or variations of these, as in saxosus (yellowish-

brown, reddish-brown, or greenish-brown). Head capsule light brown, except in saxosus, kanıi, and

rousselae, dark brown; occipital margin and mouthparts darker. Eyespots both fused and bipartite, ex-

cept in rivicola, saxosus, and thienemanni, fused. Mentum convex, with 13 teeth, except in roussellae,

15teeth, and in rivulorum, 17—21 teeth. Mandible with 3 inner teeth, 4th tooth either separate or fused;

length of apical tooth not longer than combined lengths of next 3 inner teeth except for rivnlorum;

seta interna absent only in roussellae when present with 6 branches, apices pointed; outer margin of

mandible notched opposite seta subdentalis, rest of margin smooth or crenulate. Epipharynx with 3-

toothed pecten; 4 chaetulae laterales, except in rivulorum, 9 (Thienemann’s ”moustache”); 2 chaetulae

basales; spinules variable; ungula U-shaped; premandible usually simple, although often notched, ex-

cept in calvus, roussellae and one specimen of saxosus, bifid; premandible irregularly shaped, blunt,

wide, and with a noticeable, rounded inner lobe, except ın rivulorum, pointed and without a lobe; SI

bifid; SII robust, long, simple; SIII slender, long, simple; SIV peg-like, with base; SV peg-like, with-

out base. Antenna 5-segmented; AR 1.38—3.08; Lauterborn organs robust, except in rivulorum and

roussellae, weak to moderately developed; blade usually extended to 4th or 5th segment, except in

some saxosus, extended beyond the tip. Procercus with 6 terminal and 2 lateral setae. Anal tubules

long, rounded, and subequal in abiskoensıis, rivulorum, roussellae, and thienemanni; dorsal pair shor-

ter and thicker than ventral pair in /uteipes, rivicola, and saxosus.

Variation. The color of the head capsule deteriorates after long storage in alcohol; brown head capsules may ap-

pear light yellow. Younger instars may have lighter head capsules, especially after molting. The shade or intensity

ofthe head capsule is also geographically variable. Larval skins of ashei and rıvicola reared from Norway have much

darker head capsules than larval skins of ashei reared from Ireland and larval skins of rivicola reared from Canada.

The length of the apıcal tooth of the mandible depends on the orientation of the specimen. The apical tooth can

appear greatly or slightly reduced in the same species, and there appears to be no trend within or between species.

Whether the 4th tooth of the mandible appears to be true (separate) or false (fused) is a character that is both con-

sistent and variable, depending on the species. In ashei, rıvicola, and thienemanni, the 4th tooth is fused to the man-

dible (false). In abiskoensis and rıvulorum, the 4th tooth is separated from the mandible by a groove or space (true).

Both conditions occur in saxosus. In the type mateial and in reared material from Montana, the tooth is fused; in

reared material from Alberta, the tooth is separate. In specimens of roussellae collected in the same sample from

Wyoming, some specimens have the 4th tooth fused, others separate. Chernovskii (1949) originally used this cha-

racter in his key to orthoclad larvae, and Pankratova (1970) and Soponis (1977) used this character in Orthocladius.

The large and oddly-shaped premandible was rarely orientated in a favorable position to draw. It was difficult

to determine whether or not the premandible was notched.

Setae on the larval body appear in different patterns (Fig. 57) and these may be taxonomically useful. Many of

the larvae examined, however, had either no setae or only a few short, simple setae. Setae can be lost when larvae

are not handled carefully during collection and preservation. Mounting media also affect the retention of setae dif-/

ferently, e.g. setae are retained better in Hoyer’s than in either Euparalor balsam. Setae are commonly lost from the

procercus. In rivulorum, 3 to 6 setae per procercus were observed.

The keys

To use the following keys effectively, good slides are essential. The Palearctic Orthocladius (Orthocladins) need

to be revised, and there may be difficulties keying these species to subgenus. See Soponis (1977) for labelled struc-

tures.

Key to Adult males of Orthocladins

1. Hypopygium without well developed dorsal part of inferior volsella (Fig.2)............

GBA IETE IE e Aere EIERN I Orthocladius (Eudactylocladius)

Hypopygium with well developed dorsal part of inferior volsella (Figs.1,3,4)........... 2

10.

12:

16.

Eyesiextended'dorsamedially, mälelike'(EiE112). 00 FREE

BEREE EHE EEE NASSEN VERLEIHEN, Orthocladins (Orthocladius) (part) (Soponis 1977)

Eyes widely separated, temale-Iike (Fıs- 13)... =. a0 0 ne anne a Sage ar a

Anal lobe of wing strongly produced; fore tarsal beard present; scutellar setae uniserial or

biserial; hypopygium as in Fig. 3... . Orthocladius (Pogonocladius) consobrinus (Holmgren)

With:anothercombinätion.ot. charäctersun lau. ea een alea nase

Seutellar setaetunisental Ko nous N

Seutellar.setae biserial,or. multiserial . :...... cn serie 20 ae

Gonostylus-wichirobust eristaidorsalis. (Eigs 32,33) er: 2 Han 2 2

EIER TÄLER SSPERTEI DE OT TERSPERER) Orthocladius (Enorthocladius) rıvicola Kieffer (part)

Gonastylkas with weakor nocHisea dorsalisi(Figs!7,8). HEILER

Vıirea present orsuperior volsellacollar-fike_ __ .._..... „..._..... ee

EEE EN ERE Orthocladius (Orthocladıus) (part) (Soponis 1977)

Virga absent and superior volsella triangular (Figs. 2,8). .1...4:1u.......5 26 22

EEE Orthocladius (Euorthocladius) abiskoensis Thienemann & Krüger (part)

Antennae reduced and gonostylus with large projection on dorsal edge proximally (Fig. 5)...

a a ER A Are N A LARAH EN a Orthocladius (Orthocladius) ferringtoni Soponis

Antennae not reduced and gonostylus without large projection on dorsaledge...........

Superior voksellateollarkkeifkig. 2 un U De ee ee

Superior volsella.not eollarlike.... .... „see... Orthocladius (Orthocladius) (part)

Dorsocentral setae biserial to multiserial Orthocladius (Orthocladius) smolandicus Brundın

Dprsocentraksetae uniserial.. - zu 4.2.00. 1: 2 se a ege ei 2 er

Gonostyluswith;weak or.neieristadorsalisı(Figs.7 8) na:

EHEN Sa Orthocladius (Orthocladius) abiskoensis Thienemann & Krüger (part)

Gonestylus wıthrobust.cristadorsahs 9.0. 0e 00 000.0 22.0200000 0 A

Genostylus complex; hypopygium (Fig: A). 2.0.0.0. 3 22232. I82 187 ABER BE

I ROAUERIIT ICH SEINE IE RN ZEHAWB! Orthocladius (Orthocladius) trigonolabis Edwards

Gonestylus;simple; hypopygıum otherwise... . 2.0... Sr 20 aus 5 rare at eh. 2 ee

Inferior volsella appearıng doubled, dorsal part subequal to ventral part as in some Ortho-

eladıus, (Orthocladius).(Bigs. 9,510). une Bra a: 3 ee ee ee

Inferior volsella not appearing doubled, dorsal part not subequalto ventralpart..........

Palpal,segment3>4; anal point weak (Eis. a a a es Mr le: coffmanin.sp.

Palpalsegment3<4analpointrobust (PRITNFFIIFN EIN DI ER anteilis (Roback)

Lateral antepronotals >8; hypopygium as in Figs. 11,14,15............ roussellae n. sp.

Laterallantepronetals<8;hypopysiumnotasabeve..2...... une eeucun eu

Inferior volsella with ventral part extended ventrally and laterally to dorsal part (Figs. 16, 18)

Inferior volsella with ventral part not extended, or only extended ventrally below dorsal part

Anal point with apical seta (Fig. 16); squamals >23............... telochaetus Langton

Anal point without apical seta (Figs. 18, 19);squamals<23.......... saxosus (Tokunaga)

Inferior volsella with dorsal part arched convexly (Figs. 20,21)....... rivulorum (Kieffer)

Inferior volsella with dorsal part notarched convexly.................- rivicola-group

NR IsPolackın a So a a re TE Er A Far RP

NR GES Ve ee Le ee ee wen ee ae

Interiorvolsella withidorsalparewıde,squared. (Figs. 22,23)... m... een.

Interionvolsella wıth dorsalparenarrow, long\(Eigs.25, 28)... ... 0...

Europe, North America; hypopygiumasınFig.23.......... Iuteipes Goetghebuer (part)

IepanshypopyzumasınEis Dan en kanü (Tokunaga) (part)

Balpalisesment 3>44hypopyzıumasın Eigs25, 26... .0.....0.2.0 calvus Pınder

Palpal segment 3<4; hypopygium as in Figs. 27,28. .............. thienemanni Kieffer

Inferior volsella with most of ventral part covered by dorsal part (Figs. 17,22,23).........

Inferior volsella with most of ventral part extended below dorsal part (Figs.29—33).......

Balpalsesment3=4;hypopysiumasın Fie.1Z... 22... .2 u... suspensus (Tokunaga)

Balpallsesment3>=4:hypopygaumasın E19S222,23.... rss Se

Europe, North America; hypopygiumasınFig.23.......... Iuteipes Goetghebuer (part)

japanıhy,pepySiumasınErs pam ner esse. kanı (Tokunaga) (part)

Inferior volsella with ventral part extended prominently below dorsal part (Fig. 31); Green-

amel a ee ae A Er ARE BE EEE difficıhis (Lundbeck)

Inferior volsella with ventral part less prominently extended below dorsalpart...........

Sensilla chaetica absent on ta 1 of p 2; hypopygium as in Figs.29,20.......... ashein.sp.

Sensilla chaetica present on ta 1 ofp2; hypopygium as in Figs. 32,33. .rivicola Kieffer (part)

Key to Pupae (Exuviae) of Orthocladius

Nmallobewith3anallmaerosetaen nk nn. Orthocladius (Orthocladius)

2 ee A AT ed EEE Orthocladius (Pogonocladius)

Re RN ET Orthocladius (Eudactylocladius)

Anallobewithoutanalmacrosetae. .e..... 2... Orthocladius (Euorthocladius)

Tergite II with median patch of hooklets along posterior margın (Figs.38a,39)..........

N RE N u rıvulorum-group

Tergite II usually bare (Figs. 48, 49); if wıth median patch along posterior margin, then patch

wiehstraishtspines andnothookletsi((Fıg.37a). ....... ie zn zuerse: rivicola-group

Tergite III with central round patches of strong spines anteriorly (Figs.38a,39..........

Tergite III without central round patches of strong sspinesanteriorly..................

Tergite III with large (>200 um wide) round patch of spines anteriorly that reaches

midlimeioßtersitel(Eien3g) ran ee ee u coffmani.n. sp.

Tergite III with small (<150 um wide) round patch of spines anteriorly that does not reach

midlinssoktersitel(E 19-38. a) elseA ea eg Vers Arsen (a air

Thoracice horn bubbled (Fig. 34 b); tergites IV-VI with rows of spines along posterior

REIT EB er en EEE ER EI EN rivulorum Kieffer

10.

KT.

12.

Thoracic horn smooth; tergites IV—-VI without rows of spines along posterior margins....

al a N a ie Meleabe Be Be suspensus (Tokunaga)“

Analllobe,withspines ontips (ig AD.r user a roussellae n. sp.

Analilobe withoutspineson ups sr... u ine er el ee

Frontal warts robust (Fig. 43); thoracic horn present (Fig. 34 e); hooklets on tergite II with

>100 spines; hooklets in a large patch of 3-5 rows (Fig. 44).......... saxosus (Tokunaga)

Frontal warts weak or absent; thoracic horn absent; hooklets on tergite II with <50 spines;

hooklets in a small patch of 1-2 rows (Fig. 45)........ abiskoensis Thienemann & Krüger

Tergite II with median patch of strong thorn-like spines along posterior margin (Fig. 37 a)

BREEANESUN nn ni nr at EIERN. TREND PELMEN PEN E A calvus Pinder

Terstellibaret Mill. a... ee Me tar 75 MS

Pedes spurii A present on sternite VI; spines individually slender (Fig. 37 b) in rows on

posterior margins.of tergıtes,(Figs- 46, AY ea ne Bee ensure. Iuteipes Goetghebuer

Pedes spurii A absent on sternite VI; spines individually robust (Figs. 37 c, d, e) in rows on

POStEHIOr MAargıns OLtergites Zr... cyan a a rauch ae er

Tergite III with rows of spines on posterior margin (Fig. 42)........ thienemanni Kieffer

Tergite III wıthout rows of spines on posterior margın .. e.2.2.2.02. Kersin

Dorsocentrals thick, robust (Fig. 36); spines in spine rows on posterior margins of tergites

IV—-VIll individually robust (Fig. 37 e); spines on VI<40 (Fig.48)........... ashein.sp.

Dorsocentrals normally developed (Fig. 35); spines in spine rows on posterior margins of

tergites IV—VIII individually normally developed (Fig. 37 c); spines on VI>40..........

Europe, North America, abdomen asın He Age. a........02... rivicola Kieffer

aa ee ER kan (Tokunaga)*

Mentumswich reiht! vs. san: Orthocladius (Orthocladius) lignicola Kieffer

Mentum wath 3 or moreteeth.. 2.1 ee a

Mentum wich 13teeth. 2... 2... 00 Be Orthocladius (Enorthocladius) (part)

Mentumswith I3xteeth.s... 245 aa a ea Ra ee ee Fe re

Mentum with 15 teeth (Fig. 50 e) mandible without seta interna (Fig.50c).. .roussellae n. sp.

Mentum with >15 teeth (Fig. 51 e); mandible with seta interna .. ......... u... .enaes-

Burspes@Noxth. America; Eig; 51.4... EINE FIT rivulorum Kieffer

Japan;...... Ms erllsorunae erlassenen suspensus (Tokunaga)“

Head capsuleiyellov. su Hp nen aA: Orthocladius (Orthocladius)

Head zapsuleibrown.: „iin. tesa teen ieh a Dr) u Dr Be

Antenna 4-sesmented; Lauterborn organs weak... .n.. u. un a ee

aa on rl rn HR Orthocladins (Pogonocladins) consobrinus (Holmgren)

Antenna5-segmented; Lauterborn organszebust. ......u.c nen. ae

* based on literature; specimens not seen

leadicapsuleldarksreddishibrownee sr. 20 Orthocladius (Eudactylocladius)

Head capsule light to dark brown, notreddish........ Orthocladius (Enorthocladins)part 8

8. Mentum with median tooth >1.5X width of Ist lateral (MR >1.5)................... 9

Mentum with median tooth <1.5X width of Ist lateral (MR <1.5).................... 12

DRBEINRE 1:89 2.2 1.8 272.2. RE ET ET NL NE NE EBFRHRITN U 10

ENBER SS 0, See a a re ERNRNEN NHE PIERRE. RABEN SEHEN. Tb BANN 11

IDErremandibleibitid. wessen ers Aansersatar Aelen: calvus Pinder

Bremandiblesimpleie ns 2 en 0. 2 RER NDR ER thienemanni Kietfer

BSR urope, N orthr’Americasjheadkcapsulebrowne 2... 2... 20.0... Inteipes Goetghebuer

apansiheadkeapsuleidarkbrownerman ne ee kanı (Tokunaga)*

IDREENR SO were 1:.80). Den a RER ashei nn. sp., rivicola Kieffer

INRehichten (1480) were ar ee rn er Eh ee Re Ace TE 13

IEET-Teackcapsuledark browaen me a en saxosus (Tokunaga)

Kleadieapsulellischrbrowntonbrown 2 a abiskoensis Thienemann & Krüger

Orthocladius (Euorthocladius) abiskoensis Thienemann & Krüger

Figs. 6-8, 45, 60

”Orthocladins” abiskoensis Thienemann & Krüger, 1937: 257—265, 267, figs. 1a, 3a, 4a, 5, 6a, 8a, 9a [pupal, larval

description].

Lapporthocladius abiskoensis (Thienemann & Krüger), 1937: 266.

Orthocladius (s. str.) abiskoensis Edwards, 1937: 144—145 [adult description].

Lapporthocladius abiskoensis (Edwards). Zavrel, 1938: 8, 9 [comparative analysis of larvae]. Thienemann, 1941: 66,

68, 82, 150, 180 [ecology, distribution]. Thienemann, 1944: 564, 647, figs. 20, 21, 197a, 198a [in pupal, larval

keys]. Thienemann, 1954: 182, 188, 355, 357 [notes]. Brundin, 1956: 103 [systematic placement]. Fittkau et al.,

1967: 358 [checklist]. Fittkau & Reiss, 1978: 418 [checklist].

Orthocladıius (O.) abiskoensis Edwards. Goetghebuer, 1942: 35, 37 [in male key, adult description].

”Orthocladius” abiskoensis Edwards. Saether, 1969: 65 [note].

Orthocladius (Lapporthocladius) abiskoensis Edwards. Pankratova,.1970: 173, 174, 182, 183, fig. 110 [pupal, larval

description, in pupal, larval keys].

Orthocladius (Enorthocladius) abiskoensis Edw. Säwedal, 1978: 85, 86 [ecology].

Orthocladius (Euorthocladius) Type III Soponis, 1977: 15-17, fig. 122 [pupal, larval diagnosis, in pupal, larval

keys].

[non] Orthocladins (Enorthocladins) Type III sp., Simpson & Bode, 1980: 52 [misidentification of /uteipes and rıvı-

cola].

Type Locality: Sweden, Lappland nr. Abisko.

Type Material: Lectotype: Male pupa, Lappland, Sweden, 10 VI 1936, A. Thienemann, labelled by Thienemann

as Orthocladius abiskoensis Edw. Lappland 1936 3d (ZSM). On a slide with paralectotypes, lectotype indicated in

ink as in Fig. 7. Paralectotypes (23): same data as lectotype, 8Ex (7M, IF), 1 larval head capsule, 3MP, 1FP, 1MP

abdomen. Same data as lectotype except 3c, 4FP, 5L. The specimens are mounted in balsam on a total of 6 slides and

kept at the ZSM. According to notes of Thienemann (pers. comm. F. Reiss), 3d indicates that diverse chironomid

larvae were reared, but only 2 specimens hatched and the others died (as pupae); 3c indicates that isolated larvae of

abiskoensis were reared. The specimens described above are hereby designated lectotype and paralectotype.

* based on literature; specimens not seen

Diagnosıs

Orthocladius abiskoensis can be distinguished from other Holarctic species of ©. (Euorthocladius)

by a combination of characters. Adult Male: details of the hypopygium (Figs. 6-8). Pupa: thoracic

horn absent, frontal warts weak or absent, tergites II-V with hooklets on posterior margins, hooklets

in small patches of 1-2 rows; tergite III without central round patches of strong spines anteriorly.

Larva: mentum with 13 teeth, MR<1.5, AR>1.80, head capsule brown.

Derivation of Name: Abisko; L. ensis, denoting place, locality.

Description

Adult Male (n=17)

Brown. Small to medium species. Head. Verticals 9— 14, postorbitals 1-3. Palps long with 3>4. AR

1.22—1.71. Thorax. Lateral antepronotals 2-6. Acrostichals 1-13, robust, begin within 2AW. Dor-

socentrals 4-12. Prealars 4-7. Scutellars 7-14, usually uniserial, less often biserial (20%). Wing.

Length 1.38—2.32 mm. R with 2-10 setae. Squamals 14—33. In one specimen 2 setae on R4+5. VR

1.03—1.12. Anal lobe moderately produced. Legs. LR1 0.57—0.67. LR2 0.44—0.53. LR3 0.48—0.59.

Sensilla chaetica on tal of p2, 6-22 (15), and p3, 0-8 (15). Hypopygium (Figs. 6-8). Virga absent.

Superior volsella triangular with pointed or blunt apices. Inferior volsella with dorsal part nose-like,

covering most of ventral part. Crista dorsalis weak.

Variation. The material examined contains variants in 2 or 3 populations. Males of high arctic populations from

Isachsen, Northwest Territories, are large specimens, with a robust anal point, squared inferior volsella, and full su-

perior volsella (Fig. 8a). Males from more temperate populations of Caribou Bar Creek, Yukon Territories,

(Fig.8c) more closely resemble Edward’s original material (Fig. 6) from Abisko in the superior volsella and slender

anal point. At least one specimen from Abisko has a robust anal point. The immature stages of rearings from three

Canadian sites (Hazen Camp, Banks Island, Caribou Bar Creek) agree with each other in diagnostic features. The

scutellars can be either uniserial or biserial. The number of sensilla chaetica varies in this species. The crista dorsalis

is not evident in all specimens.

Edwards (1937) separated abiskoensis by the relative lengths of palpal segments 5 to 4, aratio of 1.5. Here (n=14),

the ratio varies from 1.2 to 1.7. The color of the thorax or the patterns of the scutal stripes in abiskoensis, characters

used by Edwards, were not analysed here because these characters cannot be accurately assessed in slide-mounted

material. The shading of the scutellum is also unreliable in slide-mounted specimens, since it is essentially the same

in all the material.

Pupa (Exuviae)

Light brown, with dark apophyses on II-VII (variable); length about 3.0—4.25 mm (10). Cephalo-

thorax. Frontal warts weak and cephalic tubercles absent. Precorneals clumped; 2 median anteprono-

tals, 1 lateral antepronotal, moderately developed; 4 dorsocentrals, slightly shorter than precorneals

but thicker; arrangement varies. Thoracic horn absent. Thorax dorsally slightly wrinkled along eclo-

sıon line.

Abdomen (Fig. 45). Tergites: I bare; II-V with small central patch of recurved hooklets in 1-2

rows along posterior margin; III-VIII with large central patch of spinules separated from posterior

spine patch. Sternites: I bare; II- VII with spinules anteriorly in varying amounts; VIII with 2 off-cen-

ter patches of spinules anteriorly.

Setae on segments I— VIII:

2 I

SUR PETE,

Aynase daran 4

Anal lobe developed as slender processes with tendency for tips to curve inwards; 2 robust or fine

setae, one on proximal half and one at midpoint; genital sheaths extended beyond lobe in male, not in

female. Pedes spurii A, pedes spurii B absent.

Variation. Spine patterns are varıable: the sternites may appear bare, and anterior spine patches on the tergites

may be less developed than described here.

Larva (Fourth Instar)

Body yellow or brown. Head capsule brown. Eyespots bipartite or fused. Head capsule (Fig. 60c)

widest midway between eyespots and postoccipital margin. Mentum (Fig. 60b) convex with 13 teeth,

median tooth about as wide as 1st lateral; MR 1.0—1.3 (3); median tooth as high as Ist lateral. Ventro-

mental plates extended anteriorly between 2nd and 3rd laterals. Epipharynx (Fig. 60a) with premandı-

ble simple, narrowed before enlarged apex; apex is notched in Thienemann material. Chaetula laterales

sparse. Mandible (Fig. 60g) with apical tooth as long or longer than Ist inner tooth; outer margin

notched opposite seta subdentalis, rest of margin smooth except for occasional notch posteriorly; seta

interna present. Antenna (Fig. 60h) with robust Lauterborn organs; blade extended to 4th segment.

AR 2.00--2.50 (3). Body bare except in one specimen, haired. Anal tubules (Fig. 60f) subequal.

Variation. Pankratova (1970) described the body as bare, greenish-brown, and the head capsule as dark brown.

She described the premandible with 3 blunt teeth. In material examined here, the premandible appears simple.

Biology. Larvae and pupae were collected near Abisko in the moss of a spring. Pupae in gelatinous,

half-ellipsoid cases were also collected on bare stones, without vegetation. The species occurs in cold

rivers, streams, and springs. Males swarm beside Micropsectra (?) brunippes Zett. (Thienemann &

Krüger, 1937, Thienemann 1941, 1954). Orthocladius abiskoensis was previously recorded only from

high latitudes, but exuviae have also been collected in Kansas. Adults emerge in June and July in the

arctic, and in March in Kansas.

Distribution. Palearctic: Sweden. Nearctic: Canada, USA.

Material Examined. Type material. Non-type Material: Canada (CNC). Northwest Territories: Isachsen, 14-VII-1960,

J. F. Mc Alpine, CH1075, 10M, 3F; Hazen Camp, 81° 49’N 71° 18’W, 13-VII-1961, D. R. Oliver, CH1133, 1M w/Ex; same

data except 27-VII-1961, Ch1047, 2MP, 2Ex; Banks Is., Masik R., 4-VI-1968, W. R. M. Mason, CH2063, M, F in copula;

Harris River, 61° 52’N 121° 19°W, 18-V-1973, FWI Pipeline Proj., CH803.12, 1M w/Ex: CH803.23, 1MP; Bathurst Is., 75°

24’N, 100° 24’W, 25-VII-84, B. Hayes, 1M w/Ex, Yukon Territory: Carıbou Bar Creek, 67° 28’N 140° 37’W, 11-VI-1973,

D. R. Oliver, CH874, 3M, 1P, 7Ex; same data except 20-VI-1972, CH126, 1M; 19-VI-1972, CH128, 1M; 15-VI-1973,

CH562.1, M w/Ex, LS; 18-VI-1973, J. Robillard, CH564.6, M w/Ex, LS, CH564.10, MP w/LS, 29-VI-1972, FWI Pipeline

Proj., CH6205, IL. Sweden: Lappland, 1936, 3d, Thienemann, 5P, 8Ex, ILS; Lappland, 1938, No. 125, Thienemann, 7L

(ZSM). USA. Kansas: Leavenworth Co., Plum Creek, 1.2 mi S, 0.2 mi E of Kickapoo, 24-III-1982, L. Ferrington, 2Ex

(KSBS).

Remarks. This species was collected by Thienemann as adult and immature stages in the summer of 1936 at a

spring among dwarf birches, near the road Abisko-Björkliden in Swedish Lappland (Thienemann & Krüger 1937,

Edwards 1937, Säwedal 1978). Thienemann sent the adult males to Edwards, who initially determined them as „Or-

thocladius ? rubicundus Mg. var. or decoratus Holmgren?“ (Thienemann & Krüger 1937). Thienemann questioned

the determination of the adult because of the associated immature stages. The pupa of rubicundus belonged to Thie-

nemann’s Rheorthocladius (=Orthocladins s. str.), and Thienemann thought that the unusual pupa of abiskoensis

belonged to a new species.

To provide Thienemann with a name for the new species, Edwards (1937) published a brief adult description, pri-

marily distinguishing abiskoensis from rubicundus and decoratus by the shading of color on the thorax. He also used

the lengths of palpal segments 3 and 4 (here 4 and 5) to distinguish abiskoensis from rubicundus, adding that the cha-

racter ıs individually variable. Edwards regarded abiskoensis as a pupal species: ”the pupae are so strikingly diffe-

rent, but the adults scarcely if at all distinguishable.”

The same year Thienemann & Krüger (1937) provided a detailed description including figures of the pupa and

larva of abiskoensis, comparing it with the immature stages of rubicundus. They cited Edwards’s pending adult

description, but their own paper was published first. Thienemann & Krüger (1937) was published 15 March 1937

and received in the BM(NH) 8 April 1937. Edwards (1937) was published July 1937 and received in the BM(NH)

16 July 1937 (pers. comm. P. S. Cranston). According to Article 50 of the rules of the International Code of Zoolo-

gical Nomenclature, ı.e. the author of a name is the person who first validly publishes it, the authors of abiskoensis

are Thienemann & Krüger.

Thienemann & Krüger (1937) used ”Orthocladius” abiskoensis in the title and referred to the binomen O. abisko-

ensis once again in the paper. However, inthe last paragraph of the summary, they erected the genus Lapporthocla-

dius to accommodate abiskoensis. Because the authors referred twice to abiskoensis as Orthocladius, and because the

original specimens are labelled in Thienemann’s handwriting as Orthocladius abiskoensis, this is interpreted to mean

that the authors described abiskoensis in Orthocladius, and then erected Lapporthocladins for the species. Thiene-

mann (1941, 1944, 1954) and others (Fittkau et al. 1967, Fittkau & Reiss 1978) placed abiskoensis in the monotypic

genus Lapporthocladius. Säwedal (1978) synonymyzed Lapporthocladins with Orthocladins.

Hamilton et al. (1969) recommended that a genus should have all three life stages in a relatively discernible group.

Because the male of abiskoensis is not distinguishable from other adult Orthocladius at the generic level, abiskoensis

is not placed in another genus, ı.e. Lapporthocladius. Also, abiskoensis is not placed in its own subgenus because the

immature stages belong to O. (Euorthocladins) as defined here.

Zavrel (1938) considered the generic placement of abiskoensis in a description of the immature stages of Orthocla-

dius frigidus. He concluded that the larva of abiskoensis is closer to the larvae of Thienemann’s Euorthocladius (ex-

cepting rivulorum) than to the larva of frigidus, and that on the whole, abiskoensis belongs to Euorthocladius.

Pankratova (1970) redescribed the pupa and larva of abiskoensis, reproducing the figures of Thienemann & Krü-

ger (1937). She placed abiskoensis in the genus Orthocladius, subgenus Lapporthocladius, and stated that the species

would probably occur in the USSR.

Goetghebuer (1942), treating adults, followed Edwards (1937) and separated abiskoensis from rubicundus prima-

rıly on AR, thoracic color, and palpal proportions. He provided no figures.

Brundin (1956) reviewed the systematic position of abiskoensis as an example of incongruity of adult and imma-

ture chironomids. He examined two males (one a pupa) of Edwards’s original material, specimens in poor condition

due to long storage in alcohol. Brundin observed strewn scutellar setae that would place the adult in his subgenus

O. (Euorthocladius). However, he concluded that the position of abiskoensis was still unstable. Saether (1969) refer-

red to the incongruity, and both Soponis (1977), as Type III, and Säwedal (1978) placed abiskoensis in O. (Enortho-

cladius).

Until now, no figure of the hypopygium of abiskoensis has been available. It was impossible to identify the adult

of this species without the associated pupal skin. The species is still difficult to identify in the adult male, but this

situation occurs in other Orthocladius, not just abiskoensis.

Orthocladius (Euorthocladius) anteilis (Roback)

Fig. 10

Hydrobaenus anteilis Roback, 1957b: 14, figs. 41-45 [description of female]. Roback, 1959: 2—3, figs. 7-10

[description of male and female]. Cole, 1969: 101 [notes].

Orthocladius anteilis (Roback). Sublette & Sublette, 1965: 155 [checklist].

Type Locality: USA: Utah, Provo River.

Type Material: Holotype. Female, remounted by M. E. Roussel Dillon, in Canada balsam under 5 coverslips,

genitalıa in lateral view. Original white label: 8. ant. wing Hydrobaenus anteilis Roback 6803 det. $.S. Roback. Red

label: Utah, Summit Co., South Fork of Provo River on Stewart’s Ranch, 20 Feb. 1954, Gerald D. Brooks (ANSP).

Paratype. Female, same data, not seen, reportedly in University of Utah collection.

Diagnosis: The male of Orthocladius anteilis can be distinguished from other males of O. (Euortho-

cladius) by palpal segment 3<4 and details of the hypopygium (Fig. 10). The pupa and larva are

unknown.

Derivation of Name: ant; L. ilis, having the nature or quality of.

Description: See Roback (1957b, 1959).

Biology: Adults have been collected along rivers in western USA.

Distribution: Nearctic: USA.

Material Examined: Type Material. Non-type Material: Idaho: Freemont-Teton Co. border, north Fork of Teton R.,

Hwy. 32, 6-III-1965, A. v. Nebeker (MINN), 1 M. Photographs of males and females of Montana specimens of Roback

(1959), supplied by J. E. Sublette.

Remarks: Roback (1957b) described this species from two females collected along the Provo River in Utah. Later

Roback (1959) described the male and gave a further description of the female based on five males and two females

collected along the Blackfoot and Clard Fork Rivers in Montana.

The holotype is an adult female, but the female of anteilis has never been associated through rearing or by copu-

lation with the male. The males and females from Montana, also unassociated, do not convincingly belong to antei-

his. Other species of ©. (Enorthocladius), e.g. saxosus and coffmani, occur in mountain rivers of the western USA

so that locality and habitat cannot determine species in this subgenus. Although the study of females has advanced

(e.g. Saether 1977) since Roback’s description, it ıs still not possible to distinguish females of O. (Enorthocladius)

to species.

To define the limits of anteilis, it is necessary to study reared females and reared males. Until such specimens be-

come available, the males described by Roback (1959) are tentatively recognized as anteilıs.

Orthocladius (Euorthocladius) ashei n.sp.

Figs. 29, 30, 36, 37 e, 48, 54

Orthocladius rivicola Kieffer. Thienemann, 1911: 637 [pupa].

Orthocladius (Euorthocladius) cf. thienemanni (Kieffer). Halvorsen et al., 1982: 119 [record ın Norway].

Orthocladius (Euorthocladius) rivicola ß Rossaro, 1982: fig. 31 [pupa]. Langton, 1984: 142, fig. 49 [pupa].

Orthocladius (Euorthocladins) ? rusticus Goetghebuer. Murray & Ashe, 1983: 224, 225, 230 [checklist, pupa].

Type Locality: Ireland: River Flesk.

Type Material: Holotype. Reared male with larval skin and exuviae. Ireland, Kerry Co., Sta. 6, Clydagh Br., Ri-

ver Flesk, 16-V-1978, Drift D7, W114826, reared 16th- 19th, P. Ashe, ın gelatinous case. On slide, deposited in the

National Museum of Ireland, Type No. NMI 106: 1984. Paratypes (83). Same data as holotype, 1 reared M w/LS,

l reared M with LS, Fx Bulgaria, r. Bistriza, outfall, 14-IV-1971, N. Natchev, 1 Ex (NAT). Germany. River Isar,

ca. 500 m oberhalb Loisach-Mündung, 3-IV-1986, F. Reiss, 5 Ex; Stauseen, Unterer Inn, drift, 6-IV-1978, Egla, F.

Kohmann, 1 Ex; Westfalia, Urf-Talsperre, Sta. I, 4+-IV-1910, A. Thienemann, 5 Ex; Westfalia, Glor, 7-VI-1908, A.

Thienemann, 2 Ex; Westfalia, Fulbecke-Talsperre, 14-IX-1909, Oberfl., A. Thienemann, 1 Ex Ireland. Kerry Co.,

River Flesk, Sta. 1a, V964894, Ranunculus, 14— 17-V-1978, P. Ashe, 1 reared F w/Ex (slide), 2 reared F w/Ex (alco-

hol); same data except 14—-19-V-1978, 1 F w/Ex (alcohol); River Doddler, above Bohemabroinne Bridge, 3-IV-

1978, P. Ashe, 1 F w/Ex, probable LS$ (alcohol). Italy. Po River, 1975, 1976, B. Rossaro, 12 Ex (slides), 40 Fx (alco-

hol). Norway. Ekse, HOi: Vaksdal, “The Weir Project,“ 11-VI-1979, E. Wilassen, 1 reared M w/Ex, LS (ZMB)

(data in Halvorsen et al., 1982). Sweden. Lappland, Thienemann: River Abisko, 24-IV-1936, 48a, ex mosses, 2 Ex;

Lake Abisko, 23-IV-1936, 40, ex moses, 3 Ex; Lke Kanevegge, 17-VII.1937, 98c, surface drift, 2 Ex; delta of River,

17-VII-1937; Abisko, 19-VII-1937, 101, surface drift, 1 Ex (ZSM). Paratypes will be deposited as follows: 2 reared

F, Ireland, alcohol (NMI, No. NMI 106: 1984); 1 reared F, Ireland, alcohol (BMNH); 1 reared M, Ireland, slide,

l reared F, Ireland, alcohol (DUB); 1 reared M, slide Ireland (FSCA); 1 reared M, Norway, slide (ZMB); 1 reared

F, Ireland, slide, (ZSM); Ex, Bulgaria, slide, (NAT); Fx, Italy, alcohol, (ROSS); Ex, Italy, slides (CNC, BMNH,

FSCA, COFF); Ex, Sweden and Germany (ZSM).

Diagnosıs:

Orthocladius ashei can be distinguished from other Holarctic O. (Euorthocladins) by acombination

of characters. Adult male: low AR (<1.80),'absence of sensilla chaetica on tal of p2, and details of the

hypopygium (Figs. 29, 30). Pupa: tergite II bare, robust dorsocentrals and robust spines on margins

of tergites IV—- VIII. Larva: mentum with 13 teeth, MR<1.5; cannot be distinguished from rivicola.

Derivation of Name: this species is named after Patrick Ashe, for his help in providing reared material and for

his interest in this new species.

17.

Description

Adult Male (n=4)

Dark brown to light yellow. Small species. Head. Verticals 12-13, postorbitals 1—2 (3). Palps nor-

mal with 3>4. AR 1.18-1.62. Thorax. Lateral antepronotals 5-7 (3). Acrostichals 5-8, robust, be-

ginning within 1-2 AW. Dorsocentrals 8-11. Prealars 5-7. Scutellars 7-15 (3), weakly biserial

(Norway) to multiserial. Wing. Length 1.68-1.90 mm (3). R with 6-10 setae. Squamals 8-18. VR

1.03—1.22 (2). Anal lobe slıghtly produced. Legs. LR1 0.69—0.77 (3). LR2 0.48—0.56 (2). LR3

0.53—0.59 (3). Sensilla chaetica absent. Hypopygium (Figs. 29, 30). Virga present, moderately well

developed. Superior volsella collar-like. Inferior volsella with dorsal part nose-like, or squared, and

ventral part weakly extended below. Crista dorsalis long.

Pupa (Exuviae)

Light brown; length 3.1—3.4 mm (5).

Cephalothorax. Frontal warts absent; cephalic tubercles weak. Precorneals clumped, PC1, thick

and almost 2x as long as PC2 and PC3; 1 median antepronotal, 1 lateral antepronotal, 3 dorsocentrals

(Fig. 36), slightly longer than PC1 and thicker. Thoracic horn ellipsoid, light brown, clear, stalked,

length 30-60 um (5). Thorax dorsally wrinkled to granulose along eclosion line and anterior to wing

base.

Abdomen (Fig. 48). Tergites: I-III bare; IV-VIII with rows of individually heavy spines along

posterior margin (Fig. 37e); V-VIII with central patches of robust spinules anteriorly. Sternites: I,

VIII bare; II, III with large central patch of spinules; IV—VII with anterior patch of spinules.

Setae on segments I-VIII (robust):

Sie ui zul > int a ar E DE ze 13. 3 m Z

a Ad 4‘ DE 20, 07.0.7 0,0

Anal lobe greatly reduced; 2 setae on distal half. Pedes spurii B and pedes spurii A absent. Genital

sheaths extended beyond lobe in male, not in female.

Variation. Dorsocentrals can be forked. The spines in the posterior rows on tergites IV— VIII are individually ro-

bust (Fig. 37e), much more robust than those in rivicola (Fig., 37c). The number of spines in these rows are higher

in females than in males, and are not significantly different.

ashei females (n = 5) ashei males (n = 5) Student’s t

IV 30.60 + 11.04 (20-45) 22.60 + 8.20 (14-33) 1.3009

V 33.20 # 8.87 (27-48) 25.60 + 5.86 (20-34) 1.5988

VL 128.20, 2,832 (22.40) 22.20 + 6.26 (18-33) 1.2887

VII 22.20 + 7.22 (16-30) 18.60 + 2.97 (14-22) 1.0307

VII 16.00 + 5.20 (12-25) 14.60 + 3.36 (11-20) 0.5058

The number of spines in these rows can be used to distinguish most specimens of ashei from rivicola ‚and are signi-

ficantly different.

ashei (n = 10) rivicola (n = 10) Student’s t

IV 26.60 + 10.09 (14-45) 64.10 + 25.30 (36-104) 4.3544

V 29.40 + 8.14 (20-48) 69.30 + 21.15 (47-100) 5.5676

VI 25,20, 22277:630.(118409) 62210822209 12241103) 5.2428

Vu 20.40 # 5.54 (14-30) 50:70.+.18:874337- 190) 4.8725

VII BSIEHIE 25) 35A0rE 16245, 2079) *3.7893

(p >.001 except for *,p >.01)

Exuviae from Italy show variation in size, shade of color, and thickness and length of dorsocentrals. Generally

ashei is lighter and more weakly chitinized than rıvicola, and the cephalothorax of ashei is as light as the abdomen,

whereas in rivicola the cephalothorax tends to be darker than the abdomen. Separation of exuviae of ashei from

rivicola in alcohol is not foolproof, and slides should be made for positive determinations.

Larva (Fourth Instar)

Body green with blue tinge when live (Thienemann notebooks). Head capsule brown; preserved,

yellow. Eyespots absent in reared specimens. Mentum (Fig. 54e) with 13 teeth, median tooth about as

wide as 1st lateral; MR 1.3 (3); median tooth as high as Ist lateral. Ventromental plates extended ante-

riorly between 2nd and 3rd laterals. Premandible simple, with blunt or squared apex; apex may appear

notched. Chaetula laterales sparse. Mandible (Fig. 54c) wıth apıcal tooth as long or longer than Ist

inner; outer margin notched opposite seta subdentalis; rest of margin smooth except for occasional

notch posteriorly; seta interna present. Antenna (Fig. 54d) with robust Lauterborn organs; blade ex-

tended to 4th segment. AR 1.80 (1). Body with simple setae, some short and stiff, some long and

curved, apparently arranged like that of saxosus. Anal tubules not distinguishable.

Biology: The larvae live in gelatinous tubes in running water and are associated with plants. Ashe

reared larvae from Ranunculus, and Thienemann collected exuviae from mosses in Lake Abisko and

River Abisko. The species occurs with rivzcola in Brehm, Italy, and River Isar, Germany (Ex). Adults

emerge in April, May, and June.

Distribution: Palearctic: Bulgaria, France, Germany, Ireland, Italy, Norway, Sweden.

Remarks: This species was reared by Patrick Ashe from the River Flesk, Ireland. Originally Ashe (pers. comm.)

suspected that this species might be Orthocladius rusticus Goetghebuer, based on a slide of an exuviae labelled rustz-

cus in the Humphries collections (Murray & Ashe 1983). Professor Humphries reared larvae to adults, then sent the

adults to Goetghebuer or Freeman for 3 positive determination to species. When the name was provided she labelled

the associated immature material with the corresponding determination. The holotype male of Orthocladius rusti-

cus Goetghebuer belongs to Chaetocladins (Soponis 1986) and is not conspecific with Orthocladius ashei.

Dr. Declan Murray (Murray & Ashe 1983) has seen material of O. ashei in the Humphries collection collected

by Thienemann in “Norway”, identified by Thienemann as Euorthocladius thienemanni. Murray has collected

ashei in Norway and France.

Orthocladius ashei is morphologically similar ın all stages to rivicola: the larvae of these two species could not be

distingushed. These two species can most easily be distinguished as pupae.

Orthocladius (Euorthocladius) calvus Pinder

Figs. 25a, 26, 37a, 37f

Orthocladius (Euorthocladius) calvus Pinder, 1985: 235—241, figs. 1-3 [description of male, female, pupa, and

larval].

Orthocladius (Enorthocladius) calvus Pinder. Ladle et al., 1985: 243—254 [biology]].

Orthocladius (Euorthocladius) Pel. Langton, 1984: 140, figs. 48b [in pupal key].

Spaniotoma (Orthocladius) thienemanni Kieffer. Edwards, 1929: 344, 345, fig. 6m.

Type Locality: England: Dorset, Waterston.

Type Material: Holotype (not seen). Male with associated exuviae, Dorset, Waterston experimental channel,

7 May 1981, J.A.B. Bass (BMNH). Assorted paratypes, BMNH and FBA (Pinder 1985).

Diagnosıs:

Orthocladius calvus can be distinguished from other Holoarctic species of ©. (Enorthocladius) by

acombination of characters. Adult Male: high AR (1.73—2.08), palpal segment 3>4, and details of the

hypopygium (Figs. 25, 26). Pupa: patch of straight thorn-like spines on tergite II (Fig. 37a) and rows

of spines on posterior margins of tergites III— VIII. Larva: mentum with 13 teeth, MR>1.5, high AR

(>2.00), and premandible bifid.

Derivation of Name: L. calvus, hairless. Pinder (1985) named this species for the absence of dorsal setae on the

anal lobe of the pupa.

Description: See Pinder (1985).

Biology: Larvae are early colonızers of artificial recirculating streams where they inhabit gravel

(Pinder 1985). For a detailed account of growth, development, and production of calvus see Ladle et

al. (1985). Based on adult males, Orthocladius calvus occurs with thienemanni in the River Schwen-

tine.

Distritution: Palearctic: England, Germany.

Material Examined: Paratypes. England, Dorset, Waterston Experimental Channel, 7-V-1981, coll. J.A.B. Bass, L.C.V.

Pinder, IM w/Ex, IF w/Ex, IL. Nontypes. England. Hitchin, Herts, 28-IV-1916, F. W. Edwards, 1916-105, 1M (BMNH)

(misident. of thienemanni); Gloucester, Minchinghampton, 16-IV-1893, Miss G. Ricardo, B. M. 1920-126, 1M (BMNH)

(misident. of thienemanni). Germany. River Schwentine, East Holstein, 1935, leg. A. Thienemann, Schwentine 1935 S. 4,

1 M (ZSM) (misident. of thienemanni).

Remarks: Pinder (1985) described this species from several males and females with associated exuviae, and from

larvae collected in an artificial recirculating stream system in southern England.

The adult males of calvus are very similar to thienemanni and will present problems in identification unless asso-

ciated exuviae are available. The male of calvus can be distinguished from that of thienemanni by therrelative lengths

of palpal segments 3 and 4, and by the relatively straight margin of the dorsal part of the inferior volsella. Although

this hypopygial character holds for the type material and one male from River Schwentine in Germany (Fig. 26),

the margin in the calvus from Gloucester, England looks rounded (Fig. 25a) as in all thienemanni (Figs. 27, 28), and

the margin in thienemanni from River Schwentine (Fig. 25b) looks like that in calvus (Fig. 26). The Herts material

was identified as O. thienemanni by Edwards, but this material belongs to calvus based on the relative length of pal-

pal segment 3 and 4. Brundin’s (1956) figure of O. thienemanni looks like thienemanniı.

The exuviae of calvus and Inteipes (Figs. 46, 47) are morphologically similar; calvus can be distinguished by the

central spine patch on the posterior margin of tergite Il (Fig. 37a) and the less robust spines in the tergal spine rows

(Fig. 37f) and shagreen. The bifid premandible of the larva is distinctive in calvus. However, whether or not the pre-

mandible is bifid is difficult to determine in O. (Enorthocladius), and has not been clearly established for most spe-

cies.

Additions to Pinder’s (1985) description include: Male. Head female-like, scutellars, biserial, sensilla chaetica

absent in paratype male, present on tal of p2 in nontype male. Pupa. Pedes spurii A on IV or V to VII.

Orthocladius (Euorthocladius) coffmani n. sp.

Figs. 9, 34a, 39

Orthocladius (Enorthocladius) species 2, Coffman & Ferrington, 1984: figs. 25.394, 25.395 [pupa].

Type Locality: USA Alaska, Portage.

Type Material: Holotype. Male pupa, USA, Alaska, Portage Glacial Pool, 20-VII-1977, #23, D. Wartinbee.

Specimen dissected and parts placed in Canada balsam under 6 coverslips on a slide. Deposited in the FSCA. Para-

types (10). Canada. Alberta, Waterton Park, 21-VII-1967, A. L. Hamilton, A.3.1., 4 Ex (FWI). USA. Colorado,

Gunnison Co., Beaver Dam on East R. 3.1 mi. N. of Gothic, 13-VII-1982, L. Ferrington No.-Co. #19, 3 Ex

(KSBS). Idaho, East Fork Salmon River, 11-IV-1977, #PE 122, J. Sedell, 2 Ex (COFF). Montana, Beartooth-Absa-

roka Wilderness Area, 31-VII-1979, E.A. Wells, CH6965.1, 1FP (CNC).

Diagnosıs:

Orthocladius coffmani can be distinguished from other Holarctic species of O. (Enorthocladius) by

a combination of characters. Adult Male: palpal segment 3>4, multiserial scutellars, and details of the

hypopygium (Fig. 9). Pupa: hooklets on II, large round patches of spines on III-VII, seta on anal

lobe. Larva: Unkown.

Derivation of Name: This species is named after William P. Coffman, for providing associated material of this

species and exuviae of other Orthocladius over the years.

Description

Adult Male (n=1)

Brown. Medium species. Head. Verticals 12, postorbitals 1. Palps long with 3>4. AR cannot be de-

termined. Thorax. Lateral antepronotals 3. Acrostichals absent. Dorsocentrals 15. Prealars 5. Scutel-

lars 16, multiserial. Wing. Squamals 20. Other characters cannot be determined. Legs. Measurements

cannot be determined. Hypopygium (Fig. 9). Virga absent. Superior volsella collar-like. Inferior vol-

sella with dorsal part squared and ventral part protruding like dorsal part, appearing double-lobed.

Anal point weak with pointed apex. Crista dorsalis robust.

Variation. The virga is not visible, but it may be concealed in a mass of tissue.

Pupa (Exuviae)

Brown with darker apophyses on all tergites, and darker shading on cephalothorax and anal lobe.

Length 3.7—4.8 mm (3). Cephalothorax. Frontal warts absent; cephalic tubercles weak; protuberan-

ces between bases of antennal sheaths below cephalic tubercles. Precorneals clumped, almost 3x as

long as dorsocentrals; 1 median antepronotal, weak, O lateral antepronotals; 4 dorsocentrals, reduced

spaced in arow. Thoracic horn (Fig 34a) long, tubular, bubbled, light brown, length 220-310 um.

Thorax dorsally wrinkled along eclosion line.

Abdomen (Fig. 39). Tergites: I bare; II with large patch of recurved hooklets in about 15 rows; re-

duced patch of hooklets on IIT-V; IIT-VII with large cırcular patch of posteriorly-directed spines

along anterior margin; patches of spinules on II—-VIII. Sternites: I, IV, V bare, or at most with some

spinules anteriorly; II, III with central patch of spinules along anterior margin; VI-VII with 2 off-

center patches of spinules anteriorly.

Setae on segments I- VII:

D 4

ar or) DRS IG ie 713

V NE ZN) 303 1

5 I ID

3 1 xl 0 @ Oh lgelereluhl

Anal lobe strongly developed into large, circular lobes; usually margin is smooth, at most margın is

wrinkled along distal ?/;; 1 ventral robust seta, on distal half. Pedes spurii A absent; pedes spurii Bon

II, weak. Genital sheaths extended beyond lobes in male, not in female.

Variation. Females appear larger than males.

Biology: Exuviae were collected from cold waters of mountain rivers and glacial pools. Adults

emerge in April and July.

Distribution: Nearctic: Canada (Alberta); USA (Alaska, Colorado, Idaho, Montana)

Remarks: This species is a member of the rivulorum-group. The weak and pointed anal point is a distinguishing

feature of the male, found only in one other species of O. (Enorthocladius), abiskoensis. The hypopygium resembles

that of O. (Orthocladius) more than that of O. (Enorthocladins). The pupa may be confused with rıvulorum, but

the spine patches on tergites III-VIII are much larger in coffmanı. In Coffman & Ferrington (1984), this species

will key to couplet 45. The female and larva remain unknown.

Orthocladius (Euorthocladius) difficilis (Lundbeck)

Fig. 31

Chironomus dıfficilis Lundbeck 1898: 282 [description of male, female].

Orthocladius difficilis Lundbeck. Kertesz, 1902: 217 [catalogue]. Johannsen, 1905: 267, 277 [in adult key;

description of male, female].

Orthocladius (Orthocladius) difficilis Lundbeck. Andersen, 1937: 63 [in adult key].

Orthocladius difficılis (Lundbeck). Sublette & Sublette, 1965: 156 [checklist].

Orthocladius (Euorthocladius) difficilis (Lundbeck). Oliver, 1970: 103, 104, figs. 4-6 [designation and description

of lectotype male].

Type Locality: Greenland; Kangersuak.

Type Material: Lectotype: Adult Male, Greenland, Kangersuak, 22/9/1890. Mounted in balsam under 5 cover-

slips on slide (UCOP). Typed label Chironomus difficilis Ldbk. Written LECTOTYPE, D. R. Oliver 1969. Red

label with Type written; also date written.

Diagnosıs

The male of Orthocladius difficilis can be distinguished from other males of O. (Enorthocladius) by

the low AR and details of the hypopygium (Fig. 31). The pupa and larva are unknown.

Derivation of Name: L. difficılis, not easy, troublesome.

Description: See Oliver (1970).

Biology: Unknown.

Distribution: Nearctic: Greenland.

Material Examined: Lectotype.

Remarks: This species was reported in the literature primarily as occurring in Greenland, and was not identifiable

until Oliver’s (1970) redescription, where he also designated a lectotype and provided a figure of the hypopygium.

The male is a typical O. (Enorthocladius) with female-like eyes, biserial scutellars, crista dorsalis on gonostylus,

and collar-like superior volsella. Acrostichals are absent.

This species is part of the rivicola-group, and the male can be distinguished from thienemanni by the lower AR

(1.24 in difficılis, >1.80 in thienemanni) and from rivicola by salient features of the hypopygium. Both rivicola and

thienemanni occur in Greenland, and specimens from Greenland were examined: a single male of rivicola (acrosti-

chals present, AR=1.35) and exuviae of thienemanni.

The immature stages of difficılis are unknown. Rearings of difficilis from southwest Greenland are needed to bet-

ter understand the species.

Orthocladius (Euorthocladius) kanii (Tokunaga)

PiE722

Spaniotoma (Orthocladius) kanıi Tokunaga, 1939: 315—318, figs. 13, 36, 53, 54, 68, 76, 86, 91, 104, 114, 121, 131,

142, 157 [description of male, female, pupa, and larva]. Tokunaga, 1959; 1973: 641 [pupa, larva, fide Sasa & Ya-

mamoto, 1977]. Thienemann, 1954: 345 [note].

”Spaniotoma (Orthocladius)“ kanıı Tokunaga. Thienemann, 1944: 567, 649 [in pupal, larval keys].

Orthocladius (sen. str.) Ranıi Tokunaga. Tokunaga, 1964: 17, fig. 4 [male, female].

Orthocladius kanıi (Tokunaga, 1939). Sasa & Yamamoto, 1977: 310 [checklist].

Orthocladius (Euorthocladius) kanii (Tokunaga, 1939). Sasa, 1979: 26—28, figs. 40—43 [description of male, female,

pupa, and larva]. Sasa, 1981:87 [survey record].

Type Locality: Japan: Kyoto.

Type Material: Holotype. Male, Japan, Kyoto, Nishigamo, Jan 1936, M. Tokunaga (2 white labels, printed). Ab-

domen from segment Il to hypopygium mounted on a slide in Canada balsam under one coverslip by A.R. Soponis.

Paratypes (?8). Same data as holotype, parts of males and females mounted in Canada balsam on 4 slides. Slide 1:

hypopygium with abdominal segments VII, VII. Slide 2: hypopygium with abdominal segment VII. Slide 3: 4

coverslips, with 4 thoraces, 3 thoraces, 8 heads (4M, 4F), and parts of antennae, legs, thoraces. All type materials

are retained in the Entomological Laboratory of Kyushu University. The only locality data in the vials were those

of the holotype, and paratypes were assumed to have the same data. Paratypes from other dates and localities were

not located.

Diagnosıs

Orthocladius kanıi can be distinguished from other species of ©. (Euorthocladius) by acombination

of characters. Adult Male: details of the hypopygium (Fig. 22); distinguishable from /uteipes by distri-

bution. Pupa: tergite II bare; rows of spines on margins of tergites IV VIII; distinguishable from r:-

vicola by distribution. Larva: mentum with 13 teeth, MR>1.5, low AR (<1.85); head capsule brown;

distinguishable from /uteipes by distribution.

Derivation of Name: This species was named for Mr. T. Kani, who collected the type specimens with M. Toku-

naga (Tokunaga 1939: 318).

Description: See Tokunaga (1939) and Sasa (1979).

Biology: The larvae are widely distributed in torrential streams throughout Japan (Sasa, 1979: 28).

The larvae live in oval, clear, gelatinous cases, 8-10 mm long, 4-5 mm wide, and 3-5 mm high, often

covered with diatoms, and closely adhering to surfaces of stones on these streams (Tokunaga, 1939).

Distribution: Japan.

Material Examined: Type material.

Remarks: Tokunaga (1939) described this species from an unspecified number of males, females, pupae, and lar-

vae collected in torrential streams in the suburbs of Kyoto, Japan. Sasa (1979) gave another detailed description of

the species in all stages.

Orthocladius kanıi is morphologically similar to /uteipes in the adult male and larva. However the pupae are dif-

ferent: weaker spines occur in rows on IV—VIIl in /uteipes, whereas stronger spines occur in rows on IV- VIII in

kanıi; pedes spurii A are present on sternites V-VII in /uteipes, and entirely absent in kanii. There was no oppor-

tunity to examine pupal material of kanıi; however, detailed illustrations of the pupal abdomen (Sasa 1979, fig. 42 b)

and the pupal sternite VII (Tokunaga 1939, fig. 76) show no pedes spurii A. It is unlikely that both authors would

have missed these prominent structures in their drawings. In addition, the tergal spine patterns of kanıı more closely

resemble those of rivicola than of Iuteipes. Thienemann (1954) remarked that he knew of no European parallel of

kanii. f

The extant type series, which contains only parts of male and female adults, was slide-mounted and examined.

Variation of the hypopygium can be seen here (Fig. 22), and in Sasa (1979, fig. 41), Tokunaga (1939, fig. 36), and

Tokunaga (1964, fig. 4). The figure in Tokunaga (1964) looks distorted and the specimens have amuch higher AR

(1.5—2.2) than that recorded for the species (1.4—1.7).

Orthocladius (Euorthocladius) luteipes Goetghebuer

Figs. 1, 23, 24, 37b, 46, 47, 53

Orthocladius luteipes Goetghebuer, 1938: 457, 458, fig. 6 [description of male]. Reiss, 1983: 176 [checklist]. Ros-

saro, 1984: table 2 [record].

Euorthocladius Iuteipes Goetghebuer. Thienemann, 1939: 6, 7 fig. 2b [description of pupa, larva]. Thienemann,

1944: 559, 648, fig. 14b [in pupal, larval key]. Thienemann, 1954: 347 [ecology]].

Orthocladius (Orthocladins) Inteipes Goetghebuer. Goetghebuer, 942: 34, 49, fig. 75 [description of male; in key

to males].

“Orthocladius” Iuteipes G. Fittkau et al., 1967: 363 [checklist]. Fittkau & Reiss, 1978: 422 [checklist].

Orthocladius (Euorthocladius) luteipes (Goetghebuer). Rossaro, 1978b: 184, 185 fig. 1 [record, notes on males and

species]. Langton, 1984: 144, fig. 49b [in pupal key]. Sahin, 1984: 81, figs. 203—205 [in larval key].

Orthocladius (Euorthocladius) Type Ill sp. Simpson & Bode, 1980: 13, 52 [partim] [larval description, photograph,

in larval key].

Orthocladius (Euorthocladius) cf. Inteipes. Coffman, 1973: Table 1 [ecology].

Orthocladius (Euorthocladius) species 4 Coffman & Ferrington, 1984: figs. 25.410, 25.411 [pupa].

Type Localıty: Austria.

Type Material: Holotype. Male, Basse-Autriche, Dr. Mitis, TN18 (1938) (2 original ink labels of Goetghebuer),

R.1.Sc. N. B. 18.073. Mounted on slide in Canada balsam under 2 coverslips by A. R. Soponis; in poor condition,

body still pressed between celluloid.

23)

Diagnosıs

Orthocladius Iuteipes can be distinguished from other Holarctic species of O. (Enorthocladius) by

a combination of characters. Adult Male: details of the hypopygium (Figs. 23, 24). Pupa: Pedes spurii

A on sternites V-VIII; tergites IV-VIII with rows of slender spines on posterior margins. Larva:

mentum with 13 teeth, MR>1.5, AR<1.85 (separable from Kanii by distribution).

Derivation of Name: L. luteus, yellow; most likely refers to the color of the adult male as described by Goetghe-

buer (1938).

Description

Adult Male (n=3)

Dark brown (OD: yellow with black bands, abdomen brownish). Medium species. Head. Verticals

16-20 (2), postorbitals 2-3 (2). Palps normal with 3>4. AR 2.00 (1) (OD: 2.04). Thorax. Lateral an-

tepronotals 2-5. Acrostichals 0-4, robust, begin within 2 AW. Dorsocentrals 6-11. Prealars 6-8

(2). Scutellars 15-18, biserial or multiserial. Wing. Length 2.15—2.58 mm (2). (OD: 2.15). R with

7—11 setae. Squamals 24—31. Anal lobe cannot be determined in my material (OD: produced). Legs.

LR1 0.69 (1). LR2 0.48 (1). LR3 0.53 (1). (OD: LR1 0.78). Sensilla chaetica on tal of p2 (7-11) (2).

Hypopygium (Figs. 23, 24). Virga present. Superior volsella collar-like. Inferior volsella with dorsal

part squared, covering ventral part. Crista dorsalis long.

Variation. The acrostichals are missing on the type, and a single male specimen has sockets but no setae. Acrosti-

chals are present on the male pupa. Rossaro (1978b) reported that /uteipes has an average AR of 1.6 and an average

wing length of 2.8 mm.

Pupa (Exuviae)

Light brown to brown with darker apophyses on I-VI, variable; some parts of conjunctives V-VI

and VII-VIII darker. Length 3.50-5.00 mm (20). Cephalothorax. Frontal warts weak and cephalic

tubercles absent. Precorneals weak, 1'/,x as long as dorsocentrals; 2 median antepronotals, 1 lateral

antepronotal, weak to strong; 3 dorsocentrals, robust, thicker than procorneals. Thoracic horn ellip-

soid, light brown, filled or clear, stalked but not often seen; length 80— 110 um (20). Dorsum of thorax

usually smooth, may be wrinkled or sculptured, expecially along eclosion line with granular pattern

mesad of wing base.

Abdomen (Figs. 46, 47). Tergites: I-III bare; IV-VII with rows of straight, slender spines along

posterior margin (Fig. 37b); V-VIII with central patch of spinules, more extensive on each suc-

ceeding tergite; VII, VIII with spinules along anterior margin. Sternites: I bare; II-V with spinules

on anterior half; VI-VIII with 2 off-center patches of spinules anteriorly.

Setae on segments I— VIII:

De ee JE I MR 109RS 1000. a SIR € Dagre Was

VE WO Or uieet 1 7

Anal lobe greatly reduced, setae absent. Genital sheaths extended beyond lobe in male and female. Pe-

des spurii B absent; pedes spurii A robust, on V- VII.

Variation. Tergites may have sculpturing; pedes spurii A may be absent on V, VII, always present on VI. Setae

are weak (Germany) to strong (Italy). Shagreen may be a small to large patch (Figs. 46, 47). Specimens are small

(North America) to large (Europe). European specimens have more spines in spine patch IV-VIII than North

American specimens. North American specimens have more dark apophyses than European. Posterior spine rows

are absent from IV in one specimen from Germany.

Larva (Fourth Instar)

Body yellow. Head capsule brown. Eye spots bipartite or fused. Mentum (Fig. 53c) with 13 teeth,

median tooth about 2x as wide as Ist lateral; MR 1.6—2.4 (6); median tooth as high as 1st lateral. Ven-

tromental plates extended anteriorly between 1st and 2nd laterals. Premandible simple, with notched

apex. Chaetula laterales sparse. Mandible (Fig. 53a) with apıcal tooth as long or slightly longer than

1st inner tooth; outer margin notched opposite seta subdentalis, rest of margin smooth except for

occasional notch posteriorly; seta interna present. Antenna (Fig. 53b) with robust Lauterborn organs;

blade extended to 4th segment. AR 1.50-1.88 (6). Body with simple setae, most likely arranged like

that of saxosus. Anal tubules long, rounded, with dorsal pair shorter and thicker than ventral pair.

Biology: This species has been collected from rivers and creeks. It occurs with rivicola in habitats

in Oregon (South Santiam River) and Pennsylvania (Delaware River, Big Bushkill Creek, and Lines-

ville Creek). Thienemann (1939, 1954) reported free-living larvae and pupae in gelatinous cases on

stones. Adults emerge in April in Pennsylvanıa (Coffman, 1973), and generally from February

through May.

Distribution: Palearctic: Austria, Germany, Italy, Turkey. Nearctic: USA (Georgia, New York,

North Carolina, Oregon, Pennsylvanıa).

Material Examined: Type material. Non-type Material: Germany: Lunzer — 1940, 2-IV-40, A. Thienemann, 25 Ex; Par-

tenkirchen 1933 No. 72, A. Thienemann, IL, Mitis No. 19, 3L (ZSM). Italy: t. Ticiur, 26-3-1973, Rossaro, IM; Brembo 2,

1-III-1980, Rossaro, 2Ex; T. Piovena, 3-II-1976, Rossaro, 1MP (ROSS). USA: Georgia, Fanın Co., Noontootla Cr. at

Newport Rd., IV-24-1979, B. A. Caldwell, IL (CALD). New York, Lewis Co., Black River nr. Port Leyden, 7-VII-1976,

1L; Niagara Co., NiagaraR. nr. Youngstown, 6 Oct. 1976, 5L (NYSH). North Carolina, Iredell Co., Buffalo Shoals Creek,

Jan 1981, K. Dechart, I1L(NCNR). Oregon, South Santiam River, 18 May 1977, PE462, W. P. Coffman, 1Ex (COFF); Echo

Creek, 3 Oct. 1978, W. P. Coffman, #25, 4Ex (COFF). Pennsylvanıa, Monroe Co., Delaware River, n. of Party’s Beach,

15 April 1976, #1, D. Wartinbee, 3Ex (COFF); same data except 9 May 1976, #3, 9Ex (COFF); Lineville Creek, 13-IV-

1971, W. P. Coffman, 2Ex (CNC). Monroe Co., Big Bushkill Creek, Resica Falls, 17-IV-1976, 3Ex (COFF).

Remarks: Goetghebuer (1938) described this species in Orthocladıns from a single male collected ın Austria. The

figure ofthe hypopygium, with ahaired, pointed anal point and squared inferior volsella, resembles males of Ortho-

cladıns (Orthocladins). Goetghebuer (1942) later reproduced the figure and description, and placed /uteipes in his

heterogeneous O. (Orthocladins). He separated /uteipes from similar species of the subgenus by the VR and exten-

sion of the R4+5 and Costa, characters that are usually too variable for species determination in Orthocladius (So-

ponis 1977). However, having to use these characters showed the difficulty of separating the adult male of this spe-

cies, as does the key given here.

Not only is the male difficult to ıdentify, but the type was virtually impossible to view because it was melted by

Goetghebuer between 2 pieces of celluloid. Although the celluloid is not completely dissolved, the type can now

be examined clearly, but it is in poor condition. The type does agree with Goetghebuer’s (1938, 1942) ıllustrations.

Thienemann (1939) described the larva and pupa of /uteipes, and distinguished them from rivicola, a species with

which it ıs still confused today. Thienemann distinguished the pupa of /uteipes from rıvicola by the more slender

spines of the more numerous rows on tergites IV— VIII, and the larger thoracic horn. In the material examined here,

the thoracic horn of Iuteipes (80-110 um, n=20) is larger than that of rivicola (45-70 um, n=20). It is easiest to

identify /uteipes by the pedes spurii A on sternites V- VII, which Thienemann described but did not use in his key.

Later, Thienemann (1944) included /uteipes in his larval and pupal keys.

Simpson and Bode (1980), in their diagnosis of ©. (Enorthocladius) type Ill sp. larva, provided a photograph of

alarva of /uteipes with MR>1.5 and AR of 1.73. Examination of their material showed that rivicola was also present.

Easily identifiable only in the pupal stage, /uteipes has not been recorded frequently in the literature. Fittkau et

al. (1967) and Fittkau & Reiss (1978) recorded /uteipes in Limnofauna Europeae as an uncertain ”Orthocladins”.

Rossaro (1978b) illustrated the hypopygium and provided notes on /uteipes. In Coffman & Ferrington (1984), the

pupa will key to couplet 55.

Orthocladins luteipes belongs to the rivicola-group and is most easily determined by the exuviae. Males will be

difficult to determine without associated exuviae. Altough the type and the male from Italy have a high AR (2.00,

2.04), Rossaro (1978b) reported an average AR of 1.6 for /uteipes. The pupae are easily separable from rıvicola with

the characters provided, and become distinctive when many specimens of both species are examined. The larvae are

close to thienemanni, and Inteipes can be distinguished by the wide median tooth (high MR) and lower AR.

This species occurs with other O. (Euorthocladins), including rıvicola, and has probably been misidentified as rz-

vicola (pupa) and thienemanni (larva). Undoubtedly /uteipes occurs more widely than documented here, although

perhaps not as widely as rivicola, for example.

Orthocladius (Euorthocladius) rivicola Kieffer

Figs. 32, 33, 348, 37, 49, 55

Orthocladius rivicola Kieffer, 1911: 181 [original description; adult in key]. Thienemann, 1911:637 [notes on pupa,

locality data]. Thienemann, 1912: 74 [notes]. Potthast, 1914: 264, figs. 6-9 [pupa, larva]. Goetghebuer, 1932: 74,

88 [female, ın key to females]. Thienemann, 1935: 203—205 [in pupal, larval key; synonymy]. Pankratova, 1970:

173, 174, 178, fig. 106 [in pupal, larval keys; pupal, larval descriptions]. Reiss, 1983: 176 [checklist]. Rossaro,

1984: table 2 [record]. Bitusik & Ertlova, 1985: 603, 606, table 2 [ecology]].

Orthocladius (Chaetocladius) rivicola Kieffer. Goetghebuer, 1934: 89, 90, fig. 4 [male description].

Euorthocladius rivicola (Kieffer). Thienemann, 1936: 191 [record]. Thienemann, 1939: 7, fig. 2a [pupa]. Thiene-

mann, 1941: 65, 68, 78, 79, 82, 153, 180 [ecology, distribution]. Thienemann, 1944: 559, 648, fig. 13, 14a, 195 [in

pupal, larval keys]. Dittmar, 1955: 470, 481, 482, 484, table 30 [ecology]. Romaniszyn, 1958: 82 [in larval key].

Thienemann, 1954: 23, 31, 48, 49, 288, 301, 303, 333, 346, 347, 349, 355, 357, fig. 133.

Orthocladins (Orthocladius) rıvicola Kieffer. Goetghebuer, 1942: 32, 53, fig. 87 [male description, in male key].

Orthocladius ex gp. rivicola Kieff. Chernovskii, 1949: 205, 282 [in larval key, synonymy].

Orthocladins (Euorthocladius) rivicola Kieffer. Brundin, 1956: 101 [record]. Fittkau et al., 1967: 362 [checklist].

Saether, 1968: 463 [ecology ]. Saether, 1969: 61 [record]. Lehmann, 1971: 486 [ecology]. Kloet & Hincks, 1975:

(V)15 [checklist]. Rossaro, 1977: 122 [notes]. Rossaro, 1978a: 290, table 1 [distribution]. Rossaro, 1978b: 185

[distribution]. Säwedal, 1978: 87 [record]. Fittkau & Reiss, 1978: 421 [checklist]. Prat, 1979: 67, 68, fig. 19 [male

description]. Kownacki & Zosidze, 1980: 75, 79—81, table 2 [ecology]. Halvorsen et al., 1982: 119 [record]. Ros-

saro, 1982: 42—44 [in pupal, larval keys]. Mason & Lehmkuhl, 183: 207, fig. 19 [ecology]. Murray & Ashe, 1983:

230 [checklist]. Mason & Lehmkuhl, 1985: 878, table 1 [distribution]. Caspers & Schleuter, 1986: 323 [checklist].

Orthocadius (Enorthocladius) rivicola «. Rossaro, 1982: fig 31 [pupa].

Orthocladius (Enorthocladius) Thienemann type I. Soponis, 1977: 15-17, figs. 84c, 90, 101, 120 [pupal, larval de-

scription; in pupal, larval keys].

Orthocladius (Euorthocladius) species 6 [partim]. Coffman & Ferrington, 1984: fig. 25.415 [pupa].

Orthocladius (Euorthocladius) cf. thienemanni-saxosus |partim]. Coffman, 1973: table 1 [ecology].

Orthocladius fusiformis Goetghebuer. Goetghebuer and Dorier, 1939: 30-32, fig. 1-5.

Type Locality: Germany.

Type Material: Could not be located, believed lost.

Diagnosis

Orthocladins rivicola can be distinguished from other Holarctic species of O. (Euorthocladius) by

a combination of characters. Adult Male: lower AR (0.08- 1.76), sensilla chaetica on midleg, and de-

tails of the hypopygium (Figs. 32, 33). Pupa: absence of pedes spurii A and hooklets on tergite II, pre-

sence of posterior spine rows on tergites IV— VIII, and normally developed dorsocentral setae; can be

distinguished from kanıi by distribution. Larva: mentum with 13 teeth, MR<1.5, AR<1.8; cannot be

distinguished from asbhei.

Derivation of Name: L. rıvus, stream; L. cola, dweller, inhabitant.

Description

Adult Male (n = 27)

Small to medium species. Head. Verticals 9—20 (26), postorbitals 1-2 (21). Palps long with 3>4.

AR. 1.00-1.76 (82). Thorax. Lateral antepronotals 1—9 (26). Acrostichals 0-10, weak, begin within

l1or more AW. Dorsocentrals 7-16. Prealars 4-7. Scutellars 8-26, often multiserial, also biserial, or

multiserial. Wing. Length 1.30—2.80 mm (81). R with 2-9 setae. Squamals 10-36. VR 1.02-1.16

(25). Anal lobe slightly produced. Legs. LR1 0.61—0.75 (26). LR2 0.43—0.54 (26). LR3 0.47—0.59

(26). Sensilla chaetica (25) on tal of p2 (3-16). In addition, 3 specimens have sensilla chaetica on tal

of p3, 2 (25). Hypopygium (Figs. 32, 33). Virga present or absent. Superior volsella collar-like or

sightly triangular. Inferior volsella with dorsal part squared or rounded, and ventral part slightly ex-

tended below. Crista dorsalis long, robust.

Variation. The length of the terminal flagellomere is highly positively correlated with wing length in rivicola

(r=+0.943, p>.001, n=80) as it is in most Orthocladius (Soponis 1977). However, wing length is not as highly cor-

related with either the length of the basal flagellomeres 1-12 (r=+0.747, p>.001, n=80) or with AR (r=+0.753,

p>.001, n=80). Prat (1979) found a positive correlation between AR and wing length (WL 2.24 mm, AR 1.25; WL

3.00 mm, AR 1.6 to <1.8), and these were related to geographical locality. However, correlations with rations

should be interpreted carefully and are not usually biologically meaningful (Soponis 1977).

In the 80 specimens measured, no obvious relatinship exists between wing length and locality. Considerable va-

riation was found between specimens collected at the same locality, e.g. Trails Pond, Idaho, and at a generalllocality,

e.g. Alaska. But little variation was also found between specimens collected at the same locality, Baffin Island.

The palps appear longer in some specimens, and for a small sample there is a fairly high positive correlation be-

tween interocular distance and total palpal length (r=+0.810, p>.001, n=14). In rzvicola, at least, males with eyes

more wıdely separated tend to have longer palps. Again, there was no apparent relation between locality and either

of these two measurements.

Values from other studies of rıvıcola fall within the variation recorded in this material. AR: 1.40—1.50 (Rossaro

1978b); 1.50 (Lehmann 1971); 1.25—<1.80 (Prat 1979). Lower values of AR have been reported for high altitude

specimens: 0.80 at 1100 m (Goetghebuer 1934) and 0.90, altitude presumably high (Rossaro 1978b). Rossaro

(1978b) also reported 4—11 sensilla chaetica on tal of p2.

The hypopygium is highly variable, as seen here (Figs 32, 33), in Prat (1979), and in Goetghebuer (1934). In ma-

ture pupae (n=14), acrostichals were present or absent, and scutellars were uniserial, biserial, or multiserial.

Pupa (Exuviae)

Brown to pale brown, cephalothorax darker. Length 2.5—4.0 mm. Cephalothorax. Frontal warts

and cephalıic tubercles absent. Precorneals clumped, Pc1 usually longer and thicker than Pc2 and Pc3;

2 median antepronotals, 1 lateral antepronotal, 3 dorsocentrals, strong to weak, about as long as Pc1.

Thoracic horn (Fig. 34g) small, ellipsoid, dark brown, filled or clear, stalked; length 45-70 um (20).

Thorax dorsally wrinkled along eclosion line.

Abdomen (Fig. 49). Tergites: I bare; IV-VIII with central rows of moveable straight spines along

posterior margin (Fig. 37 c); II-VIII with small patches of spinules anteriorly; IV—VIII with central

patch of spinules anteriorly. Sternites: I bare; II-VIII with central patch of spinules anteriorly.

Setae on segments I—- VIII:

Den 4.4 5 1 IL I 39.533 3

9423 0) Odessa 0. 02, 92 009

SR Wer Non 12) 2

V 2 2402. 490,8 0)

Anal lobe greatly reduced, with two seta, one on distal half and one at midpoint. Genital sheaths

extended beyond lobe in males and females. Pedes Spurii A and B absent.

Variation. There is considerable variation in this widespread species. Spines in rows IV— VIII may be individually

weak to robust; these spines may be in 1-5 rows on the tergites; spines may be spaced close together or far apart.

The number of spines in the rows is higher in females than in males, but is not significantly different (t-test,

p>.001”).

rıvicola females (n = 5) rıvicola males (n = 5) Student’s t*

IV 80.20 # 21.19 (54-104) 48.00 + 18.51 (36-80) 2.5590

V 79.60 + 18.61 (60-100) 59.00 + 19.87 (47-94) 1.6918

VI 69.80 + 25.29 (44-103) 54.40 + 13.99 (41-76) 1.19135

VII 57.40 + 20.84 (39-90) 44.00 + 15.95 (33-72) 1.1417

VIII 41.20 + 19.80 (23-70) 29.60 + 10.83 (20-47) 1.1491

Lehmann (1971) found two distinct types of rivicola pupae in the Fulda. One type had 2 or more rows of spines

on the posterior margins of tergites IV VIII, the other had only a single row of spines. In the Plön (now ZSM) col-

lection he found many transitional forms between his two basic types, and he attributed these differences to intra-

specific varıation. It is possible that ashei was confused with rivicola, and ashei was the species with the single row

of spines.

27,

Setae on the abdomen are generally robust and easy to see in this species. Dorsocentrals can be forked or branch-

ed, weak to robust, but never as robust as in ashei. The thoracic horn can be filled or clear, small or large. The size

of the thoracic horn is unreliable for distinguishing rivicola from thienemanni or ashei. However, all Iuteipes have

larger thoracic horns than ashei, rivicola, and thienemanni.

There is considerable variation in the exuviae of rivicola. Several undescribed species may exist, including one

from northwestern North America with individually robust spines on IV—- VII.

Larva (Fourth Instar)

Body yellow or brown. Head capsule brown. Eye spots fused. Mentum (Fig. 55d) with 13 teeth,

median tooth about as wide as Ist lateral; MR 1.00-1.50 (7); median tooth as high or lower than Ist

lateral. Ventromental plates extended anteriorly between 2nd and 3rd laterals. Epipharynx (Fig. 55a)

with premandible simple. Chaetula laterales sparse. Mandible (Fig. 55b) with apical tooth as long or

slightly longer than Ist inner tooth; outer margin notched opposite seta subdentalis, rest of margin

smooth except for occasional notch posteriorly; seta interna present. Antenna (Fig. 55c) with robust

Lauterborn organs; blade extended to 5th segment. AR 1.38— 1.80 (7). Body with simple setae, some

short and stiff, some long and curved, and arranged like saxosus. Anal tubules moderately long, round-

ed, with dorsal pair thicker than ventral pair.

Variation. The variation of the larvae is underestimated in this material. Consequently, larvae associated with pu-

pae will be most accurately determined.

Biology: The larvae live on stones in currents of springs, brooks, streams, and rivers (Thienemann

1935, 1941, 1954; Dittmar 1955; Lehmann 1971). Larval and pupal tubes are similar to those of thiene-

manni (Fig. 40). Larvae usually live in individual gelatinous tubes covered with sand grains and detri-

tus, cemented along their lengths to the stone. Pupae usually live in individual, clear, half-ellipsoid, ge-

latinous tubes with holes at both ends for the current. Larvae are rheobionts and eurytherms (Thiene-

mann 1912, fide Dittmar 1955). Illies (1952, fide Dittmar 1955) found larves of rivicola in the mud, but

such finds may be accidental for rheobionts like rivicola. Thienemann (1941, 1954) observed large

numbers of free-living mature larvae and prepupae in submerged reeds (Hydrurus) collected in Swit-

zerland.

Based on exuviae, rivicola occurs with thienemanni in at least 7 different sites: Ottawa River, On-

tarıo, Canada; River Fulda, Germany; Linesville Creek, Pennsylvania; East Fork of Chattoga River,

South Carolina; Fall Creek, South Carolina; Seneca Creek, South Carolina; and, Pigeon River, Ten-

nessee. Bitüsik and Ertlovä (1985) found rivicola and thienemanni in the River Rajcıanka. They con-

cluded that rivicola occurs in small numbers in every lotic zone studied, but thienemanni is largely

confined to the 2. zone of high diatom density, occurring in high numbers. Kownacki & Zosidze

(1980) found rivicola dominant in certain zones of rivers and streams of the Little Caucasus Moun-

tains.

The adults emerge from November to May in Italy, and can be found at higher altitudes in July and

August (Rossaro 1977, 1978a, b). Dittmar (1955) found adults in Germany from January to April.

Thienemann (1954) found two generations in the Lunzer Gebiet: at the beginning of June, and from

August to October. Lehmann (1971) recorded two emerging generations from the Fulda in Germany;

the first from March to May/June, and the second in October/November. In Canada, Mason and

Lehmkuhl (1938) found rivicola emerging from April to October, with only slight differences be-

tween emergence upstream and downstream of a hydroelectric development. In South Carolina,

adults emerge from December to May.

This species also occurs at high altitudes and high latitudes (Thienemann 1936, 1941, 1954; Rossaro

1978b; Goetghebuer 1934).

Mites have been associated with rivicola (Thienemann 1954).

Distribution: Palearctic: England, France, Germany, Ireland, Italy, Poland, Sweden, Switzerland.

Nearctic: Canada, Greenland, USA.

Material Examined: Non-type material: Austria. Lunz, Thienemann, 2M, 3Ex (ZSM). Bulgaria. Blagoevgrad, r. Bistriza,

28-II-1978, N. Natchev, 1Ex (NAT). Canada. Alberta: Waterton Natl. Park, 21-VII-1967, A. L. Hamilton, A.3.1, 4Ex

(FWI); A. L. Hamilton and ©. A. Saether, 1M, 1M w/Ex (FWI); Calgary, 4-VIII-1970, J. Martin, 1M (CNC). Manitoba:

Duck Mtns, South Duck, 14-V-1980, IM; 16-V-1980, 1M; Duck Mtns, Cowan G.,24-V-1980, 1M (FWT); Edwards Cr. Stn.

1, Riding Mtn. Natl. Pk., 50°59°15”, 100°04°00“, 4-VII-1975, 1M (CNC). Ontario (CNC): Ottawa, 21-X-1971,J. R. Dow-

nes, IM; Ottawa, Central Expmntl Farm, 28-X-1966, J. Martin, 4M; Ottawa, Britannıa Flitration Plant, 5-V-1971, D.R.

Oliver, IM; Ottawa, Ottawa R. Beach at Woodrufe, 9-V-1972, A. R. Soponis, 1M; OttawaRR. at Ottawa, 22-IV-1966, J.

Martin, IEx; Ottawa, OttawaR. at Rumic Rapids, drift, 16-IV-1985, P. S. Cranston, 13Ex; Green Creek, coll. 31-V-1979,

em. 2-VI-1967, DRO, LHS, RDM, 1F w/Ex. Quebec (CNC): Ile Ste. Helene, Montreal, 2-3-VII-1964, A. Nimmo, Shadfly

Project, 1M; 16-17-VI-1964, 1M. Northwest Territories (CNC): Oscar Creek, 25-V-1972, FWI Pipeline Project, 2FP,

5MP; Trail R., 11-VIII-1972, FWI Pipeline Project, 1MP; Ft. Laird, 60°15’N 123°28’W, 5-VI-1973, D. R. Oliver, IM; Ba-

thurst Is., J. Bissett, 4-VIII-1977, 2M; 5-VIII-1977, 2M; 6-VIII-1977, 3M; 12-VIII-1977, 5M; Bathurst Is., 75°24’N

100°24’W, 25-VII-1984, B. Hayes, 1M w/Ex; Martin R., FWI Pipeline Project, 10-VIII-1972, 3M; 18-VIII-1972, 1M; Ma-

sik R., Banks Is., W. R. Mason, 18-VII-1968, 1M; Head of Clyde Inlet, Baffın Is., 7-VIII-1958, G. E. Shewell, IM.

Yukon-Territory: Carıbou Bar Creek, FWI-Pipeline Project, 19-VI-1972, 4M; 2FP, 1MP; 20-VI-1972, 2MP, 3FP, 15M;

12-VII-1972, 1M; J. Robillard, 18-VI-1973, 1M w/Ex, LS; Driftwood River, 19-VII-1972, 1L (CNC); Little Bear Creek,

Mile 1022 Alaska Hwy, 3-VI-1978, D. R. Towns, 1M w/Ex (FSCA). Germany. Partenkirchen 125, 167, 94e, 1934, A. Thie-

nemann, 5Ex (ZSM); Fulda, 10-III-1964, E. J. Fittkau, IM, 1Ex, 1FP; Rhuhr, 8-V-11, bei Oloberg, A. Thienemann, 2MP,

1FP; Pullach, 26-11-1978, E. Ott, 1M w/Ex; River Isar, ca. 500 m oberhalb Loisach-Mündung, 3-IV-1986, F. Reiss, 5Ex

(ZSM). Greenland. Nedre Midsommer So, 10-VII-1966, Can. Pearyland Expd., IM (CNC). Italy. Po River, 1975, 1976, B.

Rossaro, Ex; Brembo, 1-IX-1980, B. Rossaro, 3Ex, 1MP (ROSS); 21-VII-1980, 2M (ROSS). Norway. Ekse, HOi: Vaksdal,

9-VII-1979, E. Willassen, 1 reared M; same data except 11-VI-1979, 2 reared M; same data except 9-VII-1979, 1 reared M

(ZMB). Sweden. Stordalen Sta., 7-VII-1958, D. R. Oliver, 3Ex (CNC); Lappland, 1936-1937, A. Thienemann, 2Fx, 1MP,

3FP (ZSM). Switzerland. Nationalpark Nadig, No 417, in Hydranus, A. Thienemann, 3FP, 1MP (ZSM). USA. Alaska:

K. M. Sommermann, jeep trap (USNM): Palmer, VI-1964, 4M; Kenai Pen., Johnson L.-Soldatna, 19-VI-1965, IM; Mata-

nuska Eklutna Hwy, 22-VI-1964, 4M; Palmer-Anchorage Hwy., 22-VI-1964, 3M; Anchorage-Potter-Mt. Alyeska, 21-IX-

1966, 2M; Anchorage-Eagle R.-L. Susitna R., 22-IX-1966, 7M; 24-IX-1964, 4M (USNM); Unnamed creek above Galbraith

Camp 208075, 9-VII-1976, drift net, USGS, 2L, 1Ex (CALD). Arkansas: Benton Co., Prairie Cr., NW !/,, Sec. 2. TI9N,

R29W, 4-I-1963,O. A. Hiteand L. K. Aggus, on bridge, 1M (SUB). Colorado: Delta Co., 1 mı. N Hotchkiss, 9-VIII-1971,

M. Beard, at light, 1M (SUB). Georgia: Fannin Co., Noontootla Creek at Newport Rd., 24-IV-1979, B. A. Caldwell, 2L,

1M w/Ex (CALD); Stekoa Creek at Wolf Creek Rd. (Savannah R. Drng.), 13-X-1973, E.P. D., IL(CALD). Idaho: Latah

Co., Trails Pond, 7-III-1969, J. M. Gillespie, found on ice and vegetation, 8M (MINN). Kansas: Kiowa Co., Rezeau Ranch,

spring fed creek, 19-III-1982, No. 31, B. G. Coler, J. K. Gelhaus, 17Ex (SBSK). Minnesota: Minneapolis, 4-VI-1969, D. E.

Maschwitz, at light, 1M; Cook Co., Min. F. $. Hovland, 9-VI-1969, E. F. Cook NJ Mosquito Trap, 1M; Ramsey Co., St.

Paul, 21-V-1968, R. A. Hellenthal, U of M vacuum trap, 1M; St. Louis Co., U of M Duluth, 6-VIII-1968, E. F. Cook, NJ

Mosquito Trap, 1M (MINN). Montana: Hamilton, outside lab bldg., 19-III-1960, C. B. Philip, 2M (SUB); Rock Creek,

17-VIII-1974, W. P. Coffman, 2Ex (COFF). New York: Erie Co., Cazenovia Creek at East Aurora, 80 m upstream Mill

Rd. bridge, multiplate sample, 29-VII-1976, K. W. Simpson, L (NYSH); Green Co., Gooseberry Creek nr. Tannersville,

coll. 10-VI-1978, em. 11-VI-1978, R. W. Bode, scraped from rock, IEx w/wing (NYSH); Niagara Co., Niagara R. nr.

Youngstown, 6-X-1976, L, 1Ex (NYSH); Rensselaer Co., Cropseyville, Route 2, Quackenkill Creek, 23-IV-1985, R. Bode,

3Ex (NYSH); St. Lawrence Co., St. Lawrence River nr. Waddington, 4-X-1977, L (NYSH); Ithaca, Apr., IM (SUB).

Tompkins Co., Ellis Hollow, 15-VI-1963, ©. ©. Berg, LT, 4M (USNM); Wash. Co., Hudson R. at Hudson Falls, 1km.

upstrm. Bakers Falls dam, 8-VI-1976, K. W. Simpson, L (NYSH). North Carolina: Orange Co., Little River, II-1979, D.

Lenat, S. Mozley, 1L(NCNR); Transylvanıa Co., Horsepasture R., 20-11-1976, P. Hudson, 2M (HUD); Wake Co., Cane

Creek, II-1980, D. Lenat, S. Mozley, 1L (NCNR). Oregon: Aumsville, 22-II-1963, K. Goeden, light, 3M (USNM). Penn-

sylvanıa: Crawford Co., Lindesville Creek, 7-IV-1971, W. P. Coffman, 4Ex (CNC); same date except 13-IV-1971, 5Ex

(CNC); stream nr. PA 285/179 btwn. Cochranton and Geneva, 28-V-1975, #4, W. P. Coffman, 2Ex (COFF); Shawnee,

stream, 4-IV-1976, #2, D. Wartinbee, 2Ex (COFF); Monroe Co., Big Bushkill Creek, Resica Falls, 17-IV-1976, 4Ex

(COFF). South Carolina (HUD): Oconee Co., East Fk. Chattooga R., Nat. Fish Hatchery, 28-I-1981, P. L. Hudson, 4M;

same data except 27-XII-1979, 1Mw/Ex; 15-II-1976, 1M w/Ex, IM; Oconnee Co., Seneca, Fall Creek, Lake Keowee, 30-

IV-1974, P. L. Hudson, 1Fx; same data except 10-X-1975, 1Ex; Oconee Co., Seneca Creek, 22-I-1976, 1FP, 2M, 1L; same

data except 10-11-1977, 1MP; 26-III-1977, 1MP; 1-V-1977, 1MP; Oconee Co., Salem, Horsepasture River, 20-II-1976, P.

L. Hudson, 3Ex; 18-XI-1977, 1M w/Ex; Pickens Co., Six Mile Creek, 4-II-1976, P. L. Hudson, IM. South Dakota: Clay

Co., Missouri R., Vermillion, 26-IV-1976, P. L. Hudson, channel, 1MP (HUD). Tennessee: Pigeon R., Gattlinburg, 3-V-

1977, P. L. Hudson, IEx (HUD). Virginia: Falls Church, Holmes River, 17-VI-1960, W. W. Wirth, lighttrap, IM (USNM).

Washington: Benton Co., Hanford, Columbia R., D. R. Oliver, 20-III-1952, 1M w/Ex; 27-III-1952, 2MP; 3-IV-1952, 1M

w/Ex (CNC); Yakıma Co., 17-18-XII-1971, B. J. Landis, ex yellow water traps, 1M (SUB).

Remarks: Kieffer (1911) described this species in a key to adults of Orthocladins. He distinguished rivicola by the

greenish-white abdomen, bare wings and veins, and cubitus not extended. He stated that the specimens were

collected in Germany and were obtained by rearing the immature stages, which would subsequently be described

by Thienemann. Kieffer did not mention the sex or number of specimens, nor did he designate a holotype. In cor-

respondence between Thienemann and Kieffer, Kieffer wrote Dactylocladus rivicola nsp O’P for specimens collect-

ed 8.IX.09 from Lenne bei Schmallenberg. This same material was cited by Thienemann (1911/12:74) as Orthocla-

dius rivicola Kieffer, so most likely this was the original material that Kieffer used to describe rivicola. A search of

the ZSM collection resulted in no specimens from this locality that could qualify for lectotype designation. Speci-

mens from other localities mentioned by Thienemann (1911/12:74) were also not found. Consequently a neotype

for rivicola will not be designated here because this is not an exceptional circumstance (Rules of ICZN, Article 75a),

and suitable (reared) material from the type locality in Germany does not exist.

Goetghebuer (1932) briefly described the color and wings of the female of rivicola, and included the species in a

key to females. Later Goetghebuer (1934) described the male of rivicola, and placed it in the subgenus Orthocladius

(Chaetocladius). He included characters of body length, wing variation, AR, and LR. The figure of the hypopygium

shows the dorsal lobe of the gonocoxite rounded and in a low position, which closely resembles a male from Idaho

(Fig. 33a). Goetghebuer noted the similarity of the males of rıvicola and thienemanni, and separated them by the

high AR of thienemanni (2.00). The AR of 0.80 is low for rıvicola, and this may be linked to high altitude. Later

Goetghebuer (1942) reproduced essentially the same figure and description of the male, and placed it in Orthocla-

dius (Orthocladius).

Prat (1979) provided the most recent figure of the hypopygium of rivicola, which he distinguished from the male

of thienemanni by the lower AR (1.50). Most Nearctic specimens have amore squared dorsal part of the basal lobe

of the gonocoxite (Fig. 32, 33b) than Prat’s figure.

The immature stages of rıvicola have been recorded more often than the adults. Potthast (1914) described and fi-

gured the larva and pupa, and included the distribution. Potthast could not separate the larva of rıvicola and thiene-

manni, but separated the pupae by differences in the spine rows and the larger thoracic horn of rivicola. In the ma-

terial examined here, both species have similarly-sized thoracic horns (30-60 um for rivicola, 30— 70 um for thie-

nemannı). -

Thienemann (1944) separated rivicola larvae from other Euorthocladius by the equally long anal tubules, low AR,

small body length, and distribution. Chernovskii (1949) distinguished the rivicola group from the thienemanni

group ın his larval keys by the lower AR (1.10 versus 2.00) and the smaller body size (5 versus 8 mm). Romaniszyn

(1958) uses AR and body length in his larval keys. Pankratova (1970) used Thienemann’s figures, and separated the

larvae of rivicola and thienemanni on the AR alone (1.40 versus 2.00) She also mentioned that the premandible was

bifurcate and showed a crenulated margin on the mandible, characters not seen in this material. Soponis (1977)

briefly described rıvicola as O. (Enorthocladius) Type land included it in subgeneric larval and pupal keys. Rossaro

(1982) provided larval keys but did not separate rivicola from Iuteipes.

Thienemann (1939) distinguished the pupae of /uteipes and rivicola by the stronger, shorter, darker, and denser

spines in the posterior spine rows and the smaller thoracic horn of rıvicola. Later, in his pupal keys, Thienemann

(1944) used these same characters but he could not distinguish the pupae of rivicola and fusiformis. An exuviae of

fusiformis determined by Dorier deposited in the ZSM collection was examined, and it is a good rivicola.

Pankratova (1970) did not distinguish between the pupae of thienemanni, rivicola, and saxosus.

Rossaro (1982) provided an excellent review of the Italian species in the subgenus with workable keys and figures

of the species. He distinguished two forms of the pupae of rivicola, called @ and 8. Some of this material has been

examined, and form « is rivicola, whereas form ß is the new species asheı.

Coffman and Ferrington (1984) included rivicola in their pupal keys; rivicola will key to couplet 55.

Orthocladius (Euorthocladius) rivulorum Kieffer

Figs 20, 21, 34b, 38a, 51

Orthocladius rivulorum Kieffer, 1909: 48 [adult description]. Kieffer & Thienemann, 1909: 32 [notes]. Potthast,

1914: 264— 266, figs 10— 14 [larval, pupal descriptions]. Thienemann, 1935: 203, 204, fig. 1 [in pupal, larval keys,

synonymy]. Pankratova, 1970: 173, 174, 178, 179, fig. 107 [larval, pupal descriptions]. Brennan et al., 1981: 149,

table 2, 4 [ecology]. Reiss, 1983: 176 [checklist]. Rossaro, 1984: table 2 [record].

Orthocladius (Dactylocladius) rıivulorum Kieffer. Goetghebuer, 1933: 215, 216, 218, figs 7, 7a [description of male,

female; in keys to males, females].

Spaniotoma (Orthocladius) rivulorum Kieffer. Johannsen, 1937: 56, 58, 62, 72, fig. 240 [in larval, pupal keys].

Euorthocladius rivulorum Kieffer. Thienemann, 1935: 201—204, fig. 1 [in pupal, larval keys, distribution, syn-

onymy]. Thienemann, 1936: 191 [ecology]. Thienemann, 1944: 558, 648, fıgs 9, 11, 201 [in pupal, larval keys].

Thienemann, 1954: 23, 49, 108, 147, 191, 301, 303, 309, 344, 345, 347, 349, 356, 360, 670, figs 30, 142 a. Romanis-

zyn, 1958: 27, 82 [in larval key].

Orthocladius (Orthocladius) rivulorum Kieffer. Goetghebuer, 1942: 33, 53, fig. 88 [description of male, female, in

key to males].

Orthocladins ex gp. rivulorum Kieffer. Chernovskii, 1949: 205, fig. 129a [in larval key].

Orthocladius (Euorthocladius) rivulorum Kieffer. Brundin, 1956: 101, fig. 64 [record]. Fittkau et al., 1967: 362

[checklist]. Saether, 1968: 464 [record]. Lehmann, 1971: 486 [ecology]. Lindegaard-Petersen, 1972: 482 [eco-

logy]. Fittkau & Reiss, 1978: 421 [checklist]. Pinder, 1978: 70, figs 35G, 111 B [in key to males]. Rossaro, 1978a:

290, table 1 [ecology]. Rossaro, 1978b: 185 [ecology]. Kownacki & Zosidze, 1980: table 2 [ecology]. Cranston,

1982: 102, fig. 39e [in larval key]. Drake, 1982: 234, fig. 6 [ecology]]. Rossaro, 1982: 42, figs 30, 31 [in pupal, larval

keys]. Murrary & Ashe, 1983: 230 [checklist]. Langton, 1984: 142, fig. 49a [in pupal key]. Sahin, 1984: 80, figs

198, 199 [larval description, ın larval key].

Hydrobaenus rivulorum ? (Kieffer). Roback, 1957 a: 76, 80, figs 183, 184 [in pupal, larval keys].

Orthocladins (Euorthocladius) sp. 1 Oliver et al., 1978: 18, fig. 167 [in larval key].

Orthocladius (Euorthocladius) cf. rivnlorum-suspensus. Coffman, 1973: table 1 [ecology]].

Orthocladius (Euorthocladius) Alaska sp. III Tilley, 1979: 138, 139, fig. 8 [larva).

Orthocladius (Euorthocladius) species 1 Coffman & Ferrington, 1984: figs 25.391— 25.393 [in pupal key].

Orthocladius (Euorthocladius) sp. Coffman & Ferrington, 1984: fig. 25.203 [in larval key].

Chironomus (Orthocladius) sordidellus Zetterstedt sensu Taylor, 1903: 521-523, figs 1, 2 [ecology] [misidentifica-

ton].

Orthocladius sordidellus Zetterstedt sensu Kieffer & Thienemann, 1906:148, 152, 153, figs 7—9 [misidentifikation].

Type Locality: Germany, Ennepe, in Westphalıa.

Type Material: Lectotype: A single female exuviae, labelled by Thienemann as Ennepe, Orthocladius sordidellus,

then later labelled by Thienemann as Orthocladius rivulorum n.sp. These two labels were the only labels on the

slide. The lectotype is circled on the slide as the lower exuviae under the right cover slip when the labels are on the

left side. A paralectotype female exuviae is under the same coverslip: adamaged male exuviae (non-type material)

is under a broken cover slip on the same slide.

Based on Thienemann’s correspondence it’s certain that this is the material associated with the adult females that

Kieffer (1909) used to describe rıvulorum. In a letter sent to Kieffer dated 17-VI-1908 Thienemann wrote: „Ortho-

cladius sordidellus, Ennepe, dicht unter der Sperre 5.VI.08, Gallertgehause wie sie von Taylor u. Lauterborn be-

schrieben“. Kieffer’s answer written in the same letter, in his own handwriting, was ”Orthocladius rivulorum n.sp.

Q”. According to Opinion 1147, Ruling 1 (ICZN 1981), this material qualifies for lectotype designation. I am

hereby designating this material described above as lectotype and paralectotype. There was no other material from

Ennepe collected by Thienemann in the ZSM.

Diagnosis

Orthocladius rivulorum can be distinguished from other Holarctic species of O. (Euorthocladius)

by acombination of characters. Adult Male: lower AR (1.30) and details of the hypopygium (Figs 20,

21). Pupa: hooklets along posterior margin of tergite II, small round spine patches on tergites

IIT-VII, and bubbled thoracic horn. Larva: 17—21 teeth on mentum, weak Lauterborn organs,

squared head capsule, isolated 4th tooth of mandible, slender shape of mandible, and distribution.

Derivation of Name: L. rivulus, a small brook.

Description

Adult Male (n = 4)

Brown (O. D.: yellow body with black bands). Medium to large species. Head. Verticals 12-16,

postorbitals 0-4. Palps long with 34 (1). AR 1.29—1.30 (2). Thorax. Lateral antepronotals 5-9 (3).

Acrostichals absent. Dorsocentrals 5-9. Prealars 4-5 (3). Scutellars 15-27, biserial to multiserial.

Wing. Length 2.05-2.38 mm (2). R with 6-9 setae (2). Squamals 21-31 (3). VR 1.11-1.12 (2). Anal

lobe slightly produced. Legs. LR1 0.73 (2). LR2 0.50—0.53 (2). LR3 0.55—0.56 (2). Sensilla chaetica

on tal of p2, 7-8 (2) and p3, 7-8 (2). Hypopygium (Figs 20, 21). Virga present but difficult to see.

Superior volsella collar-like. Inferior volsella with dorsal part arched convexly, nose-like, and ventral

part covered or slightly extended below.

Variation. The varıation in the adult male is not understood. Apparently the immature stages of rivulorum are

recognized more often than the adults. The reared male from Ireland (Fig. 20) is typical of this species.

Pupa (Exuvia)

Light brown; apophyses on I-VI, variable; length 2.5-4.3 mm. Cephalothorax. Frontal warts

weak or absent; cephalic tubercles absent. Precorneals spaced from each other evenly, 2—3x as long

as dorsocentrals; 2 median antepronotals, O lateral antepronotals, 4—5 dorsocentrals, weak but often

with large sockets; spacing variable (1-1-1-1, 1-1-2, 2-1-1, 2-2). Thoracic horn (Fig. 34b)

long, tubular, filled, with bubbled surface; length 170-260 um. Thorax dorsally smooth with some

sculpturing and/or rugosity mesad of wing base and posteriorly along eclosion line.

Abdomen (Fig. 38a). Tergites: I bare; II-V with median patch of recurved hooklets along posterior

margin; III-VII with small anterior circular patch of posteriorly-directed spines; on VIlIa patch ıs ın-

dicated; [IV—-VI with strong spines in horizontal patch on either side of medial patch; anterior patch

of spinules on II-VIII, most extensive on VI—- VIII. Sternites: I, VIII bare; II-III with central patch

of spinules anteriorly; IV, VII with spinules anteriorly; VIII with 2 off-center patches of spinules an-

teriorly.

Setae (weak) on segments I- VII:

Diva STR Pe 72 IE 1 PR, TRORR JOnBB:. BIN BE PURE C9P & WORR..

Wu A E FEpE SE: N Ode te tt

Anal lobe strongly developed into large, circular lobes; setae absent. Genital sheaths slightly extended

beyond lobe in male, not in female. Pedes spurii B on II; pedes spurii A absent.

Variation. The patches of hooklets are moveable. The arrangement of the dorsocentrals is variable. The thoracic

horn can be weakly or strongly bubbled.

Larva (Fourth Instar)

Body yellow, brown, or green. Head capsule brown; preserved, yellow. Eye spots bipartite or fu-

sed. Head capsule (Fig. 51f) squared. Mentum (Fig. 5le) with 17—21 teeth, usually 19; median tooth

more than 4x as wide as 1st lateral; MR 4.8-8.5 (10); median tooth much higher than Ist lateral. Ven-

tromental plates extended anteriorly between 2nd and 3rd laterals, less commonly between Ist and

2nd or between median and Ist. Epipharynx (Fig. 5la) with premandible simple, slender, and similar

to that in Orthocladius (Orthocladins). Cheatula laterales full, moustache-like. Mandible (Fig. 51c)

with apıcal tooth longer that combined length of 3 inner teeth; 4th tooth separated by space from rest

of mandible; outer margin notched opposite seta subdentalis; rest of margin smooth except for small

notch posteriorly; seta interna present. Antenna (Fig. 51 b) with moderately developed or weak Lau-

terborn organs; blade extended to 5th segment. AR 1.88—2.22 (8). Body with simple, single setae and

possibly arranged like that of saxosus. Anal tubules long, rounded, subequal, with dorsal pair thicker

than ventral pair.

Variation. Chernovskii (1949) and Pankratova (1970) reported an AR of 2.5.

The number of teeth on the mentum are highly variable. Based on figures in the literature, these numbers of teeth

were counted: 17 (Tilley 1979; Sahin 1984); 18 (Romaniszyn 1958; Coffman & Ferrington 1984; Cranston 1982);

19 (Potthast 1914, reproduced in Thienemann 1944, Chernovskii 1949, Pankratova 1970; Johannsen 1937); 21 (Oli-

ver etal. 1978, Rossaro 1982); and 23 (Kieffer & Thienemann 1906). In 18 examined specimens, the number of teeth

on the mentum were: 21 (3), 20 (1), 19 (10), 18 (2), and 17 (2). Sometimes the number of teeth are not symmertrical

(here, and cf. Coffman & Ferrington 1984). Whether the variation in mental teeth is due to instar age, species diffe-

rences, or other factors remains to be determined.

Biology: The larvae usually live on stones and sometimes on moss in fast flowing waters of brooks,

streams, and rivers (Thienemann 1935, 1936, 1954). The gelatinous, cylindrical larval tube is attached

to the substrate at one end, and is often overgrown with one or more species of diatoms (Taylor 1903,

Lauterborn 1905, Thienemann 1954). The tube is transformed into a pear-shaped pupal case, suspend-

ed by an anchor at one end (Fig. 51d). Mites and mermithids have been found associated with rıvu-

lorum (Taylor 1903, Thienemann 1954).

The adults emerge during the winter or early spring. They emerge March to May in the Fulda, Ger-

many (Lehmann 1971); May in Denmark (Lindegaard-Petersen 1972); November, February and

April in Italy (Rossaro 1978a, b); and April and May in Pennsylvanıa, USA (Coffman 1973).

This species also occurs at high altitudes; e.g. alpine brooks (Thienemann 1936, 1954).

Distribution: Palearctic: Austria, Denmark, England, Germany, Ireland, Poland, North Africa

(Lehmann 1971), Norway, Sweden, Switzerland, Turkey. Nearctic: Canada, USA.

Material Examined: Lectotype and paralectotypes. Non-type material: Canada (CNC). Quebec: R. Blanche, $. of Per-

kins, 9-V-1972, A.R.Soponis and J. Robillard, 1MP. New Brunswick: Kouchibouguac Nat’] Park, 30-V-1978, D. R. Oliver

and M.E. Roussel, 1M. N. W.T.: Oscar Creek, 25-V-1972, FWI Pipeline Project, 1L. Yukon Territory: (FWI Pipeline Pro-

ject) Bluefish River, 14-VIII-1972, 2L; Old Crow River, 25-V-1972, 2L; Driftwood River, 16-VIII-1971, 1L; Lord Creek,

19-VII-1972, 1L; Carıbou Bar Creek, 20-VI-1972 1L; 10-VIII-1972, 1L. Denmark: Jutland, Linding Ä., 14-V-1964, Claus

Lindegaard: 14-V-1964, 1L; 22-V-1965, 1F; 3-VII-1968, 1F; V-1968, 1Ex; 20-V-1969, 2L (UCOP). England: East Sussex,

Marsh Green, 17-IV-1978, P. S. Cranston, 5L (BMNH); Tadnoll Brook, 17-V-1976, L. C. V. Pinder, 1 reard M, 1 reared

F (FBA). Germany. A. Thienemann: Bach Kossau, Ostholstein, IV-1936, 1Ex; Partenkirchen, Oberbayern, 3L (ZSM). Ire-

land. River Flesk, Clydagh Bridge, drift, 16-19-V-1978, P. Ashe, 1M w/Ex (DUB). Italy. (B. Rossaro, ROSS): Brembo, 16-

III-1981, 3L, 1Ex; 23-X11-1975, 3L;9-I-1980, 1Ex; Lot Entraygnes passerelle, 22-IX-1977, 1Ex; 26-11-1979, 1Ex. USA. Ar-

kansas: Wash. Co., Tuttle Branch off Rt. 74, 10-I-68, Allen and Fuller, 1M (INHS). Colorado: Arapahoe Co., S. Platt River,

5-XI-1981, M. W. Heyn, 2L (HEY); same locality, 6-XI-1981, P. Guthrie, 1L (GUT). Minnesota (MINN): Cook Co., Min.

F. $. Hovland, N. J. Mosquito Trap, 13-V-1968, E. F. Cook, IM; Mississippi River, Montecello, 19-II-1976, IM; same

locality, 23-II-1978, 1M North Carolina: Mitchel Co., N. Toe River, 2-II-1978, K. Dechart, 3L (NCNR). Oregon: Meto-

lius River, 3-X-1978, W. P. Coffman, 4Ex (COFF). Pennsylvanıa: Crawford Co., Linesville Creek, 13-IV-1971, W. P.

Coffman, 2Ex, (CNC); 7-IV-1971, 2Ex (CNC); 6-V-1971, 2Ex (CNC); Delaware River, 9-V-1976, D. Wartinbee, 6Ex

(COFF); Monroe Co., Big Bushkill Creek, Resica Falls, 17-IV-1976, 1Ex (COFF). South Carolina: Oconee Co., Horse

Pasture River, Salem, 17-III-1977, P. L. Hudson, IM (HUD).

Remarks: Kieffer (1909) described this species from Germany using adult specimens reared by Thienemann. He

separated rivulorum from O. pedestris by the wings, yellow body with black legs and black markings, cubitus not

extended, and non-branched antennal sensillae, adding that the larvae live in gelatinous tubes. He did not provide

figures, he did not mention how many specimens he had, and he did not designate a holotype.

Goetghebuer (1933) placed rıvulorum in Dactylocladius based on the adult characters, and provided the first fi-

gure of the hypopygium and female antenna, as well as amore complete description. He used characters such as

body length, AR, LR, wing venation, and hypopygium. In his material the AR was about 1.00. In his figure of the

hypopygium, the anal point ıs not haired, although the drawing looks like rıvnlorum. However, the examination

of several Goetghebuer types shows that Goetghebuer did not always draw setae on the anal point when present.

Later, Goetghebuer (1942) essentially reproduced the same hypopygial figure and description, and included the

male in akey with AR of 1.00.

Brundin (1956) included rzvulorum in his subgenus Euorthocladins and provided a more accurate figure ofthehy-

popygium, with haired anal point and slight crista dorsalis. Pinder (1978) provided the most recent figure of the hy-

popygium, and included rıvulorum in his key to males of Britain.

The tubes of the larvaand pupa of rivulorum were described as belonging to Chironomus (Orthocladius) sordidel-

lus by Taylor (1903, fıgs. 1, 2) and to an unnamed chironomid by Lauterborn (1905, fig. 15). Taylor’s figures were

reproduced in Thienemann (1954, fig. 30) and here (Fig. 51d).

Potthast (1914) described the larva and pupa of rıvulorum, along with its habits, and provided figures of the man-

dible, mentum, proleg claws, exuviae, and thoracic horn. In his figure of the exuviae, tergite II appears bare. The

patch of recurved spines on II most likely overlapped with the rounded spine patch of III (cf. Fig. 38a).

Thienemann, in Kieffer & Thienemann (1906), provided a figure of the mentum, mandible, pupal spines, and se-

tae of rivulorum. He treated the immature stages as sordidellus, based on the description of the larval and pupal tu-

bes by Taylor (1903). Thienemann compared his material with the descriptions of the immature stages of sordidellus

by Johannsen (1905), who actually described a species of Orthocladins (Orthocladius). Within a few years, Kieffer

& Thienemann (1909) knew they had rivulorum and not sordidellus.

Thienemann (1935) placed rivulorum in his genus Euorthocladius, and gave synonymy, distribution, and keys to

larvae and pupae. He distinguished the larvae by the 8-9 pairs of lateral teeth on the mentum, unequal apices of SI,

moustache-like epipharynx, and unique tubes. He distinguished the pupae by the spines on tergites III- VIII, tho-

racic horn, and unique tubes. Thienemann (1944) used these characters in his keys. He distinguished the larvae of

rivulorum from those of suspensus by distribution; the pupa, by the thoracic horn and spine patterns.

Johannsen (1937) used the thoracic horn and 8—9 pairs of lateral teeth on the mentum to distinguish the immature

stages in his keys, and Roback (1957 a) largely followed him, and figured the simple premandible.

Chernovskii (1949) included rivulorum in his larval keys, and noted an AR of 2.5, green body color, and body

length of 5 mm. Romaniszyn (1958) reproduced the figures of Potthast (1914) and distinguished rivulorum by the

8—10 pairs of lateral teeth on the mentum. Pankratova (1970) also reproduced Potthast’s (1914) figures and used the

9 lateral teeth on the mentum and the long thoracic horn to distinguish rivulorum in her larval and pupal keys. She

recorded the larval body as brown, the AR as 2.5, and the premandible as bifurcate.,

Oliver et al. (1978) included rivulorum as O. (Euorthocladius) sp. 1 in a key to larvae, with a photograph of the

mentum.

Rossaro (1982) included the larva and pupa of rivulorum in his keys, and provided figures. He distinguished the

larvae by the 8-10 lateral teeth on the mentum and the moustache-like epipharynx, and the pupae by the spines on

III— VII. Sahin (1984) included rivulorum in his larval keys, along with figures of the mentum and mandible. Coff-

man & Ferrington (1984) included the pupa and larva in their keys; the pupa keys to couplet 45, the larva keys to

couplet 43.

Orthocladius (Euorthocladius) roussellae n.sp.

Figs. 11, 13—15, 34c, 34d, 41, 50

Orthocladius (Euorthocladius) type II Soponis, 1977: 15, 17, figs. 20, 84e, 92, 100, 121, 107b [larval, pupal descrip-

tions; in larval, pupal keys].

Orthocladius (Euorthocladius) species 3 Coffman & Ferrington, 1984: figs. 25.406, 25.407 [in pupal key].

?Orthocladins (Euorthocladius) sp. Ferrington, 1984: table 7 [drift].

Type Locality: Canada, Northwest Territories, Axel Heiberg Island.

Type Material: Holotype. Male, Canada, NWT, Axel Heiberg Island, 79°25’N, 90°45’W, Gypsum Hill, 20-VII-

1963, H. K. Rutz, CH3635 (CNC). Paratypes (78). (From CNC unless indicated otherwise). Canada. Alberta,

Highwood Pass, 16-VII-1977, D. R. Oliver, CH7131, 1FP w/LS, 1 M w/Ex,LS, 1 F w/Ex,LS; Marmot Creek, 29-

VI-1977, D. R. Oliver, CH7079, 1MP w/LS, 2FP w/LS. Northwest Territories. Melville I., Bailey Point, J. E.H.

Martin: 27-VII-1965, CH25, 3M; 24-VII-1965, CH23, 1M; 20-VII-1965, CH21, 3M; 25-VII-1965, CH27, 1M.

Baffın I., Head of Clyde Inlet, 7-VIII-1958, G. E. Shewell, CH1162, 3M; Frobisher Bay, 5-VIII-1948, F. G. Dil-

abio, CH3653, 1M; Banks I., Masik R., 9-10-VII-1968, W. R. Mason, CH265, 1IFP w/LS; Axel Heiberg I.,H.K.

Rutz, 21-VII-1963, CH 1164, 1M; Slop E, 7-8-1963, CH3656, 3M; Expedition R., 26-VII-1963, CH3634, 4M;

4-VIII-1963, CH1281,3M; Creek SE Gypsum Hill, 21-VII-1963, CH 1164, 2M; Hazen Camp, 81°49’N, 71° 18’W,

D. R. Oliver: 14-VIII-1961, CH3619, 7M 5-VIII-1961, CH3329, 6M; 1M (ZMB); CH3631, 1 male; NE217,

11-VIII-1961, CH3649, 1M, 1MP; 31-VII-1961, CH3594, 1M; Alert, 24-VIII-1963, D. R. Oliver, CH3627, 4M;

Alert, Parr Creek, 25-VIII-1963, D. R. Oliver, CH3627, 3MP, 6Ex. Greenland. Nedre Midsommer So, Can.

Pearyland Expd.: 16-VII-1966, CH 3632, 2M, 19-VII-1966, CH1102, 2M, CH1417, IM. USA. Alaska. Portage

Glacier-Pool, 20-VII-1977, #16, D. Wartinbee, 3Ex (COFF). Wyoming, Inlet Run to Frozen lake, algal mats, 8-

VIII-1981, W. P. Coffmann, 43, 3Ex, #8, 4P, 4LS (Coff).

Diagnosis

Orthocladius roussellae can be distinguished from other Holarctic Euorthocladius by acombination

of characters. Adult Male: low AR (1.02-1.56), numerous and multiserial scutellars, numerous lateral

antepronotals and prealars, and details of the hypopygium (Figs. 11, 14, 15). Pupa: spines on the tips

of anal lobe, hooklets along posterior margin of tergite II, and the long, tubular thoracic horn. Larva:

mentum with 15 teeth, premandible bifid, Lauterborn organs weak, and mandible without seta in-

terna.

Derivation of Name: This species is named after my friend Mary E. Dillon, formerly Mary E. Roussel.

Description

Adult Male (n = 24)

Dark brown. Large species. Head. Verticals 11-30, postorbitals 1-5. Temporals doubled or

clumped medially by coronal suture (Fig. 13). Palps long with 3>4. AR 1.02—1.56. Thorax. Lateral an-

tepronotals 9-27. Acrostichals 1-23, robust, beginn within 1 or 2 AW. Dorsocentrals 6-21, some-

times biserial. Prealars 6-18. Scutellars 25-60, multiserial. Wing. Length 2.52-3.35 mm. R with

6-12 setae. Squamals 19-37. In one specimen, 1 seta on R4+5. VR 1.00-1.08. Anal lobe produced.

Legs. LR1 0.68-0.72. LR2 0.46—0.56. LR3 0.55—0.60. Sensilla chaetica on tal of p3 (6-18, 21). Hy-

popygıum (Figs. 11, 14, 15). Virga absent or, if present, weakly developed, often difficult to see. Supe-

rıor volsella collar-like. Inferior volsella with dorsal part long or short, rounded or square, covering

ventral part. Crista dorsalıs long.

Variation. There is not a high correlation between interocular distance and total palpal length (r=+0.641, p»0.02,

n=15).

Pupa (Exuviae)

Brown, with darker apophyses on II-VI, variable. Length 4.6-6.5 mm (10). Cephalothorax.

Frontal warts moderately developed, cephalic tubercles weak or absent. Precorneals clumped, 2x as

long as dorsocentrals; 2 median antepronotals, 1 lateral antepronotal, 4 dorsocentrals, weak, arranged

1-1-2 or 2-2. Thoracic horn tubular, bare, brown, with expanded base usually smooth (Fig. 34 d);

sometimes collapsed (Fig. 34 c) or partially bubbled; length 230-440 um. Thorax dorsally extensively

rugose.

Abdomen (Fig. 41). Tergites: I bare; II with large central patch of recurved hooklets in 5-6 rows

along posterior margin; III-V with central patch of spines anteriorly, separated from small horizontal

patch of spines along posterior margin; VI as V but without posterior patch; large patch of spinules

on III-VI, small patch of spinules along anterior margin on VII, VIII. Sternites: I with spinules late-

rally; II-IV with large central patch of spinules; V-VIII with 2 off-center patches of spinules ante-

riorly.

Setae on segments I-VIII:

Anal lobe slightly extended with heavy spines on tips; 3 dorsal setae, 2 at midpoint, often branched,

and one on inner margın of distal half. Genital sheaths extended beyond lobe in male, not in female.

Pedes spurii B on II, III, developed with tubercles; pedes spurii A on IV—-VI.

Variation. Sculpturing occurs on tergites VII, VIII. On one male pupa there are 5 L setae on VII. In the three ex-

uviae from Alaska (COFF) the thoracic horns have the surface partially collapsed and bubbled, but not bubbled as

uniformly as in rivulorum (Fig. 34b).

Larva (Fourth Instar)

Body yellow or brown. Head capsule dark brown. Eye spots bipartite or fused. Mentum (Fig. 50e)

with 15 teeth, median tooth as wide or slightly wider than 1st lateral (Fig. 50d); MR 1.2-1.5 (7); me-

dian tooth as high or lower than Ist lateral. Ventromental plates extended anteriorly between 1st and

2nd laterals. Epipharynx (Fig. 50a) with premandible bifid. Chaetula laterales sparse. Mandible (Fig.

50c) with apical tooth as long or longer than Ist inner; outer margin notched opposite seta subdentalis

and crenulated on posterior half on margin; rest of margin smooth, except for notches posteriorly; seta

interna absent. Antenna (Fig. 50b) with weak to moderately developed Lauterborn organs; blade ex-

tended to 5th segment. AR 2.08-3.08 (14). Body with simple setae, short and stiff and long and cur-

ved, at least on I, II; arranged most probably like those in saxosus (Fig. 57). Anal tubules long, roun-

ded, subequal.

Variation. The anterior extension of the ventromental plates is difficult to determine in this species. It merges into

thickenings of the mentum and appears to end between the Ist and 2nd laterals.

Biology: Reared larvae were collected from creeks (Alberta) and rivers (NWT), and from algal mats

in an inlet run to a lake (Wyoming).

Distribution: Nearctic: Canada, Greenland, USA.

Additional Material Examined: Northwest Territories: Axel Heiberg Island, 4M; no label, 1M. Yukon Territory: Cari-

bou Bar Creek, 5L. Alaska: 2L.

Remarks: This species could be confused in the adult male with two sympatric species possessing remarkably si-

milar hypopygia, ©. (Orthocladius) frigidus and ©. (Pogonocladius) consobrinus. The male of roussellae can be di-

stinguished from frigidus by the female-like eyes, and from consobrinus by the numerous, multiserial scutellars, ab-

sence of a fore tarsal beard, and the normally produced anal lobe of the wing. The clumped temporals will separate

males of roussellae from most males of frigidus and consobrinus. However, at least one frigidus and one consobrinus

were examined that have clumped, multiserial temporals instead of the normal uniserial arrangement.

Orthocladius roussellae occurs with frigidus primarily in mountains of temperate regions, and with consobrinus

in the high arctic. The immature stages of these three species are easily separable.

Soponis (1977) incorrectly stated certain characters of the new species. For pupae, PSA occur on IV—-VI, noton

IV-VIJ; frontals are absent, not present. For larvae, the seta interna of the mandible is absent, not present.

Coffman and Ferrington (1984) included the pupa in their keys; roussellae will key to couplet 53.

Ferrington (1984) collected an unnamed species of O. (Euorthocladius) from Inlet Run, Wyoming, where both

roussellae and saxosus have been collected.

Orthocladius (Euorthocladius) saxosus (Tokunaga)

Fıigs. 18, 19, 34 e, 43, 56-59

Spaniotoma (Orthocladius) saxosa Tokunaga, 1939: 326-329, figs. 16, 39, 61, 77, 97, 109, 117, 125, 134, 146, 153

[description of male, female, pupa, and larva]. Tokunaga, 1959; 1973: 642 [pupa, larva, fide Sasa & Yamamoto,

1977).

Euorthocladius sp. Thienemann, 1941: 180 [record from Lappland].

Euorthocladius saxosus (Tokunaga). Thienemann, 1944: 558, 649 [in pupal, larval keys]. Thienemann, 1954: 301,

303, 32,51

Orthocladius (Euorthocladius) saxosus (Tokunaga). Brundin, 1956: 101, fig. 65 [male]. Fittkau et al., 1967: 362

[checklist]. Fittkau & Reiss, 1978: 421 [checklist]. Kownacki & Zosidze, 1980: table 2 [ecology].

Orthocladius saxosus (Tokunaga, 1939). Pankratova, 1970:173, 174, 180, 181, fig. 108 [pupal, larval descriptions; in

pupal, larval keys]. Sasa & Yamamoto, 1977:310 [checklist]. Rossaro, 1984: table 2 [record].

Orthocladius (Euorthocladius) sp. Säwedal, 1978:87 [record of Abisko].

Orthocladius (Euorthocladius) species 5 Coffman & Ferrington, 1984: figs. 25.412— 25.414 [pupa].

®Orthocladius (Euorthocladius) sp. Ferrington 1984: table 7 [drift].

Type Locality: Japan, Kyoto, Kibune.

Type Material: Holotype. Male, Japan, Kyoto, Kibune, Mar 25, 1936, M. Tokunaga (two white labels, one print-

ed). Only the hypopygium and abdominal segments VI- VIII exist, mounted on a slide under one cover slip in Ca-

nada balsam by A. R. Soponis. Paratypes (?8). Same data as holotype. Parts of male and female, including genitalia,

and exuviae; mounted in Canada balsam under eight coverslips on two slides by A. R. Soponis. Previously mount-

ed, two larvae on two slides and parts of female on two slides.

Diagnosıs.

Orthocladius saxosus can be distinguished from other Holarctic species of O. (Euorthocladius) by

a combination of characters. Adult Male: details of the hypopygium (Figs. 18, 19). Pupa: hooklets

along posterior margin of tergite II, with >100 spines, and frontal warts robust. Larva: head capsule

dark brown, mentum with 13 teeth, MR<1.5, and AR>1.80.

Derivation of Name: L. saxum, rock; L. osus, having the nature of, usually indicating abundance. This is probably

a reference to the larvae that aggregate on rocks.

Description

Adult Male (n = 2)

Brown to black. Medium species. Head. Verticals 15-19, postorbitals 1. Palps long with 3>4. AR.

1.19-1.20 (OD: 1.3). Thorax. Lateral antepronotals 4-5. Acrostichals absent. Dorsocentrals 8-9.

Prealars 3-4. Scutellars 13—19, biserial to multiserial. Wing. Length 2.08-2.20 mm. R with 6 setae.

Squamals 18-19. VR 1.00-1.03. Anal lobe not produced. Legs. LR1 0.76 (OD: 0.80). LR2

0.55—0.56. LR3 0.60 (OD: 0.61). Sensilla chaetica on tal of p2 (5) and p3 (8-11). Hypopygium (Figs.

18, 19). Virga present, weak to well developed. Superior volsella collar-like. Interior volsella with dor-

sal part squared, ventral part prominently extended laterally and ventrally below.

Pupa (Exuviae)

Dark to pale brown. Length 2.5—4.0 mm. Cephalothorax. Frontal warts (Fig. 43) large to small; ce-

phalic tubercles absent. Cephalothoracic setae weak and difficult to see. Precorneals short, about

11/,% as long as dorsocentrals, with light sockets; 1 median antepronotal, O lateral antepronotals, 0-4

dorsocentrals, weak; arrangement of dorsocentrals varies. Thoracic horn (Fig. 34) ellipsoid, dark to

light brown, filled, stalked; length 50-100 um. Thorax dorsally granulose anteriorly and along eclo-

sıon line.

Abdomen (Fig. 44). Tergites: I, II bare. II-V with central patch of recurved hooklets along poster-

ior margin; patch on II almost 2X as large as other patches; VI, VII with patch of straight spines along

posterior margin; IIT- VIII with patch of spinules anteriorly. Sternites: I, VIII bare; II—-VI with cen-

tral horizontal patch of spinules anteriorly; VI with spinules along posterior margin; VI, VII with 2

off-center patches of spinules anteriorly.

Setae on segments I-VIII:

D 4

So en VarOr2Z PR

L Da aa as Er 2072) OdHrO ar

Anal lobe greatly reduced, setae absent. Genital sheaths extended beyond lobe in male and female. Pe-

des spurii B on I, II, III, and sometimes IV; pedes spurii A on VI, VI.

Variation. The size and color of the thoracic horn, and the size and shape of frontal warts vary. Setal counts here

do not agree with the original diagnosis. Hooklets are moveable, but the direction of hooklets on tergites ıs not

diagnostic. The arrangement of the DC is variable, and the number of DC varies from 0—4. The body setae are

weak. Small frontal tubercles are found in the type material, and in the material from Wyoming and Sweden. Large

frontal tubercles are found in the material from Alberta, Montana, and Alaska (glacial). The dimorphism does not

appear to be sexual.

Larva (Fourth Instar)

Body yellowish-brown, reddish-brown, or (OD) greenish-brown. Head capsule dark brown; pre-

served, yellow. Eye spots fused. Mentum (Figs. 58c, 59c) with 13 teeth, median tooth about as wide

as Ist lateral; MR 1.2-1.5 (6); median tooth as high as Ist lateral. Ventromental plates extended ante-

riorly between 2nd and 3rd, 3rd and 4th, or 4th and 5th laterals; less commonly, between 1st and 2nd

laterals. Epipharynx (Fig. 56) with premandible simple, blunt, with enlarged apex. Chaetula laterales

sparse. Mandible (Figs. 58a, 59a) with apical tooth as long or longer than Ist inner tooth; outer margin

notched opposite seta subdentalis; rest of margin smooth except for occasional notch posteriorly. Seta

interna present. Antenna (Figs. 58b, 59b) with robust Lauterborn organs; blade extended to 5th seg-

ment or beyond. AR 1.80—2.22 (6). Body with setal arrangement in 4 different patterns (Fig. 57):

those on I, II, III and IV-X. Anal tubules moderately long, rounded, with dorsal pair shorter and

thicker than ventral paır.

97,

Variation. The apical tooth of the mandible is variable, appearing more reduced in Nearctic than in type material,

but this may be due to orientation. Also, the mandible has 5 true teeth in the type material and in the Montana ma-

terial, but 4 true teeth and a false tooth in the Alberta material. The premandible is deeply bifid in a single reared

larva from Montana.

Distribution: Palearctic: Italy, Sweden. Japan. Nearctic: Canada (Alberta). USA (Alaska, Colo-

rado, Montana, North Carolina, Oregon, Wyoming).

Material Examined: Type material. Non-type material: Canada. Alberta: Marmot Creek, 29-VI-1977, D. R. Oliver,

7079, 5MP w/LS; 1FP w/LS (CNC). Japan. Mt. Hıkosan, Kyushu, III-18-1980, L. T., M. Yamamoto, 2M, 1F (YAM). Swe-

den. Lappland 1936, 41, orig. Thienemann material, 4Ex (ZSM). USA. Alaska: Portage glacial pool, 20-VII-1977, #19, D.

Wartinbee, 2Ex (COFF). Colorado: Gunnison Co., Beaver Dam on East R., 3.1 mi. N of Gothic, 13 July 1982, L. Ferring-

ton, No.Co. #9, 6Ex, IP (SBSK). Montana: Beartooth-Absaroka Wilderness area, 31-VII-1979, E. R. Wells, CH6965.1, 1

MP w/LS (CNC). Glacier Nat’] Park, small stream in west meadow w of Logan Pass (Continental Divide), 11-VIII-1975,

R. W. Lichwardt, MBL-13, 27a, 1Ex, 1FP w/LS$, 14a, 1FP w/LS (ANSP). North Carolina. Richmond Co., Forest Creek,

24-I-1981, D. Lenat, 1L; Wake Co., Cane Creek, 9-II-1980, D. Lenat and $S. Mozley, 1L; Wake Co., Reedy Creek at US

40, 28-XII-1979, D. Lenat and K. Eagleson, 2L (NCDNR). Oregon: Deschutes Co., Century Drive, Goose Creek, Head-

water Springs, 20-VIII-1983, B. Wisseman, gelatinous sheat, wood, 4L; Lane Co., H. J. Andrews Exp. For., Mack Cr., 11

mi. NE of Blue River, 20-V-1982, 1M; 26-V-1984, 1M; 18-19-II-1977, B. B. Frost, drift sample, 6pm—9am, clearcut, 1Ex

(OSU). Wyoming: Trib. to Beartooth Lake, 19-VIII-1974, S1.2, W. P. Coffman, 4Ex (COFF); Park Co., ”Inlet Run”-Fro-

zen Lake nr. Beartooth Pass, 5-VIII-1978, L. Ferrington, 20Ex (SBSK).

Biology: Larvae lıve in clear, irregular gelatinous tubes 10— 16 mm long and 3—5 mm wide, closely

adhering to stones along small crevices in rapid mountain streams. Pupal tubes are more oval, 7—10

mm long and 4-6 mm wide, with 3 mm long stems. Larvae are common in winter (Tokunaga 1939).

This species occurs with suspensus. Mites have been associated with saxosus (Thienemann 1954). Fer-

rıngton (1984) collected an unnamed species of O. (Euorthocladius) from Inlet Run, Wyoming, where

both saxosus and roussellae have been collected.

Remarks: Tokunaga (1939) described this species from an unspecified number of males, females, pupae, and lar-

vae collected in a rapıd stream at Kibune, Kyoto, Japan. Type material of all stages still exists.

Thienemann (1944) recognized saxosus as belonging to his genus Euorthocladins and included the pupae and lar-

vae in his keys. He separated the pupae primarily by the patches of hooklets on tergites II-V, and the larva by the

anal tubules and labrum. He also recognized that his pupal skins in the 1941 Lappland work belonged to this species.

Brundin (1956) placed saxosus in the subgenus Orthocladius (Euorthocladius) and provided the first complete il-

lustration of the hypoygium. Pankratova (1970) provided a description of the pupa and larva with figures reprodu-

ced from Tokunaga (1939). In the pupal keys she did not separate saxosus from thienemanni and rivicola, but did

so in the larval keys using the dorsal pair of anal gills which are shorter than the ventral pair.

Pankratova (1970) mentioned that the larval premandible is bifurcate, seen only in one examined specimen from

Montana.

This species has been included in several checklists (Fittkau et al., 1967, Fittkau & Reiss 1978, Sasa & Yamamboto,

1977), probably because it could be easily identified. The male and pupa of saxosus are very distinctive. In addition,

Brundin’s (1956) illustration of the hypopygium and Thienemann’s (1944) pupal key have contributed to the rela-

tive ease of identification of the species.

Coffman & Ferrington (1984) included the pupa in their keys; saxosus keys to couplet 55.

Orthocladius (Euorthocladius) suspensus (Tokunaga)

Fig. 17

Spaniotoma (Orthocladius) suspensa Tokunaga, 1939: 323—326, figs. 15, 38, 63, 64, 70, 80, 85, 100, 118, 122, 135,

145, 151 [description of male, female, pupa, and larva]. Tokunaga, 1959; 1973: 642 [pupa, larva, fide Sasa & Ya-

mamoto, 1977].

Euorthocladius suspensus (Tokunaga), Thienemann, 1944: 558, 649 [in pupal, larval key.]. Thienemann, 1954: 345

[note].

Orthocladins (sen.str.) suspensus (Tokunaga). Tokunaga, 1964: 17, fig. 2 [notes on adult].

Orthocladius suspensus (Tokunaga, 1939). Sasa & Yamamoto, 1977: 310 [checklist].

[non] Orthocladius suspensus (Tokunaga, 1939) sensu Ree & Kim, 1981: 176, 177, plate 27 [misidentification].

Type Locality: Japan, Kyoto, Kibune.

Type Material: Holotype. Male, mounted on a slide under 5 coverslips in Canada balsam by A. R. Soponis. Japan,

Kyoto, Kibune, Mar. 25, 1936, M. Tokunaga (typed label). Preserved in Kyushu University.

Diagnosis

Orthocladius suspensus can be distinguished from other Holarctic species of O. (Euorthocladius) by

a combination of characters. Adult Male: low AR (<1.8) and details of the hypopygium (Eier17).

Pupa: hooklets along posterior margin of tergite II, small patches of spines on tergites IIT- VIII,

smooth thorcic horn, absence of spine rows on posterior margins of tergites IV—-VI. Larva: mentum

with about 19 teeth; distinguishable from rivulorum by distribution.

Derivation of Name: L. suspendere, to hang up. This is probably a reference to the larval tube, which is suspended

from stones by one end.

Description: (See also Tokunaga 1939)

Adult Male (n = 1, holotype)

Brown (OD: black). Large species. Head. Verticals 12. postorbitals 0. Palps long with 3<4. AR 1.68

(OD: 1.5-1.7). Thorax. Lateral antepronotals 3. Acrostichals absent, but sockets may be present.

Dorsocentrals 10. Prealars 4. Scutellars 23, multiserial. Wing. Length 3.28 mm. R with 6 setae. Squa-

mals 34. VR 1.08. Anal lobe not produced, almost right-angled. Legs. (OD: LR1 0.69. LR3 0.57). Sen-

silla chaetica could not be determined. Hypopygium (Fig. 17). Virga absent or vestige present. Super-

ior volsella collar-like. Inferior volsella with dorsal part nose-like, covering most of ventral part. Cri-

sta dorsalis long.

Biology: The larvae occur in mountain streams in Japan. They live ın cylindrical tubes, 20-37 mm

long, 2.6—-3.5 mm wide, coated with a thick growth of diatoms that makes the tubes look green. Pupal

tubes are smiliar in size to larval tubes, with an oval chamber at the free and. The oval chamber is

pointed at the distal end, with one respiratory opening at either end of the chamber, 7 mm long and

3.2 mm wide (Tokunaga, 1939). Both Taylor (1903) and Lauterborn (1905) have described tubes of

rivulorum smilarly. This species occurs with saxosus.

Distribution: Japan.

Material Examined: Holotype.

Remarks: Tokunaga (1939) described this species from an unspecified number of males, females, pupae, and lar-

vae collected in amountain steam at Kibune, Kyoto, Japan. Only the holotype, acomplete male, was located among

the original type material at Kyushu University (Dr. Y. Hirashima, pers. comm.).

It has not been easy to identify saspensus in the adult male. Tokunaga (1939, fig. 38) illustrated only part of the

hypopygium. Later, Tokunaga (1964, fig. 2) illustrated ahypopygium that was dorsoventrally compressed with the

ventral part of the inferior volsella pushed below the dorsal part. He had placed suspensus in Orthocladins (s.s.)

without comment, probably influenced by the distorted inferior volsella. In addition, these specimens had an AR

of 3.00, a record for the species and for the genus Orthocladius. This may prove to be a misidentification, but be-

cause the figured specimen was distorted, and only the holotype has been examined, it is difficult to decide.

Ree & Kim (1981), most likely following Tokunaga’s (1964) work, described as suspensus the male and female of

a species of Orthocladins (Orthocladius). This species belongs to the subgenus Orthocladius because of the uniserial

scutellars, presence of acrostichals, pointed anal point, and double-lobed inferior volsella.

The immature stages of suspensus are morphologically similar to those of rıvulorum, and the larvae of both species

live in similar tubes. The adult male of suspensus has a distinctly different hypopygium (Fig. 17) from that of rivu-

lorum (Figs. 20, 21), particularly in regard to the inferior volsella. Tokunaga (1939) provided characters to separate

the pupae of rivulorum from suspensus, and so did Thienemann (1944), based on Potthast’s (1914) description of

rivulorum. Characters used in the keys are based on the original description.

Thienemann (1954) briefly commented on the similarity of suspensus and rivulorum.

Orthocladius (Euorthocladius) telochaetus Langton

Fig. 16

Orthocladius (Euorthocladius) telochaetus Langton, 1985: [description of male].

Type Locality: Spitzbergen: Advent Bay.

Type Material: Holotype. Male, Spitzbergen, Advent Bay, Holmgren (two white labels), 116 79 (pink label),

Riksmuseum Stockholm (green label), ”These 2 ©’ apparently = decoratus-F. W. E.” (tan label), ”Ch. limbatellus

Holmgren 1869 OÖ PARALECTOTYPE Orthocladius (Euorth.) telochaetus n. sp. Langton P. H. 1985 HOLO-

TYPE” (white label). Paratype. Male, similar green and white labels with 115 79 on pink label.

Diagnosis

The male of Orthocladius telochaetus can be distinguished from other males of O. (Enorthocladins)

by details of the hypopygium (Fig. 16), primarily the apical seta on the anal point. The pupa and larva

are unknown.

Derivation of Name: Gr. telos, end; chaite, seta. Langton named this species for the apical seta on the anal point

of the hypopgyium.

Description: See Langton (1985). Additional characters include: head female-like; VR 1.09; acrosti-

chals absent; >5 dorsocentrals, uniserial; 3 prealars; anal point with microtrichia.

Biology: Unknown.

Distribution: Palearctic: Spitzbergen.

Material Examined: Type material. Non-type material: Spitzbergen: Gipsdalen, 17-VII-1954, Tage Roos,

CH3659, 1M (CNC).

Remarks: Langton (1985) described this species from two males in the mixed type series of Chironomus limbatel-

Ius Holmgren. One aspect of the hypopygium makes this species unique in Orthocladins: a single apical seta on the

anal point. Microtrichia on the anal point, while rare, occur in other O. (Euorthocladius) from high latitudes.

The male of telochaetus is morphologically similar to saxosus, as Langton (1985) pointed out. These two species

can be distinguished by these characters of telochaetus: the more numerous squamals (27—31, 3; saxosus 13—19, 2);

absence of sensilla chaetica on tal of p3, lower LR’s, and more robust virga. Both species occur at high latitudes,

although saxosus is found in low arctic (Lappland) and telochaetus is found in high arctic (Spitzbergen).

Orthocladius (Euorthocladius) thienemanni Kieffer

Figs. 25b, 27, 28, 34f, 37d, 40, 42, 52

Orthocladius thienemanni Kieffer. Kieffer & Thienemann, 1906: 143, 144, 146— 149, figs. 1-5 [description of adult,

pupa, larva]. Kieffer & Thienemann, 1909: 32 [ecology]. Potthast 1914: 263, figs. 1-5 [pupa, larva]. Thienemann,

1935: 203— 205, fig. 2[synonymy, in pupal, larval keys, notes, distribution]. Pankratova, 1970: 173, 174, 177,178,

fig. 105 [pupal, larval descriptions; in pupal, larval keys]. Illies, 1971: 46, table 5 [ecology]. Drake, 1982: 231, 234,

240, fig. 6 [ecology]. Reiss, 1983: 176 [checklist]. Rossaro, 1984: table 2 [record]. Bitüsik & Ertlovä, 1985: 604,

606, table 2 [ecology].

Orthocladius (Orthocladius) thienemanni Kieffer. Goetghebuer, 1932: 75, 89, fig. 144 [adult description, in adult

key]. Goetghebuer, 1942: 34, 36, 55, fig. 95 [male, female descriptions; in male, female keys].

Spantiotoma (Orthocladius) thienemanni Kieffer. Johannsen, 1937: 56, 60, 62, 72 [in pupal, larval keys; notes].

Enorthocladius thienemanni Kieffer. Thienemann, 1944: 559, 648, fig. 12 [in pupal, larval keys]. Romaniszyn, 1958:

27, 82, fig. 122, 126, 127 [in larval key]. Thienemann, 1954: 146, 182, 345, 349, 356, 360, 364, fig. 102 [notes].

Hydrobaenus (Bryophaenocladius) thienemanni (Kieffer). Kloet & Hincks, 1945: 337 [checklist].

Hydrobaenus thienemanni (Kieffer). Roback, 1957: 76 [in pupal, larval keys].

Orthocladius ex gp. thienemanni Kieffer. Chernovskii, 1949: 205, 282, fig. 129b [synonymy, in larval key].

Orthocladius (Euorthocladius) thienemanni Kieffer. Brundin, 1956: 95, 96, 101, fig. 63 [notes, distribution]. Fittkau

et al., 1967: 362 [checklist]. Lehmann, 1971: 486 [ecology]. Kloet & Hincks, 1975: (V)15 [checklist]. Rossaro,

1977: 122 [distribution]. Rossaro, 1978a: 290 [distribution]. Rossaro, 1978b: 185 [distribution]. Fittkau & Reiss,

1978: 421 [checklist]. Pinder, 1978: 70, fig. 35E, 111C [in key to males]. Cranston, 1982: 102, figs. 39a, c, f [in

larval key]. Moubayed & Laville, 1983: 223 [distribution]. Murray & Ashe, 1983: 230 [checklist]. Langton, 1984:

144, fig. 49d [in pupal key]. Sahin, 1984: 82, figs. 200-202 [in larval key]. Michailova, 1985: 149, 158, 159, 163,

164, pl. v, viii [cytology].

Orthocladius (Euorthocladius) ct. thienemanni-saxosus [partim]. Coffman, 1973: table 1 [ecology].

Orthocladius (Euorthocladius) species 6 |partim]. Coffman & Ferrington, 1984: fig. 25.415 [pupa].

[non] Spaniotoma (Orthocladius) thienemanni Kieffer sensu Edwards, 1929: 344, 345, fig. 6m [partim] [in key to

males] [misident. ].

[non] Orthocladins (Enorthocladius) cf. thienemanni (Kieffer). Halvorsen et al., 1982:119 [record].

Type Locality: Germany: Insel Rügen: Thuringen (see Cranston 1984).

Type Material: Lectotype. Male, original label hand-written, Orthocladius thienemanni K.; printed label, R. 1.

Sc. N.B. 18.073, coll et det M. Goetghebuer; typed label. Boites no 6 Types Kieffer. Previously mounted in balsam

under two coverslips on a slide; overcleared. Abdomen III-IX dissected from body; hypopygium intact. Wings

crumpled and folded over; head squashed; only antennal flagellomeres 1 and 2 present; parts of legs missing.

Hereby designated as lectotype.

Presumably original labels have been replaced, and other labels added to the lectotype in routine curation, be-

cause ”coll et det M. Goetghebuer” makes no sense in a species reared by Thienemann and described by Kieffer.

Presently there appear to be no other sepcimens that unquestionably belong to the original type series.

Diagnosıs

Orthocladius thienemanni can be distinguished from other Holarctic species of ©. (Euorthocladius)

by a combination of characters. Adult Male: high AR (>1.75), relative lengths of palpal segments

(3=4), and details of the hypopygium (Figs. 27, 28). Pupa: rows of spines on anal margins of tergites

HI —- VIII; thoracic horn present; absence of hooklets on II and absence of pedes spurii A. Larva: 13

teeth on the mentum, wide median tooth (MR>1.5), and high AR (>1.8).

Derivation of Name: Kieffer named this species after A. Thienemann.

Description

Adult Male (n = 5)

Dark brown. Large species. Head. Verticals 10-23, postorbitals 1-3 (4). Palps long with 3<X4 (4)

(Type: 3=4). AR 1.75—2.15 (2). Thorax. Lateral antepronotals 2-7 (4). Acrostichals absent. Dorso-

centrals 4-16. Prealars 3-7. Scutellars 15—26, biserial to multiserial (Type: 22, multiserial). Wing.

Length 2.80-3.18 mm (4). R with 4-10 (3) setae. Squamals 30—40 (4). VR 1.06-1.14 (4). Anal lobe

moderately produced (OD: right-angled). Legs. LR1 0.68-0.74 (3). LR2 0.45—0.53 (4). LR3

0.53—0.56 (4). Sensilla chaetica on tal of p2 (7-13) (4), absent on p3. Hypopygium (Figs. 27, 28).

Virga present. Superior volsella collar-lıke. Inferior volsella with dorsal part slender, inner margin

rounded, apex rounded or squared, and ventral part covered or extended below. Crista dorsalis long,

robust.

Variation. Only six specimens, two originally collected and identified by Thienemann, have been examined. Leh-

mann (1971) reported the AR of thienemanni as 1.80—2.00 from his Fulda material.

Pupa (Exuviae)

Light brown, with darker apophyses on I-VIII, variable. Length 3.15—4.29 mm (10). Cephalotho-

rax. Frontal warts weak to moderately developed; cephalic tubercles absent. Precorneals clumped,

weak to strong, 1'/,x as long as dorsocentrals; 2 median antepronotals, 1 lateral antepronotals, 3 dor-

socentrals in a row, thicker than precorneals. Thoracic horn (Fig. 34f) ellipsoid, light brown, filled or

clear, stalked; length 30-70 um (10). Thorax dorsally wrinkled to granulose anteriorly along eclosion

line.

Abdomen (Fig. 42). Tergites: I bare; II— VII with 2—4 rows of strong spines along posterior mar-

gin (Fig. 37d); II-VIII with central patch of spinules anteriorly. Sternites: I, VIII bare; I—VII with

2 off-center patches of spinules anteriorly.

Setae on segments I-VIII:

D 4

Su u, Da 2 BR: SE ae LE

V Ba a Fa @d=720270=0>°0)0 0 0.0

Anal lobe greatly reduced, one seta at midpoint. Genital sheaths extended beyond lobe in male and fe-

male. Pedes spurii A and B absent.

Variation. Associated exuviae from the Fulda (collected by Lehmann) and from Bathurst Island, NWT, as well

as pupae from Kansas were examined here. The European material is more robust than the North American mate-

rial. Anterior shagreen may be absent on tergite II.

Larva (Fourth Instar) (n = 11)

Body yellow, brown, or green. Head capsule brown; preserved, yellow. Eye spots fused. Mentum

(Fig.52c) with 13 teeth, median tooth about 2X as wide as Ist lateral; MR 1.5—2.3; median tooth

higher than 1st lateral. Ventromental plates extended anteriorly between 2nd and 3rd laterals. Preman-

dible simple. Chaetula laterales sparse. Mandible (Fig.52a) with apical tooth as long or longer than first

inner tooth; outer margin notched opposite seta subdentalis, rest of margin smooth except for occasio-

nal notch posteriorly; seta interna present. Antenna (Fig.52b) with robust Lauterborn organs; blade

extended to 4th segment. AR 1.85—2.56. Body with simple, long setae, most likely arranged like that

in saxosus (Fig. 57). Anal tubules subequal, moderately long, all same length and thickness, weakly

pointed (Thienemann 1944: rounded).

Variation. Cranston (1982) recorded the head capsule as yellow-brown, and figured the mandible without the

margin notched opposite the seta subdentalis. In one specimen excamined here, the premandible appears notched.

Material from Sabina Creek, Arızona, shows considerable varıation in the width of the median tooth. With a lo-

wer MR, theese larvae will key to saxosus. The larvae of both of these species occur gregariously on stones.

Thienemann (Kieffer & Thienemann 1906) described the larva as green. In the material examined here, all

preserved on slides, the body is either yellow, green, or brown.

Biology: Larvae live on the surfaces of larger stones in fast-flowing brooks, streams, and rivers

(Kieffer & Thienemann 1906, Thienemann 1935, 1944, 1954, Lehmann 1971). Larvae live in clear ge-

latinous tubes encrusted with sand grains, often in fissures and depressions of the stone. Larvae are

also associated with algal growth on stones (Thienemann 1954) and with the common bulrush, Schoe-

noplectus lacustris (Drake 1982). In a chalk stream of southern England, Drake (1982) found that thie-

nemanni was the most abundant species of larval chironomid in the cold months. In his study, larvae

were present during high discharge, but absent in low flow.

Pupae live in the enlarged larval tubes: half-ellipsoid, 6 mm long, 3 mm wide, 2 mm high, clear

gelatinous tubes covered with small particles. The pupa undulates, causing water to flow through

openings at both ends of the tube, as figured here (Fig.40) and in Thienemann (1954, fig.102) after

Miall and Hammond (1900, fig.5). The pupal stage lasts 3-7 days (Kieffer & Thienemann 1906, Thie-

nemann 1954).

Gregarious pupation of thienemanni in a spring-fed tributary of Sabina Creek near Pigeon Springs,

Arızona, was observed by Jan Doughman (pers. comm.). In February, 1984, water temperature was

4°C with ice on the surface. Larvae were feeding on a thin film of diatoms on rocks. Larvae congregat-

ed in small (0.5 cm dıam) vertical depressions on a 1’ X 1’ granite rock, then stopped feeding, and spun

a gelatinous sheath over themselves (up to eight larvae under one sheath). Mature pupae alone, or pu-

pae with mature larvae were found under some sheaths.

This species occurs with rivicola (based on exuviae) in seven localities (see rıvicola); and with calvus

(based on adults) in Germany.

Adults swarm in large numbers (Thienemann & Kieffer 1906). Thienemann (1935) reported adults

appearing in the first of spring. Illies (1971) recorded emergence of adults from April to August in

Breitenbach, Germany. Lehmann (1971) reported two generations in the Fulda: the first from January

to May, the second in October. In southern England, Drake (1982) found two generations in 1976 and

1977, the first emerging January to April, the secound in March and April. In South Carolina, adults

emerge from January through April.

Doughman (pers. comm.) has observed that the adults of thienemanni hold their wings rooflike

over the abdomen.

Distribution: Palearctic: Denmark, England, Germany, Ireland, Lebanon, Switzerland, and Tur-

key. Nearctic: Canada, Greenland, USA.

Material Examined: Lectotype. Non-type material: No locality: coll. et det. M. Goetghebuer, R.I.Sc.N.B. 18.073, 1M

(BRUX). Canada. Northwest Territories: Oscar Creek, 25-V-1972, FWI-Pipeline Project, CH6475, 2P. Ontario: Ottawa,

Ottawa River, 22-IV-1966, Jon Martin, 0122-1, 2PEx (CNC); Rushing River, 9-V-1978, W.P. Coffmann, 2PEx (COFF).

Denmark. Zealand, Lellinge Ä, 20-V-1968, C. Lindegaard, 3Ex, 2L (? COP). England. Fast Sussex, nr. Forest Row,

51414347, 17-IV-1978, P. S. Cranston, BM1978-197, 1L (BMNH); Bucks, River Chess, 16-11-1982, W. R. Karsteter, IL

(FSCA). Germany. River Schwentine, East Holstein, 1935, A. Thienemann, IM; Insel Rügen, A. Thienemann, 1M, 2P,

Baumberge, bei Münster, Westfalen, 2-II-1908, Thienemann, IM; Fulda, Hessen 10-I-1969, Nr. 27b, J. Lehmann, IM w/

PEx; Fulda, Br. Sandlofs, 17-X-1952, E. J. Fittkau, 12 PEx (ZSM). Greenland. Tilloe Narssag Elv. hole 740, 24-VIII-1981,

C. Lindegaard, 3PEx; 9DR, 11-12-VII-1981, M w/Ex (? COP). Switzerland. Stein am Rhein, 20-III-1966, F. Reiss, 1M

(ZSM). USA. Alaska. N.Fk. Chena R., 11-VII-1978, D. Wartinbee, 1Ex (COFF); Jım River above Prospect Camp, left

bank, 095073, USGS, 1Ex (CALD). Arızona. coll J. Doughman: Pima Co., Sabino Creek at Summerhaven (0.6 mi up forest

rd.) on Mt. Lemmon nr. Tucson, from rocks, 5-II-1984, 840205, 1FP, 1MP, 10L; 26-11-1984, 840226, 1MP, 1FP, 1L; Pinal

Co., Boyce-Thompson Arb., Queen Cr., Cladophora zone, 840201, 1FP, 4L (USGS). Georgia. Fannin Co., Noontootla

Creek at Newport Rd., 24-IV-1979, 1L, 1FP(CALD). Kansas. Johnson Cop., Cedar Creek, 29-XII-1977, P.L., 4FP,4MP;

Douglas Co., Deer Creek, 0.5 mı $ of Stull, 20-III-1981, L. Ferrington, 13PEx; Atchinson Co., stream 5.5mi S, 0.2 mi At-

chinson, 24-III-1982, L. Ferrington, 1F w/Ex, IM w/Ex (SBSK). North Carolina. Yancey Co., Cane R., Sta. 4, XI-1980,

K. Dechart, 1L; Macon Co., Calor Fork, 15-I-1981, K. Dechart, 1L; Mitchel Co., N. Toe R., 2-II-1978, K. Dechart, IL;

Iredell Co., Buffalo Shoals Creek, Jan 1981, K. Dechart, 1L; Haywood Co., Pigeon R., May 1980, site #4, IL, Surry Co.,

Ararat R., 14-IV-1981, K. Eagleson, 1L; Transylvanıa Co., French Broad R., nr. Rosman, 25-IV-1978, D. Penrose, IL

(NCNR). Pennsylvania: Linesville Creek, 7-IV-1971, W.P. Coffmann, 1Ex (CNC). South Carolina. Oconee Co., Seneca

Cr., Seneca, 22-I-1977, P.L. Hudson, 1Mw/Ex (HUD); 10-11-1977, 1M w/Ex; Oconee Co., East Fk, Chattoga River, Natl

Fish Hatchery, 9-III-1978, P.L. Hudson, 1MP (HUD); Oconee Co., Seneca, Flat Shoals River, 9-II-1977, P.L. Hudson,

1M, 2Ex (HUD); Oconee Co., Fall Creek, Lake Keowee, 30-IV-1974, P.L. Hudson, 1Fx (HUD). Tennessee. Pigeon River,

Gatlinburg, 3-V-1977, P.L. Hudson, 1 Ex (HUD).

Remarks: In a joint publication (Kieffer & Thienemann 1906), Kieffer named and described this species from the

adult male and female. He did not designate a holotype, nor give any information on his material. He did describe

"general body characters, but gave no figures or details of the genitalia. In the same paper, Thienemann described the

pupa and larva of the named species. The description of the immature stages was sufficiently complete to allow the

species to be understood, particularly in the pupal stage.

Edwards (1929) included thienemanni under his Group C, or Orthocladius (s.str.) of his subgenus Spaniotoma

(Orthocladius). He separated the males of thienemanni and O. glabripennis by the absence of afore tarsal beard and

an AR of 2.00 in thienemanni. His figure 6m was the fırst available reproduction of the hypopygium. Until recently

this figure was interpreted as representing O. thienemanni. However, I have examined some of Edwards’s material

from Herts and Gloucester that he used for his 1929 work. Some of these specimens belong to arecently described

species, O. (Euorthocladins) calvus Pinder, which can be distinguished from thienemanni in adult male essentially

by the relative lengths of palpal segments 3 and 4.

Goetghebuer (1932) reproduced Edwards’s (1929) figure of O. thienemanni and separated it from other ©. (Or-

thocladıus) by color, AR, LR, and hypopygial details. Later Goetghebuer (1942) essentially reproduced the same

figure and description, and included females in his keys to adults.

Brundin (1956) provided a figure of the hypopygium and listed thienemanni as type of the subgenus O. (Ortho-

cladius) without providing other characters of the species. Pinder (1978) provided the most recent illustration of the

hypopygium of thienemanni, and included the species in a key to males on the British chironomids.

The immature stages, particularly the pupa, have received more attention in the literature than the adults. Thiene-

mann’s larval description (Kieffer & Thienemann 1906) was brief and without drawings. He compared the larva of

thienemanni to the larva of O. sordidellus as described by Johannsen (1905), probably an Orthocladius (Orthocla-

dius). Ithad an AR of 1.66, an MR of 1.5, and the Lauterborn organs were either weak or absent, based on Johann-

sen’s figures. The pupa of sordidellus was certainly not O. (Euorthocladius), but Thienemann made no mention of

this. His pupal description of thienemanni was more complete, but Thienemann mistakenly recorded the diagnostic

spine rows as present on tergites II-VII, although his figure 2 correctly shows them on tergites III-VIII.

Thienemann (1935) erected the genus Euorthocladius with thienemanni as type. He gave a synonym for thiene-

manni and its known distribution as Germany, England, and Switzerland. In his larval keys he separated thiene-

manni from rivicola by the higher AR (1.85 versus 1.28) and the longer body. In his pupal keys Thienemann sepa-

rated thienemanni from rivcıcola by the spine rows on III, smaller thoracic horn (see remarks under rivicola), and

longer body.

Johannsen (1937) treated thienemanni in the group Euorthocladius under his subgenus Spaniotoma (Orthocla-

dius). He distinguished the larva of thienemanni from the larva of rivulorum by the equally long forks of the SI (after

Thienemann 1935, figs. 1, 2), and from obumbratus by the robust Lauterborn organs and sparser pecten. He distin-

guished the pupa of thienemanni from the pupa of rivulorum by the thoracic horns.

In keys, Thienemann (1944) separated the pupa of thienemanni from other Euorthocladins pupae by the arrange-

ment of spines on the abdominal tergites, the structure of the thoracic horn, and the body length. He separated the

larva from other Euorthocladius by the length of the anal tubules, AR (1.85), teeth on the mentum, SI, and pecten.

Chernovskii (1949) largely followed Thienemann (1935) and treated the thienemanni group in alarval key, distin-

guished from rivicola by higher AR (2.00 versus 1.10) and longer body (8 mm).

Roback (1957 a) included’ thienemanni in his keys and separated the larvae (on a subgeneric level) from other Or-

thocladius (as Hydrobaenus) by the large Lauterborn organs. He distinguished the pupae by the thoracic horns and

the spine patterns of the abdominal tergites.

Pankratova (1970) separated the larvae of thienemanni from rivicola by the higher AR (2.00 versus 1.40), but did

not distinguish the pupa of thienemanni from rıvicola or saxosus. She gave a complete description of both larva and

pupa, and reproduced figures from Chernovskii (1949), and mentioned that the premandible was bifurcate. A

notched premandible was seen in only one specimen here, but this is a difficult character to assess in O. (Euortho-

cladins). A morphologically similar species, calvus, can be distinguished from thienemanni by the bifid preman-

dible.

Cranston (1982) separated thienemanni from frigidus and rivicola by the Lauterborn organs and SI of the mandi-

ble, and provided figures of the mentum (MR>1.5) and the antenna (AR=1.8).

Romaniszyn (1958) used Potthast’s (1914) figures and separated thienemanni from rivicola by AR and body

length. Coffmann & Ferrington (1984) included the pupa in their keys; thienemanni will key to couplet 55. Halvor-

sen et al. (1982) referred to their material of O. ashei as cf. thienemanni.

References

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Ir - a -transverse

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dorsal part of

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Fig. 1. Orthocladins (Euorthocladins) luteipes Goetghebuer, hypopygium, dorsal. Fig. 2. Orthocladius (Eudac-

tylocladius) sp., hypopygium, dorsal. Fig. 3. Orthocladius (Pogonocladius) consobrinus (Homgren), hypopy-

gium, dorsal.

Fig. 4. Orthocladius (Orthocladius) trigonolabis Edwards, hypopygium, dorsal.

Fig. 5. Orthocladius (Orthocladius) ferringtoni Soponis, hypopygium, dorsal.

3 |

lectotyped'

Figs. 6-8. Orthocladins (Enorthocladius) abiskoensis Thienemann & Krüger. 6. Hypopygium, dorsal, Edwards

type material. 7. Lectotype slide. 8. Hypopygia, dorsal. a. Isachsen, NWT b. Hazen Camp, NWT ce. Carıbou Bar

Creek, NWT.

Big. 9,

Orthocladius (Euorthocladius) coffmani n.sp., hypopygium, dorsal, holotype.

Fig. 10.

Orthocladius (Enorthocladius) anteılis (Roback), hypopygium, dorsal, Idaho.

ANErE

Figs. 11-15. 11. Orthocladins (Enorthocladius) roussellae n.sp., hypopygia, dorsal, partypes. a. Baffın Island,

NWTb. Axelheiberg Island, NWT c. Greenland d. Melville Island, NWT. 12. Orthocladius (Orthocladins) frigidus

(Zetterstedt), head, frontal. 13-15. Orthocladius (Euorthocladins) roussellae n.sp. 13. Head, frontal. 14, 15. Hypo-

pygia, dorsal, paratypes, Hazen Camp, NWT.

Fig. 16. Orthocladius (Euorthocladins) telochaetus Langton, hypopygium, dorsal, holotype.

Figs. 18-19. Orthocladius (Euorthocladius) saxosus (Tokunaga), hypopygia, dorsal. 18. holotype. 19. a, b. Japan,

non-type material.

Figs. 20-21.

Orthocladins (Enorthocladius) rivnlorum Kieffer, hypopygia, dorsal. 20. Ireland. 21

. England.

Fig. 22.

Orthocladius (Euorthocladius) kanıi (Tokunaga), hypopygium, dorsal, paratype.

Fig. 23.

Othocladins (Enorthocladius) luteipes Goetghebuer, hypopygium, dorsal, Italy.

Fig. 24.

Orthocladius (Euorthocladins) Inteipes Goetghebuer, hypopygium, dorsal, holotype.

Figs. 25-26. Orthocladius (Euorthocladius) calvus Pinder, hypopygia, dorsal. 25. a. Gloucester, England. 26. Ri-

ver Schwentine, Germany. 25. b. Orthocladius (Euorthocladius) thienemanni Kieffer, hypopygium, dorsal. River

Schwentine, Germany.

Figs. 27—28. Orthocladius (Euorthocladius) thienemanni Kietfer, hypopygia, dorsal. 27. a. Switzerland b. det.

Goetghebuer. 27. lectotype.

Figs. 29-30. Orthocladius (Euorthocladins) ashei n.sp., hypopygia, dorsal, paratypes. 29. Norway. 30. Ireland.

Fig. 31. Orthocladius (Euorthocladius) difficılis (Lundbeck), hypopygium, dorsal, lectotype.

Figs. 32-33. Orthocladius (Enorthocladius) rivicola Kietfer, hypopygia, dorsal. 32. Ottawa. 33. a. Idaho b. South

Carolina c. NWT.

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Figs. 34-37. Orthocladius (Euorthocladius), pupal thoracic horns. a. coffmani n.sp. b. rivnlorum Kieffer c-d.

roussellae n.sp. c. Alberta d. NWT e. saxosus (Tokunaga) f. thienemanni Kieffer g. rivicola Kieffer. 35-36. Ortho-

cladius (Enorthocladius), pupal dorsocentral setae. 35. rıvzcola Kieffer. 36. ashei n.sp. 37. pupal spines. Tergite II: a.

calvus Pıinder. Tergite IV: b. /uteipes Goetghebuer c. rıvicola Kieffer d. thienemanni Kieffer e. ashei n.sp. f. caluns

Pinder.

_ Sharon ;

N hy: \

Figs. 33-41. Orthocladius (Euorthocladius), pupal abdomen, dorsal. 38a. rıvunlorum Kieffer. 39. coffmani n.sp.

Anal lobe. 38b. ?n.sp. nr. rvnlorum. Orthocladius (Euorthocladius), pupae. 40. thienemanni Kieffer, pupa in tube,

after Miall and Hammond (1900). 41. rousselae n.sp., pupal abdomen, dorsal.

ve I ö a &

\ NEN,

MUST uaiinner

Figs. 42, 44,45. Orthocladius (Euorthocladius), pupal abdomen, dorsal. 42. thienemanni Kieffer. 44. saxosus (To-

kunaga). 45. abiskoensis Thienemann & Krüger, frontal warts, lateral. Fig. 43. saxosus (Tokunaga).

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Figs. 46-49. Orthocladius (Enorthocladins), pupal abdomen, dorsal. 46. Inteipes Goetghebuer, Pennsylvanıa,

USA. 47. Goetghebuer, Italy, segments IV-IX. 48. ashei n.sp. 49. rivicola Kieffer.

Fig. 50. Orthocladius (Enorthocladius) roussellae n.sp., larva. a. epipharynx b. antenna c. mandible, with variation

d. variation of mental teeth e. mentum.

Fig. 51 Orthocladius (Euorthocladius) rivulorum Kieffer, larva. a. epipharynx b. antenna c. mandible d. larval

(left) and pupal (right) tubes, after Taylor (1903) e. mentum f. head capsule, dorsal.

52a N | j ) , U

N AR

2 c © { c e

Figs. 52-54. Orthocladius (Enorthocladius), larvae. 52. thienemanni Kieffer. a. mandible c. antenna c. mentum.

53. Iuteipes Goetghebuer. a. mandible b. antenna c. mentum. 54. ashei n.sp. a. maxilla, dorsal b. maxilla, ventral c.

mandible d. antenna e. mentum.

Fig. 55. Orthocladius (Euorthocladius) rivicola Kieffer, larva. a. epipharynx b. mandible c. antenna d. mentum.

Fig. 56-59. Orthocladius (Euorthocladius) saxosus (Tokunaga), larva. 56. epipharynx 57. body, segments I-IV,

ventral (left), dorsal (right). 58. Alberta: a. mandible b. antenna c. mentum. 59. Type series: a. mandible b. antenna

c. mentum.

Fig. 60. Orthocladius (Euorthocladius) abiskoensis Thienemann & Krüger, larva. a. epipharynx b. mentum c. head

capsule, dorsal d. maxilla, dorsal e. premento-hypopyaryngeal complex, dorsal (left) and ventral (right) f. posterior

body, anal tubules and procercus g. mandible h. antenna.

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SPIXIANA Supplement 14

ISSN 0177— 7424

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ZEITSCHRIFT FÜR ZOOLOGIE

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Gesamtherstellung: Gebr. Geiselberger, Altötting

Lars Brundin

zum 80. Geburtstag

Jeder, der heute mit Chironomiden arbeitet, ist im Grunde genommen ein Schüler von

Lars Brundin. Er war es, der die Chironomidenforschung mit einem neuen Konzept in die

2. Hälfte unseres Jahrhunderts führte. Er setzte neue Maßstäbe auf diesem Gebiet der

aquatischen Entomologie. Seine Zeichenkunst erschloß der Chironomidensystematik bis-

lang entbehrte Grundlagen. Seine Arbeitskraft, Konzentrationsfähigkeit und Zielstrebig-

keit demonstrierten, zu welch umfassenden Aussagen und Leistungen man bei Benthos-

untersuchungen kommen kann. Seine große wissenschaftliche Begabung machte es ihm

möglich, seinen taxonomischen Erfahrungsschatz und nicht zuletzt seine tiefe Naturliebe

in phylogenetische und biogeographische Grundlagenforschung einzubringen. Er war

wohl der erste Zoologe, der das Werk von W. Hennig: „ Grundzüge einer Theorie der

phylogenetischen Systematik“ durchgearbeitet und seine grundlegende Bedeutung für die

biologische Forschung erfaßt hat. Er wurde der überzeugendste Interpret Hennigs bis

zum späteren weltweiten Durchbruch dieser Theorie.

Wir schätzen uns sehr glücklich, daß Prof. Brundin immer noch aktiv an der Chirono-

midenforschung und der Diskussion innerhalb der Phylogenie und Biogeographie teil-

nımmt. Zu seinem 70. Geburtstag widmete ihm die schwedische Entomologische Gesell-

schaft eine Festschrift. Dort erschien auch eine Würdigung seiner bisherigen vielseitigen

und umfangreichen wissenschaftlichen Arbeit, einschließlich eines Verzeichnisses seiner

Veröffentlichungen. Inzwischen liegt eine Reihe weiterer bedeutender Publikationen von

ihm vor, bzw. sind im Druck.

Mit dieser Sammlung von chironomidologischen Beiträgen unterschiedlicher For-

schungsrichtungen wollen seine Freunde, seine „Schüler“, Lars Brundin danken für seine

wegweisenden Arbeiten, seine anregenden Gespräche, seinen gewichtigen Beitrag zur mo-

dernen Chironomidenforschung. Diesem Dank und dem Glückwunsch zum 80. Geburts-

tag schließen sich zahlreiche Freunde und Fachkollegen an, die sich an dieser Festschrift

nicht beteiligen konnten.

30. Maı 1987

Ernst Josef Fittkau

INHALT — CONTENTS

ARMITAGE, P. D. & J. Tuiskunen: Thalassosmittia atlantica (Stora) comb.

nov. Description of adult female and immature stages

from Tenerife, Canary Islands (Diptera, Chironomidae)

CaspErRs, N.: Zwei neue Smittia-Arten aus dem süddeutschen Raum

(WWiptera@hironomidae)Ere ern,

COoFFMANN, W.P., Yurasıts, L. A. & C. pe a Rosa: Chironomidae of South

India. I. Generic composition, biogeographical relation-

ships and descriptions of two unusual pupal exuviae (Dip-

tenay @hironomidae) nt za ea

CONTRERAS-LICHTENBERG, R.: Tanytarsus curvicristatus spec. nov., eine neue

Chironomidenart aus Kolumbien (Diptera, Chironomi-

N) Level he be AR

CransToN, P. $S. & D. R. OLıver: Aquatic xylophagous Orthocladiinae —

systematics and ecology (Diptera, Chironomidae)......

Devaı, G.: Emergence patterns of chironomids in Keszthely-basın of

Lake Balaton (Hungary) (Diptera, Chironomidae).....

Errer, L.H.: A reconsideration of the genus Apedilum Townes, 1945

(Bipseray@hironomidae)er re

Facnanı, L. P.& A. R. Soponis: The occurrence of setal tufts on larvea of

Orthocladius (Orthocladins) annectens S&ther (Diptera,

Ehironomidae)ir Ri MOMAIDEHEERUIR SRRSEIGINEEMNMDE

Fırrkau, E. J. & D. A. Murray: Bethbilbeckia floridensis: anew genus and

species of Macropelopiini from the South Eastern Ne-

arenie(Wiprtera,Chironomidae)ar rear

Haıvorsen, G. A.: Redescription of Paratrissocladins acuminatus (Ed-

wards) comb. nov. (= Cardiocladius acuminatus Ed-

wards) from Southern Chile (Diptera, Chironomidae)...

HırvenoJa, M. & E.: Corynoneura brundini spec. nov. Ein Beitrag zur Sy-

stematik der Gattung Corynoneura (Diptera, Chironomi-

BERN: Wald. eilt. bas-ry rer...

Kress, B. P. M.: Some Records of two rare Chironomid species in the Ne-

therlands (Diptera,@hironomidae)n... 2: ren.

LavıLte, H. & F. Reıss: Rheomus, un nouveau genre du complexe Harni-

schia avec deux nouvelles especes d’Afrique du Nord

(Dipteray@hironomidaeppea ne

MAKARCHENKO, E. A., KıKNADZE, J. J. & J. E. Kerkıs: Morphokaryological

description of Euryhapsis subviridis (Siebert) from the

South of the Soviet Far East (Diptera, Chironomidae)...

Seite

25— 28

175-181

155-165

101-104

143—154

201-211

105-116

139—142

253—259

85— 89

213—238

29— 33

183—190

129—137

MicHai.ova, P. VL.: A Review of the Genus Polypedilum Kieffer. The cy-

totaxonomy of Polypedilum aberrans Tshernovskji (Dip-

tera, Chronemidaee ra:

Murray, D. A. &E. J. Fırrkau: Schineriella schineri gen nov. comb. nov.,

placement of Tanypus schineri Strobl 1880 (Diptera, Chi-

FOROIMdAR,, „er ee ee

Reiıss, F.: Die Gattung Kloosia Kruseman, 1933, mit der Neube-

schreibung zweier Arten (Diptera, Chironomidae).....

Rosack, $. $. & R. P. Rurrer: Denopelopia atrıa, anew genus and species

of Pentaneurini (Diptera: Chironomidae: Tanypodinae)

item Elanıda vice re

Rossaro, B.: A contribution to the knowledge of chironomids in Italy

(Diptera, Chironomidae) a... 2.2 u. ee

SATHER, O. A.: Diplosmittia recisus spec. nov. from Peru (Diptera, Chiro-

nomidae)y. So a 2 0 u

SETHER, ©. A. & O. A. Schnerı: Heterotrissocladius brundini spec. nov.

from Norway (Diptera, Chironomidae).............

SATHER, OÖ. A. & O. A. SchneLL: Two new species of the Rheocricotopus

(R.) effusus group (Diptera, Chironomidae)..........

SATHER, OÖ. A. & E. Wırrassen: A review of Lappodiamesa Serra Tosio,

with the description of L. boltoni spec. nov. from Ohio,

USA (Diptera, Chironomidae).. ... - 2.0lzrerzgersge ge

SCHNELL, A. & O. A. SarHer: Vivacricotopus, anew genus of Orthocladii-

nae from Norway (Diptera, Chironomidae)..........

WIEDERHOLM, T.: Changes in the profundal Chironomidae of Lake Mälaren

during 17 years u Ri in re

Wırrassen, P. E. & B. Serra-Tosıo: Description de trois femelles de Dia-

mesa Meigen dont D. cinerella Meigen (Lectotype et Pa-

ralectotype) (Diptera, Chironomidae)..............: =.

Wırson, R.S.: A survey of the zinc-polluted River Nent (Cumbria) and

the East and West Allen (Northumberland), England,

usingichironomid.pupalewıyiae 3. sb en a

WÜLKER, W. & J. Herrmann: Die weiblichen Gonodukte in normalen und

parasitierten Chironomus Imagines (Diptera, Chironomi-

daele a a un

239-246

247-252

35—44

ee,

191-200

45— 47

57— 64

65— 74

75— 84

49— 55

7-15

91-100

167-174

17-24

SPIXIANA | “ Supplement 14 | 7-15 | München, 15. Juli 1988 ISSN 0177-7424

Changes in the profundal Chironomidae of Lake Mälaren

during 17 years |

By T. Wiederholm

Abstract

The profundal benthos of Lake Mälaren was sampled in September/October each from 1970 and onwards. Con-

siderable fluctuations in population densities of the most abundant taxa occurred. In many cases the temporal pat-

terns were similar between stations, indicating common causal mechanisms. The role of food, climatic conditions

and biotic interactions is discussed.

Introduction

Lake Mälaren is the third in sıze of Sweden’s lakes. The lake serves as a water supply to Stockholm

and other surrounding communities, supports a significant fishery and is used for various recreational

activities among the population of 1.1 million within its drainage area.

Limnological monitoring of Lake Mälaren and ıts main trıbutaries has been performed since 1966.

Studies of the bottom fauna were started in 1969. Sampling network and methods have been the same

since 1970. Earlier results were reported in WIEDERHOLM (1974, 1978). In the present paper I describe

the changes that have taken place among the major taxa of profundal Chironomidae and discuss the

importance of trophic conditions and other factors to these changes.

Methods and material

Five Ekman samples (15x 15x30 cm) were taken in September/October each year from 19 stations throughout

the lake. Eight of these stations, representing the major subareas of the lake, are discussed here (Fig. 1). The sedi-

ments were washed through a 0.6 mm net and preserved in 70% ethanol in the field. The samples were stained using

Bengal Rose, sorted under microscope and the animals identified to species or genus (Chironomidae and others) or

higher taxonomic level (Oligochaeta, Hydracarina). Data were stored and treated with the use of codes in MAIT-

LAND (1977) (amended by myself and co-workers), Uppsala University’s computer (SAS, own programmes) and

PC:s (statistics and diagrammes in this paper).

The chemical and biological conditions in Lake Mälaren are described in detail by WILLEN (1984). The lake is

composed of several more or less isolated basins. The western and northern parts are the most eutrophic, with con-

siderable variation within and between years in phytoplankton biomasses and water chemistry. The central basins

are deeper, somewhat colder and more homogenous temporally.

Results

General

The profundal Chironomidae of Lake Mälaren was predominated for the most part by seven taxa.

Chironomus anthracinus Zett., C. plumosus L. and Procladius spp. dominated in the western and

$ ,

E Wi n zZ

: Swede: G

60° N 60°N Bes

"LAKE MÄLAREN

Lake Hjälmaren

Denmark

@ GG Sigtunafjärden

{} Sy

R N

Ü +

nr i 2 N. Prästfjärd

= astijarden

= Blacken Granfjärden x ° J

OD TG Ge

z TG

) DD Ga

. TG

0 10 20 km >

= S. Björkfjärden G

Fig. 1. Lake Mälaren with sampling stations.

northern basins (Blacken, Granfjärden, Ekoln and Sigtunafjärden in Fig. 1). C. neocorax Wülker &

Butler occurred in western Lake Mälaren. Micropsectra spp., Tanytarsus spp. and Procladins spp. do-

minated in the central part of the lake, where Chironomus was absent. Sergentia coracına and Stictochi-

ronomus spp. were also part ofthe fauna of the central basins but in lower numbers than the other taxa.

It has not been possible to identify all larval material to species. Imagines of Micropsectra caught by

hand-net at the central basıns were identified as M. insignilobus Kıieffer by Säwepaı (1976). Several

species of Tanytarsus and Procladius have been found and at least two species of Stictochironomus, one

of which was S. rosenschoeldi (Zett.), have been reported to occur in hand-netted material (WIEDER-

HOLM 1974). Chironomus larvae of the salinarius type from western Lake Mälaren were described as

C. neocorax by WüLker & Butter (1983) and none of the other species with thıs larval type are likely

to occur ın the lake. The identity of ©. plumosus and C. anthracinus has not been established from

chromosome identifications, but these species are the most likely ones to occur in great numbers at the

depth that has been sampled here, viz 15 m (cf. LINDEBERG & WIEDERHOLM 1979).

Considerable temporal variation in numbers have been noted for most groups of benthic organısms

during the study period. Oligochaeta and Chironomidae decreased in numbers at most stations

throughout the period. Chaoborus flavicans became more abundant during later years (station Blac-

ken in Fig. 2). Crustaceans occurred in high numbers in the central part of the lake during some few

years (station N. Prästfjärden in Fig. 2).

Chironomus anthracinus

C. anthracinus occurred in particularly high numbers ın Blacken and Sigtunafjärden during the first

part of the period (Fig. 3). The species then decreased at all stations and population density was rather

low throughout the 1980’s. Considerable variation occurred between individual years, and the corre-

Oligochaeta

Pisidium spp

P. affinis

C. flavicans

Chironomidae

ODNSESH

Oligochaeta

Pisidium spp

P. affinis

Chironomidae

Oligochaeta

Pisidium spp

P. affinis

Chironomidae

year

Fig. 2. Occurrence of major groups of benthic organisms 1970— 1986 (ind. m?) in western (stn. Blacken) and cen-

tral Lake Mälaren (stn. N. Prästfjärden).

lation coefficients between the various data sets for single years were low, except between Blacken and

Sigtunafjärden. The general pattern of development was quite similar at all stations, however, as appe-

ars from the figures and the correlation coefficients for running means (Fig. 3, Table 1).

C. plumosus

C. plumosus fluctuated in numbers at most stations without any clear general trend or correlation

between stations (Fig. 3, Table 1). High numbers occurred in Ekoln during the first part of the study

period, but the population density decreased strongly at the end of the 1970’s and remained low

throughout the remainder of the study period.

C. neocorax

This species has a rather uneven distribution within Lake Mälaren. Similar to the other Chironomus

species it is absent from the central basıns, and it occurs in high numbers only in some of the modera-

tely eutrophic western basıns. Of the stations dealt with here, only Granfjärden had high population

densities of C. neocorax, and there was no clear temporal pattern or correlation between stations

(Fig. 3, Table 1).

Micropsectra spp.

Large numbers of Micropsectra were found during only afew years (Fig. 3). Distinct peaks occurred

at all stations between 1973 and 1976 and the running means of the 30 and 50 m depths in both

N. Prästfjärden and $. Björkfjärden were strongly correlated (Table 1). The overall occurrence of

Micropsectra was low throughout the 1980’s.

Tanytarsus spp.

The occurrence of Tanytarsus was similar to that of Micropsectra (Fig. 3). Peaks occurred at all sta-

tions in 1974 and 1976, and very low numbers were found after 1979. The data sets from the various

stations were strongly correlated (Table 1).

Sergentia coracına

This species was rare or absent most years, but distinct peaks occurred in 1976 or 1977 in three out

of four stations in the central part of Lake Mälaren (Fig. 3).

Stictochironomus spp.

Stictochironomus (not shown in figure or table) had its main occurrence in 1976-79, i. e. somewhat

after the peaks of Micropsectra and Tanytarsus. Significant numbers were found only in N. Präst-

fjärden.

Procladius spp.

Two peaks occurred at the 30 m stations in central Lake Mälaren — one around 1976 and the other

one in 1982-83 (Fig. 3). The running means from these stations were strongly correlated (Table 1).

At 50 m depth in N. Prästfjärden the second peak was much less pronounced, but the general pattern

was similar. The 50 m station in S. Björkfjärden showed much less variation, but the highest numbers

occurred in the first part of the study period.

In the western and northern basins temporal variation was seemingly greater, without much sımi-

larity between stations (Fig. 3, Table 1). A general tendency to decreasing abundances may be noted,

however.

The occurrence of Procladins was similar to that of Micropsectra and Tanytarsus. Running means of

Procladius were strongly correlated to those of Micropsectra (r = 0.91—0.97) (not shown in table) at

the 30 m stations, though less so at the 50 m stations (r = 0.17—0.38). There was also a positive corre-

lation with running means of Tanytarsus or Tanytarsus plus Micropsectra (r = 0.65—0.91) atthe 30 m

stations, though again this trend was weaker at the greater depth.

Discussion

Considerable variation occurred between years. However, for most taxa trends or regular patterns

of varıation are clearly discernible. In many cases the similarity between stations, as indicated by high

correlation coefficients between the data sets, indicates acommon causal mechanism for the observed

variation. For example, the high numbers of Micropsectra, Tanytarsus, S. coracına and Stictochirono-

mus at most stations in central Lake Mälaren during the mid—1970’s would seem to have acommon

reason. Decreasing numbers of C. anthracınus were common to the stations of western and northern

Lake Mälaren and one may suspect acommon structuring factor here also. The numbers of Procladius

were strongly correlated at stations in the central part of the lake, but not at the western and northern

stations, and this may indicate that different factors are predominating. No pattern could be seen in

the numbers of C. plumosus and C. neocorax, except for the marked decrease of the former species at

station Ekoln.

Food, weather conditions and biotic factors are important determinants acting independently or

collectively in governing population density. Improved sewage treatment in all major communities

around the lake has brought about a 50 % reduction of the phosphorus loading on Lake Mälaren since

the middle of the 1960°s. The occurrence of a series of dry years up to 1976 added to this and the total

phosphorus loading decreased to less than half of its maximum during the last 20 years. Evidently as

a result of this, the excessive blooms of bluegreen algae in the western and northern parts of the lake

decreased in intensity and duration (Wirren 1987). Small species of algae (e. g. flagellates) became more

common and the number of species increased. The average total biomass of algae did not decrease to

any great extent, however, except for the most eutrophic parts of the lake.

The significance of these changes to the bottom fauna is not immediately clear. Both quantitative

and qualitative changes in profundal food supply may have resulted. Because many of the small, non-

colonial species of algae are more effective producers than large, colonial species, there is reason to be-

lieve that the total planktonic primary production per unit area is about the same as before or even

higher. Measurements to support this conclusion are lacking, however. One might assume that a grea-

ter proportion of the phytoplankton is metabolized in the water column when small and mobile forms

are more predominant. Bluegreens such as Anabaena are also metabolized in the water phase (FaLLon

& Brock 1980), but others sink and decompose on the lake bottom. This may result in increased sedi-

ment oxygen demand, but the increased microbial production favours many invertebrates that tolerate

low oxygen levels. Among these are several species of Oligochaeta. Hence the reduced numbers of

Oligochaeta in many parts of Lake Mälaren (cf. Fig. 2, station Blacken) may therefore be understood

as reflecting a reduced nutrient flow through bacteria and sediments, resulting from the reduced oc-

currence of bluegreen algae.

Little information is available on food utilization of aquatic insects — certain types of ingested mat-

ter are probably of little value, whereas small amounts of other material may be critical (LAMBERTINI &

Moore 1984). C. anthracinus and C. plumosus are both detrivores, but algae sometimes make up a

considerable part of the gut content (Jönasson 1972, LAMBERTINI & MooRrE 1974). Johnson (1985) con-

cluded that C. anthracinus is more of a deposit feeder, ingesting particulate matter scraped from the

recently deposited surface sediments, whereas C. plumosus is a filter feeder with the nutritional qualı-

ty of ingested matter depending primarily on pelagic inputs. This would explain the decreasing num-

C. anthracinus C. plumosus C. neocorax Procladius spp

Blacken

Granfjärden

Bra

Ze, :

N |

BNEo.2

Bas

Sigtunafjärden

Micropsectra spp Tanytarsus Spp S. coracina

200

N. Prästfjärden 30 m N. Prästfjärden 30 m

vr DEREN

- o

1000

S. Björkfjärden 30 m

O abe of a

Ha BR 4 Bere Pa

eu Be TEenken

Fig. 3. Occurrence of major profundal Chironomidae 1970-1986; yearly and 5 yr running means (ind. m).

Please observe differences in scales.

bers of C. anthracinus if, as argued above, one accepts that the changes in phytoplankton species com-

position has increased the relative importance of the pelagic metabolism relative to that of the benthic

one.

The decrease of C. plumosus at station Ekoln is probably more of a local change than part of agen-

eral pattern. Hypereutrophic conditions occurred at thıs station before effective sewage treatment was

installed in the nearby city of Uppsala, and C. plumosuns endured this better than C. anthracınus. It is

surprising, however, that both species have decreased to virtual extinction during later years.

Contrary to the other species dealt with here, Procladius spp. are predominantly carnıvorous, al-

though algae and detritus may also be found in their gut content (TArwın 1969, BAKER & MCLACHLAN

1979). Oligochaetes, other chironomid larvae and small benthic crustaceans are the preferred prey.

Data on the ocurrence of crustaceans are not available. There was a positive correlation between the

running means of Procladins spp. and those of Oligochaeta at two of the four stations in western and

northern Lake Mälaren. The correlation was negative and very weak at the central stations, where Mi-

cropsectra and Tanytarsus were better correlated with Procladius. This may indicate that the fluctua-

tions in numbers of Procladins depended on fluctuations of their prey organisms, but it may also indi-

cate that some of these taxa responded to some other factor that led to acommon temporal pattern (cf.

below).

Micropsectra, Tanytarsus, Sergentia coracıina and Stictochironomus had their maximum occurrence

during the mid—-1970’s. All are presumably browsers or filter feeders and typical inhabitants of the

mesotrophic or moderately oligotrophic lake types as characterized by Brundin (1956). They were

characteristic members of the profundal chironomid communities in the central part of Lake Mälaren

throughout the study period dealt with here, but high population densities occurred only during a few

years. It seems unlikely that this pattern would have been due to adirectional change toward more me-

sotrophic or oligotrophic conditions that would have presented optimal conditions during these years

only. Thus underlying factors with a more stochastic varıation should be sought to explain the occur-

rence of these taxa. Weather conditions coincident with emergence may be one such factor. Calm and

warm weather should be favourable to swarming, egg-laying, and hence the recruitment of young lar-

vae, particularly in species with a short and well synchronized flight period. Phenological data is

scarce, however, and the available information indicates rather long or even several flight periods.

Imagines of M. insignilobus are common in May and June at the central basins of Lake Mälaren (Wır-

DERHOLM 1974). Stictochironomus rosenschoeldi and another species of Stictochironomus occur in June

and July. Records of adult Sergentia coracina from southern and central Sweden include late June and

September (Brunnın 1949); no adults have been found from Lake Mälaren. If one assumes that the 4th

instar larvae found in September— October were born in the early summer the same year, weather con-

ditions ın May and June should be important. Weather records show that these months were particu-

larly warm in 1976, with 230 day degrees above the average for 1970-85 and nearly 350 day degrees

above the minimum during the period, which occurred in 1982. Micropsectra, Tanytarsus and Sergen-

tia coracina had population peaks in 1976 at some stations. Peaks did occur in other years also, howe-

ver, when temperatures were below average (e. g. 1985). Nevertheless it is still possible that the peaks

in larval abundance that occurred during these years were related to weather conditions, but more de-

tailed life history information is needed to support this conjecture.

Between 5000 and 10000 ind. m ?, occasionally as much as 20000 ind. m?, of Pontoporiea affınis

may be found at 50 m depth at station S. Björkfjärden (WIEDERHoLM unpubl.). The physical disturban-

ce by such numbers of crustaceans should have a negative influence on the sessile, tube-living and

rather fragile larvae of Tanytarsus and Micropsectra, and a strong negative correlation did occur

between running means of P. affinis and the two chironomid taxa (r = —0.85 and —0.91, respec-

tively). Hence, the absence of large numbers of P. affinis might be a prerequisite for Tanytarsus and

Micropsectra to occur in reasonably large populations densities even during years when weather

conditions and other circumstances were favourable to recruitment of young.

C. anthracinus

Blacken Granfj. Sigtunafj. Ekoln

Blacken 0.48/0.91 0.72/0.92 0.39/0.63

Granfj. 0.08/-0.51 0.60/0.98 0.34/0.78

Sigtunafj. 0.01/0.61 -0.22/-0.77 0.60/0.83

Ekoln 0.17/0.60 0.12/-0.32 0.08/0.51

C. plumosus

C. neocorax

Blacken Granfj. Sigtunafj. Ekoln

Blacken 0.15/-0.47 ORSIELAOMTEI

Granfj. 0.46/0.60 0.21/-0.43

Sigtunafj. -0.04/-0.28 -0.11/-0.27 -/-

Ekoln 0.34/0.55 0.08/0.26

Procladius spp

Micropsectra spp

N. Prästfj. 30 S. Björkfj. 30 N. Prästfj. 50 S. Björkfj. 50

NerPrast£je 30 0.30/0.76 0.47/0.89 0.2570275

S-«#BJorkf]j= 30 0.68/0.89 0.20/0.59 0.95/0.99

NesPrastrn.,50 0.67/0.92 0.80/0.98 0.19/0.57

S2B1)0rkl 2250 0.75/0.85 0.92/0.99 0.82/0.97

Tanytarsus spp

Sergentia coracina

N. Prästfj. 30 SEMBJOrkf. 30 N. Prästfj. 50 S» BJörkf3.350

N. Prästfj. 30 0.05/0.59 0.28/0.64 0.06/0.60

5“, BIockE 230 0.67/0.97 0.32/0.75 0.94/0.99

Ne=PrastrRj-50 0.65/0.87 0.73/0.94 0.60/0.82

SS BJOrKRIJ 2 50 0.60/0.75 0.46/0.80 0.53/0.91

Procladius spp

Table 1.

Correlations between stations for the dominant chironomid species; yearly data/5 yr running means.

The deep profundal of temperate lakes is sometimes thought of as a zone of stable environmental

conditions with little variation in population densities and community composition. The studies re-

ported here are not the first to demonstrate that considerable fluctuations do occur (cf. Jonasson 1972,

HorLoramen & Jönasson 1983). Few time series exist, however, that describe long-term variation ın

profundal Chironomidae. Observations from such series may be used to formulate hypotheses on the

role of abiotic factors, food, competition and predation to variation in population density and com-

munity structure. In particular, long series of field observations are the only practical way to assess the

significance of climatic varation to population densities of profundal Chironomidae. To understand

the role of biotic interactions, more information is needed, in particular, on life histories of Chirono-

midae and how these are linked to seasonal variation of phytoplankton production and deposition of

organic matter on lake bottoms.

Acknowledgements

Lars Eriksson identified and counted the Chironomidae. Richard Johnson read the manuscript.

References

BAKER, A. $. & MCLAcCHLAN, A. J. 1979. Food preferences of Tanypodinae larvae (Diptera: Chironomidae). —

Hydrobiologia 62: 283-288.

BRUNDIN, L. 1949. Chironomiden und andere Bodentiere der südschwedischen Urgebirgsseen. Fin Beitrag zur

Kenntnis der bodenfaunistischen Charakterzüge schwedischer oligotropher Seen. — Rep. Inst. Freshwater

Res. Drottningholm 30: 1— 914.

—— 1956. Die bodenfaunistischen Seetypen und ihre Anwendbarkeit auf die Südhalbkugel. Zugleich eine Theorie

der produktionsbiologischen Bedeutung der glazialen Erosion. — Rep. Inst. Freshwater Res. Drottningholm

37: 186—235.

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in Lake Mendota, Wisconsin. — Limnol. Oceanogr. 25: 72—88.

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via) in the profundal of Lake Esrom, Denmark. — Oikos 41: 99-117.

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nomidae) in mesotrophic Lake Erken. Freshwater Biology 15: 605-612.

JONAssonN, P. M. 1972. Ecology and production of the profundal benthos in relation to phytoplankton in Lake Es-

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D.M. (Eds.) The ecology of Aquatic Insects. — Praeger Publishers, New York.

LINDEBERG, B. & WIEDERHOLM, T. 1979. Notes on the taxonomy of European species of Chironomus (Diptera:

Chironomidae). — Ent. scand. Suppl. 10: 99— 116.

MAITLAND, P. $. 1977. A coded checklist of animals occurring in fresh water in the British Isles. — Institute of Ter-

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SÄWEDAL, L. 1976. Revision of the notescens-group of the genus Micropsectra Kieffer, 1909 (Diptera: Chironomi-

dae). — Ent. scand. 7: 109— 144.

TARwıD, M. 1969. Analysis of the contents of the alimentary tract of predatory Pelopiinae larvae (Chironomidae).

= Eko]a Bol Ser 2241721261511:

WIEDERHOLM, T. 1974. Studies of the bottom fauna of Lake Mälaren. — The National Environmental Protection

Board Report 415. (In Swedish with summary in English.)

—— 1978. Long-term changes in the profundal benthos of Lake Mälaren. — Verh. Internat. Verein. Limnol. 20:

818-824.

WILLEN, E. 1984. The large lakes of Sweden: Vänern, Vättern, Mälaren and Hjälmaren. In: TAus, F. B. (Ed.) Lakes

and Reservoirs. — Elsevier Science Publishers, Amsterdam.

—— 1987. Phytoplankton and reversed eutrophication in Lake Mälaren, Central Sweden, 1965-1983. — Br. phy-

col. J. 22: 193—208.

WÜLKER, W. F. & BUTLER, M. G. 1983. Karyosystematics and morphology of Northern Chironomus (Diptera:

Chironomidae): Freshwater species with larvae of the salinarins-type. — Ent. scand. 14: 121-136.

Dr. Torgny Wiederholm

National Environmental Protection Board

Environmental Quality Laboratory

Box 8005

75008 Uppsala, Sweden

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SPIXIANA | Supplement 14 | 17—24 | München, 15. Juli 1988 ISSN 0177 — 7424

Die weiblichen Gonodukte in normalen und parasitierten

Chironomus Imagines*

(Diptera, Chironomidae)

Von Wolfgang Wülker und Iris Herrmann

Abstract

The histological and functional differences of the four sections of female Chironomus gonoducts are characteri-

zed by light- and ultramicroscopical techniques. The foremost part of the oviductus communis with its high (pos-

sibly glandular) epithelium may produce secretions with importance for egg transport or sperm activatıon. The se-

cond broad part with flat epithelium is able to store eggs after deliberation from the ovary. The third part, charac-

terized by a muscular valve, restricts the egg passage to only one egg at atime. The genital chamber (vagina) ıs re-

sponsible for guiding the penis to the „Spermathekenmündungsplatte“, insemination of ripe eggs, addition of mu-

cus from the gluten gland and extrusion of the inseminated eggs. With regard to these functions, the actions of the

„Spermathekenmündungsplatte“, the attached coxosternapodemes and their musculature, and the processes pro-

truding into the genital chamber from the lateral and ventral sides are discussed. Effects of mermithid worms (Ne-

matoda) on Chironomus gonoducts are variable. Nevertheless, as a rule, the parasitogenic deficiencies begin from

the oral components (oviduct, spermathecae). The „Spermathekenmündungsplatten“ are often misshaped and re-

stricted to the caudal part. In consequence, the coxosternapodemes have arıgid median connection and can not be

moved in the normal way. Histologically, inhibition of development (e. g. oviduct), excess growth (e. g. intima) and

disintegration (e. g. musculature) can be observed. Thus, the sterility of parasitized females is not only a matter of

undeveloped ovaries, but also of the incompleteness, aberrant histological structure and functional incompetence

of the gonoducts.

Einleitung

Die Änderung des Geschlechtsapparates von Chironomus durch parasitäre Mermithiden (Nema-

toda) ist lange bekannt (Remreı 1940, WüLker 1961, Remper etal. 1962). Die weiblichen Imagines wer-

den „kastriert“, d. h. ihre Gonaden werden gar nicht ausgebildet (Remper 1940) oder erreichen nur

etwa !/oo der normalen Größe und enthalten unentwickelte oder degenerierende Keimzellen (WÜüLkEr

1961,:1971). Das 8. Abdominalsternit, in dem die weiblichen Gonodukte ausmünden, verliert ım pa-

rasitierten @ mehr oder minder die typischen Strukturen (z. B. geteiltes Borstenfeld, Subgenitalbucht,

Notum, Öffnungen der Gonodukte) und wird damit scheibenartig wie beim & (Würker 1961, Götz

1964); die zugehörige Genitalimaginalscheibe des 8. Segmentes der Larve ist dementsprechend un-

vollständig (WüLker 1975).

Über die innen liegenden sekundären Geschlechtsmerkmale ist weniger bekannt. Die aus dem

9. Abdominalsegment des @ hervorgehende Schleimdrüse fehlt der parasitierten weiblichen Imago

* Der größte Teil der Untersuchungen wurde vom Erstautor bei der 64. Jahrestagung der Central States (Kansas)

Entomological Society in Lincoln/Nebraska April 1985 vorgetragen.

oder ist winzig (Y/ıoo der normalen Größe, Görtz 1964), ihre Anlage ist bei parasitierten Larven nur in

Form eines ziemlich ungeordneten Zellhaufens vorhanden (Würker 1961, 1975, 1976). Spermatheken,

die aus der Genitalimaginalscheibe des 8. Segmentes hervorgehen, fehlen im parasitierten Tier (REmpeı

1940, WÜLKER 1961, 1978, GöTz 1964). Von den Geschlechtsausführgängen sagt WÜLker (1961), daß

sie meist nur in Andeutungen vorhanden sind, „die aber nur kurz verfolgt werden können und beim

weiblichen Intersex keine Verbindung zum rudimentären Ovar haben“. Nach boraxkarmingefärbten

Totalpräparaten sind sie auf eine Zellmasse am Hinterrand des 8. Segmentes der Imago beschränkt

(Würker 1978), einige zugehörige Muskeln (M. dilatator vaginae, M. dil. oviducti) können identifi-

ziert werden, sind aber unvollständig. Histologische und funktionelle Betrachtungen sind bei norma-

len Imagines selten (z. B. Wenster & Remper 1962, Photographien in SAETHER 1977), für parasitierte

fehlen sie völlig.

Wir haben bei histologischen Untersuchungen an C. anthracinus Zettund C. riparius K. gefunden,

daß schon die Gonodukte normaler weiblicher Imagines unbekannte Strukturen bieten, die der mor-

phologischen und funktionellen Betrachtung wert sind. Bei Parasitierung von C. anthracinus durch

Limnomermis anthracini Kaiser, Wülker & Skofitch 1987 wollten wır feststellen, welche der Gono-

duktabschnitte und Strukturen im parasitierten @ zumindest fragmentarisch gebildet werden, ob sie

typische oder aberrante Form haben, und wieweit die Gonodukte noch zu ihrer normalen Funktion

geeignet erscheinen.

Material und Methoden

Es wurde hauptsächlich an Imagines von Chironomus anthracinus Zett. aus Freilandfängen vom Schluchsee/

Hochschwarzwald (930 m über NN) gearbeitet. Zur Ergänzung ist der leicht im Labor züchtbare C. riparius K.

herangezogen worden. Beide Arten unterscheiden sich in den hier betrachteten Strukturen nur unwesentlich.

Zur Analyse des zellulären Aufbaus bestimmter Strukturen eignet sich alk. Boraxkarminfärbung nach Fixierung

in Carnoy (ROMEIS 1968). Selektive Darstellung der chitinisierten Teile mit der üblichen KOH-Behandlung (10%).

Die mazerierten und gründlich gewässerten Hinterenden können dann zur Untersuchung bestimmter Teile (z. B.

Spermathekenmündungsplatte) mit Mikronadeln präpariert und für rasterelektronenmikroskopische Darstellung

vorbereitet werden. Dazu werden die Objekte in üblicher Weise in Acetonstufen entwässert, critical point-getrock-

net (Balzers Union, CPD 020) und mit Gold beschichtet. Betrachtung in einem Nanolab 7 ZEISS. Die histologi-

schen Schnittserien sind nach Fixierung der Objekte in Glutaraldehyd (6%) in Cacodylatpuffer, Nachfixierung in

1% OsO4, Entwässerung über Alkohol/Propylenoxid und Einbettung in Epon mit 1 um Dicke auf einem Reichert

Om U2 Ultramikrotom geschnitten und mit Toluidinblau gefärbt. Lebendbeobachtungen der Imagines nach leich-

ter Betäubung mit Äther. In der Terminologie der beobachteten Strukturen folgen wir SAETHER 1980.

Ergebnisse und Diskussion

Gonodukte normaler Imagines

Am caudalen Ende der beiden Ovarien findet sich in Querschnittserien unreifer QP der zunächst

zweigeteilte, dann einheitliche Zellpfropf, der den Eiern den Eintritt ins Ovar versperrt (vgl. WENSLER

& Rempeı 1962, Fig. 45). Es gibt, wie schon Azur. Nasr (1950) festgestellt hat, keine paarigen lateralen

Ovidukte. Vielmehr beginnt das Lumen des Ovidukts (odc) gleich unpaar und ist von voluminösen

Zellen (Abb. 2a) umgeben. Dieser Anfang des ersten Oviduktabschnittes liegt noch im 7. Abdominal-

segment und ist durch das große Ganglion (g), in das dasjenige des 8. Segmentes eingeschmolzen ist,

von der Ventralwand des Segmentes getrennt. Es gibt Hinweise auf drüsige Zellkomplexe und sekre-

torische Funktion in diesem Abschnitt, was in Übereinstimmung mit dem Vorhandensein von Drüsen

im Ovidukt bei Culiciden (Cıements 1963) und der Produktion von Gleitsekreten oder Sekreten zur

Spermienaktivierung bei anderen Dipteren (z. B. Drosophila, Übersicht Sanper 1985) stünde.

-sngojfesssefonua\ = [a rulsery = 31 ‘9po sap nua\ = A ‘auejdsdunpunurusyayreunsds = duıs

‘Zunuyousyaweunadg = sd ‘snepdjenusdisog = d3d ‘srunuuros smInpAQ = >po ‘union = OU ‘swopodeu1mısoxoy sap snsI9asuen 'W = X NONPIAO

JOWLNSUO9 snInISNN = O9W ‘uarge] = [| "ewnuruny) = ul “(puej3 usınj3) asnıpunajyag = 83 “(sassa901d ayı]-1adurg) 9Z1es1107 odrunop1adurr = dy “1dıoy2,] =}

‘puemunee] = Mp Zuedusyayreunsdg = sp ‘sngoppesswosiog = [UP “yangjesrusdgng ‘p pueımpioy) usdog = q ‘“(IIIA »sAydodeuon,) sngojumpody = jde

;(e ur aım Bunogo13Ia A Syd1a]9 "nonisered “snumvugıur ‘I (q

‘ogejqy Iop 104 sarzg soul UOISUaWIL] usIun SIy99Yy 'TeUNOU smuvdi4 snwouoag)y (e "IynPou0oH uaysıglM aıp yaınp anuysssäue] spentsesereg :I 'qqV

19,

Der zweite Oviduktabschnitt (nicht abgebildet) ıst kurz und besteht aus flachen Zellen, die ein

ebenfalls sehr flaches Lumen zwischen sich haben und der Außencuticula unmittelbar benachbart

sind. Auf rasterelektronenmikroskopischen (REM) Präparationen ist dieser Abschnitt oft auf- oder

abgerissen und hinterläßt auf der Cuticula einen etwa rhombenförmigen Abdruck von beträchtlicher

Breite (ca. 120 um). Vermutlich werden in diesem Abschnitt im reifen Weibchen die aus dem Ovar

ausgetretenen Eier gespeichert. Unmittelbar dorsal von ihm liegen die Spermatheken.

Der dritte Oviduktabschnitt (Abb. 1a, 2b) ist durch Vorhandensein der voluminösen Ovidukt-

muskulatur (M. constrictor oviducti, mco, M. dilatator oviducti, mdo) gekennzeichnet, die an einem

im Querschnitt T-förmigen Apodem, dem Notum (no, Gabelapodem), ansetzen. Zunehmende

Mächtigkeit des Epithels läßt eine Art Ventil (v) entstehen (Abb. 1a), das den Durchrritt einzelner

Eier reguliert. Lebendbeobachtungen am narkotisierten Tier machen diese Einzelabgabe von Eiern

unmittelbar deutlich. Da der M. constrictor oviducti schräg am Notum ansetzt (WENSLER & REMPEL

1962, Morısch & WüLker 1987), wird bei seiner Kontraktion nicht nur der Ovidukt verengt, sondern

gleichzeitig das Eı nach hinten gezogen.

Als Vagina (vg, Genitalkammer, Abb. 1a) kann schließlich derjenige Oviduktabschnitt bezeichnet

werden, in dem sich die Übertragung des Samens in die Spermathek (evtl. mit Hilfe einer Spermato-

phore, NiErsen 1959, SAETHER 1977), die Besamung der reifen Eier sowie ihre Versorgung mit Schleim

und ihre Austreibung abspielt.

Die dorsale Wand der Vagina beginnt mit der Spermathekenmündungsplatte smp (pelvis-like struc-

ture Nıeısen 1959, small flat plate WensLer & Rempeı 1962, zeichnerische Darstellung bei Görtz 1964

©.»

Abb. 2: C. anthracinus, Querschnitte durch die weiblichen Gonodukte. a) In Höhe des ersten Oviduktabschnittes,

b) in Höhe des dritten Oviduktabschnittes, c) in Höhe der fingerförmigen Fortsätze (Vagina), d) in Höhe des hin-

teren Teiles der smp (Labien). Schleimdrüse entsprechend ihrem Knick nach hinten zweimal getroffen.

g = Ganglion, h = Hämolymphe, mdo = Musculus dilatator oviducti, mdv = M. dilatator vaginae, ml = M. late-

ralis, mv =M. ventralis,n = Nerv, ra= Rami des Notums, sgb = Subgenitalbucht, tr = „Trichter“. Andere Abkür-

zungen wie in Abb. 1.

und SAETHER 1977). Mit der Dorsalfläche der smp sind oral die Rami des Notums (ra, „Äste des Gabel-

apodems“ Götz 1964), seitlich die Spangen des Coxosternapodems (csa, Lateralspangen bei GöTz) fest

verbunden (Abb. 3a) und bilden mit ihr eine sklerotisierte Einheit. Die Vordergrenze der smp ist

schwer erkennbar, weil die Stärke des Chitins hier kontinuierlich abnimmt. Stärker chitinisiert ist der

beckenähnliche Teil (Abb. 3c), dessen Vorderrand zur Mitte hin dunkler und hakenartig erscheint

(hk, offenbar die „Haken“ bei GöTz, die er jedoch der Subgenitalbucht zurechnet). Nach vorn beglei-

tet aber noch eine schwächere, immer mehr verjüngte Chitinplatte den unpaaren Teil des Notums

(Abb. 3a).

In der Mitte der smp findet sich eine ovale Region, die wir als „Spermathekenmündungsfeld“ (smf,

Abb. 3a, c, d) bezeichnen. Das smf erscheint hell, weil es nur von der durchsichtigen Intima ausgefüllt

ist. Diese springt etwas in die Genitalkammer vor (se, Abb. 3b, „Spermathekeneminenz“ SAETHER

1977) und wird vom unpaaren Spermathekengang durchbrochen. Der Hinterrand des smf ist dorsad

wulstartig verstärkt (Abb. 3a, c). Die hintere Fläche der smp liegt unmittelbar vor der Schleimdrüsen-

öffnung (ggo) und ist räumlich kompliziert gestaltet. Im parasagittalen Längsschnitt (Abb. 1a) sieht

man einen schräg nach unten abgeknickten Teil, der den Ausgang der Schleimdrüse abdeckt. In einem

REM-Präparat, in dem mehr oder minder zufällig die Schleimdrüse (und die Verbindungsmembran

zur Postgenitalplatte) nach hinten abgerissen war (Abb. 3b), ist weiterhin erkennbar, daß in der Mitte

dieser schrägen Fläche ein nach ventral offener konischer Trichter (tr) liegt, dessen Ränder stellen-

weise mit kleinen Zähnchen bestanden sind.

Die Funktion des Komplexes aus smp, Notum und Coxosternapodem muß in bezug auf die Kopu-

lation einerseits, die Besamung und Austreibung des Eies andererseits analysiert werden. Bei der Be-

gattung wird der Penis dem smf zur Spermien-(oder Spermatophoren-)abgabe nahegebracht (Reıss

1966). Von hinten direkt auf das smf gerichtet ist die trichterartige Aussparung im hinteren Teil der

smp, sie könnte dem Penis die nötige Führung geben. Da über der Dorsalseite der smp ein starker

Muskel (M. transversus) die Knickstellen des linken und rechten Coxosternapodems verbindet

(WENSLER & Remper 1962, Morısch & WÜLkER 1987) und die schwach chitinisierte smp sicher elastisch

ist, dürfte der Trichter im Ventralbereich bei der Kontraktion des Muskels sich weiter öffnen

(Abb. 2b, d). Ein anderer Muskel (M. dilatator vaginae, mdv) setzt im wesentlichen an der (nach au-

ßen schrägen) Außenfläche der Rami des Notums (ra) an (Abb. 2c). Bei seiner Kontraktion wird das

smf gedehnt und damit vielleicht die Öffnung des Spermathekenganges zur Besamung des Eies freige-

geben. Möglich ist außerdem, daß bei der erwähnten Kontraktion des M. transversus die Mitte der

smp so nach vorne gestoßen wird, daß diese Bewegung zum Ausstoß des Eies beiträgt. Der M. lateralis

(ml, Abb. 2d), der vom Coxosternapodem zum Hinterrand des 9. Tergits verläuft (WEnster & Rempe

1962, Morısch & WüLker 1987), zieht Coxosternapodem und smp dorsad und macht dementspre-

chend die Genitalkammer für den Eidurchrritt geräumiger. Lebendbeobachtungen haben gezeigt, daß

die Spangen des Coxosternapodems bei der Eiablage lebhaft in deren Rhythmus bewegt werden,

ebenso wie die mit ihnen muskulös verbundenen Cercı, die sich beim Eidurchtritt weit öffnen. Vorher

wird jedoch das Ei noch mit dem quellbaren Schleim umgeben, der für Gelege von Chironomus ty-

pisch ist. Das Epithel um die breite Schleimdrüsenöffnung (ggo) ist an der schrägen Hinterfläche der

smp gewissermaßen ausgespannt (Abb. 3d) und wird nur von häutigen Anhängen dieser Hinterflä-

che, den Labien, median überdeckt. Durch diese Gebilde könnte der in jedem Gelege median verlau-

fende, helixartig gewundene „Zentralfaden“ (Strenzke 1959, „twisted fibres“ MıaLı & HammonD 1900)

bei seinem Austreten aus der Schleimdrüse geführt und von den beidseitigen Schleimströmen umge-

ben werden.

In der ventralen Wand des Ovidukts beginnt am sog. „Bogen“ (Görz 1964, vgl. auch b in Abb. 3c)

ein medianer Spalt, der die beiden Dorsomesalloben (dml) der Gonapophyse VIII voneinander trennt

(REM-Darstellung Morisch & WüLker 1987). In diesem Spalt sind zahlreiche fädige Gebilde, wahr-

scheinlich Sensillen, nachweisbar, deren Funktion wahrscheinlich im Zusammenhang mit der Begat-

tung wichtig ist. In diesem Spalt wird wahrscheinlich der Steg des T-Träger-artigen processus analıs

Abb. 3: Spermathekenmündungsplatte. a) C. anthracinus, Gesamtansicht mit Blickrichtung auf die dorsale

Oberfläche der smp (vom Körperinneren her!). REM nach KOH-Mazeration. Schleimdrüsenrest verdeckt hinte-

ren Teil der smp. b) C. anthracinus, Einblick von hinten in die Genitalkammer, Ventralseite unten. Schleimdrüse

und Verbindungsmembran zur Postgenitalplatte nach hinten abgerissen. c) C. anthracinus, vorderer Teil der smp

in Ventralansicht, KOH-Mazeration. Pfeil = wulstiger Hinterrand des smf. d) C. riparius, hinterer Teil der smp mit

Trichter und Labien, Ventralansicht. Boraxkarminfärbung. Man beachte, daß das vordere Epithel der Schleimdrü-

senöffnung (Pfeil) der Hinterfläche der smp fest anliegt. Gleicher Maßstab wie in c)

bf = Borstenfelder des 8. Abdominalsternites (von innen gesehen), csa = Coxosternapodem, ggo = Schleimdrüsen-

öffnung, hk = „Haken“, Ih = Leibeshöhle, ra = Rami des Notums, se = Spermathekeneminenz, sgb = Subgenital-

bucht, smf = Spermathekenmündungsfeld, tr = „Trichter“, tra = Tracheen um Spermathekengänge. Andere Ab-

kürzungen wie in Abb. 1.

des männlichen Hypopygiums geführt, während dieser in die Subgenitalbucht eindringt (Reıss 1966).

Wenn er sich dem Bogen nähert, ist im Körperinneren der Penis dem smf nahe und der Transfer von

Spermien oder Spermatophoren muß eingeleitet werden. Die fädigen Gebilde erstrecken sich nach

hinten bis zu den bisher unbeschriebenen, in Abb. 2c dargestellten „fingerförmigen Anhängern“ (fp),

auf denen zahlreiche Borsten stehen. Weiter caudad sind dann auf dem parasagittalen Schnitt der

Abb. 1a zwei Loben der Gonapophyse VIII (Dorsomesal- und Apodemlobus, dml, apl) sichtbar. Da

das Ei (Abb. 1a) größer ist als das Lumen der Genitalkammer, klaffen diese Loben bei Eiaustritt weit

auseinander (Lebendbeobachtung). Der dritte Lobus (Ventrolaterallobus, vll) der Gonapophyse, der

lange Borsten trägt, ragt von der Seite in die Genitalkammer. Unsere Arbeitshypothese ist, daß das Ei

in der Genitalkammer durch diese ventralen und seitlichen Stützen (oder Sensoren) in eine Lage ge-

bracht wird, in der die zum Spermieneintritt geeignete Region der Eihülle (Micropyle?) zur Sperma-

thekenöffnung orientiert ist. Ein solcher Vorgang würde den hohen Besamungsprozentsatz (nahe

100) bei Chironomus erklären und auch mit dem bei anderen Dipteren beobachteten Prinzip des

„alignment“ der Eier in Genital- oder Besamungskammern übereinstimmen (z. B. Musca Lzoroıp,

1980, Übersicht Sanper 1985). Auch Wenster & Remper 1962 haben die Ventrolateralloben als „egg-

guides“ angesehen.

Zusammenfassend sind wir der Meinung, daß eine Abschnittsgliederung der Gonodukte von Chr-

ronomus vor allem nach den aufeinanderfolgenden Funktionen (1. Sekretbeigabe ?, 2. Eispeicherung,

3. Durchlaß einzelner Eier, 4. Begattung, Besamung, Schleimzugabe) sinnvoll ist. Auch strukturelle

Grenzen sind, wie WÜLkEr et al. 1979 angedeutet haben, brauchbar, aber eher willkürlich. Die funk-

tionellen Vorstellungen, die wir aus den morphologischen Befunden abgeleitet haben, sind vorwie-

gend Arbeitshypothesen, die mit weiteren Untersuchungen, vor allem Beobachtungen am lebenden

Tier, Klärung der Frage, ob und wo eine Mikropyle vorhanden ist, ultramikroskopische Untersu-

chung der Borsten an den fingerförmigen Fortsätzen und Ventrolateralloben und Beobachtungen

über die zeitliche Reihenfolge von Eireifung, Begattung, Übertritt der Eier in den Oviductus commu-

nıs und Eiablage geprüft werden müssen.

Gonodukte parasitierter Imagines

Schnittserien durch das Abdomenende parasitierter Weibchen führen ebenso wie die äußere In-

spektion der 8. Abdominalsternite (WÜLKER 1961, GöTz 1964) zu sehr unterschiedlichen Ergebnissen;

kein Tier ist dem anderen gleich. Die Skala reicht von noch guter Erkennbarkeit der Gonoduktteile

bis zur Beschränkung auf eine mehr oder weniger undefinierbare Zellmasse am Ende des 8. Abdomi-

nalsternites (vgl. WüLker 1978). Als Beispiel für unsere Befunde zeigt Abb. 1b noch relativ gut struk-

turierte Gonodukte: Der Ovidukt ist verkürzt und erreicht nur etwa die Mitte des 8. Abdominalseg-

mentes. Ein Oviduktlumen ist nur auf manchen Schnitten sichtbar. Der Ovidukt ist in mehrere Aus-

läufer aufgeteilt, der Musculus constrictor oviducti im Umfang reduziert, jedoch in normaler Posi-

tion. Weiter caudal sind linkes und rechtes Coxosternapodem durch die hier verstärkte smp in der

Mitte starr verbunden. Dorsal ist ein kräftiger M. transversus aufgelagert, der hintere Teil der smp ist

meist vorhanden. Bei Betrachtung im REM ist manchmal auch seine trichterförmige Einsenkung

(S. 21) noch erkennbar. Der vordere Teil der smp ist dagegen meist mißgestaltet oder fehlt. Die

Schleimdrüse ist ein begrenzter Zellkomplex. Ventral der Genitalkammer schließlich liegt eine hyper-

trophierte Gonapophyse VIII, deren Dorsomesal- und Apodemlobus nicht mehr klar voneinander

abgrenzbar sind. Bemerkenswert ist die aberrant starke Entwicklung der Chitinintima.

Trotz der Variabilität der Befunde an parasitierten Weibchen ergibt sich als Regel, daß die morpho-

genetischen Mängel der Gonodukte oral beginnen (Ovidukt, Spermatheken) und caudad fortschrei-

ten. Ähnlich hat Baumerr-BenriscH (1960) bei Strepsipterenbefall der Zikade Calligypona nur bei 7

von 44 Imagines den Ovidukt, aber bei 32 die Vagina ausgebildet gefunden.

Histologisch ist erstens davon auszugehen, daß in den wachstumsgehemmten Bereichen der Zellzy-

klus verlangsamt ist (WüLker 1978). Zweitens scheint das normale entwicklungsphysiologische Pro-

gramm der Zellen in Richtung auf aberrante hypertrophe Bildungen geändert. Drittens wird der nor-

male Zellverband gelockert und irregulär. Befunde an Totalpräparaten haben gezeigt (unveröff.), daß

z. B. die Muskulatur von Ovidukt und Vagina, wenn sie beim Wachstum median eine nur noch un-

vollständig eingesenkte Subgenitalbucht mit mangelhaft ausgebildeten Gonodukten „antrifft“, sich

regellos und ohne nennenswerten Zellverbund von einer Segmentseite zur anderen ausspannt („criss-

cross“-Muster der Muskulatur). Zelldegeneration schließlich wurde in unseren Befunden nicht direkt

nachgewiesen, ist aber nach Erfahrungen am geschädigten Ovar (Würker 1971) früher oder später

auch in den geschädigten Gonodukten zu erwarten.

Danksagung

Wir danken Frau R. Rössler für viel technische Hilfe und fruchtbare wissenschaftliche Diskussionen.

Literatur

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BAUMERT-BEHRISCH, A. 1960: Der Einfluß des Strepsipteren-Parasitismus auf die Geschlechtsorgane einer Ho-

moptere. — Zool. Beitr. (N. F.) 6: 85-126, 291—332

CLEMENTS, A. N. 1963: Physiology of mosquitoes. — Pergamon Press, Oxford

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nomus (Dipt.). — Z. Parasitenkd. 24: 484-545

KaıseR, H., WÜLKER, W. & SKOFITSCH, G. 1987: Limnomermis anthracinin. sp. and L. bathophili n. sp., sympatric

species of Mermithidae (Nematoda) in different Chironomid (Diptera) hosts. — Zool. Jb. Syst. 114:141—156

LEOPOLD, R. A. 1980: Accessory reproductive gland involvement with the sperm-egg interaction in muscoid flies.

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land, Amsterdam, pp. 253-270

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don Press, Oxford

MORISCH, U. & WÜLKER, W. 1987: Formation of the cerci, abdominal segment X and postgenital plate in the genital

imaginal discs of female larvae and pupae in Chironomus (Diptera). — Ent. Scand. Suppl. 29:91 —96

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1252-1253

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Anz. 176: 440-449.

REMPEL, J. G. 1940: Intersexuality in Chironomidae induced by nematode parasitism. — J. exp. Zool. 84: 261—289

—— NAYLOR, J. M., ROTHFELS, K. & OTTONEN, B. 1962: The sex chromosome constitution of chironomid inter-

sexes parasitized by nematodes. — Can. J. Gen. Cytol. 4: 92-96

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A.eds) Vol. 2, 409—430

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Bull. Fish. Res. Bd. Can. 197: 209 pp

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STRENZKE, K. 1959: Revision der Gattung Chironomus Meig. I. Die Imagines von 15 norddeutschen Arten und Un-

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—— 1971: Untersuchungen über die Ultrastruktur der Gonaden von Chironomus (Dipt.) 2. Ovarstruktur nach

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Prof. Dr. Wolfgang Wülker,

Irıs Herrmann,

Institut für Biologie I (Zoologie) der Universität

Albertstr. 21a,

D-7800 Freiburg ı. Br.

SPIXIANA | Supplement14 | 25-28 München, 15. Juli 1988 | ISSN 0177-7424

Thalassosmittia atlantica (Storä) comb. nov. Description of adult

female and immature stages from Tenerife, Canary Islands

(Diptera, Chironomidae)

By P.D. Armitage and J. Tuiskunen

Abstract

Descriptions of adult female, pupa and larva of the species Eukiefferiella atlantica Storä are presented. On theba-

sis of these new data and re-examination of adult males the species is transferred to Thalassosmittia.

Introduction

Arnrtace (1986) redescribed the male of Eukiefferiella atlantica Storä from type material and speci-

mens collected in December 1983 over rock pools at El Medano, Tenerife. Generic placement of the

species was however still in doubt and further data were required to confirm its identity. In December

1985 collections at the same site were made by P. D. A. in and around rock pools. These collections

contained E. atlantica males, together with associated females which keyed-out to Thalassosmittia

Strenzke & Remmert with Sarther (1977) although the Tenerife specimens had finely pubescent eyes.

In addition, pupal exuviae and last ınstar larvae associated with the adults were also identified as Tha-

lassosmittia from descriptions given in STRENZKE & REemmerT (1957). Further evidence for thenew com-

bination ıs provided by adult males of E. atlantica which key-out to Thalassosmittia with the most re-

cent key to holarctic adult male Orthocladiinae (Cransrton et al., in prep.). The species atlantica Storä

is therefore transterred to Thalassosmittia.

Thalassosmittia atlantıica (Stora) comb. nov.

(Figs. 1-11)

Eukiefferiella atlantıca Storä (1936). Lectotype (©* imago, specimen A) Tenerife, Puerto Cruz, coll. R. Frey

28.7.1931. Slide mounted lectotype deposited in Zoological Museum, Helsinki, Finland.

Adult female (n = 6)

Colour: Abdomen and ground colour of thorax greenish grey. Scutal stripes well separated, brown.

Wings pale brown.

Head: Temporal setae 4-5, clypeals 17-25. Eyes not produced dorsally, with pubescence. Palps

5-segmented, lengths of 4 ultimate segments: 18—26 um, 38—50 um, 44—62 um, 74—88 um. Sensilla

clavata of segment 3 lacking. Antennal flagellum (Fig. 1) 5-segmented, lengths of segments:

48-60 um, 30—40 um, 32—42 um, 32—42 um, 68-86 um. AR 0.46—0.52. Sensillar setae present on

all tlagellar segments.

500

Figs 1-11. Thalassosmittia atlantica (Storä): adult female, 1 Antenna, 2 Thorax lateral, 3 Wing. 4 Genitalia; pupa,

5 Tergites II-VIII. 6 Tergite IX and genital sac (C’); larva, 7 Antenna, 8 Seta interna of mandible, 9 Premandible,

10 Mandible, 11 Mentum. (Bar lines in um.)

Thorax: (Fig. 2). Antepronotals 0 (rarely 1). Some very weak and curved acrostichals present in an-

terior /s—1/4 of thorax. Dorsocentrals 7—11, uniserial, arising from pale spots. Prealars 3, scutellars

6-8 in one row.

Wing (Fig. 3): Membrane without macrotrichia. Granulation fairly coarse, visible at a magnifica-

tion of X100. Veins bare except R wıth 4-8, R, with 2-4 and R, ‚; with 7-10 setae. Brachiolum with

1 seta. Squama bare. Costa extending beyond R,; ;, free end about 70-100 um. R,,, poorly separated,

reaching wing margin near the tip of Ry; 5. Cu; slightly curved. Anallobe moderately developed. Wing

length 1.05-1.24.

Legs: LR (P,) 0.44—0.49, P, 0.44—0.45, P; 0.52—0.57. Alltibiae with one spur. Length of front tibial

spur 20-22 um, mid tibial spur 20-22 um and hind tibial spur 22-26 um. Hind tibial comb with

9—11 setae. Pulvilli lacking. BR of hind tibıa 2-3.

Genitalıa (Fig. 4): Gonocoxite with 3—6 longer and 3-7 shorter setae. Cercus 50-80 um long. Se-

minal capsule oval, with indistinct neck, 75—80 um long and 46-56 um broad. Seminal duct with a

loop.

Pupa (n = 2)

Total length 2.0-2.3 mm. Colour: pale brown, thorax somewhat darker than abdomen.

Cephalothorax: Thoracic horn absent. Frontal setae obviously lacking (not identifiable on both

slides examıned). Frontal warts normally developed. One postorbital seta present. Verticals absent.

2 moderately developed medial antepronotals, 3 precorneals of about equal length. 4 dorsocentrals,

2 of which very short. Thorax somewhat rugulose dorsally. Wing sheaths smooth.

Abdomen: Tergite I without shagreenation. Spinules of tergites II—VIIl as in Fig. 5. Medial spinule

group very weak, lacking on tergite II-IV but present on V-VIII. Spinules of anterior group directed

backwards but those of posterior group forward, consisting of 2-4 rows of spinules. Spinules of an-

terior and posterior groups distinctly weaker in segment VIII than in other segments. Sternites I-IV

without shagreenation. Sternites V- VIII with 2-3 rows of moderately strong spinules near posterior

margin of segment, and a group of very weak, irregularly arranged spinules on the anterior part of ster-

nites. CoFFMAN etal. (1986) refer to conjunctives of both tergites and sternites. These do not occur in

the present specimens and are not shown in the figure of Thalassosmittia in LanGgron (1984). Poorly

developed pedes spurii A present on sternites II-VI. Pedes spurii B absent.

Chaetotaxy of abdomen: Dorsal setae of segment II-VII as in Fig. 5. I—VIII with 3 weak, VIII

with 2 somewhat longer setae. Setae of tergite I not visible on examined exuviae. Lateral setae ob-

viously 1 on each segment. Ventral setae 3 in each sternite except VIII with 1. Tergite IX (Fig. 6) more

or less rectangular, with posteriorly directed spinules on anterior half and with 2 moderately long se-

tae. Genital sac with ventral groups of well developed spinules.

Larva (n = 2) (fourth instar)

Body length 2.4-2.7 mm. Head capsule 280-360 um long. Body colour (after preservation) pale,

head capsule brown with mentum, mandibles and caudal margins darker brown.

Head: Antenna (Fig. 7) 5-segmented, lengths of segments: 16 um, 8-10 um, 4—6 um, 5—6 um.

AR 1.63. Antennal blade 20 um long, antennal style 7 um long. Labrum: SI setae with about 6 bran-

ches, other S-setae simple. Pecten epipharyngis indistinct in preparations. Premandible (Fig. 9)

54-66 um long, dark brown apically, pale basally. Mandible as in Fig. 10, 100-120 um long, with 5

teeth. Seta interna of mandible as ın Fig. 8, with 4-5 branches. Mentum (Fig. 11) with single pointed

(orrounded) median tooth and 4 pairs of more or less pointed lateral teeth. At sides of the mentum a

pair of toothlike structures arısing from the ventral surface. Width of mentum 84—90 um.

Body: Parapods present divided and short. Anterior parapods with both simple and serrate claws.

Posterior parapods with simple claws only. Procercus absent. Remaining features of anal segment not

clear in specimens available.

Discussion

Thalassosmittia was erected by STRENZKE & RemMmERT (1957) for the species thalassophila Bequaert &

Goetghebuer. The genus also includes 3 Nearctic species formerly in Camptocladius van der Wulp,

pacıificus Saunders 1928, marinus Saunders 1928 and clavicornis Saunders 1928, and a species from Ja-

pan nemalione Tokunaga 1936 formerly in Spaniotoma (Smittia). To these five holarctic species can

now be added T. atlantıca (Storä) which is known at present from the Canary Isles, the Azores and

Madeira. Male adults of atlantica may be separated from the other palaearctic species T. thalassophila

on the hypopygium which ıs distinctive in T. thalassophila. Other points of difference are the A.R.

(0.68 ın thalassophila, 0.29 ın atlantica) and the eyes which are bare in thalassophila and finely pubes-

cent in atlantica (ArmıtaGe 1986).

The ecology and lıfe histories of the 3 Nearctic species are discussed in Mor£y & Rıng (1972b) and

a key to these species (as Saunderia Sublette 1967, spp.) is presented in Morı£Y & Rıng (1972a). Toku-

naGA (1936) provides detailed descriptions of all life history stages of T. nemalione which is found in

algal mats in the tidal zone of rocky shores in Japan. STRENZKE & REmMERT (1957) give similar informa-

tion for the western palaearctic species T. thalassophila. All6 Holarctic species are found in the marine

littoral zone. The genus is widely distributed in the Holarctic region and further collections in the

southern hemisphere may reveal a worldwide distribution.

Acknowledgements

We are most grateful to Dr. P. Cranston (BMNH), Dr. C. Pinder (FBA) and Bernhard Lindeberg (Zoological

Museum, Helsinki) for their help and encouragement.

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Dr. Patrick D. Armitage, Dr. Jarı Tuiskunen,

Freshwater Biological Association, River Laboratory, Zoological Museum, University of Helsinki,

East Stoke, Wareham, Dorset BH20 6BB, Great Britain. P. Rautatiekatu 13, SF-00100 Helsinki 10, Finland.

| SPIXIANA Supplement 14 | 2953 | München, 15. Juli 1988 ISSN 0177— 7424

Some records of two rare chironomid species in the Netherlands

(Diptera, Chironomidae)

By B. P.M. Krebs“

Abstract

Recent finds of two rare chironomid species in the Netherlands are described. Dicrotendipes pallidicornis Goetgh.

has been found in two small brackish waters in the southwest of the Netherlands. Parachironomus mauricü Kruse-

man was collected from a freshwater ditch in the north of the Netherlands. Both finds are compared with reports

from elsewhere in and outside Europe.

Introduction

In this article the Dutch records of the rare Chironomid species Dicrotendipes pallidicornis Goet-

ghebuer and Parachironomus mauricii Kruseman are considered. D. pallidicornis ıs a new find for the

Netherlands. This species was found at two places ın the Delta-region in the south-west of the Nether-

lands.

Since the description of P. mauricu by Kruseman ın 1933, this species was not found in the Nether-

lands until 1979 when it was rediscovered by dr. W. van Vierssen in Friesland in the northern part of

the Netherlands (Van Vıerssen, 1982). Both species will be dealt with here.

Dicrotendipes pallidicornis Goetgh.

D. pallidicornis was first described by GoETGHEBUER (1934) on the basis of material collected on 13

to 15 April 1926 at Basra, Iraq, close to the Persian Gulf. Since then the species has been reported from

several European countries. In the collection of the Zoologische Staatssammlung in Munich, FRG, I

found some specimens from several parts of Europe, as indicated below (a to c):

a) One came from Lake Kurnas on Crete, Greece, collected by Dr. H. Malicky (Biological Station

Lunz, Austria) on 16-5-1971. Dr. Malicky kindly supplied some chemical information about Lake

Kurnas: it ıs an oligotrophic brackish water lake with a chlorinity that varıed between 4.8 and 8.0 %oo

CI during the period 1972-1980. The find of an adult doesn’t necessarily indicate that it came from

this lake.

b) The second specimen was one collected by Setta (Mendl) on 30—-9— 1975 at Lake Pineto, near Ba-

stia, Corsica. About this find no further details are known.

c) On 7-8-1978 R. Kühbandner caught one male on the surface of a freshwater pond near Novalja

on the island Pag, Yugoslavia. As above, this doesn’t indicate that the midge grew up in this lake as a

larva.

* Communication nr. 387, Delta Institute for Hydrobiological Research, Yerseke.

The following references were found in the literature. PrAar (1980) reports the species from the Cala

reservoir near Sevilla in the south of Spain. It is a male, caught on 16-3-1974 and was by that the first

known Spanish record of this species. Birk£rr (1984) described some finds of England. On 19 August

and 1 September 1975 he visited the sandhills at Sandscale Hawes Warren in Cumbria and caught ma-

les and females. The species is established in this part of north-west England, shown by frequent sub-

sequent captures. Since then the species has been found by Langton in Swan Pool, near Falmouth,

Cornwall, south-west England in a partly brackish pool. Birkett’s conclusion is that these records sug-

gest a discontinuous distribution at present, and he is expecting that further collecting in and around

coastal dune slacks may reveal the species to be more widespread.

The finds for the Netherlands are listed below:

a) The first sampling-station was the Doolman, a “weel” on Zuid-Beveland in the Delta-region. A

“weel” or “wiel” is around pool which is a result of a dikeburst. The Doolman has a diameter of ap-

prox. 50 meters and its largest depth is 5 meters. It contains brackish water; the chlorinity in 1978 fluc-

tuated between 1.70 and 1.90 Yo Cl . In 1975 fluctuations were somewhat larger: 2.60-5.90 Yo CI .

In the deeper parts of the pool a vertical salt gradient existed. Locally, the shallower parts were grown

with reeds and the substrate consisted of soft mud. D. pallidicornis was found while sampling this

shallow litoral zone on 6-4-1978 and 11-7-1978. On these sampling days the chlorinity was 1.90 and

1.70 Yo Cl ‚respectively. Collected larvae were reared in the laboratory. Identifications of the midges

were confırmed by dr. R. Lichtenberg (Naturhistorisches Museum, Vienna). D. pallidicornis from 6-

4-1978 was found in combination with larvae of Procladins choreus (Meigen) and Glyptotendipes bar-

bipes (Staeger). For the 11-7-1978 sample Chironomus salinarius Kieffer, Chironomus annularius Mei-

gen, Chironomus halophilus Kieffer and Cricotopus ornatus (Meigen) were additional to the combina-

tion mentioned before.

b) The second sampling-station was a in 1975 newly created pool in the recreational area “Het Poel-

bos”, also on Zuid-Beveland. This pool, which was sampled on 15-3-1978, is rather shallow with reeds

Phragmites australis (Cav.) Trin. ex Steud, salt-marsh club-rush Scirpus maritimus L. and bulrush

S. lacustris L. growing on its shore. The submerged vegetation consisted largely of Ranunculus sp., Po-

tamogeton pectinatus L. and Myriophyllum spicatum L. Chlorinity at that time was 0.92 %o. Again,

collected larvae were reared in the laboratory. Besides D. pallidicornis, we identified G. barbipes,

C. halophilus, C. annularius, C. salinarius, C. plumosus (L.), Microchironomus deribae (Freeman),

C. ornatus and P. choreus, a species composition that shows close resemblance to the combination

found in the Doolman.

Table 1

Species Brackish Freshwater Freshwater Reference

water- species with species with

species moderate to- slight tole-

lerance for rance for

brackish brackish

water water

Chironomus halophilus PARMA & KREBS, 1977

PARMA & KrEBs, 1977

PARMA & KREBS, 1977

PARMA & Kress, 1977

PARMA & KrEBs, 1977

KreEps, 1979

KreEps, 1982

KRrEBs, 1982

Chironomus annularius

Chironomus salinarius

Chironomus plumosus

Glyptotendipes barbipes

Microchironomus deribae

Cricotopus ornatus

Procladius choreus

Both the additional species from the Doolman, as well as those from the Poelbos are rather common

in the southwestern part of the Netherlands and typical for oligohaline-mesohaline waters (Table 1).

For the species composition of the accompanying macrofauna see Table 3.

From the above, the tentative conclusion can be that D. pallıdicornis shows a preference for slightly

brackish water. The question, whether it is a true brackish water species or a freshwater species with

a tolerance for brackish water has to remain unanswered due to a lack of sufficient data. The Dutch

and English data do suggest the former.

Fırrkau & Reıss (1978) classified D. pallidicornis as being a species of stagnant lakes. On basıs of the

Dutch records, we could add to this the smaller (brackish) waters. D. pallidicornis is likely to have at

least two generations a year, because larvae of the species were found both on spring (March) and on

summer (July).

Parachironomus mauricii Kruseman

W. van Vierssen (Landbouwhogeschool, Wageningen, The Netherlands) sent me some chirono-

mids reared from larvae collected in several pools and ditches in Friesland in the norther part of the

Netherlands. One of these samples was especially interesting since ıt contained P. manricı.

This species has originally been described by GoETGHEBUER (1931) as Cryptochironomus lıttorellus.

The description being based on a specimen collected on 8-9-1931 at “de Panne”, a dune area in the

southwest of Belgium. The species name “littorellus” gave rise to some confusion since Meigen used

it also for another species. Therefore Kruseman (1933) renamed the species Parachironomus manrıcn,

“mauricii” referring to Goetghebuer’s Christian name.

For the Netherlands, Kruseman reports 2 finds. One concerns a specimen from his own collection

and collected in the dunes near Vogelenzang in May 1931; the other one being a specimen from the

“de Meijere-collection” and caught near Diemen, Noord-Holland, in September 1920.

LeHmann (1970) reports the species ın his revision of the European Parachironomus species. Besides

the finds from Belgium and the Netherlands, Lehmann reports the species for Germany as well

(though with a questionmark) and concludes that nothing is known about its lifecyle.

In the collection of the Zoologische Staatssammlung Munich two male specimens were found. One

of these came from the Netherlands and was identified by Kruseman as Parachironomus varus limnaei

Guibe, but renamed by Reiss as P. maurıcii. About the origin of this material no further details are

known. The second specimen, collected by Prof. Feliksiak in 1954 came from the region of Lodz, Po-

land. About this find we have no additional information either.

P. mauricü is also known from England. Langton found this species in Flood’s Ferry, near March

in Cambridgeshire (Cranston, 1974).

The first record for the Sovietunion comes from SHıLovA (1976) who reared 4 males from larvae col-

lected in an artificial pond with a depth of 70 cm and amuddy, sandy bottom. The pupae hatched by

the end of May and July.

Table 2

Potamogeton pectinatusL. 25 %

Zannichellia pedunculata Rchb. 20%

Lemna gibbaL. 20%

Lemna trisulca L. 10%

Potamogeton pusıllus L. 10%

Green algae 10%

As stated above, the most recent Dutch find was done in Friesland by W. van Vierssen on the 22rd

of August, 1979. Larvae of P. mauricii were collected from a small ditch, running through arable land.

Its breadth was 2.5 m, and depth approx. 40 cm. The substrate consisted of clay. The water was very

clear and its level did not fluctuate. The vegetation is given in Table 2 (after van Vierssen, 1982). For

the species composition of the accompanying macrofauna see Table 4.

Unfortunately, the chlorinity is unknown, but regarding the species composition of the vegetation,

the water will have been fresh or slightly brackish at the most. Rearing of larvae resulted in 2 males and

6 females. Other species either were not present or did not hatch. After identification the midges were

compared with specimens from the collection in Munich.

Like D. pallidicornis, Fırrkau & Reıss (1978) consider P. mauricii to be a stagnant lake species. On

basis ofthe concise data given above it would be more likely to consider P. mauricii as a species typical

Table 3

Species composition of the accompanying macrofauna at the sampling locations of Dicrotendipes pallidicornis.

Doolman Doolman Poelbos

64-1978 11-7-1978 15-3-1978

Plea leachi McGreg. & Kirk x

Callicorixa concinna (Fieber) x x x

Callicorixa praeusta (Fieber) x

Sıgara stagnalıs (Leach) x x x

Sigara lateralis (Leach) x x x

Sıgara striata (L.) x x x

Corixa punctata (lllig.) x

Halıplus lineatocollis Marsh. x

Anacaena globulus (Payk.) x

Procladius choreus (Meigen) x x x

Chironomus salinarius Kieff. x x

Chironomus halophilus Kieff. x x

Chironomus annularius Meig. x x

Chironomus plumosus (L.) x

Glyptotendipes barbipes (Staeger) x x x

Microchironomus deribae (Freeman) x

Cricotopus ornatus (Meigen) x x

Limnephilus affınis Curtis x

Ischnura elegans v.d.L. x x

Electra crustulenta (Pallas) x

Hydrobia sp. x

Lymnaea palustris (Müller) x

Neomysis integer (Leach) x x

Corophium sp. x

Orchestia gammarellus (Pallas) x

Gammarus duebeni Lilljeborg x x

Gammarus zaddachi Sexton x x x

Palaemonetes varians (Leach) x "x

for small fresh waters (Kleingewässer). P. mauricü ıs mostly found in the northern-German lowlands

and their offshoots. Regarding the finds of Kruseman in May and September, of van Vierssen in Au-

gust and of Shilova by the end of May and July, the species can at least be considered bivoltine.

Table 4

Species composition of the accompanying macrofauna at the sampling location of Parachironomus mauricü

Coleoptera Heteroptera Mollusca

dominant Sigara striata (Fıeb.) Lymnaea peregra (Müller)

Hygrotus inaequalıs (Fabr.) Sigara stagnalıs (Leach) Lymnaea palustris (Müller)

Laccobius minutus (L.) Sıgara falleni (Fieb.) Lymnaea stagnalıs (L.)

frequent Corixa punctata (11l.) Planorbis planorbis (L.)

Haliplus lineatocollis Marsh. Corixa affınıs Leach Planorbis corneus (L.)

Haliplus immaculatns Gerh. Corixa panzeri (Fieb.) Bithynia tentaculata (L.)

Hesperocorixa linnei (Fıieb.) Physa fontinalıs (L.)

scarce

Laccophilus hyalınus (Deg.)

Gerris thoracicus Schumm.

Notonecta glauca L.

References

BIRKETT, N. L. 1984: Some unusual brackish water chironomids (Diptera) from Cumbria. — Entomol. Gaz. 35 (3):

197-198

CRANSTON, P. S. 1974: Corrections and additions to the list of British Chironomidae (Diptera). — Entomologist’s

Mon. Mag. 110: 87—95

FITTKAU, E. J. & F. Reıss 1978, in Illies, J.: Limnofauna Europa, Chironomidae: 404—440. — Gustav Fischer Ver-

lag, Stuttgart and Swets & Zeitlinger B. V., A’dam

GOETGHEBUER, M. 1931: Ceratopogonidae et Chironomidae nouveaux d’Europe. — Bull. Annls Soc. r. ent. Belge

71: 211-218

—— 1934: Zur Erforschung des Persischen Golfes, Beitrag Nr. 15; Ceratopogonidae et Chironomidae. — Arb.

morph. taxon. Ent. Berl. 1: 36-39

Kress, B. P. M. 1979: Microchironomus deribae (Freeman, 1957) (Diptera, Chironomidae) in the Delta region of

the Netherlands. — Hydrobiol. Bull. 13: 144—151

—— 1982: Chironomid communities of brackish inland waters. — Chironomus 2: 19—23

KRUSEMAN, G. 1933: Tendipedidae Neerlandicae. Pars I. Genus Tendipes cum generibus finitimis. — Tijdschr. Ent.

76: 119-216

LEHMANN, J. 1970: Revision der europäischen Arten (Imagines O’C’) der Gattung Parachironomus Lenz (Dipt.,

Chironomidae). — Hydrobiologia 33: 129—158

PARMA, S. & B.P. M. KreBs 1977: The distribution of chironomid larvae in relation to chloride concentration in a

brackish water region of the Netherlands. — Hydrobiologia 52: 117-126

PRAT, N. 1980: Quironömidos de los embalses Espanoles (2.° parte). — Graellsia 34: 59—119

SHILOVA, A. I. 1976: Khironomidy Rybinskogo Vodokhranilishcha. (Chironomiden des Rybinsker Stausees). —

Izd. Nauka, Leningrad 1976, 249 pp.

VAN VIERSSEN, W. 1982: The ecology of communities dominated by Zannichellia taxa in Western Europe. II. Di-

stribution, synecology and productivity aspects in relation to environmental factors. — Aquatic Botany 13:

385—483

Bernard P.M. Krebs

Delta Institute for Hydrobiological Research

Vierstraat 28

4401 EA Yerseke

The Netherlands

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SPIXIANA Supplement 14 | 35—44 | München, 15. Juli 1988 | ISSN 0177-7424

Die Gattung Kloosia Kruseman, 1933 mit der Neubeschreibung

zweier Arten

(Diptera, Chironomidae)

Von Friedrich Reiss

Abstract

Through the description of Kloosia africana spec. nov. (C’ imago and pupa) from Kenya, Kloosia koreana spec.

nov. (© imago) from North Korea and additional unassociated exuviae of a probable new species from southwest

China, the previous Holarctic distribution of the genus Kloosia is extended south to tropical Africa and to the

Oriental Region. The collection of O’ pupae allows positive association between imagines and pupal exuviae and in-

direct association with larvae. New synonymies and records presented herein document that Xloosza pusilla (Linne)

is amember of the sand-bottom fauna of large rivers. Larvae of other congeners quite likely occur in the same habı-

tat.

Einleitung

Seit ihrer Beschreibung stellte die monotypische und kaum zitierte Chironomini-Gattung Kloosia

Kruseman, 1933, von der nur ©’ Imagines bekannt waren, ein recht fragliches Taxon mit ungeklärten

Verwandtschaftsbeziehungen dar. Erst ın allerjüngster Zeit konnte SAETHER (1986, 1987) durch die

Synonymisierung von Kloosia mit der ebenfalls monotypischen, nearktischen Gattung Oschia Saet-

her, 1983 belegen, daß Kloosia einerseits dem Harnischia-Gattungskomplex angehört und anderer-

seits mit nunmehr 2 Arten holarktisch verbreitet ist.

Eine weitere offene Frage waren die unbekannten Jugendstadien von Kloosza und ihre ökologischen

Bindungen. Den ersten Hinweis hierzu gab kürzlich Kıınk (1985) mit der Vermutung, daß K. pusilla

eine potamale Art mit einer von Pacast (1936) beschriebenen Larve und Puppe sei.

Im Verlauf der vorliegenden Untersuchungen erwiesen sich die vermuteten und morphologisch

auffälligen Jugendstadien von K. pusilla als mehrfach beschrieben und konnten durch © Puppen der

Art sicher zugeordnet werden. Das dabei gewonnene Gattungskonzept für ©’ Imagines und Puppen

erlaubte es zudem, unbeschriebene Taxa fraglicher Gattungszugehörigkeit aus den Beständen der

Zoologischen Staatssammlung München der Gattung Kloosia zuzuordnen.

Herrn Prof. ©. A. Saether, Bergen, danke ich für die Einsicht in das zitierte unveröffentlichte Manuskript, Herrn

Dr. H. Laville, Toulouse, für die Bereitstellung © Puppen von Kloosza pusılla.

Kloosia Kruseman, 1933

Differentialdiagnose

Imago Cd:

Untere Volsella des Hypopysgs nicht verkürzt, schmal und schlank, das Distalende der Analspitze

erreichend oder dieses weit überragend. Bei allen anderen Gattungen des Harnischia-Komplexes fehlt

die untere Volsella völlig oder ist unterschiedlich stark verkürzt und endet weit vor dem Distalende

der Analspitze. Von den anderen Chironomini-Gattungen unterscheidet sich Kloosia durch das völ-

lige Fehlen von Makrosetae auf der unteren Volsella.

Puppe: |

Die posteriore Hakenreihe auf Tergit II ist median um etwa die Länge einer Hälfte unterbrochen.

Eine solch weit unterbrochene Hakenreihe besitzen unter den Chironomini nur noch die Gattungen

Chernovsküa Saether und Beckidia Saether (vgl. Pınper & Reıss 1986, Tafel 10.4 und 10.5). Von diesen

unterscheiden sich Kloossa-Puppen durch je eine anteriore und posteriore, zum Teil unterbrochene

Querreihe langer heller Dornen auf den Abdominalsterniten I-III. Außerdem hat Kloosia ein schma-

les posteriomedianes Feld dicht stehender, grober, kurzer oder langer Spitzen auf Tergit VI, an das

sich oralwärts ein breiter werdendes Feld zerstreut stehender Kurzspitzen anschließt, so daß ein Cha-

grinfeld in der Form eines umgekehrten Dreiecks entsteht. Die beiden anderen Gattungen haben ein

rechteckiges Chagrinfeld, das sich analwärts nicht verjüngt. Außerdem können bei Kloosia auf den

Tergiten II-V auffällig kammartige, posteriore Querbänder mit sehr langen Spitzen auftreten. Becki-

dia-Puppen sind zudem einfach an den gut entwickelten Pedes spurii B der Segmente II, die dicht mit

kurzen Dornen besetzt sind, zu erkennen. Bedornte Pedes spurii B kommen bei keiner anderen Chi-

ronomini-Gattung vor. Hinzu kommt, daß bei Chernovskiia am Cephalothorax sowohl die Frontal-

als auch die Dorsocentralborsten fehlen, die bei den beiden anderen Gattungen vorhanden sind.

Weitere geringe, differentialdiagnostisch fragliche Unterschiede finden sich bei den drei genannten

Gattungen in der Zahl der lateralen Setae an den Abdominalsegmenten II-IV, in der Chagrinierung

der Tergite I, VII und VIII sowie im Auftreten der dorsalen Setae der Schwimmplatte.

Larve:

Da kein Material zur Untersuchung verfügbar war, wird auf SAETHER (1987) verwiesen.

Kloosia koreana spec. nov.

Chironomini gen. K 1 (Reıss 1980: 147, 149)

Imago Cd:

Größe: Sehr kleine Art, Flügellänge 1,1— 1,2 mm.

Färbung: Körper durch Alkoholfixierung ausgebleicht, gelbbraun. Thoraxfärbung nicht klar zu er-

kennen, vermutlich Vittae und Postnotum rotbraun. Tarsenglieder 2—5 aller Beinpaare vermutlich

angedunkelt.

Antenne: Mit 11 Flagellomeren. AR = 1,25— 1,34.

Kopf: Augen dorsal kräftig stegartig verlängert. Mit 8-10 Clypeusborsten und 8—9 Vertexborsten.

Stirnzapfen fehlen. Länge der Palpenglieder 2-5 in um (Holotypus): 30, 68, 98, 130.

Thorax: Mit 3-4 Acrostichal-, 6 Dorsocentral-, 3 Praealar- und 4 Scutellumborsten.

Flügel: Alle Adern, mit Ausnahme von 2 proximalen Setae auf R, nackt. Squama ebenfalls nackt.

Beine: Länge der Glieder in um (Holotypus):

Fe Ti Ta, Ta, Ta; Ta; Tas

P| 430 350 480 235 180 115 70

Pıı 410 360 175 90 75 50 40

Pın 420 470 260 145 125 85 60

LR = 1,37— 1,42. Kämme der Mittel- und Hintertibien schmal getrennt und mit je 1 Sporn. Pulvil-

len fast so lang wie die Klauen.

Abb. 1: Kloosia koreana spec. nov., Hypopyg dorsal.

Hypopyg (Abb. 1): Analtergitbänder verschmolzen, Y-förmig, medianer Ast die Basıs der Anal-

spitze erreichend. Mediane Analtergitborsten fehlen, apıkale Analtergitborsten lang, lateralwärts kür-

zer werdend. Analspitze mittellang, schlank, distal verbreitert und stumpf gerundet; nur schwach

ventralwärts gekrümmt. Obere Volsella schlank, lang, schwach S-förmig gebogen und die Analspitze

deutlich überragend; nackt bis auf 2 mediane, weit auseinanderstehende und oralwärts weisende Setae.

Untere Volsella schlank, gerade, subapikal leicht verbreitert und die Analspitze ebenfalls deutlich

überragend; dicht mit auffallend langen Mikrotrichien bestanden, Makrosetae fehlen vollständig.

Obere und untere Volsella etwa auf gleicher Höhe endend. Gonostylus schlank, lang, medianwärts

gekrümmt und distal nicht verdickt; Innenkontur mit 8-9 zarten, geraden Setae besetzt.

Imago 9, Puppe und Larve: Unbekannt.

Material: Holotypus 1 © Imago, Nordkorea, Provinz South Pyongan, Pyongyan, Pyongyan-Hotel-Garten,

Malaise-Falle, 2.9.1971, leg. S. Horvatovich & J. Papp (Probennummer 234 in Papp & HORVATOVICH 1972); das

Euparal-Dauerpräparat befindet sich im Ungarischen Naturwissenschaftlichen Museum Budapest. Paratypen der-

selben Herkunft befinden sich, teils als Dauerpräparate, teils alkoholkonserviert, im Museum Budapest und in der

Zoologischen Staatssammlung München: 4.-5.8. (Probennummer 142), 7.-8.8. (150), 9.-10.8. (156) und

31.8.1971 (225).

Differentialdiagnose

Die neue Art Kloosia koreana unterscheidet sich von den anderen Gattungsvertretern durch Hypo-

pygmerkmale: Obere und untere Volsella etwa gleich lang, die Analspitze deutlich überragend; obere

Volsella auf der Innenkontur mit 2 weit auseinander stehenden, oralwärts gerichteten Setae. Die ande-

ren Arten haben unterschiedlich lange Volsellae, die obere Volsella ist anders geformt, ihre beiden me-

dianen Setae stehen eng zusammen und zumindest eine davon ist nicht oralwärts gerichtet.

Verbreitung und Ökologie

Kloosia koreana ist bisher nur vom locus typicus bekannt geworden. Die vorliegenden relativ zahl-

reichen ©’ Imagines verteilen sich über die gesamte Aufenthaltszeit der 2. Ungarischen Nordkorea-

Expedition am Fundort, die von Anfang August bis Mitte September reichte. Demnach ist anzuneh-

men, daß die Art eine lange, diesen Zeitraum überschreitende Flugzeit besitzt. Das gleichzeitige Auf-

treten von potamalen Arten, wie Rheopelopia ornata (Meigen) und Robackia pilicanda Saether in den

Proben macht es wahrscheinlich, daß auch die Jugendstadien von X. koreana im nahegelegenen Fluß

Te-dong siedeln, was der allgemeinen ökologischen Charakterisierung der Gattung Kloosia als pota-

males Faunenelement entsprechen würde.

Kloosia afrıcana spec. nov.

Imago d:

Größe: Sehr kleine Art, geschätzte Flügellänge 1,1 mm (nach Vergleich mit der pupalen Flügel-

scheidenlänge von K. pusilla).

Färbung: Körper in alkoholfixiertem Zustand bei schlüpfreifer Imago gelbbraun, Scutum mit scharf

getrennten braunen Vittae, braunem Postnotum und Praeepisternum. Scapus ebenfalls braun. Beine

ohne erkennbare Farbmuster.

Antenne: Mit 11 Flagellomeren. AR = 1,05.

Kopf: Augen dorsal kräftig stegartig verlängert. Mit 10 Clypeusborsten und ca. 5 Vertexborsten.

Stirnzapfen fehlen. Länge der Palpenglieder 2-5 in um: 20, 57, 71, 93.

Thorax: Mit 6 Acrostichal-, 7 Dorsocentral-, 3 Praealar- und 6 Scutellumborsten.

Flügel: Nicht entfaltet.

Beine: Länge der Glieder in um (in den pupalen Beinscheiden):

Fe Ti Ta, Ta, Ta; Tas Ta;

P} 300 265 325 150 140 100 75

Pı —310 290 150 75 60 40 40

Pın -350 370 —190 120 210) 65 65

LR = 1,23. Kämme der Mittel- und Hintertibien schmal getrennt und mit je 1 Sporn. Pulvillen lap-

penförmig, etwa 2/3 so lang wie die Klauen.

Hypopys (Abb. 2): Analtergitbänder kurz, weit getrennt. Zwei mediane lange Analtergitborsten,

die nicht deutlich von den etwa ebenso langen 13 apikalen Analtergitborsten getrennt sind. Analspitze

fast paralellseitig, mittellang, distal stumpf gerundet und nur schwach ventralwärts gekrümmt. Obere

Volsella nackt bis auf 2 apikale Setae, schlank, Distalhälfte medianwärts geknickt und damit im Umriß

golfschlägerförmig. Die Analspitze überragt die obere Volsella. Untere Volsella ebenfalls schlank,

leicht einwärts gekrümmt und dicht mit auffallend langen Mikrotrichien bestanden; Makrosetae feh-

len gänzlich. Analspitze auf gleicher Höhe mit unterer Volsella endend. Gonostylus schlank, lang,

medianwärts gekrümmt und distal deutlich keulenförmig verdickt; Innenkontur mit 9 zarten, geraden

Setae.

Imago P: Unbekannt.

Puppe:

Größe: Sehr klein, Länge 2,8—4,0 mm (n = 6).

Färbung: Cephalothorax und Abdominalsegmente I-V blaßbraun, restliches Abdomen farblos.

Cephalothorax (Abb. 3c): Oralhörnchen flach gerundet, apıkal mit einem kurzen spitzen Dorn.

Frontalborsten zart, einfach und mittellang. Frontalwarzen fehlen. Thorakalhorn büschelförmig,

Einzeläste glatt. Basalring oval, mit 1 Tracheenast. Thorax dorsal kräftig granuliert, Höhe und Zahl

j ; ; 2

“Li we Kr on l

A “u

hihi, „

üü a HN u,

in Y

hy Mu ui

Au hal 1

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ey an ae

“r wi

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IR N

TANZ TUR

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METALL AUTRURNN

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25 um //

Abb. 2 links: Kloosia afrıcana spec. nov., Hypopyg dorsal. 3 (rechts): Puppe. a) Abdominaltergite I-VIII; b)

Abdominalsternite I-III; c) Frontalplatte.

der Granula anal- und ventralwärts abnehmend. Ein flach gerundeter, länglicher praealarer Tuberkel

vorhanden. Scutumtuberkel fehlt. 2 schlauchförmige, sehr lange Antepronotal- und 2 Humeralbor-

sten sowie 4 einfache, kürzere, paarig stehende Dorsocentralborsten.

Abdomen (Abb. 3a, b): Tergit I mit anteriolateralen Feinchagrinfeldern, II-V mit einem posterio-

ren, transversalen, mehrreihigen Band kurzer kräftiger, etwa gleich großer Spitzen; oralwärts schließt

sich ein Feld locker stehender Spitzen gleicher Größe an, das durch spitzenfreie Flecken unterbrochen

ist. Tergit VI mit der prinzipiell gleichen Spitzenanordnung, jedoch ist das posteriore Band stark ver-

kürzt. VII und VIII mit anteriolateralen Feinchagrinflecken. Schwimmplatte mit einem anteriolatera-

len schmalen Streifen Feinchagrin. Posteriolaterale Ecken der Tergite V und VI mit einem Fleck sehr

feiner Spitzchen.

Sternite I- III mit je einer anterioren und posterioren transversalen Reihe heller, mäßig langer Spit-

zen; anteriore Reihe auf I und III median unterbrochen. Außerdem tragen die Sternite I-Ill einen zu-

sätzlichen Langspitzenfleck in den anterioren Ecken. Sternite IV—-VI chagrinfrei, VII- VIII mit 2 an-

terioren Feinchagrinflecken.

Posteriore Hakenreihe auf Tergit II median um etwa die Länge einer Hälfte unterbrochen; jede

Hälfte mit 11-17 Haken. Konjunktive nackt. Pedes spurii A vorhanden, Pedes spurii B fehlen. Seg-

ment VIII ohne Kämme oder Sporne.

Segment I ohne L-Borsten, II-IV jederseits mit 3 kurzen L-Borsten, V-VIII mit 4 LS-Borsten.

Analloben jederseits mit 1 langen dorsalen Schlauchborste. Schwimmhaarsaum aus 24-29 einzeilig

stehenden Schlauchborsten bestehend. © Gonopodenscheiden die Analloben mit der Hälfte ihrer

Länge weit überragend. Q Gonopodenscheiden entsprechend kurz, apikal mit einem kleinen warzen-

förmigen Fortsatz.

Larve: Unbekannt.

Material: Holotypus 1 schlüpfreife & Puppe, Kenia, Meru National Park, Rojewero-Flußdrift, 900 m Höhe,

März 1983, leg. E. J. Fittkau. Paratypen 5 Exuvien von derselben Lokalität. Die Typenserie befindet sich als Eupa-

ral-Dauerpräparate in der Zoologischen Staatssammlung München.

Differentialdiagnose

Die O Imago von Kloosia africana unterscheidet sich von den anderen Gattungsvertretern durch

Hypopygmerkmale: Obere und untere Volsella überragen die Analspitze nicht; obere Volsella kürzer

als die untere Volsella, golfschlägerförmig medianwärts gebogen, die beiden apikalen Setae eng zu-

sammenstehend. Die beiden anderen Arten haben lange, die Analspitze weit überragende untere Vol-

sellae, die oberen Volsellae sind nicht golfschlägerförmig und mindestens so lang oder länger als die

Analspitze.

Die Puppe von Kloosia africana ist durch posteriore Kurzspitzenbänder der Abdominaltergite

II—VI gekennzeichnet. Kloosia pusilla und Kloc sia sp. tragen dort ein Querband langer bis sehr langer

Spitzen.

Verbreitung und Ökologie

Kloosia africana ist nur vom locus typicus, einem Flüßchen in Nordkenia bekannt. Das Fließgewäs-

ser ıstan der Fundstelle etwa 5 m breit, 0,7 m tief, rasch strömend und kommt von den 2500 m hohen

Nyambeni Hills. Das Bett bestand überwiegend aus Sand mit gelegentlichen Kiesbeimengungen. In

Anlehnung an die Fließgewässerzonierung gemäßigter Breiten kann die Probestelle mit Einschrän-

kung dem oberen Potamal zugeordnet werden.

Kloosia pusilla (Linn&, 1767)

Tipula pusilla LinN£ 1767: 975

nec Chironomus pusillus LUNDSTRÖM 1910: 13

nec Chironomus pusillus LUNDSTRÖM & FREY 1916: 8

Kloosia pusilla KRUSEMAN 1933: 152— 154

Cryptochironomus sp. PAGAsT 1936: 273— 275

? Kloosia pusilla SOOT-RYEN 1943: 16

Cryptochironomus sp. (Pagast) CHERNOVSKI 1949: 52

Cryptochironomus lv. vytshegdae sp. n. ZVEREVA 1950: 271—272. Nov. syn.

Cryptochironomus serpancus sp. n. KIRPICHENKO 1961: 780. Nov. syn.

Cryptochironomus vytshegdae PANKRATOVA 1964: 192— 196

Kloosia pusilla ALgu 1980: 173— 174

Cryptochironomus vytshegdae SROKOSZ 1980: 199; table III

Cryptochironomus vytshegdae PANKRATOVA 1983: 166— 168

Cryptochironomus sp. Pagast ROssARO 1984: 131

Cryptochironomus sp. Pagast KLınk 1985: 2— 3

Chironomini Genus E PıNDER & REıss 1986: 364

Taxonomische Bemerkungen

Die Typusart der Gattung, K. pusilla, war bisher nur als O’ Imago bekannt, bis Kıınk (1985) wahr-

scheinlich machen konnte, daß die unter Cryptochironomus sp. (Pacast 1936) schon seit langem be-

schriebene Larve und Puppe dieser Art zugehören. Durch den aktuellen Fund J Puppen in Nord-

afrıka (Dr. H. Laville, unveröffentlicht) hat sich die Vermutung Klink’s voll bestätigt. Ein Vergleich

des nordafrikanischen Materials mit Exuvien aus Griechenland und der Puppenbeschreibung bei Pan-

KRATOVA (1964, 1983) lassen keinen Zweifel, daß auch Cryptochironomus vytshegdae — und damit

auch Cryptochironomus serpancus — jüngere Synonyme von Kloosia pusilla sind. Das Chironomini

Genus E bei Pınper & Reıss 1986 ist ebenfalls identisch mit K. pusilla.

Abb. 4: Kloosia pusilla (Linne). Hypopyg dorsal. a) Obere Volsella, Variation.

Differentialdiagnose

Die © Imagines von Kloosia pusilla lassen sich nach Hypopygmerkmalen (Abb. 4) von X. afrıcana

undK. koreana (Form und Länge der oberen und unteren Volsellae in Relation zur Analspitzenlänge)

und von der sehr ähnlichen nearktischen Art X. dorsenna (Saether) durch die Struktur des Tentoriums

(SAETHER 1987) unterscheiden. Die bekannten Puppen sind artlich ebenfalls gut zu trennen (vgl. Dif-

ferentialdiagnose von Kloosza afrıcana sowie Tafel 10.90 bei Pınper & Reıss 1986).

Ökologie und Verbreitung

Ein Vergleich der vorliegenden Daten zeigt, daß die Larven von Kloosia pusılla ein potamales Fau-

nenelement sind. Sie leben vorwiegend in Sandsedimenten größerer Flüsse, wobei die höchste durch-

schnittliche Larvenabundanz von Sroxosz (1980) mit 938 Ind./m? im Unterlauf der Nida, Polen, ge-

funden wurde. Eine gute Habitatcharakterisierung findet sich bei Pacast (1936): „...ım Flußsand mit

wenig organischer Substanz bei schwacher Strömung in !/4 m Tiefe.“

Die Art ist in der Westpalaearktis auch heute noch, trotz hoher Gefährdung ihres Lebensraumes,

den Flußsanden, offenbar weit verbreitet.

Funde liegen vor aus der UdSSR (Flüßchen in Livland, Oka, Pechora, Vychegda, Wolga, mittlerer Dnepr), Polen

(Nida-Unterlauf), Italien (mittlerer Po), diversen Lokalitäten in Holland (Kıınk 1985), Rumänien (ALBU 1980) und

eventuell Norwegen (SOOT-RYEN 1943; Nachbestimmung notwendig). Neunachweise liegen vor aus dem Fluß

Axios (Vardar) O Chalkidon, Griechenland, 5.5.1983, zahlreiche Exuvien aus Oberflächendrift, leg. M. Baehr so-

wie aus dem Aras-Tal W Karakurt, Provinz Hakkari, Osttürkei, 1300 m Höhe, 4.7.1985, 2 SC Imagines, leg. W.

Schacht. Das Material befindet sich in der Zoologischen Staatssammlung München. Außerdem sind JG Puppen und

Exuvien aus Tunesien bekannt (Coll. H. Laville, Toulouse; det. F. Reiss): Fluß Medjerda bei Jendouba, 146 mNN,

I Exuvie, 12.4.1985, max. Wassertemperatur 24° C; Bourheurtma, linker Zufluß des Medjerda, 133 m NN, 2 Ex-

uvien, max. Wassertemperatur 23°C; Oase Tozeur, Thermalquelle in einem Palmenhain, 3 0° Puppen aus Sandbo-

den sowie 2 Exuvien, 10.4.1985, Wassertemperatur 30,5°C; Oase Nefta, Thermalquelle in einem Palmenhain, san-

dig-kiesiges Substrat, 1 Exuvie, 10.4.1985, Wassertemperatur 28,3°C.

Die finnischen Funde von Kloosia pusilla, zitiert bei LUNDSTRÖM 1910 und LUNDSTRÖM & FREY 1916 sind Fehl-

bestimmungen, wie die kürzliche Durchsicht des kompletten Originalmaterials aus dem Zoologischen Museum

Helsinki durch den Autor ergeben hat.

Kloosia dorsenna (Saether, 1983)

Oschia dorsenna SAETHER 1983

Von dieser nearktischen Art aus South Carolina, USA, sind derzeit nur O’ Imagines bekannt. Sie

sind Kloosia pusilla sehr ähnlich. Unterschiede finden sich in der Tentorium-Struktur und in 1—2 zar-

ten Setae an der Squama, im Kontrast zur ausnahmslos nackten Squama bei pusilla.

Bezüglich der Kloosia-Gattungsdiagnose läßt sich am untersuchten Typenmaterial von dorsenna er-

gänzend feststellen, daß die obere Volsella, wie bei allen anderen Gattungsvertretern, keine Mikro-

trichien trägt und daß das Analtergit median getrennte Analtergitbänder besitzt. In der Originalbe-

schreibung muß die Abb. Ic entsprechend korrigiert werden.

Kloosia sp.

Puppe:

Entspricht in Größe (Länge 3,0—3,4 mm), Färbung und Chaetotaxie (Analflosse mit 23—26 Setae)

weitgehend der Art pusilla. Ein Unterschied besteht in der Länge der posterioren Nadelspitzen auf

den Abdominaltergiten II-V. Sie sind bei Xloosia sp. merklich länger als bei X. pusilla. Eine spezifi-

sche oder subspezifische Wertung dieses Merkmals kann ohne zugehörige Imago nicht erfolgen.

Verbreitung und Ökologie

Der vorliegende Fund aus Südwestchina ist der erste Nachweis der Gattung Kloosia aus der Orien-

talis: Prov. Yünnan, 7 km O Mung Lun, Schutzgebiet Tri Ping Fung, Flußdrift, 28.5.1980, leg. E. ].

Fittkau. Der Fluß, ein Zufluß des Mekong, ist an der Fundstelle und am Ende der Trockenzeit ca.

50-70 m breit und weist vorwiegend grobkiesigen Untergrund auf. Das mit 3 Individuen ausgespro-

chen seltene Auftreten von Kloosia sp. in einer sonst sehr arten- und individuenreichen Driftprobe läßt

sich wohl mit dem Mangel an Sandsedimenten als geeignetes Larvenhabitat erklären. Soweit über-

haupt bestimmbar, enthielt die Probe an weiteren potamalen Faunenelementen Exuvien von Robackia

sp., Sublettea sp. und von Chironomini Genus D (Pınper &Reıss 1986), einem in anderem Zusammen-

hang zu behandelnden Taxon.

Diskussion

In zunehmendem Maße erweist sich ein beträchtlicher Teil der Arten des Harnischia-Gattungs-

komplexes weltweit als bedeutende Komponente der Chironomidenfauna großer Fließgewässer.

Aufgeführt sei in diesem Zusammenhang die Zusammensetzung der Chironomidenfauna einer klei-

nen Oberflächendriftprobe aus dem griechischen Fluß Axios östlich Chalkidon vom 5.5.1983 (leg.

M. Baehr). An der Probestelle wies der Fluß etwa 150 m Breite, 2 m Tiefe, Sand- und Schlickbänke bei

kräftiger Strömung auf. Von den insgesamt 162 Exuvien gehörten 48 (29,6%), von den 19 Arten 6

(31,6%) zum Harnischia-Komplex, wobei Kloosia pusilla mit 30 Individuen dominant war.

Ein hoher Anteil dieser potamalen Komponente innerhalb des Harnischia-Komplexes ist auf Art-

und Gattungsniveau taxonomisch nicht oder erst in jüngster Zeit erfaßt worden. Eines der Beispiele

dafür ist die Gattung Äloosia, die — bis vor kurzem nur mit einer ökologisch bezugslosen europäi-

schen Art vertreten — nunmehr mindestens 4 Arten ın 3 zoogeographischen Regionen, der Holarktis,

Afrotropis und Orientalis, aufzuweisen hat. Zudem darf man annehmen, daß damit noch nicht das ge-

samte Arteninventar dokumentiert ıst.

Einer der Gründe für das späte Erkennen und verzögerte Erfassen dieses potamalen Faunenanteils

bei Chironomiden ist in methodischen Schwierigkeiten zu suchen, die eine faunistisch-biologische,

vor allem aber ökologisch-quantifizierende Bearbeitung von Fluß- und Stromzoozönosen mit sich

bringt. Große, physikalisch und biologisch extrem dynamische Fließgewässer lassen sich ungleich

schwieriger bewältigen als die relativ statischen stehenden Gewässer aller Größenklassen. Hinzu

kommt die vergleichsweise frühzeitige, zivilisatorisch bedingte Degradierung und Kontaminierung

der meisten großen Fließgewässer gemäßigter Breiten, die die Untersuchung ihrer naturnahen Fauna

in den vergangenen Jahrzehnten nicht mehr zuließen. Jedoch zeigt gerade die Erfahrung der letzten

Jahre, daß viele in Europa als erloschen gegoltene Potamobionte lokal in Klein- und Kleinstpopulatio-

nen offensichtlich überlebt haben und damit, wenn auch nicht dem Ökologen, so doch dem Taxono-

men zur Bearbeitung verfügbar blieben.

Bei den Chironomiden finden sich potamobionte Arten des Harnischia-Komplexes vor allem ın

den Gattungen Acalcarella, Beckidia, Chernovsküa, Cyphomella, Gillotia, Kloosia, Robackia, Saethe-

ria und Chironomini Genus D. Hinzu kommen zahlreiche Arten ungeklärter Gattungszugehörigkeit,

von denen zum Teil nur das Larvenstadium beschrieben ist. Etwa ein Dutzend davon ist unter Cryp-

tochironomus bei PankrarovA (1983) zu finden. Mindestens dieselbe Zahl noch unbeschriebener Lar-

ventypen unbekannter Gattungszugehörigkeit sind mir aus dem Stromsystem des Amazonasbeckens

bekannt. Hinzu kommen weitere, offenbar an größere Fließgewässer gebundene und ebenfalls unbe-

schriebene Taxa aus Nordafrika, dem tropischen Afrika, Madagaskar, Nordindien, Südwestchina und

Neukaledonien.

Global gesehen, entsteht bei der Betrachtung der potamalen Komponente des Harnischia-Komple-

xes zunehmend der Eindruck einer generisch ausnehmend stark differenzierten und phylogenetisch

alten Entwicklungslinie, die ihre Anpassung an einen archaischen aquatischen Lebensraum durch

zahlreiche Apomorphien, speziell bei Larven, dokumentiert.

Literatur

Augu, P. 1980: Fam. Chironomidae — Subfam. Chironominae. — Fauna Republ. Social. Romania, Insecta, Diptera

11: 320 p.

CHERNOVSKI, A. A. 1949: Identification of larvae of the midge family Tendipedidae. — Izd. Akad. Nauk, SSSR 31:

186p.

KIRPICHENKO, M. Ya. 1961: A new form of the larvae Cryptochironomus serpancus sp.n. — Zool. Zh. 40: 780-781

KLink, A. 1985: Een inventarisatie van volwassen Chironomidae bij Kampen (Ijssel). — Rapp. Mededel. 21: 1-5

KRUSEMAN, G. 1933: Tendipedidae Neerlandicae. I. Genus Tendipes cum generibus finitimis. — Tijdschr. Ent. 76:

119-216

LUNDSTRÖM, C. 1910: Beiträge zur Kenntnis der Dipteren Finnlands. VI. Chironomidae. — Acta Soc. Fauna Flora

fenn. 33: 1-46

LUNDSTRÖM, C. & FREY, R. 1916: Beiträge zur Kenntnis der Dipteren Finnlands. — X. Suppl. 4. Bibionidae, Chi-

ronomidae, Tipulidae. — Acta. Soc. Fauna Flora fenn. 44: 5—25

PAGAST, F. 1936: Chironomidenstudien I. — Stettin. ent. Ztg. 97: 270-278

PANKRATOVA, V. Ya. 1964: The larvae of Tendipedidae (Chironomidae) of the Oka river. — Trudy zool. Inst., Le-

nıngr. 32: 189— 207

PANKRATOVA, V. Ya. 1983: Larvae and pupae of midges of the subfamily Chironominae of the USSR fauna (Dip-

tera, Chironomidae = Tendipedidae). — Izd. Nauka, Leningrad, 295 p.

Papp, J. & HORVATOVICH, $. 1972: Zoological collectings by the Hungarian Natural History Museum in Korea.

2. A report on the collecting of the second expedition. — Folia ent. Hung. 25: 187—227

PinDEr, L. C. V. & Reıss, F. 1986: 10. The pupae of Chironominae (Diptera: Chironomidae) of the Holarctic re-

gion — Keys and diagnoses. — Ent. scand. Suppl. 28: 299-456

Reıss, F. 1980: Zur Zoogeographie der Chironomidenfauna (Diptera, Insecta) Nordkoreas. — In: MURRAY, D. A.

(ed.): Chironomidae. Ecology, systematics and physiology. — Proc. 7th Int. Symp. Chironomidae. Dublin,

Aug. 1979, Pergamon Press: 145— 149

RossARO, B. 1984: The chironomids of the Po river (Italy) between Trino Vercellese and Cremona. — Aquat. In-

sects 6: 123—135

SAETHER, O. A. 1983: Oschia dorsenna n. gen., n. sp. and Saetheria hirta n. sp., two new members of the Harnischia

complex (Diptera: Chironomidae). — Ent. scand. 14: 395—404

SAETHER, O. A. 1986: On the systematic positions of Dolichprymna, Amblycladius and Kloosia (Diptera: Chirono-

midae). — Abstr. First Int. Congr. Dipterology, Budapest: 215

SAETHER, O. A. 1987: On the systematic position of Dolichprymna, Amblycladius and Kloosia (Diptera: Chirono-

midae). — (im Druck)

SooT-RyeEn, T. 1943: A preliminary list of Norwegian finds of Heleidae and Tendipedidae. — Tromsö Mus. Arsh.

(Nat. Avd.) 64: 1-24

SROKOSZ, K. 1980: Chironomidae communities of the river Nida and its tributaries. — Acta Hydrobiol. 22:

191215

ZVEREVA, O. S. 1950: New forms of Tendipedidae larvae from the rivers Pechora and Vychegda. — Ent. Obozr. 31:

262—284

Dr. Friedrich Reiss,

Zoologische Staatssammlung,

Münchhausenstr. 21,

D-8000 München 60

q SPIXIANA Supplement 14 | 45—47 | München, 15. Juli 1988 ISSN 0177-7424

Diplosmittia recisus spec. nov. from Peru

(Diptera, Chironomidae)

By Ole A. Szther

Abstract

The male of Diplosmittia recısus spec. nov. is described from Peru. It differs from D. harrison: Szther from the

Eastern Lesser Antillean Islands, and D. carınata S&ther from Michigan, the two previously known members ofthe

genus, in having only 9 flagellomeres and a strongly extended costa.

Introduction

SATHER (1981) described four new genera from the British West Indies. Three of these genera later

were described also from the southeastern parts of the U.S.A. (S£THer 1982). The remaining genus,

Diplosmittia, recently was described from Michigan (S£THEr 1985). Recently Dr. F. Reiss, Zoologi-

sche Staatssammlung, Munich, sent me a male imago collected in Peru. It proved to belong to a new

species of Diplosmittia closely related to the previously described species.

Methods and morphology

The general terminology follows SETHER (1980) with the exception that the apical spine of the gonostylus is ter-

med the megaseta. In the figure of the male hypopygium the dorsal aspect is shown to the left, the ventral aspect

and the apodemes to the right.

Diplosmittia Szther, 1981: 29, emended

Antenna with 9-13 flagellomeres, flagellomeres 2-8 or possibly sometimes only 2-5, and ultimate

with sensilla chaetica. Costa not or barely to relatively strongly extended, with or without additional

false vein extending beyond costa nearly to wing tip. Otherwise as in SETHER (1981: 29).

Diplosmittia recisus spec. nov.

(Fig. 1)

Type material: Holotype, male, Station Koepke Panguana, about 260 m a. s. |., 9°37'S, 74°56’W, Peru,

17.2.1974, W. Kurz (Nr. 10), in the collection of Zoologische Staatssammlung, Munich.

Diagnostic characters

Nine flagellomeres, costal extension moderately long (about 68 um), LR, 0.36, LR, 0.36, LR; 0.43.

Fig. 1. Diplosmittia recisus spec. nov., male imago. — A. Wing. — B. Hypopygium.

Etymology: From Latin, recisus, cut back, cut short, referring to the reduced number of flagellomeres.

Description

Male imago (n = 1)

Total length 1.33 mm. Wing length 0.79 mm. Total length/wing length 1.68. Wing length/length of

profemur 2.98. Coloration brown.

Head (as in D. harrisoni, Szther 1981 fig. 13 A). Nine flagellomeres, ultimate 244 um long. AR

0.80. Temporal setae apparently consisting of 1 inner vertical. Clypeus apparently with 4 setae. Tento-

rıum and stipes not measurable. Palp segments length (micrometers): 15, 19, 41, 45, 45.

Thorax (as in D. harrisoni, Szrner 1981 fig. 13B). Antepronotum apparently with 1 lateral seta

only. Dorsocentrals 4, prealars 3. Scutellum with 2 setae.

Wing (Fig. 1A). VR 1.44. C extension 68 um long. Brachiolum with 1 seta.

Legs. Spur of front tibia 26 um long, spurs of middle tibia 15 um and 11 um long, of hind tibia

32 um and 15 um long. Width at apex of front tibia 26 um, of middle tibia 23 um, of hind tibia 23 um.

Comb with 8 setae, 17—23 um long. Lengths (micrometers) and proportions of legs:

fe ti ta, ta, ta, ta, ta, LR BV SV BR

Pl 266 341 124 79 56 30 3010236275 321547 498. 223

P, 323 356 129 79 58 30 327 7053687 71077 5225 er.

PJ 293 328 143 90 94 24 30 0.43 3.08 4.36 303

Hypopygium (Fig. 1B). Anal point with 6 lateral and basal setae, laterosternite IX with 2 setae.

Phallapodeme 63 um long, transverse sternapodeme 54 um long. Virga 38 um long. Gonocoxite

92 um long. Gonostylus 49 um long, with lobe 57 um long, apical megaseta 6 um long. HR 1.91, HV

PRT2.

Acknowledgement

I am much indebted to Dr. F. Reiss, Zoologische Staatssammlung, Munich, for the type material.

Literature

SETHER, ©. A. 1980: Glossary of chironomid morphology terminology (Diptera: Chironomidae). — Ent. scand.

Suppl. 14: 1-51

-— 1981: Orthocladiinae (Diptera: chironomidae) from the British West Indies, with descriptions of Antillocla-

dius n. gen., Lipurometriocnemus n. gen., Compterosmittia n. gen. and Diplosmittia n. gen. — Ent. scand.

Suppl. 16: 1-46

—— 1982: Orthocladiinae (Diptera: Chironomidae) from SE U.S.A., with descriptions of Plhudsonia, Unniella

and Platysmittia n. genera and Atelopodella n. subgen. — Ent. scand. 13: 465-510

Prof. ©. Szther

Museum of Zoology

University of Bergen

N-5007 Bergen, Norway

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SPIXIANA Supplement 14 49 —55 München, 15. Juli 1988 ISSN 0177— 7424

Vivacricotopus, a new genus of Orthocladiinae from Norway

(Diptera, Chironomidae)

&yvind A. Schnell and Ole A. S&ther

Abstract

A new genus and species of Orthocladiinae, Vivacricotopus ablusus, is described as male imago and pupa. The

imago has hairy eyes; well developed pulvilli; coarse punctation of microtrichia on wing membrane; two median

acrostichals; long, bare anal point; and a virga; which is a unique combination. The pupa has thoracic horn and anal

fringe, and a unique arrangement of simple, branched and broadly filamentous L-seta. The genus apparently occu-

pies a plesiomorphic position within the Rheocricotopus group of genera.

Introduction

During the investigations of the area around the Jostedal Glacier in connection with the future

building of hydroelectric power stations and dams, some peculiar pupal exuviae were found in the

partly glacier fed river Jostedola. Closer examination showed that also a male belonging to an un-

known genus with the same affinities was present in the same sample. The new genus and species is

described below.

Methods and Morphology

The mounting procedure used is outlined by S£ETHER (1969: 1). The general terminology follows SETHER (1980)

with the exception that the vannal fold is called the postcubitus, and the apical spine of the gonostylus the megaseta.

The measurements are given as ranges followed by a mean, followed by the number measured in parenthesis (n).

Vivacricotopus gen. nov.

Type species: Vivacricotopus ablusus spec. nov. by present designation.

Diagnostic characters

The combination of hairy eyes; large pulvilli; two central acrostichals; relatively coarse punctation

on wing membrane; nearly straight Cu, ; long, narrow and bare anal point; and presence of virga will

separate the male imago from all other orthoclads. The absence of antepronotals, the low number of

dorsocentrals and scutellars, and the absence of a crista dorsalıs also are characteristic.

The pupa has a unique pattern of shape of L-setae with L, split in 2-6 branches on segments

III—- VI; L, on VI filamentous; 4 filamentous L-setae on VII and 5 filamentous L-setae on VIII, with

the two posterior on each segment very broad. Also the caudolateral extension of segments VII and

VII and the very strong anal macrosetae are unique features.

Fig. 1. Vivacricotopus ablusus gen. nov., spec. nov., male imago. — A. Head. — B. Thorax. — C. Cibarial pump,

tentorium, and stipes. - D. Hypopygium, dorsal aspect to the left, ventral aspect to the right. — E. Wing.

Etymology: From the type locality Viva, near the river Jostedola, and Cricotopus, a common genus name and

ending among Orthocladiinae meaning ringed legs.

Description

Male imago

Medium sized species, wing length about 2 mm. Coloration brown with ringed tibia and pale tarsı.

Eyes hairy with short, wedge-shaped dorsomedial extension.

Antenna with 13 flagellomeres; antennal groove beginning at flagellomere 3; flagellomeres 2, 3, 4

and 13 with thin sensilla chaetica; AR lower than 1.0. Temporals divided into very short inner verti-

cals, longer outer verticals and longest postorbitals.

Clypeus as wide as pedicel. Anterior margin of cibarial pump deeply concave. Five palpal segments,

third and fifth subequal in length, third palpal segment with weak apical projection and apparently

without apical sensilla clavata, fourth segment with indication of similar apical projection.

Antepronotum well developed; median lobes not medially narrowed, gaping, separated in front of

scutal projection; no lateral setae. Two acrostichals in centre of scutum, dorsocentrals and prealars

few, supraalars absent. Scutellum with few setae.

Wing membrane with relatively coarse punctation of microtrichia, free of setae. Anal lobe well de-

veloped, slightly projecting. Costa slightly extended; R,,, running in the middle between R, andR,;;,

ending at !/s the distance between end ofR; and R, ,5; Ry;s ends clearly distal to end of M,,,; FCu lies

distally of RM, Cu, nearly straight, posteubitus ends far distally of FCu, anal vein ends slightly distally

of FCu. R with a few setae, R, bare, R,;; with seta(e) at apex. Sensilla campaniformia in normal num-

bers (about 12 at base, 3 below seta and 13 at apex of brachiolum, 2 on subcosta, 1on FR, and 1 at base

of R,). Squama with few setae in fringe.

Fig. 2 Vivacricotopus ablusus gen. nov., spec. noV., pupa. 7 A. Tergites. — B. Sternites. —- C. Frontal apotome and

ocular field. - D. Thoracic horn and precorneal setae. — E. Caudolateral corner of segment V.

Tibıal spurs and hind tibial comb normal. Tibia with pale rings. Pseudospurs absent. Sensilla chae-

tica present in basal !/2 of tarsomere 1 of hind leg. Pulvilli well developed, about 3/4 as long as claws.

Tergites with setae in 2-3 rows of median setae and 2—3 rows of lateral and posterior setae, or more

scattered. Anal point long, free of microtrichia, with several setae at base on tergite IX. Phallapodeme

normal, transverse sternapodeme curved with well developed oral projections. Virga consisting of clu-

ster of 2-3 spines. Gonocoxite with low inferior volsella and no superior volsella. Gonostylus evenly

wide for most of its length, with well developed megaseta, without crista dorsalıs.

Pupa

Median sızed pupae about 4 mm long. Frontal seta moderately long, on frontal apotome. Frontal

apotome with weak to moderately developed warts, slightly wrinkled. Antennal sheath at most with

2—4 pearls above pedicel. Ocular field with 2 postorbitals. Antepronotum with 2 median and 2 lateral

setae, all well developed. Thoracic horn cylindrical, tapering towards apex, covered with not very

dense spinules. Two anterior setae subequal in length and about twice as long as posterior seta. Second

dorsocentral longer than the others, anterior 3 dorsocentrals grouped or third equidistant from second

and fourth. Thorax slightly wrinkled, wing sheath nearly smooth. Sheath of coxae with 1 minute seta

each.

Tergite I without shagreen, II-IX with weak anterior and narrow median shagreen grading over

into caudal spines on II- VII. Sternites I and IX without shagreen; II- VIII with weak anterior group

shagreen, more extensive on posterior sternites and grading over into weak caudal spines. Tergites

II- VIII with about 4 rows of weak caudal spines, weaker on VIII. Tergite II without caudal hooklets.

Caudal margin of sternites II-VII with 3—4 rows of similar, but weaker caudal spines, very weak or

reduced to a few spinules on II. No spinules on conjunctives. Pedes spurii A and B absent. Segment I

with 4 D setae, 1 L seta and 2 V setae. Segments II-VII with 5 D setae, 4 L setae, 4 V setae, and O

setae in pattern B (Corrman 1979, 2 dorsal and 1 ventral pair of O setae). Segment VIII with 1 D seta,

5 Lsetaeand 1 V seta. L, on segments III- VI split into 2—6 branches, L; on segment VIandallL setae

on VII and VIII filamentous, L, and L; very broad on segments VII and VIII. Segments VII and VIII

with conspicuous, rounded, darkened caudolateral projections. Apophyses well delineated. Anal lobe

with sparse fringe of lamelliform setae and 3 very strong macrosetae about 2 as long as the lobe with

lateral macroseta slightly weaker then the median ones. Male genital sac not reaching apex of anal lobe.

Systematics

Especially the pupa but also the male imago of the new genus shows an unusual combination of

characters. The hairy eyes, the large pulvilli, the median acrostichals, and the pupal horn, spine pattern

and anal lobe fringe apparently place the genus in the Rheocricotopus Thienemann & Harnisch group

of genere (Brunpın 1956: 118; SETHER 1977 fıg. 36, 1980b: 131, 1981: 224, 1983 a fig. 5, 1985: 63).

However, Rheocricotopus, Paracricotopus Thienemann & Harnisch, Mesocricotopus Brundin, Nano-

cladius Kieffer, Doncricotopus Szther and Psectrocladius Kieffer all lacks a virga and the 5 first have a

platelike superior volsella not found in Vivacricotopus. The male imago also resembles Sublettiella

Szther (1983b) in the absence of lateral antepronotals, the low thoracic chaetotaxy, the wing puncta-

tion and venation except for the somewhat less curved Cu,, the hairy eyes, the presence of pulvilli, the

virga, and the low inferior volsella. Vivacricotopus, however, differ from Sublettiella for instance in the

absence of pseudospurs, the presence of sensilla chaetica on hind leg, the presence of 2 median acrosti-

chals and the long bare anal point.

The pupa of Vivacricotopus will key to Unniella Szther ın Corrman et al. (1986). However, it does

show few similarities with that genus. Among genera with anal lobe fringe and thoracic horn no other

genus is at the same time lacking pedes spurii A and B and caudal hooklets on tergite II. Caudal hook-

lets are absent only in Parametriocnemus Goetghebuer and Paratrissocladius Zavtel, pedes spurii A

may be absent in some Zalutschia Lipina, while Paracricotopus, Nanocladius subgen. Plecopteracolu-

thus Steffan, Zalutschia, many Psectrocladins, some Rheocricotopus, and some Heterotanytarsus

Spärck lack pedes spurii B. The shape oftheL setae with L, branched and the posterior L setae on seg-

ments VII and VIII very broad is unique. The B pattern of O setae is not very common in the Ortho-

cladiinae, but is the pattern found in Psectrocladius, Rheocricotopus and Unniella.

Although the genus most likely belong near or in the Rheocricotopus group it probably occupies a

plesiomorphic position relative to the other genera of the group. It also show similarities with Szblet-

tiella and the genera near Bryophaenocladius Thienemann and partly with Unniella. As most likely for

the last genus in which the pupa appear to belong to the Rheocricotopus group, the larva to the Para-

kiefferiella group and the male imago to the Mesosmittia group, the similarities of Vivacricotopus with

the Rheocricotopus group may be based on plesiomorphies. In that case a placement near Sublettiella

is the most likely one.

Vivacricotopus ablusus spec. nov.

(Figs 1, 2)

Type locality: Norway, Sogn & Fjordane, Luster, Jostedola river, Viva.

Type material: Holotype, male, Viva, Jostedola river, Luster, Sogn & Fjordane, Norway, 23. VII. 1986. Leg.:

&.A. Schnell and A. Fjellheim (ZMBN No. 107). Paratypes, 6 pupal exuviae, same data as holotype. Types in coll.

Mus. Zool., Ent. coll. Univ. of Bergen (ZMBN).

Diagnosis: See diagnosis of the new genus.

Etymology: From Latin ablusus, meanıing different, unlike, referring both to the unique combination of charac-

ters and to the very different types of L-setae on the pupae.

Description

Male Imago

Total length 3.10 mm. Wing length 1.93 mm. Total length/wing length 1.61. Wing length/length of

profemur 2.91. Coloration brown, halterers pale brown, tarsı and middle 2/3 of tibiae pale.

Head (Figs 1A + 1C). AR 0.72. Ultimate flagellomere 364 um long. Temporal setae 7, including 3

inner verticals, 2 outer verticals, and 2 postorbitals. Clypeus with 7 setae. Cibarıal pump, tentorium

and stipes as ın Fig. 1C. Tentorium 150 um long, 32 um wide. Stipes 139 um long, 56 um wide. Palp

segments length (micrometers): 30, 53, 120, 88, 116. Third palpal segment with weak apical projection,

apparently no sensilla clavata; fourth segment with similar projection.

Thorax (Fig. 1B). Antepronotum bare. Humeral pit weak, normal. Dorsocentrals 3, acrostichals 2,

prealars 3. Scutellum with 2 setae.

Wing (Fig. 1E). Wing membrane with punctation of microtrichia visible at 150X. C extension

45 um long. R with 6 setae, R, bare, R,,; with 1 apical seta, C extension with 1 non-marginal seta.

Squama with 7 setae.

Legs. Spur of front tibia 49 um long, spurs of middle tibia 24 um and 23 um long, of hind tibia

49 um and 19 um long. Width at apex of frontttibia 38 um, of middle tibia 36 um, ofhind tibia 41 um.

Comb of hind tibia with 12 setae, 19—49 um long. Sensilla chaetica 8 at 0.19—0.49 on ta, of hind leg.

Lengths (micrometers) and proportions of legs:

fe ti ta, ta, ta, ta, bar LR BV SV BR

PJ GEBE ze Al a 85 BL Bee Bf 2

P, 695 709 312 189 142 ie Ah Fo ee 26

P, 709 832 416 2a ae AO Te 0 Fe Se Bo

Hypopygium (Fig. 1D). Anal point 41 um long, with 14 setae at base on tergite IX;; laterosternite

IX with 5 setae. Phallapodeme 79 um long, transverse sternapodeme 109 um long. Virga 24 um long.

Gonocoxite 193 um long, inferior volsella weak. Gonostylus 77 um long, megaseta 19 um long. HR

2.51, EILV.4.03:

Pupa (n = 6, except when otherwise stated).

Total length 3.70—4.03, 3.88 mm. Exuviae pale brownish grey.

Cephalothorax. Frontal setae on frontal apotome (Fig. 2C), 75-116, 99 um long. Postorbitals

75—116, 95 um and 56-94, 70 um long. Median antepronotals both 75-113, 96 um long. Thoracic

horn (Fig. 2D) 188-221, 209 um long; 36—43, 40 um wide; 1.06-1.19, 1.14 times as long as anal ma-

crosetae. Anterior precorneal seta 79—131, 94 um long; 4— 17, 10 um in front of median seta. Median

precorneal seta 105-135, 126 um long; 4-15, 8 um in front of posterior seta. Posterior precorneal

seta 45—86, 61 um long; 38—45, 41 yum in front of horn. Second dorsocentral (Dc,) 68-105, 82 um

long; other dorsocentrals 45—64, 55 um long. Distance between Dc, and Dc, 15—23, 17 um; between

Dec, and Dec; 15—38, 24 um; between Dec; and Dec, 39—68, 50 um. Setae on coxal sheaths 2-11, 6 um

long on front leg; 11—26, 20 um long in middle leg; 23—30, 26 um long on hind leg.

Abdomen (Fig. 2A, B). Shagreenation, caudal spinesand chaetotaxy as in generic diagnosis. Ma-

ximal length (micrometers) of caudal spines on TI— VIII as: 6-9, 7;8— 11,10; 11-15, 13; 15-17, 16;

17— 23,21; 13—21, 19; 4—8, 6. L, on segment III split into 2 branches; on IV and V ın 3—4,4 branches

(Fig. 2E); on VI in 3-6, 4 branches. Anal lobe 263—278, 268 um long; with 8-12, 10 setae in fringe;

macrosetae 176-193, 183 um long. Genital sac of male ending 11-23, 17 um (5) short of apex of anal

lobe; of female ending 98 um (1) short of anal lobe.

Ecology and Distribution.

The pupal exuviae and the drowned male imago were collected in an eddy where the river Jostedola runs into a

small lake at Viva (alt. 890 m a. s. |., UTM ref. 32V MP 474184) in the uppermost part of the valley Jostedal. The

type locality is situated above the timber line only a few kilometers east of the Jostedal Glacier, which is the largest

glacier on mainland Europe, covering an area of approximately 486 sq. km.

The water temperature varies between near 0°C during the period of ice cover; from middle of the november to

end of may; to 11.7°C the day the specimens were sampled, the highest measured at Viva in the years —85 and —86.

The pH varies between 5.0 to 6.0, the conductivity (uS/cm) between 5.7 to 9.5 during —85 and —86. (A. Fjellheim

pers. comm.). The river ıs heavily loaded with silt from the nearby glacier.

Some other chironomids found at the type locality were: Diamesa lindrothi Goetghebuer, Psendodiamesa cf. ni-

vosa (Goetghebuer), Eukiefferiella minor (Edwards), several new species of Limnophyes SETHER 1988, Mesocrico-

topus thienemanni (Goetghebuer), Orthocladius (Eudactylocladius) grampianus (Edwards), Orthocladius

(Euorthocladius) frigidus (Zetterstedt), Orthocladius (Euorthocladius) rivicola Kieffer, Orthocladins (Enortho-

cladius) thienemanni Kieffer, Psectrocladius (Allopsectrocladius) sp., Rheocricotopus (Rheocricotopus) effusus

(Walker), Rheocricotopus (Rheocricotopus) reduncus Sether & Schnell 1988, Micropsectra recurvata (Goetghebuer).

The new species is known only from the type locality.

Acknowledgements

We are indebted to the Norwegian Water Resource and Electricity Board for financial support and to Arne Fjell-

heim at Museum of Zoology, University of Bergen, for providing us with information on thetemperature and water

properties of the type locality.

Literature

BRUNDIN, L. 1956: Zur Systematik der Orthocladiinae (Dipt., Chironomidae). — Rep. Inst. Freshwat. Res. Drott-

ningholm 37: 5—185

COFFMAN, W.P., P. $S. CRANSTON, D. R. OLIVER & O. A. SETHER. 1986: 9. The pupae of Orthocladiinae (Diptera:

Chironomidae) of the Holarctic region. — Keys and diagnoses. — Ent. scand. Suppl. 28: 147—296

SETHER, O. A. 1969: Some Nearctic Podonominae, Diamesinae, and Orthocladiinae (Diptera: Chironomidae). —

Bull. Fish. Res. Bd Can. 170: 1-154

1977: Female genitalia in Chironomidae and other Nematocera: Morphology, phylogenies, keys. — Bull. Fish.

Res. Bd Can. 197: 1—209

1980a: A glossary of chironomid morphology terminology (Chironomidae: Diptera). — Ent. scand. Suppl.

1980b: The females and immatures of Paracricotopus Thienemann & Harnisch, 1932, with the description of

a new species (Diptera: Chironomidae). — Aquatic Insects 2: 129— 145

1981: Doncricotopus bicandatus n. gen., n. sp. (Diptera: Chironomidae) from the Northwest Territories, Ca-

nada. — Ent. scand. 12: 223—229

1983 a: The canalized evolutionary potential — inconsistencies in phylogenetic reasoning. — Syst. Zool. 32:

343—359

1983 b: A rewiew of Holarctic Gymnometriocnemus Goetghebuer, 1932, with the description of Raphidocla-

dius subgen. n. and Sublettiella gen. n. (Diptera: Chironomidae). — Aquatic Insects 5: 209-226

1985: A review of the genus Rheocricotopus Thienemann & Harnisch, 1932, with the description of three new

species. — Spixiana Suppl. 11:59 —108

1988: A revision of the genus Limnophyes Eaton from the Holarctic and Afrotropical regions (Diptera: Chi-

ronomidae, Orthocladiinae). — Ent. scand. Suppl.

SETHER, ©. A. & SCHNELL, ©. A. 1988. Two new species ofthe Rheocricotopus (R.) effusus group (Diptera: Chiro-

nomidae). — Spixiana Suppl. 14: 65-74

©. A. Schnell

Prof. ©. A. Sxther,

Museum of Zoology, University of Bergen,

N 5007 Bergen, Norway.

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SPIXIANA Supplement 14 57-64 München, 15. Juli 1988 ISSN 0177-7424

Heterotrissocladius brundini spec. nov. from Norway

(Diptera, Chironomidae)

By Ole A. S&ther and Oyvind A. Schnell

Abstract

Heterotrissocladins brundini spec. nov. is described in all stages and both sexes. It apparently form the plesiomor-

phic sister species of A. maeaeri Brundın.

Introduction

Langron (1984: 88, fig. 28a) illustrated a pupa of Heterotrissocladius with imbedded spines on seg-

ments VI-VIIl and very large frontal warts. During an investigation of some acıidified lakes in the sou-

thernmost part of Norway the junior author found all stages of apparently the same species. Later it

was found also in western and northwestern Norway.

Methods and morphology

Morphological nomenclature follows S£THER (1980) with the exception that the apıcal spine of the gonostylus

is termed the megaseta. The measurements are given as ranges followed by amean when four or more measurements

are made, followed by the number measured in parentheses (n).

The type material is kept at the Museum of Zoology, University of Bergen (ZMB).

Heterotrissocladius Spärck 1922, emended

Imago — Eyes completely naked, with a few microtrichia between marginal ommatids or pubes-

cent, i. e. with microtrichia between all ommatids, with reticulation between central ommatids.

Otherwise as in SETHER (1975: 3).

Pupa — Frontal warts weak to strongly developed. Frontal setae on frontal apotome (most species)

or on prefrons (at least A. latilaminus Sether). Caudolateral corners of segments VI-VIll occasionally

with inner sclerotization resembling imbedded spines (H. brundini spec. nov), these strong on VII,

weaker on VI, and only indicated on VIII. Otherwise as ın SETHER (1975: 4).

Larva — Anterior parapods with apical claws grading over into basal hair-like spines anteriorly and

on acommon base. Otherwise as in SETHER (1975: 4).

Heterotrissocladius brundini spec. nov.

(Figs 1-3)

Heterotrissocladins Pe 1, Langton 1984: 88

Type locality: Norway, Aust-Agder, Birkenes, Lake Repstadvatn.

Type material: Holotype, male, Lake Repstadvatn, Birkenes, Aust-Agder, Norway, 19/9/86, leg. H. Segrov &

©. A. Schnell, in coll. Mus. Zool. Univ. of Bergen (ZMB No. 115).

Paratypes: 5 males, 4 females, 2 male pupae, 147 pupal exuviae, 6 larval exuviae, as holotype; 5 pupal exuviae,

Lake Store Hovvatn, Birkenes, Aust-Agder, Norway, 6/9/86, ©. A. Schnell mature female pupa, Lake Jolster, Jol-

ster, Sogn & Fjordane, Norway, 18/8/87, H. S&grov; mature male pupa, Lake Litlebovatn, Volda, More & Roms-

dal, Norway 26/7/86, ©. A. Schnell; Other material (?): Male, Nordseter, Lillehammer, Oppland, Norway, 31/8/

86, R. Bergo.

Diagnostic characters

The normal clypeus and cıbarıal pump combined with the moderately short acrostichals starting

some distance from the scutal projection and the few setae on the wing membrane will separate the

species from other Heterotrissocladius except H. maeaeri Brundin. However, the stipes isreduced, the

wing slightly darker, the AR lower (0.7-1.1), the LR, higher (0.76—0.78) and the number of setae in

cell r,;; usually lower (7-18, except in one mature male pupa apparently with about 50 setae and an-

other possible specimen with 45 setae, see remarks) in 4. brundini spec. nov. The pupa can be separat-

ed from other members of the genus by means of the large frontal warts and the imbedded spines on

segments VI-VIII. The larva can be distinguished by the narrowly separated two median teeth of the

mentum with distinct lateral notches, the brownish black submentum which is conspicously darker

than the surrounding areas of the head capsule, and the VM ratio of 1.1-1.5.

Etymology: Named in honour of Prof. emer. Lars Z. Brundin to his 80 year birthday and to signify his eminent

contribution to chironomidology and lake typology where Heterotrissocladius has occupied a central position.

Description

Male imago (n = 7-9, except when otherwise stated).

Total length 3.93—4.22, 4.03 mm (6). Wing length 2.10-2.19, 2.14 mm (4). Total length/wing

length 1.82—1.96, 1.90 (4). Wing length/length of profemur 2.44—2.55, 2.50 (4). Coloration brown

with central parts of scutellum conspicuously pale. Wing pale yellowish brown.

Head. Eye with weak microtrichia between all ommatids. AR 0.92-1.05, 0.98 with one antenna of

one specimen with an AR of 0.68. Ultimate flagellomere 424—484. 460 um long with one antenna of

one specimen with ultimate flagellomere 381 um long. Pedicel 129-160, 139 um wide. Temporal

setae 10-17, 14, including 3-5, 4 inner verticals; 2—5, 4 outer verticals; and 5-8, 7 postorbitals.

Clypeus 123-138, 132 um wide; with 6-13, 10 setae. Cibarial pump, tentorium and stipes as in

Fig. 1A, of doubtful specimen from Nordseter as in Fig. 1B. Tentorium 150-176, 162 um long,

35-55, 42 um wide. Stipes with reduced sclerotization in basal half and reduced median plate;

105-139, 120 um long; 19—38, 26 um wide (normal in possible specimen from Nordseter, see re-

marks). Palp lengths (micrometers): 29—45, 32; 41—62, 51; 109-149, 134; 86— 103, 98; 106-138, 125.

Third palpal segment with 2-4, 3(6) sensilla clavata at apex.

Thorax (Fig. 1C). Antepronotum with 6-14, 11 setae. Dorsocentrals 15—25, 18; acrostichals 5-9,

7, starting some distance from scutal projection ranging from !/3—!/2 the length of scutum, longest

acrostichal 38—56, 45 um long; prealars 7—9,8. Scutellum with 16-23, 19 setae.

Wing (Fig. 1D). VR 1.13—1.19, 1.16 (4). Brachiolum with 1—2, 2 setae; R with 11-15, 13 (6): R,

with 4-11, 7 (6); Ry,; with 7-10, 9 (6); M, ,, with 0-8, 2 setae; other veins bare. Cell r,,; with 7—18,

14 (5) setae, apparently with about 50 setae in mature male pupa from Volda; cell m, ,, with 0-3,

1 seta; cell an with 0-1, 0 setae. Squama with 18—21, 19 setae.

Legs. Spur of front tibia 47—56, 50 um long; spurs of middle tibia 35—38, 36 um and 21-34, 25 um

long; of hind tibia 50-67, 62 um and 18—26, 22 um long.

Width at apex of front tibia 44-50, 47 um (4); of middle tibia 47—53, 49 um (4); of hind tibia

56-59, 56 um (4). Comb with 11-12, 11 setae; shortest seta 21—29, 24 um long; longest seta 53—70,

61 um long. Sensilla chaetica absent. Lengths (micrometers) and proportions of legs (n = 4):

P 849-860

857

Br 801-860

837

IP 884-956

935

884-920

905

777-848

822

920-1004

980

681-717

696

406-418

412

568-6.21

599

341-382

360

227

311-335

323

245-263

258

167-173

170

251-263

260

ta,

179-185

181

131

155-167

158

ta,

126-132

127

114-120

116

126-132

131

0.76-0.78

0.77

0.49-0.52

0.50

0.60-0.62

0.61

2.58-2.70

2.66

3.10-3.27

3.22

2.80-2.94

2.89

2.47-2.56

2.53

3.83-4.12

4.02

3.15-3.28

3.20

Fig. 1. Heterotrissocladins spp, male imago: A, C-E. H. brundini spec. nov; B,F. H. brundini or maeaeri (Brun-

din); A, B. Cibarial pump, tentorıum and stipes; C. Thorax; D. Wing; E-F. Hypopygium.

Hypopygium (Fig. IE). Anal point 53-70, 63 um long; tergite IX including anal point with

20-27, 23 setae; laterosternite IX with 4-8, 7 setae. Phallapodeme 117-139, 125 um long; transverse

sternapodeme 106-116, 111 um (6) long. Virga very weak and inconspicuous, consisting of one or

two clusters of all together 6-7 spines; 11—15, 13 um (5) long. Gonocoxite 255—278, 268 um long;

inferior volsella rounded, with long setae on distal side. Gonostylus 117-141, 127 um long; outer

margin rounded; crista dorsalis long, relatively pronounced, rounded near apex; megaseta 10-15,

12 um long. HR 1.96-2.28, 2.12; HV 2.95— 3.46, 3.18.

Female imago (n = 9-11, except where otherwise stated).

Total length 3.28—4.00, 3.61 mm. Wing length 2.11—2.33, 2.18 mm. Total length/wing length

1.51—1.80, 1.66. Wing length/length of profemur 3.01—3.24, 3.12 (8). Coloration as in male.

Head. Flagellomere lengths (micrometers): 83-109, 98; 45-60, 55; 53-64, 59; 56-69, 64;

120-156, 136. Pedicel 68-82, 73 um wide. AR 0.44—0.55, 0.50. Temporals 9— 14, 12; including 4—6,

5 inner verticals; 2-5, 3 outer verticals; and 2-5, 3 postorbitals. Clypeus 138-161, 151 um wide;

1.85— 2.22, 2.06 times as wide as pedicel; with 7—15, 11 setae. Tentorium 138— 154, 146 um (8) long;

18-26, 22 um (8) wide. Stipes 86-149, 118 um (8) long; 9—23, 16 um (8) wide. Palp segments length

(micrometers): 26-35, 31; 38—49, 43; 82-103, 94; 68—88, 78 (7); 98-135, 118 (7). Two specimens

with fourth and fifth palp segments fused; 120-135 um long. Third palpal segment with 3-5, 4 sen-

silla clavata at apex. Coronal suture incomplete; 64—98, 82 um long.

Thorax. Antepronotum with 8-17, 11 lateral setae. Dorsocentrals 14—29, 19; acrostichals 4—7, 5;

prealars 6-10, 8. Scutellum with 16-24, 19 setae.

Wing. VR 1.11-1.16, 1.13. Brachiolum with 2 setae; R with 12-25, 17; R, with 5-13, 8; R,,; with

12-33, 24; M, ,, with 0—2, 1 setae; other veins bare. Cell r,,; with 23—54, 36; cell m, ,, with 4-17,

13 (8) setae; cell an with 2-14, 6 (8) setae. Squama with 15—25, 21 setae.

Legs. Spur on front tibia 41-56, 50 um long; spurs on middle tibia 30—41, 37 um and 23—34,

28 um long; on hind tıbia 60-71, 66 um and 18—26, 24 um long. Width at apex of front tibia 38—47,

43 um; of middle tıbıa 41—53, 48 um; of hind tıbia 49—59, 53 um. Comb of 7-10, 8 setae; shortest

seta 23—34, 29 um long; longest seta 49—67, 56 um long. Sensilla chaetica 1-7, 4 at 0.20—0.30, 0.28

(8) to 0.40—0.60, 0.53 on ta, of hind leg; apparently absent in mid leg. Lengths (micrometers) and pro-

portions of legs (n = 7-9):

fe ti ta, ta, ta, ta, ca, LR BV SV BR

PJ 669-753 753-848 548-624 274-321 194-217 137-151 99-131 0.72-0.76 2.68-2.92 2.47-2.63 2.1-3.1

697 793 582 291 200 143 106 0.74 2.87 2.54 2.9

PJ 690-807 717-807 359-384 191-227 142-167 95-120 85-114 0,.49-0,51 3.23-3.47 3.76-4.10 2.1-3.0

716 749 Shit 203 149 106 99 0,50 3081 3.89 ED

pP, 784-896 884-992 548-624 293-335 227-263 135-161 99-134 0.61-0.65 2.78-2.98 2.88-3.10 2.9-3,9

817 921 576 - 303 242 144 114 0.63 2.88 3.02 Sr3

Abdomen. Number of setae on tergites I-VII as: 33—53, 42; 38—46, 42; 26-40, 25; 24—44, 34;

24—38, 31; 19-35, 28; 19—35, 24; 9—26, 16. Number of setae on sternites I-VIII as: 0; 3-7, 5;5-10,

7:8-17, 1131220, 15; 14-24, 17; 1523, 18, 4371, 56.

Genitalıa (Fig. 2). Gonocoxite with 13—20, 16 setae. Tergite IX well divided, with 20-31, 25 setae.

Cercus 114—142, 133 um long. Seminal capsule 76-101, 86 um (8) long; excluding 11-26, 17 um

long neck; 64-79, 71 um wide. Notum 105-161, 132 um long.

Pupa (n = 11, except when otherwise stated)

Total length 3.76—4.78, 4.31 mm (9). Length of thoracic horn/length of anal macrosetae 1.10— 1.50,

1.34. Thorax of exuviae pale brownish grey, abdomen nearly transparent.

Cephalothorax. Frontal warts (Fig. 3A) conspicuous, 82-117, 97 um high; 35-53, 40 um (10)

wide at base. Frontal setae on frontal apotome; 59—83, 73 yum (8) long. Postorbitals 59— 73, 66 um (4)

and 41-56 um (2) long. Median antepronotals 117—147, 140 um (9) and 103—132, 120 um long; la-

teral antepronotals 73—88, 80 um and 0—44,9 um long (when absent reduced to setal mark). Thoracic

horn (Fig. 3B) 290-396, 349 um long; 44—70, 54 um (9) wide; 5.44—8.00, 6.57 (9) times as long as

wide. Anterior dorsocentral 44—88, 65 um long; Dc, 53— 117, 88 um long; Dec; 32—129, 68 yum long;

Apl

Fig. 2. Heterotrissocladius brundini spec. nov, female genitalia: A. Dorsal view; B. Ventral view; C. Lateral view;

D. Lobes of gonapophysis VII (DmL dorsomesal lobe; VIL, ventrolateral lobe; ApL, apodeme lobe).

Dc, 64-88, 78 um long. Distance between Dc, and Dc, 18-67, 42 um; between Dec, and Dec; 15-53,

28 um; between Dc, and Dec; 23—82, 58 um. Prealar observed in one specimen, 18 um long. Wing

sheath with fine marginal lines.

Abdomen (Fig. 3C, D). Shagreen absent on tergite 1 (T I) and T IX, weak median on T II, more ex-

tensive and stronger posterior on TIII—-VI, weak anteriomedian on T VII- VIII. Sternites I (S I) and

IX bare, S II-II with weak anterior and median shagreen, S TV- VII with weak anterior shagreen and

a few posterior spinules asS VI. Pedes spurii A present on sternites IV-VIland occasionally VIII (3 in-

dividuals). Pedes spurii B wider than high. Caudal hooklets 20-40, 32 on T II. Sternite VIII of male

with 19-33, 27 (7) posterior spines. Caudolateral corners of segments VI-VIII with imbedded spı-

nes, distinct on VII, only indicated on VI and VIII. L setae on segments I-VIII as: 1—2 (4): 3:3: 3:

3: 3: 3-4, 3: 5; 3 lJamelliform on VII, 4 lamelliform on VIII, occasionally also fıfth L seta on VIII

slightly lamelliform. Anal lobe 269-337, 302 um long; with 19—25, 22 setae in fringe; anal macrosetae

234—293, 261 um long. Genital sac of male overreaching anal lobe by 88-124, 109 um (4).

Larva (n ='6, except when otherwise stated)

Total length about 6 mm (larval exuviae). Head capsule length 447-502 um (3).

Head. Antenna as in Fig. 3E. Lengths of antennal segments (micrometers): 82—88, 86; 35—41, 36;

3—4, 4; 18-22, 19 (4); 6-9, 7 (4); 4—6, 4 (4). AR 1.07-1.16, 1.12 (4). Basal antennal segment 18-21,

20 um wide; 4.05—4.32, 4.24 times as long as wide; distance from base to annular organ 9-18, 15 um;

to distal mark of seta 21-35, 31 um; blade at apex 50-64, 50 um long; accessory blade 7-9, 9 um

long. Subapical style of second segment 6-9, 7 um (5) long. Labrum and epipharyngeal areas as ın

Fig. 3. Heterotrissocladins brundini spec. nov, pupa: A. Frontal apotome; B. Thoracic horn; C. Tergites of male;

D. Sternites of male. — larva: E. Antenna; F. Labrum and epipharyngeal area; G. Mandible; H. Maxilla; I. Mentum.

Fig. 3F. S I broad, finely plumose. Labral lamellae rounded apically. Median spine of pecten epipha-

ryngis narrower than lateral spines. Premandible 79-85, 83 um (5) long. Mandible (Fig. 3G)

158-176, 168 um (4) long. Maxilla as in Fig. 3H. Mentum (Fig. 31) with two close median teeth each

with distinct lateral accessory tooth, width of one median tooth including accessory tooth 18—21,

19 um; ventromental plates 21-26, 25 um (4) wide; 1.08-1.50, 1.32 times as wide as one median

tooth including accessory tooth. Postmentum distinctly darker than surrounding areas of head cap-

sule; 208-229, 222 um long.

Abdomen. Procercus 38-47, 43 um high; 29—32, 31 um wide; anal setae 647-747, 706 um (5)

long. Supraanal seta 234—264, 242 um long; 0.32—0.35, 0.34 (5) times as long as anal setae. Anal tubu-

les and posterior parapods not measurable.

Remarks

A male imago from Nordseter near Lillehammer is not included in the type material. Judging from

the leg proportions and the antennal ratio the specimen belong to A. brundini. The normally

sclerotized stipes (132 um long, Fig. 1B) and the more numerous setae on the wing membrane (45

setae in cell r,,;, 7 setae in each of m}, and m; , 4), however, indicate that ıt belongs to H. maeaeri. The

hypopygium (Fig. 1F) could belong to any ofthetwo species. Associated pupal material is needed to

decide whether the stipes and the wing setation is more variable within 7. brundini or whether the

specimen represents FH. maeaerı or an additional new species.

The pupa described by Langton (1984: 88) do not quite fit 77. brundini. The thoracic horn of Hete-

rotrissocladius Pe 1 of Langton is wider (length/width 4.0-5.1), and there are more numerous setae in

the anal lobe fringe (26-30). These variations, however, probably ıs not more than can be expected

between various populations.

Ecology and distribution

H. brundini has been found in two acidified lakes in Southern Norway, two lakes in the provinces

of More & Romsdal and Sogn & Fjordane, and ın Lake Assynt in the Sutherland District of Scotland.

It could conceivably be an indicator species of oligotrophic acid lakes.

Systematics

SETHER (1975: 57—62, fig. 15) does a phylogenetic analysis of Heterotrissocladius and related genera.

In the scheme of argumentation H. brundini will be synapomorphous for trends 57, 58, 59, 60; 9, 10,

11; and 51, 52, showing that the species is a good Heterotrissocladius. It is symplesiomorphous for

trends 41 and 44, apomorphous for trends 42 (reduced acrostichals) and 43 (reduced number of setae

on the wing membrane). The last two trends show parallelisms between the subpilosus and the maeaeri

groups. However, there are no pearl rows on the pupal wing sheaths (trend 37), only 4 filamentous se-

tae on segment VIII of the pupa (trend 39), and no setae dorsad of the genital sac (trend 40), showing

that the species belong with the maeaeri and marcidus groups. However, the pedes spurii A which may

occur on sternite VIII and the fifth L seta which in some specimen is somewhat filamentous, indicate

a plesiomorphic position within these groups.

The species is synapomorphous with A. maeaeri for trends 32 (short acrostichals) and 35 (distribu-

tion of L setae in pupa). It is plesiomorphous for trends 33 (PSB), 34 (PSA), 36 (median teeth of men-

tum), and apomorphous for trends 30 (setae on male tergite IX) and 31 (AR of larva), indicating a pla-

cement in the marcidus group. However, trends 30 and 31 are not very significant, representing con-

tinuous variation and probably good only as specific autapomorphies. Also several additional synapo-

morphies between H. maeaeri and H. brundini can be found in the male hypopygium in the shape of

the inferior volsella, the inner margin of the gonocoxite and in the shape of the gonostylus. We feel re-

latively certain about the phylogenetic position of the species as the sister species of H. maeaeri.

There are few problems fitting the imagines and the pupa within the concept of the maeaeri group

(SETHER 1975: 22); the stipes of the imagines and the pedes spurii B of the pupa have to be deleted

as dıstinguishing characters. However, the larva will key to H. marcidus (Walker) in the larval key

(SETHER 1975: 9) because of the two median teeth of the mentum. The median teeth, however, are

closer together than in other described species except for Heterotrissocladius sp. E from Lake Superior

(Sather 1975: 55) which is mentioned as intermediate between the marcidus and maeaeri groups.

Heterotrissocladius sp E almost certainly is close to 7. brundini. A redefinition of the maeaeri group

to include larvae with median teeth close together and distinct accessory teeth will make a separation

between the maeaeri and marcıdus groups still feasıble.

The imbedded spines found in the pupa is nearly unique within the orthoclads, otherwise found in

some members of Zalutschia Lipina only. S#rHer (1976) did not attempt to classify Oliveridia Szxther,

Hydrobaenus Fries, Trissocladius Kieffer and Zalutschia combined as a monophyletic group. Accor-

ding to S£rHer (1977: 82), however, the genera apparently form a monophyletic unit based on the

shape of the female gonapophysis VIII. This trend, however, is rather insecure as similar gonapophy-

ses occur elsewhere including in the Heterotrissocladins group of genera. Itthus is possible that Zalut-

schia is more closely related to the Heterotrissocladius group than to Hydrobaenus, and that the im-

bedded spines represent an underlying synapomorphy occurring in some, bot not allmembers of both

Zalutschia and Heterotrissocladius. We have observed indication of imbedded spines also in some ex-

uviae of A. marcidus. On the other hand the imbedded spines merely represent a strengthening of the

paratergital margin and could easily be a result of parallel selection.

The larvae of Heterotrissocladins all have hair-like spines at base of the anterior parapods similar to

those illustrated by Srrenzke (1950 fig. 11) for Paraphaenocladius Thienemann, but not found in Pa-

rametriocnemus Goetghebuer or apparently in the same form in other orthoclads. However, other

synapomorphies seem to confirm the relationship shown by Sruer (1975 fig. 15) and these spines

must be regarded either as secondarily reduced in Parametriocnemus or as an underlying synapomor-

phy for the whole group. |

Acknowledgements

We would like to thank Laboratory for Freshwater Ecology and Inland Fisheries Bergen for financial support.

We also are indebted to Mr. Tien van Trieu for doing the illustrations and to Mrs. N. Waagsberg for typing the ma-

nuscript.

Literature

LANGTON, P. H. 1984: A key to pupal exuviae of British Chironomidae. — P.H. Langton, March, Cambridgeshire,

324 pp

SETHER, OÖ. A. 1975: Nearctic and Palaearctic Heterotrissocladius (Diptera: Chironomidae). — Bull. Fish. Res. Bd

Can. 11931167

—- 1976: Revision of Hydrobaenus, Trissocladius, Zalutschia, Paratrissocladius, and some related genera (Diptera:

Chironomidae). — Bull. Fish. Res. Bd Can. 195: 1—287

—— 1977: Female genitalia in Chironomidae and other Nematocera: morphology, phylogenies, keys. — Bull. Fish.

Res. Bd Can. 197: 1- 209

—- 1980: A glossary of chironomid morphology terminology (Chironomidae: Diptera). — Ent. scand. Suppl. 15:

1-51

STRENZKE, K. 1950: Systematik, Morphologie und Ökologie der terrestrichen Chironomiden. — Arch. Hydrobiol.

Suppl. 18(2): 207—414

Prof. Ole A. S&ther

&yvind Schnell

Museum of Zoology

University of Bergen

N-5007 Bergen, Norway

SPIXIANA Supplement 14 | 65-74 | München, 15. Juli 1988 ISSN 0177-7424

Two new species of the Rheocricotopus (R.) effusus group

(Diptera, Chironomidae)

By Ole A. Szther & ©yvind A. Schnell

Abstract

Two new species belonging to the Rheocricotopus (Rheocricotopus) effusus group are described; R. (R.)reduncus

spec. nov. as male imago and pupa; R. (R.) unidentatus spec. nov. as male and female imago, pupa and larva. R. (R.)

reduncus is intermediate between R. (R.) paucıiseta Szther and the rest of the effusus group. R. (R.) unidentatus ıs

intermediate between R. (R.) effusoides Sether and R. (R.) effusus (Walker).

Introduction

The genus Rheocricotopus Thienemann & Harnish, 1932, recently was revised (S£THER 1985). It was

divided into two subgenera, Psilocricotopus Sether and Rheocricotopus s. str. Of the three species

groups in Rheocricotopus s. str. only the effusus group clearly was shown monophyletic.

Recently Michael Bolton from the Ohio EPA in Columbus, Ohio, sent the senior author some rear-

ed specimens collected in a spring stream in Ohio. These specimens showed a puzzling combination

of characters. The pupa appeared to belong to AR. (R.) effusoides SETHER, while the adults apparently

belonged in R. (R.) effusus (Walker). The associated larva, however, had a single median mental tooth

and thus resembled R. (R.)tuberculatus Caldwell. A closer examination showed that also the pupa and

the adult could be separated from the closest related species although perhaps only in insignificant de-

tails. The species is described below as R. (R.) unidentatus spec. nov.

The junior author during investigations around the Jostedal Glacier, found some males and pupal

exuviae of a species of the effusus group in the partly glacier fed rıver Jostedola. The males appeared

quite similar to R. (R.) panciseta Szther except for the quite distinct superior volsella, while the pupa

showed similarities to R. (R.) tuberculatus with L-setae on segments V—-VIII as 3:3:3:4—5. The spe-

cies ıs described below as R. (R.) reduncus spec. nov.

Methods and morphology

Morphological nomenclature follows SETHER (1980), with the exception that the apical spine of the gonostylus

istermed the megaseta. The measurements are given as ranges followed by amean when four or more measurements

are made, followed by the number measured in parentheses (n). In the figures of the male hypopygia the dorsal

aspect is shown to the left, the ventral aspect and the apodemes to the right.

The type material is kept at the Museum of Zoology, University of Bergen, Norway (ZMB).

Rheocricotopus (Rheocricotopus) reduncus spec. nov.

(Eigsal; 2A AGHE)

Type localıty: Jostedola River, Luster, Sogn & Fjordane, Norway.

Type material: Holotype, male, Jostedola River at inletto smalllake, Luster, Sogn & Fjordane, Norway, 23/7/86,

A. Fjellheim and ©. A. Schnell (ZMB No. 116). Paratypes: 3 males, 1 male pupa, 7 pupal exuviae, as holotype.

Diagnostic characters

The strongly curved, narrow-based, tooth-like projection of the superior volsella combined with

the small and indistinct humeral pit will separate the male from other members of the genus. The me-

dian spine patches on the tergites grading over in shagreen, presence of pedes spurii B on segment II,

combined with L setae of segments V— VIII as 3:3:3:4—5 will separate the pupa from other known pu-

pae of the genus.

Etymology: From Latin, reduncus, curved (hooked) backwards, referring to the shape of the superior volsella.

Description

Male imago (n = 4, except when otherwise stated).

Total length 2.69—2.90, 2.80 mm. Wing length 1.58-1.73, 1.94 mm. Total length/wing length

1.41—1.47, 1.45. Wing length/length of profemur 3.01—3.11, 3.06. Coloration dark brown.

Head. AR 0.83—0.90, 0.87. Ultimate flagellomere 402-442, 426 um long. Temporal setae 5-7, 6;

including 0-1, 1 inner vertical; 0-2, 1 outer vertical; and 3—5, 4 postorbitals. Clypeus with 10—20,

16 setae. Cibarial pump, tentorium and stipes as in Fig. 1A. Tentorium 149-155, 151 um long;

29—31,29 um wide. Stipes 149— 161, 154 um long; 53—64, 60 um (3) wide. Palp segments length (mi-

crometers): 23—29, 27; 59—64, 62; 88-104, 99; 94— 105, 100; 155-177, 167. Third palpal segment

with 1-3, 2 sensilla clavata.

Thorax (Fig. 1B). Antepronotum with 6-9, 8(5) lateral setae. Humeral pit small and indistinct.

Dorsocentrals 7—9, 8(5); acrostichals 7—9, 8, all situated in front except for in one specimen with an

additional median seta, longest acrostichals 44—47, 46 um (5) long; prealars 2-3, 3 (5). Scutellum with

610,85) setae.

Wing (Fig. 1C). VR 1.10-1.16, 1.13. Wing membrane with punctation visible at 250X. Anal lobe

reduced. C extension 35—59, 52 um long. Brachiolum with 1-2, 1 seta; R with 6-8, 7; R, with 0-1,

0; R,;; with 0-1, 1; C extension with 0-3, 1 non-marginal setae. Squama with 2-3, 3 setae.

Legs. Spur of front tibia 43—47 um (2) long; spurs of middle tibia 18—25, 21 um (5) and 15 um (2)

long; of hind tibia 44-50, 47 um and 16-18 um (2) long. Width at apex of front tibia 38-41, 40 um;

of middle tıbia 41 um; of hind tıbia 47 um. Comb of 12-14, 13 setae; shorter seta 21—23, 23 um (5)

long; longest seta 44—56, 49 um long. Sensilla chaetica0—1,0(5) at 0.25 of ta, of middle leg; 1-3, 2 (5)

at 0.20—0.35 of ta, of hind leg. Lengths (micrometers) and proportions of legs:

de ige ta, ta, ta, ta, u LR BU SV BR

PJ 609-681 735-807 490-526 311-335 221-227 137-149 84-90 0,65-0.67 2.42-2,.52 2.74-2.83 2.4-2.7

645 777 511 325 226 143 89 0.66 2.48 2.78 2.6

PJ 681-741 681-735 347-376 197-125 143-155 72-84 72-84 0.50-0.51 3.42-3.52 3.92-4.02 3.0-3.3

sul zııl 360 208 148 80 8l 0,50 3.48 3.95 BR

pP, 645-729 789-878 454-478 245-263 191-209 90-108 71-84 0.53-0.58 3.11-3.20 3.16-3.39 3.4-4.1

689 843 466 254 200 98 78 0.56 3.16 3.29 3.9

Hypopygium (Fig. 1D). Anal point 38 um (2) long; with 6-11, 9(5) setae; laterosternite IX with

3-5,4(5) setae. Phallapodeme 67 —76, 72 um long; transverse sternapodeme 79—98, 88 um long. Go-

nocoxite 202-220, 208 um long; with strongly developed, digitiform, narrowbased, caudomesal pro-

jection of superior volsella, superior volsella 59—67, 64 um (5) long; inferior volsella single, blunt-

Fig. 1. Rheocricotopus (Rheocricotopus) reduncus spec. nov., maleimago: A. Cibarial pump, tentorium and stipes;

B. Thorax; C. Wing; D. Hypopygium.

tipped. Gonostylus 105-111, 109 um long; crista dorsalis low, very long and relatively well devel-

oped, but appearing as absent in some views; megaseta 19—21, 20 um (3) long. HR 1.82—2.00, 1.90;

HV 2.48-2.63, 2.56.

Pupa (n = 8, except when otherwise stated).

Total length 3.38—3.80, 3.57 mm (7). Length of thoracic horn/length of anal macrosetae 1.00-1.17,

1.12 (7). Exuviae relatively dark greyish brown.

Cephalothorax. Frontal seta 44—73, 64 um (7) long; on frontal apotome (Fig. 2A). Postorbitals

44— 73,63 um (7) long. Median antepronotals 147— 190, 172 um (7) and 88— 176, 134 um long; lateral

antepronotal 50-73, 59 um long. Thoracic horn (Fig. 2C) 264—308, 295 um (7) long; 47—56, 53 um

(4) wide. Anterior precorneal seta 132-176, 154 um (6) long; median seta 88-147, 129 um (4) long;

posterior seta 44—59, 48 um 15 long. Anterior dorsocentral (Dc,) 26-82, 47 um long; De, 32-103,

75 um long; Dc, 26-53, 33 um long; De, 59— 100, 72 um long. Distance between De, and De, 23—38,

28 um; between Dc, and Dc; 21-41, 31 um; between Dc; and De, 50-11, 70 um.

Abdomen (Fig. 2E). Shagreen absent on tergitel(T I), weak anterior on T II, stronger and extensive

on T II-VI, reduced laterally on T VII- VIII, present anterior on T IX. Sternites1(S I) and IX bare,

shagreen anterior on S II; S III-IV with anterior and lateral S V-VII with extensive, S VIII with an-

terior shagreen. Pedes spurii A on S III-VI, sometimes indicated on S VII. Pedes spurii B on segment

II, well developed. T II with about 120-170, 49 caudal hooklets. Conjunctives IIV/IV, TV/V and so-

metimes V/VI with anteriorly directed spinules. Number of spines in median patches on T IV—Vlas:

10-45, 29; 35—80, 68; 55—80, 70. Maximal length (micrometers) of spines as: 6-12, 9; 12—18, 15;

9—18, 14. L setae on segments I-VIIl as: 2,3, 3, 3,3, 3, 3, 4-5; all lamelliform on segments VII and

VIII. Anal lobe with 16-20, 18 setae in fringe; anal macrosetae 249-270, 263 um long. Genital sac

of male folded and not measurable in males, of female ending 64 um (1) short of apex of anal lobe.

Larvae unknown

Remarks

The systematic positions of this and the following species are treated in the systematics part at the

end of this paper.

Ecology and distribution

The male imagines, pupa and pupal exuviae all were collected in a back eddy where the river Jostedola enters a

small lake at Viva (alt. 850 ma.s. |.) inthe uppermost part of the valley Jostedal. The type of locality is situated above

the timber line only a few kilometers east of the Jostedal Glacier, which is the largest glacier on mainland Europe,

covering an area of approx. 486 sq. km.

Some environmental parameters and a list of some other chironomids found at the locality is given by SCHNELL

& SA£THER (1987). At least three species, Vivacricotopus ablusus Schnell & Szther, a new species of Limnophyes and

R. (R.) reduncus spec. nov. appear to be endemic to the area.

Rheocricotopus (Rheocricotopus) unidentatus spec. nov.

(Bigs2B, DIE: 3:2)

Type locality; Camp Lazarus, Delaware Co., Ohio, U.S.A.

Type material: Holotype, male with pupal and larval exuviae, spring stream, Camp Lazarus, Delaware Co.,

Ohio, U. S. A., 18/4/86, M. J. Bolton, in coll. Mus. Zool. Univ. of Bergen (ZMB No. 117).

Paratypes: 2 males reared from larva, 2 females, reared from larvae, 3 larvae, as holotype; 3 larvae as holotype ex-

cept 1/4/86.

Diagnostic character

The high number of dorsocentrals (18—22 in males, 18—33 in females) and acrostichals (30-36 ın

males, 43—47 in females) will separate the imagines from R. (R.) effusus and R (R.) effusoides, the clo-

sest relatives. The pupa has more numerous spinules (65— 120) in the spine patches of T V and VIthan

in R (R.) effusus, and a pale greyish brown coloration of the exuviae separating it from the darker co-

lour of R. (R.) effusoides. The single median tooth of the mentum combined with the lack of ventral

tubercles on the head capsule will separate the larva from other members of the genus.

Etymology: From Latin, »ni-, one, and dentatus, toothed, referring to the single median tooth of the larval men-

tum.

Description

Male imago (n = 3, except when otherwise stated).

Total length 3.08-3.42 mm. Wing length 1.51-1.83 mm. Total length/wing length 1.87— 2.04.

Wing length/length of profemur 2.59— 2.69. Coloration brown.

Head. AR 1.10-1.36. Ultimate flagellomere 394-563 um long. Temporal setae 4—8, including

0-1 inner verticals, 1-4 outer verticals and 3 postorbitals. Clypeus with 10-14 setae. Cibarial pump,

tentorium, and stipes as in Fig. 3A. Tentorıum 146-169 um long, 38-47 um wide. Stipes

135—158 um long, 45—53 um (2) wide. Palp segments length (micrometers):38—41,68—81, 90-116,

101-124, 69—184. Third palpal segment with 2-3 sensilla clevata.

Thorax (Fig. 3B). Antepronotum with 4—8 lateral setae. Humeral pit very large, elongate ellipsoid

with smaller rounded pit below. Dorsocentrals 18-22; acrostichals 30-36, the longest reaching

SS SS

NIISSN

— N

II AS

\N h N \ ERKTILLLLNET \ Ler) Y

N \ < \ SS - S — ANA TR [d =

ze) km : 3,1, > > Sum E e} FEFZET

— R

2 > l IS 5

n IL ENS SS

Fig. 2. Rheocricotopus (Rheocricotopus) spec., pupae: A-B. Frontal apodeme; C-D. Thoracic horn; E-F. Ter-

gites. A, C, E. R. (R.) reduncus spec. nov.; B, D, F. R. (R.) unidentatus spec. nov.

Fig. 3. Rheocricotopus (Rheocricotopus) unidentatus spec. nov., male imago: A. Cibarial pump, tentorium and stıi-

pes; B. Thorax; C. Wing; D. Hypopygium.

26-38 um in length (15—26 um in R (R.) effusus and R (R.) effusoides); prealars 4. Scutellum with

6-9 setae.

Wing. (Fig. 3C). VR 1.08-1.14. Wing membrane with fine punctation of microtrichia visible at

150X. Anal lobe protruding. C extension 38—49 um long. R with 6-10 setae. Squama with 3-11 se-

tae.

Legs. Spur of front tibia 49—64 um long, spurs of middle tibia 21-26 um and 19—23 um long, of

hind tibia 45-58 um and 19—23 um long. Width at apex of front tibia 34—41 um, of middle tibia

38-49 um, of hind tibia 41-53 um. Comb of 11-12 setae, shortest seta 30—34 um long, longest seta

49—60 um long. Sensilla chaetica absent. Lengths (micrometers) and proportions of legs:

fe ti ta, ta, ta, ta, ta, LR BV SV BR

PJ 567-709 662-841 493-595 265-340 180-236 123-151 61-80 0.71 2.60-2.71 2.56-2.60 2.2-2.8

E 576-747 586-747 302-395 „1512-208 115-151 57- 80 47-66 0.52-0.53 3.58-3.97 3.72-3.84 2.5-2.7

P, 558-709 671-869 397-491 203-274 165-222 80-113 57-71 0.57-0.59 3.04-3.22 3.02-3.21 3.2-4,4

Hypopygium (Fig. 3D). Anal point 41—60 um long, with 8-12 setae, laterosternite IX with 5-6

setae. Phallapodeme 79-109 um long, transverse sternapodene 99—154 um long. Gonocoxite

218-251 um long; with well developed tooth-like caudomesal projection on superior volsella,

62-71 um long; inferior volsella single with pointed apex. Gonostylus 90—101 um long; crista dor-

salis low and long, relatively well developed, but not visible in some views; megaseta 13—15 um long.

FHR2422250; HV 3.393,47.

Female imago (n = 2, except when otherwise stated).

Total length 3.34—4.20 mm. Wing length 1.81-2.07 mm. Total length/wing length 1.85—2.03.

Wing length/length of profemur 2.61-2.65. Coloration yellowish brown with dark brown separate

vittae and thoracic markings, scutellum yellowish brown ın central area.

Head. Flagellomere lengths (micrometers): 98-105, 56, 53-60, 60-64, 113-116. AR 0.41—0.45.

Temporals 7-18, including 1-4 inner verticals, 2-8 outer verticals and 4-6 postorbitals. Tentorium

180 um long, 38-41 um wide. Stipes 161-180 um long, 49—68 um wide. Palp segments length (mi-

crometers): 41-49, 79—86, 113-114, 128-133, 218—225. Third palpal segment with 2-3 sensilla cla-

vata at apex. Coronal suture complete.

Thorax. Antepronotum wıth 8-10 lateral setae. Humeral pit as in male. Dorsocentrals 18-33;

acrostichals 43-47, longest 38—49 um long; prealars 6-7. Scutellum with 19—22 setae.

Wing. VR 1.01-1.16. C extension 79—90 um long. Brachiolum with 1-2 setae, R with 22-37, R,

with 10-17, R,,; with 26-51, extended part of costa with 5-8 non-marginal setae. Squama with

9722’ setae.

Legs. Spur of front tibia 30-38 um long, spurs of middle tıbia both 23-26 um long, of hind tibia

53-56 um and 21-23 um long. Width at apex of front tıbia 43—53 um, of middle tibia 45-56 um,

of hind tıbia 53-73 um. Comb of 12-14 setae, 30-60 um long. Sensilla chaetica apparently absent.

Lengths (micrometers) and proportions of legs:

fe zat ta, ta, ta, ta, ta, LR BV SV BR

P} 680-784 765-954 468-586 255-302 170-217 113-142 66-80 0.61 3.15-3.16 2.98-3.09 1.5-2.0

P, 718-803 713-851 317-425 161-217 113-161 57- 85 57-76 0.44-0.50 3.79-4.51 3.89-4.52 1.5-1.8

pP, 671-794 784-959 421-539 208-279 161-227 76-109 61-85 0.54-0.56 3.26-3.71 3.25-3.46 1.3 (1)

Abdomen. Number of setae on tergites I- VIII as: 27—48, 34—66, 32-55, 39—55, 40-59, 39 —49,

43, 36-47. Number of setae on sternites I-VIII as: 0, 0-3, 2-6, 7-11, 19-21, 23—29, 20-27,

28—46.

Genitalıa (Fig. 4A-C). Gonocoxite wıth 13— 17 setae, including 7—8 strong and 5-10 weak setae.

Tergite IX strongly divided with 22-26 setae. Cercus 90-128 um long. Seminal capsule 101-116 um

long, including 23-30 um long neck; 77—90 um wide. Notum 139 um long.

Pupa (n = 5, except when otherwise stated)

Total length 3.31—4.83, 3.92 mm. Length of thoracic horn/length of anal macrosetae 0.87—1.22,

1.00. Exuviae pale greyish brown, with darker apophyses and thoracic markings.

Cephalothorax. Frontal seta 90-120 sum (2) long, on frontal apotome (Fig. 2B). Vertical not mea-

surable; postorbitals 23—49, 35 um (4) long. Median antepronotals both 120-199, 163 um long; late-

ral antepronotals 41-88, 62 um long and small peg, 11-19 um (3) long. Thoracic horn (Fig. 2D)

259-390, 303 um long; 38—53, 44 um wide. Anterior precorneal seta 131—233, 186 um long; median

seta 83-218, 165 um long; posterior seta 49—161, 95 um long. Anterior dorsocentral (Dc,) 75-105,

82 um long; other dorsocentrals each 41-101, 65 um long. Distance between De, and Dec, 64-116,

92 um; between Dc, and Dec, 15-83, 50 um; between Dc; and Dec, 23-53, 37 um.

Abdomen (Fig. 2F). Shagreen absent on tergite I (T I), weak median on T II, stronger and more ex-

tensive on T III- VI, median on T VII- VIII, anterior on T IX. Sternites I (S I) and IX bare; shagreen

on $ II-IIl anterior, median and laterally; on S TV—VI posterio-laterally, not very strong; on S VII—

VIII anterior group shagreen. Pedes spurii A present on S IV-VI with indications also on S VII-

VIII in most specimens. Pedes spurii B well developed on segment II and present also on segment

III. About 140-260, 150 caudal hooklets on T II. Conjunctives IIV/IV, IV/V and usually V/VI with

rows of anteriorly divided spinules, medially interrupted or (in one specimen) absent on V/VI. Num-

ber of spinules in median patches on T IV-VI as: 34-75, 49; 65-118, 79; 68-120, 91. Maximal

Fig. 4. Rheocricotopus (Rheocricotopus) unidentatus spec. nov., female imago and larva: A-C. Female genitalia,

ventral (A) and dorsal (B) aspect and lobes of gonapohysis VIN (C) (DmL, dorsomesal lobe; VIL, ventrolateral

lobe; ApL, apodeme lobe); D. Larval mentum; E. Posterior end of larva.

lengths (micrometers) of spines as: 11—23, 15; 15—26, 19; 15— 26, 20. L setae on segments II- VIII as:

4,4,4,4,4, 4, 4-5; all lamelliform on segments VII-VIH. Anal lobe with 21—26, 23 setae in fringe;

anal macrosetae 281-319, 301 um long. Genital sac of male not measurable, of female ending

86—124 um (2) short of apex of anal lobe.

Larva (n = 10-11, except when otherwise stated)

Total length 3.97—7.24, 5.15 mm. Head capsule length 397—473, 438 um.

Head. Antenna as in R. (R.) effusoides Sther 1985 (fig. 21 A). Lengths of antennal segments (micro-

meters): 69— 87,78; 16—23, 18; 9—14,12;7—9,8;7—9,8. AR 1.55—1.96, 1.70. Basal antennal segment

15—24, 18 um wide, distance from base to ring organ 8-15, 10 um; to basal mark of seta 7-13, 9 um

(8); to distal mark 41—48, 44 um (7). Lauterborn organs and apical style of second segment each 6-9,

8 um long. Labrum as in R. (R.) effusoides (Szther 1985 fig. 21 B). Premandible 69— 79, 74 um long.

Mandible 116-158, 138 um long; with 7 branches in seta interna. Mentum (Fig. 4D) with one median

tooth with lateral notches; width of median tooth including notches 27-38, 32 um; ventromental pla-

tes 19—25, 22 um wide; with 28-40, 33 setae underneath. Postmentum 223—249, 233 um long.

Abdomen (Fig. 4E). Procercus 30-38, 34 um high; 21-28, 24 um wide; with 5-6 anal setae

450-563, 520 um long. Supraanal seta 98-131, 112 um long; 0.90—0.23, 0.21 times as long as anal se-

tae. Anal tubules 98-158, 118 um long; 38-56, 49 um wide at base. Posterior parapods 263—319,

290 um (8) long.

76%

Systematics

SETHER (1985) erected a scheme of argumentation delineating the cladogenesis of the genus Rheocri-

cotopus. Both species treated here are synapomorphous for trends 17, symplesiomorphous for trends

18, showing that they both belong in the nominal subgenus.

Trends 16 contain sıx different trends of which all except one for each species are symplesiomor-

phous for both species. In R. (R.) unidentatus, however, there is a single median tooth of the mentum

as in R. (R.)tuberculatus; and R. (R.) reduncns has L setae of the same distribution as in R. (R.)tuber-

culatus. However, none of these two trends are very significant since there is a tendency to reduction

to one median mental tooth in for instance R. (R.) effusoides, and to reduction of theL setae in the fs-

cipes group. Trend 15 is synapomorphous for R. (R.) unidentatus; unknown, but probably also syn-

apomorphous for R. (R.)reduncus. Both species are synapomorphous for trend 9, a highly significant

trend. It is thus clear that both species belong in the effusus group.

R. (R.)reduncus is somewhat intermediate for trends 8, while the species ıs synapomorphous for the

first of trends 7 (with tooth-shaped projection of the superior volsella), symplesiomorphous for the

second (with small humeral pit). The phylogenetic placement thus ıs relatively obvious. It form the si-

ster species of the rest of the effusus group minus R. (R.) pauciseta, while the latter form the sister spe-

cies of R. (R.) reduncus plus the rest of the group. An alternative placement could be as the sister spe-

cies of R. (R.) pauciseta alone, something further indicated by the similarity of the anal points of the

two species.

The placement of R. (R.) unidentatus also ıs relatively simple. Trends 7 and 3 are synapomorphous,

8 and 4 symplesiomorphous, meaning that R. (R.) unidentatus belong in a group with AR. (R.) effusus

and R. (R.) effusoides. Trends 1 and 2 indicate that R. (R.) effusus is the closest species.

R. (R.) reduncus and R. (R.) unidentatus ıs compared with their closest relatives in Table 1.

Table 1. Comparisons of some members of the Rheocricopus (R.) effusus group. Lengths in um.

R. pauciseta R. reduncus R. effusoides R. unidentatus R. effusus

Male:

AR 0.60 - 0.83 0.83 - 0.90 1.39 - 1.61 1.10 - 1.36 0.99 - 1.33

LRı 0.62 - 0.74 0.65 - 0.67 0.64 - 0.67 0.71 0.70 - 0.74

No. dorsocentrals 5-7 7-9 12-16 18 - 22 9-16

No. acrostichals 12-17 7-9 14 - 21 30 - 36 18 - 26

Max. 1. acrostichals 25 44 - 47 15 - 26 26 - 38 15 - 26

Huneral pit small small large, ellipsoid large, ellipsoid large, ellipsoid

Anal point setae 6-8 6-1 15 - 20 8-12 5-13

HR 2.61 - 2.69 1.82 - 2.00 1.92 - 2.07 2.42 - 2.50 -

Fenale:

No. acrostichals - - 21 - 2 43-47 18 - 26

Pupa:

Coloration pale yellow- dark greyish dark yellow- pale greyish pale greyish

ish brown brown ish brown brown brown

L. frontal seta 84 - 100 4-73 56 - 101 90 - 120 38 - 64

W. thoracic horn 48 - 58 47 - 56 49 - 86 38 - 53 34 - 53

Spines patch T IV ca. 0 - 15 10 - 45 6-43 34 - 75 22 - 46

N a TV ca. 25 35 - &0 36 - 79 65 - 118 28-47

a a ca. 30 55 - 80 35 - 74 68 - 120 26 - 55

No. setae in fringe 11-13 16 - 20 18 - 27 21 - 26 1-21

Larva:

Med. mental tooth double - double single double

L. basal antennal seen. 62 - 64 - 70 - 86 69 - 87 55 - 69

Acknowledgements

We would like to thank the Norwegian Water Resource and Elektricity Board for financial aupport. We also are

indebted to Mr. Michael Bolton, Ohio EPA, Columbus, Ohio, for the type material of R. (R.) unidentatus spec.

nov., to Mr. Tien van Trieu for doing the illustrations, and to Mrs. I. Wiese-Hansen for typing the manuscript.

Literature

SETHER, OÖ. A. 1980: A glossary of chironomid morphology terminology (Chironomidae: Diptera). — Ent. scand.

Suppl. 15: 1-51

—— 1985. A review of the genus Rheocricotopus Thieneman & Harnisch, 1932, with the description of three new

species (Diptera, Chironomidae). — Spixiana Suppl. 11: 59— 108

SCHNELL, ©. A. & O. A. SETHER 1988: Vivacricotopus, a new genus of Orthocladiinae (Diptera, Chironomidae).

— Spixıana Suppl. 14: 4955.

Prof. Ole A. Szther

&yvind A. Schnell

Museum of Zoology, University of Bergen

N-5007 Bergen, Norway

| SPIXIANA Supplement 14 75—84 | München, 15. Juli 1988 ISSN 0177-7424

A review of Lappodiamesa Serra Tosio, with the description of

L. boltoni spec. nov. from Ohio, USA

(Diptera, Chironomidae)

By Ole A. Sxzther and Endre Willassen

Abstract

The genus Lappodiamesa Serra-Tosio is emended. Lappodiamesa brundini Serra-Tosio is regarded as a junior

synonym of. vidua (Kieffer) comb. nov. The female of L. vidua and the male and female imagines, pupa and larva

of Lappodiamesa boltoni spec. nov. are described in full detail. The genus apparently forms the sister group of Psen-

dodiamesa Goetghebuer.

Introduction

The previously monotypic genus Lappodiamesa was described by SErrA-Tosıo (1968) from two ma-

les collected by L. Brundin ın N.-Sweden, close to the Norwegian border. The immature stages were

unknown when the keys and diagnoses to the Holarctic Diamesinae were elaborated by OLıver (1983,

1986). After the initial description of L. brundini Serra-Tosio no additional records were made of the

species until MAKARCHENKO (1983) described the male, pupa and larva from the Chukotskii Peninsula.

Having the opportunity to describe all stages of a recently discovered Nearctic species, we decided to

examine putative types of Syndiamesa vıidua Kieffer, a species which for a while has been suspected

to represent a synonym of L. brundini. Through the courtesy of Dr. E. Makarchenko, we also were

able to reexamine some specimens used in his description.

Methods

The material was mounted in Canadabalsam (Sx&ther, 1969) or in Euparal. Terminology follows S&ther (1980).

In the descriptions measurements are given as ranges. When more than 4 specimens have been measured, the range

is followed by a mean and the number of specimens measured in parenthesis.

Lappodiamesa Serra-Tosio, 1968, emended

Diagnosıs

Male antenna plumose, female antenna with six or seven flagellomeres. Eyes with weak or strong

pubescence, in male moderately extended medially. Frontal and orbital setae absent. Maxillary palp

about as long as width of head, with weak indication of sensilla capıtata on segment three or devoid

ofsensilla capitata. Antepronotum deeply notched dorsally. Acrostichals present or absent, dorsocen-

trals uniserial or biserial posteriorly. Wing membrane almost smooth (Makarchenko, 1983) or punc-

tated with microtrichia, R, distinctly arched, FCu clearly proximal to FR, MCu slightly proximal of

RM, R,;; with a few setae distally, anal lobe prominent. Fourth tarsal segment of legs cylindrical and

subequal in length to fifth segment. Male hypopygium with anal point and pars ventralis, with or with-

out rounded and setose superior volsella, and with weekly delineated inferior volsella. Female genita-

lia with 3 rounded seminal capsules, gonocoxite IX with moderate projection and tergite IX clearly

divided into two setigerous protrusions. Pupa with rugulose cephalothorax, thoracic horn absent; ab-

domen reticulate with strong and dark apophyses on tergites and sternites, L-setae simple or bifid and

mostly subequal; anal lobe overreaching genital sac, with 3 anal macrosetae, without fringe or median

setae, with or without pointed apical tubercle. Larva with 2 pairs of serrate labral lamellae, labral sen-

silla $S T-S III simple, pecten epipharyngis with 7 scales, premandible with 4-5 teeth, mentum with

1 median and 8-9 lateral teeth, body setae pale and moderately long, procercus about as long as wide

with 8-9 anal setae and 2 subapical setae.

Lappodiamesa vidua (Kieffer) comb. nov.

Fiıgs 1A-C

Syndiamesa vidna Kieffer, 1922: 23—24, fig. 15.

Lappodiamesa brundini Serra-Tosio, 1968: 140— 145, pl. 5. syn. nov.

Diagnosıs

Female with 6 flagellomeres. Acrostichals present. R,;; of wing ending close to R,; male without

superior volsella; female apparently without sensilla chaetica on front and middle legs. Pupa with dis-

tinct apical tubercle on anal lobe. Larva with first lateral teeth of mentum about equally wide as median

tooth.

Male imago

Scutellum with more than 30 setae in 2-3 irregular rows. LR, 0.64, LR;, 0.47, LR; 0.57; BV, 3.72,

SV, 2.72, SV, 2.20, SV; 3.38. Tergite IX (Fig. 1 A) with 22 setae, laterosternite with 14 setae; transverse

sternapodeme 131 um long, with or without distinct orolateral corners; phallapodeme 120 um long,

pars ventralis 34 um long, gonocoxite 304 um long, gonostylus 165 um long. Otherwise about as in

descriptions by Kırrrer (1922), SERRA-Tosıo (1968) and MAKARCHENKO (1983).

Female imago (n = 2)

Total length 4.53—4.58 mm. Wing length 3.37—3.58 mm. Total length/wing length 1.28—1.34.

Wing length/length of profemur 3.64—3.68.

Head: Pedicel with 3-4 setae. Length of flagellomeres (um): 83, 56-64, 53-56, 45-49, 49,

150-165. AR 0.57—0.59. Flagellomeres 1-3 with 1-2 apical sensilla coeloconica dorsally, ultimate

flagellomere with about 20 sensilla chaetica and a few sensilla coeloconica. Ultimate flagellomere with

1—2 subapical setae. Coronal suture faint or reduced. Temporal setae 17— 20, including 6-8 postorbi-

tals, 7—9 inner- and 3—4 outer verticals. Eyes with strong pubescence (sensu S£THER, 1980). Clypeus

with 5-7 setae. Palp segments 4 and 5 fused in one specimen; palp segments length (um) in normal

specimen: 45, 64, 131, 116, 154; about 4—5 sensilla clavata observed on palp segment 3. Tentorıium

180—214 um long. Stipes length/width 191/60—68 um.

Thorax: Antepronotum with 9-12 lateral setae. Dorsocentrals 16-17, acrostichals 13—16, supra-

alars absent, prealars 10-13. Scutellars 38—43, in 3—6 rows.

Wing: VR 0.92-0.96. Costa produced 83—90 um beyond R, ;;. Anal lobe with well developed pro-

jection. Microtrichia of wing membrane visible under 100X magnification. R with 10-13 setae, R,

with 7—9, R,;; with 6-9 setae. Alula with 4—5 setae. Squama with 40-50 setae. Subcosta with 3 sen-

silla campaniformia, R, with 1, R,,; with 2 and R,,; without sensilla campanıformia.

Legs: Spur of front tibia 68-71 um, spurs of middle tibia 49 um and 53—58 um, of hind tibia

45—51 um and 77-85 um long. Width at apex of front tibia 64— 71 um, of middle tibia 60 um, of

hind tibia 79—86 um. Comb on hind tibia with 7—9 setae 45—60 um long. Middle and hind legs with

following numbers of pseudospurs on ta, +; respectively: 4—9 plus 0-2 apical, 0, 0 (P,); 6-8 plus 2

Fig. 1. Lappodiamesa vidua (Kieffer) comb. nov., imago. — A. Male hypopygium. — B-C. Female genitalia, la-

teral (B) and ventral (C) view (cerci damaged).

apical, 0, 0 (P;). Front and middle ta, without sensilla chaetica; hind ta, with 18 sensilla chaetica

distributed from 0.29—0.39 to 0.83—0.88. Lengths (um) and proportions of legs:

fe ti ta, ta, ta, ta, ta, LR BV SV BR

Pl 926- 973 1106-1181 728-751 383 (1) 236 (1) 161 (1) 137 (1) 0.64-0.66 3.17 (1) 2.79-2.87 2.6 (1)

P, 1096-1143 1001-1158 576-595 293-321 208-217 132 113-123 0.51-0.52 3.65-3.72 3.81-3.87 2.9 (1)

P, 1247 1351-1427 756-832 389-444 265-288 142 (1) 142 (1) 0.56-0.58 3.59 (1) 3.22-3.44 2.2 (1)

Genitalia (Figs 1B-C): Sternite VIII with a total of 28-49 setae in amore or less contiguous distri-

bution; gonocoxapodeme strongly sclerotized. Seminal capsules rounded, 98-132 um long inclusive

19—23 um long neck, width 79-86 um; surface with possible indication of scattered, tubercle-like

microtrichia. Notum 109-120 um long, ramı long and curved towards midline. Flap (WırLassen,

1982) poorly developed. Ventrolateral lobe large with dense lanceolate microtrichia. Apodeme lobe

ill-defined. Gonocoxite rounded with 15—16 setae. Tergite IX clearly dıvided with a total of 35—50

setae.

Remarks

Kieffer based his original description on two males and one female collected by Dr. Okland on No-

vaja Semlja. According to Kırrrer (1922: 24) one male and the female were collected on 10th August

and labelled No. 239. The second male was collected 23th August and was labelled No. 258. After

completing his first description of Syndiamesa vidna, Kieffer received a second batch of specimens

which he published on later (see Kırrrer, 1923: 4, 11): two males plus one female collected on 19th Au-

gust, labelled No. 239, and three females with No. 228 and collection date 17th August.

We have studied one male and two females which according to the labels seem to belong to the se-

cond batch of Kırrrers specimens. However, they are all labelled “Types” and the male carries two ad-

ditional labels: one reading “Diamesa vidua” det. D. R. Oliver, the other “lectotypus det. D. R. Oli-

ver”. According to curator Mr. J. E. Raastad (personal communication to E. W.) four males (No. 239,

19th August) and one female (No. 228, 17th August) in addition to the ones examined here remain in

Museum of Oslo, and he indicates that “10th August” is an error which should read “19th August”.

Unfortunately, this does not seem to solve the problem of authenticity satisfactory and thus our ques-

tion of the validity of the lectotype-designation remains open. Nevertheless, based on the available

specimens and the detailed description of Serra-Tosıo (1968) it seems clear that Zappodiamesa brun-

dini ıs asynonym of S. vidna.

Material studied

Lectotype, male: Diamesa vidua (Kieffer) (D. R. Oliver det.) USSR: Novaj. Semlja, Ökland leg., Pankratjef Pe-

ninsula, 19. Aug. No. 239, (with additional numbers: Gl 2772 and 12463); Litselustina Bay, 17. Aug., No. 228

(with additional numbers 12461 and 12462), 2 females; in coll. Mus. Zool., Oslo. Chukotka, Chegitun River,

4. Aug. 1981, E. Makarchenko, 2 male hypopygia, 3 pupal exuviae, 3 larvae; 8. Aug. 1981, 1 larva; in coll. Mus.

Zool. Bergen.

Lappodiamesa boltoni spec. nov.

Figs 2-4

Type locality: USA, Ohio, Franklin County, Sharon Woods Park.

Holotype: Pharate male pupa labeled: USA, Oh., Franklin Co. Sharon Woods Pk, 3/25/86, leg. M. J. Bolton,

ZMBNNGo. 114.

Paratypes: 5 males, 2 females, 1 pharate female pupa with associated larval exuviae, 4 pupal exuviae, 1 larva; as

holotype. Types in coll. Museum of Zoology, University of Bergen, Norway.

Diagnosıs

Female with 7 flagellomeres. Acrostichals absent. Wing punctated with microtrichia visible under

30x magnification, R,,; ending in middle between R, and R,;;, hind leg without distinct tibial comb,

female with sensilla chaetica on ta, of all legs. Pupa without apical tubercle on anal lobe. Larva with

first lateral teeth of mentum clearly narrower than median tooth.

Descriptions

Male imago (n = 5, unless otherwise stated)

F-

“

SS N

Fig. 2. Lappodiamesa boltoni spec. nov., imago. — A. Male head. — B. Female head. — C. Cibarıal pump. — D.

Stipes. — E. Thorax. — F. Male wing. - G-L. Male hypopygium with varıation of anal point (H-]), details of su-

perior volsella (J), and gonostyli (K-L).

Total length 4.37—4.82, 4.58 mm. Wing length 2.63—2.84, 2.69 mm. Total length/wing length

1.68—1.81, 1.72. Wing length/length of profemur 2.55— 2.75, 2.65.

Head (Figs 2A, C-D): AR 2.32— 2.53, 2.46. Ultimate flagellomere 732—860, 819 um long. Coro-

nal suture complete. Temporal setae 12-21, 15 (6) including 6-11 postorbitals, 3—6, 5 outer verticals

and 1-5, 3 inner verticals. Eyes pubescent (not hairy) between allommatids. Clypeus with 0-8, 3 (6)

setae. Palps with long setae, sensilla clavata not apparent. Palp segments length (um): 38—43, 40;

90-113, 99; 150-169, 156; 116-137, 122; 124— 154, 142. Tentorium 203—244, 231 yum long. Cibarial

pump as in Fig. 2C. Stipes (Fig. 2D) length/width 158-188, 168/49—68, 57 um.

Thorax (Fig. 2E): Antepronotum deeply notched; with 3— 10, 6 lateral setae. Dorsocentrals 14— 26,

17 (4); supraalars and acrostichals absent; prealars 4—11, 6; Scutellars 14— 17 (2), more or less biserial.

Wing (Fig. 2F): VR 0.81—0.89, 0.84. Costa produced 94— 113, 99 um beyond R;;;. Ry,; in middle

between R, and R,,;. Microtrichia of wing membrane visible under 30% magnification. Brachiolum

with 1-2, 2 (4) delicate setae. R with 8-10, 9 setae; R, with 2-6, 4; R,,; with 2—3, 2 setae. Squama

with 22-55, 40 (6) setae. Subcosta with 3 sensilla campaniformia, R, with 1, R,,, with 3 and R,;;

without sensilla campaniformia. Anal lobe with well developed projection.

Legs: Spur of front tibia 71-83, 77 (4) um long; spurs of middle tibia 45-68, 54 and 44-49, 44

(4) um; of hind tibia 64—68, 65 and 41-47, 43 (4) um long. Width at apex of front tibia 53—73, 66

(6) um; of middle tıbia 60— 71, 65 um; of hind tibia 68-79, 74 um. Apex of hind tibia with somewhat

irregularly dispersed setae, but without distinct setal comb. Middle and hind legs with the following

numbers of pseudospurs on ta,_, respectively: 8-15, 11 plus 2 apical; 0-4, 2 plus 2 apical (P,); 6-14,

11 plus 2 apical; 1-4, 3 plus 0—2 apical (P;). Other tarsal segments without pseudospurs. Posterior

ta, with 6-13, 10 sensilla chaetica distributed from 0.66—0.83, 0.77 t0 0.93—0.95,0.94. Pulvilli weak.

Lengths (um) and proportions of legs:

fe ti ta, ta, ta, ta, ta, LR BV SV BR

Pl 983-1049 1210-1266 888-945 369-397 255-284 142-161 104-123 0.72-0.75 3.32-3.64 2.39-2.53 3.0-4,4

1015 1249 910 384 267 148 113 0.73 3.49 2.49 3.9

P, 1088-1134 1134-1200 633-661 279-302 208-217 123-132 98-113 0.54-0.58 3.87-4.03 3.41-3.66 2.7-4.0

1101 1164 647 294 213 128 106 0.56 3.94 3.50 3.

P, 1229-1380 1436-1539 805-841 416-444 265-321 142-175 104-118 0,.52-0.56 3.60-3.77 3.32-3.66 4.6-5.2

1304 1500 814 428 289 158 112 0.54 3.67 3.44 4,9

Hypopygium (Figs 2G-L): Tergite IX with 20-30, 25 setae. Laterosternite with 8-14, 10 setae.

Anal point (Figs 2G-I) 53-105 um, (apparently broken in 3 of 6 specimens), slender with delicate

apıcal hair sensillum or more stout with double to triple apical spines. Transverse sternapodeme ap-

proximately 95—150 um wide, without oral projections. Aedeagal lobe broad, rhomboid to spatulate;

phallapodeme well sclerotized, 101-180, 151 um long; pars ventralis (Fig. 2G) small and knoblike.

Gonocoxite 296—360, 338 um long; basally with rounded, setigerous superior volsellae (Figs 2G, J);

inferior volsellae weakly delineated. Gonostylus (Figs 2G, K-L) 150-210, 173 um long; with well

developed crista dorsalis, macroseta relatively short. HR 1.71—2.20, 1.97. HV 2.27—2.95, 2.67.

Female imago (n = 2)

Total length 5.06-5.39 mm. Wing length 2.86-2.94 mm. Total length/wing length 1.77—1.83.

Wing length/length of profemur 3.46.

Head (Fig. 2B): Pedicel without setae. Length of flagellomeres (um): 79-83, 45-53, 53—60,

45—53, 45-56, 49—62, 120-161; 7th flagellomere partially divided 45—68 um from base. AR 0.32.

Flagellomeres 1-6 with 1 pair of apical sensilla chaetica dorsally and 0-2 sensilla chaetica ventrally;

ultimate flagellomere with 18 sensilla chaetica; ringed sensilla coeloconica absent. Ultimate flagello-

mere with 1-2 subapical setae. Coronal suture incomplete or faint. Temporal setae 6-7, including

2-3 postorbitals, 2 inner- and 2 outer verticals. Eyes with weak pubescence which is mostly concealed

by overlying ommatid lenses. Clypeus with 0-4 setae. Palp segments length (um): 41-45, 83—90,

Fig. 3. Lappodiamesa boltoni spec. nov., female genitalia. — A. Ventral view. — B. Lateral view.

94-98, 83-90, 109-128; no sensilla clavata observed on palp. Tentorium 191 um (1) long. Stipes

length/width 154/71 um.

Thorax: Antepronotum with 4-6 lateral setae. Dorsocentrals 16-17, supraalars and acrostichals

absent, prealars 7. Scutellars 22-32, in biserial pattern.

Wing: VR 0.92-0.94. Costa produced 98-116 um beyond R,;;. Projection of anal lobe slightly

less developed than in male. Microtrichia of wing membrane visible under 30x magnification. Bra-

chiolum with 1—2 weak seta. R with 10-11 setae, R, with 5-7, R,;; with 5-6 setae. Alula with 5 se-

tae. Squama with 36—44 setae. Subcosta with 4 sensillacampaniformia, R, with 1, R,,, with3 and R, ;;

without sensilla campaniformia.

Legs: Spur of front tibia 60-64 um, spurs of middle tibia 45 um and 49—56 um, of hind tibia

45 um and 60-64 um long. Width at apex of front tibia 64 um, of middle tibia 64—68 um, of hind ti-

bia 71-75 um. Comb on hind tibia absent. Middle and hind legs with following numbers of pseudo-

spurs on ta,_, respectively: 7—10 plus 2 apical, 3-5 plus 2 apical, 0-1 (P,); 7—8 plus 2 apical, 0—3 plus

0-2 apıcal, 0 (P;). Front ta, with 4-9 sensilla chaetica distributed from 0.51—0.64 to 0.84—0.85;

middle ta, with 17-21 sensilla chaetica from 0.23—0.38 to 0.86—0.91; hind ta, with 8-18 sensilla

chaetica from 0.52—0.54 to 0.90—0.94.

Lengths (um) and proportions of legs:

fe ti ta, ta, ta, ta, ta, LR BV sv BR

PL 827- 851 973-1002 628-643 260-284 180-194 118-132 104-113 0.64-0.65 3.45-3.67 2.86-2.88 2.0-2.1

P, 983-1011 1044-1063 473-501 208-246 146-170 104-128 95-113 0.45-0.47 3.92-4.52 4.14-4.29 1.9-2.0

P 1096-1125 1257-1323 647-652 340 227-246 137-151 113-123 0.49-0.52 3.60-3.67 3.64-3.75 2.7

Genitalia (Figs 3A-B): Sternite VIII with 12-16 setae to each side of midline; gonocoxapodeme

narrow. Seminal capsules rounded, 139— 154 um long inclusive 8 um long neck, width 113—128 um;

surface covered with dense, very fine microtrichia (500X) and scattered, tubercle-like microtrichia.

Notum 94— 101 um long, ramı long and curved towards midline. Flap (Wırrassen, 1982) poorly devel-

oped. Ventrolateral lobe large with dense lanceolate microtrichia. Apodeme lobe with microtrichia.

Gonocoxite rounded with 14-16 setae. Tergite IX divided with 30-48 setae.

Pupa (n = 5, unless stated otherwise):

Total length 5.98—6.51, 6.20 mm. Exuviae brownish grey. Thorax rugulose. Abdomen with dark

brown, strongly delineated apophyses.

Cephalothorax: Frontal apotome weakly rugose, without frontal warts or cephalic tubercles. Fron-

tal setae 71—98, 80 um long (4); postorbital seta 38—90, 58 yum long (4).

Median antepronotals 75—95, 86 um and 41-68, 54 um long; lateral antepronotal seta 75-90 um

long (3). Anterior precorneal seta 45—83, 68 um long; median precorneal 79—98, 90 um; posterior

precorneal seta 49-68, 59 um long. Distance between anterior and median precorneals 19—28,

26 um; between median and posterior precorneals 23-56, 41 um. Thoracic horn absent. Dorsocen-

tral setae Dc, 41-53, 41 um long; Dec, 23—56, 37 um long; distance between Dc, and Dc, 143—236,

189 um. Metanotal seta (only one?) 15—38 um long (3). Supraalar seta 23—26 um long (3). Wing

sheath nearly smooth.

Abdomen (Figs 4 A—B): Tergite Imore or less bare, but reticulate; T Il with weak median shagreen,

T III-IX with anterior, median and more extensive posterior shagreen composed of slightly coarser

spinules; conspicuous polygonous reticulation laterally. Sternites land IX bare, S II-VIII with ante-

romedian shagreen. O-setae absent. Some tergal connectives with a few anteriorly directed spinules.

T I—-VII with 5 pairs of subequal, thin and simple D-setae. Segment I with 3 L-setae, segment II-VII

with 4 ventral subequal, simple or bifid L-setae, L-setae of segment VIII displaced dorsomedially.

Anal lobe 488-525, 499 um long (6); overreaching genital sac by 30-56 um in male (2); 90-101,

94 um in female, without apical tubercle. Length of anal macrosetae 289—308, 297 um.

Fig. 4. Lappodiamesa boltoni spec. nov., immatures. — A-B. Pupa, tergites VII-IX (A) with detail of shagreen

near apophysis (B). — C-I. Larva; labrum and epipharynx (C) with detail of SI — SIV (D), mandible (E), maxilla

(F), mentum (G), posterior body segments (H) and detail of procercus (I).

Fourth instar larva (n = 3, unless stated otherwise):

Total length 8.96—-9.92 mm. Head capsule length 0.44—-0.52 mm.

Head: Antenna with 5 segments. Length of antennal segments (um): 60-73, 17-22, 9-12, 3—4,

3-4. AR 1.74—1.85. Basal antennal segment 20-21 um wide; distance from base to ring organ

13—20 um, to basal mark of seta 17-34 um, to distal mark 31-53 um; blade 28—29 um long, acces-

sory blade 15 um (1). Apical style of second segment 8-9 um long. Third segment with about 7 an-

nuli. Lengths of setae S 1-S 10 respectively (um): 45 (1), 56 (2), 45, 75-105, 120-131, 86-120,

120-135, 158— 184, 139— 150. Labrum and epipharynx as in Figs: 4C-D; labral lamella consisting of

2 pairs of pectinate lobes, each pair respectively with about 16-25 and 21-30 branches; SI simple and

hairlike, SII (Fig. 4 D) stouter with scalelike base, SIII very weak and simple, STV a short and conelike

with cuplike base, SI Vb small and digitiform without cuplike base; 14 simple chaetae, some very

short, and 8 spinulae present. Pecten epipharyngis consisting of 7 elongate scales, chaetulae laterales

4, chaetulae basales 2. Premandibles (Fig. 4C) 41-49 um (2) long, with 5 teeth; lateral spine not appa-

rent. Mandible (Fig. 4E) 116-128 um long, seta interna with 17—18 branches including 10 plumose

branches; seta subdentalis short. Maxilla as in Fig. 4F. Mentum (Fig. 4G) with broad median tooth

and 8 (9?) lateral teeth, first lateral teeth equally high as median tooth, but distinctly narrower.

Abdomen: Shorter claws of anterior parapods with 2-4 inner teeth. Body setae moderately long.

Procercus 68-79 um long, 45-53 um wide; with 8-9 numbers of 533—555 um long anal setae; late-

ral setae 49-56 um long. Supraanal setae 338-356 um long. Posterior parapods 469—488 um long.

Longest anal tubulus 248-281 um long, 94 um wide at base, 53—60 um wide at middle.

Remarks:

The peculiar furcate anal point in two males of the material studied apparently is adimorphism. The

few males available appear to fall into two groups regarding the numbers of setae on the squama

(52-55 and 22-30 respectively) and the HR ratio (1.71-1.88 and 2.00—2.20 respectively). Themore

normal-looking anal point is somewhat similar to those of Pseudodiamesa Goetghebuer and Pseudo-

kiefferiella parva (Edwards), ı. e with a delicate apical hair sensillum. The hypopygium is similar to

L. vidua apparently differing only in the slightly more massive and less acute shape of the gonostyli,

but the diagnostic characters will otherwise suffice to separate the two species.

The female genitalia of Z/. boltoni are quite “normal-looking”. The pattern of microtrichia on the

genitalia is somewhat similar to some Potthastia species. The microsculpture on the seminal capsules

may be unique, and the presence of sensilla chaetica on the front leg will separate the species from all

other known female Diamesinae.

The pupa differs most conspicuously from L. vıdna by lacking apical tubercles on the anal lobe.

Also, the L-setae are apparently much longer.

According to MAKARCHENKO (1983) the labral sensillum S I is pectinate in the larva of /. vidua. Re-

examination of his material, however, shows that he misidentified S I and that this structure actually

belongs to the labral lamella, thus making both S I, S II and S III simple in this species, just like ın

L. boltoni. The scale at the base of S II both in /. boltonı and L. vidua is similar to that found onS II

in Pseudokiefferiella parva and in the Diamesa latitarsis group (see FErRARESE & Rossaro, 1981: figs 15,

35). Reexamination of Z. vidna also shows that this species has 7 scales in the pecten epipharyngis. The

larva of L. vidua apparently differs from /. boltoni by having the first lateral teeth of the mentum

about equally wide as the median tooth. The median tooth area, however, is highly variable and asym-

metric in all the specimens.

Lappodiamesa was regarded as the sister group of Diamesa Meigen, Sympotthastia Pagast and Pot-

thastia Kieffer combined by Serra-Tosıo (1968). Later the genus was placed provisionally as the sister

group of Diamesa (Serra-Tosıo, 1973). The immature stages as well as the male imagines, however,

will key out near Psendodiamesa in the keys of OLıver (1983, 1986, 1988). A good synapomorphy for

Lappodiamesa plus Pseudodiamesa appears to be the seven elongate scales of the pecten epipharyngis

and we are convinced that Zappodiamesa at least belongs in the same group as Pseudodiamesa together

with Arctodiamesa Makarchenko, Pagastia Oliver, Potthastia, and Sympotthastia. The overall simila-

rities of the larvae with Pseudokiefferiella Zaviel probably are symplesiomorphies.

Acknowledgement

We thank Mr. M. J. Bolton who provided the specimens of L. boltoni, Mr. J. E. Raastad for arranging a loan of

putative types of Syndiamesa vidua, and Dr. E. A. Makarchenko who sent us additional specimens of L. vidua for

comparison. Prof. emeritus H. Kauri spent some time revealing the contents of Russian text for us.

Literature

FERRARESE, U. & B. RossARO 1981: 12. Chironomidi, 1. (Diptera, Chironomidae: Generalitä, Diamesinae, Prodia-

mesinae). — In: Ruffo, S. (ed.): Guide per il riconoscimento delle specie animali delle acque interne italiane.

Consiglio Nazionale delle Ricerche AQ/1/129, Verona, XIX + 97pp

KIEFFER, ]. J. 1922. Chironomides de la Nouvelle-Zemble. — Rep. sc. Results Norw. Exped. Novaya Zemlya 1921

2124

—— 1923: Nouvelle contribution ä l’etude des Chironomides de la Nouvelle-Zemble. — Rep. sc. Results Norw.

Exped. Novaya Zemlya 1921 9: 3—11

MAKARCHENKO, E. A. 1983: K sıstematike ı raspostraneniyu Lappodiamesa brundini Serra-Tosio (Diptera, Chiro-

nomidae). Pp. 52-56 in Ekologija i sistematika presnovoliykh organinemov dal’nego vostoka. — Akad. Nauk

SSSR, Vladivostok

OLIVER, D. R. 1983: The larvae of Diamesinae (Diptera: Chironomidae) of the Holarctic region — Keys and dia-

gnoses. — Ent. scand., Suppl. 19: 115—138

—— 1986: The pupae of Diamesinae (Diptera: Chironomidae) of the Holarctic region — Keys and diagnoses. —

Ent. scand., Suppl. 28: 119—137

—— 1988: The male imagines of Diamesinae (Diptera: Chironomidae) of the Holarctic Region. — Ent. scand.,

Suppl. (in press)

SETHER, OÖ. A. 1969: Some Nearctic Podonominae, Diamesinae, and Orthocladiinae (Diptera: Chironomidae). —

Bull. Fish. Res. Bd Can. 170: 1—154

—— 1980: Glossary of chironomid morphology terminology (Diptera: Chironomidae). — Ent. scand., Suppl. 14:

SERRA-TOSIO, B. 1968: Taxonomie phylogenetique des Diamesini: les genres Potthastia Kieffer, Sympotthastia Pa-

gast, Parapotthastia n. g. et Lappodiamesa n. g. (Diptera, Chironomidae). — Trav. Lab. d’Hydrobiol. Piscic.

Univ. Grenoble 59—60: 117—164

—- 1973: Ecologie et biogeographie des Diamesini d’Europe (Diptera, Chironomidae). — Trav. Lab. d’Hydobiol.

Piscic. Univ. Grenoble 63: 5— 175

WILLASSEN, E. 1982: Descriptions and redescriptions of female Diamesa Meigen (Diptera: Chironomidae); with a

comparative morphology, keys, phylogenetic and biogeographical notes. — Dr. scient. thesis, Dept. Syst.,

Mus. Zool., Univ. Bergen, Norway, V + 316 pp

Prof. O. A. Szther

Dr. E. Willassen

Museum of Zoology

University of Bergen

N-5007 Bergen, Norway

j SPIXIANA | Supplement 14 | 85—89 | München, 15. Juli 1988 | ISSN 0177-7424

Redescription of Paratrissocladius acuminatus (Edwards) comb. n.

(= Cardiocladius acuminatus Edwards) from Southern Chile

(Diptera: Chironomidae)

By Godtfred A. Halvorsen

Abstract

The holotype of Cardiocladins acuminatus Edwards, 1931: 274, is redescribed and transferred to the emended ge-

nus Paratrissocladius Zavrel, 1937. The terminology ofthe chironomid wing venation is discussed and posterior cu-

bitus — PCu, and second anal vein — An, ıs used instead of vannal fold and Ax-vein respectively.

Introduction

While revising the genera in the Cardiocladius group, the holotype of Cardiocladius acuminatus Ed-

wards (1931: 274) from Southern Chile was examined. As mentioned by Brunpin (1956: 66) the pre-

sence of astout anal point with setae suggests that the species does not belong in Cardiocladins Kieffer.

Lacking knowledge of other stages than the male makes the generic placement of the species somewhat

uncertain. However, the species will in recent keys (Pınper, 1978; OLıver et. al. [MS]) key out to Pa-

ratrissocladins Zavrel, as defined by S£THEr (1976: 253).

Morphology

The morphological terminology follows S£THEr (1980) with some additions discussed below. Mea-

surements and ratios follow SCHLEE (1966) with additions and modifications given by S£THEr (1969,

1976).

The presence of a second anal vein (An,) is a plesiomorphous character in the Diptera (Hennig 1954,

1969). In the chironomids only the first anal vein (An,) is well developed, while An, usually is strongly

reduced. However, in several genera it ıs possible to trace remnants of the vein as often is indicated in

the figures. Epwarps (1929: 285, text to fig. 1; 317) used the term Ax-vein, and so did also Brunpın

(1956: 24, fig. 5; 66). An, is especially well developed in Cardiocladins Kieffer and was used by Ed-

wards as a diagnostic character for the genus. In Brunpın’s (1956) figure of the wing of C. capucınus

(Zetterstedt), An, is clearly forked with astem, ashort and weak anterior branch and a stronger poste-

rior branch. This condition is present in all species of Cardiocladins I have examined, as well as in Pa-

ratrissocladius acuminatus (Fig. 1C). However, it is not so clearly developed in the latter species. In

other genera where An; is indicated, including the two other species of Paratrissocladins, ıt always ap-

pears unbranched. Edwards stated that when present, the Ax-vein was lying between two anal folds.

According to Worron (1979: 88) and LinpegerG (1983: 168) the anterior fold (a. f. 1 of Edwards) pro-

bably is the claval furrow, a flexion line which runs posterior to An, in the Diptera. Thus, the anterior

branch of An; clearly present in Cardiocladins may be interpreted as this flexion line, Iying very close

to the true vein and appearing as a part ot it.

Following LinneserG (1983) and Worron (1979) the term vannal fold as used by Hansen & Cook

(1976) and S£THer (1980) is erroneous, and posterior cubitus (PCu) has to be adopted in its place.

Systematics

Eowarps (1931: 274) apparently placed P. acuminatus in Cardiocladins due to the presence of a cor-

diform ta,. This character is at least not agood synapomorphy. Brunpin (1966: 363) made a review of

the occurrence of a cordiform ta, in the Chironomidae, and he regarded the character as a plesiomor-

phy and a probable adaptation to a life in strong currents. Furthermore, the degree of cordiformity va-

ries inside Cardiocladius. C. capucinus (Zetterstedt) has for instance an almost cylindrical ta, at least

on the front leg, slightly shorter than ta; only, while other species in the genus have a more or less dis-

tinctly cordiform ta,. In addition, ta, of P. acuminatus ıs slightly different from that in Cardiocladius

in that ıt is stronger produced beneath the base of tas. These different types of cordiformity are also

found inside the Diamesinae (WırLassen, pers. com.), and may be interpreted as different trends. Ho-

wever, the tendency to get a cordiform ta, can be interpreted as an underlying synapomorphy inside

the Orthocladiinae. S£THER (1977: 86), Sether & Halvorsen (1981: 283) and Szther (1983 a: 284) indi-

cates that the sistergroup of the Cardiocladins group is the Heterotrissocladius group, and that the

Parakiefferiella group is the sistergroup of these two combined. The tendency to get a cordiform ta,,

interpreted as an underlying synapomorphy, would support the hypothesis of these genus groups

being amonophyletic unit, possibly including Psilometriocnemus S&ther, and with one parallel devel-

opment only, the Corynoneura group.

One of the reasons for transferring the species to Paratrissocladins is the shape of the anal point.

S&THER (1983 b: 355) made a phylogenetic analysis of the anal points in the Heterotrissocladius group,

stating that his Type 1 present in all species of Paratrissocladius and Heterotrissocladius, and in some

species of Parametriocnemus Goetghebuer and Paraphaenocladius Thienemann, was unique. This

Type 1 anal point is present in P. acuminatus also, and is regarded as an underlying synapomorphy for

the Heterotrissocladius group.

SETHER (1975: 62) regarded the presence of strong microtrichia only on the wingmembrane asassyn-

apomorphy for Paratrissocladius, and the presence of both microtrichia and setae as the plesiomor-

phous character alternative for the rest of the Heterotrissocladins group. The situation in the Cardio-

cladius group where the microtrichia are very weak, may be seen as the apomorphous alternative in a

three step trend, and exclude P. acuminatus from being a member of that group.

The presence of a virga consisting of a group of minute spines also shows affinity with the Hetero-

trissocladius group. This is Type 5 in SzrHer (1983 b: 356) and is also found in some Paraphaenocla-

dius. SETHER (1976: 253) states that the spines of the penis cavity in Paratrissocladius apparently are

very small. Only Tokunagaia Szther and Tvetenia Kieffer possess virga in the Cardiocladius group,

the first one has Type 4, the latter one a derivation of Type 4 which possibly is a sixth type of virga in

the orthoclads.

The presence of tarsal pseudospurs may indicate a closer relationship with the Cardiocladins group.

These are absent in the Heterotrissocladius group, but present in the Cardiocladius group except in the

Eukiefferiella claripennis and coerulescens groups, in Dratnalia S&ther & Halvorsen, and in Tvetenia

where they are both present and absent.

The strong Parakiefferiella-lıke bend of the gonostylus is unique for P. acuminatus. However, an

indication of a bend is at least present in P. natalensis (Freeman). Until additional knowledge of the

other stages of P. acuminatus is available, the species can best be placed in Paratrissocladins.

Diagnosis

The following corrections and emendations have to be included in the diagnosis given by S£THER (1976: 253):

Eye scarcely to moderately elongated dorsally. Flagellomere 2 and 3, or 2-6 with sensilla chaetica.

Dorsocentrals numerous to normal, with or without humerals. Acrostichals numerous to few. An,

distinct. ta, cylindrical or cordiform. Pseudospurs absent or present. Sensilla chaetica present on mid

or hind legs. Anterior margin of transverse sternapodeme convex or straight. Virga present as a small

cluster of minute spines. Gonostylus with more or less Parakiefferiella-like bend.

Paratrissocladius acuminatus (Edwards) comb. nov.

Eisler E

Cardiocladius acuminatus Edwards 1931: 274, male described.

The male is characterızed by an AR of 1.40; eyes scarcely elongated dorsally; 6 temporal setae; ta,

cordiform; pseudospurs present; sensilla chaetica on mid ta, only; humerals absent; gonostylus with

marked bend.

Male imago (n = 1)

Total length 3.67 mm. Wing length 2.23 mm. Total length/wing length 1.65. Wing length/length of

profemur 2.72.

Antenna: (Fig. 1 A) Flagellum with 13 flagellomeres, last flagellomere 534 um long. AR 1.40.

Head: (Fig. 1B) Temporal setae 6, including 1 inner vertical, 1 outer vertical, and 4 postorbitals.

Clypeus with 8 setae. Tentorıium 183 um long, 28 um wide at sieve pore. Stipes 151 um long, 50 um

wide. Palp missing.

Thorax: (Fig. 1C) Antepronotum with 11 setae. Dorsocentrals 16; acrostichals 5 (some probably

lost in preparation); prealars 4. Scutellum with 7 setae.

Fig. 1. Paratrissocladius acnminatus (Edwards) comb. n.; A — antenna; B — head; C — thorax; D — wing; E —

male hypopygium, dorsal view; F — male hypopygium, ventral view.

Wing: (Fig. 1D) Anal lobe well developed. R,;; ending slightly before the middle of the distance

between R, and R,;;. VR 1.13. Brachiolum with 2 setae; R with 14 setae; R, with 5 setae; and R,;;

with 1 seta. Squama with 16 setae.

Legs: Spur on front tibia 65 um long. Spurs on mid tibia 48 um and 33 um long; on hind tibia

65 um and 38 um long. Width at apex of front tibia 50 um; of mid tıbia 45 um; and of hind tibia

50 um. Comb on hind tibia with 10 setae; longest seta 40 um long; shortest seta 23 um long. Mid leg

with 2 pseudospurs on ta, and ta,, and 1 on ta;; hind leg with 2 on each. Mid leg with 10 sensilla

chaetica apıcally on ta,; hind leg without. Leg lengths (micrometers):

fe at ta] ta, ta, ta, er LR BV SV BR

PJ 819 1051 859 440 252 78 88 0.82 3.18 2.18 243

P, 890 971 516 272 138 55 88 0253 4.30 are 222

pP, 950 1183 748 390 163 60 95 0.63 4.07 2.85 ---

Abdomen: Setae on tergites numerous, more or less uniformly scattered. Sternites 5—8 with median

patches of large setae in addition to lateral lines with weaker setae.

Hypopygium: (Figs. 1 E, F) Ninth tergite with 27 setae, including setae on anal point; laterosternite

with 7 setae. Phallapodeme 110 um long. Transverse sternapodeme 68 um long. Gonocoxite 282 um

long; gonostylus 124 um long. HR 2.28. HV 2.96.

Female, Pupa and Larva unknown.

Material examined and distribution: Holotype, male, slide labelled “Cardiocladius acuminatus Edw., F. W. Ed-

wards det. 1931, Peulla: 12-13. XII. 1926, Southern Chile: Llanquihue Prov., F.& M. Edwards., B.M. 1927-63”.

(In the British Museum.) The species is known from the type locality only. This is the first record, of the genus from

South America.

Acknowledgement

I am much indebted to Dr. P. S. Cranston, British Museum (Nat. Hist.), London, for lending me the holotype,

and to Prof. O. A. Sether, Museum of Zoology, Bergen, for valuable discussions.

Literature

BRUNDIN, L. 1956: Zur Systematik des Orthocladiinae (Dipt., Chironomidae). — Rep. Inst. Freshwat. Res. Drott-

ningholm 37: 5—185

—— 1966. Transantarctic relationships and their significance, as evidenced by chironomid midges. With a mono-

graph of the subfamilies Podonominae and Aphroteniinae and the austral Heptagyiae. — K. Svenska Vetensk.

Akad. Handl. Ser. 4, 11: 1-472

EDWARDS, F. W. 1929: British non-biting midges (Diptera, Chironomidae). — Trans. R. ent. Soc. Lond. 77:

279—430

—— 1931. Diptera of Patagonia and Southern Chile. — II, 5, Chironomidae: 233—331, British Museum, London

HaNSsEN, D. C. & E. F. Cook 1976: The systematics and morphology of the Nearctic species of Diamesa Meigen,

1835 (Diptera: Chironomidae). — Mem. Am. ent. Soc. 30: 1— 203

HENNIG, W. 1954: Flügeläder und System der Dipteren unter Berücksichtigung der aus dem Mesozoikum be-

schriebenen Fossilien. — Beitr. Ent. 4: 246—388.

—— 1969. Die Stammesgeschichte der Insekten. — Frankfurt am Main, 436 pp.

LINDEBERG, B. 1983: Terminology of the wing-veins in Chironomidae (Diptera). — Mem. Am. ent. Soc. 34:

165-168

PiNDER, L. C. V. 1978: A key to the adult males of British Chironomidae. — Freshwat. Biol. Assoc. Scient. Publ.

37: 169pp. + 189 fig.

SETHER, O. A. 1969: Some Nearctic Podonominae, Diamesinae, and Orthocladiinae (Diptera: Chironomidae). —

Bull. Fish. Res. Bd Can. 170: 1-154

—— 1975: Nearctic and Palaearctic Heterotrissocladins (Diptera: Chironomidae). — Bull. Fish. Res. Bd Can. 193:

1-67

—— 1976: Revision of Hydrobaenus, Trissocladius, Zalutschia, Paratrissocladius, and some related genera (Diptera:

Chironomidae). — Bull. Fish. Res. Bd Can. 195: 1—287

—— 1980: Glossary of chironomid morphology terminology (Diptera: Chironomidae). — Ent. scand. Suppl. 14:

1-51

—— 1983. a: Three new species of Lopescladins Oliveira, 1967 (syn. “Cordites” Brundin, n. syn.), with aphylogeny

of the Parakiefferiella group. - Mem. Am. ent. Soc. 34: 279—298

—— 1983b: The canalized evolutionary potential: Inconsistencies in phylogenetic reasoning. — Syst. Zool. 32:

343—359

SCHLEE, D. 1966: Präparation und Ermittlung von Meßwerten und Chironomiden (Diptera). — Gewäss. Abwäss.

41/42: 169-193

WOTTON, R. J. 1979: Function, homology and terminology in insect wings. — Syst. Ent. 4: 81-93

ZAVREL, J. 1937: Eine neue Trissocladius Art. — Spisy. vydav. prir. Fak. Masaryk. Univ. 139: 1-12

Yersabuih Tan 3 chem uaKd ben ch Mer

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[ SPIXIANA ä Supplement 14 | 91-100 | München, 15. Juli 1988 ISSN 0177-7424

Description de trois femelles de Diamesa Meigen dont D. cinerella

Meigen (Lectotype et Paralectotype)

(Diptera, Chironomidae)

Par Endre Willassen et Bernard Serra-Tosio

Abstract

A lectotype has been selected and a full description is given of the types of the genus Diamesa: D. cinerella Mei-

gen, 1835. Examination of the types confirms that our present concept of the genus is correct. The female of D. ka-

symovi Kownacki & Kownacka is described for the firsttime and the female of D. dampfi (Kietfer) is redescribed.

Introduction

Au cours des ans, une considerable confusion de nomenclature a regne dans la litterature consacree

aux chironomides ä propos de la date de creation du genre Diamesa et de son auteur. Les causes de

cette confusion sont multiples. Certaines d’entre elles ont et€ passees en revue par Hansen & Cook

(1976) et par Ashe (1982). Il semble maintenant Evident que la premiere description valıde du genre

Diamesa doit Etre mise au credit de Meigen (in Gistı, 1835) (cf. FREEMAN & CransTon, 1980) et que

Diamesa cinerella est le type de genre par monotypie originale. La description posterieure de D. cine-

rella par Meigen (in Wartı, 1837) n’etait evidemment qu’une simple repetition de la description origi-

nale (comme l’indiquait le sous-titre de l’article: „Aus dem ‚Faunus‘ von GistL“). Meigen consid£rait

les specimens examines comme des mäles, mais on peut &mettre quelque doute sur cette opinion quand

on lit sous sa plume: „Verdient genauere Beobachtung, besonders des Geschlechtsunterschiedes we-

gen“ (in GistL, p. 67).

Plus tard, et pour des raisons inconnues, MEIGEN (1838) rebaptise tout simplement l’espece Diamesa

waltlır. Il est tres probable que la description de D. waltlıı est basee sur les m&mes specimens qui

avaıent Et€ anterieurement appeles D. cinerella (BERGROTH, 1887; Hansen & Cook, 1976).

GOETGHEBURR (1923) examina lestypes de D. waltlii et constata qu’il s’agissait de femelles qu’il asso-

cia de facon erron&e avec les mäles de Tanypus praecox Meigen (= Prodiamesa olivacea [Meigen]). Par

voie de consequence, il declara que le quatrieme article du tarse de l’esp£ce £tait cylindrique. Plus tard

cependant (GOETGHEBUER, 1932), il considera D. waltlii comme une esp£ce valide et fit une courte des-

_ cription du mäle et de la femelle. En m&me temps, il ne donnait aucune reference sur la synonymie an-

terieure de D. waltlii avec T. praecox et dEcrivait le quatrieme article du tarse comme cordiforme.

Pasasr (1947), qui fit la premiere grande revision des Diamesa et genres voisins, supprima le nom

D. waltlii et le remplaca pour des raisons de priorite par D. cinerella. Cependant, comme les types du

genre Diamesa £taient des femelles, il s’abstint apparemment de les examiner, de sorte qu’il n’est pas

sür que le concept actuel du genre Diamesa corresponde ä P’ıdentite des types (Hansen & Cook, 1976).

Sur la base d’une &tude comparee des imagos femelles de Diamesa faite par l’un de nous (WırLassen,

1982), nous estimons que le moment est venu de reexaminer le type du genre et d’en presenter une des-

cription detaillee. En m&me temps, nous en profitons pour decrire les femelles, inconnues jusqu’icı, de

Diamesa kasymovi Kownacki & Kownacka et de D. dampfi (Kieffer), cette derniere espece &tant im-

possible ä identifier d’apres sa description originale.

Diamesa cinerella Meigen

(Fig. 7)

Diamesa cinerella Meigen in GIsTL, 1835: 66; MEIGEN in WALTL, 1837: 283—284 (description de la femelle).

Diamesa waltlii Meigen, 1838: 13 (femelle).

? Diamesa cinerella Meigen: PAGAST, 1947: 482—484 (description du mäle).

Imago mäle

Cf. SErRA-Tosıo, 1971.

Imago femelle (lectotype)

Diagnose

Antenne avec dorsalement une sensille coeloconique ä microtriches sur les flagellomeres 1 et 2.

Flagellomere 7 avec 132 soies pr&apicales et 2 soies apicales. Yeux a pubescence longue. Ta; de chaque

patte cordiforme. Capsules spermathecales de taille moyenne, au nombre de deux. Gonapophyse VIII

avec un pan («flap» de Willassen, 1982) recourb& ventralement. Gonocoxite en forme de plaque con-

vexe et plus ou moins triangulaire. Tergite IX avec deux plaques arrondies. Segment X avec, en vue

ventrale, un tubercule digitiforme.

Description

Longueur/largeur de l’aile: 5,20/2,06 mm. Longueur de l’aile/longueur du femur anterieur: 3,04.

Antenne: scape sans soies. Pedicelle avec 2 soies. Flagellomeres 1 & 6 avec respectivement lenombre

de soies suivant: 6,5, 6,4, 4, 1. Dernier flagellomere avec 1 a 2 soies preapıcales et 2 soies apıcales. Lon-

gueur maximale des soies antennaires: 147 um. Longueur/largeur du pedicelle: 72/120 um. Lon-

gueur/largeur des flagellomeres 137: 112/49, 68/41, 68/41, 52/38, 49/41, 41/34, 196/45 um. AR 0,46.

Une sensille coeloconique a microtriches dorsalement sur chaque flagellomere 1 et 2.

Tete: suture coronale ne depassant pas le nıveau des sensilles frontales. Sensilles frontales & 132 um

l’une de l’autre, et chacune A environ 40 um de l’apex de la protuberance frontale. Protuberances fron-

tales bien saillantes. Soies temporales: 43 au total, dont 1 soie frontale et 11 soies postorbitales. Micro-

triches oculaires denses, depassant tres largement les facettes laterales. Clyp&us avec 15 soies. Lon-

gueur/largeur du clypeus: 135/233 um. Longueur/largeur des articles du palpe: 38/56, 105/64, 169/

72, 177/45, 252/45 um. «Palpal stoutness» (Hansen & Cook 1976) (= rapport longueur du palpe ä

partır de l’article 2 sur somme des largeurs des articles 245): 3,11. Tentorium avec un processus ante-

rolateral bien marque.

Thorax: antepronotum depassant un peu la protuberance scutale, avec 21 soies pronotales laterales.

Soies acrosticales absentes. 17 A 18 soies dorsocentrales. 13 ä 14 soies prealaires disposees sur 1A 3

rangs. Protuberance de l’epımeron avec 4 soies. Preepisternum sans soies. 50 soies scutellaires.

Aile: coloration brunätre. VR 0,96. R, peu arqu£e, faiblement recourbee vers l’avant et prolongee

du cöte distal, presque parallele ala costa A partir de son milieu. RM bien courbee. R,,; arqu&e. Micro-

triches de la membrane alaire visibles des le grossissement 30x. Brachiolum avec 3 soies distales et 1

soie basale. R avec 19 soies, R, avec 20 soies, R,,, sans soies, R,,; avec 20 soies et RM avec Oäl soıe.

Alula avec 12 soies. Squame avec environ 70 soies. Sensilles campaniformes: 4 sur Sc, 2 sur R,, 2 sur

R,;3, 4 sur R,; >.

Pattes: eperons des tibias: 64 um (P,); 68 et 64 um (P;); 109 et 72 um (P;). Largeur de l’extremite

du tibia de P, A P; respectivement: 98, 90, 117 um. Peigne du tibia de P; avec 19 soies (45 A 87 um). 2

soies spiniformes apicales sur seulement chacun des deux premiers articles de chaque tarse. Soies spi-

nıformes preapicales des tarses: 4,0,0 (P,); 18,9,4 (P5); 26,8,4 (P;). Sensilles chetiformes: environ 220

disposees sur 1 A3 rangs depuis le 0,06 jusqu’au 0,61 du metatarse de P;. Pattes ä 4e article du tarse cor-

diforme (cf. Fig. 2.3). Pulvilles petites, visibles au grossissement 300x.

Longueurs (en um) et proportions des articles des pattes:

fe ol ta, ta, ta, ta, ta, LR BV SV

pP] 1710 1971 1140 624 422 163 172 0.58 3.49 SorS)

P, 1971 1781 760 451 307 154 181 0.43 4.12 4.94

P, 2018 2256 1235 784 428 144 182 0.55 3.58 3.46

1.8

2.0

Hypopyge (Fig. 1): sternite VIII avec de chaque cöte 17 & 20 soies. Gonocoxapodeme distinct, for-

tement courbe en vue ventrale. Pan (= «flap») recourbe ventralement, formant de la sorte, en vue ven-

trale, une saillie arquee ä l’avant du lobe ventrolateral. Capsules seminales de taille moyenne, avec une

ponctuation superficielle visible des le grossissement 300X. Conduits seminaux de longueur moy-

enne. Gonocoxite en forme de plaque convexe et plus ou moins triangulaire, A partie moyenne faible-

ment protuberante; 17 soies sur chaque gonocoxite. Tergite IX avec deux plaques arrondies, comple-

tement separees l’une de l’autre; chacune avec ä peu pres 25 soies longues. Partie posterolaterale du

segment X avec, en vue ventrale, un tubercule digitiforme dirige vers l’arriere. Cerques prolonges ven-

tralement, a face laterale un peu plissee.

Fig. 1. Diamesa cinerella Meigen (lectotype), femelle; hypopyge en vue laterale.

Nymphe

Cf. Serra-Tosıo, 1971; FERRARESE & Rossaro, 1981.

Larve

Cf. Rossaro, 1980; FERRARESE & Rossaro, 1981.

Remarque

L’affirmation par GOETGHEBUER (1923) que Ta; a une forme cylindrique est erronee. La morphologie

de la femelle decrite ci-dessus confirme l’appartenance reelle du type de genre au genre Diamesa tel que

nous le concevons actuellement. D’ailleurs le pan recourb& de la gonapophyse et la saillie digitiforme

du segment X montrent que cette espece appartient au groupe cinerella. Cette appartenance &tait ä

l’origine fondee sur des caracteres du mäle et de lanymphe (SerrA-Tosıo, 1971). Le groupe cinerella

est donc facile aussi ä caracteriser gräce aux genitalia femelles (Wırrassen, 1982). Mais malgre le large

eventail de variation des caracteres distinctifs specifiques constate A l’interieur du groupe, on n’est pas

parvenu jusqu’ä present ä des resultats probants pour l’identification des especes en utilisant les imagos

femelles. Il est pourtant interessant de noter que le gonocoxite possede une ebauche de saillie arrondie

du cöte medıan. Il est possible que cette caract£ristique soit sujette ä variation, puisque cette saillie est

moins apparente chez les specimens suppos&s appartenir ä l’espece D. cinerella. Une saillie arrondie

assez developpee se rencontre chez Diamesa lavillei Serra-Tosio et ce caractere a ete utilise par Wırras-

sen (1982) pour separer cette espece des autres femelles du groupe cinerella. Bien que l’espece D. cine-

rella ait et& associee avec le mäle decrit par Pacasr (1947) et par des auteurs ulterieurs, il n’y a pas jus-

qu’a present de preuves Evidentes d’une telle association.

Repartition

Massifs montagneux d’Europe moyenne (arc alpin, For&t Noire, Massif Central, Pyrenees) (SerRA-

Tosıo, 1971).

Materiel examine

Lectotype et paralectotype (par designation actuelle) femelles en pr¶tions au baume du Canada, portant des

etiquettes A texte original suivant: Diamesa waltli, Meigen (et derriere la m&me &tiquette le numero 267/40), Mu-

seum National d’Histoire Naturelle, Paris, France, No 133.

Diamesa kasymovi Kownacki & Kownacka

(Fig. 2.1-6)

Diamesa kasymovi Kownacki & Kownacka, 1973: 32—36 (description du mäle, de la nymphe et de la larve).

Imago mäle

Cf. Kownackı & Kownacka, 1973.

Complements ä la description originale: dans la publication de Kownackı & Kownacka (1973), la tete

du mäle de D. kasymovi est dessinee fig. 8. L’antenne y est representee avec 9 flagellom£res, donc 10

articles apparents si l’on compte le pedicelle, alors que le texte correspondant indique 9 articles, c’est-

a-dire le pedicelle plus 8 flagellomeres (come D. sakartvella decrit par les memes auteurs dans lam&me

publication). Nous avons examine un specimen mäle de D. kasymovi provenant du Caucase et aussi

plusieurs specimens de la Turquie et du Liban. Nous avons pu constater qu’une des antennes de quel-

ques specimens comptait bien 8 flagellomeres, mais l’autre 7 seulement. Nous estimons cependant que

le nombre normal d’articles du flagelle antennaire de D. kasymovi doit &tre de 8. La figure de Kow-

NACKI & Kownacka est donc erron&e, comme nous l’a confırme A. Kownackı ın litt.

Dans la me&me publication, Kownackı & Kownacka representent l’hypopyge mäle de D. kasymovi

(fig. 9) avec de longues soies dorsales sur la partie proximale de l’appendice du coxite. En fait ces soies

(correspondant au «basimedial setal cluster» de Hansen & Cook, 1976) sont ventrales et portees par

un tubercule, ce que nous a confirm& aussi A. Kownackı (in litt.).

Imago femelle

Diagnose: cf. D. cinerella

Fig. 2.1-6. Diamesa kasymovi Kownacki & Kownacka, femelle. 2.1 aile, 2.23 Ta, des pattes en vue laterale et

en vue dorsale, 2.4 hypopyge en vue laterale, 2.5 lobe ventrolateral en vue ventrale, 2.6 hypopyge en vue ventrale.

Description (n=1a5)

Longueur totale: 4,00 mm. Longueur/largeur de l’aile: 3,00/1,17 mm. Longueur totale/longueur de

l’aile: 1,33. Longueur totale/longueur du femur anterieur: 2,27.

Antenne: scape sans soies. Pedicelle avec 3 soies. Flagellomeres 1 & 6 avec respectivement lenombre

de soies suivant: 4,4, 4, 2,4, 1. Dernier flagellomere avec 1 soie preapicale et 2 soies apıcales. Longueur

maximale des soies antennaires: 98 um. Longueuer/largeur du pedicelle: 64/94 um. Longueur/lar-

geur des flagellom£res 1 a 7: 98/42, 49/33, 51/29, 43/25, 43/29, 25/23, 130/29 um. AR 0,42. Une sen-

sille coeloconique A microtriches dorsalement sur chaque flagellomere 1 et 2. Remarque: chez le spe-

cimen decrit, le flagellom£re 6 est incompletement separe du flagellomere 7; chez les autres specimens

examines, les flagellomeres 6 et 7 sont au contraire bien separ&s; le coefficient AR reste dans tous les

cas a peu pres le m&me.

Tete: suture coronale complete. Sensilles frontales ä 90 um l’une de l’autre, et chacune environ

35 um de l’apex de la protuberance frontale. Protub£rances frontales aplaties, peu saillantes. Soies

temporales: 33 & 36 au total (dont 1 &2 soies frontales et 7 & 8 soies postorbitales). Microtriches oculaı-

res denses, depassant tres largement les facettes laterales. Clypeus avec 5 soies. Longueur/largeur du

clypeus: 140/160 um. Longueur/largeur des articles du palpe: 39/60, 95/54, 125/49, 130/39, 230/35

um. «Palpal stoutness» (Hansen & Cook, 1976): 3,47. Tentorium avec un processus anterolateral peu

marque.

Thorax (n = 5): antepronotum depassant un peu la protuberance scutale, avec 10 & 21 soies prono-

tales laterales. Soies acrosticales absentes. 13 ä 17 soies dorsocentrales, disposees sur 1 a 2 rangs en

avant et en arriere. 8& 12 soies prealaires disposees sur 1A 2 rangs.Pas de soies supraalaires. Protube-

rance de l’epimeron avec 8ä 15 soies. Preepisternum sans soies. 34 & 68 soies scutellaires.

Aile (Fig. 2.1): coloration brunätre clair. VR 0,91. R, arquee, faiblement recourbee vers l’avant et

prolongee du cöte distal, rapprochee de la costa et presque parallele ä elle a partir de son milieu. RM

fortement courbee. Microtriches de lamembrane alaire visibles des le grossissement 50x. Brachiolum

avec 3 soies distales, 1 soie mediane et 1 forte soie basale. R avec 20 soies, R, avec 19 soıies, Ry;; avec

22 soies. RM sans soies. Alula avec 7 soies. Squame avec 37 soies. Sensilles campaniformes: 2-3 sur

Sc,2surR,1a3 sur R,,,436sur R;;;.

Pattes (Fig. 2.2—3): eperons des tibias: 52 um (P,); 46 et 56 um (P;); 58 et 90 um (P;). Peigne du

tıbıa de P; avec 16 soies (68 A 83 um). 2 soies spiniformes apıcales sur seulement chacun des deux pre-

miers articles de chaque tarse. Soies spiniformes preapicales des tarses: 0,0,0 (P,); 11,4,0 (P,); 17,6,0

(P;). Sensilles chetiformes: environ 235 reparties depuis le 0.06 jusqu’au 0.63 (soit environ les 2) du

metatarse de P;. Pattes a 4e article du tarse cordiforme (Fig. 2.2-3). Pulvilles petites, visibles au gros-

sissement 350xX.

Longueurs (en um) et proportions des articles des pattes:

fe ti ta, ta, ta, ta, ta, LR BV SV BR

P] 13207 21657 710557 7450 295 110 150 OS 72 22823017.364 1%

P, 1377022 41335 62525320 200 110 140 0.47 4.32 4.33 19

P, 1615 1690 1100 595 295 215 1507 70.65 773.812 73500 1.8

Hypopyge (Fig. 2.4—6) (n = 5): sternite VIII avec de chaque cöte 16 a 28 soies. Gonocoxapodeme

distinct, fortement courbe& en vue ventrale. Pan (= «flap») recourbe ventralement, formant de la sorte,

en vue ventrale, une saillie arquee en avant du lobe ventrolateral. Capsules seminales de taille mo-

yenne, a ponctuation superficielle visible des le grossissement 300x. Conduits seminaux de longueur

moyenne. Gonocoxite en forme de plaque convexe et plus ou moins triangulaire, partie moyenne avec

une petite protuberance arrondie; 16 ä 23 soies sur chaque gonocoxite. Tergite IX avec deux plaques

arrondies, completement separees l’une de l’autre; chaque plaque porte 16 ä 29 soies longues. Partie

posterolaterale du segment X avec, en vue ventrale, un tubercule digitiforme dirige vers l’arrıere. Cer-

ques prolonges ventralement, & face laterale un peu plissee.

Remarque

La femelle de Diamesa kasymovı, dont c’est ıcı la premiere description, est tres semblable aux femel-

les de la plupart des especes du groupe cinerella (sauf D. lavillei Serra-Tosio).

Le faible nombre d’exemplaires de D. kasymovi disponibles presente un nombre significativement

plus eleve (U-Test de Mann-Whitney) de soies sur le sternite VIII que les femelles appartenant proba-

blement ä l’espece D. tonsa (Halıday) provenant de Turquie. Cependant l’analyse statistique des ca-

racteres dans le groupe cinerella est actuellement basee sur des donnees incompl£tes, et on ne sait pas

si le nombre de soies sur le sternite VIII peut Etre utilise comme caractere general pour separer les fe-

melles de D. kasymovi et celles de D. tonsa.

Repartition

Versants asiatique et europeen du Caucase (Kownackı & Kownacka, 1973, 1974), Liban (MousaveEn

& Lavirıe, 1983), Turquie (Reıss, 1985).

Materiel examine

Liban: Mont Liban, 1100 ä 1450 m; leg. Z. Moubayed, 1981 et 1982; 7 mäles, 9 femelles, 2 nymphes femelles;

coll. B. Serra-Tosio, Grenoble, France. Turquie: Prov. Rize, Ovit-Paß, südlich Ikizdere; 2600 m; leg. W. Schacht;

10.7.1986; 4 mäles, Zoologische Staatssammlung, Munich, R. F. A.

Diamesa dampfi (Kieffer)

(Fig. 3.1-6)

Syndiamesa dampfi Kieffer, 1924: 50.

Imago mäle

Cf. Serra-Tosıo, 1970.

Fig. 3.1-6. Diamesa dampfi (Kieffer), femelle. 3.1 aile, 3.23 Ta, des pattes en vue laterale et dorsale, 3.4 hypo-

pyge en vue ventrale, 3.5 lobe ventrolateral et pan (= «flap») en vue ventrale, 3.6 hypopyge en vue laterale.

Imago femelle

Diagnose

Antenne avec dorsalement une sensille coeloconique a microtriches sur les flagellomeres 1 & 3.

Flagellom£re 6 sans soies et flagellomere 7 sans soies preapicales. Yeux A pubescence courte. Ry;; de

l’aile fortement courbee. Ta; de chaque patte un peu aplati et @largı ä l’extremite distale, mais pas

nettement bilobe. Capsules spermathecales grosses. Bord posterieur du pan (= «flap») Echancre au

niveau du lobe ventrolateral. Tergite IX avec deux plaques £troites transversales. Gonocoxite digiti-

forme. Cerques arrondis, un peu allonges du cöte posterieur.

Description (n = 1)

Longueur totale: 5,01 mm. Longueur/largeur de l’aile: 4,04/1,43 mm. Longueur totale/longueur du

femur anterieur: 2,62.

Antenne: scape sans soies. Pedicelle avec 2 soies. Flagellomeres 1 a 6 avec respectivement lenombre

de soies suivant: 6,5, 5, 4, 4, 0. Dernier flagellomere avec 2 soies apıcales. Longueur maximale des soies

antennaires: 169 um. Longueur/largeur du pedicelle: 45/102 um. Longueur/largeur des flagellomeres

127:83/45,53/38, 60/34, 60/26, 41/15, 146/30 um. AR 0,39. Une sensille coeloconique a microtriches

dorsalement sur chaque flagellomere 1-3.

Tete: suture coronale complete. Sensilles frontales a 130 um l’une de l’autre, et chacune environ A

25 um de l’apex de la protuberance frontale. Protuberances frontales tres saillantes. Soies temporales:

environ 40. Microtriches oculaires ne depassant pas les facettes laterales. Clypeus avec 4 soies. Lon-

gueur/largeur du elypeus: 140/150 um. Longueur/largeur des articles du palpe: 45/45, 87/49, 169/49,

154/41, 245/30 um. «Palpal stoutness» (Hansen & Cook, 1976): 3,86. Tentorium avec un processus

anterolateral bien marque.

Thorax: antepronotum depassant un peu la protuberance scutale, avec 9 soies pronotales laterales.

Soies acrosticales absentes. 13 a 15 soies dorsocentrales. 11 soies prealaires disposees sur 1 & 2 rangs.

Protuberance de l’epimeron et preepisternum sans soies. 26 soies scutellaires.

Aile (Fig. 3.1): coloration brunätre clair. VR 0,90. R, peu arquee, faiblement recourbee vers l’avant

et prolongee du cöte distal. RM tres courbee. R,;; relativement fortement arquee. Microtriches de la

membrane alaire visibles des le grossissement 50x. Brachiolum avec 1 soie distale et 1 soie basale. R

avec 24 soıes, R, avec 19 soies, R,,;, avec0ä 1 soıe, R,;; avec 20 soies, et RM avecOä1soie. Alula avec

1 soie. Squame avec environ 30 soies. Sensilles campanıformes: 4 sur Sc, 2 sur R,, 1 sur R,,,,3& 4 sur

Ry;;.

Pattes (Fig. 3.2—3): eperons des tibias: 86 um (P,); 83 et 79 um (P,); 128 et 75 um (P;). Peigne du

tibia de P; avec 16 soies (414 79 um). 2 soies spiniformes apicales sur seulement chacun des deux pre-

miers articles de chaque tarse. Soies spiniformes pr£apicales des tarses: 0,0,0 (P,); 13,6,0 (P,); 22,7,4

(P;). Sensilles chetiformes: environ 160 reparties depuis le 0,13 jusqu’au 0,92 du metatarse de P;. 4e ar-

ticle du tarse un peu aplati dorsoventralement, en vue dorsale faiblement elargi ä l’extremite (Fig. 3.3).

Pulvilles petites, visibles au grossissement 300x.

Longueurs (en um) et proportions des articles des pattes:

ge ti ta, ta, ta, ta, ta, LR BV SV BR

P| IS43T lesbar 2ser586 374 135 165° 0.65: 3.66 2.80 Sal

P, 1663 1734 8l6 432 256 113 162707 Et ell6 2.4

pP, 18527 OST 23562 374 128 ea ef Seile 2.4

Hypopyge (Fig. 3.4—6): sternite VIII avec de chaque cöte 8 & 12 soies. Gonocoxapodeme distinct,

plus ou moins parallele au bord posterieur du sternite. Gonapophyse VIII triangulaire en vue ventrale,

a bords median et posterieur formant un angle aigu; bord median tres allonge ventralement, formant

une saillie parallele a l’axe du corps; bord posterieur Echancre au niveau du lobe ventrolateral

(Fig. 3.4—5); lobe ventrolateral presque entierement entoure par le pan (= «flap»), sauf dans sa partie

mediodistale (Fig. 3.5). Capsules seminales grosses, ovoides; longues d’environ 240 um, apparem-

ment sans ponctuation superficielle. Conduits spermathecaux tres longs. Gonocoxite ä limite basale

indistincte; partie distale droite, digitiforme et dirigee vers l’arriere, portant 9 soies sur son extremite.

Tergite IX avec deux plaques transversales, assez &troites et completement separe&es l’une de l’autre;

chaque plaque porte 17 & 18 soies longues. Protuberance du segment X faiblement developpee. Cer-

ques allonges du cöte posterieur.

Nymphe

Cf. Serra-Tosıo, 1970; FERRARESE & Rossaro, 1981.

Larve

Cf. Rossaro, 1980; FERRARESE & Rossaro, 1981.

Remarque:

La femelle de D. dampfı a Ete insuffisamment d&crite par Kieffer (1924) et en pratique elle est restee

inconnue jusqu’ä maintenant. Par rapport aux autres femelles de Diamesa, on remarque dans legroupe

dampfi (Serra-Tosio, 1970) une synapomorphie nette dans la configuration des spermatheques, c’est-

a-dire des capsules tr&s larges et des conduits longs. On remarque aussi des differences distinctives

entre les trois especes du groupe dampfi. Cette espece se distingue de D. permacra (Walker) et de

D. thomasi Serra-Tosio en particulier par le gonocoxite digitiforme de son hypopyge.

Repartition

Alpes frangaises, Foret-Noire, Tyrol (Serra-Tosio, 1973).

Materiel examine

France: Revel (38), bords du Domenon en amont de l’Oursiere; 1610 m; leg. B. Serra-Tosio; 10.9.1982; 1 mäle,

1 femelle. Huez (38), affl. du lac Besson; 2110 m; leg. B. Serra-Tosio; 8.9. 1983; 2 exuvies nymphales mäles, 3 nym-

phes femelles.

Remerciements

Nous remercions les Drs. A. Kownacki (Cracovie), L. Matile (Paris), F. Reiss (Munich) qui nous ont aimable-

ment prete certains des specimens utilises dans cette Etude.

Travaux cites

AsHE, P. 1983: A catalogue of chiıronomid genera and subgenera of the world including synonyms (Diptera: Chiro-

nomidae). — Ent. scand. Suppl. 20: 1-68

BERGROTH, E. 1887: Entomologische Parenthesen. — Ent. Nachr. 10: 147-153

FERRARESE, U. & B. Rossaro 1981: Chironomidi, 1 (Diptera, Chironomidae: Generalitä, Diamesinae, Prodiame-

sinae). — Guide per il riconoscimento delle specie animalı delle acque interne 12. Consiglio Nazionale delle Ri-

cerche AQ/1/129, Verona, XIX + 97pp

FREEMAN, P. & P. S. CRANSTON 1980: Family Chironomidae, pp. 175—202. In: CROSSKEY, R. W. (Ed.): Catalogue

of the Diptera of the Afrotropical Region. — British Museum (Natural History), London, 1437 pp

GisTL, J. 1835: Neue Arten von Diptern aus der Umgegend von München, benannt und beschrieben von Meigen,

aufgefunden von Dr. J. Waltl. — Faunus 2: 66-72

GOETGHEBUER, M. 1923: Etude critique des Chironomides de la collection Meigen conservee au Museum d’Hi-

stoire Naturelle de Paris. 2e note, genre Tanypus Meigen. — Bull. Soc. ent. Belg. 5: 120-129

GOETGHEBUER, M. 1932: Dipteres Chironomidae IV (Orthocladiinae, Corynoneuriinae, Clunioninae, Diamesi-

nae) — Faune de France 23: 1— 204

GOETGHEBUER, M. 1939: Tendipedidae — Diamesinae. A. Die Imagines. — In: LINDNER, E. (Ed.). Die Fliegen der

Palaearktischen Region 3(13d): 1-28

HANSEN, D. C. & E. F. Cook 1976: The systematics and morphology of the Nearctic species of Diamesa Meigen,

1835 (Diptera: Chironomidae). — Mem. Am. ent. Soc. 30: 1— 203

KIEFFER, J. J. 1924: Chironomides nouveaux ou rares de l’Europe centrale. — Bull. Soc. Hist. nat. Moselle 30:

11-110

Kownackı, A. & M. Kownacka 1973: Chironomidae (Diptera) from the Caucasus. Diamesa Waltl group steın-

boecki. — Bull. Acad. pol. Scı. Cl. II Ser. Sci. biol. 21: 131-138

Kownackt, A. & M. KownackA 1974: Relation of Chironomidae from Tatra and the Caucasus Mts. — Ent.

Tidskr. Suppl. 95: 129— 138

MEIGEN, J. W. 1838: Systematische Beschreibung der bekannten europaeischen zweiflügeligen Insekten. 7: IV +

401 pp, Pls 55-66. Hamm

MOUBAYED, Z. & H. LAVvILLE 1983: Les Chironomides (Diptera) du Liban. I. Premier inventaire faunistique. —

Ann. Limn. 19: 219-228

PAGAST, F. 1947: Systematik und Verbreitung der um die Gattung Diamesa gruppierten Chironomiden. — Arch.

Hydrobiol. 41: 435-596

Reıss, F. 1985: A contribution to the Zoogeography of the turkish Chironomidae (Diptera). — Israel J. Entomol.

19: 161-170

Rossaro, B. 1980: Description of some unknown larvae of Diamesa genus and corrections to previous descriptions

(Diptera, Chironomidae). — Arch. Hydrobiol. 90: 298-308

SERRA-TOSIO, B. 1970: Les Diamesa du groupe dampfi. Description d’une espece nouvelle (Diptera, Chironomi-

dae). — Trav. Lab. Hydrobiol. Grenoble 61: 107— 146

SERRA-TOSIO, B. 1971: Contribution A l’&tude taxonomique, phylogenetique, biogeographique et Ecologique des

Diamesini (Diptera, Chironomidae) d’Europe. — These Univ. Scient. Med. Grenoble, T. I: 1-303, T. II:

304—462

SERRA-TOSIO, B. 1973: Ecologie et biog&ographie des Diamesini d’Europe (Diptera, Chironomidae). — Trav. Lab.

Hydrobiol. Piscic. Univ. Grenoble 63: 5—175

WALTL, J. 1837: Neue Gattungen von Mücken bei München. — Isis 21: 283—287

WILLASSEN, E. 1982: Descriptions and redescriptions of female Diamesa Meigen (Diptera: Chironomidae); with a

comparative morphology, keys, phylogenetic and biogeographical notes. — Dr. scient. thesis, Dept. Syst.,

Mus. Zool., Univ. Bergen, Norway, V + 316pp

Endre Willassen

Museum de Zoologie

Museplass 3

N-5007 Bergen

Norvege

Bernard Serra-Tosio

Universite Joseph Fourier (Grenoble I)

Boite Postale No 53 X

F-38041 Grenoble Cedex

France

100

| SPIXIANA Supplement 14 | 101-104 | München, 15. Juli 1988 | ISSN 0177— 7424

Tanytarsus curvicristatus spec. nov. eine neue Chironomidenart

aus Kolumbien

(Diptera, Chironomidae)

Von Ruth Contreras-Lichtenberg

Abstract

A new species of Tanytarsini is described on the morphological characters of the male imago from Colombia,

South America. The specimens have been collected in the Central Andes, south of the Nevado del Ruız ın 2115 m

above sea-level. Larvae, pupae and females of the new species are unknown, as well as its ecology.

Einleitung

Im Juni 1976 konnten in Kolumbien schwärmende Männchen einer Tanytarsus-Art gesammelt

werden. Die Tiere stammen aus der Zentralkordillere, südlich des Nevado del Ruiz, aus dem Bereich

des Rio Otün und wurden bei einer Forellenzuchtstation „El Cederal“ 25 km östlich von Pereira ın

2115 m Seehöhe gefangen. Der Fundort liegt in der subtropischen Klimaregion Kolumbiens.

Aus Südamerika sind nur wenige Tanytarsini-Arten beschrieben (Kırrrer 1917, 1925, Remper 1939,

Rosack 1960, Reıss 1972, 1985, Reıss & SUBLETTE, 1985).

Tanytarsus curvicristatus spec. nov.

Etymologie: Latein, der Name bezieht sich auf die deutlich median- und dorsalwärts gekrümmten Analkämme.

Material: Holotypus, ©, 9 Paratypen, CC’. (Holotypus und 7 Paratypen Naturhistorisches Museum Wien,

1 Paratypus Zoologische Staatssammlung München, 1 Paratypus Instituto de Ciencias Naturales, Universidad Na-

cional Bogotä, Kolumbien), alles Mikropräparate, Fuparal; El Cederal, Forellenzuchtstation ım Bereich des Rıo

Otun, 2115 m SH, Lichtenberg leg. 17.6.1976.

Imago CO

Färbung: dunkelbraun, Thorax im mazerierten und präparierten Zustand: längs der Mitte heller als

lateral, Scutellum und Postnotum dunkel, Postnotum mit heller Mittellinie, Halteren braun, Beine

braun, Tibien der PII und PIII mit dunklen Kämmen und Spornen.

Größe: größere Art, Flügellänge 2,02 (1,8-2,3) mm; n = 5

Kopf: keine Stirnzapfen vorhanden, 8 (8-10) Vertexborsten (n = 5), 13 (10-15) Clypeusborsten

(n = 4), Augen unbehaart. Länge der Palpomeren in um: Pm 1:36 (32—44); Pm 2: 54 (48-60); Pm

3:119 (116-120); Pm 4: 89 (80-100); Pm 5: 154 (140-160); n = 3.

Antenne: mit 13 Flagellomeren, AR = 0,69 (0,51—0,86);n = 5.

Flügel: (Abb. 1 E) Costa nicht über R,,; hinaus verlängert, Subcosta ın Falte proximal von RM en-

dend. R,,; und R,,; eng nebeneinander parallel verlaufend, nur kurz vor der Mündung von R;,; ın

101

Abb. 1: Tanytarsus curvicristatus spec. nov. A, B, Thorax dorsal und lateral; C, P,-Tibia. D, P,-Tibia; E. Flügel.

Maßstab: A, B, E — 100 um. C, D - 50 um.

die Costa divergierend. M,,;;, knapp vor der Mündung in den Hinterrand des Flügels deutlich ge-

schwungen. An auf der Höhe von FCu endend.

VR = 1,20 (1,16-1,23);n = 5.

Macrotrichien auf R,, R,,; und R,,; sowie auf M, ;, weiters auch an der Flügelspitze zwischen den

Längsadern und parallel dem Flügelhinterrand, proximales Drittel des Flügelhinterrandes ohne Ma-

crotrichien. Squama ohne Haarsaum.

Thorax: (Abb. 1 A, B) Pronotum stark reduziert, von dorsal nicht sichtbar. Humeralborsten feh-

lend, 8-9 Dorsocentralborsten, 4 Acrostichalborsten, jederseits 2 Präalarborsten, 6 Scutellumbor-

sten.

Beine: (Abb. 1C und D) Tibien der P, mit einem Sporn, Vordertarsen ungebartet, Tibien der P, und

P; mit zwei voneinander getrennten schmalen Kämmen, jeder der beiden Kämme mit einem Sporn.

Länge der Beinglieder in um

Fe Ti Ta Ta Ta Ta Ta LR

PJ 880(n=5) 419(n=5) 1004(n=2) 569(n=2) 450(n=2) 332(n=2) 163(n=2) 2,4 (n=2)

(821-949) (386-455) (940-1068) (534-603) (425-475) (307-356) (158-168)

P, 880(n=7) 674(n=7) 407(n=6) 206(n=6) 155(n-=6) 104(n-=6) 79(n=6)

(831-940) (613-712) (366- 455)(188-227) (148-168) ( 99-109) (69-89)

P, 980(n=7) 838(n=7) 527(n=3) 353(n=3) 304(n=3) 191(n=3) 99(n=3)

(910-1058) (762-910) (504-544) 346-356) (297-307) (178-198) (89-109)

Hypopyg: (Abb. 2) Analtergitbänder lang, konvergierend, gegen die Analspitze zu parallel verlau-

fend, am Ende der Analtergitbänder je zwei Setae. Analspitze deutlich ventralwärts gekrümmt, mit

einem oralwärts gerichteten stangenförmigen Fortsatz, jederseits der Analspitze fünf Setae. Ein Paar

102

Abb. 2: Tanytarsus curvicristatus spec. nov. A, Hypopyg, dorsal; B, Hypopyg, lateral; C, Anhang 2, lateral;

D, Anhang 2a; E, Analspitze. Maßstab: A, C, D. E — 50 um; B — 100 um.

deutliche median- und in Lateralansicht dorsalwärts gekrümmte Analkämme vorhanden. Jederseits

der Analspitze ein Zahn. Anhänge 1 medianwärts mit einem Lobus und fünf in einer Reihe stehenden

Setae, an der Basıs zwei weitere, starr medianwärts gerichtete Setae. Digitus schmal, nicht über den

Anhang 1 hinausragend, auf einem breiten basalen, eine Borste tragenden Höcker sitzend. Anhänge 2

keulenförmig, distal schwach zweilappig, mit über die gesamten Anhänge verbreiteten Microtrichien

und distalen, dorsalwärts gekrümmten Setae. Anhänge 2a kurz, gerade, distal mit schlanken, spitzen,

gespaltenen Lamellenborsten.

Imago 9, Larve und Puppe: unbekannt.

Differentialdiagnose

Fehlende Stirnzapfen, knapp vor der Mündung in den Hinterrand des Flügels deutlich geschwun-

gene M;,,. Hypopyg: je 2 Setae am Ende der Analtergitbänder. Die Art unterscheidet sich von allen

anderen Tanytarsus-Arten durch die median- und dorsalwärts gekrümmten Analkämme, die sich bei-

derseits der stark ventralwärts gebogenen Analspitze befinden sowie die Kombination dieser mit fol-

genden Merkmalen: querorientierte in einen medianwärts gerichteten Lobus endende Anhänge 1,

Analspitze mit oralwärts gerichtetem stangenförmigem Fortsatz schmaler, nıcht über den Anhang 1

hinausreichender Digitus, distal keulenförmig erweiterte Anhänge 2 mit schwach angedeuteter Zwei-

teilung, kurze Anhänge 2a mit distalen, schlanken, spitzen, gespaltenen Lamellenborsten.

Die auffällig gekrümmten Analkämme könnten eine Zugehörigkeit zum Genus Paratanytarsus ver-

muten lassen (Reıss & Säwepar, 1981). Die Art wird aber dennoch zu Tanytarsus gestellt, da sie sich

103

von den Arten des Genus Paratanytarsus Thienemann & Bause, 1913 durch die lange Analspitze und

den kurzen, schmalen Digitus unterscheidet. Weitere Arten mit einer oralwärts gerichteten Stange auf

der Analspitze sind die vermutlich holarktische Art Tanytarsus signatus v. d. W. (Reıss & Fırrkau,

1971) sowie die aus Australien beschriebene Art Tanytarsus liepae GLovEr 1973.

Danksagung

Herrn Dr. F. Reıss (Zoologische Staatssammlung München) danke ich für die Hilfe und wertvolle Hinweise bei

der Diskussion der neuen Art.

Literatur

BAUSE, E., 1913: Die Metamorphose der Gattung Tanytarsus und einiger verwandter Tendipediden. — Arch. Hy-

drobiol. Suppl. 2: 1-126

GLOVER, B., 1973: The Tanytarsını (Diptera, Chironomidae) of Australia. — Austr. J. Zool. Suppl. 23: 403-478

KiEFFER, J. J., 1917: Chironomides d’Amerique conserves au Musee National Hongrois de Budapest. — Annls.

hist.-nat. Mus. natn. hung. 15: 292— 364

—— 1925: Chironomides de la Republique Argentine. — Annls. Soc. scient. Brux. 44: 73—92

Reıss, F., 1972: Die Tanytarsını (Chironomidae, Diptera) Südchiles und Westpatagoniens. Mit Hinweisen auf die

Tanytarsını-Fauna der Neotropis. — Stud. neotrop. Fauna 7/1: 49—94

—— 1985: Die panamerikanisch verbreitete Tanytarsini-Gattung Skutzia gen. nov. (Diptera, Chironomidae). —

Spixiana, Suppl. 11: 179—193

Reıss, F. & Firtkau, E. J., 1971: Taxonomie und Ökologie europäisch verbreiteter Tanytarsus-Arten (Chironomi-

dae, Diptera). — Arch. Hydrobiol. Suppl. 40: 75—200

Reıss, F. & SÄwEDAL, L., 1981: Keys to males and pupae of the Palaearctic (excl. Japan) Paratanytarsus Thienemann

& Bause, 1913, n. comb., with descriptions of three new species (Diptera: Chironomidae). — Ent. scand.

Suppl. 15: 73-104

Reıss, F. & SUBLETTE, J. E., 1985: Beardius new genus with notes on additional Pan-American taxa (Diptera, Chi-

ronomidae). — Spixiana, Suppl. 11: 179— 193

REMPEL, J. G., 1939: Neue Chironomiden aus Nordostbrasilien. — Zool. Anz. 127: 209— 216

ROBACK, $. S., 1960: New species of South American Tendipedidae (Diptera). — Res. Cath. Per.-Amaz. Expedi-

tion. — Trans. Amer. ent. Soc. 86: 87-107

Dr. Ruth Contreras-Lichtenberg,

Naturhistorisches Museum Wien,

Burgring 7, A-1014 Wien

104

SPIXIANA | Supplement 14 | 105-116 | München, 15. Juli 1988 | ISSN 0177 — 7424

A reconsideration of the genus Apedilum Townes, 1945

(Diptera: Chironomidae)

By J. H. Epler

Abstract

The taxonomy of the genus Apedilum Townes, 1945 is reviewed. The adult males and females, pupae and larvae

of the two Nearctic species, A. elachistus Townes and A. subcinctum Townes, are redescribed. A key for both spe-

cies in all life stages is provided, and notes on the biology of the genus are given.

Introduction

The genus Apedılum was established by Townes (1945) for 2 species, A. elachistus Townes and

A. subcinctum Townes (the type species). Chironomus nigrohalteris Malloch, 1915 was also included;

this species was considered by Townes (1945: 33) to be a senior synonym of Ch. (Lauterborniella) bra-

chylabis Edwards, 1929. Rosack (1957: 97) noted that Lenz (1941:48) had established the genus Para-

lauterborniella for Ch. (L.) brachylabıs. Since then, all 3 species have been treated as members of Pa-

ralauterborniella. However, as demonstrated in this paper, A. elachistus and A. subcinctum are not

congeneric with P. nıgrohalteris. The genus Apedilum ıs resurrected for the species elachistus and sub-

cinctum, and Paralauterborniella becomes a monotypic genus.

The larva and pupa of A. elachistus (as Paralanterborniella) were briefly described by Beck & Beck

(1970); the larva and pupa of A. subcinctum (as Paralauterborniella) were fiıgured and keyed by Dargy

(1962). Portions of the pupal morphology of A. subcinctum (as Paralauterborniella) were illustrated

and discussed in Pınper & Reıss (1986). In this paper the adult males and females, pupae and larvae of

both species are described/redescribed.

Methodology

Terminology follows EPLER (1987) and SAETHER (1980); methodology follows EPLER (1987). The female notum

was measured from the anterior apex to its posteriormost point. Means are not given for samples of less than 3; all

measurements are in micrometers unless otherwise stated. Abbreviations for collections from which material was

borrowed are: BC = B. A. Caldwell, Georgia Dept. Natural Resources, Atlanta, GA, USA; CN = Canadian Na-

tional Collection, Ottawa, Ontario, Canada (D. R. Oliver); FS = Florida State Collection of Arthropods at Florida

A &M University, Tallahassee, FL, USA; GG = G. Grodhaus, California Dept. of Health, Vector Biology and

Control Section, Berkeley, CA, USA; JE = J. H. Epler; KS = State Biological Survey of Kansas, Lawrence, KS,

USA (L. €. Ferrington, Jr.); ZS = Zoologische Staatssammlung, Munich, West Germany (F. Reiss).

105

Biology

Apedilum larvae most often inhabit submerged vegetation. Darsy (1962: 145— 146) and Macr etal.

(1970) give more detailed information on biology and emergence phenology for A. subcinctum. Mr.

Broughton A. Caldwell (pers. comm., 1987) has been kind enough to provide the following informa-

tion for A. elachistus collected ın Georgia:

On collecting dates in June and August, water temperatures ranged from 25.0° to 39.0° C. The following water

quality parameters were recorded from the collection site in June: flow = 0.10 cfs; D. ©. = 7.2 mg/l; pH = 6.8;

conductivity = 10lumho/cm; BOD, = 3.0 mg/l; fecal coliform = 4900 MPN/100 ml; total solids = 88 mg/l; su-

spended solids = 10 mg/l; total ammonia = 0.06 mg/l (as N); NO, + NO; = <0.02 mg/l (as N), and total P =

0.53 mg/l.

Collection data for A. elachistus ın Florida indicate that adult emergence takes place year round. In

addition to occurring in freshwater, I have collected A. elachistus larvae from submerged vegetation

in a brackish pond in Florida.

Systematics

Apedıilum ıs distinguished from Paralauterborniella by the following characters (Paralanterbor-

niella ın parentheses):

adult male: superior volsella globose (digitiform), gonostyli well developed (strongly reduced);

adult female: labia without microtrichia (with microtrichia), seminal capsule with distinct neck

(without distinct neck);

pupa: cephalic tubercles absent (present), no nose present on wing sheath (present), 5 lateral lamel-

lar setae on T VIII (4);

larva: mentum with deeply bifid median tooth (dome-shaped single median tooth), S II with short

basal segment (long basal segment), pecten epipharyngis a single plate (2 plates), ventromental plates

finely striated (coarsely striated), maxillary plate with well defined striae (weakly defined striae), man-

dible with 2 dorsal teeth and well developed pecten mandibularis (no dorsal teeth and reduced pecten

mandibularis) and supraanal setae approximately equal to length of anal tubules (much longer than

anal tubules).

Apedilum is probably most closely related to Zavreliella Kieffer and Paralauterborniella, and pos-

sıbly Oukuriella Epler, as suggested by Erı£r (1986). The sclerotized neck of the seminal capsule is a

synapomorphy shared with Zavreliella (as Lauterborniella in SAETHER 1977).

At least one other species of Apedilum occurs in the Neotropics. I have examined associated larvae

and pupae from the Rio Marauiä, Amazonas, Brazil which represent a new species. Only pharate

adults within their pupal exuviae were available. The genitalia were similar to those of A. elachistus and

A. subcinctum, but the larvae and pupae were distinctive. The undescribed larva has 52-55 maxillary

plate striae and 2 sclerotized areas on the anteromedial portion of the frontoclypeal apotome. (A. ela-

chistus and A. subcinctum larvae have the anterolateral portions of the frontoclypeal apotome sclero-

tızed.) The pupa has 5 lateral lamellar setae of equal width, only 11-20 T II hooklets, the T VIII cau-

dolateral comb is a cluster of small spines, and the thoracic horn has only 3 branches. Because com-

pletely developed, associated adults are not available, acomplete description is not offered at this time.

Genus Apedilum Townes

Apedilum TowNnss, 1945: 32. Type-species: Apedilum subcinctum Townes, 1945, by original designation (misspell-

ed as succinctum).

Paralauterborniella Lenz, 1941 (partim): ROBACK 1957: 97 (synonymized Apedilum with Paralauterborniella);

PıiNDER & Reıss 1983: 330 (larva); PINDER & Reıss 1986: 341 (pupa).

106

Adult male.

Small chironomids, light brown to greenish-brown in general coloration. Wings with or without

spots, wing length 1.1-1.8 mm.

Eyes bare. Temporal setae in single row beginning mesad to dorsomesal extension of compound

eye, ending behind approximate middle of eye. Frontal tubercles minute (<2 micrometers). Antennal

flagellum with 13 flagellomeres. Maxillary palp 5-segmented, basal segment weakly sclerotized. Cly-

peus subquadrate, setose. Cibarial setae present.

Antepronotum bare, narrowed and weakly notched dorsomesally. Thoracıc scar small; humeral

“pit” present as a single tubercle or 1 large tubercle with smaller tubercles, dorsocaudally to thoracic

scar. Scutal tubercle absent. Acrostichal setae present in double row, anteriorly beginning close to an-

tepronotum and running posteriorly to approximate mid-scutum. Dorsocentral setae in single row/

side, with scattered sensilla campanıformıa along setal row. Scutellar setae usually in 1 row. Supraalar

seta 1/side. Prealar setae about 1-4/side.

Wing membrane without macrotrichia; squama without setal fringe. Brachiolum with 1 seta and 2

groups of campanıform sensilla; R, R, and R,;; usually with setae, R,,; occasionally bare; costa ends

at Ry,;; FCu distal to RM; RM oblique to RR, ; ;.

Metatarsal beard absent on foreleg. Foretibia without apıcal scale or spine. Middle and hind tibiae

each with 2 combs, apparently fused; outer comb bearing one larger spine which projects beyond

others. Sensilla chaetica present on metatarsus of middle leg, confined to apıcal !/5; also sometimes pre-

sent on apical !/s of hind metatarsus. Pulvilli vestigial; empodıum thın, with sparse ventral fringe.

Gonostylus moderately inflated, short, slightly longer than gonocoxite, with straight inner margin;

bearing several long setae along inner margın. Superior volsella well developed; globose-pediform,

with microtrichia dorsally and ventrally; often membranous apically; bearing several to many large

sensilla chaetica on mesal and dorsal surface. Inferior volsella well developed, elongate-digitiform,

slightly arched dorsoventrally, with preapıcal and apical sensilla chaetica. T IX with small group of se-

tae on each side posteriorly; anal point present or absent.

Female.

Generally similar to male; abdomen and wings stouter, and overall generally more setose than male.

Antennae with 5 apparent flagellomeres. Frontal tubercles vestigial or absent.

Mid and hind metatarsı with 1-2 rows of sensilla chaetica on distal /a—27.

Genitalia with well developed dorsomesal and ventrolateral lobes; a narrow, bare apodeme lobe

present. Posteromesal angle of S VIII with a shelf which extends partially over ventrolateral lobe. La-

bia without microtrichia. Seminal capsules spherical with distinct, more heavily sclerotized neck;

spermathecal ducts without major loops or bends.

Pupa.

Small pupae, 3-5 mm long. Exuviae colorless to pale brown with light yellow-brown margins, ce-

phalothorax darker. Cephalic tubercles absent, frontal setae well developed, lamellar. Dorsum of tho-

rax moderately granulose, without scutal tubercle. Thoracic horn with 4 (6?) partly spinose or serrate

branches; base of thoracic horn circular, with 1 apparent tracheal bundle. Thoracic setae/side: precor-

neal 2; dorsocentral 4 in 2 groups; median antepronotal 1; lateral antepronotal 1. Wing sheath without

“nose” or at most a slight protuberance.

Abdominal segment I without lateral setae; segment II with 4 lateral hairlike setae, III with 4 lateral

hairlike setae or 3 haırlike setae and 1 lamellar seta or 2 haırlike setae and 2 thin lamellar setae; IV with

2 hairlike and 2 lamellar lateral setae or 3 haırlike and 1 lamellar lateral setae; V-VII with 4 lateral lamel-

lar setae; VIII with 5, rarely 4, lateral lamellar setae. Anal lobe with a pair of thin dorsal caudolateral

setae and a fringe of 15-30 ventral lateral lamellar setae on each lobe. An uninterrupted median row

of caudal hooklets on T II.

107

Intersegmental conjunctiva of T IIV/IV/V with fine to coarse spinules (usually poorly developed or

absent on T III).

Caudolateral corners of VIII with one large spine or a comb of several smaller spines. Shagreen ab-

sent on T I. Dorsal shagreen on T II-VI subquadrate in outline, slightly wider anteriorly; shagreen

points larger anteriorly. Segments VIland VIII with rounded anterolateral shagreen areas. Anal lobes

without shagreen. Ventral shagreen present only on S VII and VIII as paired, weakly developed,

round anterolateral areas. Pedes spurii A present on $ IV, pedes spurii B present on segments I and II.

Segments II-VIII with one dorsal and one ventral pair O-setae.

Larva.

Small larvae, 3-5 mm long. Body greenish-gold (Darsy 1962: 145) in life. Head capsule pale yel-

low, with 2 pairs of eyespots.

Antenna with 6 segments, segment 4 approximately equal to 2. Antennal blade approximately as

long as flagellum, with small accessory blade. Lauterborn organs well developed, alternately placed at

apices of 2 and 3; a well developed style present near apex of segment 3. Ring organ present at appro-

ximate middle of basal segment.

Frontoclypeal apotome narrowed anteriorly, wıth anterolateral corners produced and more heavily

sclerotized. Labrum with setae I-IVA+B present; S I broad, plumose, with toroidal bases joined me-

dially; S II large, fringed, with short basal segment; S III hairlike, with small basal pedestal; S IVA

small, two-segmented; S IVB smaller than S IVA, inflated, simple. Labral lamella with fringe. Pecten

epipharyngis a distally broader, quadrilateral plate with 8-15 distal teeth. Premandible distally bifid,

the inner blade wider than outer blade, with 1 smaller inner medial tooth; a medial premandibular

brush present.

Mandible with apical tooth, 2 dorsal preapical teeth, and 3 inner teeth. Pecten mandibularis

composed of 8-11 strong setae. Seta subdentalis simple, well developed, 8-15 times longer than wide.

Seta interna with 4 main branches, united basally.

Mentum with deeply bifid pale median tooth and 6 pairs of darker lateral teeth; Ist lateral teeth small

and appressed to 2nd lateral teeth. Ventromental plates about twice as wide as long, with smooth an-

terior margin and continuous fine striae. Setae submenti simple. Maxillary plate with well developed

striations. Triangulum occipitale not visible in ventral view.

Lateral and ventral tubules absent; 2 pairs of conical anal tubules. Procercus about as wide as long,

with 6—8 large apical setae and 2 small, fine anterolateral preapical setae. Supraanal setae subequal to

anal tubules.

Key to all stages of Nearctic Apedilum

Wanduhr aka eene ee Vre ena yore Bohr erfor Fe 2

=: Iulmmature stagesit.\aryse nl uk me IS BR RAN RER DIS 3

2. ‚Wing wich’ pale spots;male never withanal’poine WI. A ENTE A. elachistus

— Wingimmaculate;male withor withoutanalpoint .........2.2220.. A. subcinctum

SealaPupaeıpal: a5 gta ap a Blog dägrstaege Dre Jap zuayla 8 Inn 2 4

rl rBatwe.: VA Nee rer A Fre 5

4. Caudolateral angle of T VIII with 1 large spur or large spur with smaller basal spur (s); T VIII

Iateralserae ofequal’wideh (Fries adden Roms Ne ee A. elachistus

— Caudolateral angle of T VIII with group of smaller spurs; T VIII with lateral seta thinner than

atkter 4 (K1es Dr eh) Tee er En DAR EN E Ru u Eau) Rn A. subcinctum

5:905105,masüllargjplase striag se... dan ae as en ae A. elachistus

= 9110-123maxıllary pltesärraet en. BEI FI N ER A. subcinctum

108

Apedılum elachistus Townes

Apedilum elachistus TOWNES, 1945: 33 (adult description).

Paralauterborniella elachista (Townes): SUBLETTE & SUBLETTE 1965: 173 (placement); BECK & BECK 1970: 30

(pupa, larva description).

nec Paralauterborniella elachista (Townes): DARrBY 1962: 47, 64, 88, 143 (misdetermination of A. subcinctum).

Male Imago (n = 5)

Color. Head and thorax light brown to dark brown; abdomen light brown to brown suffused with

green, or completely light green with light brown areas on dorsum. Legs cream to pale brown, apices

of femora sometimes darker brown. Wings with pale gray spots surrounding base of R,,;; in center

and apex of r,;; (these 2 spots sometimes joined by a narrow bridge); along R, ,; from approximate

midpoint to apex; at FCu and extending along Cu,; and below An (see Townes 1945: fig. 205).

Length. Total 2.12—2.97, 2.69 mm. Thorax 0.69—0.84, 0.80 mm. Abdomen 1.43—2.13, 1.39 mm.

Head. Setae: temporal 20-32, 26; clypeal 12-16, 14; cibarıal 4. Palpomere lengths (4): 28-40, 32;

D0.159951069—90,72799 105310 123 37146 AR 0,85 — 1237206:

Thorax. Setae: acrostichals 10-12, 11; dorsocentrals 16-24, 20; scutellars 6-9, 7; prealars 1—4, 4.

Wing. Length 1.13-1.48, 1.38 mm; with 0.35—0.49, 0.43 mm. VR 0.74—0.78, 0.76. Setae: R+R,

BSR. 0 10,8,

Fig. 1. Male genitalia. A. elachistus: a. hypopygium; b-d. variations of superior volsella. A. subeinctum: e-h. va-

riations of caudal apex T IX, anal point; ı—k. variations of superior volsella.

109

Legs. Palmate sensilla chaetica: 4-11, 7 on middle metatarsus, 0-1 on hind metatarsus. Lengths

and proportions of legs:

d fe ti ta, ta, ta, ta, var LR BV sV

pP 500-650, 415-550, 590-660, 280-330, 220-255, 135-180, 90-100, 1.07-1.27, 2.12-2.40, 1.77-1.95,

1 586 513 618 (3) 297 (3) BER) 152 (3) 93 (3) 1.16 (3) 2.29 (3) 1.85 (3)

P 560-725, 440-570, 290-335, 130-169, 90-115, 50-70, 40-70 0.57-0.66, 3.67-4.16, 3.45-3,.89,

2 637 5ıl 308 147 107 65 60 0.60 3.85 Salz

pP 580-750, 500-640, 380-475, 190-260, 170-220, 90-110, 60-80, 0.73-0.76, 2.63-3.01, 2.80-2.93,

3 672 6 444 (4) 235 (4) 204 (4) 103 (4) 73 0.74 (4) 2.79 (4) 2.85 (4)

Hypopygium (Figs 1a-d). As in generic description. Anal point absent.

Female Imago (n = 4)

Color. Sımilar to male.

Length. Total 1.85—2.17, 2.04 mm (3). Thorax 0.65—0.76, 0.71 mm (3). Abdomen 1.20-1.46,

1.33 mm (3).

Head. Setae: temporal 23—24, 24; clypeal 19-25, 23; cıbarıal 3-5, 4. Palpomere lengths: 22—28,

24, 23—33, 27,53-58, 55; 70-90, 81; 113—128, 120 (3). AR 0.41-—-0.49, 0.45.

Fig. 2. Female genitalia. A. elachistus: a. apex of abdomen, ventral aspect; b. DmL, Apl, VIL. A. subcinctum:

c. apex of abdomen, ventral aspect; d. DmL, ApL, VIL.

110

Thorax. Setae: acrostichals 6-11, 9; dorsocentrals 26-32, 29; scutellars 7—8, 8; prealars 4.

Wing. Length 1.26-1.43, 1.36 mm; width 0.48— 0.54, 0.51 mm. VR 0.70—0.74, 0.72. Setae: R+R,

DD OR 10 105

Legs. Palmate sensilla chaetica: 18-25, 21 on middle metatarsus; 17—22, 20 on hind metatarsus.

Lengths and proportions of legs:

ij \B BV SV

je} fe ti ta, ta, ta, ta, 3; LR

P] 460-520, 415-470, 470-520, 195-220, 140-160, 90-110, 65-75, 1.09-1.17, 2.65-2,74, 1.80-1.94,

494 444 504 2ıl 150 103 73 1.14 2.69 1.86

PR, 520-585, 420-480, 240-270, 105-130, 70-85, 40-55, 45-55, 0.56-0.57, 4,17-4.54, 3.89-3.94,

561 463 261 118 79 48 53 0.57 4.35 3.92

P_ 560-630, 510-580, 350-400, 185-220, 160-190, 70-90 60-70, 0.68-0.69 2.85-2.99, 3.03-3.11,

603 553 378 204 174 8l 68 0.68 2.92 3.06

Abdomen. Notum 162-175, 168; cercı 73-87, 81 long. S VIII with 5-9, 7 setae/side; T X with

12-21, 15 setae; Gc IX with 1-2 setae/sıde. DmL, VIL and Apl as ın Figs. 2a, b.

Pupa: (n=7)

Color. Clear with pale yellow-brown borders.

Fig. 3. Pupa. A. subcinctum: a. frontal apotome; c. abdomen, dorsal; fh. lateral margin, T VIII. A. elachistus:

b. thorax, lateral aspect; d-e. lateral margin, T VIII.

Length. Total 1.98—2.34 mm (3). Cephalothorax 0.66 mm (1). Abdomen 1.68 mm ().

Abdomen (similar to A. subeinctum, Fig. 3c). Shagreen as in generic description. Posterior margin

of T II with transverse row of 26-30, 28 (4) hooklets. T VIII with 5 equally sızed lamellar lateral setae.

Caudolateral spurs on T VII (Figs. 3d, e) single or with small basal spurs. Anal lobes with 16-28, 20

Setae, DR4165 3,20,223:

Fourth instar larva: (n = 9)

Color. Head capsule light yellow.

Head. Postmentum length 113— 125, 118 (8). Mandible (Fig. 4c) length 103— 113. Pecten mandibu-

larıs composed of 8-11, 10 setae. Mentum width 80-90, 84 (5). Ventromental plate with smooth an-

terior margin; width 69-80, 74; length 32-37, 35; VPR 1.86— 2.41, 2.13; IPD 15-18, 17 (3); PSR

4.11—5.33, 4.61 (3); 90-105, 100 (8) maxillary plate striae. Length of antennal segments (7): 60—66,

62; 13-16, 15 (6); 16-28, 17; 12—14, 1356-9, 7;4=6, 5. AR0,95-—-1.15, 1.05 (6). Frontocypese po

tome as in Fig. 4b.

Remarks.

There are 2 apparent forms of adult male A. elachistus. Those with alow AR (0.83—0.85) have high

R+R;, (23-25) and R,,; (17-19) setal counts and a low number (4-5) of palmate sensilla chaetica on

the mid metatarsus. Those with a higher AR (1.19—1.21) have 8-12 R+R, and 0-2 R,,; setae and a

higher number (7— 11) of palmate sensilla chaetica on the mid metatarsus. These differences were not-

ed in California and Florida populations.

I have also examined 2 males from Piedra de los Indios, Colonia, Uruguay which probably are this

species. The genitalia are indistinguishable from A. elachistus and the wing markings, although much

fainter, are sımilar.

Townes (1945) also recorded A. elachistus from Alabama, Mississippi and its type locality ın Galve-

ston, Texas.

Material examined (BLT = black, light trap): U.S.A.: CALIFORNIA: Imperial Co., Wister Wildlife Manage-

ment Area nr Niland, swarming along canal, 26-XII-1981, leg. J. H. Epler, 30 males (JE). FLORIDA: Broward Co.,

Plantation Canal, 9-II-1960, W. Beck, 1 larva (FS); Dade Co., Gould’s Monkey Jungle, BLT, 24-IV-1968, R. E.

Woodruff, 1 male (FS); Homestead, BLT, R. Baranowski, I male (FS); Duval Co., Jacksonville, BLT, 30-IX-1969,

R. King, 12 males (FS); Hamilton Co., roadside ditch nr Occident, 6-IX-1967, W. Beck, 1 male/Pex/Lex (FS);

Leon Co., Tallahassee, 2112 Faulk Dr., at light, 5-VI-1980, Leg. A. R. Soponis, 1 male (FS); Marion Co., Sharpe’s

Ferry Field Sta., malaise trap, 8-13-X-1975, W. R. H., 1 male (FS); Monroe Co., Big Pine Key, light trap, 10-IV-

1970, W. W. Wirth, 7 males, 1 female (FS); Big Pine Key, BLT, 25-V-1978, L. Stange, 1 male (FS); Stock Island,

BLT, 25-I-1967, F. A. Buchanan, 1 male (FS); Palm Beach Co., Palm Beach (residence), 4-X1II-1979, R. P. Toma-

sello, 9 males, 3 females (FS); St. John’s Co., St. John’s River, W. Beck, 1 male/Pex/Lex (FS); Taylor Co., Hickory

Mounds Impoundment, 20-11-1983, leg. J. H. Epler, 51 males (JE); Wakulla Co., St. Marks Natl. Wldlf. Ref., light-

house pool, 19-TV-1980, swarms totaling approx. 600 males & females (JE); same locality & collector, 30-IV-1980,

37 males, 1 female (JE); same locality & collector, BLT, 30-V-1980, 66 males, 3 females (JE); same locality & collec-

tor, 6-11-1983, 80 males (JE); same locality & collector, 8-III-1986, 1 larva (JE); St. Marks Natl. Wldlf. Ref., Stony

Bayou Pool, 23-IX, 27-X-1984, leg. J. H. Epler, 75 males (JE); St. Marks Natl. Wldlf. Ref., Nature Trail, 25-XII-

1984, leg. J. H. Epler, 14 males (JE). Also probably from FL: 5 males/Pex/Lex — no data — (FS). GEORGIA:

Pierce Co., Sixty Foot Branch 1.0 mi downstream of confluence “Patterson Creek”, 18-VI-1986, leg. B. A. Caldwell

& V. Barnes, 2 pupae, 5 larvae (BC); same locality, 20-VIII-1986, leg. B. A. Caldwell & C. Stevens, 1 pharate female

pupa/Lex (BC).

Apedilum subcinctum Townes

Apedıilum subcinctum TOwNEs, 1945: 33 (adult description).

Paralauterborniella subcincta (Townes): SUBLETTE 1960: 202 (adult description, placement); DARBY 1962: 47, 64,

88, 143 (figures of larva, pupa, adult; biology); SUBLETTE & SUBLETTE 1965: 173 (placement); May, et al. 1970:

112

115 (biology ; although often cited as pp. 116-119, a note from Grodhaus on the reprint indicates original pagi-

nation was 115-119).

Paralauterborniella subcincta alamedensıis Sublette, 1960: 203 (description of variant subspecies).

Male Imago (n = 5)

Color. Head and thorax brown to almost black; abdomen light brown to dark brown, sometimes

suffused with green, or completely green with light brown areas on dorsum. Legs pale brown, base of

femora sometimes lighter; femora darker than remainder of leg. Wings immaculate.

Length. Total 2.43—3.20, 2.74 mm (4). Thorax 0.68-0.90, 0.70 mm (4). Abdomen 1.75—2.30,

ie95 mm.

Head. Setae: temporal 18-38, 28 (4); clypeal 13—22, 18 (4); cıbarıal 4-5, 4 (3). Palpomere lengths

(4): 40-49, 44; 33—58, 43; 85-161, 116; 88-150, 116; 140-180, 164 (3). AR 0.66--1.59, 1.15 (4).

Thorax. Setae: acrostichals 9-12, 11; dorsocentrals 15—30, 21; scutellars 6-10, 8; prealars 3—4, 4.

Wing. Length 1.25-1.73, 1.48 mm; wıdth 0.37—0.46, 0.41 mm. VR 0.78-0.81, 0.79 (3). Setae:

R+R, 13-16, 15 3); R,,; 1-22, 9. w

Legs. Palmate sensilla chaetica: 4-7, 6 on middle metatarsus, 0-2 on hind metatarsus. Lengths and

proportions of legs (4):

fe jest ta ta ta ta, ta, LR BV SV

P 600-740, 460-680, 570-650, 280-360, 215-270, 140-185, 85-105, 0.96-1.28, 2.16-2.29, 1.80-2.18,

655 583 603 320 243 165 96 1.06 2.24 2.03

P, 650-800, 515-665, 310-360, 140-205, 105-160, 65-100, 60-80, 0.53-0.60, 3.31-3.99, 3.76-4.09,

718 595 329 178 136 88 74 0.55 3.49 3.99

P, 670-890, 580-790, 410-560, 220-330, 190-290, 95-160, 65-95, 0.71-0.72, 2.55-2.91, 2.96-3.05,

u) ‚696 490 (3) 280 (3) 243 (3) 132 (3) 83 (3) 0.71 2.68 (3) 3.00 (3)

Hypopygium (sımilar to A. elachistus, Fig. 1a). As in generic description; with or without anal

point (Figs. 1e-h)

Female Imago (n = 5)

Color. Similar to male.

Length. Total 1.67—2.19 mm (2). Thorax 0.74—0.87, 0.82 mm (4). Abdomen 0.93— 1.35 mm (2).

Head. Setae: temporal 20-23, 22 (3); clypeal 15-16, 16 (3); cibarial 4 (3). Palpomere lengths:

28-35, 31 (3); 38-42, 39 (3); 105 (2); 110-118 (2); 167 (1). AR 0.47—0.53, 0.49 (4).

Thorax. Setae: acrostichals 9-12, 10 (4); dorsocentrals 24—40, 34; scutellars 7—11, 10; prealars 4.

Wing. Length 1.56-1.73, 1.63 mm; wıdth 0.57—0.61, 0.59 mm. VR 0.76—0.79, 0.78. Setae: R+R;

EDER. 254,31.

Legs. Palmate sensilla chaetica: 19—24, 22 on middle metatarsus; 17—24, 20 on hind metatarsus.

Lengths and proportions ot legs (4):

fe Bat ta] ta, ta, ta, ta, LR BV SV

P, 550-620, 460-580, 495-575, 240-280, 170-215, 110-145, 80-90, 0.95-1.22, 2.15-2.71, 1.80-2.33,

580 531 543 263 189 121 86 1.03 2.52 2.09

P, 660-760, 545-645, 300-330, 150-170, 110-125, 70-85, 60-75, 0.51-0.55, 3.81-3.90, 4.02-4.26,

709 591 313 158 116 75 69 0.53 3.86 4.16

P, 680-820, 620-750, 430-525, 245-355, 200-230, 105-120, 80-100, 0.69-0.70, 2.47-2.88, 2.95-3.02,

751 691 483 289 216 111 89 0.70 2.73 2.99

Abdomen. Notum 163-173, 168 (3); cerci 85-93, 90 (3) long. S VIII with 5 setae/side (3); T X with

15-22 (2) setae; Gc IX with 1 seta/side (2). DmL, VIL and ApL as in Figs. 2c, d.

Pupa: (n = 9)

Color. Clear with pale yellow-brown borders.

115

Fig. 4. Larva. A. elachistus: a. mentum, maxillary plate striae (left), ventromental plate striae (right); b. frontocly-

peal apotome; c. mandible; d. pecten epipharyngis; e.S I; f. S II; g. premandible; h. antenna.

Length. Total 2.56-3.14, 2.88 mm (5). Cephalothorax 0.74—0.93, 0.80 mm (5). Abdomen

1.81-2.79, 2.20 mm (6).

Abdomen (Fig. 3c). Shagreen as in generic description. Posterior margin of T II with transverse

row of 20-35, 28 hooklets. T VIII with 5 lamellar lateral setae, one of which is usually narrower than

the others; occasionally some setae are bifid (Figs. 3f-h). Caudolateral spurs on T VIII (Figs. 3f<h)

a comb of small spines. Anal lobes with 20-28, 24 setae. DR 1.63—3.29, 2.22.

Fourth instar Larva: (n = 6)

Color. Head capsule light yellow.

Head. Postmentum length 116-148, 133 (5). Mandible length 108-133 (3). Pecten mandibularis

composed of 9-11, 10 setae. Mentum with 68-96, 80 (4). Ventromental plate with smooth anterior

margin; width 71-87, 82 (5); length 37—44, 42 (4); VPR 1.97—2.21, 2.06 (4); IPD 16-25, 22 (4); PSR

3.28—4.44, 3.72 (4); 110-125, 118 maxillary plate striae. Length of antennal segments (5): 59-79, 69;

15-17, 16; 16-18, 17; 14—19, 16; 8-10, 9 (4); 5-6, 6 (4). AR 0.91-1.13, 1.00 (4). Frontoclypeal apo-

tome similar to A. elachistus (Fig. 4b).

Remarks.

Townes (1945: 33) stated that the genitalia of A. subcinctum were similar to A. elachistus except for

the presence of an anal point and “the slightly longer superior appendage”. Measurements of the

length and width of the superior volsellae revealed no significant differences between the 2 species in

the length or width of the superior volsellae.

The male’s anal point may be present or absent. SusL£ttE (1960: 202) believed that the presence or

absence of an anal point was due to differences in mounting technique. I have found that this generally

is not the case with this species. Although the anal point, when present, may be dislocated laterally due

to cover slip pressure (Fig. 1g), many specimens I examined possessed no anal point (Figs. le, f). No

differences were noted in the immature stages associated with males with or without anal points.

Darsy (1962) encountered pointless males in his study and considered them to belong to A. elachistus;

114

however, the lack of wing spots, as noted by Darby, and the illustrated pupal T VIII spurs (Darsy

1962: fig. 137) identify his specimens as A. subeinctum.

Although Darsy (1962: fig 136) illustrated a pupal thoracic horn with 6 branches, I have not been

able to discern more than 4 branches on the thoracic horn of either A. subcinctum or A. elachistus.

Macy et al. (1970) have noted that at times of peak emergence, A. subcinctum can be considered a

pest species.

Townes (1945) also recorded this species from its type locality in Reno, Nevada.

Material examined: Canada: ONTARIO: Marmora, 1-VII-1952, J. Vockeroth, -3’males (CN). Mexico: JA-

LISCO: Guadalajara, Zucht aus Springbrunnen, 15-V-1981, leg. H. Fittkau, 1 male/Pex, 2 Pex (ZS). U.S.A.: ARI-

ZONA: Coconino Co., Lake Elaine nr Flagstaff, light trap, 15-X-1986, K. Brenneman & H. Speidel, 2 males (JE).

CALIFORNIA: Imperial Co., Hot Mineral, pool from hot spring overflow, 29-III-1967, leg. R. Soroker &

G. Grodhaus, 1 male/Pex, 1 pharate male pupa/Lex, 2 larvae (GG); Lassen Co., 3 mi. SE Johnstonville, reared

from Typha leaf, leg. G. Grodhaus, 1 male, 1 female, 1 pharate male pupa/Lex, 2 Pex, 1 larva (GG); Merced Co.,

Los Banos Reservoir, 1-V, 28-VII-1968, E. W. Mortenson, 3 males (GG); San Luis Obispo Co., San Luis Obispo,

Laguna Lake, 30-VII-1964, 1 female (GG); same locality, 20-VII-1966, leg. G. Grodhaus, 1 male, 1 pharate male

pupa (GG); same locality & collector, 26-VIII-1965, 1 pharate male pupa, 1 larva (GG); same locality, light trap,

8-IX-1965, leg. J. Montez, 2 males (CN); Solano Co., Vallejo, Lake Dalwigk, 2 males, 3 females, 2 larvae (GG).

KANSAS: Barber Co., North Elm Creek, 12.0 mi. N, 7.0 mi. W of Medicine Lodge, 10-VIII-1981, leg. J. Gelhaus,

1 male (KS); Montgomery Co., Verdigris River, 3.0 mi. E, 2.5 mi. Sof Independence, 23-VIII-1980, leg. J. Gelhaus,

1 male (KS). NEW MEXICO: Albuquerque, drainage ditch nr Rio Grande, 20-IV-1967, 1 male/Pex/Lex gCN).

Acknowledgements

I wish to thank Mr. B. A. Caldwell, Dr. L. C. Ferrington, Jr., Mr. G. Grodhaus, Dr. D.R. Oliver and Dr. F. Reiss

for the loan of specimens. Dr. M. D. Hubbard provided helpful comments. This research was partially supported

by a USDA/CSRS grant to Florida A & M University.

Literature

BECK, W.M. & E. C. Beck. 1970. The immature stages of some Chironomini (Chironomidae). — Q. Jl. Fla Acad.

Sc1. 33: 2942

Dargy, R. E. 1962. Midges associated with California rice fields, with special reference to their ecology (Diptera:

Chironomidae). — Hilgardıa 32: 1—206

EDwARDs, F. W. 1929. British non-biting midges (Diptera, Chironomidae). — Trans. R. ent. Soc. Lond. 77:

279430

EPLER, J. H. 1986. Oukuriella, a new genus of Chironomidae (Diptera) from South America. — Ent. scand. 17:

157— 163

—— 1987. Revision of the Nearctic Dicrotendipes Kieffer, 1913 (Diptera: Chironomidae). — Evol. Monogr. 9:

1-139

LENZ, F. 1941. Die Jugendstadien der Sectio Chironomariae (Tendipedini) connectentes (Subf. Chironominae =

Tendipedinae). Zusammenfassung und Revision. — Arch. Hydrobiol. 38: 1-69

MAcy,H. I., G. GRODHAUS, J. D. GATES & J. MONTEZ. 1970. Pondweed — a substrate for chironomids, especially

Paralauterborniella subcincta. — Calıf. Mosq. Control Ass. 37: 115-119

ROBAcK, S. S. 1957. The immature tendipedids of the Philadelphia area. - Monogr. Acad. Nat. Sci. Philad. 9:

1-152

SAETHER, ©. A. 1977. Female genitalia in Chironomidae and other Nematocera: morphology, phylogenies, keys.

— Bull. Fish. Res. Board Can. 197: 1—209

—— 1980. Glossary of chironomid morphology terminology (Diptera: Chironomidae). — Ent. scand. Suppl. 14:

1-51

SUBLETTE, J. E. 1960. Chironomid midges of California. I. Chironominae, exclusive of Tanytarsini (= Calopsec-

trını). — Proc. U. S. natn. Mus. 112: 197—226

115

SUBLETTE, J. E. & M. S. SUBLETTE. 1965. Family Chironomidae. In: A catalogue of the Diptera of America north

of Mexico. — U. S. Dept. Agric. Handb. No. 276: 142-181

Townes, H. K. 1945. The Nearctic species of Tendipedini [Diptera, Tendipedidae (= Chironomidae)]. — Am.

Midl. Nat. 34: 1— 206

Dr.)]. HliEpler,

Entomology, Florida A & M University,

Tallahassee, FL 32307, U.S.A.

116

| SPIXIANA | Supplement 14 | 1171277 | München, 15. Juli 1988

I" ISSN 0177-7424

Denopelopia atria, a new genus and species of Pentaneurini

(Diptera: Chironomidae: Tanypodinae) from Florida

By Selwyn S. Roback and Robert P. Rutter

Abstract

The larva, pupa and J’ and 9 adults of anew genus of Pentaneurini, Denopelopia n. gen. and Denopelopia atrıa

n. sp. are described from Florida. Comparison is made with the genera Pentaneurella, Monopelopia, Telmatopelo-

pia, Krenopelopia, Xenopelopia, and Zavrelimyıa. To date, this species is known only from the type locality, a 7y-

pha sp. — choked drainage ditch in southwest Florida. The larvae are tolerant of extended periods ot low dissolved

oxygen and relatively high iron concentrations, and consume chironomid larvae, naidid worms, ploimate rotifers

and diatoms.

Introduction

The new genus and species herein described were collected and reared by Mr. Rutter from a small ditch near the

Florida Department of Environmental regulation (FDER) laboratory in Punta Gorda, Florida. The specimens

listed are in the collections of the Academy of Natural Sciences of Philadelphia. Additional material is in the Punta

Gorda FDER laboratory. Most of the specimens are slide mounted, those in ethanol are indicated by (Alc.) under

specimens examined.

Abbreviations and Terminology

The following abbreviations and terminology are used in the descriptions, figures and tables. With some excep-

tions they follow SAETHER (1980). The ® genitalic terminology is after SAETHER (1977). The adult wing venation

and pupal abdominal setae follow FITTKAU (1962), the pupal thoraic setae follow COFFMAN (1983) and the ventral

head setae, KOWALYK (1985). The measurements of the ligula and the ratio of the inner and mesal teeth to the outer

teeth are illustrated in ROBACK (1985, Fig. 86). Land W are used to indicate length and width.

Adult Ta} _;, tarsal segments

Ti, tibia

A.1I-X, abdominal segments

I-II, pro, meso-, metathoraic legs

AR, antennal ratio

Ce, cerci 5

Csa, coxosternapodeme A. IX, ® genitalia EDS

F, femur A.I-VIII, abdominal segments

Gc, gonocoxite, © genitalia

GCa, gonocoxapodeme VII, Q genitalia

Gp, gonapophysis A. VIII. Q genitalıa

Gs, gonostylus, ©’ genitalia

HR, hypopygium ratio Gc/Gs

LR, leg ratio Ta,/Ti

NO, notum, ® genitalıa

R, ramı, 9 genitalia

SCa, seminal capsules, Q genitalia

SDu, spermathecal duct, Q genitalia

Aet, aeropyle tube of respiratory organ

AL, anal lobes

ANi, apical nipple, respiratory organ

D,_;, dorsal abdominal setae

Des 1, 2, 4, dorsocentral thoracic setae

L, , lateral abdominal setae

LS, _;, lateral filaments, abdomen, AL

GS, genital sacs

O , O,, anterior setae, abdomen

V,, ventral abdominal setae

117

Larva

a-e, apical palpal sensillae

aa, antaxial seta of lacınıa

A,_,, antennal segments

ABl, accessory blade

AeT, aeropyle tube

ANi, apical nipple

AR, antennal ratio A,/A;_;

Bl, blade

BR, basal ring of Bl

CS, campaniform sensillum

IC, head ratio Y/L

Lc, lacınıa

LCh, lacinıal chaeta

Li, ligula

pa, paraxial seta of lacinia

Pl, paraligula

PH, pecten hypopharyngis

S|_3, lateroventral mandibular setae

Sy_0, genal setae

SP,, sensory pegs A,

SSm, setae submenti

VP, ventral pore

Denopelopia gen. nov.

Type species, Denopelopia atria spec. nov., by present designation

Diagnosıs

Adult — AR 1.6-1.7; CO antenna with 14 flagellomeres, @ with 11; verticals and orbital setae in

simple row (Fig. 9); wing densely haired with slight pattern in both sexes (Figs. 11, 12); costa not ex-

tended beyond R,,;; (Figs 11-13) ends between apices of M and CU, (Figs. 11, 12); m-cu close to

r—m (Fig. 14); tibial spurs with elongate apıcal tooth (Figs. 1-3) more than half the length of the

Comparison of some characters of Denopelopia (D) with those of Pentaneurella (P), Monopelopia (M),

Krenopelopia (K), Xenopelopia (X), Telmatopelopia (T) and Zavrelimyia (Z). C = concave, E = even,

Table 1.

I = inner, OU = outer, + = present, — = absent.

B M K

Adult

AR 1.0 1.0 1.4

Produced Costa 2 = +

end of Costa % Cu,>M >M

T.IX setae = + ir

Ge, basidorsal lobes - = -

Pupa

RO L/W 3.0 Je2 3.0

distinet ANi = = =

long Aet = = =

no. LS.VII 4 4 3

IGS/AL 0.66 >1.00 0.75

AL spines I-OU OU =

Sticky sheath LS,

AL = = =

Larva

SPa sessile + + +

CS,A, position <0.5 025-026 0.607

apices of Li Teeth C (& c

no. Pl teeth 2 2 2

no. PH teeth 15 1 1

118

1:5-148,1...1:6 RE ı

- + - +

Eur SMI, u=ıM EuToMunelM

- + + +

+ + & -

5.0 5.0 11.0 4.0-5.0

£ E + 3

\ 2 + e

4 3 4

0.80 0.66 0.52 1.00+

OU OU ou I-oU

+ - + +

+ + + -

0.5 0.5 0.6 0.6

E E

2 2 3

10 9 11 15

entire spur in length; claws sharp; © genitalia simple (Fig. 18); tergite A. IX with transverse row of

setae; anal point conical; hypopygium ratio about 1.6; goncoxite without internal lobes; L/W >2.00;

DO genitalia (Figs. 15, 16); anterior end of notum slightly expanded; notum/rami 1.4; coxosternapo-

deme 1X curved, broadened mesally; gonocoxapodeme VIII very poorly developed; gonapophysis

VIII rounded somewhat trıangular; labıa bare.

Pupa — Wholly brown; respiratory organ elongate with apical nipple (Figs. 20—22) and small plas-

tron plate; respiratory atrıum with constrictions, about 11 times as long as greatest width; distinct tho-

racıc comb present (Fig. 32) Dcs 1 very short with apparent apıcal points (Fig. 25); scar A. I distinct,

pigmented in shape of inverted T; shagreen of abdomen, short scattered spinules (4 um); venter, A. II

with arcs of combs (Fig. 28); D setae appear shaped as in Fig. 24; A. VII with three LS filaments, A.

VIII with five (Fig. 29); anal lobes elongate-triangular (Fig. 29) with arow of spinules on outer margin

only (Fig. 30); ©’ genital sacs about half as long as anal lobes (Fig. 29); LS filaments of anal lobes with

sticky sheaths (Fig. 29).

Larva — Body of instar IV red, ıinstar III more orange; head IC ratio about 0.50; lauterborn organs

(SP;) on A, sessile (Figs. 35, 36); ligula with all teeth fairly even in length (Fig. 41); paraligula trifid

(Fig. 41); posterior prolegs with two smaller claws with large inner spurs (Fig. 39) and some outer pec-

tination; medium claws (Fig. 39) with a single small inner spine; all claws (14) pale; anal tubules

(Fig. 45) shorter than posterior prolegs.

Discussion

On the basıs of its overall characters Denopelopia appears most closely related to Telmatopelopia

Fittkau and Zavrelimyia Fittkau. The adult head and thoracic chaetotaxy (Figs. 6, 9) closely resemble

that of T. nemorum (Goetg.) as figured in Fırrkau (1962) Figs. 218, 219, 220. The tibial spurs (Figs.

1-3) are also close to those of T. nemorum as are the O’ genitalia (Fig. 18). The wing macrotrichial

pattern (Fig. 11) ıs similar to that of Z. signatipennis (Kieffer) as figured (Fig. 240) by Fırrkau (1962).

The pupa is very close to that of T. nemorum in the form of the Des setae (Figs. 25—27) the form of

the D setae (Fig. 24) and the form and setae of the AL (Figs. 29, 30). The shape of the unusual RO of

Denopelopia (Fig. 20) can be possibly derived by elongating the RO of T. nemorum (Fırrkau 1962,

Fig. 226) with its suggestion of an ANi. The pupa of Denopelopia differs in having a distinct thoracic

comb (Fig. 32) and only three LS on A. VII (Fig. 29). The larval Li (Fig. 41) is close to that of Zavre-

limyia and Telmatopelopia but the trifid Pl is distinct from either of those genera. The two claws of

the PP with large inner spurs (Fig. 39) resemble those of Zavrelimyia. The latter genus has none to

only one claw of that form. Telmatopelopia possesses several pectinate claws on the PP.

The sessile SP, (Fig. 36) would superficially appear to relate Denopelopia to Pentaneurella Fittkau

& Murray, Monopelopia, Fittkau, Krenopelopia Fittkau, Xenopelopia Fittkau, and Telmatopelopia

Fittkau. However, as noted above, the only one of those genera to which it appears more clearly rela-

ted on the basis of other characters is Telmatopelopia. Zavrelimyıa lacks the sessile SP,. Table 1 com-

pares some of the characters of all those genera with those of Denopelopia and will serve to help differ-

entiate them.

The ® genitalia of Denopelopia (Figs. 15-17) will key to Zavrelimyıa in SAETHER (1977), differing

in the slightly expanded anterior end of NO and the broader R of Gp IX. The other characters in

Saether’s description of the Zavrelimyıa Q genitalia fit Denopelopia closely. The Q genitalia of Telma-

topelopia as described by RopovA (1971) and cited by SAETHER (1977) possess only two seminal capsu-

les.

Geographically Denopelopia poses somewhat of a problem. Telmatopelopia has not been to-date re-

corded from either the Nearctic or Neotropical Regions. The presence of Denopelopia in southern

Florida would suggest a Neotropical derivation. This is possible, inasmuch as the Neotropical fauna

is still very poorly known. T. nemorum has been found only in the western Palaearctic.

111%)

Denopelopia atria spec. nov.

In the following descriptions, unless otherwise indicated, the n for th x in the CO’ adult = 4 and for the pupa and

larva = 5.

Adult ©’ — head brown; eyes bare; outer verticals 107 um long, uniseriate; orbitals uniseriate ex-

tend to center of dorsal eye extension (Fig. 9), 63 um long; pedicel with five setae, 141 um diameter

(Fig. 9), pedicel 1.41% interocular space; AR 1.63—1.71 (X 1.683); flagellomere 14, 80 um long with

Figs 1-10. Adult. 1. spur, tibia I; 2. spurs, tibia II; 3. spurs, tibia III; 4. comb, tibia III; 5. apex of Ta, II;6. scutum,

male, dorsal view; 7. apices of dorsocentrals and acrostichals, female; 8. antepronotum, lateral view; 9. anterior view

of male dorsal eye extensions and head setae; 10. flagellomeres 8 and 9, female antenna.

120

Table 2. The range of and x of the leg segment measurements um and LR for adult Q’ and ? Denopelopia atrıa,

x ın parentheses, n = 3.

d F Te! Ta, Ta, Ta, Ta, Ta, LR

P] 636-722 677-833 592-707 318-370 266-322 181-204 89-107 0.85-0.88

(673.3) (762.2) (661.3) (Se) (299.7) (192.3) (96.3) (0.867)

PJ 699-792 629-751 496-555 237-278 162-222 103-130 73-89 0.74-0.79

(698.0) (701.7) (534.0) (261.7) (192.0) (117.0) (82.7) (0.763)

P, 629-722 803-940 666-755 352-407 248-300 148-181 78-104 0.80-0.83

(674.7) (879.3) ASS) (386.3) (281.3) (169.0) (91.7) (0.813)

[) I abal Ta, Ta, Ta, Ta, Ta, LR

P 592-622 703-777 570-599 307-315 233-252 155-167 93-100 0.77-0.82

\ (608.3) (734.0) (581.0) KR) (244,3) (167.7) (96.3) (0.793)

P, 685-770 681-740 459-496 241-266 163-167 107-118 89-93 0.66-0.67

(725.7) AO) (477.3) (249.3) rESENT) (112,0) (90.3) (0.667)

B 629-692 870-932 629-648 333-370 241-274 148-167 89-100 0.70-0.72

3 (655.0) (909.0) (640.3) (354.0) (260.3) (159.3) (94.0) (0.707)

preapical seta 61 um L; clypeus with 22 long setae; palpus segments ın ratio 26, 78, 141, 170,301 um;

campanıform sensillum ventral on segment 3; antepronotum (Fig. 8) with 2-3 antepronotals and an

upper (15 um L) and lower (32 um long) membranous projections; scutum (Fig. 6) pale brown; vit-

tae, episterna, postnotum, bare, dark brown; scutellum paler brown; 9— 12 setae on disc + 15 anterior

smaller setae; three humerals; dorsocentrals irregularly uniseriate (Fig. 6); acrostichals biseriate, ex-

tend to caudal end of median vittae; a single seta mesal and behind dorsocentrals; six-nine prealars and

one supra-alar; wing (Fig. 11) with overall hair pattern, slightly paler mesally; length 1.6—- 1.7 mm; co-

sta not produced (Fig. 13) ends between M and Cu,; R, not present; m—cu 0.30 arculus to wing tip;

crossvein area of wingas in Fig. 14; legs pale, unmarked; leg measurements and ratios in Table 2; ratio

of longest hairs of Ti, Ta, , to mesal segment width in Table 3; slight beard present on Ta, ; I; hairs on

Ti, Ta, II, III longer than on I; spur Ti I 48-54 um L with three lateral teeth (Fig. 1); spurs Ti II

413

Figs. 11-14. Adult. 11. male wing; 12. female wing; 13. detail of apex of costa, female wing; 14, detail base of ra-

dial sector and r-m, m-cu crossveins.

127

57-58 and 31-35 um long with three lateral teeth (Fig. 2); spurs Ti III 73-77 and 34—35 um long

with three lateral teeth; comb Ti III (Fig. 4) of seven setae; pseudospurs on Ta,, Ta; (Fig. 5) and Ta; ,

Il; none could be discerned on leg III; abdomen with A. I-III pale; basal third of A. IV brown; A.

V-VIII wholly brown; A. IX with 8-10 setae; A. X conical, membranous; genitalia as in Fig. 18,

pale; Gc 118-126 um L (x 124.0); Gc length/width 2.00-2.27 (x 2.133); Gs 74-81 um L (x 77.8);

HR 1.45—1.70 (x 1.598); megaseta of Gs, 11 um L; only strut 3 clearly discernible (Fig. 18).

Figs. 15-18. Adult genitalia. 15. female, ventral view; 16. female, lateral view; 17. detail of seminal capsule and

duct; 18. male, dorsal view.

Adult Q — Head verticals and orbitals as in CO’; pedicel with 6-7 setae; scape with 4—6 setae;

pedicel 81 um diameter, equal to interocular space; antenna with 11 flagellomeres; flagellomere 1

73-76 um L; flagellomeres 2-4, 44—51 um long; flagellomeres 5-8, 55-61 um L; flagellomeres

9—10, 64-70 um long; flagellomere 11 145—146 um L with 115 um preapical seta; flagellomeres 8,

9 as in Fig. 10; basiconic sensillum 73 um long; clypeus and palpus as in O’; antepronotum and scutum

marked as in ©; 8-11 humerals; 8-12 prealars; dorsocentrals extended caudad as are acrostichals

(Fig. 7); 10 scutellars across disc; postnotum as in ©’; wing 1.30—1.33 mm; wing venation as in CO;

hairs in basal third of wing denser and darker (Fig. 12); apex of C and R, ;; (Fig. 13); legs as in CO’, un-

marked; tibial spurs and comb III as in O’; leg measurements and ratios in Table 2; longest hairs on

Ta, ; I not longer than those on Il and III (Table 3); hairs on Ti and Ta, II, III longer than those on

Tı, Ta, I as ın J (Table 3); abdomen wholly brown; genitalia as in Figs. 15, 16; SCa (Fig. 17) pale

Table 3. Ratio of longest hairs/mesal segment width for O’ and 9 Ti, and Ta,_; all legs, Denopelopia atrıa.

o* Tat Ta, Ta, Ta, [) rat Ta, Ta, Ta,

Pl 4,40 4.62 133 6.50 E12 3275 ST 3.43 2.85

P) 6.00 5.29 4.50 3.83 2056400 5.00 3.14 er 1

P2 8.00 7.69 3.883 6.00 P2 6.67 6.00 5.43 3

122

brown, 63x55 um; L/W 2.25; SDu (Fig. 17) thın walled with special secretory cells; NO 108 umL,

slightly widened anteriorly; R 77 um (Fig. 15); NO/R 1.40; Gca VIII poorly developed; Csa IX cur-

ved, slightly broadened mesally (Figs. 15, 16); gonocoxite IX small; gonotergite IX well developed,

bare, appears transversely striate; Gp VIII apically rounded, somewhat triangular (Fig. 15) labia bare;

X with three-four setae on each side (Figs. 15, 16), Ce (Figs. 15, 16) 63x55 um; L/W 1.15; Sca/Ce

1.40; Sca/NO 0.58.

Pupa — Exuviae including wing pads medium to dark brown; frontal apotome as in Fig. 19; respi-

ratory organ (Fig. 20) elongate, narrow with a distinct apıcal nıpple (Figs. 21, 22); overall length of re-

spiratory organ 480-533 um (x 508.0); respiratory atrıum with varıable constrictions; 432—480 um

long (x 460.8); L/W 10.7—11.5 (x 11.14); greatest width of respiratory atrıum 0.37 from its base; apical

nipple, lightly sclerotized, 66-76 um L (x 70.0); L/W 2.22—3.46 (x 2.586); aeropyle tube with bend

toward base (Figs. 21, 22); overall 61-67 um_L (x 64.2); plastron plate (Fig. 21) ovoid, 15-16 um L

(x 15.3, n = 4); L/W 0.53—0.79 (x 0.643, n = 4); base of respiratory organ with small surface spines

(Fig. 23); a distinct thoracic comb of 10-11 short conical tubercles present (Fig. 32); precorneal setae

not clearly discernible; dorsocentral setae in Figs. 25-27, Dcesl, short with 3,4 points, 9 um L (Fig.

25); Dcs2 (Fig. 26) 20 um L; Des4 (= Sas) 27 umL (Fig. 27); scar A. I distinct, pigmented in the form

of an inverted T; abdominal shagreen of scattered fine spinules about 4 um L; venter of A. I with arcs

of spinule combs, 5-12 um_L (Fig. 28); A. I with three D setae; A. I and A. VII (Fig. 34) with four

D setae; A. III-VI (Fig. 33) with five D setae; L, appears present on A. I-VI; L, on A. I-VII; V, ,

on A. III-VII; almost aligned and close together on A. VII (Fig. 34); arrangement of D, L, V setae of

A. V shown in Fig. 33; D setae appear shaped as ın Fig. 24; most D setae distorted in mounting; on

* dr

r nn ANNDADP =

Figs. 19-34. Pupa. 19. frontal apotome; 20. respiratory organ; 21. detail of apex of respiratory organ, dorsal; 22.

detail of apex of respiratory organ, lateral; 23. detail of base of respiratory organ; 24. seta D;, A. IV; 25. seta Desl;

26. seta Dcs2; 27. seta Dcs4 (= Sas); 28. ventral combs of A. II; 29. A. VII, VII and anal lobes; 30 detail of anal

lobe; 31. detail apex of anal lobe; 32. thoracic comb; 33. setal pattern of A. V; 34. setal pattern of A. VII.

123

A.V D, 73 um L; D, (Fig. 24) 5l um L; D; 24 um L; Op and Oy in anterior segment margins

(Fig. 33); on A. VIL (Fig. 34) Op, Oy distinctly caudad of anterior margins; anterior of A. III-V pro-

duced strap-like (Fig. 33) with transverse grooves; A. VII with three LS filaments about 348 um L

(Fig. 29); filaments placed at 0.47, 0.74, 0.94 from base to apex of segment; A. VIII with five LS fila-

ments about 481 um L (Fig. 29); filaments placed at 0.28, 0.54, 0.72, 0.87; 0.98 from base to apex of

segment; on some specimens L, of A. VL ıs enlarged into filament 222 um L; anal fins elongate. trian-

gular (Fig. 29) with row of 9-13 spinules (Fig. 30) on outer margin; spinules 8-9 um L; anal lobes

344—418 umL (x 387.8); L/W 3.21—3.64 (x 3.492); lateral filaments of anal lobes with sticky sheaths

(Fig. 29); LS, about 222 umL placed at 0.39—0.42 (x 0.408) from base to apex of anal lobes; LS, about

259 um L placed at 0.49—0.52 (x 0.506) base to apex of anal lobes; apex of anal lobes knob-like

(Fig. 31); ©’ genital sacs, slightly narrowed apically; 0.49—0.55 (x 0.523, n = 3) length of anal lobes;

Q genital sacs 0.26 (x 0.275, n = 2) length of anal lobes.

Figs. 35—45. Larva. 35. apıcalantennal segments; 36. detail of apex of A}; 37. maxillary palpus and apical sensillae;

38. detail of lacinia of maxilla; 39. smaller claws of posterior prolegs; 40. mentum; 41. ligula, paraligula and pecten

hypopharyngis; 42. ventral head setal pattern; 43. detail of lateroventral mandibular setae; 44. detail of apex of man-

dible; 45. anal tubules, lateral view.

124

Larva — Head pale brown; length 739-749; IC 0.51; depth/length 0.52; labrum similar to Abla-

besmyia (Roback, 1985, Figs. 11, 12); So, Sjo, SSm and VP (Fig. 42) similar to Monopelopia (Kowauyk,

1985); A, 300-318 um (& 307.8); CS, A, 0.55-0.61 (x 0.570) from base; A, 94-100 um (Fig. 35)

(x 96.7,n = 3); L/W A, about 12; A; 4 um; A,4—-5um; AR 2.75—2.91 (x 2.846); Bland ABl (Fig. 35)

subequal in length to A,_4; BV/A,_,0.95—0.98 (x 0.963, n = 4); Bl 100-111 um (x 104.8,n = 4); ABl

102-111 um (x 105.8,n = 4); basal ring (BR) 24 um L; '/W 3.00 (Fig. 35); sensory pegs (Lauterborn

organs) of A, 4-5 um L; sessile on apex of A; (Figs. 35, 36); blade of A, (Fig. 35) about 7 um L; man-

dible 107-122 um; lateroventral setae (S,_;) as in Fig. 43; S, short and peg-like; inner teeth and seta

subdentalis (Fig. 44); apıcal tooth of mandible about 0.30 length of mandible and 2.75 as long as width

at base; A,/mandible 2.57—2.80 (x 2.656); palpus (Fig. 37) 44-47 um L (x 45.0); 1/W 4.43—5.17

(x 4.758); CS of palpus 0.64—0.71 from base (x 0.690); apical segments ot palpus (b) and sensillae (a,

c,d, e, f) shown in Fig. 37; lacınıa as ın Fig. 38 with border of LCh; antaxial seta and paraxial setae on

membranous projections, simple (Fig. 38); A,/palpus 6.45—7.07 (x 6.848); ligula (Fig. 41) 77-78 um

(x 81.8); inner and mesal teeth appear slightly paler; inner teeth and mesal tooth 1.00-1.02 length of

outer tooth (x 1.012); paraligula trıfid (Fig. 41); 44—45 um L (x 44.4); first inner tooth apex at about

0.75 base to apex; second inner tooth apex 0.65 base to apex; pecten hypopharyngis with 11-12 teeth

(Fig. 41); mentum as ın Fig. 40; pseudoradula slightly broadened basally, pustules random; M appen-

dage and ventromentum clearly separated, laterally separation appears partially sclerotized with al-

veoli (Fig. 40); two dorsomental teeth present; body red in fourth ınstar, more orange ın third; ante-

rior prolegs about 469 um L, fork about 0.76 of their length from base; claws 29-58 um L, simple;

those under 29 um with 1-3 fine inner spines; procercus 159-170 um (x 163.8, n = 4); L/W

4.74—5.38 (x 5.000, n = 4); with seven apical elongate setae; two lateral setae below procercus; upper

simple, 38 um; lower multibranched, 92 um; supraanal setae about 350 um L; three lateral setae be-

low supraanal seta, upper bifid 114 um; mesal seta simple 27 um; lower bifid 54 um; subbasal seta of

posterior prolegs simple, about 230 um L; claws of posterior prolegs pale brown; two shortest claws

(Fig. 39) with strong inner teeth and some spinules on outer margins, three slightly larger claws with

one small internal spine (Fig. 39); nıne longer teeth narrow, finely pectinate; anal tubules as ın Fig. 45;

upper about 192 um L; lower 155 um; both slightly more than 4 times as long as greatest wıdth.

Specimens examined

Holotype — ©’ pupal exuviae-adult, shallow ditch near FDER laboratory, Punta Gorda FL, VII-1986, coll. Rutter.

Allotype — ? pupal exuviae — adult, same data

Paratypes - 50°C, 39 pupal exuviae — adult, same data; 3PP pupal exuviae — adult, same data (Alc.)

Non-types — 12 larvae; 400°, 3 Q pupae, same data: 19 larvae; 200°, 3 Q pupae, same data: (Alc.).

Habitat, Ecology and Rearing

To date this midge is known only from the type locality, a shallow drainage ditch surrounding the

Punta Gorda Branch Office of the Florida Department of Environmental Regulation (FDER) in

Charlotte County. Charlotte County lies in the transition zone between the humid subtropical clı-

mate that prevails over much of the southeastern United States and the quasi-tropical climate of

southernmost Florida. Freezing temperatures are rare. Rainfall averages between 127 and 140 cm per

year. Although some rainfall normally occurs every month, there is a distinct rainy season extending

from May through September and a low rainfall season from October through April. About 60 to 65

percent of the annual rainfall occurs during the late spring-summer rainy season (FERNALD & PATTON

1984, p. 179).

The 3 m wide ditch, at least 12 years old, was constructed to aid in stormwater drainage but has wa-

ter yearround due to groundwater seepage. Ditchbank vegetation includes cinnamon fern (Osmunda

125

cinnamomea), royal fern (O. regalıs var. spectabilis), water prımrose (Ludwigia sp.), wax myrtle (My-

rica cerifera), exotic Brazilian pepper (Schinus terebinthifolins), swamp willow (Salıx caroliniana),

southern fox grape (Vitis munsoniana), and south Florida slash pine (Pinus elliottii var. densa). Emer-

gent species are cattail (7’ypha sp.), present throughout the ditch, duck-potato (Sagıttaria lancifolia)

and pickerelweed (Pontederia lanceolata). Overstory canopy is restricted to the shoreline, but the

dense growth T'ypha sp. effectively shades much of the open water areas.

The pupae and adults were reared from fourth instar larvae collected in mid-July and early August

1986. A D-frame aquatic dip net was pulled swiftly backward along the bottom among the cattails pro-

ducing currents which suspended the surficial detritus, then the direction was rapidly reversed and the

mouth of the net bag was pushed forward through the incoming water. In this manner about 24 larvae

were collected with a minimum of debris from an area of approximately 3 m’.

Water depth in the area where D. atria larvae were collected was 613 cm and water movement was

barely perceptible. The bottom was 34 cm of accumulated leaf and needlefall from rıparian and emer-

gent vegetation, overalain by an orange floc. Values for selected physicochemical parameters measu-

red at this location at a depth of 4 cm on 14 July 1986 were: temperature 27.5C, specific conductance

370 umhos/cm, pH 7.1, dissolved oxygen 0.3 mg/l, Fe 1.2 mg/l. On 19 December 1986 values were:

21.5 C, 345 umhos/cm, 7.1, 0.6 mg/l and 2.8 mg/l, respectively. An iron concentration of 108 mg/l

was recorded for a floc sample collected from the bottom on 22 December 1986.

Midges coexisting with D. atrıa in August were Chironomus sp., Goeldichironomus holoprasinus

(Goeldi) and Zavreliella varipennis (Coquillett); in December were Chironomus sp., Einfeldia austini

Beck & Beck and Tanypus carinatus Sublette. Larvae of the phantom cranefly, Bittacomorpha sp.,

were also present in both months. Gut contents of 15 slide-mounted D. atria larvae included Chirono-

mus sp. larvae, naidid worm setae, unidentified arthropod parts, the ploimate rotifer Lecane sp., a va-

riety of pennate diatoms (Achnanthes sp., Cymbella sp., Gomphonema sp., Navicula spp., Nitzchia

spp., Pinnularıa sp.) and undifferentiated detritus.

Fourth instar larvae were held communally in a 5.0X 8.5 cm clear glass container filled with 75 ml

of ditchwater, several detrital willow leaves and floc. In the rearing container larvae clung either to the

underside of the willow leaves or to the small irregular clumps of floc/fine detritus. The larvae readily

pupated and each pupa was placed in to a 17x60 mm 2dram clear glass vial containing 4 ml (depth

25 mm) of ditchwater and either a toothpick or willow leaf extending through the air/water interface.

Pupae typically assumed a “C” configuration against the submerged portion of the leaf or stick, and

often times were observed with the respiratory organs against the water surface. When prodded they

were strong swimmers. However, all pupae died within 36H, some in the process of eclosion. Subse-

quently all but 3 mm of water was drained from the vials and some willow leaf fragments were placed

on the bottom. This worked nicely and several adults emerged successfully. It appears that the pupa

of this species may leave the water partially or entirely at some time prior to ecolosion.

A multitude of habitats and water quality conditions in southwest Florida rivers, streams, canals,

lakes and ponds have been sampled for benthic macroinvertebrates since 1980 as part ofthe FDER wa-

ter quality monitoring program, but not a single specimen of D. atrıia has been collected. This suggests

restrictive ecological requirements for the species, most likely the larval stage. FDER sampling in

ditches is rate, but such habitats are not uncommon in southwest Florida. Ditching to improve

drainage for pastureland, cropland and residential developments has been and still is extensively

employed. Southwest Florida has an extensive surficial aquifer system (FernaLp Parron 1984, p. 36),

and in Charlotte County the water table may be at or above land surface for months in poorly drained

areas. Many wells that tap this aquifer in the central portion of the county produce water with more

than 2 mg/l of iron (Surcuirre 1975). According to Woransky (1978) a significant but unknown

amount of discharge from this unconfined aquifer is into canals and drainage ditches. Further

collecting in ironrich groundwater seeps may reveal additional populations of Denopelopia atria.

126

Acknowledgements

We would like to thank Andy Barth who identified gut content diatoms, Robert A. Christianson who supplied

literature on Charlotte County groundwater, Douglas L. Fry who identified the rotifers, Don E. Sessions who mea-

sured some physicochemical parameters, and Gail M. Sloane who identified some ditchbank vegetation.

Literature

COFFMAN, W.P. 1983. Thoracic chaetotaxy of chironomid pupae (Diptera: Chironomidae). — Mem. Amer. Ent.

Soc. 34: 61-70.

FERNALD, FE. A. & D. J. PATTON (eds.). 1984. Water resources atlas of Florıda. — Florida State Univ., Tallahassee.

291 pp.

FITTKAU, E. J. 1962. Die Tanypodinae (Diptera: Chironomidae). — Abhandl. z. Larvalsyst. der Insekten. 6: 453 pp.

KOowALYK, H. E. 1985. The larval cephalic setae in the Tanypodinae (Diptera: Chironomidae) and their importance

in generic determinations. — The Can. Ent. 117: 67-94.

ROBACK, S. S. 1985. The immature chironomids of the Eastern United States VI. Pentaneurini — Genus Ablabes-

myıa. — Proc. Acad. of Nat. Scı. of Phila. 137 (2): 153-212.

RoDovA, R. A. 1971. Lichinka ı samka Telmatopelopia nemorum (Geotghebuer, 1921) (Diptera; Chironomidae).

— Tr. Inst. Biol. Vnutrennikh Vod, Akad. Nauk SSR 22 (25): 144—151.

SAETHER, O. A. 1977. Female genitalia in Chironomidae and other Nematocera: morphology, phylogenies, keys.

— Bull. Fish. Res. Board Can. 197: V + 209 pp.

—— „1980. Glossary of chironomid morphology terminology (Diptera: Chironomidae). — Ent. Scand. Suppl. 14:

51 pp.

SUTCLIFFE, H., Jr. 1975. Appraisal of the water resources of Charlotte County, Florida. — Florida Bureau of Geo-

logy, Rept. of Invest. 78. Tallahassee. 61 pp.

WOLANSKY, R. M. 1978. Feasibility of water-supply development from the unconfined aquifer in Charlotte

County, Florida. — USGS, Water-Resources Invest. 7826. Tallahassee. 34 pp.

Dr. Selwyn S. Roback,

Academy of Natural Sciences of Philadelphia,

19th and the Parkway, Phila., PA 19103, USA

Mr. Robert P. Rutter,

State of Florida, Dept. of Environmental Regulation

7451 Golf Course Blvd., Punta Gorda, FL 33950-0359, USA

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SPIXIANA Supplement 14 | 129-137 München, 15. Juli 1988 ISSN 0177-7424

Morpho-karyological description of Euryhapsis subviridis (Siebert)

from the South of the Soviet Far East

(Diptera, Chironomidae)

By E. A. Makarchenko, ]J. J. Kiknadze and J. E. Kerkis

Introduction

The species Euryhapsis subviridis (Siebert) belongs to the primitive Orthocladiinae including also

chironomids of the genera Eurycnemus Wulp, Xylotopus Oliver, Brillia K. and Irisobrillia Oliver

(Ouıver, 1981, 1982, 1985; OLıver & Rousser, 1983). Out of the genera given, only species Brıllia and

mainly B. longifurca K.' and B. modesta Mg. (Pankratova, 1970; SHiLovAa, 1976) are widespread over

the USSR. In addition to representatives of this genus (B. flavifrons Joh., B. modesta, B. laculata Olı-

ver & Roussel’, we have found two species of Euryhapsis Oliver: E. cılum Oliver and E. subviridis

(Siebert) in the Soviet Far East, the metamorphosis of the latter being examined from larva to imago

and the larvae fixed for karyoanalysıs.

Chironomids of the genus Euryhapsis which is new to the USSR fauna at all stages of development

are close to Brillia, however they can be easily recognized by the structure of male hypopygium (wide

transverse sternapodeme, distomedial lobe of gonostylus with 2 terminal and few subterminal lamel-

late macrosetae and shorter than subapical lobe, by structures of pupa sternites (sternites VII—VIII

each with a posterior row of spines on low protuberance) and by antenna structure of larva (the 2nd

segment is never divided into 2 parts).

There are detailed descriptions of the preimaginal stages in E. cılium and imago of North America

and Mongolia (OLıver, 1981) whereas E. subviridis was identified by the only male from Austria (Sır-

BERT, 1979). That is why the present work deals with description of larva and pupa of this species as

well as the re-description of a male obtained in the Soviet Far East. As the larvae of Euryhapsıs are

poorly distiguished by their external morphological characters we considered it necessary to carry out

the karyological analysis of E. subviridis and to give the first description of its karyotype demonstra-

ting the photomap of the polytene chromosomes as well as to present information about the chromo-

some polymorphism in the Sakhalın population studied.

Karyotype analysis has contributed much to our current knowledge of the taxonomy and evolution

on Chironomidae (Keyı, 1961, 1962; Marrın et al., 1974; Marrın, 1979; WULKER & BUTLER, 1983; BE-

LYANINA et al., 1983). The karyotypes of more than 130 species of the subfamily Chironominae have

been most extensively studied. Based on karyological data, the taxonomic position of many species

was made more precise, anumber of new ones described (among which of particular interest are the

sibling-species), and the phylogenetic relation of the species to each other were established (Kryı,

! In our opinion B. longifurca Kieffer, 1921 should be considered as a synonym of B. flavifrons Johannsen, 1905

because the male hypopygium of these species are identical.

* Previously this species was known only from North America (OLIVER & RoUussEL, 1983).

129

1962; Marrın, 1979; WÜLKER & BUTLER, 1983; Devaı etal., 1983; Ryser etal., 1983; KıknapzE & Kerkıs,

1984, 1986). Other subfamilies, however, in particular Diamesinae and Orthocladiinae, have been less

studied on karyological grounds. At present the number of Orthocladiinae species studied is just 40

or so, Diamesinae — about 7 (KuserskayaA, 1974, 1979, 1984; MECHELKE, 1953; MıcHA1LovA, 1976, 1980,

1985, Perrova, 1983), and this is clearly insufficient for a general consideration of the karyosystema-

tics of the entire Chironomidae family.

Materials and methods

Material for morphological studies of E. subviridis: 20°C’, Primorye, Khasan Region, Barabashevka River,

17. VII.1975 (E. Makarchenko); 10°, Primorye, Dalnegorsky Region, Inza River, 1.X.1983 (E. Makarchenko);

10), ibid., Partizansk Region, Partizanskaya River, 18.V.1984 (E. Makarchenko); 1 0°, Sakhalin Island, Tym Ri-

ver, 18.1X. 1979 (E. Makarchenko); 20°0°, 1399, 8 pupae, 12 larvae, ibid., Sokol Village, 18. VII. 1986 (E. Ma-

karchenko). Larvae and pupae were fixed by 70% ethanol and imagines by Udemans solution.

Terminology and abbreviation to A. I. SHıLOVA (1976) and SAETHER (1980).

The larvae for karyological analysis were fixed in 3:1 ethanol-acetic acid and stored in arrefrigirator. Analysis was

performed on squashed spreads of salivary gland polytene chromosomes prepared by the standard aceto-orcein

technique. The gonads and the imaginal discs stained with aceto-orcein were used for the meiotic and mitotic meta-

phase chromosome spreads. Determination of larvae age was based on the morphology of the imaginal discs

according to WÜLKER & GÖTZ (1968). The total number of larvae studied was 28.

Euryhapsis subviridis (Siebert) comb. nov.

SIEBERT, 1979: 167— 168 (Brillia)

Male (n = 8).

Total color greenish-white or yellowish white. Head brownish or reddish, eyes black, antenna

brownish, thorax yellowish or greenish. Legs white or greenish, spurs of tibia black. Length,

4.5—5.9 mm; TL/WL 1.4—1.7.

Head. Verticals, 13-25; postorbitals, 6-11; clypeals, 11-28. AR 1.58—2.27. Length of last four

maxillary palp segments (gu): 55— 75.8; 250-345; 160— 244; 157.5—247.5.

Thorax. Antepronotum with 3—6 dorsomedial and 8-16 ventrolateral antepronotals. Dorsocen-

trals, 36-55; prealars, 11—21; supraalars, 1; scutellars, 34—36.

Wing with micro-and macrotrichias. Length, 2.7—3.8 mm. Squamal setae, 9-51.

Legs. LR, 0.86-0.88; LR; 0.55—0.59; LR; 0.57—0.75; SV, 2.0-2.1;SV, 3.5-3.74; SV, 2.49—2.98;

BV, 2.17-2.38; BV, 2.8—3.48; BV, 2.61—2.72. Hind tibial comb consisting of 8-9 spines.

Hypopygium (fig. 1 A—H). Tergite IX with 12-28 setae; sternite IX laterally with 4-9 setae. Go-

nocoxite parallel-sided, without stout setae on dorsomedial margin. Superior volsella with 18-30 se-

tae (about 30 u of length). Distomedial lobe of gonostylus about three-quartes as long as subapical

lobe and with 2 terminal and 6-8 subterminal lamellate macrosetae. Transverse sternapodeme square

to rectangular with a truncated apex.

Pupa (n = 6)

Colour grey or dark grey, anal lobe yellowish or greyish; length, 6.3—-8 mm. Exuviae grey.

Thoracic horn brownish-yellow or yellow; large, pointed and covered with spinules only in the up-

per three quarters (fig. 2A). Length of thoracic horn 481.4—572.7 u; ratio of length of thoracic horn

to width of thoracic horn 5.2—6.9.

Tergites II-V each with rough shagreen on all surface (fig. 3D); tergite VI medial with tender sha-

green only. Tergites II-VI each with posterior row or rows of spines (fig. 3C, E). Tergites VII-VIII

130

Fig. 1. Male of Euryhapsıs subviridis: A-C — hypopygium (A — from Sakhalın Island, B-C — from Primorye),

D-F — gonostylus, G-H -— transverse sternapodeme (G — from Primorye, H — from Sakhalın Island).

| H

Fig. 2. Pupa(A-B) and larva (C-H) of Euryhapsis subviridis: A — thoracic horn; B — segments VII-IX ofmale,

ventral; C — SI-SIV setae of labrum; D — premandible; E — frontal apotome, clypeus, and labral sclerites; E —

antenna, G — mandible, H — mentum.

without shagreen and posterior rows of spines. Sternites II-III each with very tender anterior sha-

green only. Male sternites VI-VIII each with a posterior row of spines, on low protuberance on VII

(fig. 3F) and two low protuberance on VIII (fig. 2 B); female sternites VI- VII similar to male, and two

large, flap-like protuberance on VII (fig. 3G). Number of spines of posterior row of sternites

VI-VII: 14-17; 13—14; 10-16. Conjuctives IVIII-IV/V with spinules (fig. 3A-B). Segments

I-VIII each with 4 L-setae; length of lateral setae of segments I-V 105.6—-132 u, of segments

VI-VII 161.7—-264 . Anal lobe with complete lateral fringe of 26-34 setae and 3 anal macrosetae

(fig. 2B).

Fig. 3. Structures of tergites and sternites of pupa Euryhapsis subviridis: A — posterior row of spines tergite Il and

conjuctives IV/III, X275; B — ibid, X525; C — posterior rows of spines tergite III, x525; D — shagreen of tergite

III, x525; E — posterior row of spines tergite VI, X525; F — posterior row of spines sternite VII, X275; G — poste-

rior row of spines sternite VIII of female, X 275.

Larva, fourth instar (n = 6)

Colour white or greenish; length, 8.2—8.9 mm. Head capsule yellow, width of head 0.4—0.5 mm;

eye spot, apical half of mandible and mentum black.

Labrum granular on lateral; labral sclerites as on fig. 2E. SI setae of labrum distal plumose,

S II-SIII simple, S IV present, but very small (fig. 2C).

Antenna yellowish-brown, 4-segmented; ring organ near base of first segment; antennal blade ab-

out 1.5—1.6 times as long as combined length of flagellar segments (fig. 2F); AR 1.63—1.95. Preman-

dible distal with 3 teeth (fig. 2D).

132

Mandible with 5 teeth; basal tooth blunt and shorter as other inner teeth. Seta subdentalıs long, api-

cally slightly curved, ending at level of apex of second inner tooth; seta interna with 5 plumose bran-

ches (fig. 2G).

Mentum with one median tooth and 6 pairs of lateral teeth present; sızth lateral tooth lower as other

lateral teeth (fig. 2H).

Procercus about 1.5—1.6 times as long as wide, posterior part strongly sclerotized; bearing 8 apıcal

and 2 subapical anal setae, the lowest of them about 2—3 times as long as the upper one.

Oecology

Larvae inhabit the low course of foothill rivers on gravel-pebble grounds with sandy filling at the

rate of 0.5—0.7 m/s.

Karyological analysis of larva of fourth instar

Observation of the diplotene nuclei and mitotic metaphases (of imaginal disc mitoses) in E. subviri-

dis show three pairs of very small chromosomes (fig. 4a). Because of the somatic pairing characteristic

of Diptera, the homologues lie close to each other, and the number of chromosomes discernible at the

mitotic metaphase often appears to be three. Judging by the morphology of the metaphase chromoso-

mes, two pairs are metacentrics, and the third one is submetacentric. As expected from the mitotic ar-

rangement, in the salivary gland nuclei there are three chromosomes, each of them exhibits two paired

homologues (fig. 4b). The chromosomes have been numbered I to III in order of decreasing length

and tentatively given left and rıght ends. The major division has been numbered consecutively throug-

hout each chromosome, and each major one subdivided into minor ones denoted by letters (fig. 5).

E. subviridis polytene chromosomes have distinctive banding patterns making them advantageous for

karyological analysis. However, we encountered difficulties when attemping to identify the centro-

meres because there were no associated large blocks of heterochromatin. We succeeded in defining the

Fig. 4. The entire chromosome complement of Euryhapsis subviridis. a — mitotice chromosomes; b — salivary

gland polytene chromosomes. IL-IR; IIL-IIR; IIIL—-IIIR — chromosome arms. The arrows indicate the putative

candıdates for the centromeric bands. N — Nucleolar organızer, BR — Balbıanı ring.

133

centromeric regions only after thoroughly examining of many preparations. The bands we thought to

represent the centromeric regions in all the larvae studied were distinguished from all other bands of

the polytene chromosome by their homogenous staining.

Chromosome I (175+ 5.2 u) is the longest of the complement. It is ametacentric chromosome con-

taining section 1— 21. The centromeric region is identified in section 9. A characteristic feature isthe

presence of a nucleolus at the end of arm IL in section 1. Reviable markers of arm IL are the constric-

tions in sections 3 and 8 as well as dark bands in sections 4, 6 and 7. Such markers in arm IR are the

constriction in section 10, heavy bands in sections 16, 17 and 18. A prominent puff develops in some

larvae in section 15. Heterozygous inversions in arm IL were detected only ın one larvae of all those

examined (fig. 6a).

Chromosome II (168 +8 u) ıs nearly as long as chromosome I. It is a metacentric like chromosome

I (fig. 5). It consists of 21 sections. The centromeric region shows two dark bands in section !%Yıı. Arm

IIL is distinguished by the presence of constriction in sections 2 and 4, clear-cut banding in section 5

and pale bands in section 8; arm IIR has a constrictions in section 11, bands distinct ın sections 12, 13

and 15). There is a small puff in section 20 in all the larvae studied, and also ın sections 10 and 18

in some (fig. 4-5). No nucleoli and Balbianı rings were identified in chromosome II, neither were

rearrangements.

Chromosome III (152# 4.5 u) is the shortest of the complement, and it is divided into 17 sections.

It is asubmetacentric (fig. 4-5). The centromere was deduced by the presence of a dark band in section

10. There ıs a Balbianı ring near section 9 of arm IIIR and a nucleolus in section 15. The main markers

@ HR

TB '

IL IR

Fig. 5. A standard map of the salivary gland chromosomes of Euryhapsis subviridis. The designations are the same

as in fig. 4.

134

of arm IIIL are groups of bands in sections 4, 5, 6, 7 and 8; and those of arm IIIR are groups of pale

bands in section 11 and dark ones in sections 12 and 17 (fig. 5). Heterozygous inversions were obser-

ved in both arms in 33 % of the larvae (fig. 6).

The distinctive morphological features of the salivary gland of E. subviridis deserve some com-

ments. The morphology of the salivary glands of some other Orthocladiinae species have been consi-

dered elsewhere (MEcHeık£, 1953). The salivary glands of Orthocladiinae have been described as flat

sacs composed of three lobes. The glands are paired organs asymmetrically arranged in the larval body.

The cell nuclei in the lobes differ in polyteny level, puffing and nucleolus patterns (MEcHELKkE, 1953).

The paired salivary glands of E. subviridis are sacshaped too. One gland is elongated and the other is

ovoid. In E. subviridis, the salivary gland cells are radıally arranged cells with low ploidy level lyıng

in a semicercle closer to the centre, and the level of chromosome polyteny increases with the distance

from the center (fig. 6d-e). An attempt was made to compare the polytene chromosomes from the

central and peripheral parts of the salıvary gland according to the activities of the nucleolı and Balbiani

rings. This seemed reasonable because some Orthocladiinae species were found to differ markedly in

the activities of these regions in some of the gland cells. However, no such differences were observed,

so far, for E. subviridıs.

The peripheral gland cells have the highest level of polyteny, and they are, consequently, advanta-

geous for karyological analysis.

It has been reported that ecological and ontogenetic factors affect the structure of the polytene chro-

mosomes (ILımskaya, 1984; Demin & IrımskayA, 1986). There are also indications in the literature that

compactization of the salivary gland chromosomes in summer may be manifested as loss of the fine

structural details. However, we have not observed that this might be the case. Although the E. subvi-

ridis larvae were fixed in June, the polytene chromosome retained their distinct banding patterns. The

discrepance between the observations reported in the literature and ours may be due to the fact that

they were made on the represent actives of different Chironomidae subfamilies.

IIL

Fig. 6. Arm IL (a), arm IIIR (b), arm IIIL (c) heterozygotes. The morphology of the salivary glands of Euryhapsıs

subviridis: d— magnified part of the gland near the duct; e — general appearance. Cells with nuclei at different levels

of polyteny are distinctly seen.

135

Notes

E. subviridis males are very close in the structure of hypopygium to the North American species

E. ılloba Ol., however they differ from it in the value of AR (AR = 1.06 ın E. ılloba). Individuals of

the Far East differ also from the male of the type locality in higher values of ARand LR (AR = 1.25,

LR = 0.80 in E. subviridis from Austria).

Within the genus Euryhapsis the preimaginal stages were previously known only for E. cilium. Pupa

of E. subviridis differs from E. cilinm in the form of thoracic horn which is bifurcated and totally co-

vered with fine spinules whereas in E. subviridis ıt is simple and covered with spinules only in the up-

per three quarters. Larvae of E. cilium and E. subviridis are very similar and they can be distinguished

only by certain characters. SI of labrum in E. subviridis with 7—9 lobes (more than 10 in E. cilium),

the seta interna with 5 plumose branches (7 in E. cılium), the 6th lateral tooth of mentum is rounded,

larger than the 5th and much lower as the other lateral teeth (the 5-6th lateral teeth of mentum of

E. cilium are of similar shape and size and arranged close to each other).

Comparisions ofthe major karyotypic features of E. subviridis with those of the other Orthocladii-

nae studied demonstrate similarities with respect to diploid chromosome number (2 n = 6), location

of the centromeric region (2 metacentrics, 1 submetacentric) (MicHaıLovA, 1976, 1985).

There are some karyotypic features, specific to E. subviridis to be noted. The level of chromosome

polyteny is higher in E. subviridis as compared with that established for the other members of the Or-

thocladiinae subfamily (MicHarova, 1976, 1985). This facilitates chromosome mapping in E. subviri-

dis. Another specific feature ofthe E. subviridis karyotype is the unusual telocentric localization ofthe

nucleolar organizer in chromosome I. Futhermore, its polymorphism level is 0.4 heterozygous inver-

sions per individual, higher than reported for the other representatives of the Orthocladiinae subfa-

mily (MıcnaiLova, 1985).

It would be of interest to study karyotypes of other species of the genus Euryhapsis to provide a

broader bases for comparision of the structural organization of chromosomes and also to make judge-

ments about their interrelated evolutionary history.

Literature

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mosomen. — Chromosoma 12: 26-47

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KUBERSKAYA, E. F. 1979: Karyotypical characteristic of chironomids subfamily Diamesinae (Diptera, Chironomi-

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E. A. Makarchenko,

Institute of Biology and Pedology, Far East Science Center, USSR

Academy of Sciences, Vladivostok, USSR,

1. I. Kiknadze, I. E. Kerkis,

Novosibirsk University,

Novosibirsk, USSR

137

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SPIXITANA Supplement 14 | 139-142 | München, 15. Juli 1988 ISSN 0177-7424

The occurrence of setal tufts on larvae of

Orthocladius (Orthocladius) annectens Saether

(Diptera, Chironomidae)

By J. P. Fagnani and A. R. Soponis

Abstract

Setal tufts are described and illustrated for larvae of Orthocladius (O.) annectens Saether. The usefulness of this

character for distinguishing genera must be re-examined.

Introduction

This is the first report of setal tufts on larvae ofthe genus Orthocladins. The occurrence of setal tufts

has been a traditional character for distinguishing larvae of some species of Cricotopus from larvae of

all species of Orthocladius.

A setal tuft is a group of setae arısing from a single base and occurring laterally in 1 to 4 pairs on ab-

dominal segments in larvae of some Chironomidae. Although the fourth lateral seta is most often de-

-veloped as a setal tuft, lateral setae 1 to 3 can also occur as setal tufts. SAETHER (1980) did not define setal

tufts in his glossary.

Orthocladius (Orthocladius) annectens Szther

Larval exuviae of O. (O.) annectens reared from Otsego Lake, New York, possess distinct setal tufts

(Fig. 1). One pair occurs on each of segments IV to IX, situated posterolaterally in place of the L4 se-

tae. The tufts can be long and robust (Fig. 1a, b) to short and weak (Fig. 1c). Each tuft is composed of

approximately 9 to 16 setae which can sometimes appear to be a single thick seta (Fig. 1a, Ic).

Setal tufts have been noted previously in three genera of Orthocladiinae: Cricotopus, Stackel-

bergina, and Symposiocladins (THIENEMANN 1944 ; CHERNOvsKI 1949; HırvEnoJA 1973; SHıLova and ZeEı.-

ENTSOV 1978; CrAnsTon 1982a; CransTon et al. 1983; Orıver and Rousser 1983). Similar plumose setae

have been described in Parorthocladius and Synorthocladius. Setal tufts have been referred to as bristle

tufts (JOHANNSEN 1937); hair pencils (JoHAnnsen 1937; Rogack 1957; Darsy 1962; Mason 1973; BECK

1975, 1976; Sınrson and Bone 1980); setal tufts (THIENEMANN 1944; CHERNOVSKII 1949; CRANSTON

1982a; Cranston et al. 1983); setal brushes (THIENEMANN 1944; Cranston 1982b; CraAnsTon et al.

1983); many-branched brush setae (CHernovsku 1949); hair tufts (Mason 1973); plumose setae (OLı-

ver et al. 1978; Cransron 1982a; OLıver and Rousser 1983); and group of setae (Corrman and FERRING-

Ton 1984). THIENEMANN (1944, and earlier) used both Büschelborsten (setal tufts) and Borstenpinsel

(setal brushes) for these setae in Crzcotopus.

To our knowledge there is no published record of the occurrence of setal tufts in larvae of Orthocla-

dius. SAETHER (1969) originally described O. annectens, but he did not include body setae in the larval

139

description. Soronis (1977), in her redescription, did not examine body setae on the larvae, but the pre-

sence of setal tufts on those specimens was verified by D. R. Oliver (pers. comm.). Soronis (1977) cor-

rectly synonymized Hydrobaenus sp. 2 0f Rogack (1957) with O. annectens, but Rosack reported that

the unassociated larvae lacked hair pencils on the body. Sımpson and Bope (1980) provided photo-

graphs of the larva of O. annectens (as O. [O.] prob. annectens), and stated that the larval abdomen

lacked haır pencils. Presumably, the setal tufts were obscured or destroyed on the specimens exami-

ned, or these authors overlooked them. Setal tufts vary in size on larvae of O. annectens, and they are

difficult to see on larval exuviae and larvae that have been mounted in self-clearing media for some

time. In alcohol-preserved larvae, prominant setal tufts are generally visible at 25 to 50x under a dis-

secting microscope, but they may be lost or obscured after slide-mounting. Rosack (1957) pointed out

that, „These haır pencils are often broken off in handling and mounting and must be searched for with

great care“.

Figs 1-2. Orthocladius annectens Szther. 1 a-c: Setal tufts in slide-mounted larvae (scale = 0.2 mm). 2: Larval

mentum.

Orthocladius annectens ıs Nearctic and is found throughout most of North America (Soronis 1977).

This widely distributed species lives in small streams, large rivers, and lakes (Soronis 1977; Sımpson and

Bope 1980). The larvae of O. annectens can be distinguished from all morphologically similar species

of Orthocladius and Cricotopus by the unusual 13-toothed mentum with the 5 light median teeth ex-

tended anteriorly (Fig. 2). However, because of setal tufts, the larva of O. annectens will key to Crico-

topus in most recent keys (e. g., OLıver etal. 1978; Simpson and Bone 1980; Cranston 1982a; CRANSTON

etal. 1983; OLıver and Rousseı. 1983; Corrman and FERRINGToN 1984).

The functions of setal tufts are not clear. Menzıe (1978, 1981) theorized that setal tufts on Cricotopus

sylvestris enhanced the larva’s ability to stay on aquatic plants in a strong tidal current. The setal tufts

could adhere to leaves, stems and filamentous algae, preventing the larva from being washed out ofthe

vegetation. Hershey and Dopson (1984 and pers. comm.) found that larvae of Cricotopus sylvestris,

which possess large setal tufts, were less susceptible to predation from Hydra than Cricotopus bicinc-

tus, which possess smaller setal tufts. In addition to attachment and anti-predation, other functions of

setal tufts may include floating, swimming, feeding, and antifouling.

140

As stated above, the occurrence of setal tufts has been a traditional character for separating the larvae

of some species of Cricotopus from all species of Orthocladins. Whether setal tufts occur in other lar-

vae of Orthocladins has yet to be determined.

Material examined: Rat Cove, Otsego Lake, Cooperstown, Otsego Co., New York, J. P. Fagnani: 1 male with

lar. and pup. ex., Ekman grab, 10° water, 24.1V.1982; 1 pharate female pupa with lar. ex., Ekman grab, 3’ water,

24.1V. 1982; 1 pharate female pupa with lar. ex., Ekman grab, 8.1V. 1981; pup. ex. and assoc. lar. ex., rock in 0.5° wa-

ter, 25.V1.1980; 1 pharate female pupa with lar. ex., ventral surface of Nuphar varıegatum leaves, 24. VI. 1980; 2

pre-pup. larvae. Ekman grab, 23.V.1980; 2 mature larvae, Ekman grab, 8.1V. 1981.

Acknowledgements

We would like to thank D. R. Oliver and P. S. Cranston for verifying the occurrence of setal tufts on additio-

nal larvae of Orthocladius annectens. This research was supported by the Biological Field Station, State University

of New York, College at Oneonta; the Wadsworth Center for Laboratories and Research, NYSDH; and CSRS,

USDA (FLAX 79009).

Literature

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organisms of the southeastern United States. - USEPA, Environmental Monitoring and Support Lab., Cin-

cinnati, Ohio. 195 pp.

—— 1976: Biology of larval chironomids. — Florida Dept. of Environ. Reg. Tech. Ser. 2. 58 pp.

CHERNOVSKII, A. A. 1949: Identification of larvae of the midge family Tendipedidae. (Trans. E. Lees, ed. K. E.

Marshall). — Nat. Lending Lib. for Sci. and Tech., Boston Spa., Yorkshire, England, 1961.

COFFMAN, W. P. and L. C. FERRINGTON 1984: Chironomidae, pp. 551-652. In: R. W. Merritt and K. W. Cum-

mins (Eds.). — An Introduction to the Aquatic Insects of North America, 2nd Ed., Kendall/Hunt, Dubuque,

Iowa. 722 pp.

CRANSTON, P. $. 1982a: A key to the larvae of the British Orthocladiinae (Chironomidae). — Freshwatr. Biol. As-

soc. Scı. Pub. 45. 152 pp.

—— 1982b: The metamorphosis of Symposiocladius lignicola (Kieffer) n. gen., n. comb., awood-mining Chirono-

midae (Diptera). — Ent scand. 13: 419—429

CRANSTON, P. S., D. R. OLIVER, and ©. A. SAETHER. 1983: 9. The larvae of Orthocladiinae (Diptera: Chironomi-

dae) of the Holarctic region — Keys and diagnoses, pp. 149-291. In: T. Wiederholm (Ed.). — Chironomidae

ofthe Holarctic Region. Keys and Diagnoses. Part 1. Larvae. — Ent. scand. Suppl. 19. 457 pp.

Dargv, R. E. 1962: Midges associated with California rice fields, with special reference to their ecology (Diptera:

Chironomidae). — Hilgardıa 32: 1—206

HERSHEY, A. E. and $. I. Dopson. 1984: Predator avoidance in Cricotopus: the importance of being big and hairy

(Abstract). — Bull. Ecol. Soc. Amer. 65: W185

HIRVENOJA, M. 1973: Revision der Gattung Cricotopus van der Wulp und ihrer Verwandten (Diptera, Chironomi-

dae). — Ann. Zool. fenn. 10. 363 pp.

JOHANNSEN, O. A. 1937: Aquatic Diptera. Part III. Chironomidae: Subfamilies Tanypodinae, Diamesinae, and

Orthocladiinae. — Mem. Cornell Univ. agr. Exp. Stn. 205. 84 pp.

Mason, W. M. 1973: An introduction to the identification of chironomid larvae. — USEPA, National Einviron-

mental Res. Center, Cincinnati, Ohio. 90 pp.

MENZIE, C. A. 1978: Productivity of chironomid larvae in alittoral area of the Hudson River Estuary. — Ph. D. the-

sis. The City University of New York. 127 pp.

—— 1981: Production ecology of Cricotopus sylvestris (Fabricius) (Diptera: Chironomidae) in a shallow estuarine

cove. — Limnol. Oceanogr. 26: 467-481

OLIVER, D. R., D. MCCLyMoONT, and M. E. RousseL 1978: A key to some larvae of Chironomidae (Diptera) from

the MacKenzie and Porcupine River watersheds. — Can. Fish. Marine Ser. Tech. Rpt. 791. 73 pp.

OLIVER D. R.andM. E. RousseL 1983: The Insects and Arachnids of Canada. Part 11. The genera of larval midges

of Canada (Diptera: Chironomidae). — Agric. Can. Pub. 1746. 263 pp.

141

ROBACK, $. $. 1957: The immature tendipedids of the Philadelphia area (Diptera: Tendipedidae). — Acad. Nat. Scı.

Philad. Monogr. 9. 152 pp.

SAETHER, O. A. 1969: Some Nearctic Podonominae, Diamesinae, and Orthocladiinae (Diptera: Chironomidae). —

Bull. Fish. Res. Bd. Can. 170. 154 pp.

—— 1980: Glossary of chironomid morphology terminology (Diptera: Chironomidae). — Ent. scand. Suppl. 14.

151 pp.

SHILOVA, A. I.and N. I. ZELENTSOV 1978: (A new genus and species of the subfamily Orthocladiinae [Diptera, Chi-

ronomidae]). — Zool. Zh. 57: 1584—1588

Sımpson, K. W.and R. W. BoDE. 1980: Common larvae of Chironomidae (Diptera) from New York State streams

and rivers with particular reference to the fauna of artifical substrates. — Bull. N. Y. St. Mus. 439. 105 pp.

Soronis, A. R. 1977: A revision of the Nearctic species of Orthocladius (Orthocladius) van der Wulp (Diptera: Chi-

ronomidae). — Mem. Ent.Soc. Can. 102. 187 pp.

THIENEMANN, A. 1944: Bestimmungstabellen für die bis jetzt bekannten Larven und Puppen der Orthocladiinen

(Diptera: Chironomidae). — Arch. Hydrobiol. 39: 551 —564

Joseph F. Fagnanı

Wadsworth Center for Laboratories and Research

New York State Department of Health

Albany, NY 12201 U.S.A.

AnnelleR. Soponis

Department of Entomology

Florida A & M University

Tallahassee, FL 32307 U.S.A.

142

SPIXIANA | Supplement 14 143 — 154 München, 15. Julı 1988 ISSN 01777424

Aquatic xylophagous Orthocladiinae — systematics and ecology

(Diptera, Chironomidae)

By P. S. Cranston and D.R. Oliver

Abstract

Wood-mining Orthocladiinae are reviewed. Chaetocladins ligni, the first xylophagous member of the genus, is

described as new to science. Symposiocladius Cranston, erected for lignicola Kieffer, is synonymised with, and trea-

ted as a subgenus of, Orthocladius Wulp. Extensive morphological variation in O. lignicola is recognized and dis-

cussed. Observations on the biology of xylophagous chironomids are reviewed with suggestions for further study.

Introduction

Larval Chironomidae occur in the most diverse geographic and habitat ranges of any insect family,

from high Arctic lakes, permanent montane snowfields and glaciers through temperate and tropical

ecosystems to terrestrial mosses in the Antarctic. In many aquatic biotopes, chironomids may domi-

nate in abundance and even biomass, despite their relatively small size. Studies on systematics and eco-

logy of northern hemisphere Chironomidae have elucidated their importance in lake classification,

both extant (e. g. Brunpın, 1949; SAETHER, 1979; WIEDERHOLM, 1976) and historic (e. g. WAarwıck, 1975),

and allowed recognition that chironomid community structure is an integral part of environmental

monitoring and assessment of lotic water quality.

A research area that has received rather less intensive study ıs the röle of larval chironomids in the

decomposition of aquatic immersed wood. Foremost amongst those examining invertebrates in such

ecosystems is N. H. Anderson and his associates at Oregon State University. Their work has resulted

in both research and review publications (e. g. AnDERson et al., 1978; Dupıey & Anperson, 1982) and

the provision of specimens of wood-mining Chironomidae for systematic studies. This material re-

veals that taxonomic problems remain and there are still undescribed xylophagous species of ecologi-

cal significance. We take this opportunity to describe a new species of wood-mining Chaetocladıns

Kieffer, to reassess the taxonomy of Symposiocladins Cranston, and to consider the range and diversity

of xylophagous Chironomidae. We conclude with some observations on past studies and make

suggestions for future research.

Methods and abbreviations

Allnew material examined is slide mounted in Canada Balsam or Euparal and deposited in the Canadian National

Collection, Ottawa, Canada (CNC), unless otherwise stated. Abbreviations of other Institutes are as follows:

OSU _-— Oregon State University, Corvallis, Oregon, U.S.A.

BMNH British Museum (Natural History), Cromwell Road, London, England.

MZBN - Museum of Zoology, Bergen, Norway.

USNM United States National Museum, Washington, D. C., U.S.A.

ZSBS _- Zoologische Staatssammlung, Munich, German Federal Republic.

143

Morphological terminology follows SAETHER (1980). Other abbreviations used in text: dc, _, — Pupal dorso-

central setae; Fe — Femur; L. — Larva; Le. — Larval exuviae; L. R. — Leg ratio = Length of tarsomere 1: Length

of tibia; P. — Pupa; Pe. — Pupal exuviae; pc,_; — Pupal precorneal setae; Ta,_; — Tarsomeres; Tı — Tibia.

Chaetocladius (Chaetocladius) ligni spec. nov.

Adult male (n = 5):

Body length 2.8-3.4 mm, wing length 1.48— 1.82 mm.

Antennal ratio 0.55—0.99, apical flagellomere 213-340 um long. Head with 3-5 inner verticals

continuous with 6-9 outer verticals and postorbitals; 7-10 clypeals; palp segment 4 shorter than 3rd

and 5th segments, 5th segment longer than 3rd.

Thorax with 7—9 lateral antepronotals, 13—18 uni— to partly biserial dorsocentrals, 5—8 uniserial

posterior prealars, 6— 10 uniserial scutellars.

Wing wıth obtuse anal lobe, brachiolum with 1 setae, R with 5—21 setae, R, with 5-9 setae, R,,;

with 12—21 setae; 5-8 squamals.

Legs with pseudospurs on tarsomere 1 and 2 of mid and hind legs; tarsomere 1 of mid leg with 0 sen-

silla chaetica, hind leg with 0-3 sensilla chaetica on apical one-quarter and sometimes also on middle

third; pulvilli small. Fore leg ratio, 0.71—0.76; mid leg ratio, 0.42—0.47; hind leg ratio, 0.55—0.58.

Hypopygıum (Fig. 1) with long, narrow anal point. Virga consisting of cluster of long, dark spines.

Gonocoxite with dorsal part of inferior volsella narrow, bare, apically rounded and slightly curved

posteriorly; ventral part variable in shape, usually rounded, sometimes almost absent. Gonostylus

with rounded outer corner and low, broad, crista dorsalıs.

Figs. 1-5. Chaetocladins ligni spec. nov. Adult male, — 1 Hypopygium . Adult female, — 2 genitalia, — 3 tergite

IX. Male pupa, — 4 segments I and II, left tergites, right sternites, — 5 segments VIII and anal end, left dorsal, right

ventral.

144

Adult female (n = 2):

As male, except, wing length 1.66-1.82 mm.

Antenna with 5 flagellomeres, apical flagellomere with strong apical seta. Head with 11-12 tempo-

rals, 10-13 clypeals.

Thorax with 9-12 lateral antepronotals, 26-33 dorsocentrals, 8-9 posterior prealars, 9 scutellars.

Wing with obtuse (almost rıght-angled) anal lobe; R wıth 18-20 setae, R, with 12-16 setae, R,;;

with 29—30 setae; 6-9 squamals.

Legs. Tarsomere 1 of mid leg with 27 (n = 1) sensilla chaetica on distal two-thirds, hind leg with

29—32 sensilla chaetica on distal two-thirds. Fore leg ratio, 0.74—0.75; mid leg ratio, 0.41—0.45; hind

leg ratio, 0.56.

Genitalıia (Fig. 2) very similar to those illustrated for the genus by S£THer (1977, fig. 54 A—C) ex-

cept concavity of distal margin of tergite IX deeper (Fig. 3). Also 2 long, strong setae arise from each

cercus.

Pupa (n = 12):

Length, 2.9-3.7 mm.

Frontal apotome with small frontal seta on small cephalic tubercle.

Thoracic horn cylindrical with spinose apex (Fig. 9), 112-165 um long. Narrow, pointed to

bluntly rounded tubercle on scutum dorsomedial to base of thoracic horn (Fig. 9).

Tergites II-VIII covered wıth coarse spinules grading into posterior spine band; sternites with

weak shagreen on II becoming more extensive and stronger on successive posterior sternites; posterior

margin of sternite VIII straight with spines (C') or bilobed without spines (9) (Figs. 4-6). All L-setae

hair-like. Anal lobe broad with outer margin up-curved; megasetae about 10x as long as wide, with

middle megaseta located close to posterior megaseta (Figs. 5, 6).

Figs. 6-13. Chaetocladins lıgni spec. nov. Pupa, — 6 female, segments VI & VIII and anal end, left dorsal, right

ventral, 9 thoracic horn, precorneal setae and scutal protuberence/spine. Larva, — 7 mentum, — 8 antenna, — 10 la-

brum and premandibles, — 11 maxilla, — 12 mandible, — 13 anal end.

145

Larva, Fourth instar (n = 15):

Antenna (Fig. 8) five-segmented with segments consecutively shorter or segments 3 and 4 subequal;

antennal blade ending at level of segment 5; Lauterborn’s organs large, subequal in length to segment

3. Antennal ratio, 1.06—-1.43 (n = 13).

Labrum (Fig. 10) with anterior part folded under frontoclypeus; SI smooth, lamelliform, $ II and

S III smooth, S IV consisting of 2 simple pegs; labral lamellae absent; labral chaetae and spinulae

smooth with chaeta media serrate on one margin; pecten epipharyngis consisting of 3 small spines;

median pair of chaetulae laterales large, rectangularly rounded, covering rest of chaetulae; chaetulae

basales with bifid apex; basal sclerite large.

Premandible with 1 apical tooth and low-rounded accessory tooth; brush weak.

Mandible (Fig. 12) 144—166 um (n = 11) long with short apical tooth and 4 inner teeth; seta interna

with 5 serrate branches and 1 long smooth to serrate branch.

Mentum (Fig. 7) with long apically bifid median tooth and 4 pairs of shorter lateral teeth; lateral

notch present at level of base of 4* lateral tooth; ventromental plates small.

Maxilla (Fig. 11) with weak pecten galearis.

Abdomen (Fig. 13) with preanal segment curved over remaining posterior segments; procercus

with 6 anal setae, directed posteriorly.

Type-material: Holotype O’ slide-mounted in Canada Balsam: U.S.A., Oregon: Benton County, Berry Creek,

15.X.1984 (mass reared from immersed wood) (N. H. Anderson) (CNC No. 19701). Paratypes: U.S.A., Oregon:

same locality as holotype, 12.X.1982—12.XI. 1984 (mass reared from immersed wood) (N. H. Anderson) (19,20

P. with associated Le., 19 P., 11 Pe., 20L.); Crook County, Allen Creek, 26. VIII.1978 (T. Dudley) (19 P.),

Ochoco Creek, IX. 1978 (T. Dudley) (1 0°); Lake County, Mack Creek, 7. VII.—28.1X. 1982 (mass reared from ım-

mersed wood) (N. H. Anderson) (20°C, 10° with associated Pe., 19,1 P.,2L.)(BMNH, CNC, MZBN, USNM,

ZSBS).

Remarks:

Adult males of C. ligni are similar to those of C. melaleucus (Meigen) (Epwarps, 1929; PINDER, 1978),

differing by having a lower antennal ratio and a smaller inferior volsella. The presence of two long cer-

cal setae will distinguish the adult females from all other Chaetocladins including C. melaleucus (see

GOETGHEBUER, 1942). Also the immature stages do not equate with the adult male resemblance (see

Pınper & ArMITAGE, 1985, for detailed descriptions of the immature stages of C. melaleucus).

The distinctive scutal tubercle is not known to occur on other Chaetocladius pupa except on asingle

pupal exuviae mass reared from immersed wood (Quebec, Gatineau Park, stream nr. Beech Grove,

5.V1.1985, P. S. Cranston & M. E. Dillon). This exuviae, otherwise similar to C. ligni, has an almost

smooth thoracic horn and short (c. 3X as long as wide), thorn-like megasetae. An adult male obtained,

at a later date, from the same rearing container cannot be distinguished from C. ligni. At present, it is

not possible to determine if the pupal differences are due to variation because of the absence of larvae

and positive association of the two life history stages. Therefore, eastern specimens, including an adult

male from New Hampshire (White Mountain National Forest, Ammonoosuc River, 26. V.1981, D.

R. Oliver & M. E. Roussel) are not included in the type series of C. ligni.

The larva differs from the generic diagnosis of Cranston et al. (1983) plus the addition by Pinder

and Armrrace (1985). SI is simple not serrated, plumose or branched, the premandible has 1 tooth api-

cally, not 2, the basal sclerite is large, and the preanal segment is curved over the posterior segments.

Furthermore, the mentum has only 4 lateral teeth although the lateral notch may be the remains of the

notch between the 4" and 5" lateral teeth.

Orthocladius Wulp subgenus Symposiocladius Cranston stat. nov.

Orthocladius WuLp, 1874: 132. Type-species: Tipula stercoraria DEGEER, 1776: 388 sensu Wulp (misident.) (des.

COQUILLETT, 1910: 581) = Chironomus oblidens WALKER, 1856.

146

Symposiocladius CRANSTON, 1982: 419. Type-species: Orthocladius lignicola Kieffer in POTTHAST, 1915: 273 (orig.

des.). Syn. nov.

Symposiocladins was erected for the xylophagous Orthocladius lignicola Kieffer, a species with a

highly distinctive larva, but closely resembling Orthocladius (Orthocladius) in pupa and adult (Cran-

ston, 1982). The major reason given for the erection of anew genus was the presence of the larval ab-

dominal l, seta developed as a setal brush, which Cranston described as an unique inside parallelism/

underlying synapomorphy for the lineage Synorthocladius to Cricotopus (1. e., excluding Orthocla-

dius). Two recent discoveries expose the fallacy of this reasoning: Orthocladius annectens Saether has

previously unrecognized larval setal tufts and, since there is no doubt that annectens is truly an Ortho-

cladius, setal tufts do occur in the Orthocladins lineage. Furthermore, lignicola larvae with only simple

abdominal 1, setae are now known (see below). The case well illustrates the erroneous nature of phy-

logenetic reasoning based upon ambiguous characters, particularly the use of non-unique derived cha-

racter states (”"underlying synapomorphies”).

However, since the larva is highly distinctive and the pupa and female can be distinguished from ot-

her subgenera of Orthocladius, Symposiocladius can be retained as a subgenus of Orthocladius s. |. un-

til the phylogeny ıs assessed and the ranking of other taxa presently given generic status, such as

Stackelbergina Shilova and Zelentzov, is resolved.

Orthocladius (Symposiocladius) lignicola Kieffer

Orthocladius lignicola Kieffer ın POTTHAST, 1915: 273; CRANSTON, 1982: 421. Type-locality: West Germany, Sau-

erland, Haspersperre. Lectotype pupa examined CRANSTON, 1982: 421 (ZSBS).

Cransrton (1982) described all stages of O. lignicola — “typical” lignicola in this discussion refers to

this description. All specimens are closely associated as larvae with wood immersed in running water.

Localities referred to in the discussion are cited in detail below.

In Berry Creek, O. (Symposiocladius) larvae (n = 100+) virtually always have a simple, or at most

2 branched abdominal l, seta and have more or less fused lateral mental teeth (Fig. 15a) but otherwise

resemble typical lignicola, with all measurements falling into the lower end of the range cited (Cran-

ston, 1982: 426). This larval type (“A”) is associated with pupae (type “A”) lacking frontal setae, with

a weakly spinose thoracic horn (Fig. 14a) 200-280 um long (n = 40), and with a blunt spinose apex

to the anal lobe (Fig. 16a). Adult males from Berry Creek fall towards the small end of the ranges cited

by Soronis (1977: table ZZ, as tryoni) except for the terminal antennal flagellomere (450-550 um

long) and antennal ratio (0.98-1.18,n = 9) which fall at or below the cited range. Females cannot be

distinguished from typical lignicola (Cranston, 1982: 421). Despite the apparent identity of the geni-

talıa of both sexes, the low male antennal ratio, simple larval I, seta, absence of pupal frontal setae and

differences in pupal anal lobe conventionally would indicate specific distinction. However, from

Berry Creek we have seen a single ©. (Symposiocladius) larva with fused lateral mental teeth, but with

abdominal l, setal tufts with 6-8 branches about 80 um long (typical lignicola have 20+ branches, up

to 160 um long) and one typical ignicola pupa with pharate adult (terminal flagellomere 620 um long,

antennal ratio 1.4). Furthermore a single prepupal larva oftype “A” from Yukon contains a pupa with

typical lignicola curved spinose apex to the anal lobe, but the frontal setae and thoracic horn cannot

be seen.

In Mack Creek only typical lignicola larvae have been found (n = 25). Pupal exuviae derived from

mass rearing from colonised immersed wood (not directly associated) all have a curved, spinose anal

lobe apex typical of lignicola, a rather densely spinose cigar-shaped thoracic horn (Fig. 14c), length

165-201 um, but lack frontal setae (n = 17). This pupal type is associated with typical lignicola larvae

associated with prepupae and a pupa from Starvation Creek. Mass rearings from immersed wood from

Mack Creek give rise to males with a relatively long apical flagellomere (c. 600-650 um) but rather

1574

a b 16 c d

Figs. 14-16. Orthocladius (Symposiocladius) lignicola Kieffer. Pupa, — 14 thoracic horn (a — Berry Creek,

b — Beech Grove, c — Mack Creck), — 16 apex of anal lobe (a — Berry Creek, b and c — Beech Grove, d — Mack

Creek). Larva, — 15 mentum (a — Berry Creek, b — Beech Grove).

low antennal ratio (1.2—1.3). This Mack Creek pupal type also occurs in Flynn Creek, together with

type “A” pupae. Both typical /ignicola and type “A” larvae were present, but no associations are avai-

lable.

Re-examination of most of the material studied by Cranston (1982) confirmed that none of these

“aberrant” types mentioned above were present, unrecognized, and showed that frontal setae (omit-

ted from the description) were invariably present in earlier examined Nearctic and Palaearctic speci-

mens. However, the low antennal ratio (relative to the mean value) cited by Soronis (1977: table ZZ)

suggests that some adults associated with the larval types above are included, since adults associated

with typical lignicola larvae and pupae tend to have an antennal ratio of at least 1.5.

In order to resolve the status of the specimens discussed above, immersed wood, mostly Alnus sp.,

known to contain O. (Symposiocladius) larvae was collected in mid-June and again in mid-July from

a stream in Gatineau Park near Beech Grove. This was kept in an oxygenated aquarium cooled to ap-

proximately 12°C and emergent adults, cast skins and pharate adults were collected at intervals over

the ensuing four months. Individual rearings were not possible since lignicola larvae are easily dama-

ged in handling and die very readily. However, some larval/pupal and pupal/adult associations were

made through failed emergence at the aquarium water surface. Alllarvae examined (n = 29) are typical

lignicola, with well developed setal tufts, although the lateral mental teeth division is somewhat varıa-

bly developed (Fig. 15b). Most pupae (n = 38 of 40) have frontal setae, amoderately spinose thoracic

horn 220-275 um long (Fig. 14b) and an anal lobe intermediate (Figs. 16b—c) between that of type

“A” and typical lignicola (Figs. 16a, 16d). One pupa, otherwise similar to this, has only 1 scar of a

frontal seta on one side of the frons, with no trace on the other side. One pupa has no trace of frontal

148

setae, a weakly spinose thoracic horn 160 um long and a blunt spinose apex to the anal lobe, closely

resembling pupal type “A”. The pharate adult within this pupa has an apical flagellomere length

415 um and antennal ratio of 1.13. Adult males associated with the dominant pupal type in this mass

rearing could not be distinguished from typical lignicola, having an apical flagellomere 540-640 um

long and antennal ratio of 1.4—-1.6 (n = 10). Adult females cannot be distinguished from typical ligni-

cola.

Clearly, from the evidence above, it is impossible to recognize separate species which can be confir-

med from all life history stages. The adult males could be divided into those with an antennal ratio be-

low 1.2 and a relatively short apical flagellomere (for example, below 550 um) and those with a higher

antennal ratio and longer apical flagellomere, but there is virtually continuous variation in these cha-

racters. There is remarkably little variation in both absolute size and wing length in all specimens ex-

amined, with the range less than 10 % each side of the mean, thus it is unlikely that the increased length

ofthe apical flagellomere and antennal ratio is explained by allometry. Although the low antennal ratio

male is apparently associated with the type “A” larva and pupa, this ıs not exclusively so. The highest

values of antennal ratio and apical flagellomere length are found in association with typical Jignicola

larvae and pupae in Nearctic and Palaearctic populations. Intermediate values of antennal ratio and

apical flagellomere length are found in Mack and Flynn Creek and Quebec populations which have

typical lignicola larvae but at least 3 different combinations of pupal characters. However, there is no

detectable correlation between pupal characters of the anal lobe, frontal setae and thoracic horn and

the complex tergal and sternal spine patterns, pedes spurii A and B and thoracic setation. Adult fema-

les, increasingly recognised as valuable in the detection of cryptic species, are relatively invariate and

indicate a single species.

In each intensively sampled habitat more than one pupal type occurs, but this is not always correla-

ted with the occurrence of more than asingle type of larva or male. The variation evident in the Quebec

pupae, for example is not evident ın larvae or adults. We conclude by suggesting that on present evi-

dence O. lignicola ıs a single species and cannot yet be divided.

At present, it is not possible to explain the variation seen in Jignicola, although several possibilities

can be raised. There may be a clinal varıation in the length of the apical antennal flagellomere and an-

tennal ratio. The highest values for both are found in the easternmost populations examined in Quebec

and the lowest values in Oregon. Superimposed on this, there is some evidence that winter (Oregon)

or spring (Quebec) adults have a lower antennal ratio and shorter apical antennal flagellomere than

adults emerging later. This must be analysed in more detail, with more individual associations of all

life history stages and with control over the effect of rearing in laboratory cultures.

The apparently excessive varıation in O. lignicola, ın characters that are believed to be of specific

importance in other taxa, might suggest a faulty species concept or inability to discriminate closely re-

lated species. However, recognition is growing that some taxa do show high variation in terms of tra-

ditional characters used in delimitation of chironomid species. For example, the hygropetric Ortho-

cladıus (Eudactylocladius) fuscimanus Kieffer and O. (E.) dubitatus Johannsen both show as much va-

ration as does O. lignicola. In all cases the varıation is demonstrated to be intraspecific through recog-

nıtion of high variation present within a single population.

New material examined: CANADA, Yukon Territory: Alaska Highway, Watson Lake Campground, 60°07'N,

128°48’ W, 23.V.1982 (E. Fuller) (1 prepupal L.); Quebec: Pontiac County, Gatineau Park, 8 km N. Beech Grove,

45°35’N, 76°8'’W, 18.V.1983 (D. R. Oliver & M. E. Roussel) (3 Pe.), 17. VI.1985 (mass reared from immersed

wood) (P. S. Cranston and M. E. Dillon) 1090, 8 P9,1C'and 19 with associated Pe., 1 P. with associated Le.,

44Pe., 29L., slide mounted, many more in spirit); Missisquoi County, Im. N. Abercorn, (reared ex Acer)

19.-27. VIII. 1980 (A. Borkent) (30'0',1Q P. with associated Le., 9Pe.,). U.S.A., Oregon: Benton County, Berry

Creek, 14.-28. VII. 1982 (mass reared from immersed wood) (N. H. Anderson) (9 0'C', 10° with associated Pe.,

1099,1QP. with associated Le., 40 Pe., 100+L); Lincoln County, Flynn Creek, 30. VII.-28. XII. 1982 (mass rea-

red from immersed wood) (N. H. Anderson) (20°C, 1C°’P., 1QP., 8Pe., 20+L.); Lane County, Mack Creek,

7.VII.—28.1X.1982 (mass reared from immersed wood) (N. H. Anderson) (3 0°C’, 17Pe., 20+L.); Hood River

149

County, w. of Hood River, Starvation Creek, 27. V1.1981 (A. Borkent) (1 Le. with associated prepupa, 5 L.). Some

Berry, Flynn and Mack Creek specimens in collection of N. H. Anderson (OSU), remainder in CNC.

Discussion

A major impediment to the elucidation of the röle of Chironomidae in aquatic wood decomposi-

tion, even in the best studied north temperate regions of the world, has been taxonomic. For example,

Cranston (1982) traced the first discovery of a distinctive larva (called “acutilabis”) to Konstantınov

(1948), yet it was 30 years before its xylophagy was established, despite Thienemann’s knowledge that

the eventually associated pupa and adult belonged to a xylophage (hence the specific name lignicola gi-

ven by Kieffer). Similarly, although the genus Stenochironomus Kieffer has been known to include

wood-mining larvae since ZagLotskt (1939), it was not until BORKENT’s (1984) monograph that its great

taxonomic diversity and the wood-mining of its relatives was revealed.

Figs. 17—22. Larva, outline of anterior margin of mentum. — 17 Orthocladius (Symposiocladins) lignicola Kieffer,

— 18 Chaetocladins ligni sp. nov., — 19 Limnophyes sp. ident., — 20 Xylotopus par (Coquillett), — 21 Polypedilum

(Polypedilum) fallax (Johannsen), — 22 Harrisius pallidus Freeman, — 23 Stenochironomus (Stenochironomus) hila-

rıs (Walker), — 24, Xestochironomus subletti Borkent.

There are several reasons for the relative paucity of taxonomic studies, including failure to include

woody substrates in routine aquatic invertebrate surveillance, difficulties in sampling immersed wood

and the fragile nature of many xylophagous larvae. However, there is no doubt that suitably located

immersed wood of an appropriate age, especially Alnus spp., may have dense mixed populations of

wood-mining Chironomidae, contributing substantially to the diversity and biomass of the aquatic

biota. For example, Kaurman (1983) reported an annual standing biomass of 5000 mg/m? of Xylotopus

par (Coquillett) and Anderson (in prep), a lower figure of 57 mg/m? from a mean of 618 C. lignı and

O. lignicola/m?. Sampling can pose difficulties and assessment of adult emergence from rearings from

aquarium retained wood may avoid many problems. However this method, widely used by Anderson

and his colleagues, only allows, at best, circumstantial larval/adult associations. Furthermore, wood

held in the laboratory gives rise to adult emergence from both xylophages and insects using the wood

surface as a substrate for other feeding methods. High pressure hosing of the wood after recovery from

the aquatic habitat may remove many non-stenotopic invertebrates, but ambiguity remains. How-

ever, obligate wood feeders starve if removed from wood. Thus the optimum method to obtain asso-

ciations of xylophagous larva, pupa and adult is to mass rear in natal wood, obtaining larval head

capsules by recovery from mines that contain pupae whose identity can be established through pupal/

adult associations either as pharate adults or through partial emergence.

150

Mass rearıng, as discussed above, does not readily discriminate between obligate xylophages and

other benthic insects associated in a less stenotopic manner with woody substrates. Discussions con-

cerningxylophagy are handicapped by our still limited knowledge of the precise nature of associations

with wood. A major criterion is the making of mines or galleys in wood, and/or the domination of the

larval gut contents by wood. Even observations on gut contents may be ambiguous, since insects de-

riving epilignic nutrients (from superficial components on immersed wood) will ingest non-dietary

wood fibres, particularly from extremely soft and decomposing woods. Analyses of xylophagous chi-

ronomid guts to establish the presence either of cellulases or a symbiont cellulose decomposer fauna,

required to confirm stenoxylophagy, have been made only in X. par (Kaurman et al., 1986) and even

this study failed to demonstrate cellulase activity of gut microflora. Therefore, our assessments of ha-

bit are of necessity somewhat circumstantial, but we present the following list and categories of xylo-

phagous Chironomidae:

Taxa strongly suspected of being obligate xylophages, only found mining in immersed wood, and

with guts consistently filled with wood fibres, include: Chironominae: Stenochironomus (Stenochiro-

nomus), Harrisins Freeman and Xestochironomus Sublette & Wirth species (BorkENT, 1984); Polype-

dilum (Polypedilum Kiefter), including at least Nearctic fallax (Johannsen) (JOHANNsEN, 1937, ROBACK,

1953); Orthocladiinae: Chaetocladins ligni sp. nov.; Orthocladins (Symposiocladius) lignıcola Kieffer

(Cransron, 1982); Xylotopus par (Coquillett) (OLıver, 1982; 1985).

Taxa which may be obligate xylophages but confirmation is lacking include: Chironominae: Glyp-

totendipes (Phytotendipes) testaceus Townes (reared from a water soaked log in Arkansas, Derreoux

Creek, W. Pine Bluff, route 270, 12.111. 1972, H. Teskey, C.N.C.); other perhaps misidentified Glyp-

totendipes, including gripekoveni (Kieffer) from the Netherlands (Grirekoven, 1914) glaucus (Meigen)

(= pallens [|Meigen]) from England (Burrr, 1940) and “dendrophila” from the Soviet Union (ZvErEvA,

1950); Orthocladiinae: ZLimnophyes sp. indet. unreared from Oregon streams, perhaps associated with

L. pilicistulus Saether adults emerged from laboratory mass rearıngs of immersed wood (N.H. Ander-

son).

Taxa with distinct associations with immersed wood, but either unsubstantiated as miners and/or

having a wood diet: Chironominae: Stelechomyia perpulchra (Mitchell) from S. E. USA (Reiıss, 1982);

Polypedilum (Polypedilum) including palaearctic pedestre (Meigen); Orthocladiinae: Brillia, particu-

larly flavifrons group species in Palearctic and Nearctic regions; Diamesinae: Pagastıa larval type A

of OLıver and Rousser, 1982 (Nearctic observations by CRAnsToN).

Many other Chironomidae have been recorded from immersed woody substrates but the nature of

the wood association is unclear. For example, many of the genera listed by Dupıry and AnDERSON

(1982) appear to be no more than casual visitors using the wood as either resting sites or for feeding.

Our categories above reflect our knowledge of mining: we have too little evidence to consider the röle

of these epiphytic taxa ın wood degradation.

_ The wood mining taxa listed above show no close phylogenetic relationships to one another, with

the exception of internal relationships within the Stenochironomus complex. Thus several unrelated

(or at least very distantly related) taxa have come to occupy aquatic woody microhabitats. Three sub-

families are represented: the Diamesinae, Orthocladiinae and Chironominae. Only within the Steno-

chironomus complex (BorkeEnT, 1984) is there evidence of species formation in amonophyletic grou-

ping probably following ancestral adoption of the wood mining habit (BorkENT, 1984). In other taxa

here considered the mining habit has been independently acquired. Phyletically close relatives of

mining taxa tend to be rheophilic and, where known, appear to be Aufwuchs grazers. Taxa in which

mining is probable, but not established with certainty, include some species that otherwise burrow in

immersed leaves (Brillia, Stelechomyia, Polypedilum). Since there are examples of presumed wood

mining in which the immersed wood is so soft that it is friable by hand, there may be little distinction

between grazing on immersed leaf surfaces and on soft wood. In either case, the source of nutrition

may be other than wood derived cellulose since the chironomid gut contents frequently contain algae

and fungi in addition to wood fibre.

151

Given that wood mining Chironomidae are not related, are there morphological features (conver-

gences) associated with the larval habit? One of the few characteristics that unite many miners, irre-

spective of subfamily, is the very flimsy body cuticle, which cannot support the flaccid larva outside

the mine. This is particularly striking in O. (Symposiocladins) and Stenochironomus, less so in Xyloto-

pus, and explains the fragility of the larva leading to larval death if extracted live from wood for rearing.

Within the Orthocladiinae, structures of the wood miners’ head capsule show some convergence, with

the mental teeth characteristically elongate (Figs. 17—20) and heavily sclerotised relative to non-mi-

ning congeners or close relatives. Other similarities include the broadening and shortening ofthe man-

dible and the narrowing of the anterior labrum, with a folding of the anterior part beneath the fronto-

clypeus in such a way that it cannot be flattened readily for microscope examination. In C. ligni, O.

(S.) lignicola and the undescribed mining Limnophyes species referred to above, the SI seta is simple

compared to the condition in congeners, but there is little evidence of any further reduction in head

capsule appendages such as the antenna, maxilla, and labral chaetae, or in the abdominal appendages

(prolegs, procerci, and anal tubules). In contrast, the obligate wood miner X. par lacks these orthocla-

diine character state changes associated with the mining habit, with the exception of the lateral com-

pression and strong folding of the labrum beneath the frontoclypeal apotome.

In contrast to the Orthocladiinae, mining Chironominae do not have an anteriorly projecting me-

dian part of the mentum (Figs. 21-24). Stenochironomus do indeed have a modified mentum, but this

mentum is transverse with very reduced teeth. The very great reduction of the striae of the ventromen-

tal plates is also characteristic, is perhaps associated with a loss of silk making and is part of a transfor-

mation series that includes non-wood mining species. Polypedilum miners appear little different from

non-mining congeners and neither does Glyptotendipes testaceus differ greatly from its non-(wood)

mining relatives. The diamesine Pagastıa (larval type A) has a most curiously modified mentum with

the teeth obscured by lying ventral (and posterior) to a continuous ventromental plate (OLıver &

Rousser, 1982, fig. 4). It is impossible to speculate on how this unusual structure may be related to fee-

ding, let alone to wood mining.

Although much work remains to be done, some ecological generalisations can be made concerning

wood-mining Chironomidae. Firstly, mining Chironomidae may be locally abundant but by no me-

ans all immersed wood contains larvae. In lotic systems, suitable wood must be anchored in the cur-

rent, but not such that it becomes covered in sediment. Mining Orthocladiinae are not found in poorly

oxygenated running waters, although at least in Britain, O. ($.) lignicola larvae can tolerate high orga-

nic loadings in fast current. In contrast to the situation in lotic systems, Orthocladiinae are much less

common in lentic habitats, where Chironominae (notably Stenochironomus) dominate the wood-mi-

ning chironomids. Once again, immersed wood must be free from sediments for successful colonisa-

tion (A. Borkent, pers. comm.).

The quality and texture of wood is important in relation to the frequently observed differences in

colonisation of different wood species. For example, it isacommon observation that alder (Alnus spp.)

in the Holarctic, and maple (Acer) in the Nearctic, are strongly preferred substrates for many miners.

Unquantified observations in Britain show that in a stream known to support O. lignicola, freshly pla-

ced Alnus branches were colonised after five winter months and larvae were not found in any other

immersed wood, ranging from oak (Quercus) to pine (Pinus). O. lignicola is most abundant in wood

so recently immersed that a knife is required to excise larvae and is not found in soft wood. In contrast,

X. par and Pagastia larval species A appear to be most abundant in wood friable enough to crumble

by hand. However, faunal succession in relation to temporal changes in wood quality are littleknown

at present. The röle of chironomids in wood breakdown and their dietary use of apparently refractory

wood and/or associated microbial flora are poorly understood and elucidation through further study

is required.

Acknowledgements

We wish to acknowledge Professor N. H. Anderson and his colleagues at Oregon State University, Corvallis, for

providing much material examined in preparation of this paper. We thank M. E. Dillon for her technical assistance

in field work, rearing and preparation of materials of wood-mining Chironomidae. Part of this study was underta-

ken while P.S.C. held a postdoctoral fellowship from the National Science Environmental and Research Council,

Canada at the Biosystematics Research Institute.

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ANIC

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Agriculture Canada, Ottawa K1A 0C6, Canada

154

Chironomidae of South India. I. Generic composition, biogeogra-

phical relationships and descriptions of two unusual pupal exuviae

(Diptera, Chironomidae)

By William P. Coffman, L. A. Yurasits, Jr. and Carlos de la Rosa

Abstract

Collections of pupal exuviae were made from 25 sites (lotic, lentic and marine) in the states of Tamil Nadu and

Kerala, South India. Species-level taxa belonging to 55 named genera (35 new to India) were encountered — increas-

ing the number of named genera known to occur in India from 38 to 73. An additional 20 possibly “new” genera

were also discovered. Descriptions for two of these “new” genera, which are not referable to any known subfamily,

are given. Almost all of the named genera collected in S. India are known to occur widely in the Holarctic region,

and many are subcosmopolitan. The 20 “new” genera may represent an endemic component in $. India. However,

there is some evidence that at least a few of these “new” genera may occur elsewhere in the Oriental region.

Introduction

SUBLETTE & SUBLETTE (1973) list 56 genera of Chironomidae from the Oriental Region. Due to synonymy only

53 of the generic names they give are still considered to be valid (AsHE, 1983). Of these, only 21 had been recorded

from India (Table 1). Since 1973 anumber of investigators have added to our knowledge of the generic composition

of the chironomid fauna of India (BHATTACHARYA et al., 1985 a, 1985 b; CHAUDHURI & DEBNATH, 1985; CHAUD-

HURI & GHOSH, 1981, 1982a, 1982 b, 1986; CHAUDHURLI et al., 1984; CHAUDHURI et al., 1979; GHOSH & CHAUD-

HURI, 1983; Hirvenoja, 1985; SINGH & KULSHRESTA, 1975; SINHARAY & CHAUDHURI, 1978, 1984; SINHARAY et al.,

1978). As a result of these efforts, the number of genera known to occur in India has risen to 38 (Table 1).

Nearly all of the distribution records given for the Chironomidae of India in SUBLETTE & SUBLETTE

(1973) and all but one (Chaetocladins) of the more recent additions to the Indian fauna have come from

Indian states bordering on the Himalayas. In this paper we are presenting the results (at the generic

level) of a series of collections of pupal exuviae made in the southern Indian states of Tamil Nadu and

Kerala. Broad biogeographical affinities of this fauna are discussed and two unusual pupal exuviae,

that cannot be placed in any subfamily, are described. Forthcoming papers will treat the subfamilies

Tanypodinae, orthocladiinae and Chironominae.

Material and Methods

All of the specimens utilized in this study were collected using the surface drift pupal exuviae method (BRUNDIN,

1966; COFFMAN, 1973; PINDER, 1986; WILSON & BRIGHT, 1973). Collections were made with fine mesh drift nets

and soil sieves (125 microns) and field preserved in 70% EtOH. Collections were sorted at 12X and series of each

species from each sample were slide mounted with Euparal for identification and study. Abbreviations utilized in

the descriptions are from SAETHER (1980).

155

Table 1.

sources.

SUBLETTE & SUBLETTE (1973)

Chironomidae genera reported from India in SUBLETTE & SUBLETTE (1973) and from recent literature

Recent Literature

TANYPODINAE

Ablabesmyia Paramerina (CHAUDHURI & DEBNATH, 1985)

Clinotanypus

Procladius

Tanypus

DIAMESINAE

Diamesa

Sympotthastia

ORTHOCLADIINAE

Brıllia Bryophaenocladius (GHOSH & CHAUDHURI, 1983)

Cricotopus Chaetocladinus (CHAUDHURI & GHOSH, 1982)

Heterotrissocladius Eukiefferiella SHINHARAY et al., 1978)

Parametriocnemus Limnophyes (CHAUDHURI et al., 1979)

Metriocnemus (SHINHARAY & CHAUDHURI, 1978)

Nasuticladins (SINHARAY & CHAUDHURI, 1984)

Orthocladins (CHAUDHURI & GHOSH, 1982 b)

Paracladius (HIRVENOJA, 1985)

Paratrichocladius (HIRVENOJA, 1985)

Thienemannia (BHATTACHARYA et al., 1985 a)

CHIRONOMINAE

CHIRONOMINI

Chironomus Beckidia (BHATTACHARYA et al., 1985 b)

Cryptochironomus Cladopelma (BHATTACHARYA et al., 1985 b)

Dicrotendipes Gillotia (BHATTACHARYA et al., 1985 b)

Glyptotendipes Kiefferulus (CHAUDHURI & GHOSH, 1986)

Harnischia

Lauterborniella

Nilodorum

Paratendipes

Polypedilum

Stenochironomus

TANYTARSINI

Micropsectra Paratanytarsus (CHAUDHURI et al., 1984)

Tanytarsus (SINGH & KULSHRESTA, 1975)

Taxonomic placement

It has been established that pupal exuviae of most chironomids are easily placed at the generic level.

There are, however, a few striking exceptions; for example, Baetotendipes, Chironomus and Einfeldia

species group C (Pınner and Reıss, 1986). For this reason, as well as others, the list of chironomid ge-

nera given here should be considered to represent a minimum number of genera actually present. A

further complication in taxonomic placement was encountered frequently in this material. Many of

the species placed in named genera do not conform exactly to all aspects of the pupal diagnoses of ge-

nera given in WIEDERHOLM (1986). When the differences were only slight, the species were placed in the

156

Fig. 1. Map of extreme southern India — numbers indicate areas in which collections were made: 1—2, Ootaca-

mund-Coimbatore Road; 3-4, Kodaikanal-Madurai Road; 5, Periyar; 6, Rajapalaıyam; 7, Cape Cormorin.

appropriate named genera. However, when the differences were considered to have a greater probabi-

lity of taxonomic significance, the species were either placed in a named genus, with the name given

in quotation marks in the list, or the species were considered to be different enough to warrant place-

ment in separate, numbered genera.

Collection sites

Most of the 25 collection sites were first to fourth order streams and were visited only once. however, a few lentic

systems (ponds and reservoirs) and one marine locality were also sampled. T'wo areas were collected relatively ex-

tensively: the Palnı Hills in the vicinity of Kodaikanal and the Nilgiri Hills in the vicinity of Ootacamund. Additio-

nal collections were made from a stream in the vicinity of Rajapalaiyam, Tamil Nadu, a stream near Periyar Lake,

Kerala and tidal rock pools at Cape Comorin (Fig. 1).

Results and Discussion

Among the approximately 150 species-level taxa represented in these collections, 55 named genera

(including Chironomini Genus D and Genus E of Pınper & Reıss [1986]) were identified (Table 2). All

but one of these (Sublettea) is known from the Palaearctic, and all buttwo oftheremaining genera (Pa-

ratrissocladins and Virgatanytarsus) are Holarctic in distribution. In addition to the 55 named genera

represented in these collections, another 20 “new” genera were recognized. These categories were

established for species-level taxa that could not be placed in any of the genera for which diagnoses are

given in WIEDERHOLM (1986) or other literature, even when the generic limits were reasonably expand-

ed to allow for some differences (Table 2).

Thirty-five of the named genera represented in the collections from South India are recorded here

for the first time from India-bringing the total number of genera with known distributions in India to

Varlabler2).:

RoBack & Corrman (in press) found that the fauna of high altitude regions of Nepal, with only four

possible endemic genera (all Tanytarsini and based on larvae), was overwhelmingly Holarctic in distri-

bution. The results of other workers in the northern regions of the Indian subcontinent have included

157

Table 2. Genera of Chironomidae collected as pupal exuviae from South India. *indicates new to India.

TANYPODINAE CHIRONOMINI (continued)

Ablabesmyia Glyptotendipes

Conchapelopia” Harnischia

Hayesomyia” Microtendipes“

Larsia“ Nilodorum

Nilotanypus” Parachironomus*

Paramerina Paracladopelma*

Procladius Paralauterborniella*

Tanypus “Paratendipes”

Zavrelimyia” Phaenopsectra*

Pentaneurini Genera 1-5 Polypedilum

ORTHOCLADIINAE Bora

Saetheria”

Bryophaenocladins Stenochironomus

Cardiocladius“

“Stictochironomus””

Clunio* S Chironomini Genus D*

Corynoneura® (PINDER & Reıss, 1986)

Cricotopus Chironomini Genus E*

Eukiefferiella (PINDER & Reıss, 1986)

Heleniella* Chironomini Genera 1-6

Krenosmittia”

Limnophyes TANYTARSINI

Nanocladius“ Cladotanytarsus”

Paracricotopus” Rheotanytarsus“

Parakiefferiella Stempellinella”

Parametriocnemus” Sublettea”

Paratrissocladius” Tanytarsus

Rheocricotopus” “Virgatanytarsus””

Rheosmittia” Zavrelia”

Thienemanniella” Tanytarsıni Genera 1-3

Orthocladiinae Genera 1-4 PSEUDOCHIRONOMINI

CHIRONOMINAE Psendochironomus”

CHIRONOMINI UNKNOWN CHIRONOMIDAE

Chironomus Taroa

Cladopelma Tazem2

Cryptochironomus

Cryptotendipes*

Dicrotendipes

only two endemic genera (Asclerina, Reiss [1968] and Neopodonomus, Chaudhuri & Ghosh (1981).

Neopodonomus has, however, been shown to be a synonym of Boreoheptagyia (ROBACK & COFFMAN,

in press). These results are, perhaps, not surprising since the areas that have been most extensively col-

lected (montane regions) are ecologically more closely related to the Palaearctic than they are to the

Oriental Region. The fauna of South India, as revealed by these collections, is clearly dominated by

genera that have widespread distributions, but there is also a major component of apparently “ende-

mic” genera. It is not clear whether these genera are limited to South India (perhaps including Sri

Lanka) or aremore widespread in the Oriental Region. Unfortunately, no comprehensive study ofthe

chironomid fauna of any other part of the Oriental Region has been carried out. The limited data that

do exist would seem to indicate that at least some part of this group of “endemic” genera may occur

in other parts of the Oriental Region. Thienemann, Johannsen, Zavrel and Lenz found a chironomid

158

fauna on the Greater Sunda Islands that may be similar to that of South India. The $. Indian species

that is here placed in “Stictochironomus” bears some resemblance to Stictotendipes Lenz (1937) and the

Rheotanytarsus anomalus group of Zavreı (1934) reported from Sumatra also occurs in the South In-

dian material. There are other known connections of the South Indian fauna with other parts of the

Oriental Region. The species of Sublettea from South India is very similar (perhaps identical) to that

found by Fittkau in SW China and reported by Pınner & Reiss (1986). Chironomidae Genus 2 (de-

scribed below), or something very similar, has recently been found in China and in collections from

Burma (E. J. Fittkau, personal communication).

Descriptions of pupae of two unusual taxa

Taxon 1

Pupa (Figs. 2-8):

Size demonstrating strong sexual dimorphism, total length of females 7.0 mm — 8.3 mm, x =

7.6 mm, n = 4; total length of males 5.3 mm — 6.5 mm, x = 5.4mm,n = 6.

Cephalothorax: antepronotum, ventral structures of thorax, cephalic region and leg sheaths yellow;

dorsal parts ofthorax and wing sheaths yellow-brown; Cephalıc area (Fig. 2): frontal apotome strong-

ly sclerotized; frontal setae absent; frontal apotome without tubercles or warts; ocular field with one

postorbital seta; Thorax (Fig. 3): antero-dorsal region of mesonotum with a field of very strong spine-

like processes; fields of rugulosity dorsal of the dorsocentral setae, on the prealar lobe and on a lobe-

like process posterior to the precorneal setae — the latter somewhat stronger than the other two fields;

all leg sheaths recurved under wing sheaths; wing sheaths without pearl rows or terminal protube-

rance; thoracic horn absent; Chaetotaxy of thorax: two narrowly separated MAps; one LAps located

on a low protuberance; four Dc setae, Dc2 (apparently) located much ventral of Dei, Dc3 and Dc4

forming a posterior pair; three Pc setae of varıable lengths, but Pc1 is usually shorter than the other

two; no Pa or Mn setae present.

Abdominal segments I- VIII (Figs. 4-8): mostly yellow-brown, but armature and adjacent areas

reddish-brown to black.

Shagreenation — tergites: tergite I without, tergites II and III with extensive fields of dense fine sha-

green, slightly larger toward the posterior margıns (Figs. 4-5), tergite IV with a narrow transverse an-

terior band and posterolateral areas of dense fine shagreen (Figs. 4 and 6), tergite V with two narrowly

separated anterior groups and a transverse posterior band of fine dense shagreen (Figs. 4 and 6), tergite

VI with most of the central area covered by fine, but less dense, shagreen, expanded laterally along the

posterior margin (Figs. 4 and 7), tergite VII with central area covered with weak shagreen (Fig. 4), ter-

gite VIII with a weak transverse anterior band or without shagreen (Figs. 4 and 8).

pleurites: pleurite I without, pleurites II and III with small anterior areas of fine shagreen, usually

much weaker on III (Figs. 4-5), pleurites IV and V with extremely fine shagreen on posterior halves

(Fig. 4), pleurites VI-VIII with extremely fine shagreen over much of surface, sometimes absent on

VII and much of VIII.

Sternites: sternites I- VI with at most very fine and very sparse shagreen spinules, sternite VII with

a weak anteromedian group of shagreen spinules, sternite VIII without shagreen.

Armature and other structures of abdomen: rows of approximately 50-60 Hl on posterior margıns

of tergites Il and III (Figs. 4-5); tergites III (II)—IV with transverse anterior rows of sharp dark poste-

riorly directed spines (Figs. 4-5), tergite IV with a posteromedian transverse group of sharp, dark,

anteriorly directed spines of varying lengths (Figs. 4 and 6); tergite V with a sinuous row of large,

dark, posteriorly directed spines (Figs. 4-6); tergite VI with a dense, anteromedian, semicircular

patch of spines (Figs. 4 and 7); tergites VII and VIII without armature; PSB absent on II, but a PSB-

like structure present on segment III (Figs. 4-5); PSA absent on all segments.

199

anni,

oo?

= os oO?

Figs. 2-6. Taxon 1, pupa, — 2. cephalic region, — 3. thorax, lateral, — 4. abdomen, dorsal, — 5. segment 3, dorsal,

— 6. posterior armature of tergite 4 and anterior armature of tergite 5.

160

Figs. 7-10. Taxon 1, pupa. — 7. Anteromedian armature of tergite 6, — 8. posterior margin of segment 8 and anal

lobes, dorsal; Taxon 2, pupa. — 9. cephalic area, — 10. thorax and adjacent structures of cephalic region, lateral.

Chaetotaxy — tergites (Figs. 4-5 and 8): tergite I with 4 D setae; tergites II-V with 7 D setae; ter-

gites VI and VII apparently with 6 D setae; tergite VIII with 1 D seta.

Pleurites: pleurite I with 1 L seta; pleurites II-VII with 3 L setae; pleurite VIII with one relatively

large posterior L seta (Figs. 4-5 and 8).

Sternites: sternite I with V setae apparently absent; sternites II-VII with 4 V setae; sternite VIII

with 2 V setae.

Anal lobes and segment IX (Figs. 4 and 8): tergite IX with lateral and median fields of shagreen; AL

without marginal fringe of setae, but with several rows of small spine-like protuberances; each AL

with a dorsal and ventral seta on the disc; inner margins of AL with small groups of weak spinules;

genital sheaths of male extend to tip of AL.

Specimens examined:

Kodaikanal area, Palni Hills, Tamil Nadu, South India — small stream along Madurai — Kodaikanal Road, at

milepost 36, 1 female pupal exuviae, 18 Mar. 1978; 3rd order stream along Madurai — Kodaikanal Road, about

161

4 km east of Kodaikanal, Tiger Forest stream, 2 female pupal exuviae, 20 Mar. 1978; 2nd order stream along Madu-

rai — Kodaikanal Road, between mileposts 17/5 and 17/6, 1 male and 1 female pupal exuviae, 18 Mar. 1978; Oota-

camund area, Nilgiri Hills, Tamil Nadu, South India — Vellappalam stream (2nd or 3rd order) along Coimbatore

— Ootacamund Road, 1 male pupal exuviae (fragment), 5 May 1979; 2nd order stream along Coimbatore — Oota-

camund Road, 3 male pupal exuviae, 8 May 1979; Kallar River (4th order stream) at base of Nilgiri Hills along Co-

imbatore — Ootacamund Road, 2 male pupal exuviae, 8 May 1979.

Taxon 2

Pupa (Figs. 9-15):

Moderate sexual dimorphism in size, total length of females 3.0 mm — 3.5 mm, x = 3.2 mm, n =

6: total length of males 2.5 mm — 3.25 mm, x = 2.9 mm, n = 12.

Cephalothorax: All thoracic structures yellow to yellow-brown; Cephalic area (Fig. 9); frontal

apotome strongly sclerotized; frontal setae absent; frontal apotome without tubercles or warts; ocular

field with one postorbital seta; Thorax (Fig. 10): dorsal surface of thorax not rugulose, but with very

weak sculpturing; all leg sheaths recurved under wing sheaths; wing sheaths without pearl rows or ter-

minal protuberance, but with an angular process on the inner margin near the base, adjacent to and fit-

ting closely against the prealar lobe; thoracic horn absent; Chaetotaxy ofthorax: two widely separated

MAps; one LAps located on a low protuberance; four Dc setae, Dc2 or Dei located more ventrally

than others, Dc3 and Dc4 forming a posterior pair; three relatively strong Pc setae of approximately

equal size; no Pa or Mn setae present.

Abdominal segments I- VIII (Figs. 11-15): tergites light reddish-brown, armature of tergites dark

reddish-brown; tergites II-VII with lateral, longitudinal dark brown lines; pleurites and sternites

with little or no pigmentation.

Shagreenation — tergites: tergites II-VIII with lateral and/or anterior groups of very weak sha-

green spinules (Figs. 11-15).

Pleurites: all pleurites without shagreen.

Sternites: sternites II-VIII with lateral, moderately dense groups of very fine shagreen spinules in

irregular arching rows.

Armature and other structures of abdomen: posterior margins of tergites II and III with rows of ab-

out 25-30 strong Hl (Figs. 11-12); tergites III-V with anterior rows of strong Hl, 25—30 on tergite

III, about 18 on tergite IV and about 12 on tergite V (Figs. 11-13); tergite IV with a posterior median

group of anteriorly directed spines, the median spine in this group much larger than the others

(Figs. 11 and 13); tergite VI with a pair of very large anteromedian Hl (Fig. 11); tergites VII and VIII

without armature; PSB and PSA absent on all segments.

Chaetotaxy — tergites (Figs. 11-13): tergite I with 4 D setae; tergites II- VII with 7 D setae; tergite

VIII with 2 D setae.

pleurites: pleurite I with 1 L seta; pleurites II-VII with 3 L setae; tergite VIII with 1-2 L seta.

sternites: sternite I without V setae; sternites II-VII with 4 V setae; sternite VIII with 1—2 V setae.

Anal lobes and segment IX (Figs. 11, 14 and 15): tergite IX without shagreen; AL without marginal

fringe, but margin weakly crenulate; each AL with a dorsal and ventral seta on the disc; AL and seg-

ment IX forming a Telmatogetoninae-like concave, oval plate; the posterior margin of tergite VIII

heavily sclerotized and crenulate, forming an arching ridge above the AL.

Specimens examined:

Ootacamund area, Nilgiri Hills, Tamil Nadu, South India — Kallar River (4th order), at base of Nilgiri Hills

along Coimbatore — Ootacamund Road, 16 male and 10 female pupal exuviae, 8 May 1979.

162

\ AI NNUR 2

= ee

Figs. 11-15. Taxon 2, pupa. — 11. abdomen, dorsal, — 12. segment 3, dorsal, — 13. segment 4, dorsal, — 14. seg-

ment 8 and anal lobes, dorsal, — 15. posterior margin of segment 8 and anal lobes, lateral.

163

Comments on taxa 1and 2

Although the pupal exuviae of these unusual taxa differ from each other in size and many structural

features, they have a number of basic similarities that, most probably, indicate a close relationship.

Among these features are: 1). the absence ofa TH; 2). the absence of FS; 3). the ventral position of one

of the anterior De setae; 4). the unusual position of the LAps on a low protuberance; 5). the absence

of LS on all abdominal segments; 6). the presence of only one L seta on segment VIII; 7). the presence

of seven D setae on most tergites; 8). the absence of an AL fringe; 9). the presence of dorsal and ventral

setae on the disc ofthe AL; and 10). the presence of unusual armature on the abdominal tergites.

Each of these taxa undoubtedly represents anew genus and, since the combination of characters that

they possess is not consistent with any described subfamily, they most likely represent a new higher

taxon as well, perhaps a new subfamily. The number and distribution of thethoracic setae ofthese taxa

would seem to indicate a relationship with Orthocladiinae. However, the abdomen of Taxon 1 shows

a number of relationships with Pseudochironomus, e. g., the shape of the AL and the presence of a

nearly circular patch of spines on tergite VI. Some species of Pseudochironomus also have no fringe on

the AL. The abdomen of Taxon 2 does not, however, show this possible relationship. Instead, the

unusual arrangement of segment VIll and IX, with the AL, appears to represent a remarkable conver-

gence with Telmatogetoninae. Clarification of the taxonomic position and phylogenetic relationships

of these taxa must await discovery of the larvae and/or adults.

Acknowledgements

This work was supported by Smithsonian Institution Grant FC-70117000 to Dr. George Saunders, whose en-

couragement and interest are gratefully acknowledged. We are indebted to Professor $S. Krishnaswamy, Dr. George

Michael and Mr. Navaneethakrishnan for their considerable logistic help in making the field work possible. Appre-

ciation is expressed to numerous postdoctoral and graduate students at Madurai Kamaraj University for field assi-

stance. Lastly, thanks are given to Stephan Coffman, whose enthusiasm and companionship aided greatly.

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—— 1982b: Record of Chaetocladius Kieffer (Diptera: Chironomidae) from India. — Folia entomologica Hunga-

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scand. Suppl. 28: 1482

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Dr. William P. Coffman,

L. A. Yurasiıts, Jr.

Dr. Carlos de la Rosa,

Department of Biological Sciences,

University of Pittsburgh

Pittsburgh, PA 15260, USA

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SPIXIANA Supplement 14 167—174 München, 15. Juli 1988 | ISSN 0177-7424

A survey of the zinc-polluted River Nent (Cumbria) and the

East and West Allen (Northumberland), England, using

chironomid pupal exuviae

By RR.S. Wilson

Abstract

Collections of chironomid pupal exuviae from zinc-enriched sites on the Rivers Nent, and East and West Allen

in the English Pennines, showed consistently lower diversities than those from low-zinc sites. The dominant species

found at zinc-enriched sites differ between the Nent and the Allens, being Krenosmittia camptophleps in the Nent,

and Eukiefferiella clypeata and Tvetenia calvescens in the West Allen. The low-zinc sites may contain chironomid

communities close to those which would live in normal unpolluted rivers of similar types to those investigated.

These communities will vary as conditions change between different sites or rivers, but will form the basic set of

species from which a zinc-tolerant sub-set will be selected in each case.

Introduction

The area of the North Pennine orefield in Britain, is notable for the extensive mining operations that

were carried out principally during the eighteenth century (Dunham, 1949). Currently there is very

little mining activity in the area, but previous workings and accumulations of mine spoil have left aleg-

acy of acıd and heavy metal enriched streams. The Rivers Nent and the East and West Allen, rise in

the high moors near Alston, at the intersection of the three counties, Cumbria, Durham and North-

umberland. This paper reports an investigation carried out in 1983, into the chironomid fauna of the

zinc-enriched Nent and the Allens, using collections of chironomid pupal exuviae. It was hoped to

gain some understanding of the distribution of the chironomid fauna in relation to the stream condi-

tions, and ın particular to the concentration of zinc in the water.

The rivers are affected by acid mine drainage from old mine workings, and are particularly rich in

zinc. Certain locations also receive organic effluents of domestic and agricultural origin. ArMmITAGE

(1980), studied the macroinvertebrates of an extensive series of sites on the Nent and its tributaries,

and has related their distribution and abundance to broad categories of zinc concentrations. He also

gives figures for certain chemical measurements related to organic and metal concentrations in the wa-

ter. ABEL & GREEN (1981) have reported on the ecology and distribution of macroinvertebrates on the

Allens in relation to the zinc concentrations, which are significantly higher in the West, as opposed to

the East, Allen.

In the Nent, ArnıtAce (1980) showed that there was a significant negative correlation between the

zinc level and the number of taxa per site, although the situation was complicated by localised inflows

of calcium-rich water. Summer growths of Stigeoclonium tenue may also have affected the faunal dis-

tribution. AsEL & GREEN (1981) showed that the fauna in the zinc-enriched West Allen was quantitati-

vely and qualitatively restricted as compared to that in the East Allen, and suggest that this is directly

due to the toxic effects of the zinc. They also showed a significant negative correlation between num-

bers of species of macroinvertebrate and the concentration of zinc in the water.

167

Haltwhistle

10 km

Fig. 1: Diagrammatic sketch-map of the section of the River South Tyne and its tributaries the River Nent and

the East and West Allen. Sampling sites are shown in circles.

Other work on the effects of heavy metals on river invertebrates includes the classical studies on the

Rivers Ystwyth and Rheidol in North Wales (see eg. CarrENTER, 1924; Jones, 1940), and on the Willow

Brook by Sors£ (1977), while a general discussion may be found in Wnrrton (1975). Little of the data

refers in detail to the Chironomidae, however, and the work reported in this paper is an attempt to re-

medy this omission by examining data from a series of collections of chironomid pupal exuviae made

in July 1983 in the same area as worked by Armırace (1980), and by Assı & Green (1981).

Locality and Methods

The River Nent rises south-west of Nenthead (NGR NY 801421) at an altitude of 656 m and flows north-west-

ward into the River South Tyne at Alston after a course of only 16 km. The East and West Allens are a pair of very

Table 1: Sampling sites and physical and chemical data.

Chemical ranges* -

Site River and site National Altitude Gradient Width PH ca zn Pb

code Grid Ref. m o/coo range mg/l mg/l mg/l

Nl Nent, u/s Dowgang Level NY782434 433 33 4-5 6-7 5-10 0.5-1 <0.05

N2 Nent, d/s Dowgang Level NY781435 432 33 4-5 7-8 10-20 1-2 <0.05

DL Dowgang Level adit NY782435 433 - 1 7-8 50-100 5-10 <0.05

DB Dowgang Burn NY780435 428 125 2-3 3-4 5-10 0.1-0.5 0.1-0.2

FB Foreshield Burn NY750467 338 50 2-3 7-8 20-50 <o.1 <0.05

ST South Tyne, Bardon Mill NY781643 90 3 80-100 7-8 10-20 <o.1 <0.05

WAL West Allen, Corryhill Br. NY782524 245 16 10-20 7-8 20-50 1-2 trace

WA2 West Allen, Whitfield Weir NY782570 175 14 10-20 7-8 20-50 1-2 trace

EAl East Allen, Rye Close Ford NY842509 294 11 10-20 7-8 20-50 <0.2 trace

EA2 East Allen, Thornley Gate Br. NY831567 193 10 10-20 7-8 20-50 <0.2 trace

* data modified from ARMITAGE (1980), ABEL & GREEN (1981)

168

Table 2: List of samples from the River Nent, West and East Allen,

and South Tyne, showing numbers of exuviae and taxa, and certain

exuvial and diversity indices.

Percentages to the nearest

whole percent. KEY: Nl and N2, Nent; DL, Dowgang Level adit;

DB, Dowgang Burn; FB, Foreshield Burn; ST, South Tyne;

WA, West Allen; EA, East Allen; %It, %Intolerant taxa;

%Ii, % Intolerant individuals; %Sedt, % Sediment-dwelling taxa;

%Sedi, % Sediment-dwelling individuals; Men D, Menhinick

Diversity; S-W D, Shannon-Weaver Diversity.

No. of ko. of

Station exuviae taxa

Nl 303 13

N2 477 18

DL 1 1

DB 51 4

FB 118 23

ST 225 3l

WA 45 13

EA 315 37

ITaxa indices

%It %Sedt

69 3l

56 39

(0) 25

52 17

71 45

69 15

62 22

Men

Exuvial indices

sw ı

$3Ii Sei D I

6 9%

ES,

= zZ ı

2 an

epras!

TOR 2!

24 2.6 |

similar rivers which rise im mediately east of the Nent (NGR: NY802442, NY860432), and flow northwards paral-

lelto each other for 30—40 km, before joining and entering the South Tyne approximately 33 km downstream from

the Nent junction.

Fig. 1 is a diagrammatic sketch-map of the area, showing the rivers and the sampling stations. The two sites on

the Nent were both near the village of Nenthead, N1 approximately 200 m above and N2 300 m below the zinc-rich

output from an old mine adit, the Dowgang Level; DL was on the outflow from the Dowgang Level, which flows

for only about 3 m before entering the Nent; DB was on the Dowgang Burn tributary which is highly acidic

(pH c.4) with effluent from an old coal mine; FB was on the Foreshield Burn tributary, which is organically en-

riched from a local farm but has few heavy metals; and ST was on the South Tyne. Two samples were taken from

both the East (EA1, EA2) and West Allen (WA1, WA2).

taxa

exuviae

—

0 50

% similarity

100

N2 FB DB ST WA

% similarity

Fig. 2: Dendrograms of percentage similarity derived from average linkage clustering of Sorensen’s and Czeka-

nowski’s coefficients of similarity; A. taxa presence/absence; B. number of exuviae. The matrices of similarıty from

which the dendrograms are derived are also shown.

169

Sets of exuvial collections were made at each site over atwo day period (18th and 19th July 1983) by both hand-

net and surface drift-nets (similar to Brundin nets but with lateral floats, BRUNDIN, 1956; WıLson & MCGiLL,

1977), and all collections were lumped for each site. On the Nent and its tributaries, small drift-nets were set out

overnight so as to increase the numbers of exuviae collected.

Table 3. List of species and percentage abundance found in chironcmid

exuvial collections fram rivers in the North Pennine orefield in July 1983.

Figures are whole percentages; + = <0.5%; . = absent from sample.

KEY TO SITES: N1,N2, Nent; DL, Dowgang Level adit; DB, Dowgang Burn;

FB, Foreshield Burn; ST, South Tyne; WA, West Allen; EA, East Allen.

Percentages (nearest whole percent) -

TAXA Nl N2 DL DB FB ST WA EA TOTAL

BUCHINOMYINAE

Buchincmyia thienemanni Fitt . . . ® En an 5

PODONOMINAE

Paraboreochlus minutissimus (Strobl) 1 1 . E o 2 ® & +

TANYPODINAE

Macropelopia nebulosa (Mg) 5 5 © o a, ıl

Ablabesmyia longistyla Fitt. Bach 02: - - ö > +

Conchapelopia viator (K) a ae Be 3

Rheopelopia maculipennis (Zett) - e 5 5 il +

Thienemannymia laeta (Mg) Pel/2 Se or er er +

Trissopelopia longimana (Staeg) Rees TR +

DIAMESINAE

Diamesa insignipes K Ste: Ist uno 0 +

Diamesa thienemanni K ö o ® - & + +

Potthastia Pela A . . & = . 5 +

Potthastia longimana K = . . & & + - 5 +

Potthastia sp.2 . B . 5 de 6 1

PRODIAMESINAE

Prodiamesa olivacea (Mg) 5 E ® = ° - ser! +

ORTHOCLADIINAE

Brillia modesta (Mg) a 3 . 6 6 h 5 2 1

Cardiocladius fuscus s o . > - R - + +

Chaetocladius Pe2 FL SRE21007 05T R E P +

Corynoneura spp. . . . . 4 . . +

Cricotopus (C) Pe2 ?similis Goet Aue A a a TE N DS +

Cricotopus (C) bicinctus (Mg) - - b c - e SIE +

Cricotopus (C) pulchripes Verr 4.107 == > Ze > Ö 4

Cricotopus (C) spp. oft Orig oki mer u In KALT. +

Cricotopus (C) tremulus (L) Se ed +

Cricotopus (C) trifascia Edw ER 5 Se SARER 5 ° +

Cricotopus (I) brevipalpis (K) a: Q we] 2 1

Eukiefferiella brevicalcar (K) . . - : : ö BD +

Eukiefferiella claripennis (Lund) Ha 02022 30, 0 12 4

Eukiefferiella clypeata (K) NS 5 - - Bo |

Eukiefferiella coerulescens (K) Hu Fre B ee | l

Eukiefferiella devonica (Edw) ER: s - SER EEG) EL 2

Eukiefferiella ilkleyensis (Edw) ae u a ee 23 1

Eukiefferiella minor (Edw) ö 2 0 . 3 . .. + +

Heleniella omaticollis (Edw) 3... e 5 1 R o o 3

Krenosmittia camptophleps (Edw) EEE ee ot 36

Limnophyes spp. 5 5 92 c B Gr 3

Metriocnemus hygropetricus (K) gp. SR + +

Nanocladius bicolor (Zett) er: Q De +

Nanocladius rectinervis (K) B B o ee: £ +

Orthocladius (Eudact) obtexens Brundd . + . 1 + +

Orthocladius (Euorth) rivicola K es 9 2

Orthocladius ?rubicundus Edw . . & . il +

Orthocladius Pel . o © . e + +

Orthocladius rhyacobius K B . . 5 Tl! B +

Orthocladius rubicundus sensu Edward . . Au 6 2

Orthocladius sp.A Pinder - 1 Br 3r

Paracladius conversus (Walk) + x e R Ö : +

Parametriocnemus stylatus (K) 5 . £ 2 10 Sa +

Paratrichocladius rufiventris (Mg) 5 e S e 2 +

Paratrissocladius excerptus Pel nt +

Pseudorthocladius sp. . 4 . > - B e a +

Pseudosmittia Pel b + 5 5 R = E + +

170

Rheocricotopus chalybeatus (Edw) R 3 0 0 1 2

Rheocricotopus dispar (Goet) Se: . o .

Rheocricotopus effusus (Walk) Sn 0.02

Thienemannia gracilis K ö ö ö ö ö ö ö

Tvetenia calvescens (Edw) en 0 SO AEO Er?!

Tvetenia discoloripes (Goet) ae 0 0 5 EA

CHIRONOMINAE

CHIRONOMINI

Microtendipes spp. . . o ö gl .

Paratendipes Pe2 . . . . . . Der

Polypedilum Pe2 . ö S . 1 es 23 ee

Polypedilum albicorne (Mg) A RO ne] :

Polypedilum cultellatum Goet ö ö o © o 0 o

Polypedilum laetum (Mg) BETT ET Et

TANYTARSINI

Cladotanytarsus ?nigrovittatus Goet . . . . . 1

Micropsectra aristata Pind He 2 OMA

Micropsectra atrofasciata K . ° o . he a cn

Micropsectra apposita (Walk) SER sh o..abl

Parapsectra nana (Mg) gt

Rheotanytarsus pentapoda K +

Tanytarsus brundini Lind pw © Ron

Tanytarsus eminulus (Walk) 2 N ee PAR

Tanytarsus palmeni Lind 4

wWO++Hr

++Pr++u Ho+n++

H+rt+tt+toHret+

TOTAL TAXA IN SAMPLE 13 18 1 4 23 31 13 37 72

TOTAL EXUVIAE IN SAMPLE 303 477 1 51 118 225 45 315 1535

TOTAL PERCENTAGE (nearest whole $) 99 102 100 % 104 1009 % 96

Results

Most of the samples contained satisfactory numbers of exuviae, but some were numerically poor.

The Dowgang Level mine adit (DL) yielded only a single exuviae, and the acıd Dowgang Burn (DB)

only 51 exuviae in 24 h drift-netting. The Allens were sampled with hand and drift nets for about an

hour at each site, and WA1 and WA2 on the West Allen, yielded only 4 and 41 exuviae respectively;

whereas the two East Allen samples EA1 and EA2 on the other hand yielded 65 and 250 exuviae. It

was therefore decided to lump EA1 and EA2, and WA1l and WA2, together for analysis, forming a

single composite sample for each of the East Allen (EA) and the West Allen (WA).

Table 1 gives some basic parameters such as altitude, slope of site, and grid reference for each site,

as well as chemical data derived from the papers by Arnırace (1980) and Ageı & Green (1981) which

illustrate the chemical conditions. Table 2 shows the numbers of exuviae and identified taxa found in

each sample together with certain exuvial and diversity indices. Table 3 gives the complete species list

found in the samples in taxonomic order, while Table 4 lists only those species which were found mak-

ing up 5% or more of the exuviae in any one sample.

Fig. 2 shows the dendrograms plotted from data from average linkage clustering of the species col-

lected from each site, using Sorensen’s and Czekanowski’s coefficients of similarity for taxa presence/

absence and numbers of individual exuviae respectively.

Discussion

The cluster analysis data (see Fig. 2) show that the two Nent sites (N1, N2) are very similar ın both

taxa and numbers of exuviae (N1:N2 taxa, 58%; exuviae, 76%). They are also closely linked to the

Foreshield Burn (FB) by taxa presence/absence (N1:N2, 58% ; N1:FB, 56 % ; N2:FB, 49%), but not

by numbers of individual exuviae (N1:FB, 8%; N2:FB, 8%). Similarly the West Allen and the East

Allen are similar in taxa (WA:EA, 48%) but not in numbers of exuviae (WA:EA, 17 %). The zinc-en-

riched sites also show a lower diversity of species (N1: Menhinick, 0.7; Shannon, 0.9; N2: Menhinick,

0.8; Shannon, 1.3) in contrast to the higher diversity in the low-zinc sites (FB: Menhinick, 2.1, Shan-

non, 2.4; EA: Menhinick, 2.1; Shannon, 2.6).

121

Table 4: Table of principal species (>5% in any sample) in the

Nent, East and West Allen, and South Tyne. Figures to the

nearest whole percent. KEY: Nl and N2, Nent; DB, Dowgang Burn;

FB, Foreshield Burn; ST, South Tyne; WA, West Allen; EA, East Allen;

+ = <0.5%; . = absent from sample.

Percentäages -—

Species N N2 DB FB ST WA _EA

Krenosmittia camptophleps (Edw) TSF TESgLzE. ES nr

Heleniella ornata (Edw) Din Korte ul nr

Cricotopus (C) pulchripes Verr A Or e E

Polypedilum albicorne (Mg) AN PS EL, WET Me

Limnophyes spp. - a. ° < ee

Eukiefferiella claripennis (Lund) E24 1 02,730, AZ

Tvetenia calvescens (Edw) dus; 15,200547395529

Micropsectra aristata Pinder oe te 2 7719,54, 9,214

Brillia modesta (Mg) ee STTHoweee en

Conchapelopia viator (K) Fr ea Zu

Microtendipes spp. . . ° 31 . .

Micropsectra apposita (Walk) “= ee) ae,

Eukiefferiella clypeata (K) 0 eye einen Cu

Eukiefferiella devonica (Edw) - - N et PR

Orthocladius (ss) rubicundus Edw = - B sa. wrTaide

Orthocladius (Euorth) rivicola K > e . - ® Be)

Total taxa in sample 33,218, 4.237 7312137237,

Total exuviae examined 303 477 51 118 225 45 315

If the taxa composition is similar between comparable zinc-enriched and low-zinc sites, but the

abundances of individual species differs, and the diversity is lower at the enriched sites, then it seems

possible that the fauna of the zinc-enriched sites (N1, N2 and WA), comprise those species that have

higher tolerance of the zinc, and represent subsets of the unpolluted chironomid communities of these

rivers. The low-zinc sites (FB and EA) may exemplify the unpolluted fauna.

It is important, however, to note that the species found in the zinc-enriched Nent were very dif-

ferent from those in the similarly zinc-enriched West Allen. The Nent samples (N1 and N2) were

dominated by Krenosmittia camptophleps (Edwards) (78% and 65 %), but also contained substantial

numbers of Heleniella ornata (Edwards) (5% and 6%), Cricotopus (C.) pulchripes Verrall (4% and

10%) and Polypedilum albicorne (Meigen) (4% and 9%): in contrast, the West Allen samples (WA)

were dominated by Eukiefferiella clypeata (Kieffer) (36%) and Eukiefferiella claripennis (Lundbeck)

(13%), and did not contain any of the four species characteristic of the Nent.

It appears therefore that the Nent and the Allens, although adjacent and similar stony rivers, differ

in some way meaningful to the Chironomidae. It is therefore not possible from these samples to de-

signate a characteristic “zinc-tolerant” community of chironomid species, which might be found

wherever zinc-enrichment occurs, in the same way thata characteristic “organic-tolerant” community

may be defined which is linked to organic enrichment in many different rivers and situations. Zinc-

tolerance may develop in anumber of different species, selected as appropriate from those available at

the particular site.

It is well-known that chironomid communities differ in different sections of a single river, and that

different communities are found associated with different gradients and substrates (THIENEMANN,

1954; Kownackı & Kownacka, 1972). The Nent sites N1, N2 and FS have gradients of from 30—50%,

whereas the sites on the Allens have gradients of from 10— 16%. It is possible therefore that the chi-

ronomid communities found in the Nent are characteristic of higher gradients than those of the Allens,

even though the rivers are basically similar.

The dominant species in the Nent in samples N1 and N2, Krenosmittia camptophleps, Heleniella or-

nata, Cricotopus (C.) pulchripes and Polypedilum albicorne are all characteristic of upland streams and

often associated with hygropetric conditions (CRAnsToN, 1982; WIEDERHOLM, 1983; LAnGTon, 1984). In

contrast, Eukiefferiella and Tvetenia spp., which dominate the samples from the West Allen, are nor-

172

Pe u

mally more abundant at lower altitudes, and especially where there is increased growth of moss (Hun-

PHRIES & Frost, 1937; THIENEMANN, 1954; CrAnsToN, 1982; LAnGTon, 1984).

The calcium concentrations in the the Nent were lower than in the Allens (see Table 1). It is widely

held that heavy metals are less toxic in calcium-rich water than in acid, low calcium water (Hynes,

1960; WARREN, 1971; Wurtton & Say, 1975; Mason, 1981), and this effect may influence the response

of the chironomids to the river conditions.

The sample from the acid Dowgang Burn (c. pH 4) is quite different from all the other samples, and

is dominated by Limnophyes spp. (92%). The single exuviae from the very short outfall from the

Dowgang Level, which has a high zinc content of 5-10 mg/l, was a Chaetocladius sp. (Pe2 in Lanc-

ton, 1984), also found in low numbers in the Nent and the Foreshield Burn. The River South Tyne at

Bardon Mill Ford is amuch larger river with more fine sediment, and has amuch lower gradient (3%)

than the Nent and Allens, and this is reflected in the dominance of sediment-living taxa such as Micro-

tendipes spp. and Micropsectra apposita (Walker) found in the sample (31% and 11 %, respectively).

In the cluster analysis, the South Tyne sample (ST) showed closest similarity with the Foreshield Burn

(ST:FB taxa, 30 % ; exuviae, 16%) and with the East Allen (ST:EA taxa, 32 % ; exuviae, 27 %), but the

linkage is weak. As the South Tyne has a low zinc content (<0.1 mg/l), it it might be expected to link

with the other low-zinc sites.

More data is required before the ideas advanced in this paper can be considered valid, but it is hoped

that they will point the way to further, more detailed, work using chironomid pupal exuviae on the

effects of heavy metals on chironomid communities.

Acknowledgements

I am grateful to Dr. P. D. Armitage, Dr. B. A. Whitton and Dr. P. J. Say for discussion and encouragement, and

to Mrs. S. E. Wilson for discussion and invaluable help with the samplıng.

References

ABEL,P.D.& D. W.]. GREEN 1980: Ecological and toxicological studies on invertebrate fauna of two rivers in the

Northern Pennine Orefield. — In: Heavy Metals in Northern England: Environmental and Biological

Aspecıs. Eds: P. J. Say and B. A. Whitton. University of Durham, Department of Botany

ARMITAGE, P. D. 1980: The effects of mine drainage and organic enrichment on benthos in the River Nent system,

Northern Pennines. — Hydrobiologia 74: 119—128

BRUNDIN, L. 1956: Transantarctic relationships and their significance, evidenced by chironomid midges, with a

monograph of the subfamilies Podonominae and Aphroteniinae and the austral Heptagyiae. — Kgl. Sven.

Vetenskapsakad Handl. 11: 1-472

CARPENTER, K. E. 1924: A study of the fauna of rivers polluted by lead mining in the Aberystwyth district of

Cardiganshire. — Ann. appl. Biol. 11: 1-23

CRANSTON, P. S. 1982: A key to the larvae of the British orthocladiinae (Chironomidae). — Freshwater Biological

Association, Scientific Publication No. 45

DunHam, K. 1949: Geology of the Northern Pennine Orefield Vol. I. Tyne to Stainmore. — Mem. Geol. Surv. 6:

1-357

HUMPHRIES, C. F. & W. E. Frost 1937: River Liffey survey. The chironomid fauna of the submerged mosses. —

Proc. R. Irısh Acad. 18: 161-181

Jones, J. R. E. 1940: A study of the zinc-polluted river Ystwyth in North Cardiganshire, Wales. — Ann. appl.

Biol. 27: 368—378

Hynes, H. B. N. 1960: The biology of polluted waters. — Liverpool University Press

Kownacka,M. & A. Kownackı, 1972: Vertical distribution of zoocoenoses in the streams of the Tatra, Caucasus

and Balkan Mts. — Verh. Int. Ver. Limnol. 18: 742—750

Langrton, P.H. 1984: A Key to pupal exuviae of British Chironomidae. — Private publication

173

Mason, C. F. 1981: Biology of freshwater pollution. — Longman, London

SOLBE, ]J. F. DE L. G. 1977: Water quality, fish and invertebrates in a zinc-polluted stream. — In: Biological Moni-

toring of Inland Fisheries, Editor J. S. Alabaster, Applied Science Publishers, London

THIENEMANN, A. 1954: Chironomus, Leben, Verbreitung und wirtschaftliche Bedeutung der Chironomiden. —

Binnengewässer, XX. Schweizerbartsche Verlagshandlung, Stuttgart

WARREN, C. E. 1971: Biology and water pollution control. — W. B. Saunders Co., Philadelphia

WHITTON, B. A. & P. J. Say 1975: Heavy Metals. — In: River Ecology, Editor B. A. Whitton, Studies in Ecology

Vol. 2., University of California Press, Berkeley

Wırson, R. $. & J. D. McGıtr 1977: A new method of monitoring water quality in a stream receiving sewage

effluent, using chironomid pupal exuviae. — Wat. Research., 11: 959—962

WIEDERHOLM, T. (Ed.) 1983: Chironomidae of the Holarctic Region. Keys and diagnoses. Part 1. Larvae. — Ent.

Scand. Suppl. No. 19

Dr.R.S. Wilson,

Department of Zoology,

University of Bristol, Bristol BS8 1UG,

Great Britain

174

SPIXIANA Supplement 14 VS München, 15. Juli 1988 ISSN 0177-7424

Zwei neue Smittia-Arten aus dem süddeutschen Raum

(Diptera, Chironomidae)*)

VonN. Caspers

Abstract

Two new species of the genus Smittia Holmgren are described from southern Germany. The description of Smit-

tıa scutellosetosa n. sp. is based on several male (and one female) imagines, that of Smittia amoena n. sp. on one male

only. Both species are characterized by the peculiar shape of the male genitalia. Larvae and pupae are unknown.

Most probably the larvae settle in dry, gravelly sediments of river banks in southern Germany.

Einleitung

Die weltweit verbreitete Gattung Smittia Holmgren umfaßt ausschließlich Arten mit terrestrischen

bzw. hygrophilen Larvenstadien. Die Gattungszugehörigkeit der Arten im Imaginalstadium ist leicht

am Besitz einer kräftigen Subapikalborste auf dem Antennenendglied der Männchen erkennbar. Le-

diglich die nearktische Saetheriella amplicristata Halvorsen besitzt ebenfalls eine Subapikalborste auf

dem Antennenendglied; die Analspitze ist jedoch bei dieser Art im Gegensatz zu den Arten des Genus

Smittia kurz und breit (Harvorsen 1982). Die infragnerische Gliederung des Taxons Smittia bereitet

zur Zeit noch außerordentliche Schwierigkeiten. Die enorm hohe Artenzahl, die oft nur subtilen Un-

terschiede in der Genitalmorphologie der Männchen, vielleicht auch die etwas ungewöhnlichen Habi-

tate vieler Smittza-Arten vereitelten bis zum heutigen Tage die Durchführung einer dringend benötig-

ten Revision des Gattungskomplexes. Die Abbildungen und Beschreibungen in der älteren Literatur

— insbesondere bei GOETGHEBUER (1940-1950) — sind aus heutiger Sicht völlig unzureichend und

nicht für eine gültige Differenzierung der Arten verwendbar. Zur Zeit sollte eine Beschreibung neuer

Smittia-Arten sinnvollerweise nur dann vorgenommen werden, wenn eindeutige und auffällige Merk-

male ihre Eigenständigkeit herausstellen. In der vorliegenden Arbeit werden zwei derartige Smittia-

Arten mit besonders markantem Hypopygbau beschrieben.

Smittia scutellosetosa spec. nov.

(Abb. 1-5)

Fundort: Isarinsel bei Aumühle (Nähe Schäftlarn in Oberbayern, Bundesrepublik Deutschland).

Material: Holotyp ©’, Barberfalle, 26.5.1982, leg. M. Baehr; in coll. Zoologische Staatssammlung München

(Objektträgerpräparat gemäß SCHLEE 1966). Paratypen: 3 0'C', mit gleichen Funddaten wie der Holotyp; 6 O’C',

Aumühle, Barberfalle, 13.5.1983, leg. M. Baehr (in sehr schlechtem Erhaltungszustand); 1 0°, Gerolfing (Nähe In-

golstadt, Bundesrepublik Deutschland), Barberfalle, 3.5.1983, leg. M. Baehr; 1 0°, Weichering (Nähe Ingolstadt),

Barberfalle, 6.5.1982, leg. M. Baehr.

Alle Paratypen werden in der Zoologischen Staatssammlung München aufbewahrt.

*) mit finanzieller Unterstützung der Deutschen Forschungsgemeinschaft

175

Diagnose: Die neue Art ist insbesondere durch genitalmorphologische Merkmale hinreichend ge-

kennzeichnet: extrem lange Analspitze ohne Mikrotrichien in der apikalen Hälfte; caudad gebogene,

fast digitiforme Basalloben; Gonostylen abweichend von allen bisher beschriebenen Smittia-Arten:

Crista dorsalis abgerundet, extrem stark vorgewölbt. Auffällig ist darüberhinaus auch die intensive

Beborstung des Skutellums.

Imago O (Holotypus), Terminologie wie bei SAETHEr (1980). Daten der Paratypen als Klammerzu-

sätze.

Körperlänge 2,72 mm (2,44— 2,83). Flügellänge 1,83 mm (1,74—1,82). Verhältnis Körperlänge/Flü-

gellänge 1,49 (1,41—1,59). Verhältnis Flügellänge/Länge des Profemur 3,02 (3,00—3,22). Färbung von

Thorax und Abdomen einheitlich dunkelbraun.

Kopf. Länge des Antennenendgliedes 507 um (432-520). AR 1,88 (1,57—1,84). Augen behaart,

dorsalwärts mäßig verlängert. Temporal- und Postorbitalborsten (nicht deutlich voneinander diffe-

renziert): 8 (8-10). 4 (2-6) Clypeusborsten. Länge der Palpenglieder (Mikrometer): 24 (22-24), 50

(44-46), 104 (88-96), 88 (80-88), 102 (84-90).

Thorax. Antepronotum wie bei anderen Smittia-Arten entwickelt, mit 2 lateralen Borsten. 21

(14-21) Dorsocentralborsten, zum Teil in doppelter Reihe. Anzahl der Acrostichalborsten (= Dor-

somedianborsten) beim Holotypus wegen Seitenlage der Thoraxpartie nicht bestimmbar; bei den Pa-

ratypen wurden bis zu 15 Acrostichalborsten gezählt, die weit vorne am Pronotum beginnen. 5 (5-8)

Präalarborsten. Anzahl der Skutellarborsten beim Holotypus wegen Seitenlage des Thorax nicht ex-

akt zu ermitteln; bei den Paratypen bis zu 21 (!) Skutellarborsten von verhältnismäßig geringer Größe

vorhanden.

Abb. 1u.2: Smittia scutellosetosa C': 1. Flügel, Maßstab = 0,5 mm; 2. Hypopygium (dorsal), Maßstab = 0,1 mm.

176

Flügel (Abb. 1). VR ca. 1,0. Verlängerung der Costa (jenseits der Mündung von R,;;): 86 um

(76-104). Brachiolum mit 1 Borste (1-2). 3 (3) Borsten auf R,, 7 (8-13) Borsten auf R,;;, 13 (3-13)

Borsten aufR.

Beine. Sporn der Vordertibia 44 um (42-52) lang, Sporne der Mitteltibien 20 um (18-22) und

22 um (20-28) lang, Sporne der Hintertibien 22 um (20-22) und 50 um (48-52) lang. Breite der

Vordertibia im distalen Bereich: 32 um (32-34); Breite der Mitteltibia im distalen Bereich: 32 um

(32); Breite der Hintertibia im distalen Bereich: 42 um (42-48). Tibialkamm mit 10 (9—12) Borsten,

die kürzesten 26 um (24-26), die längsten 40 um (40-42) lang.

Länge (Mikrometer) der Beinglieder und Längenrelationen:

Fe 4bat Ta, Ta, Ta, Ta, Ta, LR BV SV

BE 592 740 357 229 144 98 84 0,48 3,04 313

545 - 598 697 - 748 321 - 387 205 - 241 134 - 144 92 - 104 80 - 86 0,46-0,52 3.,01-3,06 3,48-3,87

P 653 669 273 158 116 80 80 0,41 3,68 4,84

626 - 700 639 - 700 261 - 289 148 - 168 112 - 120 78 - 80 76 - 84 0,41 3,63-3,74 4,81-4,85

B 700 772 405 221 172 92 90 0,53 3,26 3,64

651 - 724 700 - 780 381 - 421 196 - 225 158 - 182 88 - 100 84 - 94 0,54-0,55 3,13-3,29 3,50-3,57

Abdomen. Die Beborstung der Abdominaltergite wurde nur bei einem Paratypus überprüft. Die

Anzahl der Borsten auf den Tergiten I-VIII betrug: 28, 62, 67, 55, 52, 47, 40, 50. Beim gleichen Indi-

viduum betrug die Anzahl der Borsten auf den Sterniten I-VIII: 0, 0, 11, 14, 26, 26, 24, 23.

Hypopygium (Abb. 2). Länge der Analspitze 94 um (90-94). Länge des Gonocoxits 156 um

(140-172). Länge des Gonostylus 90 um (80-92). Griffel 13 um (12-14) lang. Virga 26 um (26-28)

lang. HR 1,73 (1,59—1,95). HV 3,02 (2,77-3,54).

Ökologie. Die Jugendstadien von Smittia scutellosetosa spec. nov. liegen nicht vor. Es ist jedoch mit

großer Wahrscheinlichkeit davon auszugehen, daß Habitate im unmittelbaren Umfeld der exponier-

ten Barberfallen, in denen die Imagines gefangen wurden, besiedelt werden.

Das Untersuchungsgebiet auf hochgelegenen Kiesbänken im Auengebiet der Isar ist in der Tat ein

ungewöhnlicher Lebensraum für terrestrische Chironomiden — oder wohl richtiger: ein bisher wenig

| Abb. 3: Smittia scutellosetosa Q: Kopf, Maßstab = 0,2 mm.

MT ae

beachteter und entsprechend wenig besammelter Lebensraum: auf engstem Raum kommt es hier un-

ter dem prägenden Einfluß der ökologischen Faktoren Temperatur und Bodenfeuchte/-trocknis zur

Bildung eines vielgestaltigen Biotopmosaiks, in dem bei überraschend hohen Artenzahlen vor allem

xerophile Faunenelemente — z. B. bei den Arachniden und Carabiden (mündl. Mitt. M. Baehr) — eine

wichtige Rolle spielen. Die knapp bemessene Zufuhr von Wasser beschränkt sich auf gelegentliche Re-

genfälle und nächtliche Tauniederschläge, die von einer lückigen Moosdecke für kurze Zeit gespei-

chert werden können, ansonsten aber im kiesig-sandigen Untergrund rasch versickern. Nur bei extre-

men Hochwässern der Isar ist in größeren Zeitabständen mit kurzfristigen Überflutungen dieser

Standorte zu rechnen.

Imago

Ob die hohe Zahl der Skutellarborsten bei Smittia scutellosetosa spec. nov. differentialdiagnostische Wertigkeit

hat, kann angesichts des unrevidierten Status der Gattung Smittia zur Zeit nicht endgültig entschieden werden.

Hinweise, die dies jedoch richtig erscheinen lassen, ergeben sich bei Durchsicht der umfangreichen Belegsammlung

des Autors an westpaläarktisch verbreiteten Smittia-Sıppen: alle hierauf überprüften Arten wiesen 2 bis maximal

8 Skutellarborsten auf.

Im Rahmen von Emergenz-Studien (CAspErs 1980, 1983) gefangene Smittia-Arten, bei denen infolge überein-

stimmender phänologischer Daten eine direkte Zuordnung von Männchen und Weibchen möglich war, zeigten

darüberhinaus keine bedeutenden geschlechtsspezifischen Unterschiede in der Skutellarbeborstung an.

Es spricht somit einiges dafür, daß die Skutellarbeborstung der neuen Smittia-Art ein hochwertiges diagnosti-

sches Merkmal darstellt und daß es sich bei dem nachstehend beschriebenen Smittia-Weibchen (mit 20 Skutellar-

borsten) aus einer Barberfalle bei Aumühle um das Weibchen von Smittia scutellosetosa handelt:

Hi At 2335

Abb. 4u.5: Smittia scutellosetosa 9:4. Flügel, Maßstab = 0,5 mm; 5. Genital (ventrolateral), Maßstab = 0,2 mm.

178

Körperlänge 2,30 mm. Flügellänge 1,54 mm. Verhältnis Körperlänge/Flügellänge 1,49. Verhältnis

Flügellänge/Länge des Profemur 3,29. Färbung von Thorax und Abdomen einheitlich dunkelbraun.

Kopf (Abb. 3). Länge und Breite des Pedicellus (Mikrometer): 48/56. Länge und Breite der Flagel-

lomeren 1-5 (Mikrometer): 68/31, 46/26, 46/24, 46/24, 66/23. Temporal- und Postorbitalborsten

(nicht deutlich voneinander differenziert): 10. 6 Borsten auf dem Clypeus. Länge der Palpenglieder

(Mikrometer): Länge der Palpenglieder 1 und 2 bei dem vorliegenden Objektträger-Präparat nicht ex-

akt meßbar; Palpenglied 3: 83. 4: 64. 5: 80.

Thorax. Antepronotum wie im männlichen Geschlecht ausgebildet. 30 bzw. 38 Dorsocentralbor-

sten, 12 Acrostichalborsten. 16 bzw. 15 Präalarborsten. 22 kleine Skutellarborsten.

Flügel (Abb. 4). Verlängerung der Costa (jenseits der Mündung von R,,;): 168 um. Brachiolum mit

3 Borsten. R, mit 6 Borsten, Ry,; mit 24 Borsten, R mit 16 Borsten.

Beine. Sporn der Vordertibia 31 yum lang, Sporne der Mitteltibien 18 und 22 um lang, Sporne der

Hintertibien 12 und 30 um lang. Breite der Vordertibia im distalen Bereich: 32 um; Breite der Mittel-

tibia im distalen Bereich: 32 um; Breite der Hintertibia im distalen Bereich: 42 um. Tibialkamm mit

11 Borsten, die kürzesten 24 um, die längsten 38 um lang.

Länge (Mikrometer) der Beinglieder und Längenrelationen:

Fe Ti Ta, Ta, Ta, Ta, Ta, LR BV SV

P] 476 516 221 144 94 ed Ti 0,43 Heil

D, 570 535 205 124 96 Sg? Or ne 5,

P 582 610 331 168 146 18 Te 0,54 19,25 2,650,

Abdomen. Anzahl der Borsten auf den Abdominaltergiten I-VII: 48, 49, 40, 34, 29, 39, 47. Anzahl

der Borsten auf den Abdominalsterniten I-VII: 0, 8, 13, 21, 22, 20, 38.

Genitalien: Abb. 5.

Smittia amoena spec. nov.

(Abb. 6-7)

Fundort: Kiesbänke in der Flußaue der Donau bei Gerolfing (Nähe Ingolstadt, Bundesrepublik Deutschland).

Material: Holotyp ©’, Barberfalle, 3.5.1983, leg. M. Baehr; in coll. Zoologische Staatssammlung München

(Objektträgerpräparat).

Diagnose: Auch Smittia amoena weist ein vom üblichen Smittia-Typ stark abweichendes Hypo-

pyg-Merkmal hoher differentialdiagnostischer Wertigkeit auf: die hyaline Crista dorsalis ist deutlich

ausgeprägt und extrem stark vorgewölbt.

Imago JO’ (Holotypus)

Körperlänge 2,44 mm. Flügellänge 1,54 mm. Verhältnis Körperlänge/Flügellänge 1,58. Verhältnis

Flügellänge/Länge des Profemur 2,87. Färbung von Thorax und Abdomen einheitlich dunkelbraun.

Kopf. Länge des Antennenendglieds 374 um. AR 1,15. Augen behaart, dorsalwärts nicht verlän-

gert. I innere, 2 äußere Temporalborsten; soweit am Typusexemplar erkennbar, keine Postorbital-

borsten. Clypeus mit 2 Borsten. Länge der Palpenglieder (Mikrometer): 1 nicht meßbar am Typusex-

emplar; 40, 78, 76, 64.

Thorax. Antepronotum fehlt am Typusexemplar. 12 Dorsocentralborsten in einfacher Reihe.

Acrostichalborsten beginnen vorne am Pronotum, am Typusexemplar sind 4 deutliche Borsten er-

kennbar. 6 Präalarborsten, die vorderen schwächer ausgebildet als die hinteren. Anzahl der Skutellar-

| borsten am Typusexemplar nicht deutlich erkennbar, mindestens 4 Borsten vorhanden.

Flügel (Abb. 6). R,,; erreicht nicht ganz die Costa. Verlängerung der Costa (jenseits der Mündung

| von R,,;): 80 um. Brachiolum ohne Borsten. 6 Borsten auf R,, 6 Borsten auf R,;;, 6 Borsten aufR.

179

NEIN K

Abb. 6u.7: Smittia amoena O': 6. Flügel, Maßstab = 0,5 mm; 7. Hypopygium (dorsal), Maßstab = 0,1 mm.

Beine. Sporn der Vordertibia 42 um lang, Sporne der Mitteltibien 16 und 20 um lang, Sporne der

Hintertibien 20 und 48 um lang. Breite der Vordertibia im distalen Bereich: 30 um; Mitteltibia:

32 um; Hintertibia: 40 um. Tibialkamm mit 9 Borsten, 20 bis 40 um lang.

Länge (Mikrometer) der Beinglieder und Längenrelationen:

Fe Ti Ta, Ta, Ta, Ta, Ta, LR BV SV

PJ 538 645 297 182 130 84 84 0.46 3.08 3.98

P, 620 613 233 132 110 68 78 0.380 73.18 5.29

P 632 661 357 166 156 88 88 OS5ANEE Sn 3.62

Abdomen. Die Beborstung der Abdominaltergite und -sternite ist am Objektträger-Präparat des

Typusexemplars nicht genau zu bestimmen, entspricht aber offensichtlich etwa der Beborstung von

Smittia scutellosetosa.

Hypopygium (Abb. 7). Länge der Analspitze 88 um. Länge des Gonocoxits 162 um. Länge des

Gonostylus 96 um. Griffel 8 um lang. Innere Skleritstrukturen des Hypopygiums am Typusexem-

plar nicht in Einzelheiten erkennbar. HR 1,69. HV 2,54.

180

Ökologie. Auch bei dieser neuen Art ist zu vermuten, daß der Lebensraum der — vorerst nicht be-

kannten - terrestrischen Larven in der Nähe der Barberfallen-Standorte zu suchen ist: die männlichen

Imagines der Smittia-Arten zeichnen sich — soweit dies bekannt ist — nicht durch auffällige Flugakti-

vität bzw. einen größeren Aktionsradius aus.

Von seinen allgemeinen physiographischen Verhältnissen ähnelt der Standort „Kiesbänke/Donau-

Auwald bei Gerolfing“ dem Locus classicus von Smittia scutellosetosa am Isarufer bei Aumühle. Auf

den ersten Blick ist der im Vergleich noch stärker xerotherme Charakter an schütteren Wacholder-

und Kiefernbeständen auf weitgehend vegetationsfreiem Untergrund erkennbar. Auch bei Betrach-

tung der ökologischen Standortansprüche epigäischer Raubarthropoden (mündl. Mitt. M. Baehr) er-

gibt sich eine eindeutige Dominanz wärmeliebender Arten.

Es ist davon auszugehen, daß der Fundort von Smittia amoena — ca. 200 m Entfernung von der Do-

nau bei Normalwasserstand — auch bei extremen Hochwasserbedingungen nicht überflutet wird.

Literatur

CaspERS, N. 1980: Die Emergenz eines kleinen Waldbaches bei Bonn. — Decheniana-Beihefte (Bonn) 23: 1-175.

CASPERS, N. 1983: Chironomiden-Emergenz zweier Lunzer Bäche, 1972. — Arch. Hydrobiol. Suppl. 65: 484549.

GOETGHEBUER, M. 1940— 1950: Tendipedidae (Chironomidae). f) Subfamilie Orthocladiinae. A. Die Imagines. —

In LINDNER, E. (Hrsg.): Die Fliegen der palaearktischen Region. 13g: 1— 208.

HALVORSEN, G. A. 1982: Saetheriella amplicristata gen. n., sp. n., anew Orthocladiinae (Diptera: Chironomidae)

from Tennessee. — Aquatic Insects 4: 131-136.

SAETHER, O. A. 1980: Glossary of chironomid morphology terminology (Diptera: Chironomidae). — Ent. scand.

Suppl. 14: 1-51.

SCHLEE, D. 1966: Präparation und Ermittlung von Meßwerten an Chironomidae (Diptera). —- Gewässer und Ab-

wässer 41/42: 169— 193.

Professor Dr. Norbert Caspers,

Morgengraben 8, D-5000 Köln 80

181

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SPIXIANA | Supplement 14 | 183— 190 | München, 15. Juli 1988 ISSN 0177-7424 |

Rheomus, un nouveau genre du complexe Harnischia avec deux

nouvelles especes d’Afrique du Nord

(Diptera, Chironomidae)

Henri Laville et Friedrich Reiss

Abstract

Rheomus, a new genus of the Harnischia complex with two new species from North Africa (Diptera, Chirono-

midae)

Rheomus, a new genus of the Harnischia complex is established for two new species from North Africa. The

imago ©’, the pupal exuvia, of R. yahiae recorded in Morocco and Tunisia and the imago ©’ of R. alatus only

recorded in Morocco are described. From the imaginal characteristics, Saetheria seems the most related genus but

the pupal characteristics are rather related with the genus Cryptotendipes. Nevertheless, its throw-back anal lobe is

a very original one among Chironomini.

Introduction

L’etude des Chironomides recoltes par derive dans un affluent rive droite de l’Oued Medjerda (Tu-

nisie) par M. Boumaiza nous a permis d’isoler une nymphe CO’ originale dont 3 imagos C’, dejä connus

du Maroc et conserves A la Zoologische Staatssammlung München, se rangent dans le complexe Har-

nischia.

Les caracteres nettement regressifs du lobe anal de la nymphe autorisent la creation d’un nouveau

genre avec deux nouvelles especes connues seulement d’Afrique du Nord.

Nous remercions M. M. Boumaiza (Universite de Tunis) pour le leg du mat£riel de Tunisie.

Rheomus gen. nov.

L’espece type est Rheomus yahiae spec. nov.

Imago C:

Petit, longueur de l’aile 1,2—1,65 mm. Yeux nettement prolonges dorsalement et juste separes de la

longueur du prolongement. Tubercules frontaux cyclindriques presents. Antenne avec 11 flagellome-

res, AR = 1,2-1,3. Palpes normaux, non raccourcis. Nombre de soies thoraciques peu eleve en

rapport avec la petite taille du corps; soies acrostichales presentes. Scutum avec tuberecule. Aile avec

lobe anal tr&s peu developp&, squama seulement avec 2-3 soies. FCu Eloignee distalement de RM, C

se terminant avant M et non prolongee au-delä de R4+5. Seule R avec quelques courtes soies, les

autres nervures sans soies. Surface alaire sans soies ni taches. Pattes normales non raccourcies. Grosses

pulvilles presentes. Peignes du tibıa plats, courts et non separ&s avec seulement un court Eperon.

183

Hypopyge avec un tergite anal ä bandes assez superficielles reunies par une zone brunätre dans la

partie mediane. Pointe anale longue, en forme de raquette et nue. Tergite anal sans soies medianes, avec

des soies apicales Ecartees de la base de la pointe anale. Volsella superieure en forme de pied ou de mas-

sue, ämembrane souple, avec ou sans tubercule apical, sans soies dorsales, avec 2-3 longues soies ven-

trales issues d’une zone Amicrotriches. Volsella inferieure de taille petite amoyenne, garnie de longues

microtriches se prolongeant loin vers sa base tout le long du bord interne du gonocoxite. Gonocoxite

avec seulement 3 longues soies medianes marginales. Gonostyle effile, avec dans sa moite apicale une

lamelle dorsale ou mediane, une zone dorsobasale depourvue de microtriches, une dent apicale absente

et des soies me&dianes en partie fourchues distalement.

Nymphe:

Seule lanymphe de R. yahiae est connue. Sa description ci-apres peut egalement servir de genero-

type.

Repartition et Ecologie

Le genre Rheomus est actuellement connu d’Afrique du Nord (Maroc et Tunisie).

Il s’agit probablement d’el&ments faunistiques du Potamal dont les larves colonisent de preference

les sediments sableux des grands cours d’eau.

Diagnose differentielle

L’imago JO de Rheomus se differencie des autres representants du complexe Harnischia surtout par

la combinaison des caracteres suivants de l’hypopyge: bandes du tergite anal droites pratiquement non

reunies dans la partie mediane d&pourvue de soies; pointe anale longue, en forme de raquette et nue;

Volsella superieure en forme de pied ou de massue, dorsalement nue et avec deux longues soies ventra-

les dans une zone de microtriches; Volsella inferieure de longueur courte amoyenne; gonocoxite avec

une lamelle dorsale ou mediane dans sa moite apicale. De plus, l’imago possede des tubercules fron-

taux, de longues soies acrostichales (dorsom&dianes), un petit tubercule scutal ainsi que des peignes du

tibia non separes et avec un seul petit &peron sur Pr et Pıyı. Rheomus se distingue du genre en apparence

le plus proche, Saetheria Jackson, 1977 par la presence d’un seul Eperon tibial court & la place de deux

eperons forts, par la presence d’un petit tubercule scutal, absent chez Saetheria, par des bandes en

forme de V, pratigquement separees dans la partie mediane du tergite anal et en forme de Y chez Saethe-

ria, par l’absence de soies medianes sur le tergite anal (pr&sentes en petit nombre chez Saetheria et in-

serees ä l’extremite distale des bandes du tergite anal), par une pointe anale fortement elargie distale-

ment en forme de raquette (legerement &largie chez Saetheria) par une Volsella superieure dorsalement

nue (en 2 lobes separes: le distal couvert de microtriches, le basal sans microtriches), enfin par la pre-

sence d’une lamelle dorsale ou mediane dans la partie terminale du gonostyle, absente chez Saetheria.

Nymphe: nous renvoyons ä la diagnose differentielle de Janymphe de R. yahiae.

Rheomus yahıae spec. nov.

Imago d’:

Taille: petit, longueur de l’aile 1,35—1,65 mm.

Coloration: corps brun (dans l’alcool). Vittae (bandes m&sonotales) brun fonce ainsi que les 2/3

posterieurs du postnotum, le praeepisternum et une tache sur l’anepisternum.

Tete: yeux fortement prolonges dorsalement et separes dans la partie mediane de la longueur du

prolongement. Tubercules frontaux cylindriques, jusqu’ä 10 um de long. Antenne avec 11 flagello-

meres, AR = 1,22. Longueur des articles 2-5 des palpes en um (Holotype): 27, 106, 124, 189.

Thorax: avec 5 soies acrostichales, 5-8 dorsocentrales, 2—3 prealaires et 6-7 scutellaires. Scutum

avec un petit tubercule arrondı.

184

Aile: lobe anal peu developpe. Squame avec seulement 2-3 soies. FCu Eloignee distalement deRM,

C non prolongee au-delä de R4+5 et finissant avant M. R2+3 debouchant au !/3 proximal entre R1

et R4+5. Seule R porte 3-5 soies.

Pattes: longueur des articles en um (Holotype):

Fe Ti Ta, Ta; Ta; Ta; Ta;

Pı 600 460 630 305 235 170 100

Pıı 540 540 200 130 100 70 80

Praeio 640 370 210 175 100 90

LR = 1,32-1,37. Pulvilles presque aussi longs que les griffes. Tarses anterieurs non barbus. Sensilla

chaetica non visible sur le Ta; de Pı,. Peignes des tibias moyens et posterieurs plats, courts, non separes,

avec seulement un court Eperon.

Hypopyge (fig. 1): bandes du tergite anal en V, assez superficielles, ne se rejoignant pas dans la par-

tie mediane oü une simple zone brunätre les relie. Pointe anale a base £troite et partie terminale arron-

die en forme de disque, un peu recourbee ventralement (fig. 1a). Partie basale plus courte que la partie

terminale. Tergite anal sans soies medianes, soies apicales legerement Ecartees de la base de la pointe

et regroup£es en touffe. Volsella superieure en forme de pied lateralement deporte. Partie distale A

membrane d’ apparence delicate, sans soies dorsales, avec 2—3 soies ventrales issues d’une zone de mi-

crotriches.

Volsella inferieure arrondie distalement, tres courte, n’atteignant pas la base du gonostyle.

Fig. 1. Rheomus yahiae spec. nov., Hypopyge dorsal. a. Pointe anale, vue laterale. Fig. 2. Rheomus alatus spec.

_ nov., Hypopyge dorsal, a. Gonostyle, vue laterale.

185

Fig. 3. Rheomus yahiae spec. nov., Nymphe ®. a. Apotome frontal. b. Anneau basal de la corne thoracique. c.

Thorax. d. Tergites abdominaux I-VII. e. Sternites V-VIII, f. Corne thoracique.

186

La Volsella inferieure se raccorde tout le long du bord interne du gonocoxite. Gonocoxite avec seu-

lement 2-3 longues soies medianes et marginales. Gonostyle avec une partie basale etroite, un bord

interne tres concave, une partie distale &largie et aplatie en lame de couteau, progressivement retrecie

vers l’apex. Partie distale du gonostyle entierement couvert de microtriches, partie basale avec une

zone dorsale depourvue de microtriches; soies du bord interne partiellement fourchues a leur extre-

mite.

Imago Q: inconnue.

Nymphe 9:

Longueur 3,2—3,7 mm (n = 8). Exuvie pratiquement incolore.

Cephalothorax (fig. 3a): soies frontales courtes, fines, inserees & l’extremite de petits tubercules ce-

phaliques coniques. Tubercules frontaux absents. Corne thoracique de type Chironomini, caracterı-

stique (1 = 0,7—0,9 mm): formee par 5 branches bifurquees dans leur moitie distale en branches plus

fines, avec ä sa base une touffe de filaments courts et fins; anneau basal ovale avec une structure alveo-

laire bien visible (fig. 3b, f). En avant de l’anneau, presence d’un processus conique legerement colore.

Thorax finement granuleux dorsalement, par ailleurs lisse (fig. 3c). Tubercule prealaire developpe

en projection conique courte et plissee. Tubercule du scutum peu developpe. Fourreau alaire, aux

bords legerement colores, sans nez ni perles. Soies thoraciques: 2 precorn£ales, 2 antepronotales, 4

dorsocentrales.

Abdomen (fig. 3d, e): tergite I nu; II-VII avec une paire de tubercules medio-posterieurs garnis

d’&pines courtes et portant une forte soie plus chitinisee. Tergite II sans rangee de crochets. Conjonctif

et pleures sans chagrin; fin chagrin aux angles anterieurs des tergites VI, VII, VIII, sur la partie me-

diane des tergites III-VI ainsi que sur la moitie ant&rieure du segment anal. Pedes spurii A absents; Pe-

des spurii B aux angles anterieurs du segment I, absents du segment II. Sternites pratiquement sans

chagrin avec deux lobules posterieurs portant chacun une longue soie plus chitinisee; quelques spinu-

les seulement presentes pres des lobules des sternites V et VI. Segment VIII sans Epines nı &perons a

P’angle posterieur. Segment anal avec deux lobes allonges, termin&s par une pointe effilee et recourbee

ventralement, avec chacun deux L setae inserees au tiers anterieur. Sac genital P distalement trilobe.

Setation abdominale: segments II-VII avec 3 L setae courtes. Segment VIII avec 3 L setae plus lon-

gues. 4 L setae observees sur un cöte d’une seule exuvie. La setation des tergites et des sternites V-VII

est representee sur la figure 3d et 3e.

Mat£riel: Holotype 1 imago CO’, Maroc, S. Yahia Boka, juin 1967, leg. Choumara. Paratypes: 3 imagos JO’ du

locus typicus, 20.4.1967 et juin 1967.

Maroc: 1EQ Oued Fes, 20.12.1987, leg. Azzouzi. Tunisie: INC’,5EQ Oued Tessa, 15.5.1983; 1EQ Oued Med-

jerda (Jendouba) 12.4.1985; 2EQ Oued Medjerda (Bathan), 6.4.1985, leg. Boumaiza.

L’Holotype se trouve dans la Zoologische Staatssammlung München. Paratypes dans la collection H. Laville,

Toulouse et dans Z. S. M. Tous les exemplaires sont montes dans l’Euparal.

Repartition et Ecologie

R. yahiae est connue du Maroc et de Tunisie.

Au Maroc, la station de l’Oued Fes, petit affluent (22 km) de l’Oued Sebou, est situee 3 km en

amont de Fes et ä 380 m d’altitude: elle est large de 13 m, profonde de 0,8-1 m, avec un substrat de

gravter et de sable grossier; la temperature de l’eau Etait de 22°C le 20 avrıl 1986.

Deux stations tunisiennes se situent dans le principal fleuve du Nord de la Tunisie l’Oued Medjerda,

la troisieme dans l’Oued Tessa, 2 km en amont de sa confluence avec la Medjerda.

Il s’agit de cours d’eau de plaine (28 m—146 m), larges (8-12 m), ä fond sableux a sablo-vaseux,

avec des temperatures de 24°C en juin etde 12°C en fevrier. La vegetation bordante estrare ou absente.

Dans l’Oued Medjerda, pres de Jendouba, l’espece a Et£ r£coltee en compagnie de l’espece Kloosia pu-

silla (L.) (12.4.85) et s’avere, comme cette derniere, un nouvel exemple d’un element faunistique du

potamal au sein du complexe Harnischia.

187

Diagnose differentielle

L’imago © se differencie bien de l’imago © de alatus au niveau des caracteres de l’hypopyge: Vol-

sella superieure en forme de pied, sans tubercule (en forme de massue avec un petit tubercule apical);

gonostyle avec un bord interne nettement concave et une moitie distale elargie en lame de couteau

(bord interne droit avec une lamelle laterodistale sur la face dorsale). De plus, yahiae est de coloration

nettement plus fonc&e que alatus.

La nymphe de R. yahiae se caracterise par un segment anal d’un type original et apparement unique

chez un Chironomini: de plus, la reduction de la setation anale 4 2 simples L setae le rapprocherait de

celui d’un Tanypodinae. Seulement 3 L setae aux segments V—-VIII (normalement 4 LS setae chez le

complexe Harnischia).

Rheomaus alatus spec. nov.

Imago d:

Taille: petit, longueur de l’aile 1,2—1,35 mm.

Coloration: corps brun-jaune (dans l’alcool). Vittae brun, bien separ&es. Postnotum brun avec une

etroite bande basale plus claire. Praeepisternum, une tache sur l’anepisternum, pedicellus, articles 2—4

du palpe, egalement brun; article 5 brun clair. Balanciers clairs. Tergites abdominaux I-VI avec une

bande medio-longitudinale brune. Pattes entierement brunes excepte la base du femur anterieur plus

claire et les femurs moyens et posterieurs clairs avec toutefois une bande longitudinale brune.

Tete: yeux fortement prolonges dorsalement et separes dans la partie mediane de la longueur du

prolongement. Tubercules frontaux cylindriques, jusqu’ä 16 um de long. Antenne avec 11 flagello-

meres, AR = 1,17—1,33 (n = 3). Longueur des articles 2-5 des palpes en um (Holotype): 38, 88, 103,

133

Thorax: avec 6 soies acrostichales, 5-7 dorsocentrales, 3—4 prealaires et 6-7 scutellaires. Scutum

avec un petit tubercule arrondı.

Aile: lobe anal peu developpe&. Squame avec seulement 2 soies. FCu eloignee distalement de RM, C

non prolonge&e au-delä de R4+5 et finissant avant M. R2+3 debouchant au !/3 proximal entre Ri et

R4+5. Seule R porte 4—6 soies.

Pattes: longueur des articles en um (Holotype):

Fe Ti Ta; Ta; 1a; len Ta;

P, 480 365 460 235 170 120 7

Pır 420 415 150 95 70 55 60

Pın 490 500 265 150 126 65 70

LR = 1,26-1,40 (n = 2). Pulvilles presque aussi longs que les griffes. Tarses anterieurs non barbus.

2 sensilla chaetica subapicales sur Ta, de Pı,. Peignes des tibias moyens et posterieurs plats, courts, non

separ&s, avec seulement un court Eperon.

Hypopyge (fig. 2): bandes du tergite anal en V, assez superficielles, ne se rejoignant pas dans la par-

tie mediane oü une simple zone brunätre les relie. Pointe anale ä base £troite et partie terminale elargie

en forme de disque, un peu recourb&e ventralement. Partie basale presque aussi longue que la partie

terminale. Tergite anal sans soies medianes, soies apicales legerement Ecartees de la pointe et regrou-

pees en touffe. Volsella superieure &troite, presque droite, Elargie distalement et avec une ebauche de

tubercule apical; dorsalement sans soies et avec 2 longues soies subapicales dans une zone de microtri-

ches sur la face ventrale. Volsella inferieure triangulaire et effil&e, de longueur moyenne, atteignant la

partie basale du gonostyle et garnie de tr&s longues microtriches. La Volsella inferieure se raccorde

tout le long du bord interne du gonocoxite. Gonocoxite avec seulement 3 longues soies medianes et

188

marginales. Gonostyle avec un bord interne droit, legerement elargi au centre; la partie distale avec au

bord externe une lamelle peu apparente en vue dorsale, seulement visible en position laterale (fig. 2a);

soies du bord interne partiellement fourchues ä leur extremite.

Imago 9, Nymphe et Larve: inconnues.

Materiel: Holotype 1 imago O’, Maroc, Meknes, 553 mN.N., 23.8.1979, leg. G. Fontain. Paratypes: 1 imago JO’,

Maroc, Haut-Atlas du Sud, Oasis Meski, 1160 m N.N., 2.5.1979, leg. F. Auer & K. Werner; 2 imagos JO’, Ma-

roc, Oued Tensift, rive Sud, Palmeraie de Marrakech, 10.5.1979, leg. D. Masson.

Les preparations & l’Euparal de l’Holotype et du Paratype premier nomme se trouvent dans la Zoologische Staats-

sammlung München, les deux autres Paratypes dans la collection H. Laville, Toulouse.

Repartition et €cologie

L’espece n’est jusqu’ici connue que du Maroc. Son Ecologie n’est pas connue.

Diagnose differentielle

L’espcce se differencie du second representant du genre (yahiae) par les caracteres suivants de ’hy-

popyge: Volsella superieure en forme de massue et avec une Ebauche de tubercule apical (en forme de

pied, sans tubercule); gonostyle avec un bord interne droit et une lamelle latero-distale peu apparente

dorsalement, bien visible en vue laterale (bord interne, en partie concave, et lamelle medio-distale en

lame de couteau).

Discussion

Les caracteres imaginaux montrent (voir diagnose differentielle du genre) que Rheomus se rappro-

che le plus du genre Saetheria qui, de son cöt&, a montre de grandes affinites avec le genre Paraclado-

pelma Harnisch, 1923. Malgre la pr&sente description d’un nouveau genre au sein de ce groupe par-

ental, les limites du genre doivent rester provisoires, car d’une part, un grand nombre d’especes dont

les stades jeunes sont inconnus different beaucoup du generotype et d’autre part, plusieurs especes

non decrites existent dont l’analyse des caracteres peut amener des transformations taxonomiques.

L’appartenance douteuse au genre existe pour Paracladopelma mikiana (Goetgh., 1973) espece Ouest-

Palearctique, ainsi que pour les trois especes afrotropicales P. melutense (Freem., 1957), P. pullata

(Freem., 1957) et P. reidi (Freem., 1957). Une nymphe © de l’espece essentiellement afrotropicale

P. graminicolor (Kieff., 1929), recemment trouvee ä la Zoologische Staatssammlung, s’est revelee ap-

partenir nettement au genre Cryptotendipes Lenz, 1941. Paracladopelma tamanıpparai Sasa, 1983 du

Japon, d’apres la description originale et les figures se range sans aucun doute dans le genre Saetheria.

Ces quelques exemples justifient la necessit€ de nouvelles &tudes taxonomiques dans cette partie du

complexe Harnischia.

La nymphe presente une corne thoracique caract£ristique de type Chironomini.

L’absence de rangee de crochets au tergite II et l’absence de peignes ou d’Eperons latero-posterieurs

au segment VIII larapproche du genre Harnischia (Pinder & Reiss 1986). L’ornementation des tergites

abdominaux I-VIII ressemble davantage ä celle du genre Cryptotendipes.

L’absence de parente reelle de lanymphe avec quelques genres de la tribu des Chironomini interdit

toute speculation phylogenetique. La forte regression du segment anal de lanymphe de R. yahiae in-

cite A considerer le genre Rheomus comme autapomorphe.

L’&cologie de R. yahiae, comparable & celle des genres Kloosia et Demicryptochironomus (Irmakia)

(Reiss 1988 a, b), est un nouvel exemple d’un genre potamobionte au sein du complexe Harnischia.

189

Bibliographie

FREEMAN, P. & CRANSTON, P. S. 1980: Family Chironomidae: In Crosskey R. W. (Ed.): Catalogue of the Diptera

of Afrotropical Region. British Museum (Natural History), London: 175—202

JACKSON, G. A. 1977: Nearctic and Palaearctic Paracladopelma Harnisch and Saetheria n. gen. (Diptera: Chirono-

midae). — J. Fish. Res. Board Canada, 34: 1321-1359

PinDEr, L. C. V. & Reıss, F. 1986: The pupae of Chironominae (Diptera: Chironomidae) of the Holarctic region

— Keys and diagnoses. — Ent. scand. Suppl. 28: 299-471

Reıss, F. 1988a: Die Gattung Kloosia Kruseman, 1933 mit der Neubeschreibung zweier Arten (Chironomidae,

Diptera) — Spixiana Suppl. 14:

Reıss, F. 1988b: /rmakia, ein neues Subgenus von Demicryptochironomus Lenz, 1941, mit der Beschreibung von

4 neuen Arten (Chironomidae, Diptera). — Spixiana Suppl. 14:

SAETHER, ©. A. 1977: Taxonomic studies on Chironomidae: Nanocladius, Psendochironomus and the Harnischia

complex. — Bull. Fish. Res. Board Can. 196: 143 p

Dr. Henri Laville,

Laboratoire d’Hydrobiologie, UA 695, CNRS,

Universite Paul Sabatier, 118 route de Narbonne,

F-31062 Toulouse Cedex.

Dr. Friedrich Reiss,

Zoologische Staatssammlung,

Münchhausenstr. 21,

D-8000 München 60.

190

SPIXIANA | Supplement 14 | 191-200 | München, 15. Juli 1988 ISSN 0177— 7424

A Contribution to the knowledge of Chironomids in Italy

(Diptera, Chironomidae)

By B. Rossaro

Abstract

In recent years Chironomid research ın Italy has developed greatly. 359 species have been recorded. The subfami-

lies are represented by 42 Tanypodinae, 23 Diamesinae, 3 Prodiamesinae, 143 Orthocladiinae, 148 Chironominae,

59 belong to the tribe Tanytarsini and 89 to Chironomini. Another 45 taxa tentatively identified must be added to

these. Most species are widely distributed in the Palearctic region. Few Mediterranean (6), Afrotropical (5) or Pan-

paleotropical (3) species have been found. The list includes five previously considered to be Nearctic species, which

have Holarctic distribution according to the present evidence.

Introduction

Chironomid research in Italy is very active. In the last ten years the number of species known to be

present has gone up steeply. Information was very scanty before 1968, grew slowly up to 1976, then

quickly. One important stimulus has been a series of volumes with identification keys for the aquatic

fauna published by the Italian National Research Council (C.N.R.). Four volumes deal with Chiro-

nomids: FERRARESE & Rossaro (1981), Rossaro (1982), FERRARESE (1983) & Nocenrini (1985). Only the

larvae of all subfamilies and the pupal exuviae of all subfamilies except Chironominae are considered

in these volumes. The literature about Chironomids in Italy has been reviewed by Rossaro (1979) and

FERRARESE (1982).

The interests in Chironomid research are three. There are systematic, zoogeographic and ecological

problems that stimulate the study. Italy is in the centre of the Mediterranean area and is a natural

bridge between Central Europe and Africa. There are several very different biotopes in Italy: cold

springs and glacier streams in the Alps, large lakes in the pre-Alps (Garda, Maggiore, Como), rivers

in the Padana lowland (Po, Adige, Ticino, Adda, etc.), Mediterranean streams in central and southern

Italy. A very long list of species was to be expected and this is indeed the case, although at present it

is very incomplete. The first reason is that no extensive sampling effort has been made. Pollution is the

second reason. The capture of springs and the creation of reservoirs for water supply are also unfavou-

rable factors for the preservation of autochthonous, highly diversified fauna. Eury-topic and eury-

ecious species are favoured. The large spread of insecticides and water drainage to fight malarıa in the

Mediterranean countries after the second World War destroyed the species living in marshes. Ob-

viously, pollution is not exclusive to Italy, but in some very urbanized areas it has certainly drastically

modified the species composition, favouring the more common ones more than elsewhere. The mas-

sive development of Chironomus salinarius in the Venetian and Orbetello lagoons is an example (Arı

& Majorı 1984).

The list of species in a country must be continuously updated, as new species are captured. Genera

are submitted to revision, synonyms are created, species change names obeying the principle of prio-

rity. Some species can be easily identified thanks to peculiar morphological characters, but it is diffi-

191

Fig. 1. Location of sampling sites in Italy. Only the sites sampled most intensely indicated.

1— Cold springs and glacier streams in Aosta valley (Valle d’Aosta)

2— Ortles-Adamello group; many different biotopes, springs, streams, small lakes, all above 1000 m, etc. (Lom-

bardıa and Trentino-Alto Adige)

3— the Ticino river near Turbigo and Boffalora (Lombardia)

4— Springs near Milano, called “fontanili” and the Lambro (Lombardia)

5— prealpine streams near Bergamo (Brembo and Serina) (Lombardia)

6— the Po River near Trino Vercellese (Torino) and Casale Monferrato (Piemonte)

7— the Po River between Piacenza and Cremona (Emilia Romagna)

8— Mantova lakes, along the Mincio river (Mantova) (Lombardia)

9— Reno and other streams on the northern side of the northern Apennines (Emilia Romagna)

10— the Potenza near Macerata (Marche)

11— the Aso near San Benedetto del Tronto (Marche)

12— Springs and streams in Parco Nazionale Abruzzi (Abruzzi e Molise)

13— Occhito artificial basin between Molise and Puglia, and the Fortore in Molise

14— the Cedrino and other small stream in northern Sardinia (Sardegna)

15— Rio Mannu and brackish waters in southern Sardinia (Sardegna)

16— Dirillo artificial basin and the Dirillo (Sicilia)

192

cult to give names to species belonging to genera that requiere revision such as Chaetocladius, Bryo-

phaenocladins, Smittia, Micropsectra atrofasciata group, Microtendipes, Cryptochironomus, etc.. The-

refore, the list in this paper represents only a transitory phase in our developing knowledge.

Methods, sampling sites

The list of species is based on determinations on adult males in most cases. Sometimes determinations are made

on pupal exuviae. Larvae were obtained alive from some sites (always from sites 4 and 7, sometimes from other sites,

see Figure 1) and reared to adults in the laboratory. In laboratory-reared material one can associate larvae and pupae

with the adults and this procedure is highly recommended. For species determination in the genus Chironomaus, sa-

livary gland chromosomal analysis was carried out C. annularius, C. anthracinus, C. melanotus and C. riparins

were identified by this technique.

Sampling efforts were made in different regions in Italy, but Lombardia has been the most intensively sampled.

Figure 1 is amap showing the sites at which the most intensive sampling was carried out. Each sampling site includes

many different stations and covers a large area. In each sampling site, captures were made throughoutthe year. Sites

that were sampled only at intervals are not reported in Figure 1.

Collecting was done with hand-nets, light traps, surface drift nets.

Results and Discussion

Table 1 lists the known species in Italy. It has been necessary to separate them into the following

groups:

1— Species well described and identifiable: within this group, we further separate:

1a— species present in the author’s collection (Department of Biology, University of Milan):they are

marked with + ın Table 1;

1b-— species as 1a, new for Italian fauna: they are marked with =;

1c-species reported in the literature as present in Italy, but not in the author’s collection; this material

is deposited in different collections and it has not been possible for the author to verify the validity of

all these findings. All these species are marked with £. In this group, species determinations before

1950 are especially dubious.

2— Species whose presence is unsure because:

2a— determinations were made on ill-preserved specimens or only pre-imaginal stages are available;

2b — species belong to genera that require revision.

Species belonging to 2a—- and 2b- are marked with *.

Table 1: Chironomidae taxa from Italy (see text for explanation)

Namppadinze + M. sp. Schweiz

+ Chnotanypus nervosus (Mg.) £ Natarsia punctata (Fabr.)

£ Tanypus kraatzi (K.) + Psectrotanypus varıns (Fabr.)

+ T. punctipennis (Mg.) + Alabesmyia longistyla Fitt.

+ T. villipennis (K.) £ A. monhlıis (L.)

* Procladius choreus (Mg.) + Arctopelopia griseipennis (v. d. W.)

P. sagittalıs K. + Krenopelopia binotata (Wied.)

+ Apsectrotanypus trıfascipennis (Zett.) £_ Conchapelopia melanops (Wied.)

+ Macropelopia fehlmanni (K.) + C.pallidula (Mg.)

£ M. goetghebueri (K.) + Monopelopia tenuicalcar (K.)

+ M.nebulosa (Mg.) + Paramerina divisa (Walk.)

£ AM. notata (Mg.) + P. cingulata (Walk.)

193

PaspsA

Rheopelopia acra Roback

R. maculipennis (Zett.)

R. ornata (Mg.)

Telopelopia fascigera Verneaux

Thienemannimyia carnea (Fabr.)

T. fuscipes (Edw.)

T. geijskesi (G.)

T. northumbrica (Edw.)

T. psendocarnea Murray

T. woodi (Edw.)

Xenopelopia falcigera (K.)

Telmatopelopia nemorum (G.)

Trissepelopia longimana (Staeg.)

Zavrelimyia barbatipes (K.)

Z. hırtimana (K.)

Z. melanura (Mg.)

Z. nubila (Mg.)

Z. punctatissima (G.)

Z. signatipennis (K.)

Larsıa atrocincta (G.)

Nilotanypus dubius (Mg.)

I Se I

mm mm + m + m ı + 1 ml + |

Diamesinae

Boreoheptagyia cinctipes Edw.

Psendodiamesa branicki (Now.)

P. nıvosa (G.)

Syndiamesa nıgra

Diamesa dampfi (K.)

D. permacer (Walk.)

. steinboecki (G.)

. longipes G.

. latitarsıs (G.)

. goetghebueri Pag.

.lindrothi G.

. laticanda Ser.-Tos.

modesta Ser.-Tos.

. bertrami Edw.

. aberrata (Lundb.)

sp. A

.incallida (Walk.)

. insignipes K.

. cinerella (Mg.)

thienemanni K.

. vaillantı Ser.-Tos.

. zernyi Edw.

Pseudokiefferiella parva (Edw.)

Potthastia gaedii (Mg.)

P. longimanus K.

® + "++ +++ HtH+r HN

SESESESESESESZ SUSE SEES ESESESD

++ ++++++++

Prodiamesinae

+ Prodiamesa olivacea (Mg.)

+ P.rufovittata (G.)

194

Sympotthastia spinifera Ser.-Tos.

Monodiamesa sp.

+ Odontomesa fulva (K.)

Orthocladiinae

II Se Sell N

"+ +++ 1

E+ ++ ++++H+H+

I++++++++

IF ++ I +++ HH IHN

Diplocladius cultriger K.

Euryhapsis annuliventris (Mall.)

Brıllia longifurca K.

B. modesta (Mg.)

Cardiocladius capucinus (Zett.)

C. fuscus K.

Tvetenia bavarica (G.)

T. calvescens (Edw.)

T. discoloripes (G.)

T. verralli (Edw.)

T. scanica (Br.)

Eukiefferiella gracei (Edw.)

. coerulescens (K.)

. devonica (Edw.)

. ilkleyensis (Edw.)

. fittkani Lehm.

. minor (Edw.)

. brehmi Gow.

. cyanea Th.

. clypeata (K.)

. psendomontana G.

. dittmari Lehm.

. fuldensis Lehm.

.lobifera G.

. brevicalcar (K.)

. tirolensis G.

. hospita Edw.

Tokunagaia rectangularıs (G.)

T. tonollii (Ross.)

Psectrocladius (Psectrocladius) psilopterus K.

P. (P.) brehmi K.

P. (P.) edwardsi Br.

P. (P.) limbatellus (Holm.)

P. (Monospectrocladius) octomaculatus Wülk.

P. (M.) schlienzi Wülk.

P. (Allopectrocladius) gr. dılatatus

Rheocricotopus (Psilocricotopus) atrıpes (K.)

R. (P.) chalybeatus (Edw.)

R. (P.) glabricollis (Mg.)

R. (Rheocricotopus) effusus (Walk.)

R. (R.) fuscipes (K.)

Paracricotopus nıger (K.)

Nanocladius balticus Pal.

N. bicolor (Zett.)

N. rectinervis (K.)

N. spiniplenus Saether

Parorthocladius nudipennis (K.)

Synorthocladius semivirens (K.)

Orthocladius (Eudactylocladins) fuscimanus (K.)

O. (E.) gelidus (K.)

nyyubhbmyhmnmtmmntmnmn&m m m m &

Et t+t+t+Ht+tHtHr ++

"+ +++

N Ne NM NM Frl MM Sr ar

"rer Va Me ara

NM rel

O. (E.) olivaceus (K.)

O. (E.) sp. A

Orthocladius (Euorthocladius) ashei Soponis

O. (E.) frigidus (Zett.)

O. (E.) Inteipes (G.)

O. (E.) rivicola (K.)

O. (E.) rıvulorum (K.)

O. (E.) saxosus (Tok.)

O. (E.) thienemanni (K.)

Orthocladius (Orthocladius) excavatus Br.

O. (O.) rubicundus (Mg.)

O. (O.) tubicula K.

O. (O.) wetterensis Br.

O. (O.) sp. A Pinder

O. (O.) Pe 1 Langhton

O. (O.) Pe 10 Langhton

Symposiocladius lignicola (K.)

Acricotopus lucens (Zett.)

Paratrichocladius rufiventris (Mg.)

P. skirwithensis (E.dw.)

ESP

Paracladius conversus (Walk.)

P. alpıcola (Zett.)

Halocladius stagnorum (G.)

H. varıans (Staeg.)

Cricotopus (Cricotopus) polarıs K.

C. (C.) tibialis (Mg.)

C. (C.) fuscus (K.)

C. (C.) tremulus (L.)

C. (C.) annulator G.

©. (C.) curtus Hırv.

C. (C.) pulchripes Verr.

C. (C.) triannulatus (Macq.)

C. (C.) alindraceus (K.)

C. (C.) patens Hirv.

C. (C.) festivellus (K.)

C. (C.) bicinctus (Mg.)

©. (C.) vierriensis G.

C. (C.) trıfascia Edw.

Cricotopus (Isocladius) ornatus (Mg.)

C. (1.) sylvestris (Fabr.)

C. (I.) glacıalis Edw.

C. (I.) trıfasciatus (Mg.)

C. (I.) intersectus (Staeg.)

C. (I.) tricinctus (Mg.)

Hydrobaenus distylus (K.)

Zalutschia tatrica (Pag.)

Metriocnemus fuscipes (Mg.)

M. hirticollis (Staeg.)

M. hygropetricus K.

Thienemannia gracılis K.

Chaetocladius dissipatus (Edw.)

C. gelidus Br.

C. laminatus Br.

C. perennis (Mg.)

> | > 69

SH ++t+t+ttHHtHmm HI

"mmmMm

ı m +Mm+ mM |

C. setilobus Marc.

C. suecicus (K.)

Paratrissocladius fluviatılis (G.)

Heterotrissocladius marcidus (Walk.)

H. grimshavi (Edw.)

Parametriocnemus stylatus (K.)

P. boreoalpinus Gow.

P. eochvus Saether

Paraphaenocladins impensus (Walk.)

Parakiefferiella gracillima (K.)

P. bathophila (K.)

P. coronata (Edw.)

Epoicocladius sp. A

Rheosmittia spinicornis Br.

Krenosmittia boreoalpina (G.)

K. camptophleps (Edw.)

K. sp. A (?=hispanica Wülk.)

Stilocladins montanus Ross.

Heleniella dorieri Ser.-Tos.

H. ornaticollis (Edw.)

TISPEN

Parachaetocladius abnobaenus Wülk.

Limnophyes bidumus Saether n. sp.

(Saether in press)

L. minımus (Mg.)

L. natalensıs K.

Psendorthocladius curtistylus (G.)

Gymnometriocnemus subnudus Edw.

G. volitans (G.)

Bryophaenocladius aestivus Br.

. llımbatus Edw.

. inconstans (Br.)

. scanıcus Br.

.subvernalis (Edw.)

. tuberculatus (Edw.)

. vernalis (G.)

Smittia aterrima (Mg.)

S. edwardsi G.

S. foliacea (K.)

5. giordani-soikai Marc.

S. gridellii Marc.

S. httorella G.

S. malarodaı Marc.

S. nudipennis G.

S. pratorum G.

Parasmittia carıinata Str.

Camptocladius stercorarins (d. Geer.)

Mesosmittia flexuella (Edw.)

Psendosmittia angusta Edw.

P. d’anconai (Marc.)

P. gracilis (G.)

P. holsata T. & Str.

P. mathildae Albu

P. recta Edw.

P. subtrilobata (Freem.)

Du u u u I

196

£ P.tnlobata (Edw.)

* Georthocladius sp. A

* Thienemanniella clavicornis K.

T. obscura Br.

= T.partita Schlee

= Corynoneura coronata (Edw.)

+ C. edwardsi Br.

= C. fittkani Schlee

= C.lacustris Edw.

C. lobata Edw.

+ C. scutellata Winn.

Chironominae

Tanytarsıni

Zavrelia pentatoma K.

Zavreliella marmorata (v.d. W.)

Stempellinella brevis Edw.

Stempellina bausei (K.)

S. subglabripennis Br.

Neozavrelia fuldensis Fitt.

Tanytarsus bathophilus (K.)

T. brundini Lind.

T. buchonius Reiss & Fitt.

T. chinyensis G.

T. curticornis K.

T. eminulus Walk.

T. fimbriatus Reiss & Fitt.

T. gracilentus Holm.

T. heusdensis G.

T. horni G.

T. lestagei G.

T. lugens K.

T. miriforceps K.

T. nemorosus Edw.

T. nigricollis G.

T. pallidicornis Walk.

T. recurvatus Br.

T. sylvaticus v. d. W.

T. usmaensıs Pag.

Virgatanytarsus maroccanus (Kügler & Reiss)

V. triangularis (G.)

Stilotanytarsus inguilinus Krüger

Cladotanytarsus atridorsum (K.)

C. mancus (Walk.)

Rheotanytarsus curtistylus G.

R. montanus Lehm.

R. muscicola K.

R. nigricauda Fitt.

R. pentapoda K.

R. photophilus G.

R. ringei Lehm.

Paratanytarsus austriacus K.

P. bituberculatus (Edw.)

Ta en Es die a a Et a, ren

TI „ca > BerEGe 3> 50 \> BOTBRE ı> Bo Base Tao BE 30 ZT „0 Zur Su zZ

P. confusus Pal.

P. inopertus (Walk.)

P. lauterborni K.

P. mediterraneus Reiss & Sawedal

P. penicıllatus G.

P. tenellulus (G.)

Micropsectra (Lauterbornia) coracına K.

Micropsectra apposita (Walk.

M. atrofasciata K.

attennata Reiss

bidentata G.

clastrieri Reiss

groenlandica And.

Juncı (Mg.)

lindrothi G.

miki Marc.

notescens (Walk.)

pharetrophora Fitt.

recurvata G.

roseiventris K.

tenellula G.

Pseudochironomini

Pseudochironomus prasinatus (Staeg.)

Chironominıi

*+ +1 +mm

"++ ++++++ 1 + Pmmm +

Mm ++++mM

Pagastiella orophila (Edw.)

Paratendipes albimanus (Mg.)

P. nubilus (Mg.)

P. plebejus (Mg.)

Microtendipes britteni Edw.

M. chloris K.

M. diffinis Edw.

M. pedellus (de Geer)

M. tarsalıs (Walk.)

Nilothauma (= Kribioxenus) brayi (G.)

*Paralauterborniella nigrohalteralis (Mall.)

Phaenopsectra flavipes (Mg.)

P. punctipes (Wiedemann)

Polypedilum (Pentapedilum) nubens Edw.

P. (P.) sordens (v. d. W.)

Polypedilum (Polypedilum) acutum K.

P. (P.) albicorne (Mg.)

P. (P.) convictum (Walk.)

P. (P.) cultellatum G.

P. (P.) laetum (Mg.)

P. (P.) nubeculosum (Mg.)

P. (P.) pedestre (Mg.)

P. (P.) nubifer (?= aberrans Chern.) (Skuse)

Polypedilum (Tripodura) acıfer Townes

P. (T.) aegyptinm (?=pruina Freeman) K.

P. (T.) apfelbecki (?= elongatum Albu) (Strobl.)

P. (T.) bicrenatum K.

P. (T.) breviantennatum Chern.

£ P.(T.) pullum (Zett.) £ Einfeldia carbonarıa Mg.

+ P.(T.) quadriguttatum K. = E.longıpes (Staeg.)

+ P.(T.) scalaenum Schr. £ E. gr. pagana

+ Endochironomus albipennis (Mg.) * _ Sergentia sp.

£ E.dıispar (Mg.) + Xenochironomus xenolabis K.

£ E.lepidus (Mg.) £ Cladopelma edwardsi Krus.

+ E.tendens Fabr. + C. virescens (Mg.)

* Stictochironomus pictulus (Mg.) + C. viridula (Fabr.)

+ Stenochironomus ranzül Ross. + Cryptotendipes holsatus Lenz

£ 5. gibbus Fabr. + Microchironomus tener K.

= S$. spatuliger K. + Parachironomus arcuatus G.

* Fleuria lacustris K. + P. longiforceps K.

= Dicrotendipes lobiger K. = P. monochromus (v.d. W.)

+ D.nervosus (Staeg.) + P.panılis (Walk.)

+ D. notatus (Mg.) EN Pavaras@-

= D. peringueyanus Freem. + P. vitiosus G.

£ D.pulsus (Walk.) + Paracladopelma camptolabis K.

+ D.tritomus K. £ P.nigritula G.

* Glyptotendipes gripekonveni K. * Genus near Paracladopelma

G. pallens (Mg.) + Harnischia angularis Albu & Botn.

" G.paripes Edw. + AH. curtilamellata (Mall.)

£ _G. severmni G. + AH. fuscimana (K.)

+ Camptochironomus pallidivittatus (Mall.) £ Demicryptochironomus vulneratus (Zett.)

+ Chironomus annularius (Mg.) + Cryptochironomus albofasciatus (Staeg.)

+ C.anthracinus Zett. + C. defectus K.

+ €. calipterus K. + C.obreptans (Walk.)

£ C. cingulatus (Mg.) + C.rostratus K.

£__C. dorsalis (Mg.) + C. supplicans (Mg.)

= C.melanotus Keyl + “Cryptochironomus” sp. Pagast

+ C.obtusidens G. + Saetheria sp. 1 Jackson

+ C.plumosus L. £ Chernovskiia macrocera (Chern.)

+ C.riparius Mg. + Genus near Robackia

+ C. salinarius K. + Robackia pilicauda Saether

+ Kiefferulus tendipediformis G. = Beckidia tethys (Townes)

+ Halliella noctivaga K.

Table 2 summarizes the present knowledge of Chironomids in Italy. 76 species new to the Italian

fauna and 45 dubious determinations that require further study and additional material emphasize that

there is still room for knowledge about Italian Chironomids to progress.

Most species are widely spread in the Palearctic region, some have more restricted areas of distribu-

tion. This is due to their ecology more than to zoogeographical reasons. Cold-stenothermal species are

restricted to highlands (most Diamesinae and many Orthocladiinae): most of these species are wide-

spread in the Alps (Region 4 in Fırrkau & Reıss 1978), many are also present in cold countries in

Northern-Europe (Fırrkau & Reıss 1978). Many species found in the Alps are not present ın cold wa-

ters in the Apennines, but this may be because of insufficient sampling. A lack of glaciers and of very

cold water in the Apennines might be other reasons.

Three species (Syndiamesa nigra, Tokunagaia tonollii and Stilocladins montanus) were found on the

southern side of the Italian Alps, but not on the northern side. $. montanus is also present in the Apen-

nines (Rossaro 1984). This supports the existence of cold-stenothermal species with an endemic distri-

bution in the Alps or with a wider southern distribution, as there are known to be for other species in

the Mediterranean area. For example, Diamesa lavillei and D. thomasi are known from the Pyrenees

only (Serra-Tosıo 1973), whereas D. veletensis was previously known only in the Sierra Nevada

197

(Spain) but was then captured in the Atlas mountains (Morocco) and in Mongolıa, suggesting it has a

wider distribution in southern and oriental parts of the Palearctic Region (SErrA-Tosıo 1983). The cap-

ture of another species of Stilocladins (S. clinopecten Saether) in the southeastern United States favours

a Gondwanian origin for this genus (SAETHER 1983). The southern side of the Alps has large extensions

of Gondwanian origin, so this hypothesis has good geological support.

Table 2: Summary of the present information about Chironomids in Italy:

+ taxa well identified, deposited in the author’s collection

= species new to the Italian fauna

£ species belonging to the Italian fauna according to the literature but not present in the author’s

collection

total: sum of +, =,£

species only tentatively identified because of a lack of adult males, ill-preserved material or species

belonging to genera that require revision

+ = = total

Tanypodinae 21 8 13 42 3

Diamesinae 2a 2 0 23 4

Prodiamesinae 3 0 0 3 1

Orthocladiinae 82 47 14 143 27

Tanytarsını 26 11 22 59 1

Pseudochironomini ) ) 0 ) 1

Chironominıi 61 8 20 89 8

Total 214 76 69 359 45

Reıss (1968) did not find any differences in Chironomid species composition in lakes on the nor-

thern and southern sides of the Alps, suggesting that the Alps are not a zoogeographical barrier for

Chironomids. This does not contradict our findings. Indeed Chironomid species from the upper rea-

ches of the mountains and their springs (Brunpın 1966) have more chance of showing an endemic dis-

tribution than species from lakes. In any case the number of endemic species seems to be very low.

The percentages of Mediterranean, Afro- and Panpaleotropical species in Italy are also of interest.

The Chironomids from the Mediterranean area have been studied only recently, but rather intensively

in recent years. According to Reıss (1977), the Mediterranean fauna contains 22.7 % of Palearctic Me-

diterranean species and about 9% of Ethiopian species. PrAat (1979, 1980), MoUBAYED & LavirLe (1983)

and Rrıss (1985, 1986) give lists of species from single countries within the Mediterranean area. There

are low percentages of endemic Mediterranean species and lower percentages of Afro- and Panpaleo-

tropical species in most cases, with a dominance of Palearctic species. Even lower percentages are

known in Italy. Two species with a Panpaleotropical distribution (Reıss 1985, 1986) that extend their

presence to the Mediterranean region are restricted to southern areas of Italy. These are Chironomus

calipterus, found in Sardinia, and Dicrotendipes peringueyanus in Sicily. Another species with a Pan-

paleotropical distribution, Polypedilum nubifer, is probably present in northern Italy, in the Po and

Adige rivers (Nocenrinı, 1985), but unfortunately only larvae are available and species determination

is uncertain. On the other hand, some species with an Afrotropical distribution are definitely present

in northern Italy: they are Rheotanytarsus montanus, Polypedilum aegyptium and Stenochironomus

spatuliger. Two other Afrotropical species known to occur in Italy are Psendosmittia subtrilobata

from Sardinia and Tanytarsus horni from Sicily (Reıss 1977). R. montanus was collected from the Ti-

cino (Lombardia) and the Potenza (Marche) (stations 3 and 10, Figure 1). It fits all the details in the

description of Lenmann (1979) except for the A. R., which is larger (= 0.8). It does not fit the descrip-

tions of the other European species. P. aegyptium comes from the Po river, 5. spatuliger from the

Mantova lakes (Figure 1). The halobiont species Haliella noctivaga and Halocladius stagnorum were

198

included in the Mediterranean faunal component by Reıss (1977). Other endemic Mediterranean spe-

cies captured in Italy are: Paratanytarsus mediterraneus, Tanytarsus maroccanus, Polypedilum agıfer

and Harnischia angularıs.

It is interesting to note that five species reported as Nearctic have been found in Italy: Rheopelopia

acra, Euryhapsis annuliventris, Nanocladius spiniplenus, Parametriocnemus eoclivus and Saetheria

sp. 1 Jackson. According to this, Holarctie distribution must be assigned to them.

In any case, the lack of knowledge or uncertain information for large areas suggests caution in draw-

ing conclusions about geographical distribution. Very often new findings extend the distribution area

of species very greatly. For example, T. tonollii was recently captured in Norway (Tuiskunen pers.

comm.). More extensive knowledge ofthe Chironomids in Italy is very desiderable, because of its stra-

tegic position between the Palearctic and Afrotropical regions.

There is very scanty information for central and southern Italy at present. More detailed knowledge

about these areas will probably raise the number of Mediterranean and Afrotropical species. Another

very interesting field is the faunal composition in the cold waters on the southern side of the Alps and

in the Apennines: it is important to confirm or refute the existence of cold-stenothermal Chironomid

species with a southern distribution.

Acknowledgements

I am much indebted to Dr. Reiss, Zoologische Staatssammlung, München, and to Prof. Saether, Zoologie Mu-

seum, Bergen, for having examined species.

Literature

Aıı, A. & MAJoRI, G. 1984: A short-term investigation of chironomid midge (Diptera: Chironomidae) problem in

saltwater lakes of Orbetello, Grosseto, Italy. — Mosq. News 44: 17-21.

BRUNDIN, L. 1966: Transantarctic relationships and their significance, as evidenced by Chironomid midges. —

Kungl. Sv. Vetenskapsakad. Handl. 11: 1-472.

FERRARESE, U. 1982: Chironomid research in Italy. — Chironomus 2(4): 29— 33.

—— 1983: Chironomidi, 3 (Diptera, Chironomidae: Tanypodinae). Guide per il riconoscimento delle specie anı-

mali delle acque interne italıane. - CNR AQ/1/204 26: 1-67.

FERRARESE, U. & ROssARO, B. 1981: Chironomidi, 1 (Diptera, Chironomidae: Generalitä, Diamesinae, Prodiame-

sinae). Guide per il riconoscimento delle specie anımali delle acque interne italiane. - CNR AQ/1/129 12:

197

FITTKAU, E. J. & F. Reıss 1978: Chironomidae. — In Illies J. (ed.) Limnofauna Europea: 404440. G. Fischer, Stutt-

gart.

LEHMANN, J. 1979: Chironomidae (Diptera) aus Fließgewässern Zentralafrikas (Systematik, Ökologie, Verbrei-

tung und Produktionsbiologie). Teil I: Kivu-Gebiet, Ostzaire. — Spixiana Suppl. 3: 1—144.

MOUBAYED Z. & LAVILLE, H. 1983: Les Chironomides (Diptera) du Liban. I. Premier inventaire faunistique. —

Annls. Limnol. 19: 219— 228.

NOcENTINI, A. 1985: Chironomidi, 4 (Diptera, Chironomidae: Chironominae, larve. Guide per il riconoscimento

delle specie anımalı delle acque interne ıtaliane. - CNR AQ/1/233 29: 1— 186.

PRAT, N. 1979: Quironömidos de los embalses espanoles (1a parte). — Graellsia 33: 37—96.

—— 1980: Quironömidos de los embalses espanoles (2a parte). — Graellsia 34: 59—119.

Reıss, F. 1968: Verbreitung lakustrischer Chironomiden (Diptera) des Alpengebietes. — Ann. Zool. Fenn. 5:

INY)-125,

—— 1977: Verbreitungsmuster bei paläarktischen Chironomidenarten (Diptera, Chironomidae). — Spixiana 1:

—— 1985: A contribution to the zoogeography of the turkish Chironomidae (Diptera). — Israel Journal of Ento-

mology. 19: 161-170.

—- 1986: Ein Beitrag zur Chironomidenfauna Syriens (Diptera, Chironomidae). — Entomofauna 7: 153— 168.

199

Rossaro, B. 1979: Contributo alla conoscenza delle Orthocladiinae e Diamesinae italiane (Diptera, Chironomidae)

(seconda nota). — Boll. Museo Civico Storia Naturale Verona 6: 79— 94.

—— 1982: Chironomidi, 2 (Diptera, Chironomidae: Orthocladiinae). Guide per il riconoscimento delle specie anı-

malı delle acque interne italiane. - CNR AQ/1/171, 16: 1-80.

—— 1984: Stilocladius Rossaro, 1979 reconsidered, with descriptions of the female and larva of $. montanus Ros-

saro (Diptera: Chironomidae, Orthocladiinae). — Ent. scand. 15: 185-191.

SAETHER, ©. A. 1983: Three new Species of Lopescladius Oliveira, 1967 (syn. “Cordites” Brundin, 1966, n. syn.),

with a Phylogeny of the Parakiefferiella group. — Mem. Amer. Ent. Soc. 34: 279—298.

SERRA-TOSIO, B. 1973: Ecologie et biogeographie des Diamesini d’Europe (Diptera, Chironomidae). — Trav. Lab.

Hydrobiol. Grenoble 63: 5- 175.

—— 1983: Nouveax Diamesinae de la Palearctide meridionale et orientale. — Spixiana 6: 1—26.

Prof. Dr. Bruno Rossaro

Department of Environmental Sciences

Vıa $. Sısto 20

I-67100 L’Aquila, Italıa

200

SPIXIANA Supplement 14 201 2 | München, 15. Juli 1988 ISSN 0177— 7424

Emergence patterns of chironomids in Keszthely-basin of

Lake Balaton (Hungary)

(Diptera, Chironomidae)

By György Devai

Abstract

The author tries to give a general idea of the characteristic features and patterns of the emergence of chironomids

relying upon two series of examinations of along period between March 11 and October 18, 1980 as well as between

March 15 and August 31, 1983, which were performed on the Keszthely shoreline of the biggest shallow lake of

Europe, Lake Balaton. Relying upon the comparative analysis and computer processing of the data set of the daily

observations as well as that of 53 and 147 samples of pupal exuviae collected in 1980 and 1983, respectively, the au-

thor presents the characteristic emergence types and states the main emergence periods. He analyses in detail the

essential changes which occurred in the taxonomic composition of samples of pupal exuviae between 1980 and 1983.

Introduction

One of the decisive and obviously the most spectacular moments of the life of chironomids is their

emergence. Moreover, this process has an outstanding significance also in the settling dynamics and

matter circulation of water bodies (D£fvaı et al. 1979, Devaı 1980a, 1980). Thus it is understandable

that the rough and regular interventions performed in the environs of water bodies as e. g. mosquito

extermination by aeroplane or helicopter inevitably raise the question how the chironomids are en

dangered.

In the case of Lake Balaton there was another important viewpoint to be said for the fact that mos-

quito extermination should follow the emergence dynamics and swarming peculıarities of chirono-

mids. It is well-known that land-fauna exterminated in consequence of mosquito extermination is re-

latively soon supplemented from the neighbouring regions due to the so-called vacuum-effect (SArın-

GER etal. 1984). However, at insects flying out of water, especially in the case if extermination covers

the predominant part of the shoreline (as e. g. in the case of Lake Balaton as well), the decrease of in-

dividual number can be so drastic among the animals generally assembling here that the population

sustains permanent loss through it and the standing mass can decrease to 10 or 20% of the original

number. This condition of risk especially exists in the case of Lake Balaton, the peculiar chironomid

fauna of which rests first of all on self-revival since the fauna practically cannot or can only be slightly

supplemented from the neighbouring regions failing water bodies of a similar type and size.

Accordingly it was justified from several viewpoints to investigate the emergence dynamics of the

chironomid fauna of Lake Balaton thoroughly. In my present paper I wish to give a survey about the

results of our investigations of emergence performed in 1980 and 1983.

According to the reports of the majority of papers (see e. g. Parm£n 1958, 1962, Dans 1971, STAHL

1975, LiNDEGAARD & JOnasson 1979, BUTLER 1980) the emergence of chironomid adults from the pupal-

exuviae and at the same time their emergence from the water bodies occur in periods characteristic of

201

the different species and they do not show more than little temporal shift in the subsequent years. The

emergence can be generally observed once or maximum two or three times a year and on these occa-

sions the animals emerge in great mass. At the same time certain experiences in the field (e. g. Heın &

SCHMULBACH 1972, Jonsson & SanpLunn 1975, Arı et al. 1983, 1985) or laboratory experiments (e. 8.

Danks 1978) refer to the fact that the emergence period of certain species can be significantly long-

drawn and during this long period several generations can emerge. According to the data up to now

the emergence was synchronized with one of the abiotic environmental factors or the joint effect of

two of them. Such factors can be e. g. water temperature, light conditions, air pressure or ebb and flow

(Lenz 1962, Parm£n 1962, Danks 1971, 1978, Hein & SCHMULBACH 1972, HasHımoTo 1975, HEIMBACH

1978, Bacce et al. 1980).

Material and Method

Our examinations were performed on the biggest shallow lake of Central Europe, Lake Balaton (its detailed cha-

racterization see in papers by BiRÖ 1984, and D£vA1 et al. 1984). Our main sample area out of the regions of the lake

with different water qualities was the Keszthely-basin mostly endangered by the accelerated eutrophication of

recent years.

A series of observations during a period of 236 days was performed on the shoreline of Lake Balaton at Keszthely

in 1980 in order to examine the emergence in a pragmatical way. The regular surveys were performed on the pier

of the harbour of Keszthely each day from March 11 to October 28. This point of the shoreline was proved to be

convenient from several viewpoints: the pier projects far into the open water of the lake (to a distance of 200 m), at

its end the water depth is already near the average (about 2 m) and no reeds of a large extension can be found at the

pier (in a district of approximately 500 m). In this way this site of observation and sampling was suitable to reflect

satisfactorily the conditions of the open water of almost the whole Keszthely-basin. The second series of examına-

tions of 170 days was performed in the same place from March 15 to August 31, 1983.

Relying upon our earlier experiences collections of pupal exuviae proved to be the most suitable for establishing

the frequency and intensity of emergence. Therefore parallel with the observations on the occasions of great emer-

gences we also took samples of pupal exuviae from the floating material accumulated along the pier. From this view-

point the choice of the sample area was very favourable since the peculiar flow conditions of the Keszthely-basin

(GYÖRKE et al. 1980) ensured with the greatest possibility that the pupal exuviae material characteristic of the total

water surface could drift together.

For the sampling of pupal exuviae we used a sack-like “skimming-net” of our planning. The samples were

preserved in 70% ethylalcohol in of 50-200 ml cubic capacity depending on the quantity of the collected material.

The pupal exuviae were selected and counted by means of a stereomicroscope of Technival type and of Zeiss (GDR)

make. In order to unify the processing permanent preparations were made from the pupal exuviae of different type

for an ıdentifying and comparative collection (SCHLEE 1966).

On the occasions of great emergences several hundred or sometimes several thousand pupal exuviae could often

be found in the samples. For their processing the application of the following method seemed to be expedient. After

a careful shaking a certain number of pupal exuviae was taken out of the sample with Leonhard forceps, the genera

occurring in it were determined and the number of their pupal exuviae was in the percentage of all the counted ones.

We had only two obligations regarding the quantity taken out of the sample. One of them was that the number of

examined exuviae should be above 100 and the other was that any additional pieces taken out were also counted.

The first obligation ensured that the obtained quantities should reflect roughly the proportions in the whole sample.

And by counting all the pupal exuviae taken out we avoided the mistake of conscious choice.

Relying upon our literary studies and our field experiences we chose factors from the system of factors being able

to influence the emergence and their detailed analysis was expected to result in the knowledge of phenomena and

processes producing and synchronizing the emergence. The factors investigated were the following: the real daily

mean value, maximum and minimum of the air temperature; the sum of the daily radiation and the duration of sun-

shine; the daily most frequent direction of wind; the real daily most frequent direction of wind; the real daily mean

value of the wind speed as well as the extent of its daily fluctuation; the real daily mean value of the air pressure and

the extent of its fluctuation; the daily mean value of the air humidity; the daily sums of the evaporation and preci-

pitation; the value of the morning and evening water temperature; the strength of waviness in the morning and in

202

the evening; the values of the air temperature, wind direction, wind speed and air humidity measured at 7 p. m.

These factors were measured partly by ourselves and partly the data of the Keszthely Synoptic Station of the Natio-

nal Meteorological Service as well as those of the Research Institute of the Atmosphere Physics were utilized by us,

for the conveyance of which data we express our thanks.

The nearly 10 thousands data were fixed on a magnetic band and processed by the Robotron (GDR) computer

ofR 55 M type of the Computing Centre of the L. Kossuth University. The evaluation of the data was performed

by means of the hierarchical cluster analysis programme functioning on the basis of Euclidean distances as well as

the stepwise discriminance analysis programme of BMDP. In order to increase the certainty of the processing, we

took notice first of all of the great emergence cases (indicated with double or triple motive on Figure 1) or we drew

into the processing only those of the emergences of less intensity (indicated with one motive) which could be iden-

tified regionally or could be delimited from one another temporarily with proper certainty.

Apfil_ DE

EN ? ? ? 7?

June 1m pn nn m

July m

Aug: Bi ae mer m. nn

* Ey ”

DE Se Takko En Sue *

Eu ? 2 * x *

5 10 1b 20 25 30

x small intensity * middleintensity * great intensity ? uncertainly decidable emergence

Fig. 1. Emergences in 1980 and their intensities on the basis of pupal exuviae collections.

Results and Discussion

Although the processing procedure detailed above, could not bring absolutely exact results, it made

the present opinions about the emergence dynamics and the change in taxonomic composition of the

chironomids of Lake Balaton more exact even in its first approach to a great extent. On the basıs of

our field experiences and the results of our evaluation we can draw the following main statements.

The emergence dynamics of the chironomids in Lake Balaton is almost unprecedented considering

both the long-drawn period of the emergence and the frequency of the emergences. According to our

observations in 1980 the period of emergence lasted from the middle of March until the end of October

(Fig. 1). From May to September emergence was observed on an average every second day, it was even

of middle intensity every fifth day. The main emergence periods were May 26-27, June 20-25, July

29-August 4 and August 25—27.

203

In 1983 the emergence was considered to be even more uniform, disregarding some small inter-

ruptions it lasted almost continuously. Emergence cases of great intensity were observed on 5 oc-

casions: April 11-18, June 25-July 3, July 30-August 2, August 21-26 and September 12-18. Regar-

ding the mass of the emergence the first emergence case was specially striking. It is definitely worth

mentioning that during nearly the whole month of May emergence of medium strength went on and

no doubt it was entirely at least as significant as any other very strong emergence lasting for a short

time. As a result of our observations performed on the whole territory of the Lake Balaton in

1980— 1985 we can state that significant differences can be experienced in the frequency and strength

of the small emergence cases along the total shoreline of Lake Balaton. However, according to the

experiences up to now the great emergence cases are never limited to a smaller region or any basin but

they go on simultaneously on the whole surface of the lake.

Comparing the results of the observations performed in 1980 and 1983 it can be seen that emergence

cases became definitely more frequent by 1983. It manifested itself in not only the number but also the

strength of the emergence cases. The most conspicuous sign of it was that mass emergence cases also

increased, namely with two peaks in the middle of April and in the middle of September. In the same

periods of 1980 we observed only medium emergence. The change can be obviously explained with

the significant increase of the food supply and the change in the species composition (Devat et al.

1984). The earlier peak at the end of May — due to similar causes — changed into an emergence period

of middle strength lasting almost continuously in May, which well can be brought into correlations

with the results of larval investigations (Devaı 1985). However, the three great summer emergence

DISSIMILARITY

0,00

0,300

0.200

SEO RRRORIITIIEN SRLRNISRRREITIRÄRRTOSCTETIKAFAn EN En ERS Serigl number

SIEHSFISTEETEEHKÜTESTETKIFIETERISIFELTETITCEHEHESLETSTSSIHTEETETSTS ze

SISLRITKRITSSUTESKRKTELNLNESKTSÜSATTSSEIKTKRTNTRILEISIITEES

Fig. 2. Dendrogram of pupal exuviae samples collected on occasions of important emergences in 1980.

204

periods remained the same, disregarding a shift of some days due to the actual change in weather con-

ditions.

The field experiences showed that mass emergences mainly occurred at sudden change of tempera-

ture and they seemed to be characterized roughly with the changes of meteorological factors. This hy-

pothesis was supported by the observations that mass emergence often followed the red storm signals,

however, they did not occur if wave activity was permanently strong and especially if combing waves

were observed.

Accordingly, when making an effort to understand the effects inducing or at least promoting the

emergence we analysed the examined meteorological factors successively and compared them at first

separately and later in different combinations to the emergence pattern. Naturally, in the latter case

we took into consideration that between the factual occurrence and the observation of emergence

there were significant temporal differences, depending upon the actual weather conditions (first of all

wind conditions) amounted to at least 2-3 hours, generally 6-10 hours but sometimes even 12—24

hours.

We did not get an unambiguous and comprehensive relationship in any case, not even the discrimi-

nance analyses brought satisfactory results. We only succeeded in establishing that groupings made on

the basis of empirical facts and the discriminance function may correspond each other in an acceptable

way (in approximately 70-75 % of all the cases) if we consider the values of the daily radiation and

the average air temperature or those of the water temperature in the morning. However, this relation-

ship weakens if we take notice that for example the emergence of the Chironomus balatonicus, the

most abundant and frequent chironomid species occurred at the values of the water temperature

between 10-28°C alıke.

Relying upon all these findings we drew the conclusion that in the future we have to put the maın

stress on the examination of the relationship of emergence and front situations since the separate study

of individual meteorological factors does not reflect properly the change in the weather conditions

caused by the different meteorological fronts (Kıss 1959). At the same time the results obtained called

our attention to the fact that in the peculiar weather conditions of Carpathian basin that are characte-

ristic of the environs of Lake Balaton, it is hardly probable that the start or stop of the emergence can

be attributed to only one or 2-3 factors as well. It can be rather postulated that the emergence dyna-

mics of the chironomids do not depend upon the change of a certain meteorological element but upon

the joint effect of several, often a series of factors, they can be practically interpreted as the resultant

of them; naturally with the exception of extreme cases. However, by means of demonstrating the re-

gularities still observable in the emergences as well as elucidating their causes we see the possibility of

predicting mass emergences both in region and in time and so of successfully promoting the preven-

tion of the permanent damage of fauna.

At the samples of 1980 in the percentage distribution of exuviae belonging to different genera two

main groups could be separated on the basis of the results of both comparative analyses and cluster

analysis (Fig. 2). In one of them the Chironomus exuviae dominated, their proportion was mainly

above 70 % but it always exceeded 50 %, except one sample. Four separate subgroups occurred within

this group. It was characteristic of the first subgroup that Chironomus exuviae represented 85-98 %

of all the exuviae (samples 1-31). In the other samples the remaining proportion (15-55 %) in addi-

tion to the Chironomus exuviae was represented by mainly Procladius exuviae in the second group

(samples 25-50), by Tanypus exuviae in the third group (samples 16-49) and by Cricotopus exuviae

and those belonging to other Chironominae taxa in the fourth group (samples 46-19). The samples

being present within this group constitute the most significant proportion, approximately 53 % of all

the examined samples.

The other great group shows amuch more colourful picture than the previous one and far more sub-

groups could be separated within it mainly relying upon the exuviae belonging to genera occurring in

greater proportions. On such a basis four subgroups could be recognised with more or less certainty.

205

Sıhrinmornlarmausiniyapge

30.07.1980

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12.08.1980

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tt tt tt tt tt HH HH

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Fig. 3. Different types of emergence patterns in 1980 on the basis of pupal exuviae collections.

206

Tanypus-type

30.05.1980 27.05.1980

09.06.1980

Parachironomus-

subtype subtype

Endochironomus-

Cryptochironomus —

subtype

+++t++t++++++++

++ t+t+t+t+t+t+t++t+++++ 3

++t+t+t+t++t+++++++++

++t+t+t+t+t+t+tt+tt+t+t+++H+ +

++t+t+t+t+t+t+t+t+tt+t++++++

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22.05.1980

20.06.1980

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11.08.1980

taxa

25.08.1980

Subtype of other

Cricotopus-

subtype

Polypedilum —

subtype

23.09.1980

27.04.1980

Tanypus

Other taxa

BSR:

Cricotopus

10 =

207

In the first one the relatively high percentage (between 43—81%) of Procladius exuviae (samples

11-44), in the second one that of Tanypus exuviae (samples 8-10), in the third one that of Glyptoten-

dipes exuviae (samples 7—33) and the smaller quantity of Chironomus exuviae (in the majority of cases

under 25%) in all the three could be considered as a separative feature. These subgroups represent

11%, 6% and 11% of allthe examined samples, respectively. The other samples drawn in the fourth

group are of very varied composition. In fact this constitutes an independent subgroup because of its

separation from the former ones. The samples grouped here in addition to the Chironomus exuviae of

the proportion mainly above 30% either contain exuviae that belong to some genera not mentioned

in the foregoing in a proportion of approximately 30% or their composition according to genera

shows a very varied picture. These cases constitute about 19% of all the examined samples. The circle

diagrams shown in Figure 3 illustrate some of the typical cases of these emergence patterns.

The summarized evaluation of the frequency of occurrence of pupal exuviae belonging to genera

that were detectable in a significant amount was performed in two different ways from the data of the

53 selected samples. On the one hand we examined in what proportion the pupal exuviae were present

by genera compared to the total amount of counted pupal exuviae. For the genus Chironomus this va-

lue was 51 %, for Procladins it was 15%, for Glyptotendipes ıt was 11% and for Tanypus it was 7%

i. e. these genera constituted 84% of all the pupal exuviae. Altogether only 16% was the proportion

of the other genera under the frequency limit of 5% (Cryptochironomus, Endochironomus, Microchi-

ronomus, Parachironomus, Polypedilum, Cricotopus).

Somewhat different picture was obtained if the frequency of occurrence was evaluated only on the

basis of presence and absence independently of quantitative results. Although the order of the first

four species did not change since among the pupal exuviae found in the samples Chironomus occurred

in 96% of the cases, Procladius occurred in 79%, Glyptotendipes occurred in 75% and Tanypus

occurred in 60% of all cases. However, in this summarizing the presence of pupal exuviae belonging

to other genera was also significant since they occurred in 96 % of the samples.

An opportunity presented itself for an even more comprehensive evaluation by combining the two

procedures i. e. by establishing the presence or absence above or under certain quantitative limits. The

following results were achieved by this method:

The percentage frequency At

of occurrence A B E D E

genera (in %)

In case of a quantity above 75 % 32 4 ) 0 0

In case of a quantity above 50 % 53 11 9 4 4

In case of a quantity above 25 % 76 23 13 8 28

In case of a quantity above 10% 87 36 25 25 47

In case of a quantity above 5% 92 49 34 34 68

In case of a quantity under 5% 4 30 41 26 28

The frequency of the cases when the pupal

exuviae belonging to the given genus did

not occur in the sample 4 21 25 40 4

(A =Chironomus, B =Procladius, C = Glyptotendipes, D = Tanypus, E = Other taxa)

It can be seen from the data that already in 1980 the Chironomus exuviae could be considered as

most significant not only regarding the number of occasions but also quantitatively since they proved

to be absolute dominant in more than half of the examined cases and they were the determinants of

type (in an amount of above 25 %) in more than three-quarters of them. The Procladius exuviae could

be considered to a much less extent but still unambiguously characteristic as regards both the fre-

quency of occurrence and the quantitative conditions.

208

We got pictures fairly similar to each other with the exuviae belonging to the Glyptotendipes and

Tanypus genera, which occurred as the determinants of type in a part of the cases and as absolute do-

minants in some cases. However, all together they did not prove to be more significant than Procladius

itself. The difference between them was fırst of all that the Glyptotendipes exuviae were present more

times in great (above 50 and 25%) and small (under 5%) amounts than Tanypus exuviae, however, the

latter were more frequently absent in the samples.

The total amount of exuviae belonging to nearly 8-10 other genera could not be considered as neg-

ligible in spite of the fact that they have individually never proved to be absolute dominants, they oc-

curred as the determinants of type in only 6 cases even the number of the cases when they were sepa-

rately detectable in an amount above 10% was only 18.

At last, relying upon the data of 1980 we wished to see whether the differences in the distribution

according to genera corresponded to any temporal emergence pattern. Unfortunately relying upon

the processed samples, satisfactory answer cannot be given to this question. We could state with a

more or less certainty all in allthat Chironomus dominance could be demonstrated continuously while

a proportion exceeding the average was observed for Procladius exuviae first of all from the end of May

to the middle of June and in August, for Tanypus exuviae mainly in June and in the first half of Septem-

ber and for Glyptotendipes exuviae in the middle of May, at the end and beginning of June as well as

after that on several occasions during certain shorter periods (e. g. about July 14, 26, August 10, 25,

30, September 8). The samples in which the representatives of the other genera were present in a signi-

ficant amount, sporadically originated from the whole emergence period but the samples collected in

May and September seem to be at least relatively somewhat more frequent among them.

If we compare the data of the series of samples from 1983 to the results of 1980, the following im-

portant differences come out. In the percentage summarizing according to the amount of pupal

exuviae the proportion of Chironomus exuviae strikingly increased (by 16%) and that of the Procla-

dins, Glyptotendipes and Tanypus exuviae significantly decreased (by 4-6%). The proportion of the

exuviae belonging to other genera did not change significantly (only decreased by 2%). These shifts

of the proportion totally coincide with the experiences of larval examinations (D£vaı 1985).

The examination of the frequency of occurrence on the basis of presence and absence brought

somewhat different results compared to the foregoing. A decrease was experienced only for Glypto-

tendipes exuviae and it was only slight (altogether 7%). With the exception of that arise could be esta-

blished in the case of all the genera.

The following data obtained by combining these two methods let us conclude to the real relations

most of all.

The percentage frequency At

of occurrence A B C D E

genera (in %)

In case of an amount above 75 % 29 0) ) 0 0

In case of an amount above 50 % 90 0 (6) 0 0

In case of an amount above 25 % 100 1 1 0) 7

In case of an amount above 10% 100 57 10 5 65

In case of an amount above 5% 100 88 30 28 90

In case of an amount under 5% 0 11 38 54 10

The frequency of the cases when the pupal

exuviae belonging to a given genus did not

oceur in the sample 0 dc 32 18 0

(A =Chironomus, B =Procladius, C = Glyptotendipes, D = Tanypus, E = Other taxa)

These results unambiguously show the almost absolute dominance of Chironomus exuviae and the

significant depression, of the proportion of all other genera. For example it is characteristic that ın

209

1983 in the case of the Keszthely basın we did not find an example of the Procladius dominance remin-

ding of the picture of the original fauna in Lake Balaton and still detectable on some occasions in 1980.

The tendencies recognized here support our ideas which we formed of the transformation of the sedi-

ment-dwelling chironomid fauna of Lake Balaton and of the cases and trends of the resulted changes

(Devaı 1985).

When we wanted to determine the emergence periods for the quantitatively significant genera on

the basıs of the samples taken continuously in 1983, maybe we got into an even more difficult situation

than in the case of sporadically taken samples of 1980. Namely these data really give evidence of the

fact that emergence lasts almost continuously for the representatives of almost all the genera.

Although the strength of emergence changes and it can decrease on several occasions for periods of

some days or at most one or two weeks, however, it only rarely ceases entirely on these dates.

Acknowledgements

The sampling and the processing of the collected material were performed with the financial support of the Cen-

tral Research Fund of the Hungarian Academy of Sciences and that of the National Research Programme (sign A-

12) coordinated by the National Authority for Environment Protection and Nature Conservation. Iowe my thanks

to Professor P. Jakucs (Department of Ecology, L. Kossuth University) for the continuous support of my work. I

express my thanks to the members of our Balaton-research group (first of all D. Rönai, Dr. F. Gyulai, Z. Gyurko-

vics and I. Zsupän as well as Zs. Enyedi, Dr. A. Koväcs, J. Moldovän, E. Nagy, Ä. Petrö and I. Molnär). For their

co-operation in the evaluation of the results I am grateful to my co-workers Dr. I. Devai, Dr. G. Lörincz and B.

Töthmeresz. In arranging my paper Zs. Bälint, Zs. Dienes, M. Miskolczi and E. Szilagyi were very helpful, for

which I offer my thanks to them. Iowe special thanks Dr. I. Meszäros for the conscientious professional and lingu-

istic revision of my manuscript.

Literature

Auı, A., STANLEY, B. H. & G. Majorı, 1985: Daily abundance patterns of pestiferous Chironomidae (Diptera) in

an urban lakefront in Central Florida. — Environ. Entomol. 14: 780-784

——, STANLEY, B. H. & $S. R. STAFFORD 1983: Short-term daily emergence of adult midges (Diptera: Chironomi-

dae) from a natural lake and an artıificial reservoir. — Environ. Entomol. 12: 765-767

BAGGE, P., ILus, E. & L. PAAsıvIRTA 1980: Emergence of insects (esp. Diptera, Chironomidae) at different depths

in the archipelago of Lovisa (Gulf of Finland) in 1971. — Ann. Ent. Fenn. 46(4): 89— 100

Birö, P. 1984: Lake Balaton: a shallow Pannonian water in the Carpathian Basin. — In: TAuB, F. B. (ed.) Lakes and

reservoirs. — Elsevier Sci. Publ. B. V., Amsterdam: 231—245

BUTLER, M. G. 1980: Emergence phenologies of some arctic Alaskan Chironomidae. — In: MURRAY, D.E. (ed.)

Chironomidae — ecology, systematics, cytology and physiology. — Pergamon Press, Oxford: 307—314

Danks, H. V. 1971: Life history and biology of Einfeldia synchrona (Diptera: Chironomidae). — Can. Ent. 103:

1597—1606

—— 1978: Some effects of photoperiod, temperature, and food on emergence in three species of Chironomidae

(Diptera). — Can. Ent. 110: 289— 300

D£vaı, Gy. 1980a: Investigations of the sediment-dwelling chironomid (Diptera: Chironomidae) fauna of Lake Ba-

laton. Workshop comment. — In: KARPATI, I. (ed.) New results concerning the research of Lake Balaton I. —

MTA VEAB Monogräfiäi 12: 82-84 (in Hung.)

—— 1980b: Vorstudien zur Bedeutung der sedimentbewohnenden Zuckmücken im Stoffhaushalt des Balatonsees

(Ungarn). — In: MURRAY, D. E. (ed.) Chironomidae — ecology, systematics, cytology and physiology. — Per-

gamon Press, Oxford: 269— 273.

—— (ed.) 1984: Studies of the ecological effects of Lake Balaton and River Zala sediments on chironomids (Diptera:

Chironomidae). — Acta Biol. Debr. Oecol. Hung. 1: 1- 183, Tab. 1-7, Fig. 1-59

—— 1985: Ecological background and importance of the change of chironomid fauna in the shallow Lake Balaton

(Hungary). — IXth Int. Symp. on Chironomidae, Bergen — Museum of Zoology, Bergen, 38-39

210

——, D£vaı, I., KovAcs, A. & I. MOLNAR 1979: Preliminary studies on the significance of the sediment-dwelling

chironomids in the matter circulation of Lake Balaton. — National Congress of the Hungarian Hydrological

Society, Keszthely — MHT, Budapest, II/A/11: 1—22 (in Hung.)

GYÖRKE, O., MUSZKALAY, L. & L. RAköczı, 1980: Morphological characteristics of Lake Balaton, the movement

of lakewater. — In: BARANYI, S. (ed.) Research and regulation of Lake Balaton. — Vıruk1 Közl. 27: 175—210

(in Hung.)

HASHIMOTO, H. 1975: Seasonal emergence of Clunio aquılonius Tokunaga (Diptera: Chironomidae). — Kontyü

43: 4954

HEIMBACH, F. 1978: Sympatric species, Clunio marınus Hal. and Cl. balticus n. sp. (Dipt., Chironomidae), isolated

by differences in diel emergence time. — Oecologia 32: 195—202

HEIN, J. & J. C. SCHMULBACH 1972: Larval development and adult emergence periodicity of Chironomus pallidivit-

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Kıss, I. 1959: The high increase (mass production) of plant microorganisms, as manifestation of bioindication. —:

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PALMEN, E. 1958: Periodic emergence in some North European chironomids. — Verh. Internat. Ver. Limnol. 13:

817-822

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mus crassiforceps K. — Ann. Ent. Fenn. 28: 137—168

SARINGER, GY., TOTH, $., DEVAI, GY., FORRÖ, L., VASÄRHELYI, T., PonYı, J. & G. Körüs 1984: The experiences

concerning mosquito control around Lake Balaton. — Termeszet Viläga 115 (7): 294—297 (in Hung.)

SCHLEE, D. 1966: Präparation und Ermittlung von Meßwerten an Chironomiden (Diptera). — Gewäss. Abwäss.

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nomidae). — Ent. News 86: 69-72

Dr. György Devaı,

Department of Ecology, L. Kossuth University,

H-4010 Debrecen, Hungary

211

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SPIXITANA | Supplement 14 213—238 München, 15. Juli 1988 ISSN 0177-7424

Corynoneura brundini spec. nov. Ein Beitrag zur Systematik der

Gattung Corynoneura

(Diptera, Chironomidae)

Von Mauri und Elina Hirvenoja

Zusammenfassung

Für Corynoneura scutellata, der Typusart der Gattung Corynoneura Winn. wird ein Neotypus designiert und die

systematische Stellung der Art geklärt. Arten der Untergattungen Corynoneura s. str. und Paracorynoneura im

Sinne von Goetghebuer (1939) werden in die Corynoneura scutellata-, C. edwardsi- und C. carriana-Gruppen ein-

geordnet. Corynoneura brundini spec. nov. ist nächstverwandt mit Corynoneura edwardsi Brund.

Einführung

In der älteren Literatur findet man eine Reihe von Arten und auch höheren Taxa, die in den Verwandtschaftskreis

der Gattung Corynoneura gehören. Bis heute liegt keine umfassende Revision der europäischen Arten vor, so daß

eine Artbestimmung vielfach nur mit Vorbehalt möglich ist. In der Limnofauna Europaea sind von FITTKAU und

Reıss (1978) 28 Artnamen aufgeführt. Es handelt sich um jene „Arten“, die, meist beschrieben von Kieffer, schon

im „LINDNER“ von GOETGHEBUER (1939) aufgelistet worden sind.

Die taxonomischen Probleme mit der Gattung Corynoneura beginnen bereits mit der Originalbeschreibung der

Gattung von WINNERTZ (1846: 12):

„Fam. Tıp. culiciformes. Antennae prorectae, fıliformes; marıs novem-articulatae, plumosae, inferioribus octo

ovatıs, ultimo majore, clavato, apice pılis coronato; feminae quinque-articulatae, pilosae, inferioribus quatuor ova-

tıx, ultimo eongato, pilis brevioribus coronato quam marıs. Palpi incurvi, 4-articulati; articulo ultimo elongato.

Oculi subrotundi, intus paullulum emarginati. Ocelli nulli. Alae lanceolatae, nudae, defelexae. Costa brevissima, in

clavae formam dılatata; marıs circiter ad quartam partem, feminae ad medium marginis antıci pertinens.“

Aus der ausführlichen illustrierten Diagnose auf Deutsch geht deutlich hervor, daß es sich um eine Chironomide

handelt, die eine sehr charakteristische 10gliedrige Antenne (Pedicellus mitgezählt) mit einer großen und unge-

wöhnlichen apikalen „Rosette“ besitzt. Weiter ist zu ersehen, daß es sich bei der Gattungsdiagnose eindeutig um

die Art Corynoneura minuta Winnertz (1846: 13) handelt. EDwARrDSs (1929: 367—368) hat das einzige Männchen,

das der Originalbeschreibung von C. minuta zu Grunde lag, gesehen. Er bestätigt die Anzahl der Antennenglieder,

wie sie bei der Originalbeschreibung gegeben ist. Die Abbildungen bei Winnertz sind außerdem so gut, daß die Art

leicht erkannt werden kann. Im Naturhistorischen Museum in Wien befindet sich ein als Typus markiertes Weib-

chen, das die Identität mit dieser Art bestätigt.

COQUILLETT (1910: 528) faßte nach den damals eingeführten nomenklatorischen Regeln Corynoneura scutellata

als Typus auf, da sie die erstaufgeführte, wenn auch ganz ungenügend beschriebene Art in der Arbeit von Wın-

NERTZ (1846: 13) ıst. WINNERTZ erwähnt u. a., daß die Rosette von C. scutellata kurz ist.

Die dritte Corynoneura-Art, C. lemnae beschrieb SCHINER (in FRAUENFELD 1866: 974). Frauenfeld selbst er-

gänzt die Beschreibung dieser Art mit Angaben über deren Ökologie und Metamorphosestadien. GOETGHEBUER

(1939: 7) sieht C. lemnae als ein nomen dubium an. C. lemnae ist aber auf Grund des Originalmaterials (vgl. weiter

unten) gut definierbar. SCHINER versucht bei seiner Beschreibung von C. lemnae diese Art mit C. scutellata zu ver-

gleichen. Seine Auffassung von C. scutellata ist belegt mit einem Weibchen seiner Sammlung, heute aufbewahrt im

Naturhistorischen Museum in Wien. Dieses Exemplar ist jedoch ein Weibchen von C. coronata Edw. einer Artmit

einer deutlichen Antennenrosette.

213

Von KıEFFER (1906: 328) wurde das O’ einer weiteren Art, C. bitensis, beschrieben. Nach der damaligen Vorstel-

lung von Kieffer können die Antennen der Männchen von Corynoneura 10- oder 11gliedrig (wie bei C. bitensis)

sein, wobei das letzte Glied etwa ebenso lang ist, wie die vorhergegangenen zusammen. Die Beschreibung bei Kief-

fer ist ungenügend. Da aber das 2. Palpenglied beim Männchen von C. bitensis länger als breit sein soll, kann es sich

auch um eine Art der später beschriebenen Gattung Thienemanniella Kieff. handeln. Kieffer vergleicht die Art bi-

tensis mit ©. celeripes WINNERTZ (1852).

Kieffers Vorstellung von C. celeripes wurde von ihm (1899: 825—27) sehr gut begründet. Die illustrierte Be-

schreibung zeigt eindeutig eine Art, die in die C. scutellata-Gruppe gehört, wie sie in der vorliegenden Arbeit defi-

niert ist. Kieffer hatte jedoch eine falsche Zuordnung vorgenommen. Nach EDWARDSs (1929: 367—368) besitzt

C. celeripes eine 13gliedrige Antenne. Aus dem Text bei EDWARDS geht hervor, daß er Originalmaterial gesehen hat.

EpDwarDs (1919: 226) beschrieb aus einer parthenogenetischen Population das Weibchen von C. innupta und

später (1924: 188, 189 Fig. 11) auch das Männchen. C. innupta wurde von ihm (1929: 369) mit C. scutellata synony-

misiert, obgleich C. innupta keine Antennenrosette hat. Es hat sich gezeigt, daß C. innupta und C. lemnae artiden-

tisch sind (vgl. unten).

Das Fehlen der Antennenrosette bei gewissen Arten wurde von Edwards (1924: 188) diskutiert, als er den Bau

der rosettenlosen Antenne von C. celeripes Winn. erwähnt und bezweifelt, daß Kieffer diese Art richtig verstanden

hat. Als Edwards C. innupta beschrieb, erwähnt er, daß C. innupta C. scutellata ähnelt, aber daß die letzterwähnte

Art, ihm unbekannt ist. Edwards hatte offenbar noch vor 1929 das Originalmaterial aller von Winnertz beschriebe-

nen Corynoneura-Arten untersucht. Vermutlich ist dieses während des Entomologenkongresses in Bonn gesche-

hen (Dr. P. Cranston in litt.), wenngleich Edwards nichts davon in seinen Veröffentlichungen erwähnt.

Der Auffassung Edwards folgend wurde seine Gruppe B mit C. scutellata als Untergattung Corynoneura s. str.

von GOETGHEBUER (1939) verstanden. Die der Originalbeschreibung der Gattung zu Grunde liegende Art C. mi-

nuta müßte wegen der „Antennenrosette“, die beide Geschlechter besitzen (ein Merkmal, das auch bei den bekann-

ten Arten stets gegeben ist) in die Gruppe A von EDWARDS (1929: 368) oder zur Untergattung Eucorynoneura

GOETGHEBUER (1939: 4) gestellt werden.

Die Schwierigkeiten, die die oben erwähnten Autoren in der Klassifizierung gehabt haben, werden auch dadurch

deutlich, daß GOETGHEBUER (1939: 6) selbst C. brevipennis Goetgh. in Corynoneura s. str. gestellt hat. C. brevi-

pennis ıst eine Art mit Antennenrosette und weiteren Merkmalen der Eucorynoneura-Gruppe und sollte darum

nicht zu Corynoneura s. str. im Sinne Goetghebuers gestellt werden. EDWARDS (1929: 369) seinerseits ordnet C. fus-

cihalter Edw. zu den Arten seiner Gruppe A, obgleich diese Art zum Verwandtschaftskreis von Corynoneurella

Brund. gestellt werden muß, wie das erhaltene Originalexemplar erkennen läßt.

Unter dem Namen C. scutellata sind noch weitere Arten in der Literatur aufgefaßt worden. EDWARDS hat u. a.

das Männchen von C. gratias Schlee C. innupta zugeordnet. PıNDER (1978, 125 A) versteht unter C. scutellata die

von Schlee beschriebene Art C. gratias. Eine Varietät von C. scutellata (sensu EDWARDS 1929: 369) wurde von

BRUNDIN (1949: 833) als C. edwardsi beschrieben. SCHLEE (1968: 26) seinerseits grenzte C. gratias ab ohne zu wis-

sen, welche Art C. scutellata im Sinne von Winnertz ist. Als C. scutellata wurde von SCHLEE (1968: 107) eine Art

aufgefaßt, deren Hypopygium von LINDEBERG (1962: 8, sub C. scutellata) abgebildet worden ist. Zuvor hatten ne-

ben KıEFFER (1899, sub C. celeripes) und EDWARDS (1924, sub C. innupta) auch GOETGHEBUER (1932: 136, sub

C. scutellata) von diesem Hypopyg-Typ mehr oder weniger gute Abbildungen gebracht. Wegen der Art C. ed-

wardsi hatte LINDEBERG (1962) versucht, mit dem ihm vorliegenden finnischen Material die Art C. scutellata abzu-

grenzen. Später zeigte sich jedoch, daß die von ihm wegen ihres Hypopygiums als C. scutellata aufgefaßte Art

C. arctica Kieff. war.

In Europa kommen wenigstens 3 Arten mit einem ähnlichen Stylus vor. Der Typus von C. scutellata ist jedoch

ein Weibchen. SCHLEE (1968) konnte auf Grund genauerer Untersuchungen der Innenstrukturen des Hypopy-

giums 2 „C. scutellata-Arten“ unterscheiden. Seine Auffassung von C. scutellata gewann er nicht zuletzt auf Grund

von Exemplaren aus einem Torfstich, M,, Riihimäki, Südfinnland, und aus dem Fluß Luiro, Sodankylä, Finnisch-

Lappland, leg. HırvENOJA (vgl. SCHLEE 1968: 107). Vermutlich befanden sich in dem unpräparierten, ihm zur Ver-

fügung gestellten Material mehrere Arten. Dafür spricht, daß Schlee die abweichende Apikalborste von Ti/P; bei

C. scutellata erwähnt, jedoch ein Hypopygium ohne Coxitlobus abbildet, das zu C. arctica gehört.

Die Tatsache, daß Edwards nicht den Typus von C. scutellata bis zur Art bestimmen konnte, ist bedauerlich,

denn er war offensichtlich der letzte, dem Originalmaterial vorlag. Nach HORN & KAHLE (1935— 1937: 305) war

der größte Teil des Corynoneura-Materials 1881 an das Zoologische Museum in Bonn, weitere Exemplare auch an

das Naturhistorische Museum von Wien, und nach Senckenberg, Frankfurt/Main, gekommen. Jener Teil der Insek-

tensammlung mit den Chironomiden ist in Bonn während des 2. Weltkrieges verloren gegangen.

214

1 GRDRR 5

Fig. 1 Corynoneura scutellata. A Segmentgrenze der Puppenexuvie im Dunkelfeld mit intersegmentalen Dörn-

chen der Tergite und Sternite; B intersegmentale Dörnchen, REM-Aufnahme; C lange Spitzchen von Sternit 2;

D Chagrinierung von Tergit 4.

Bemerkungen zur Morphologie von Corynoneura

Imago

Bei der Beurteilung der Corynoneura-Imago wird in diesem Zusammenhang von der Diagnose aus-

gegangen, die Brunin (1956: 172) gegeben hat. Brundin grenzt die Gattung im Sinne von Winnertz

ab. ScHLEE (1968) faßt die Gattung Corynoneurella Brunpin (1956: 171) als ein Synonym von Coryno-

neura auf. Es wird hier verzichtet, Stellung zu dieser Entscheidung zu nehmen, weil die Metamorpho-

sestadien der in Frage kommenden Arten von Corynonerella nahezu unbekannt sind (Syn. ? Bauseia

Kieffer 1922). Es ist nur die volle Metamorphose von C. fittkani Schlee bekannt, die eventuell zur

Gattung Corynoneurella gehören könnte. Diese Art zeigt Unterschiede gegenüber den nachstehend

behandelten Arten, deren Wertung jedoch derzeit noch nicht möglich ist.

Zu den morphologischen Merkmalen, denen bisher noch nicht genügend Aufmerksamkeit ge-

schenkt worden ist, gehört die Ausbildung des Clavus beim Flügel. Brunpın (1956: 26) illustriert des-

sen Entwicklungstrend, wobei die Costa, Ri und R4+5 (vielleicht auch R2+3) zunehmend ver-

schmelzen. In den Diagnosen wird jedoch nicht erwähnt, daß R4+5 offenbar nur beim Weibchen von

Corynoneura vorhanden ist und beim Männchen mehr oder weniger reduziert zu sein scheint. Der

Grad der Verschmelzung der erwähnten Adern kann bei der Bestimmung hilfreich sein.

Puppe (vgl. Zavreı 1928, Lenz 1939).

Exuvie ohne Thorakalhorn, mit für die meisten Chironomiden charakteristischem Borstenbesatz.

An den Segmenten 3 bis 8, vier Lateralborsten. Auf Tergit 7 scheinen oft 5d-Borsten zu stehen; ihre

Zahl pflegt jedoch auf den vorderen Segmenten geringer zu sein. Zusätzlich zu den 3 Analborsten, wie

215

sie charakteristisch für Ortholadiinen sind, kommt noch ein viertes (? plesiomorphes) medianes Paar

vor. Die Beborstung zeigt eine starke Tendenz zur Entwicklung von Schlauchborsten, die jedoch

nicht immer erkennbar sein muß. Die Chagrinierung der Segmente ist ziemlich einheitlich und besteht

aus kleinen Spitzchen, die sich auch auf die Analsegmente ausdehnen; Reihen von stärkeren interseg-

mentalen Dörnchen reichen auf die Analränder der Tergite und Sternite (2) 3-7. Ihre Ausbildung

scheint zwischen den Populationen zu variieren. Sie sind im Text berücksichtigt worden, weil sie

leicht zu erkennen sind und ihre taxonomische Bedeutung überprüft werden sollte. Dieser sonst so

charakteristische Dörnchenbesatz scheint z. B. bei C. minuta zu fehlen. Diese Art hat eine abwei-

chende, wohl stärker apomorphe Chagrinierung in Form von langen Spitzchen an den Abdominalseg-

menten. Lange und schmale Spitzchen kommen bei den nachstehend behandelten Arten sonst nur auf

den Sterniten 1 und/oder 2 vor.

Larve (vgl. Zavreır 1928, Lenz 1939 und Cranston & al. 1983).

Kopf länglich, glatt oder mit verschiedenartigen Oberflächenstrukturen. Antennen nahezu so lang

oder länger als der Kopf, 4gliedrig, 3. und 4. Glied verwachsen. Nach Lenz (1939: 17) hat Goetghe-

buer bei den Larvulae ein zweiteiliges Antennenbasalglied gesehen. Erstes Glied etwa so lang wie die

Summe der Endglieder; Unterschiede zwischen den Arten scheint es u. a. in der absoluten Länge und

im Längenverhältnis der einzelnen Glieder zu geben; 2. Glied etwas geknickt. Ringorgan etwa in der

Mitte des ersten Gliedes, zwei winzige Borsten in der distalen Hälfte; apıkale Blattborste des ersten

Gliedes sehr kurz. S, einfach. Zahnleiste der Labialregion mit oder ohne kleinem, unpaaren Median-

zahn und 6 Lateralzähnen. Mandibeln neben dem Apikalzahn (bei den vorliegenden Larven) mit

4 Lateralzähnen, von denen der erste am größten ist; ssd klein, bei sı können nach Zavreı (1928: 655)

Unterschiede zwischen den Arten existieren. Prämandibeln mit mehreren kleinen Zähnen, von denen

die am weitesten ventralen am breitesten sind. Maxillen Fig. 9: 7. Thorakalsegmente 2 und 3 verwach-

sen. Ansonsten zeigen die Larven den normalen Habitus der Chironomiden. Zwei Ventralborsten von

Segment 10 sind auffallend stark und oft verzweigt. Vier Analpapillen.

Dem vorliegenden Manuskript liegen teilweise Untersuchungsergebnisse zu Grunde, die schon vor mehr als

20 Jahren zusammengestellt worden sind. Ein erheblicher Teil der Präparationsarbeiten und Messungen an finni-

schen Thienemanniella- und Corynoneura-Arten ist seiner Zeit von Mag. phil. Elina Hirvenoja vorgenommen

worden.

Für eine befriedigende taxonomische Beurteilung der Corynoneura-Arten sind die Weibchen ebenso wichtig,

wie die Metamorphosestadien. Auf Grund unserer damaligen Erfahrungen können einige Auffassungen von

SCHLEE (1968) nicht akzeptiert werden. Bedauerlicherweise waren wir seitdem nicht in der Lage, eine umfassende

Monographie der Gattung durchzuführen. Ein großes Hemnis war dabei die Schwierigkeit, die Taxonomie von

C. scutellata, der Typusart der Gattung Corynoneura zu klären. Dieses war im Herbst 1987 möglich.

SCHLEE (1968) beschreibt die Männchen, der ihm vorgelegenen Arten sehr gründlich. Nachstehend werden seine

Angaben noch ergänzt mit Meßwerten und Beschreibungen, die insbesondere an den selben Populationen von den

beiden Geschlechtern gemacht werden konnten. Die Zahlenangaben basieren zwar nur auf relativ wenigen Indivi-

duen, dürften aber einer allgemeinen Orientierung sehr dienlich sein, zumal z. B. der LR-Wert erfahrungsgemäß

höchstens um etwa 0,1 variiert. Die Beborstung (siehe BRUNDIN 1956: 172) der behandelten Arten ist von uns auch

untersucht worden, aber weil die Unterschiede zwischen den Arten nicht überzeugend sind, wird auf sie nicht wei-

ter eingegangen. Es wird ebenfalls auf die BR-Werte verzichtet, die etwa 3,5 erreichen und keine erkennbaren Ar-

tenunterschiede erkennen lassen.

Aufgrund unserer neueren Erfahrungen ist die Chagrinierung der Exuvie besser an getrockneten als an in Euparal

eingebetteten Häuten zu sehen. Es lassen sich auch Unterschiede in der Gestalt der Dörnchen, wenigstens zwischen

den Artengruppen, erkennen. Sie sind jedoch nachstehend nicht genügend berücksichtigt worden. Leider stand

nicht ausreichend genug unpräpariertes Material zur Verfügung.

Die Corynoneura scutellata-Gruppe

Zu den auffälligsten morphologischen Veränderungen innerhalb der Gattung Corynoneura gehören die Trends

bei der Ausbildung des Sternapodems X und der Phallusarmatur. Die Richtung der Anagenese ist wahrscheinlich

216

so zu verstehen, wie sie bereits von SCHLEE (1968: 145) aufgezeigt wurde. Am Ende dieser Entwicklungsrichtung

steht nach SCHLEE (1968) C. celtica Edw. in seinem Sinne (diese Art ist jedoch C. brevipennis Goetgh.). SCHLEE

(l. c.) verwendet für die Begründung der stammesgeschichtlichen Beziehungen in erster Linie die Strukturen des

Sternapodem X und der Phallusarmatur. Andere Merkmale wurden von ihm als weniger aussagekräftig interpre-

tiert. Zu den letzteren gehört u. a. der Bau der Antennen, der von den älteren Autoren als wichtig betrachtet wurde

und, wie gezeigt werden kann, nicht zu vernachlässigen ist. Ein besonders wichtiges Merkmal bei der Beurteilung

der Weibchen sind die Antennen. Sie zeigen ähnliche Tendenzen in der Entwicklung der Beborstung wie die Männ-

chen. Der Stammbaum von SCHLEE (1968: 145) weist hier schwer zu deutende Diskrepanzen auf. Immer wieder be-

finden sich getrennt von einander Arten, in denen sowohl bei den Männchen als auch bei den Weibchen Antennen

eine Apikalrosette besitzen und daneben die Hintertibien gleichzeitig auch eine stark s-förmige Apikalborste („A“

T. eAlwardsi CT. apnaca €. brundini

)

Fig. 2: Hypopygien von Corynoneura-Arten der scutellata-, edwardsi- und carriana-Gruppe.

277

bei Fig. 112 von SCHLEE 1968) tragen. Natürlich sind solche gemeinsam auftretenden Strukturen als Parallelismen

denkbar, aber ebenso wahrscheinlich ist auch eine parallele Entwicklung des Hypopygiums in verschiedenen Li-

nien.

Eine befriedigende Diskussion dieser Problematik ist derzeit mit dem vorliegenden Material noch nicht möglich.

Hennig und Brundin haben oft genug betont, daß die phylogenetische Systematik das Suchen der Schwesterarten

oder -gruppen ist. Es wird sich zeigen müssen, ob die hier zusammengestellte scztellata-Gruppe monophyletisch

ist. Sie besteht aus einander sehr ähnlichen Arten, die bisher nicht gut unterschieden werden konnten. Viele der ge-

meinsamen Merkmale scheinen Symplesiomorphien zu sein, z. B. der Bau der Antennen oder die Innenkontur des

Hypopygiums, vielleicht auch der „Doppelstylus“.

Diagnose

Imagines bräunlich, grünlich getönt. Antennen des Männchens 11gliedrig; Apikalteil des letzten

Gliedes keulig verdickt, gegen das Ende zugespitzt, distal proximalwärts bis über die Verdickung mit

Sensilla chaeticae besetzt; im Proximalteil des letzten Antennengliedes viele normale Antennenbor-

sten. Antennen des Weibchens 7gliedrig, entspricht im Bau dem der Weibchen der Familie, wobei

letztes Glied Sensillen nahezu auf der ganzen Länge bis zum Apex trägt. Auf dem Flügel erreichen die

Costa bzw. der Clavus beim ® etwa !/, beim O’ kaum die Hälfte der Flügellänge; beim @ R4+5 mit

den vorderen Teilen des Clavus wenigstens bis zur Hälfte des Clavus verschmolzen, beim J ist der

Clavus höchstens etwa zweimal länger als breit. Hypopygium des O’ mit Styli, die eine mehr oder we-

niger lobusähnliche Crista dorsalıs tragen; die proximalen Anhänge (?PV) reichen etwa bis zur Mitte

der Gonocoxite. Puppe mit langen Spitzchen auf 2. Sternit oder oft auch an den Lateralteilen vom

1. Sternit. Eindeutige Gruppenmerkmale fehlen jedoch im Puppenstadium. Larven mit einer Kopf-

kapsel, die mehr oder weniger deutliche Skulptur zeigt. Antenne länger als der Kopf. Zahnleiste der

Labialregion mit einem kleinen, unpaaren Medianzahn und 6 etwa gleichgroßen Lateralzähnen. Die

großen Ventralborsten von Segment 10 mit langen proximalen Nebenstacheln, die individuell zu vari-

ieren scheinen.

Corynoneura gratias Schlee (1968: 26-29, C')

Typus-Material ist nicht vorhanden. Die Artbeschreibung bei Schlee ist jedoch eindeutig.

Imago,

AR 0,77-1,00, LR 0,52—0,60. Als Ergänzung zur Originalbeschreibung einige Werte (n = 3; Ma-

terial aus dem Fluß Puujoki, Ryttylä, Hausjärvi, Südfinnland, Juni/Juli 1964 gezüchtet). Palpenglie-

der: 18-22, 22-27, 27—31 und 45—54 um. Flügellänge etwa 1 mm; der Clavus etwa zweimal so lang

wie breit.

Beine ın um:

Fe le Ta, Ta, Ta, Ta, =: LR

P\ 268-299 321-366 192-214 96-107 54-63 18-27 31-36 0.58-0.61

PJ 384-429 348-402 219-232 94-107 45-58 18-18 34-40 0.58-0.61

PJ 313-357 339-375 188-219 94-107 40-49 18-18 36-40 0.53-0.58

Charakteristisch für das Hypopygium ist das Gelenk des Phallapodems. Arttypisch sind auch die

im Vergleich zu anderen Arten der C. scutellata-Gruppe stärkeren Borstenbuckel am Hinterrand des

Analtergits wie auch die Kürze des Phallapodems. Gonocoxite ohne Innenlobus.

218

Imago, 9

Zusammen mit den obigen Männchen wurden viele Exemplare von C. edwardsi gezüchtet, aber ein

neugeschlüpftes Weibchen war verschieden und wegen des rundlichen dritten Palpangliedes, der

Beinmaßen und Merkmalen der Puppenexuvie konnte es als C. gratias bestimmt werden. Maße der

Palpen: 13, 17, 17 und 40 um. Flügel geschrumpft, Clavus ähnelt C. scutellata.

Beine in um:

Be Aral Ta, Ta, Ta, Ta, 2 LR

PJ 214 268 15 Ol 88 18 36 0.58

P, 304 Sg 18922168 40 18 36 0.58

P, 260 Zutat! IS a 18 36 0.57

Spermatheken oval, etwa 80 um.

Puppe

Material: Puujoki, vgl. oben.

Länge der Exuvie 2,2—2,3 mm. Vorderteil des Nahtrandes schmal, schwach gekörnelt oder ge-

dornt. An den Flügelscheiden 1-4, meist verhältnismäßig schwache Perlenreihen. Schwer sichtbare

(? kurze) Spitzchen auf Sternit 2. Dorsale Chagrinierung auf den Segmenten 2-9, fehlt lateral von den

d-Borsten. Intersegmentale Dörnchen an den Hinterrändern der Segmente 3-7, dorsal und ventral

meist weniger als 10 (4-12) in jeder Reihe. Die (?4) I-Borsten von Segment 2 einfache Spitzborsten.

Gonopodenscheiden des Männchens enden zwischen dem Hinterrand des Analsegments und den

Sockeln des medianen Analborstenpaares. Scheiden der Cerci des Weibchens enden weit vor dem me-

dianen Analborstenpaar. Schwimmhaarsaum des Analsegments mit 33—47 Borsten (n = 7).

Larve

Material: Puujoki, vgl. oben, Zuordnung nicht sicher.

C. gratias lebte in dem Fluß Puujoki u. a. zusammen mit C. edwardsi. Die Larvenhäute einer Ein-

zelzucht von C. edwardsi aus Tvärminne (wie einige aus Puujoki) hatten kürzere Antennen als die

Larvenhäute des letzten Larvenstadiums einiger Larven, die hier als zu C. gratias gehörig aufgefaßt

worden sind. Kopfkapsel schwach und unregelmäßig skulpturiert. Antenne 370-390 um lang, höch-

stens 1/4 länger als der 320-360 um lange Kopf; Längenverhältnisse der Antennenglieder:

100:39:47:3;, 100:38:44:3; 100:34:46:3.

Mundteile wie bei C. scutellata. Die großen Ventralborsten von Segment 10 nahezu einfach, nur

mit kleinen, proximalen Nebenstacheln besetzt.

Verbreitung

In Finnland ist C. gratias etwa bis zum 62°N bekannt.

Corynoneura scutellata Winnertz (1846: 13, ?)

Neotypus, 9, aus der Kollektion Winnertz, in Euparal eingebettet, Museum National d’Histoire Naturelle, Pa-

rıs. Dort auch noch ein genadeltes anderes, unpräpariertes Weibchen.

Synonymie

Corynoneura lemnae Schiner (in FRAUENFELD 1866: 974, Imago)

Syntypen, 4Q in einer Glastube im Naturhistorischen Museum Wien. Ein Exemplar davon in Euparal gebettet,

als Lectotypus hier festgelegt. Den vorliegenden Individuen fehlen die Antennen oder wenigstens das letzte An-

05 mm

EC. gpnocera €. carriana

UT. carriana

EC edWwardsi

220

€. gratias

CL. arctica

C. edWnardei

TC. edWwardsi

€. carriana

Fig. 4: Antennen der Corynoneura-Arten der scutellata-, edwardsi- und carriana-Gruppe.

tennenglied, aber die charakteristische Apikalborste (A) am Ende der Ti/P; belegt eindeutig die Zuordnung, da die-

ses Merkmal bisher von keiner Art sonst aus Europa bekannt ist.

Corynoneura innupta Edwards (1919: 226, ?)

Syntypen, 4Q aus Letchworth, Hertfordshire, England; VI. 1918, F. W. Edwards leg. Ein Exemplar davon in Eu-

paral gebettet, hier als Lectotypus festgelegt.

Corynoneura longipennis Tokunaga (1936: 50, O’ und 9)

Das Typusmaterial (siehe TOKUNAGA 1936: 51) wurde ı. J. 1964 untersucht und konnte nicht von den aus Finn-

land vorliegenden bisexuellen Exemplaren unterschieden werden. Das Material war damals nicht in Kyoto, wıe ın

der Originalbeschreibung erwähnt wird, sondern in dem entomologischen Laboratorium der Universität Kyushu,

Fukuoka, Japan. Es war jedoch nicht möglich, die Artidentität der parthenogenetischen und der bisexuellen Popu-

lationen zu beweisen. Geringe Unterschiede bei den Puppenexuvien und Imagines der vorliegenden Populationen

lassen sich nicht werten.

Allgemeine Bemerkungen

Die Maßangaben beziehen sich jeweils auf 5°’ und 5 9 von mehreren Lokalitäten in Riihimäki, Südfinnland, wo

diese Art auch, aus einem Tonstich stammend (T,), 1964 im Laboratorium gezüchtet werden konnte. Die Zuchten

ergaben sowohl Männchen als auch Weibchen. Es zeigte sich später, daß sich in den Aquarien viele Jahre hindurch

auch eine parthenogenetische Population hielt, die sich nicht von den bisexuellen Tieren unterscheiden ließ. Der ge-

naue Ursprung der parthenogenetischen Population konnte nicht geklärt werden. Das Schlüpfen der Imagines voll-

zog sich täglich.

Bzell

Nach FRAUENFELD (1866: 974) sollten die Puppen von C. lemnae zu beiden Seiten des Anallobus nur 8 Borsten

und die Larven nur 2 Analpapillen haben. Falls diese Beobachtungen der Wirklichkeit entsprächen, wäre die obige

Synonymisierung mit der bisexuellen Population fraglich. Weil das Typusmaterial aber sowohl von €. scutellata als

auch von C. lemnae aus Weibchen besteht, ist es jedoch wahrscheinlich, daß beide aus parthenogenetischen Popu-

lationen stammten. EDWARDS (1919: 226.227) schildert ein ähnliches Zuchtergebnis aus seinem Labor im Zusam-

menhang mit der Beschreibung von C. innupta (innupta = Jungfrau). Er bezweifelt allerdings, daß sich die Angabe

von Goetghebuer (siehe Lenz 1939: 17) über die Parthenogenese von C. celeripes, also auf C. scutellata, auf C. in-

nupta beziehen. Dies dürfte jedoch zutreffen, denn wie bereits erwähnt, hatte KIEFFER (1899) eindeutig eine Art der

C. scutellata-Gruppe als C. celeripes Winn. neubeschrieben und illustriert.

Imago, J

AR 0,77-1,00. Palpenglieder: 17—24, 24-29, 31—36 und 50-62 um. Flügellänge: 1,0—-1,2 mm;

der Clavus 1,5 oder höchstens zweimal so lang wie breit.

Beine ın um:

Fe rm Ta, Ta, Ta, Ta, Zr LR

P\ 238-305 333-371 195-219 105-114 57-67 29-33 42-43 0.55-0.60

P, 362-433 376-400 210-229 95-105 48-57 20-29 33-43 0.54-0.60

P, 305-362 333-429 195-248 100-114 48-48 29-29 38-43 0.57-0.59

Apikalborste (A) von Ti/P; senkrecht zur Längsachse von Ti, kurz S-förmig. Charakteristisch für

die Art ist auch ein sehr kleiner, unter dem Stylus leicht unsichtbar bleibender Innenlobus der Gono-

coxite. Gelenkzapfen des Penisapodems in dem Sternapodem X lateral gerichtet.

Imago, 9

Letztes Antennenglied etwa so lang wie 1,5—2 der vorhergehenden zusammen. Palpenglieder:

17-19, 22—26, 26-31 und 50-55 um. Flügellänge etwa 1,1 mm; die distale Hälfte des Clavus ver-

wachsen, aber an der proximalen Hälfte R4+5 weit von dem Vorderteil des Clavus getrennt.

Beine in um:

Fe Ta, Ta, Ta, Ta, Ta Ta, LR

PJ 238-286 276-357 152-190 76-95 38-57 29-29 38-43 0.51-0.58

P, 343-405 310-390 190-238 76-95 43-57 24-29 33-48 0.58-0.62

P, 295-352 290-384 171-219 86-100 38-48 24-29 38-135 20853-035577

Apikalborste wie beim Männchen, kurz S-förmig. Sie ist das wichtigste Merkmal bei der Bestim-

mung der Art innerhalb der Gruppe. Besonders bei parthenogenetischen Tieren die letzten Tarsen-

glieder und die Enden der anderen Glieder etwas verdunkelt. Spermatheken oval, etwa 80 um.

Puppe

Material: Riihimäki und Tvärminne, Südfinnland, M.H. leg., vgl. auch LANGTON 1984, sub innupta.

Exuvie 2.0—2.4 mm lang. Nahtrand schwach gekörnelt. An den Flügelscheiden 1-4 kurze Perlen-

reihen, oft sehr schwach ausgebildet. Chagrinierung auf allen Segmenten, das 1. ausgenommen. Lange

Spitzchen auf Sternit 2. Lateralteile von Sternit 1, besonders bei den parthenogenetischen Tieren, mit

Spitzchen wie bei C. arctica. Sterniten 3-5 (6) spärlich mit winzigen Dörnchen bis zum Lateralrand

der Segmente. Intersegmentale Dörnchen, etwa 10 (7-12), an den Hinterrändern der Tergite und

Sternite 3-7. Gonopodenscheiden des @ enden etwa in der Mitte zwischen den Sockeln der medianen

Analborsten und dem Hinterrand des Analsegments; ihre Gestalt ist auch unter dem Deckglas stärker

wurstförmig als bei anderen Arten. Scheiden der Cerci beim ® enden weit vor dem medianen Anal-

borstenpaar. Schwimmhaarsaum mit 31—42 Borsten.

222

Auf Segment 2 waren im obigen Material, die Borsten |; und ], oft eine Schlauchborste. Aus dem See Puruvesi lie-

gen Puppenexuvien, B. Lindeberg leg., vor, bei denen I, -1; bzw. I,-1, Schlauchborsten sind. Obgleich die Zugehö-

rigkeit der Exuvien nicht mit Zuchten belegt war, wird hier intraspezifische Variation vermutet. Die Puppe von

C. scutellata ähnelt sehr der von C. gratias. Wenn beide nebeneinander liegen, ist die Unterscheidung durch den

Habitus möglich, aber für eine Einzelbestimmung sind kaum andere Merkmale als die schwachen Unterschiede in

der lateralen Chagrinierung der mittleren Segmente und die Unterschiede am Analsegment zu verwenden. Die Ar-

ten sind jedoch bisher nie nebeneinander vorkommend angetroffen worden.

Larve

Material: Riihimäki.

Wenigstens auf dem Hinterteil der Kopfkapsel, allerdings nur mit starker Vergrößerung sichtbar,

mit netzartiger Skulptur; nur in einem Falle, bei dem größten Larvenkopf, war die Skulptur nahezu

ebenso deutlich wie auf dem Kopf von €. arctica. Antenne etwa 340-450 um lang, etwa !/4 länger als

der etwa 300-340 um lange Kopf; Längenverhältnisse der Antennenglieder: 100:42:43:3;

100:49:53:3; 100:46:51:3.

Weitere Strukturen der Larve auf Fig. 10.

Verbreitung

In Finnland ist C. scutellata aus Brackwasser, Flüssen, Seen und Teichen von der Südküste bis Lappland bekannt

(bei TUISKUNEN & LINDEBERG 1986 sub C. longipennis).

2 = €. gratias

€. scutellata

arctica

CT edlWnardgi

de : €. brundini

€. tarriana

50 um €. gpnocera

Fig. 5: Palpen der Corynoneura-Arten der scutellata-, edwardsi- und carriana-Gruppe.

225

Corynoneura arctica Kieffer (1923: 4, ©)

Lectotypus d, designiert von D. R. Oliver, aus Novaja Semlja, F. Okland leg. in den Sammlungen des Zoologi-

schen Museums der Universität Oslo.

Material: 3J und 2 9, aus dem Fluß Luiro, Sodankylä, Finnisch-Lappland, August 1960;2 0’ und 2Q aus einem

Torfstich in Riihimäki, Südfinnland, Mai— Juni 1962, M. H. leg. ; 10°, 1 P, Felsentümpel bei Tvärminne an der Süd-

küste Finnlands, B. Lindeberg leg.

Imago, ©

AR 1,02 (0,87—1,09). Palpenglieder: 22-26, 29-34, 36-46 und 67-82 um. Flügellänge

1,1- 1,4 mm; der Clavus etwa zweimal so lang wie breit. Beine in um:

i A Ta LR

Fe 7 Ta, Ta, Ta, a, 5

PJ 314-362 371-429 210-238 114-124 67-76 29-33 38-48 0,.53-0.57

P, 438-505 424-176 238-257 114-119 62-67 29-29 48-48 0,.56-0.59

PJ 376-419 429-464 250-268 123-138 57-67 29-33 43-48 0.54-0.60

Apikalborste (A) von Ti/P; etwas gekrümmt, steht in einem ein wenig größeren Winkel zur Tibial-

längsachse als bei C. gratias. Hypopygium mit Gelenkzapfen des Penisapodems, der lateral gerichtet

ist. Gonocoxite ohne Innenlobus.

Imago, 9

Letztes Antennenglied etwa so lang wie die zwei vorhergehenden zusammen. Palpenglieder:

17—22, 23—34, 30-34, 30—36 und 48-60 um. Flügellänge 1,2—1,3 mm; Clavus nahezu vollständig

(wenigstens bei typischen Exemplaren) verwachsen. Beine in um:

Fe Tr Ta, Ta, Ta, Ta, Tar LR

PJ 280-395 357-371 190-210 90-105 52-57 20-33 38-43 0.53-0.59

P, 419-429 390-414 229-252 105-105 52-57 26-33 38-48 0.57-0.64

PJ 348-376 393-429 225-241 110-119 43-48 24-33 38-48 0.56-0.60

Aus dem kleinen See Posolampi (ein Cladotanytarsus-See), nicht weit vom oligotrophen Fluß Luiro in Sodan-

kylä, schlüpften 1962 in den Fangtrichtern eine große Zahl Weibchen, die eine parthenogenetische Population oder

jedenfalls ein sehr abweichendes Q-Q-Verhältnis repräsentierten. Die Tiere sind kleiner als die ursprünglich be-

schriebenen, aber aufgrund der Puppenexuvien und des LR-Wertes konnten sie als C. arctica bestimmt werden. Es

ist nicht auszuschließen, daß die erwähnte Kleinheit und das abweichende O’-Q-Verhältnis von dem Lebensraum

abhängt. Die Arten auch anderer Chironomiden-Gattungen waren dort kleiner als in anderen Gewässern. Die ge-

nauen Werte werden hier gegeben, weil die Exemplare wegen der geringen Größe sehr C. gratias und auch C. ed-

wardsi ähnelten, von denen in demselben Biotop eine deutlich bisexuelle Population vorkam. Die Beinlängen in

um:

Fe bt Ta, Ta, Ta, Ta, ur LR

PJ 210-250 254-304 143-170 66- 85 38-39 18-18 34-36 0.56-0.57

P, 311-366 281-339 165-213 76- 98 40-46 18-19 31-40 0,.59-0.63

P, 268-339 277-348 154-192 84-105 36-45 21-19 31-40 0.55-0.55

Apikalborste (A) von Tı/P; wie beim Q. Spermatheken etwa 75-80 um. Sowohl beim J als auch beim ® sind

die drei letzten Tarsusglieder und die Apikalenden der anderen Beinglieder bisweilen etwas verdunkelt. Vielleicht

224

ist dies eine Eigenschaft, die auch bei anderen Arten mehr oder weniger deutlich ausgeprägt ist, aber vor allem bei

mit Fangtrichtern gesammelten und in Alkohol aufbewahrten Tieren nicht zu sehen ist.

Puppe

Material: Rıihimäki, Südfinnland und Sompio-Gebiet in Sodankylä, Finnisch-Lappland, M. H. leg.; Tvärminne,

B. Lindeberg leg.

Exuvie 2,1—2,8 mm lang. Nahtrand schmal, schwach gerunzelt oder gekörnelt. Die erste deutliche

Perlenreihe an den Flügelscheiden kurz, dazu kommen weitere 2—4 mehr oder weniger deutliche Rei-

hen. Chagrinierung auf allen Segmenten; auf den Sterniten 3—5 kleine Spitzchen, spärlich, bis zum La-

teralrand des Segments. Intersegmentale Dörnchen an den Analrändern der Tergite und Sternite 3—7

je mit mehr als 10 (10-18), bei britischen Exemplaren weniger; auf Tergit 2 bei Puppenhäuten aus

Finnland eine unregelmäßige Reihe mit wenigen Dornen. Lange Spitzchen auf Sternit 2. An den oral-

lateralen Ecken von Sternit 1 sind diese Spitzchen verhältnismäßig lang, in der Mitte des Sternits sehr

klein. Gonopodenscheiden des O reichen bis zum Hinterrand des Analsegments oder sind etwas län-

ger. Scheiden der Cerci beim Weibchen reichen nicht zu den Sockeln des medianen Analborstenpaa-

res. Schwimmhaarsaum des Analtergits mit 40-50 Borsten.

Larve

Material: 1 aus Tvärminne, Finnland, B. Lindeberg leg; 2 Larven mit Puppenexuvien ohne Imagines, sub innupta

Edw. aus der Sammlung des British Museum, London, $. ©. Howard leg., konnten nicht von dieser unterschieden

werden.

Kopfkapsel mit ziemlich starker netz- oder schuppenförmig angeordneter Skulptur. Antenne

495—560 um lang, etwa 1,5mal länger als der etwa 370 um lange Kopf; Längenverhältnisse der An-

tennenglieder: 100:40:36?(fehlt) (Tvärminne); 100:41:41:2; 100:42:38:2 (England).

Die großen Ventralborsten von Segment 10 entweder mit wenigen oder vielen proximalen Neben-

stacheln (die britischen Tiere mit wenigen zarten proximalen Ästen; bei der Larve aus Finnland die an-

dere Borste mit vielen Ästen besetzt).

Verbreitung

In Finnland in Schmelzwasserlachen, Tümpeln, Flüssen und Seen von den Felsentümpeln an der Südküste bis

Lappland (bei TUISKUNEN & LINDEBERG 1986 sub C. scutellata).

Die Corynoneura edwardsi-Gruppe

Die Begründung von Schwesternarten sollte nach Möglichkeit von apotypischen Arten ausgehen. Auf den ersten

Blick sehen die Arten der C. edwardsi- oder C. carriana-Gruppe ziemlich abgeleitet aus, aber bei näherer Betrach-

tung zeigen sich viele plesiomorphe Züge. Die Abgrenzung einer C. edwardsi-Gruppe mag unnötig erscheinen,

aber sie erleichtert in jedem Fall die artliche Zuordnung bei der Bestimmung.

Betrachtet man die Antennen oder die Strukturen des Hypopygiums beim Männchen, scheinen die Arten der

C. edwardsi-Gruppe das Ergebnis gewisser Reduktionstendenzen von der anagenetischen Stufe der C. scutellata-

Gruppe zu sein. Auch die Unterscheidung der Weibchen oder der Metamorphosestadien beider Gruppen vonein-

ander ist bisweilen sehr schwierig. Der Bau des Flügels, der der Puppe und der Larve in der C. edwardsi-Gruppe

zeigen ım Vergleich zu allen Arten der C. scutellata-Gruppe jedoch relativ ursprünglichere Züge.

Die C. edwardsi-Gruppe wird hier als die unmittelbare Schwestergruppe zur C. scutellata-Gruppe verstanden.

Innerhalb der C. edwardsi-Gruppe, ist C. brundini als eine apomorphe Schwesternart von C. edwardsi aufzufas-

sen.

Diagnose

Die Imagines ähneln in vielen Merkmalen denen der C. scutellata-Gruppe. Antenne 11gliedrig,

letztes Antennenglied kurz. Palpen können länger sein. Beim Flügel des & nehmen die starken Vor-

derrandadern wenigstens 25, beim ® wenigstens die Hälfte der Flügellänge ein. R4+5 deutlich er-

kennbar bis zur Hälfte, beim 9 bis 2/3 von der Länge des Clavus. Beim ©’ Clavus wenigstens zweimal

225

scutellata

=2— ——

20 um

CL. brundin

Fig. 6: Apikalenden der Hintertibien der Corynoneura-Arten der scutellata-, edwardsi- und carriana-Gruppe.

so lang wie breit. Apikalborste (die Borste „A“ bei ScHLeE 1968, Fig. 111-112) der Hintertibien nur

schwach gekrümmt. Medianteil des Sternapodems X beim Hypopygium sehr reduziert; Gelenkzap-

fen des Penisapodem lateralwärts gerichtet. Penisapodem lang und stark bogenförmig. Proximaler

Anhang der Gonocoxite kurz und distal spitz; im mediodistalen Teil der Gonocoxite sehr schwach an-

gedeutet, ein breiter, flacher Lobus. Stylus einfach, aber mit mehr oder weniger leicht sichtbarer,

schmaler, dorsaler Crista besetzt. Puppe mit langen Spitzchen auf den beiden ersten Abdominalsterni-

ten. Von den Puppen und von den Larven liegen nicht eindeutige Gruppenmerkmale vor; Antenne der

Larve so lang oder wenig kürzer als die Kopfkapsel.

Corynoneura edwardsi Brundin (1949: 698, 831-833, (')

Typusmaterial (Riksmuseum, Stockholm, Schweden).

Material: 5C und 5Q aus den Seen Sompiojärvi und Seitajärvi, nebst aus dem Fluß Luiro, Sodankylä, Finnisch-

Lappland, 1959-1960, M. H. leg.

Imago, ©

AR 0,68-0,87. Letztes Flagellomer so lang wie die 6-8 vorhergehenden zusammen. Palpenglieder:

21-23, 28-32, 38-50 und 55—71 um. Kopf und Thorax etwas, auch der Raum zwischen den Meso-

notalbinden verdunkelt. Flügellänge 1,1—1,3 mm; Clavus etwa 2mal länger als breit. Beine ın um:

Fe hal Ta, Ta, Ta, Ta, Ir LR

P\ 298-333 377-412 197-239 101-118 57-83 26-31 35-39 0.54-0.57

PJ 417-500 377-465 228-280 101-127 57-70 26-31 35-41 0.56-0.60

PJ 346-412 355-421 206-237 96-118 39-53 22-31 35-39 0.54-0.58

Abdomen etwas verdunkelt, anale Hälfte von Tergit 7 mit hellem Fleck. Hypopygium wie in Fig. 2.

Imago, 9

Farbe wie beim Männchen, aber der Raum zwischen den Mesonotalbinden heller. Das 6. Anten-

nenglied so lang wie die 2—2,5 vorhergehende zusammen. Palpenglieder: 17—21, 23—29, 38—44 und

57—67 um. Flügellänge 1,2—-1,5 mm; Costa bzw. der Clavus endet etwa in der Mitte des Flügels;

R4+5 nur im apikalen Drittel mit dem Clavus verschmolzen; wenn mehr, ist die Trennung der Adern

jedenfalls deutlich. Beine in um:

Fe Ti Ta, Ta, Ta, Ta, Ta, LR

PJ 263-333 325-412 162-211 70- 92 44-53 22-35 31-39 0.47-0.51

PJ 377-482 360-465 162-281 88-114 48-61 26-35 35-39 0.57-0.60

pP, 324-430 342-390 171-219 88-114 35-48 22-31 31-44 0,.52-0.56

Spermatheken 80-100 um lang, oval, vor der Mündung zu einem Hals verschmälert und ge-

krümmt; dieser scheint bisweilen zu fehlen oder ist wegen der Lage in Präparaten nicht immer zu

sehen.

Puppe

Material: Züchtungen aus dem Fluß Puujoki, Ryttylä, Hausjärvi, Juni/Juli 1964 und aus dem Brackwasser bei

Tvärminne, 23.— 27.7.1970, Südfinnland; Fangtrichtermaterialien aus dem See Sompiojärvi, Finnisch-Lappland,

M.nH. leg.

Exuvie 2,2—2,5 mm lang. Nahtrand schmal, schwach gerunzelt und gekörnelt. Auf den Flügel-

scheiden eine lange und etwa 3 schwächere Perlenreihen vorhanden. Alle Puppenexuvien, die aus

Finnland zusammen mit den Imagines von C. edwardsi angetroffen wurden, haben lang Spitzchen auf

den Sterniten 1 und 2, etwa wie „Corynoneura Pe 1a“ in der Arbeit von LanGron (1984: 82—83,

Fig. 25c). Nach Langton sollte C. edwardsi dagegen nu: aut Sternit 2 solche Chagrinierung haben.

Anzahl der intersegmentalen Dörnchen am Analrand der Tergite 2-7 und Sternite 3-7 etwa 10

(6-15). Beim CO enden die Gonopodenscheiden im Bereich des Hinterrandes des Analsegments. Beim

haarsaum mit 38—48 (n = 8) Borsten.

3 09° or

€. gratiag €. arctica N €. carriana

C. scutellata C. edWwardgi €. brundini

Fig. 7: Spermatheken der Corynoneura-Arten der scutellata-, edwardsi- und carriana-Gruppe.

227

Larve

Material: Einzelzuchten aus Tvärminne, vgl. oben.

Kopfkapsel teilweise schwach, unregelmäßig gerunzelt, im Hinterteil ist eine schwache netzartige

Skulptur zu erkennen. Antenne 330-350 um, ebenso lang oder etwas kürzer als der 330-340 um

lange Kopf; Längenverhältnisse der Antennenglieder (n = 2): 100:43:50:3; 100:42:42:2.

Labialregion mit einer Zahnleiste, bei der sich ein kleiner, unpaarer Medianzahn zwischen den ver-

wachsenen größeren Zähnen befindet und 5 freien, etwa gleich großen Lateralzähnen. Mandibel ne-

ben dem Apikalzahn mit 4 Lateralzähnen, von denen der apikale am größten ist. Prämandibeln mit

7-8 Apikalzähnen, die breitesten und am meisten ventral liegenden sind apikal gerundet. Die großen

Ventralborsten am 10. Abdominalsegment nahezu einfach, proximal nur mit undeutlichen, dünnen

Nebenstacheln besetzt.

Verbreitung

C. edwardsi bevorzugt offensichtlich Flüsse und größere stehende Gewässer und lebt auch im Brackwasser. In

Finnland bis Lappland bekannt.

Corynoneura brundini spec. nov.

Holotypus, ©’, und Paratypen aus dem Kleinsee Posolampi (zur Zeit ein Teil des Stausees Lokka), Sodankylä,

Finnisch-Lappland, M.H. leg., in Coll. Zoologisches Museum der Universität Helsinki, Finnland. Ein Teil der 9 ©

und 4 Q wurde vermessen.

Die neue Art ähnelt C. edwardsı, aber läßt sich von dieser u. a. durch einen kleineren AR-Wert, kürzere Palpen

und die Beinmaße gut unterscheiden. Von den zahlreichen von Kieffer beschriebenen Arten mit 11gliedrigen An-

tennen, besitzt C. tyrolensis (Kieffer 1925: 565) ein etwa ebenso niedriges AR. Darüber hinaus gibt es jedoch Ab-

weichungen im Bau der Antenne. Aus der Beschreibung von C. tyrolensis bei ALBRECHT (1924: 196) sind die abwei-

chenden Merkmale (von Eucorynoneura) deutlich erkennbar.

Imago, CO

Das ganze Tier etwas, auch der Raum zwischen den Mesonotalbinden dunkel gefärbt. Letztes Fla-

gellomer mit vielen langen, normalen Antennenborsten, so lang wie die 3—3,5 vorhergehenden Glie-

der zusammen; AR 0,39—0,47. Palpenglieder: 24-26, 29—31, 29—34 und 34—43 um. Flügellänge

1,0-1,1 mm; Costa bzw. Clavusreichen ın die Nähe der Mitte des Flügels, Clavus 3—4mal so lang wie

breit. Beine ın um:

Fe Aral Ta, Ta, Ta, Ta, Ta, LR

P\ 276-333 305-371 124-143 52-57 33-38 24-29 38-43 0.38-0.41

P, 381-429 352-381 190-210 86-95 48-52 29-29 38-48 0.53-0.57

P_ 295-343 386-436 190-218 67-76 29-38 24-29 38-38 0.49-0,54

Hypopygium etwa wie bei C. edwardsi, aber die Styli sind stärker gekrümmt.

Imago, 9

Die Pigmentierung gleicht dem des Männchens, aber der Raum zwischen den Mesonotalbinden

bleibt hell. 6. Antennenglied etwa so lang wie die 2 vorhergehenden zusammen. Palpen: 17—24,

22—24, 24—32 und 34—38 um. Flügellänge 0,9—-1,2 mm; Costa bzw. Clavus reichen etwa bis in die

Mitte des Flügels; Beim Clavus R4+5 proximal deutlich von den vorderen Adern getrennt, die distale

Hälfte vom Clavus ist geschlossen. Beine in um:

228

(E. arctica CT. scutellata FT. cacriana

Fig. 8: - Analenden von Puppenexuvien der Corynoneura-Arten der edwardsi-, scutellata- und carriana-Gruppe.

1 Anb = laterale Analborsten undm Anb = mediane Analborsten des Analsegments.

229

Fe en Ta Ta Ta Ta Ta LR

1l 2 3 4 5

PJ 229-276 276-324 114-133 48-52 29-33 24-29 29-38 0.40-0.45

PJ 333-371 | 305357 % 171-1907 57-76 38-38 24-29 29-3 0,.51-0.59

PJ 276-333 370-420 200-230 62-71 29-34 24-29 38-38 0,51-0.56

Spermatheken oval, etwa 70-100 um lang.

Puppe

Material: !!ine Puppenexuvie des Männchens liegt aus den Tümpeln, den sogenannten „Lompolos“ in der Nähe

des Sees Kilpisjärvi, Enontekiö, Finnisch-Lappland, 15.7.1969, B. Lindeberg leg., vor. Die Exuvie zeigt eine ge-

wisse Ähnlichkeit mit der von C. edwarasi, läßt sich aber deutlich von dieser durch die Länge der Gonopodenschei-

den unterscheiden. Weil in derselben Probe neben den Imagines von C. brundini auch solche von anderen Coryno-

neura-Arten vorhanden waren, ist die Zugehörigkeit nicht ganz gesichert. Aus der Typuslokalität liegt eine ähnli-

che Exuvie eines Weibchens vor, ihr fehlt das 9. Segment.

Exuvie etwa 1,3 mm lang. Nahtrand schmal, schwach spitzenartig gekörnelt und gerunzelt. Fine

lange und deutliche und 4 kürzere und schwächer werdende Perlenreihen an den Flügelscheiden. Ster-

nite 1 und 2 mit langen Spitzchen; am Medianteil von Sternit 1 scheinen die Anzahl und Länge der

Spitzchen jedoch deutlich geringer als bei C. edwardsi zu sein. Chagrinierung der übrigen Abdomi-

nalsegmente etwa wie bei C. edwardsi; sehr kleine Chagrinierung wenigstens auf 3. Sternit, auch au-

ßerhalb der d-Borsten bis zum Lateralrand des Segments. Anzahl der intersegmentalen Dörnchen am

Hinterrand der Sternite und Tergite 3-7, etwa 10 (8-14), am 2. Tergit nur 2-3. Gonopodenscheiden

überragen nicht die Sockeln des medianen Analborstenpaares. Schwimmhaarsaum vom 9. Segment

mit nahezu 40 Borsten.

Larve unbekannt.

Verbreitung

C. brundini lebt in Seen und Teichen in ganz Finnland.

Die Corynoneura carriana-Gruppe

SCHLEE (1968: Fig. 204) sieht in C. carrıana besonders aufgrund der Strukturen des Hypopygiums eine Schwe-

sterart von C. edwardsi. Ihre proximalen (?PV) Anhänge des Hypopygiums stellen jedoch nicht zwingend das Er-

gebnis derselben Entwicklungslinie dar. Vergleicht man die Flügel, so reicht bei C. carrıana die Costa bzw. der Cla-

vus (plesiomorph) weiter distal als bei allen hier behandelten Arten. Ferner hat C. carriana nach SCHLEE (1968: 30)

die Borsten auf den Metatarsen gegenüber den anderen Corynoneura-Arten in 6 (nicht in 5) Längsreihen wie bei

Thienemaniella angeordnet. Die (apomorphen) Antennen lassen sich von der Form ableiten, wie sie u. a. bei der

C. scutellata- oder C. edwardsi-Gruppe oder bei „Corynoneurella“ ausgebildet ist. Wegen der plesiomorphen

Züge ist die C. carriana-Gruppe jedoch nicht von der C. edwardsi-Gruppe abzuleiten, noch weniger von der

C. scutellata-Gruppe. Diese beiden können als die Schwestergruppen der C. carriana-Gruppe verstanden werden.

C. gynocera ist leicht als eine apomorphe Schwesterart von C. carrıana zu erkennen. Die carriana-Gruppe ent-

spricht der Gruppe C bei EDWARDS (1928: 369). Ob diese Gruppe als eigene Untergattung Paracorynoneura Goet-

ghebuer (1939: 7) anerkannt werden kann, wird sich erst klären lassen, wenn die zugehörigen Metamorphose-

stadien besser bekannt sind.

Diagnose

Imago:

Färbung ähnlich wie bei den Arten der vorhergehenden Artengruppen. Antenne beim JO’ höchstens

l1gliedrig, 6gliedrig beim P. Länge der einzelnen Haare des Flagellums höchstens 100 um (bei den

anderen Artengruppen sind sie länger, bis etwa 200 um). Das letzte Glied bei beiden Geschlechtern

230

apikal ohne Sensilla chaeticae, besonders beim ©’ spielkegelförmig. Im Proximalteil des letzten Glie-

des keine oder sehr wenige normale Antennenborsten. Costa bzw. Clavus reichen bis zur Flügelmitte

oder darüber hinaus; beim ©’ ist der Clavus wenigstens 4mal so lang wie breit. Beim Hypopygium

sind die proximalen Anhänge kurz, apikal nicht spitz. Stylus einfach. Medianteil des Sternapodems X

stark reduziert mit lateral gerichtetem Gelenkzapfen für das Penisapodem; letzteres lang und stark ge-

bogen.

Puppe nur schwer von der der C. scutellata-Gruppe zu unterscheiden. Lange Spitzchen kommen

auf 2. Sternit und sehr winzige auf 1. Sternit vor.

Larve unbekannt.

Corynoneura carriana Edwards (1924: 188-189, O', P)

Als eventuelle Synonyme von C. carrıana geben EDWARDS (1929: 369) und GOETGHEBUER (1939: 7) folgende Ar-

ten an:

C. acuticornis Kieffer (1912: 101-102, 9)

C. heterocera Kieffer (1915: 87, ©’)

C. crassipes Kieffer (1925: 564, ©’)

C. acuticornis könnte wegen der FJügeladerung mit C. carrıana artidentisch sein.

C. heterocera hat nach EDWARDS (1924: 189) ähnliche Antennen aber die Flügel sind verkürzt. Weil die Antennen

nach der Originalbeschreibung ohne Federbusch sind, hat GOETGHEBUER (1939: 7. 11) C. hbeterocera vermutlich

nicht synonymisiert. Goetghebuer faßt jedoch C. crassipes als ein Synonym auf. Die beiden letztgenannten Arten

sind wegen des Baues der Antennen und der Palpen sehr wahrscheinlich Synonyme von C. carrıana. Eine weitere

Art, die nach der Originalbeschreibung eine 11gliedrige Antenne mit kurzem Haarbusch besitzt, ist C. marına

Kieffer (1923: 43, O', 9).

Alle vier hier aufgeführten Arten werden vorerst als naomina dubia betrachtet und nicht als Synonyme zu €. car-

rıana gestellt, bis ihre genaue Zuordnung an Originalmaterial geprüft worden ist.

Typusmaterial von C. carrıana befindet sich im British Museum, London. Es wurde verzichtet Originalmaterıal

zu überprüfen, da die Zuordnung des finnischen Materials sicher erscheint.

Material: 3’ und 29 aus dem Brack wasser bei Tvärminne, Brännskär, Juni/Juli 1963, Elina Hirvenoja leg., 10°

aus Tvärminne, Jovskär, 27.7.1971, B. Lindeberg leg. und 1 aus einem Torfstich, Riihimäki, 1.7.1956, M. Hirve-

noja leg. Die Exemplare aus Rıihimäki sind deutlich kleiner als die aus dem Brack wasser.

Imago, ©

Antenne 11gliedrig; Apikalglied etwa so lang wie die 3-4 vorhergehenden zusammen, bisweilen

mit einigen normalen Antennenborsten im Proximalteil; die längsten von den locker sitzenden Feder-

buschborsten etwa 100 um; AR 0,37—0,45. Palpenglieder: 18-22, 22—24, 24—33 und 34—46 um.

Flügellänge 0,7—1,1 mm; Clavus etwa 4mal so lang wie breit. Beine ın um:

Fe eis Ta, Ta, Ta, Ta, Ta, LR

PJ 258-325 289-356 144-182 71-89 40-52 18-22 31-40 0.47-0.52

P, 338-409 307-383 169-182 71-89 37-45 18-22 35-40 0.48-0.55

P, 289-365 280-352 151-182 89-107 36-37 18-22 35-35 0.50-0.57

Hypopygium wiıe in Fig. 2.

Imago,

Bei typischen Exemplaren etwa das apikale Viertel des letzten Antennengliedes ohne Sensilla chaeti-

cae; Palpenglieder: 15-18, 18-22, 20-23 und 39—42 um. Flügellänge 1,1-1,3 mm; Clavus reicht

über die Mitte des Flügels, die einzelnen Adern stark verschmolzen. Beine in um:

23]

Fe en Ta Ta Ta Ta Ta LR

J: 2 3 4 5

PJ 285-303 338-343 160-173 71-84 40-49 20-22 40-40 0.47-0,51

P, 365-405 351-378 191-196 80-80 36-40 22-27 36-40 0.52-0.54

P, 347-365 329-356 178-187 89-90 36-36 20-20 36-45 0,53-0.54

Spermatheken 80-90 um lang; vor dem Ductus charakteristisch gekrümmt und etwas verjüngt.

Puppe

Material: Tvärmınne

Exuvie 2,0—2,4 mm (n = 3) lang. Nahtrand oral schmal, schwach gekörnelt oder gerunzelt. Apex

der Flügelscheiden mit 2-3 kurzen Perlenreihen. Auf 1. Tergit bisweilen einige Dörnchen und auf

50um

(12)

50um

(2-11)

Fig. 9: Corynoneura scutellata Winn., Larve: 1 Kopf; 2 Skulptur des Kopfes; 3 Sinnesfeld des Labrums, Epipha-

rynx und Praemandibeln. 4 Augen; 5 Maxille; 6 Praementum; 7 Mandibel; 8 Klauen der Vorderfüße; 9 Zahnleiste

der Labialregion nebst ventralen Borsten von Postgena; 10 Praeanale Borstenträger (Procerci); 11 Ventralborsten

des 10. Abdominalsegments; 12 Antenne.

232

1. Sternit an den oral-lateralen Ecken sehr kleine Spitzchen. Chagrinierung normal, nicht sehr breit,

beginnt auf 2. Segment. 2. Sternit mit langen Spitzchen. Intersegmentale Dörnchen dorsal und ventral

auf den Hinterrändern des 3.—-7. Segments, einige bisweilen auf 2. Tergit, meist etwa 10 (6-16). Go-

nopodenscheiden reichen etwa bis zum Hinterrand des Analsegments; beim O’ enden die Scheiden der

Cerci deutlich vor den Sockeln der medianen Analborsten. Analer Schwimmhaarsaum mit etwa

32-55 Borsten.

Larve unbekannt.

Verbreitung

C. carriana ist in Finnland bisher etwa bis zum 62°N Breitengrad angetroffen worden. Die Angaben von TUISKU-

NEN & LINDEBERG 1986: 366, über C. carrıana in Lappland, beziehen sich auf C. brundini spec. nov.

Corynoneura gynocera Tuiskunen (1983: 0’, Puppe).

Typusmaterial Coll. Zoologisches Museum der Universität Helsinki, Finnland.

CT erWwardgi %

un 08

wf 0S

CF. arctica en BE

(? carriana)

Fig. 10: Corynoneura spec., Larve: Antennen, Ventralborsten von Abdominalsegment 10, Apikalteile der

Praemandibeln nebst Skulpturen der Larvenköpfe von C. edwardsi Brundin, der vermuteten Larve von C. gratias

Schlee (mag die individuelle Variation von C. edwardsi repräsentieren!) und C. arctıca Kieff. Ferner Teile (die An-

tenne abgebrochen) eines Larvenkopfes aus einem Brackwasserbiotop bei Tvärminne, wo u. a. C. carrıana Edw.

angetroffen wurde.

233

Imago, d

Es ist nur das © mit der eigentümlichen 7gliedrigen Antenne (Pedicellus mitgezählt) bekannt; AR

(nach Tuiskunen) 0,58—0,72. Ergänzungen zur Originalbeschreibung: Palpen (Holotypus): 20, 28, 36

und 45 um. Die Flügel zeigen die ursprünglichste Gestalt der hier behandelten Arten; der Clavus

reicht deutlich über die Flügelmitte und ist etwa 6mal so lang wie die größte Breite.

Beine (Holotypus) in um:

Fe Abt Ta, Ta, Ta Ta, Ta, LR (nach Tuiskunen)

PJ 312120347 169 88 49 22 40 0.47-0.53

P, BSsregiT 178 80 45 27 45 0.47-0.52

P_ 338 342 178 98 45 27 45 0.47-0.53

Das Hypopygium weist eine bemerkenswerte Struktur auf. Ein geschlossenes Sklerit scheint einen

Annulus um das Analsegment zu bilden. Penisapodem verhältnismäßig kurz, etwa wie bei C. gratias,

aber sein Gelenkzapfen im X. Sternapodem ist stärker lateral gerichtet.

Puppe

Ergänzungen und Korrekturen zur Originalbeschreibung: Besonders an den oral-lateralen Ecken

vom 1. Sternit winzige Spitzchen, auf Sternit 2 sind diese lang und deutlich. Die dorsale Chagrinie-

rung fängt auf dem 3. Tergit an; an den Hinterrändern der Segmente dorsal und ventral mit etwa 10

(8-12) größere Dörnchen in jeder Reihe (n = 10).

Larve unbekannt.

Verbreitung

Die Art ist nur von der Fundlokalität in Finnisch-Lappland bekannt.

Bestimmungstabellen

Die hier behandelten Arten bzw. Artengruppen werden als eine monophyletische Einheit aufgefaßt. Dieser Ar-

tenkomplex repräsentiert zum Teil Corynoneura s. str. und Paracorynoneura im Sinne von GOETGHEBUER (1939).

Bei ihnen ist das Apikalende der Hintertibien stark angeschwollen. Das Apikalglied der Antennen hat beim O ent-

weder eine spielkegelförmige Gestalt und ist apikal ohne Sensilla chaeticae, oder es bleibt terminal verschmälert und

ist distal der subapikalen Verdickung mit Sensillae besetzt.

EIS.

1 (4) Der apikale Teil des spielkegelförmigen (Fig. 4) letzten Antennengliedes wenigstens auf

!/ı der Länge ohne Sensilla chaeticae: Apikalglied etwa so lang wie die 3-4 vorhergehen-

den Glieder zusammen; Behaarung der Flagellomeren kurz... .............. 2...

ae Me nee ke EEE. : 2: 00 aanae: Corynoneura carriana-Gruppe 2

2 96) Antenne Ileliedmer 2.22... 20 a N C. carrıana Edw.

312) Antenne zeliedusep Me... 0 2 a C. gynocera Tuisk.

4 (1) Apikalglied der Antenne anders (Fig. 4) geformt, distal von der Verdickung breit mit

Sensilla chaeticae und proximal mit vielen langen Antennenborsten besetzt (= normaler

Flaarbusch)s ge ee 0 Sr 5

5 (10) Stylus mit einer hyalinen, lobusähnlichen Crista dorsalis. Proximale (?PV) Anhänge der

Gonocoxite reichen in die Mitte der Gonocoxite. Apikalglied der Antennen nahezu so

lang wie die übrigen Flagellomeren zusammen. ............ Corynoneura-Gruppe 6

234

6 (m

7 (6)

Se)

9 (8)

10 6)

11 (12)

12 (11)

1.)

2 (0)

3.6)

” ©)

5 (6)

6 6)

7. (8)

9 (10)

10 (9)

Gelenkzapfen des Penisapodems im Sternapodem X ventral gerichtet. Vorletztes Palpen-

glied kaum länger als die ersten freien, kugeligen Glieder ........ C. gratias Schlee

Gelenkzapfen des Penisapodems im Sternapodem X lateral gerichtet. Vorletztes Palpen-

eliedmahezurzyyeimallläangeralsibreitage en.

An der distalen Hälfte der Gonocoxite ein sehr kleiner Lobus. Apikalborste (A) der

Hintertibien steht senkrecht zur Längsachse von Ti, kurz S-förmig (Fig.6)...........

rer tenls bon wassge C. scutellata Winn.

Gonocoxite ohne Lobus. Apikalborste (A) nicht S-förmig, zwar etwas gekrümmt, aber

deutlich ım spitzen Winkel zur Tibiallängsachse stehend .......... C. arctica Kieff.

Crista dorsalis nicht wie ein Lobus, sondern eine mehr oder weniger deutlich längere

Leiste. Die proximalen Anhänge (?PV) der Gonocoxite kurz, distal ziemlich spitz.

Apikalglied der Antennen höchstens so lang wie die 8 vorhergehenden Flagellomeren

ZUSAM EHE ee eg: nee Se she ter, Wasch: Corynoneura edwardsi-Gruppe

AR > 0.6; Apikalglied so lang wie die 6-8 Flagellomeren zusammen. Das letzte Palpen-

glied länger als die Summe der zwei ersten freien Glieder ...... C. edwardsı Brund.

AR < 0.5; Apikalglied so lang wie die 3-3.5 vorhergehenden zusammen. Das letzte

Palpenglied nicht länger als die beiden ersten freien Glieder zusammen. .............

BEN EEE RER RENNEN Ah LI DEE DAHIN So Sn ee C. brundini spec. nov.

(Das Weibchen von C. gynocera ist unbekannt).

Antenne apikal ohne Sensilla chaeticae. Das dritte Palpenglied kaum länger als die zwei

ersten kugeligen Glieder zusammen. Spermatheken proximal verschmälert, gekrümmt

(Eig.9.x4, SW EB IBIE = DUR. 1). nl tab SUR N C. carrıana Edw.

Wenigstens die distale Hälfte des Apikalgliedes der Antennen mit Sensilla chaeticae

Apikalborste (A) von Ti/P; kurz S-förmig und etwa senkrecht zur Längsachse der Tibia

SEe Te N C. scutellata Winn.

Apikalborste (A) von Ti/P; etwas gekrümmt, aber niemals senkrecht zur Längsachse der

Hibiasstehendi ee we en een Seiser ee ah Mar Dale

LR/P, < 0.45. Das letzte Palpenglied kürzer als die zwei vorhergehenden zusammen...

IE ee Rt se C. brundini spec. nov.

TRUE SO ASS ee ee

Letztes Palpenglied etwa ebenso lang wie die beiden rundlichen vorhergehenden Glieder

Letztes Palpenglied etwas länger als die beiden vorhergehenden zusammen. Auf P;

VE ee I EN N SER RE

LR/P, etwa 0.55. Drittes freies Palpenglied weniger als zweimal so lang wie breit. Clavus

wenigstens auf der distalen Hälfte mit völlig verschmolzenen Adern. Spermatheken etwa

ms Omslanehf ee, er ae hab: Sc C. arctica Kieff.

LR/P, ungefähr 0.50. Drittes freies Palpenglied etwa zweimal so lang wie breit. Clavus

wenigstens im distalen Drittel mit verschmolzenen Adern, im proximalen Teil ist R4+5

deutlich von den vorderen Adern zu trennen. Spermatheken größer (80-100 um).....

N NEL NEE EEE ERSTER SE ER al HE C. edwardsı Brund.

—

235

Puppe

Die weiblichen Puppen von C. gynocera und C. brundini sind unbekannt. Die Tabelle ist nur als ein Versuch an-

zusehen, da das vorliegende Material unzureichend ist.

1 (7) Mehr oder weniger lange, farblose Spitzchen auf dem 1. und 2. Sternit, die meisten von

ihnen mehimalslänger alsbrae# 1.2 u. a Erle 2

2 (3) Lange Spitzchen auf der ganzen Fläche der Sterniten 1 und 2. Gonopodenscheiden

reichen etwa bis zum Hinterrand des Analsegments. Scheiden der Cerci reichen etwa bıs

zu den Sockeln der medianen Analborsten ..........-......- C. edwardsi Brund.

3 (2) Spitzchen in der Mitte des 1. Sternits, wenn vorhanden, sehr klein, stets viel kleiner als

ari den, Lateralieilen tn... Harn a0 a ee re 4

5 (4) Gonopodenscheiden reichen bis zum Hinterrand des Analsegments oder darüber hinaus

sin de Ansteroen ers. hilden. Pers sea kartin.er C. arctica Kieff.

7 (1) Segment I ohne dorsale und ventrale Chagrinierung oder, wenn vorhanden, mit

winzigen Spitzchen oder Dörnchen, die höchstens 2mal so lang wie breit sind. Eine

sichere Bestimmung der Weibchen ist schwierig, oder nicht möglich... ............. 8

8 (13) Winzige Dörnchen bis zum Lateralrand der Exuvie auf den Sterniten3-5(6)......... 9

9 (10) Gonopodenscheiden auffallend groß, überragen den Hinterrand des Analsegments sehr

deutlich. Antennenscheiden des O ungewöhnlich kurz (etwa 200 um). Auf Tergit 2 sehr

wenige Dörnchen. Flügelscheiden mit vielen deutlichen Perlenreihen................

10 (9) Gonopodenscheiden kürzer. Antennenscheiden des O' 300-500 um lang........... 11

11 (12) Gonopodenscheiden mit parallelen Seiten, enden auf der Mitte zwischen dem Hinter-

rand des Analsegments und den Sockeln der medianen Analborsten. Flügelscheiden mit

wenigen Perlenreihen, bisweilen schwer zu erkennen. Am Hinterrand des 2. Segments

fehlen oft die großen Dörnchen. Antennenscheiden des O’ etwa 400-500 umlang.....

N N a ne ara re ee C. scutellata Winn.

12 (11) Gonopodenscheiden schwach konisch, reichen etwa bis zum Hinterrand des Anal-

segments. Am Hinterrand des 2. Segments oft große Dörnchen. Perlenreihen stets

deutlich erkennbar. Antennenscheiden des Q’ etwa 300-400 um lang. . C. carrıana Edw.

13 (8) Chagrinierung verhältnismäßig spärlich, reicht lateral oft nicht über die Dorsalborsten

hinaus. Gonopodenscheiden reichen kaum bis zum Hinterrand des Analsegments.....

BEREIT U SCI EAN. LIVEERGEN SER URTCAL DZ BER AO ANI N BL. C. gratias Schlee

Larven

Die Tabelle ist nur als ein Versuch anzusehen, da das vorliegende Material zu unvollständig ist. Es sei darauf hin-

gewiesen, daß sich die Antenne vom ersten bis letzten Larvenstadium verlängert. Bei den Corynoneura-Arten

scheinen darüber hinaus auch Längenunterschiede zwischen den Arten gegeben zu sein. Die längsten Antennen be-

sitzt C. celeripes Winn. (sensu Edwards nec. Schlee), sie sind etwa 2,5mal so lang ist wie der Kopf.

1 (8) Zahnleiste der Labialregion mit einem sehr kleinen unpaarigen Medianzahn zwischen

demerwa:gleiehgroßen F:ateralzähnen 22... autan) aan la. OR Nero re BE 2

2 (3) Antenne bis etwa 330-350 um lang, höchstens so lang wie der Kopf. Kopf schwach

skulmturiertt ne A UEIE KO IC. VIRDLREL ES NER LER PER C. edwardsi Brundin.

3-. (2) Antenne länger,als der. Kopf»... dert varna seta Aackelaae a Aaee 4

4 -(7). Antenne etwal/längerals der Kopt u PR ae. 22 EEE TEE 5

5 (6) Antenne etwa 370-390 um lang. Kopf schwach und unregelmäßig skulpturiert

N AR AN tete ns eh ebene C. gratias Schlee

6 (5) Antenne etwa 340-450 um lang. Der Kopf meist schwach und netzartig skulpturiert....

N ee C. scutellata Winn.

7 (4) Antenne bis 495-560 um lang, etwa um halbe Kopflänge länger als der Kopf. Kopf deut-

lichmerzartig;skulpsumiereee Sr ee ee. C. arctica Kieff.

8 (1) Der Medianzahn der Zahnleiste der Labialregion reduziert (Fig. 10). ..............-

NE nee 5 0 EIER Corynoneura spec. (? C. carrıana Edw.)

Danksagung

Originalmaterial für diese Studie stellten folgende Museen zur Verfügung: British Museum (Natural History),

London, England (Dr. P. S. Cranston); Institut Royal des Sciences Naturelles de Belgique, Bruxelles, Belgique (Dr.

P. Grootaert); Kyusyu University, Entomological Laboratory, Fukuoka, Japan (Dr. K. Yano und Prof. Dr. K.

Yasumatsu); Museum National d’Histoire Naturelle, Paris, France (Dr. L. Matile); Naturhistorisches Museum,

Zoologische Abteilung, Wien, Österreich (Dr. R. Contreras-Lichtenberg); Zoologisk Museum, Universiteti Oslo,

Oslo, Norge (Dr. J. E. Raastad) und auch das Zoologische Museum in Helsinki (Dr. B. Lindeberg). Bei den genann-

ten Kustoden und für wertvolle Informationen bei den Herren Dr. W. Tobias (Frankfurt am Main), Dr. H. Ulrich

(Bonn) sind wir zu großem Dank verpflichtet. Unser Dank gilt besonders auch Herrn Prof. Dr. E. J. Fittkau für

die sprachliche Überarbeitung des Manuskripts.

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TUISKUNEN, J. 1983: A description of Corynoneura gynocera sp. n. (Diptera, Chironomidae) from Finland. — Ann.

Ent. Fenn. 49: 100-102.

TUISKUNEN, J. & LINDEBERG, B. 1986: Chironomidae (Diptera) from Fennoscandia north of 68°N, with a descrip-

tion of ten new species and two new genera. — Ann. Zool. Fennici 23: 361-393.

WINNERTZ, J. 1846: Beschreibung einiger neuer Gattungen aus der Ordnung der Zweiflügler. — Ent. Zeit. Stettin 7:

1120:

—— 1852: Dipterologisches. — Ent. Zeit. Stettin 13: 49—58.

Elina und Dr. Mauri Hirvenoja

Sotilaskorventie 13

SF-01730 Vantaa 73

Finland

238

| SPIXIANA | Supplement 14 | 239-246 München, 15. Juli 1988 ISSN 0177 — 7424

A Review of the Genus Polypedilum Kieffer

The cytotaxonomy of Polypedilum aberrans Tshernovskji

(Diptera, Chironomidae)

By Paraskeva VI. Michailova

Abstract

The male, female and pupa of Polypedilum aberrans Tshernovskji are described for the first time. The species is

characterised cytotaxonomically. The banding patterns of the polytene chromosomes of P. aberrans and P. nubifer

are compared. Analysis of the banding pattern indicates that the relationships of these species are close. Sequences

in arms A, E, and F are apparently quite ancient, probably ancestral, because of their occurence in P. nubifer (A,/

Aı; Ey/Es; Fı/F,) P. nubeculosum, and P. aberrans. The species is female heterogametic. The females always being

heterozygous for the differential heterochromatic end on chromosome IV. The two species: P. aberrans and P. nu-

bifer are very similar but are readily distinguishable by the hypopygium of the males and by their cytology.

Introduction

Polypedilum aberrans Tshernovskji is a species known through a larva described by 'TsHERNovsKYı

(1949). Pınper & Reıss (1983) considered this species as a synonym of Polypedilum nubifer (Skuse).

The two species have alternate Lauterborn organs on the larval antenna.

The imago (C’) of P. nubifer has been reviewed by Freeman (1961), and the cytology has been des-

cribed by PorTEr & Marrın (1977). P. nubifer has been recorded from North Africa, Iraq, Srı Lanka

and Australia (Freeman, 1961).

A description of the imago and pupa of P. aberrans does not exist. Marker features of the chromo-

somes of this species have been given by MıcHa1Lova (in press). PANKRATOVA (1983) reviewed the larva

of this species and presented its known distribution: USSR, Bulgaria and Hungary.

The purpose of the present paper is to describe the adult and pupa stages of P. aberrans from mate-

rıal reared in the laboratory; to characterise this species cytotaxonomically and to show that the imma-

ture stages and cytology serve to separate P. aberrans and P. nubifer. They are readily recognisable

species. The karyotypic relationships between these species is also indicated.

Material and methods

Larval material of P. aberrans was collected in lake Durankulak, VII. 1979. Specimens of this species from Hun-

gary, collected in Kondor-tö VII. 1983 and VIII. 1986, were also studied. In the laboratory larvae grow to maturity

in about 3—4 weeks. Laboratory reared adults were used for keeping a stock of this species. The crossing of adult

midges was made by a method described earlier (MICHAILOVA, 1985).

18 specimens (100°C’ and 8Q 9), 5 pupae and 35 larvae have been examined. Adults were fixed in 70% alcohol

and the preparations made according to SCHLEE (1966). Larvae were fixed in 3:1 ethanolacetic acid and stored in a

deep freeze. The general terminology follows SAETHER (1980).

299

Karyological preparations of salivary glands and gonads of IVth instar larvae (35 specimens: 5 from Durankulak

and 30 from Kondor-tö) were prepared applying the well known acetorcein method. After analysis the slides were

made permanent by freezing in liquid nitrogen, to enable removal of the cover glass, and mounting in euparal.

The chromosomes of P. nubifer have been proposed as a standard for the comparative cytotaxonomy of the genus

Polypedilum (PORTER & MARTIN, 1977).

Following the homology with the chromosomes of this standard, the chromosomes of P. aberrans were designa-

ted as: Ist(AB), IInd (CD), IIIrd(EF) and IVth(G). Every chromosome has been divided into sections, beginning

from 1 in the left arm of every chromosome. This division is not the same as that of P. nubifer, done by PORTER &

Marrın (1977) (a photo map of each arm of P. nubifer was not given). The common sections of the chromosomes

between these species have been shown in outline in the text.

External morphology of Polypedilum aberrans

(Fir 1,2)

Imago: &

Colour: Thorax and abdomen dark brown, legs, light brown.

Thorax: Antepronotal and dorsocentral lobes well formed, dark brown, about 35 dorsocentral setae

in 2 rows. Scutellum dark brown with 20 setae in two rows. Postnotum brown.

Legs: With middle and hind tibial combs, each with one spur (Fig. 1A). Legs proportions: (um,

n=5).

F 7 zu T 2, T au

1 an & Ay 5

En 485 425 480 275 255 180 105

P, 550 440 300 160 130 85 75

PJ 600 515 400 135 190 115 90

Hypopygium: (Fig. 1B). Gonocoxite consists of two parts, distal part is as long as wide. Gonosty-

lus 2X as long as wide. Volsella apically curved. Anal pofnt is straight from the very beginning. Anal

field oval. Coxpodeme well developed. Phallapodeme rod-like.

Imago: ?

Colour: light brown

Genitalia. (Fig. 1C). Gonosternite oval, with long setae and darkly coloured edges. Gonapophysis

IX well formed, reaches almost the middle of SV II. Coxosternapodeme dark, dense and arched.

Pupa:

Exuvia light brown. 2. segment in a lower part with a row of setae. 3.— 7. segments with a group of

setae in an upper and lower part. Posterolateral spurs of segment 8. with 8 or 9 spines (Fig. 1D).

Larva:(Fıg.2A,B,G).

Antenna, labrum, mandible and premandible as in Tshernovskjt (1949, fig. 47) and PAnKRATOVA

(1983, fig. 201).

Fig. 1. Polypedilum aberrans. A, Hind leg with comb and spur. B, Hypopygium; Ca — coxapodeme; Pha — phal-

lapodeme; AnP — anal point; Ge — gonocoxite; Gs — gonostylus. C, Genitalia (P); Gca — Gonocoxapodeme;

Va — vaginal apodeme; Ce — cercus. D, Pupa: 8. segment, posterolateral spur with 8 spines.

240

Fig. 2. Polypedilum aberrans. A, Antenna. B, Labrum. C, Mandible. D, Heterozygous inversion ın arm B.

The karyotype of Polypedilum aberrans

(Fig.3 A,B,@,D)

2n = 8. Chromosome I (AB), and II (CD), Ist (AB), IInd (CD) chromosomes — metacentric; Chro-

mosome III (EF) — submetacentric and IV (G) — acrocentric. Every chromosome with a dark hetero-

chromatin band in the centromere region. Sex chromosomes were not found. The species is female he-

terogametic. The females always being heterozygous for the heterochromatic end on the chromosome

IV.

Chromosome polymorphism has been observed only in arm B of Ist chromosome.

The chromosome markers have been described by MıcHA1LovA (in press).

Chromosome I with arms AB similar to its counterpart in P. nubifer. Porter & Marrın (1977) re-

ported five rearrangements in arm A. Only two ofthem are in homozygous condition. P. aberrans has

only one type of band sequence. Part of which coincides with that of P. nubifer (A,/A,) and part —

with Ay/A..

Chromosome I

P.inubifer v2 3 4 5 56, 7.829 10T zes

P. aberrans) 1221 3 445.6 . 789 Pia ee

B.jnubtjerse 1, 24..32 2,559:6-.40,28,7 9.10.02 Ile, 22015

242

Arm B of P. nubifer has four rearrangements in heterozygous condition (PoRTER & Marrın, 1977)

on both populations of P. aberrans, we found only one type of heterozygous inversion with a very

low frequency (Fig. 2D).

Chromosome II, with arms CD showing considerable change from the sequences seen in P. nubi-

fer. Three rearrangements have been recognized in arm C of P. nubifer (Porter & Martın, 1977). In

arm C of P. aberrans only onetype of band sequence was found. The two groups of dark bands in sec-

tion 2-3 of P. aberrans exist in P. nubifer, described by Porter & Marrın (1977) as C,/C, and Cy/C,.

The other sequences of P. aberrans must have changed considerably in appearance compared with the

standard. Arm C shows a sequence not previously reported.

Chromosome II

Br nubi fer Nor: I 22 723722526

EHE

P. aberrans 1 ZB le Dt 8 5

Ban nubierwnlleeo lar NETTO ZN FE E24 257 26

Arm D is considerably changed in P. aberrans.

Chromosome III, with arms EF. There are four rearrangements in arm E of P. nubifer (PoRTER &

Marrın, 1977). All have been seen in the homozygous condition. In P. aberrans only one type of band

sequence was seen in which the banding pattern on section 1-6 is very similar to that of E/E; of P. nu-

bifer. The bands after this sequence appear to differ from that of P. nubifer.

There were four rearrangements in arm F of P. nubifer (Porter & Marrın, 1977). Two ofthem have

been seen in the homozygous conditions. Arm F of P. aberrans has only one type of sequence. Section

9—11 of P. aberrans ıs identical to F,/F, described by Porter & Marrın (1977) for P. nubifer.

Chromosome II

P. nubifer zZ 730 DEZE 27

NE |

P. aberrans 1 2 3 4 5 6 7

P. nubifer 31 52 335 1.35.1358

en)

P. aberrans 8 9 10 11

Chromosome IV, or arm G, appears similar to P. nubifer. When a detailed analysis of the chromo-

some is attempted the bands in section 4 of P. aberrans can be positively correlated with those in the

middle of P. nubifer.

Chromosome IV

P. nubifer SE 158,059 EERA0F Fi

P. aberrans 1 2 3 4 5 6

PoRTER & Marrın (1977) reported six rearrangements in arm G of P. nubifer. In P. aberrans there

are only two types of sequences which are in one and the same salivary gland. These differ in their

functional activity.

245

P. aberrans is female heterozygous for a heterochromatic end (Fig. 3 D, G,), while males are homo-

zygous for this. The female rearrangements are similar to G,/G; of P. nubifer.

B ”

Fig. 3. Polypedilum aberrans. A, Chromosome Ist (AB). B, Chromosome IInd (CD). C, Chromosome IlIrd

(EF). D, Chromosome IVth (G, or G,); N — nucleolus; BR — Balbiani ring; c — centromere region.

244

Discussion

Detailed study of adults of P. nubifer and P. aberrans indicated that there are two different species.

These species are very similar but are readily distinguishable by the male hypopygium. The coxite of

P. nubifer consists of two parts. The second part ıs strongly prolongated (Frezman, 1961). The second

part of coxite of P. aberrans ıs as long as wide. Between these parts in P. aberransthere isa well formed

concavity (Fig. 1B). Such a concavity is not seen in P. nubifer (Freeman, 1961). The volsella of

P. aberrans ıs terminally recurved (Fig. 1B). The hypopygium of P. aberrans differs from that of

P. nubifer by having broader gonostyli while the gonostyli of P. nubifer are more slender (Freeman,

1961). The two species differ in the shape of the anal point. That of P. nubifer is narrower basally while

the anal point of P. aberrans ıs straight from base to apex (Fig. 1B).

Cytogenetical investigations of material which keys out as P. nubifer (Porter & Marrın, 1977) and

P. aberrans revealed the existence of two readly recognisable species with four chromosomes in the sa-

livery gland. The banding patterns of P. aberrans and P. nubifer also indicate a close relationship. The

salivary gland chromosomes of P. aberrans show sufficient similarity in banding pattern to those of

P. nubifer, mainly ın arms A, E, and F. In the genus Polypedilum as in the genus Chironomus arms A,

E, and F seem to have been more stable in evolution. Arms B, C, D, and G as we have seen show more

differences between the species. However within the arms it is not yet possible to suggest homology

of more than a few of the most conspicious band groups. Every species has marker features on each

chromosome. Some sequences differ between these species by a simple inversion.

There is no cytological evidence and external morphological data to support the suggestion that

P. aberrans should be considered as a synonym of P. nubifer.

In the present case a few species of Polypedilum have been studied cytologically: P. nubifer (Porter

& Marrın, 1977), P. aberrans, P. nubeculosum and Polypedilum sp. (Chironominae genuinae N3 Li-

pina) (MicHaA1Lova, in press). This allows determination of the relative age of some sequences. Phylo-

genetically central sequences with wide distribution are more ancient than sequences with restricted

distribution. With the exception of the G chromosome, the other chromosomes of P. aberrans and

P. nubeculosum have only one type of band sequence. PoRTER & Marrın (1977) reported few rearran-

gements for arms A, C, E, F, G of P. nubifer. One of them is common for P. nubifer, P. aberrans and

P. nubeculosum. The banding pattern of P. aberrans arm A, section 2-5 and the banding pattern of

the same arm of P. nubifer, section 2-4a9 (A,/A,) are common for the species. In arm C there are also

common patterns: bands in section 2-3 of P. aberrans and 15 (C,/C, or C,/C,) of P. nubifer. The

banding pattern of arm E of P. aberrans, section 2-6 corresponds with that of P. nubifer, section

27—-30-27al (Ey/E;). Arm F: the banding pattern, section 9-11 (P. aberrans) and section

33c9—-36b12 (P. nubifer) is common for both species. The banding pattern (,)ofarm G (P. aberrans)

and that in the middle of arm G (P. nubifer) are common.

These band sequences have been found in P. nubecnlosum also (MicHa1LovA, in press). These pat-

terns could be considered as “basic patterns” of genus Polypedilum in sense of Würker (1980). Perhaps

these common patterns existed in a hypothetical stem species. Starting from the hypothetical species

these patterns have been retained in one species but in an other (P. nubifer) have undergone different

mutations. On that way P. nubifer displays a high frequency of chromosomal rearrangements. There

is an other explanations also: the common ancestor was polymorphic for anumber of sequences which

still occur as polymorphism only on P. nubifer.

PoRTER & Marrın (1977) reported a female heterogamety of P. nubifer. They have also found hete-

rozygous males. A more lıkely explanations is that the species is still in the process of changing from

the normal male heterogamety to female heterogamety (PorTEr & Marrın, 1977). In P. aberrans this

process ıs going forward. Only females always carry the large heterochromatinized differential seg-

ment.

245

Literature

FREEMAN, P. 1961: The Chironomidae (Diptera) of Australia. — Aust. J. Zool. 9: 611-737

MICHAILOVA, P. 1985: Method of breeding of the species from the Family Chironomidae, Diptera in experimental

conditions. — Compt. rendus de !’Academie Bulgare des Sciences 9: 1179— 1181

—— (in press): Polytene chromosomes and their significance for the Systematics ans Phylogeny of the family

Chironomidae, Diptera. — Izd. Bulgarian Academy of Sciences, Sofia.

PANKRATOVA, V. Ya. 1983: Larvae and pupae of midges of the subfamily Chironomidae (Diptera, Chironomidae

= Tendipedidae) of the USSR fauna. — Izd. “Nauka”, Leningr., 295 pp.

PINDER, C. V. & F. Reıss 1983: The larvae of Chironominae (Diptera, Chironomidae) of the Holarctie region —

Keys and diagnoses. — Ent. Scand. Suppl. 10: 293—435

PORTER, D. L. & J. MARTIN 1977: The Cytology of Polypedilum nubifer (Diptera, Chironomidae). — Caryologia

1:41—-62

S£THER, ©. A. 1980: Glossary of Chironomid morphology terminology (Diptera, Chironomidae). Ent. Scand.

Suppl. 14:51 pp.

SCHLEE, D. 1966: Präparation und Ermittlung von Meßwerten an Chironomidae (Diptera). —- Gewäss. Abwäss. 41/

42: 189—193

TSHERNOVSKJI, A. A. 1949: The key of larvae of midges of the family Tendipedidae. — Izd. Acad. Nauk. USSR,

Moskow and Leningr. 185 pp.

WÜLKER, W. 1980: Basic patterns in the chromosomes evolution of the genus Chironomus (Diptera). — Z. zool. Sy-

stematik Evolutionsforschung 2: 112-123

Dr. Paraskeva Michailova,

Institute of Zoology, Bulgarian Academy of Sciences, 1,

Russky Blvd., 1000-Sofia, Bulgarıa

246

SPIXIANA | Supplement 14 | 247-252 | München, 15. Juli 1988 ISSN 0177— 7424

Schineriella schineri gen. nov., comb. nov., placement of

Tanypus schineri Strobl 1880

(Diptera: Chironomidae)

By D. A. Murray and E. J. Fittkau

Abstract

The new genus Schineriella is erected for the species Tanypus schineri Strobl 1880. Diagnoses for the adult male

and pupa are given. The type species is, by monotypy, Schineriella schineri (Strobl).

Introduction

The species Tanypus schineri was recognised by StrosL (1880) from material collected and pre-

viously identified by ScHmer (1862) as Tanypus binotatus (Wiedemann). The adult male is readily re-

cognised by its distinctive abdominal pigmentation pattern and was included by Epwarps (1929) and

GOETGHEBUER (1936) in keys to the British and Palaearctic chironomid fauna as Pentaneura schineri

and Ablabesmyıa schineri respectively.

In the revision of European Tanypodinae Fittkau (1962) indicated that the adult male of this species

was sufficiently distinct to warrent its assignment to a new genus within the Pentaneurini. However,

he refrained from erecting a new genus at that time in the absence of juvenile material and referred to

the taxon as Pentaneurini gen ? Schineri. In more recent years pupae and pupal exuviae of this species

have been obtained in East Holstein and Southern Bavaria, Germany (leg. Reiss). Frrrkau and Murray

(1986) included a diagnosis for the pupal stage (as Tanypodinae Genus II) in the recently published

keys and diagnoses to chironomid pupae of the Holarctic Region (WıEDErHOLM, 1986).

The new genus Schineriella is now erected for this species and generic diagnoses are given for the

adult male and pupa.

Schineriella gen. nov.

Type and only included species: Schineriella schineri (Strobl 1880) by present designation.

Generic diagnosis

Imago C:

Small to medium sized species, winglength 2.5—3.0 mm, terminal antennal flagellomere noticably

narrower and distinctly set off from penultimate falgellomere. Scutal tubercle absent. Wings unmar-

ked, R,,; absent, costa not produced, anal lobe not developed. Anterior tarsus with beard, tibial spurs

with main tooth and 3—4 side teeth. Tibial comb of 6-7 setae on hind leg. Volsella absent, transverse

sternapodeme pointed anteriorly.

Pupa:

Thoracic horn somewhat swollen, “sausage” shaped; horn sac fills most of the lumen; plastron plate

laterally displaced, circular and only 0.1%X horn length. Shagreen spines narrow, sparse. Posterior bor-

der of tergite VIII projects backwards over anal lobe. Segment VII with only 3 LS setae, all in the distal

!/2. Anal macrosetae with adhesive sheaths.

Larva:

The larva is as yet unknown

Generic description

Imago Cd:

Small to medium sized species, wings 2.5—3.0 mm long. Body pigmentation cuticular; thorax

brownish, vittae and median anepisternum somewhat darker; legs pale yellow; abdomen varıously

pigmented.

Head: Pale; eyes with dorsal, parallel-sided, extension, minimum width of eye bridge with 4 ocelli;

temporal setae uniserial, inner verticals beginning level with apex of coronal triangle and merging im-

perceptably with outer verticals and post orbitals. Palps almost as long as the antennae and 1.5x as

long as head width; palpomeres 3 and 4 equal in length and almost 2x as long as palpomere 2. Cly-

paeus long, narrow, slightly more than 2X as long as broad. Posterior tentorial pit close to tip of ten-

torıum. Antenna with 14 flagellomeres; terminal flagellomere noticably narrower than penultimate

flagellomere, 4—5xX as long as broad and more or less cylindrical. AR about 2.0.

Thorax: Antepronotum reduced, scutal tubercle absent. Dorsocentral setae more or less uniserial

between the vittae. Acrostichals biserial, almost reaching to the pre-scutellar field; scutellar setae bise-

rıal. Scutal tubercle absent, 3—4 flagelliform setae present in mid scutal region. Postnotum round

posteriorly.

Wing: Membrane with macrotrichia, unpigmented. MCu beyond FCu, RM close to MCu, R; ;; ab-

sent, Ry;; close to Ri and costa. Costa not produced beyond end of R,,; and ending clearly before tip

of M, ,, midway between M, ‚, and M;,,. Anal lobe not developed.

Legs: Anterior tarsus with distinct beard, LR 0.91; tibial spur sinuous, with main tooth, reaching

to 0.5X spur length, and 3—4 side teeth. Outer spur small on mid and hind legs, hardly reaching 0.5x

inner spur length. Tibial comb, of 6-7 setae present on hind leg. Pulvilli absent, empodium approxi-

mately as long as the terminally spatulate claws.

Hypopygium: Anal point large, triangular and broadly rounded apıcally. Gonocoxite more or less

cylindrical, slightly swollen basally; dorsal and median surfaces setose on the distal 2/3, longest seta

equal to, or slightly longer than, the gonocoxite width; entire surface with a more or less uniform co-

vering of macrotrichia which are more dense in the anterior median field. Gonostylus slender, swollen

basally and distally narrow, reaching 3— 3/4 gonocoxite length. Terminal spur expanded medially, apı-

cally curved and pointed. Transverse sternapodeme distinctly pointed anteriorly.

Pupa:

Medium sized species, exuviae 6.0 mm in length and light, golden brown in colour.

Cephalothorax: Thoracic horn somewhat swollen, 3.0x as long as broad, slightly arched and more

or less “sausage” shaped. External membrane with solitary spines. Horn sac fills most of the horn lu-

men, narrow basally, gradually expanded and connected to a small plastron plate by a narrow, sı-

nuous, neck. Plastron plate slightly laterally displaced, set on a small tubercle, more or less circular ın

outline and only 0.1X horn length. Thoracic comb of 10-12 elongate, distally round or pointed, tu-

bercles. Basal lobe distinct, conical and round apically. Thoracicmembrane otherwise smooth. Thora-

cic setae filamentous and distally round, Dc, 5.0X as long as Dc,, Sa slightly longer than Dec.

Abdomen: Elongate, pigmented scar on tergite I. Shagreen spines narrow, solitary and relatively

sparse. Posterior border of tergite VIII projects backwards over the anal lobe. Abdominal setation; D,

V and L setae filamentous, distally round, D3 unusually long. Segment VII with only 3 LS setae,

248

Figs. 1.6. Schineriella schineri gen. nov., comb. nov., adult male: 1. head; 2. terminal antennal flagellomere;

3. flagelliform setae on scutum; 4. wing; 5. tibial spurs; 6. hypopygium.

0.75x segment length, all in the distal 1/2. Segment VIII with 5 LS setae, 1.5% segment length. Anal

lobe 1.5x as long as broad. Points evenly tapered. Outer border toothed in the distal !/2, inner border

smooth. Anal macrosetae, with adhesive sheaths, arising from the middle !/3 of the outer border.

Larva:

The larva is so far unknown.

Schineriella schineri (Strobl) comb. nov.

Tanypus schineri Strobl 1880 p. 55

Pentaneura schineri Edwards 1929 p. 294

Ablabesmyia schineri Goetghebuer 1936 p. 45

Pentaneurini gen ? schineri Fırtkau 1962 p. 275

Tanypodinae “Genus Il” Fittkau & Murray, 1986 p. 64

249

Figs. 7.—13. Schineriella schineri gen. nov., comb. nov., pupa: 7. thoracic horn, basal lobe and thoracic comb;

8. thoracic setae; 9. tergite I; 10. segment IV; 11. shagreen; 12. segments VI-VIII and anal lobe; 13. anal lobe and

female genital sacs.

250

Imago © (Figs. 1-6)

Additional to generic diagnosis: Temporal setae 18, including inner and outer verticals and post-

orbitals. Clypaeus with 21 setae. Lateral antepronotal setae 4—5, humerals 3-4, dorsocentrals 20-22,

acrostichals 36, prealars 7-8, supra alars 1. Abdominal tergites I-III and VI pale yellow, IV and V

mostly dark brown with a pale posterior band, VII and VIII entirely dark brown. Tergite IX with a

posterior transverse row of 12-15 backwardly directed setae.

Pupa:

See generic diagnosis and Figs. 7-13

Material studied: Imagines, 10°, Murnauer Moos, Oberbayern, Germany 11/7/77; 10 same localıty 21/6/78,

both leg. Reiss; 1°, Hart Saltacher See, Oberbayern, 16/6/80, leg. Plassmann; 1 © Province of Adiyaman, Turkey,

21/6/85, leg. Schacht; 10° Witherslack, Westmorland, Great Britain, leg. Edwards, V1/1929; 10° Pe, Krebssee,

Murnauer Moos, Oberbayern, 11/7/77, leg. Reiss; 19 Im and Pe (reared), Unter Ausgrabenssee, Plön, Holstein,

Germany, 20/5/74 leg. Reiss.

Distribution and Ecology

Records of S. schineri exist from Austria (Strogı 1880), Belgium (GoETGEHBUER 1936), Great Britain

(Epwarps 1929) and more recently from Holstein and Bavarıa, Germany (leg. Reiss) and the Province

of Adiyaman, Turkey (leg. Schacht). Although the larva has yet to be found it is likely to occur in se-

diments of small, nutrient rich ponds and small lakes containing much decayıng leaf litter. Reiss (pers.

comm.) has reared a pupa from such sediments taken from a depth of 1.5 m.

Systematic position of Schineriella

The new genus clearly belongs to the tribe Pentaneurini within the Tanypodinae. Fırrkau (1962)

tentatively indicated a relationship between Pentaneurini gen ?schineri and Krenopelopia on the basıs

of similarity in structure of the tibial spurs and male hypopygium. However, adult male imagines of

the new genus are clearly separable from Krenopelopia by the presence in the latter of a well developed

wing vein R,;,, which clearly divides into its constituent veins R, and R;. Moreover, the pupal stages

of Krenopelopia and Schineriella are quite different, especially with regard to the thoracic horn which

is trumpet shaped and with a well developed plastron in Ärenopelopia in contrast to the bulbous horn

and reduced plastron in Schineriella (see also Fırrkau and Murray 1986 under Tanypodinae “Ge-

nus II”).

Within the tribe Pentaneurini wing vein Ry,; is absent or reduced, together with Schineriella, in the

genera Labrundinia Fittkau, Monopelopia Fittkau and Nilotanypus Fittkau. However, adults of the

latter genera have either only a single tibial spur or no spur (Labrundinia) on the mid and hind legs ın

contrast to the two tibial spurs present on the mid and hind legs in Schineriella. The pupa of Schine-

riella most closely resembles Zabrundinia in the form of the thoracic horn but may be readily separa-

ted by the backwardly projecting corners of tergite VIII and the presence of only 3 LS setae on tergite

VII in Schineriella. The overall similarity is also apparent in the adult stage but apart from the tibial

spurs further differences are evident in the terminal antennal flagellomere which is only weakly set off

from the penultimate flagellomere in Zabrundinia in contrast to the well differentiated and distinctly

set off, narrower, terminal flagellomere in Schineriella. Additionally, the abdominal pigmentation is

cuticular in Schineriella while Labrundina has subcuticular pigmentation.

251

Literature

EDWARDS, F. W., 1929: British bon-biting midges (Diptera: Chironomidae). — Trans. Ent. Soc. London, 77, 2:

279-430.

FiTTkAu, E. J., 1962: Die Tanypodinae (Diptera, Chironomidae). Die Tribus Anatopyiini, Macropelopiini und

Pentaneurini. Abh. Larvalsyst. Insekten 6: 453 pp. Akademie Verlag. Berlin.

FITTKkau, E. J.& D. A. MuRRAY, 1986: The Pupae of Tanypodinae (Diptera: Chironomidae) of the Holarctic region

— Keys and diagnoses. Ent. scand. Suppl. 28: 31-113.

GOETGHEBUER, M., 1936: Tendipedidae (Chironomidae). a) subfamiliae Pelopiinae (Tanypodinae). A. Die Imagi-

nes. — In: LINDNER, Die Fliegen der Palaearktischen Region, 13b: 1-50.

SCHINER, 1862: Fauna Austriaca: Die Fliegen (Diptera) Il.

STROBL, 1880: Progr. Gymnas. Seitenstetten 55.

WIEDERHOLM, T., 1986: (Ed.) Chironomidae of the Holarctic region. Keys and diagnoses. Part 2, Pupae, Ent.

scand. suppl. 28: 1—482

Dr.D. A. Murray,

Department of Zoology, University College Dublin,

Belfield, Dublin 4, Ireland.

Prof. Dr. E.J. Fittkau,

Zoologische Staatssammlung,

Münchhausenstr. 21, D-8000 München 60,

Bundesrepublik Deutschland.

252

SPIXIANA Supplement 14 253—259 | München, 15. Juli 1988 | ISSN 0177-7424

Bethbilbeckia floridensis: a new genus and species of

Macropelopiini from the South Eastern Nearctic

(Diptera: Chironomidae)

By E. J. Fittkau and D. A. Murray

Abstract

The genus Bethbilbeckia ıs established for a species of Macropelopiini from the South Eastern United States. Ge-

neric diagnoses for the larva, pupa and adult male are given together with descriptions of all life stages of the single

included new species Bethbilbeckia floridensis spec. nov.

Introduction

While reviewing some types and material of tanypodine species in connection with the ongoing co-

operative work on keys and diagnoses to genera of the holarctic Chironomidae (Wiederholm 1983,

1986 et seq.) some Macropelopiini specimens were kindly donated by Elizabeth and Bill Beck, Jack-

sonville, Florida. The material in question had been reared from the larval stage and thus larval and pu-

pal exuviae and adult male imagines were available for study. Such significant differences in morpho-

logy exist, in all life stages, that the specimens cannot readily be associated with any currently recog-

nısed genera. A description of the pupal exuviae of these specimens was given as Tanypodinae “Ge-

nus I” in Fırrkau and Murray (1986). It gives us much pleasure to name the genus after Elizabeth and

Bill Beck as a gesture of appreciation for their contribution to chironomid science. A complete generic

diagnosis for the larva, pupa and male adult of Bethbilbeckia gen. nov. and descriptions of Bethbil-

beckia floridensis spec. nov. are given ın this paper.

The terminology and abbreviations used in the descriptions follow SAETHER (1980). The slide mounted holotype

and one paratype is deposited in the collections of the Academy of Natural Sciences of Philadelphia; a second para-

type is deposited in Zoologische Staatssammlung, Munich.

Bethbilbeckia gen. nov.

Type species. Bethbilbeckia floridensis spec. nov., by present designation.

Generic description

Imago C:

Medium sized species, winglength about 3.0 mm; Antennae with 14 flagellomeres, apical flagello-

mere indistinctly set off from preapical, A. R. about 2.9; eyes with dorsal extension; Temporal setae

uniserial; antepronotum well developed, lobes well separated medially ; scutal tubercle distinct; lateral

antepronotal, anepisternal, preepisternal and postnotal setae present. Wings unmarked, membrane

evenly covered with macrotrichia, MCu almost directly above FCu on M,;‚ ‚, costa strongly produced,

255

R,., present and forked, R2 distinct, anal lobe well developed; tibia I without comb, tibia III with

comb of 4-5 setae; tibial spurs with main tooth and 9—13 side teeth; pulvilli absent. Tergite IX with

strong setae posteriorly; gonocoxite more or less cylindrical, evenly setose and with pocket-like lon-

gitudinal depression anteriorly on inner border; volsellae absent; gonostylus abruptly bent through

90° near base and tapering towards apex.

Imago 9: Unknown.

Pupa:

Medium sized, approximately 7.0 mm long, brownish in colour; thoracic horn tubular, expanding

gradually from base to apex, 4.0X as long as maximum apical width; horn sac thinwalled, not quite fil-

ling the horn lumen, plastron plate oval, slightly longer than broad, 0.25% horn length; thoracic comb

and basal lobe absent; thoracic setae simple, pointed or round apically; scar on tergite I elongate and

pigmented; shagreen spines short, blunt and partially serially arranged in groups of 2-4; abdominal

setae D, on segments II-VII large, distinct and arising from very large and prominent tubercles; D,,

D,; on segments III- V arısing from small tubercles; 4 short LS setae on segment VII, 0.25% segment

length; segment VIII with 5 LS setae, 0.75% segment length; anal lobe longer than broad, with simple

spine shagreen laterally, outer border fringed with long seta-like spinules and more or less convex, spi-

nules reduced to indistinct decumbent spines at the distal end; inner borders divergent, without fringe

but with 5-7 preapical decumbant spines; anal macrosetae arise from the basal !/4.

Larva:

Antenna 5 segmented, 1.25% as long as mandible, basal segment 8x as long as basal width, ring or-

gan at apical !/, AR about 9; ring organ at basal !/s of basal palp segment; mandible slender, curved,

basal tooth distinct; ventrolateral setae 1 and 3 simple, 2 bifid; dorsomentum medially almost reaching

the pseudoradula; ligula with 5 teeth; tooth row distinctly concave; inner teeth straight; paraligula bi-

fid, pecten hypopharyngis with about 14 teeth; posterior parapods with normal and 3—4 wide claws.

Bethbilbeckia floridensis, spec. nov.

Imago J (Figs. 1-6.)

Head: Pale brown, pedicel darker; eyes with dorsal extension. Frontal setae present, inner verticals

6-7, outer verticals 7—8, post orbitals 5-6. Antennal ratio 2.9; pedicel with three anteroventral and

two lateroventral setae; flagellum with 14 flagellomeres, terminal flagellomere more or less conical,

1.5x as long as basal width.

Thorax: Brownish with vittae slightly darker; antepronotum well developed, lobes separated, with

9 lateral antepronotal setae; humerals XX; dorsocentrals irregularly biserial; supraalars 22, prealars

20-22; acrostichals biserial; preepisternals 4-6; anepisternals 2-4; scutellum with about 30 setae;

postnotals 10 on either side; scutal tubercle distinct.

Table 1. Leg measurements (t) and ratios for the holotype (a) and paratype (b) of B. floridensis spec. nov.

Leg le Tı ter a Ta 3 Ta 4 Ta5 LR BV

la 947 1026 774 379 205 190 142 0.75 2.99

b 1010 1184 900 442 300 190 142 0.76 2.88

IIa 916 1011 490 268 221 142 110 0.48 3.26

b 1105 1200 663 315 245 186 142 0.55 3.24

Illa 868 1176 781 418 300 201 142 0.66 39%

b 1026 1374 908 _ _ _ _ 0.66 =

Figs. 1-6. Bethbilbeckia floridensis gen. nov., spec. nov., adult male: 1. head; 2. terminal antennal flagellomere;

3. scutal tubercle; 4. wing; 5. tibial spurs; 6. hypopygium.

Wing: About 3.0 mm long, with macrotrichiae; MCu on M3;4 close to FCu; RM removed from

MCu by length of MCu; R,, ; present and forked, R; distinct; costa distinctly produced beyond R;;;.

Legs: Pale, with faint indication of bands on apices of femora and bases of tibiae; spur Ti I with 13

side teeth, comb absent; spurs Ti II, Ti II with 9-10 side teeth; apex Ti III with comb of 4-5 setae;

pulvilli absent; claws terminally pointed; leg ratios in Table 1.

Abdomen: Uniformly brown, densly setose; ninth tergite with posterior multiserial row of 22-27

setae. Hypopygium (Fig. 6.) anal point triangular; gonocoxite more or less cylindrical, evenly setose

and with pocket-like longitudinal depression anteriorly on inner borders; gonostylus setose, swollen

basally and bent anteriorly through 90° near base.

Female. Unknown.

Pupa (Figs. 7-11.)

Medium sized, approximately 7.0 mm long, brownish in colour.

Thoracic horn tubular, relatively narrow, expanding gradually from base to apex, 4.0X as long as

maximum apical width; external membrane smooth, with few spinules; horn sac thin-walled, not

quite filling the horn lumen, evenly expanded towards the apex and connected to a more or less oval

shaped plastron by two short necks; plastron plate slightly longer than broad, 0.25 horn length.

Thoracic comb and basal lobe absent, thoracic membrane with transverse ridges extending to the me-

dian suture. Thoracic setae Dc, simple, pointed; Dc, extremely small, rounded, approximately 0.25 as

long as Dc, ; Sa simple, long and pointed, 3.5X as long as Dc..

255

er Ze u

Bizs-2 12:

Bethbilbeckia floridensis gen. nov., spec. nov., pupa: 7. thoracic horn; 8. thoracic setae; 9. tergites

I-III; 10. segments IV-VIII and anal lobe; 11. shagreen; 12. anal lobe and © genital sacs.

256

Abdomen: Scar on tergite I elongate and pigmented; shagreen spines short, blunt and partially se-

rıially arranged in groups of 2-4. Abdominal setae D, on segments II-VII large, distinct and arising

from very large and prominent tubercles; remaining D and V setae of varying sizes, D,, D; on seg-

ments III—V arising from small tubercles; segments I-VII with 2 L setae; 4 short LS setae on segment

VII, 0.25% segment length; segment VIII with 5 LS setae, 0.75x segment length. Anal lobe longer

than broad, with simple spine shagreen laterally, outer border fringed with long seta-like spinules and

more or less convex, spinules reduced to indistinet decumbent spines at the distal end; inner borders

divergent, without fringe but with 5-7 pre-apical decumbant spines. Anal macrosetae 0.5X segment

length, arise from the basal !/4.

Larva. (Figs. 13-18.)

Head capsule yellowish; antenna 5 segmented, 1.25X as long as mandible, basal segment 8x as long

as basal width, ring organ at apıcal !/s, AR about 9; basal segment of maxillary palp about 3x as long

as wide, ring organ at basal '/3. Mandible slender, curved, basal tooth distinct with apically directed

crest extending over inner margin and smaller inner tooth; ventrolateral setae 1 and 3 simple, 2 bifid.

Dorsomentum on each side with 5—6 side teeth, inner part extending medially and almost reaching the

pseudoradula. Ligula about !/3 longer than apical width, with 5 teeth; tooth row distinctly concave,

outer tooth 2.5x middle tooth; inner teeth straight. Paraligula bifid, 0.5x as long as ligula. Pecten hy-

popharyngis with 14 teeth, inner tooth large, medially directed and with a low rounded protuberance

on outer border. Posterior parapods with normal and 3—4 wide claws.

Figs. 13.18. Bethbilbeckia floridensis gen. nov., spec. nov., larva: 13. antenna; 14. maxillary palp; 15. mandible;

16. mentum; 17. ligula and paraligula; 18. pecten hypopharyngis; 19. Claws of posterior parapod.

(

257,

Material studied: Holotype; Larval and pupal exuviae and adult male (reared) slide mounted in Euparal in the col-

lections of the Academy of Natural Sciences of Philadelphia (ANSP). Coll. E. & W. Beck, Peter’s Creek, Clay

County, Florida 6.7.68.

Paratypes: 1 associated larva-pupal-adult male, slide mounted, in coll. ANSP; 1 associated larva-pupa-adult

male, slide mounted in coll. Zoologische Staatssammlung, Munich, West Germany. Both paratypes were collected

at the same site as the holotype.

Systematic position

The new genus clearly belongs to the tribe Macropelopiini which was recently enlarged to include

the genus Radotanypus (Fırrkau and Murray 1985) and is now considered to be composed of the ge-

nera Psectrotanypus Kieffer, Derotanypus Roback, Alotanypus Roback, Brundiniella Roback, Macro-

pelopia Thienemann, Radotanypus Fittkau and Murray, Apsectrotanypus Fittkau, Fittkanimyia

Karunakaran and Bethbilbeckia. The larva of Derotanypus and Psectrotanypus differ from those of all

other genera in the tribe by having only 4 teeth in the ligula in contrast to the more usual arrangement

of5. It is thus not necessary to consider these two genera further in the present context. In Fırrkau and

Rosack (1983) the larva keys easily to couplet 12 which leads to the genera Alotanypus, Brundiniella,

Apsectrotanypus and Macropelopia. Alotanypus differs from these and the recently described larva of

Radotanypus (Erıer 1986) in having all ventrolateral mandıbular setae simple while in the remaining

above mentioned genera ventrolateral setae 1 and 3 are multibranched. In Bethbilbeckia setae 1 and 3

are simple and seta 2 is clearly bifid only. The pupa of Bethbilbeckia ıs readily separable from all other

members of the Macropelopiini. Although superficially similar to Macropelopia it is easily distinguis-

hed by the prominent, straight D, setae which arise from distinct tubercles- larger than those present

in other Macropelopiini. The very short, weak, LS setae on segment VII are also diagnostic for the ge-

nus. In the key to genera of holarctic tanypodine pupae Bethbilbeckia has been included as “Tanypo-

dinae Genus I” (Frrrkau and Murray 1986 p. 64, Fig. 5.46).

In Rosack (1972) the adult male keys to couplet 9 which includes Macropelopia decedens (Walk.

and Parapelopia serta Roback. The former is “a large northern species” in the U.S.A. while the latter

is a “small Florida species” (Rosack op. cit p. 87). Rogack (1982) has already drawn attention to the

possible synonomy between Parapelopia Roback and Fittkauimyia Karunakaran. The adult male of

Bethbilbeckia differs from all other Macropelopiini, with the exception of Fittkanimyia in the unise-

rial arrangement of the temporal (inner and outer vertical and postorbital) setae. However Bethbil-

beckia may be easily separated from Fittkauimyia in having preepisternal setae and a posterior multi-

serial row of setae on tergite IX oftheabdomen. An additional differential character is seen in the tibial

spurs where Bethbilbeckia has a maximum of 13 side teeth compared with 13—20 in Fıttkauimyıa.

Literature

EPLER, J. H. 1986: The larva of Radotanypus submarginella (Sublette) (Diptera, Chironomidae). Spixiana 9:

285-287.

FITTKAU, E. J. & S. S. ROBACK 1983. The larvae of Tanypodinae (Diptera: Chironomidae) of the Holarctic region

— Keys and diagnoses. In T. Wiederholm (Ed), Chironomidae of the Holarctic region — Keys and diagnoses.

Part I, Larvae. Ent. scand. Suppl. 19: 34— 110.

—-— &D. A. Murray 1985. Radotanypus a new genus of Tanypodinae from the Nearctic (Diptera, Chironomi-

dae). — Spixiana Suppl. 11: 209-213.

-— & —--— 1986. The pupae of Tanypodinae (Diptera: Chironomidae) of the Holarctic region — Keys and dia-

gnoses. In T. Wiederholm (Ed), Chironomidae of the Holarctic region — Keys and diagnoses. Part 2, Pupae.

Ent. scand. Suppl. 28: 31-113.

ROBACK, 5. S. 1971. The subfamily Tanypodinae in North America. Monogr. Acad. Nat. Sci. Phila. 17: 1-410.

258

—— 1982. The Tanypodinae (Diptera: Chironomidae) of Australia II. Proc. Acad. Nat. Sci. Phila. 134: 80-112.

SAETHER, O. A. 1980. Glossary of Chironomid Morphology Terminology. Ent. scand. Suppl. 14: 1-51.

WIEDERHOLM, T. (Ed.) 1983 Chironomidae of the Holarctic region — Keys and diagnoses. Part I, Larvae. Ent.

scand. Suppl. 19: 1—457.

—— 1986. ibid. Part 2, Pupae. Ent. scand. Suppl. 28: 1—482.

Prof. Dr. Ernst Josef Fittkau,

Zoologische Staatssammlung,

Münchhausenstraße 21,

D-8000 Munich 60, West Germany.

Dr. Declan A. Murray,

Department of Zoology, University College Dublin,

Dublin 4, Ireland.

259

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a ae ee

Bisher erschienene Supplementbände der SPixıana:

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Die Schwebfliege Episyrphus balteatus. Zwei C'O fliegend; auf der Blüte nach oben zwei Q® (Ph. E. Kotzke).

Zum zwanzigjährigen Bestehen der

Station Randecker Maar

Während es in der Zoologie bis heute die verschiedensten Modeströmungen gab, denen teils jahr-

zehntelang das bevorzugte Augenmerk der Forschung galt, hat die Migrationsforschung zu keiner

Zeit etwas von ihrer Faszination verloren und ist heute so aktuell wie früher.

Im Gegensatz zur Vogelzugforschung fanden aber dıe Insektenwanderungen erst spät das Interesse

der Fachwelt und das erste Fachbuch zu diesem Thema wurde 1930 publiziert. Dabei waren es vielfach

die Vogelzugforscher, die uns frühe Hinweise auf Insektenmigrationen übermittelten. Heinrich

Gätke verdanken wir Berichte aus dem vorigen Jahrhundert über Massenwanderungen von Schmet-

terlingen nach Helgoland. Schon 1872 erkannte er die Zusammenhänge von abnormen Wetterlagen

und dem Auftreten südlicher Vögel und Schmetterlinge auf der Insel: „Lepidoptera (-migrations)...

are subjects to the same meteorogical influences as birds. . .“ Neben Verdichtungen des Zuges entlang

der Küsten haben Kanalisierungseffekte an Gebirgspässen ın allen Teilen der Welt bei der Erfassung

des Vogelzugs wie auch der Insektenmigrationsforschung eine große Bedeutung.

C. W. Mackworth-Praed, der berühmte Afrika-Ornithologe, wies als einer der ersten auf Massen-

wanderungen von Schwebfliegen über den Kanal hin. Das britische Forscherehepaar D. & E. Lack

entdeckte bei Vogelzugbeobachtungen in den Pyrenäen auffälligen Insektenzug und machte die

Wissenschaft auf die dortigen Pässe aufmerksam.

Auch die Bedeutung des Bretolet-Passes in der Westschweiz, langjähriges entomologisches Ar-

beitsgebiet für J. Auberts Migrationsforschungen an Insekten, war bei ornithologischen Untersu-

chungen erkannt worden.

Schon Anfang der sechziger Jahre faszinierten uns die Admirale, die jeden Herbst mit der Zugrich-

tung der Vögel über die Schwäbische Alb nach Südwesten zogen. Als die Station Randecker Maar nach

ersten Versuchen von 1967 bis 1969 im Jahre 1970 einen regelmäßigen Stationsbetrieb aufnahm, lagen

die Arbeitsschwerpunkte „OGELZUG — INSEKTENWANDERUNGEN“ fest. Seitdem sind

zahlreiche Veröffentlichungen zu beiden Themen erschienen. Mit dem vorliegenden Schwebfliegen-

heft findet ein erstes Projekt der Station seinen Abschluß. Monographien über den Zug der Schmetter-

linge und der Vögel werden folgen.

Ursprünglich war die Stationsarbeit auf wenige Jahre konzipiert. Doch die zahlreichen Fragen, die

nach zwei Jahrzehnten blieben, zeigen immer mehr, daß manche Aspekte nur in Langzeitprojekten zu

klären sind. Neben vielen ungelösten Problemen ist die Spannbreite der Erscheinungsformen im

Migrationsbereich selbst nach 20 Jahren offenbar nicht annähernd erfaßt, wie uns die alljährlichen

Überraschungen zeigen.

Dies dürfte einer der Gründe sein, weshalb sich manche der inzwischen über 350 jungen Mitarbeiter

(Biologen, Schüler und Studenten) über Jahre hinweg, von erstaunlichem Enthusiasmus getragen,

ehrenamtlich an der oft harten Stationsarbeit beteiligen. Regen, Schnee und Wind ausgesetzt, wurden

die Beobachtungen in den ersten Jahren bis in den Dezember hinein ausgedehnt. Ein Schäferkarren

diente gleichermaßen als Küche und Büro wie auch als spartanische Schlafstatt und hatte wenig gemein

mit der bescheidenen Gemütlichkeit der heutigen Stationsunterkunft.

Mit den 70000 bisher geleisteten Beobachtungsstunden kann inzwischen ein unersetzliches Mate-

rial von Millionen Daten zur Langzeitdynamik von etwa 100 Vogelarten und zahlreichen Insekten-

arten, und damit Basismaterial für die Naturschutzarbeit geliefert werden.

Die Station hat ihre Tätigkeit während der ersten Jahre ohne jegliche finanzielle Unterstützung be-

trieben.

Meinen Freunden von der Ortsgruppe Kirchheim/Teck und dem Landesverband Baden-Württem-

berg des Deutschen Bundes für Vogelschutz (DBV) ist es zu verdanken, daß schließlich durch jährli-

che Zuschüsse ein finanzieller Grundstock geschaffen wurde. Doch letztendlich wäre der Betrieb der

Station ohne die vielen privaten Spender und Helfer nicht möglich gewesen, denen ein besonderes

Kapitel auf Seite 95 gewidmet ist. Ganz besonders möchte ich mich bei meinen Freunden Prof. Dr.

H. Mattes und K. Sill bedanken, die zeitweise die organısatorischen Geschäfte der Station führten.

Ihnen und allen anderen, die unsere Arbeit unterstützten, herzlichen Dank.

Last not least, der herzlichste Dank an meine Frau Dorothea. Sie hat die Station nicht nur von An-

beginn durch alle Schwierigkeiten mitbegleitet, sondern hat sich neben ihrer Mitarbeit als Entomolo-

gin besonders um die Betreuung der Mitarbeiter bemüht und beim Organisatorischen mitgewirkt.

Lenningen, November 1989 Wulf Gatter

Inhaltsverzeichnis

Einleitung: Überblick über die bisherige Erforschung von

Schwebrliesenwanderungennur 2... 00

Material und Methode

BagerderStationsgRandecken Maar a

Banssder Schhwebtliegen ap 10 2.00

Resistrierumegden Wetterdaten mn. ee en

KlimadatenrdewSeationiRandecker Maar. 2

ANb kürZUn Ser RE EN RN Se nee een

Ergebnisse

SchwebiliegenantensamyRandecken aa

Schwebtliegenimi#zo0ophagensllarvenee

SchwebrlresenfmititertestrischentsaprophagennParven re

Schwebfliegen mit aquatischen sapro-/microphagen Larven ........ 222222220.

SchwuebtlresentmiaphytophagenelEarvene

Kyfeitenea VanderartensmEuropawey

SchnyebHlie senimigzo0phagenalauyene

Schwebiliegenimititerrestrisehen,saprophasen Larven

Schwebtliegenimi@sapro/200pha genWllarvene

Übersicht über die Wanderschwebfliegen Europas (Tabelle) ..................

Wanderaktivität in Abhängigkeit von den Windverhältnissen .................

Episynphasibalteatus| 980 un. rot ee ee

Blasycheimusichypeatuss) Son: ee

Bpssynphasibalteatusi 981 dar. A seem ae enaeerenee

Blasycheinuste)Deatas IS Mas re N

Das tageszeitliche Windangebot und seine Auswirkungen auf die Wanderaktivität

von Episyrphus balteatus und Platycheirus chypeatus ......... 2.222222 ceceeen.

Zusammenfassung: Windabhängige Wanderstrategien Saisonaler Migranten und

SarsomalerDismmistantene ee

SaisonaleMisranten gs EN N RR NE ae

SaisonaleaDismmipramtene „3. ee ee edel le ee ne

Bopulationsdymanmike Sr ar a

DISKUSSION FR IT REN N N ni see

Vergleich der Phänologie der Schwebfliegenwanderungen am Randecker Maar und

amk@olideBretolerır Se re N a te

Veränderungen der Zahlenverhältnisse der Geschlechter im Verlauf der Migration .

BrolosieidersN/andersehwyebtlie sen

Arten mit phytophagen und terrestrischen saprophagen Larven ...............

rtenkmitzoophagentlanven a

Artenimigaquatıschenlsapto/mierophasenllasvens re

Zur Bedeutung der Wanderschwebfliegen bei der biologischen Schädlings-

bDekamplunse en ee een ee ne nem

EN SgemenmeritspekterdenSchwebtliesenwanderungenn aa

Verbreitung Nee ee Ne

DiapausesundıNisratonee ne ee

Generationenfolge und Migration ..........20.--...2nosenunesenennenee

Die Stellung der Schwebfliegen innerhalb der Wanderinsekten ................

Zusammenfassung 13. Ve NE EN ES Bene CHR

Dank'W.

EI aan fallalafıer (ade Te) lo: Snlhon.e-Menkelieila' nat m etlenhniin Mia laf,eruuiie?; era ie. (ailluifeljarieiäin Aut in Lanin zZ npiwstiniiuit nateZtozrw zn une

a 0 OO ID EC SKOBDWOID: DE LEO IL AO RO D: DABEI II ELTELMOLOBDBONT HOSEN OD I

Die Wanderungen der Schwebfliegen (Diptera, Syrphidae)

am Randecker Maar

Von Wulf Gatter und Ulrich Schmid

Abstract

Hoverfly migration (Diptera, Syrphidae) at Randecker Maar, SW-Germany

During the years 1975-1987 (except 1983) hoverfly migration was investigated at the migration research station

“Randecker Maar — Schwäbische Alb” (SW-Germany; 48.35 N, 9.31 E, 772 m). From mid July or the end of July

to the beginning of October of these 12 years, 90049 syrphids migrating southward were collected with a large net

trap which opened to the north (S-trap). During 5 years, 9815 hoverflies migrating northward were caught with a

similar trap which opened to the south (N-trap) (see Tab. 1).

Aspects of the phenology of species with more than 50 individuals collected (see Tab. 2) are described in the pre-

sent paper (3.1, 3.2), e. g. the seasonal and diurnal activity patterns of the species and sexes, differences between the

two traps, sex-ratios, captures in relation to wind directions. Earlier observations of migrations of the species are

summarized, and each species is arranged (if possible) in one of the following categories:

I) Anemomigration (drift): passive transport of flight-inclined insects by wind without direction orientation.

II) Dismigration: endogenously (dispersal) and exogenously (spacing) released active dismigrations without or

with only indirect direction orientation (selection of favourable winds).

III) Migration: migration of individuals or species programmed to endogenous migration and direction orienta-

tion.

Both dismigration and migration can be expansive (without returning) or seasonal (migration to and return from

summer, winter or dıapause habıtats).

Tab. 3 shows all known migratory species of Europe.

The phenology of seasonal migrants (species which migrate orientated into winter and summer areas; see Tab. 3)

is characterized as follows:

1. The difference between numbers in traps and supplementary field observations show that the flies move in late

summer and autumn in southerly and southwesterly directions.

2. Field observations indicate that seasonal migrants move northward in spring.

3. There are characteristic differences in the phenology of seasonal migrants between the Randecker Maar and the

Col de Bretolet (Swiss Alps, 1923 m) which is situated 350 km to the southwest: in most cases maximum south-

bound migration peaks are later at Col de Bretolet. Migratory birds show the same shift ofthe maxima. During their

S and SW orientated migration, both birds and hoverflies cross Randecker Maar some days before they reach the

more southwesterly situated Col de Bretolet (4.1).

4. At Randecker Maar and more northerly sites, males and females of most seasonal migrants appear in nearly

equal numbers. At Col de Bretolet, females predominate clearly (see Tab. 5). — Females of Episyrphus balteatus

have a much longer lifespan than males. At the beginning of the migration period, males participate ın great num-

bers; during migration an increasing proportion of them dies. At the end of the migration period, only females are

left. The later peak at Col de Bretolet causes predominance of females at thıs site (4.2).

5. Only if there is a headwind do seasonal migrants fly close to the ground in order to diminish the unfavourable

influence of this wind. They make use of the helping influence of a tailwind by flying at a greater height. Therefore

southbound migrants are caught in the trap in much greater numbers when the wind blows from the south (3.4).

6. Seasonal and diurnal migration patterns are very similar from year to year apart from the high fluctuation of

numbers.

The group of dismigrating species (see Tab. 3), which have an active dispersal migration, can be less clearly char-

acterized. The following features of the phenology of these dismigrants are not shared by all species — a conse-

quence of the differing functions, extents and ranges involved:

1. The difference between numbers of flies collected in the two traps is often low; no preferred direction of flight

is distinguishable. — In some species the difference is higher; presumably they choose favourable wind directions

for migration.

2. In most cases there is no difference between the seasonal activity patterns of Randecker Maar and Col de

Bretolet (4.1).

3. The differences of sex ratios at Randecker Maar and Col de Bretolet which are shown by seasonal migrants are

not present in seasonal dismigrants (see above; Tab. 5; 4.2).

4. In all species, females predominate at Randecker Maar. Flight activity of females close to the ground level is

probably greater than that of males due to the females’ search for favourable egg laying habitats.

5. Seasonal dismigrants do not react to different wind directions in the same manner as seasonal migrants (see

above). Migrations with tailwind take place (at least in part) close to the ground (3.4).

For some ecological groups of the family Syrphidae, seasonal migrations are an integral part of their life cycles

(4.3, 4.4):

1. Seasonal migration is known only from species with zoophagous (aphidophagous) larvae or from species with

aquatic sapro-/microphagous larvae.

2. In particular species with hibernating imagines or puparia are seasonal migrants (see Tab. 3). These species can-

not survive the winter in northern parts of their summer range, or they survive only in small numbers under optimal

conditions. So they immigrate into these areas in spring. In summer and autumn they move back to the south. —

Some species with hibernating larvae migrate in the same manner. Presumably migration to southern Europe can

also reduce winter mortality of their larvae.

3. The adaptive advantage of migration to the south in summer and autumn is an increase of their chance of sur-

viving the winter. The advantages of spring migration to the north for aphidophagous species are the much better

conditions for larval development (maximum of aphids) and imaginal feeding (richness in “fly-flowers”) in central

and northern Europe; for aquatic species the presence of larval habitats in much higher densities and with much les-

ser risk of drying up than in the summer-arid southern Europe.

The most important function of dismigration is to find habitats with favourable conditions for larval develop-

ment. Also in such migrations species with zoophagous larvae are the most common participants (4.3); they depend

on food resources which occur unpredictably in time and space (species see Tab. 3).

Aphidophagous hoverflies show different life cycles, which are partly correlated with spring and/or autumn ma-

ximum of aphid density (mono- and bivoltine species). Polyvoltine species have to integrate migration periods in

their life cycles to enhance the chances for successful reproduction during aphid minimum in summer. They are

much less specialized in habitat selection and larval diet than mono- and bivoltine species. In polyvoltine zoopha-

gous syrphids two groups compete for larval food: seasonal migrants and seasonal dismigrants (see Tab. 3). The for-

mer are better direct competitors for food, the latter are able to use a higher variety of food (4.3).

Species with terrestrial saprophagous or phytophagous larvae are not forced to migrate to find their often predic-

tably occuring larval food resources. Some saprophagous hoverflies dismigrate over short distances (see Tab. 3). No

migratory species is known among the usually highly specialized phytophagous species (4.3).

The migration behaviour of Syrphidae shows much similarity to that of other well-known migratory insects, e.g.

the butterflies (Lepidoptera) (4.5). There are strong connections between diapause and migration (species with dia-

pausing imagines migrate; 4.5.2), number of generations per year and migration (in particular polyvoltine species

migrate; 4.5.3) and between the dimension of the distribution area and migration (migrants have vast areas; 4.5.1).

Some of our common hoverfly species are the most impressive migrants amongst European insects in respect of

dimensions and regularity of their migrations.

Furthermore most of the hoverfly species which are important as aphid predators in agriculture and forestry are

migrants. If some of these species offer the possibility to act as biological pest control a precise knowledge of the

life cycle of each species is necessary. The influence of migratory behaviour has been greatly underestimated up to

now in the research on biological pest control.

1. Einleitung: Überblick über die bisherige Erforschung von Schwebfliegenwanderungen

Wandernde Schwebfliegen haben gelegentlich schon früher Aufmerksamkeit erregt, wenn sie in ge-

waltigen Massen auftraten (z. B. Eimer 1880, Prell 1925, Walker 1864) oder extrem weit außerhalb

ihrer Verbreitungsgebiete gefunden wurden (z. B. Syrphus ribesü auf Spitzbergen; Elton 1925).

Auf die Möglichkeit regelmäßiger Schwebfliegenwanderungen wiesen aber erstD. & E. Lack (1951)

nach Beobachtungen von gegen den Wind nach WSW fliegenden Episyrphus balteatus an einem Pyre-

näenpaß hin. Williams et al. (1958) bestätigten und vertieften diese ersten Kenntnisse über regelmäßige

Syrphidenzüge.

Nachdem diese Beobachtungen auf völlig neue Aspekte der Ökologie von Dipteren hinwiesen,

brachten die folgenden Jahre so wenig neue Erkenntnisse, daß zwei in den 1960er Jahren erschienene

Standardwerke über Insektenwanderungen (Williams 1961, Johnson 1969) kaum mehr als eine Seite

benötigten, um das Wissen über Syrphidenmigration zusammenzufassen.

Erst die Forschungen von Aubert und seiner Arbeitsgruppe brachten wesentliche Fortschritte (Au-

bert 1962, 1964a, 1964b, 1964c, 1969; Aubert, Aubert & Goeldlin 1976; Aubert & Goeldlin 1981; Au-

bert, Goeldlin & Lyon 1969; Aubert & Jaccard 1981; Dethier & Goeldlin 1981, Goeldlin 1975). In den

Jahren 1962-1973 betrieb die Arbeitsgruppe jeweils ab Ende Juni oder Anfang Juli bis zu den ersten

stärkeren herbstlichen Schneefällen Insektenreusen am Col de Bretolet, einem in 1923 m Höhe gelege-

nen Paß in den wallisischen Alpen. Der Col de Bretolet liegt am Schluß des vom Rhönetal in südwest-

licher Richtung heraufziehenden Val d’Illiez. Die Ergebnisse dieser zwölfjährigen Beobachtungen

sind in Form von Tagessummendiagrammen und einer sehr knapp kommentierten Artenliste bei Au-

bert et al. (1976) dokumentiert.

Der paßähnliche Einschnitt im Norden des Randecker Maars.

Die Reuse am Randecker Maar.

Neben diesen Routinefängen wurden an verschiedenen Alpenpässen Massenfangaktionen mit sehr

großen Netzreusen durchgeführt. Durch Wiederfänge von dabei markierten Schwebfliegen sind

Wanderungen einzelner Individuen über maximal 111 km belegt! Über die von NE nach SW hinter-

einander liegenden Pässe Krinnenpaß (1659 m), Col de la Croix (1778 m), Col de Cou und Col de

Bretolet (1921 und 1923 m), Col de la Goleze (1671 m) und Col du Glandon (1961 m) konnten

Schwebfliegenwanderungen über ca. 160 km verfolgt werden. Im Verlauf dieser Wanderung wird

zwischen dem Col de la Croix und dem Col de Bretolet auch das Rhönetal (430 m) gequert.

Nach der Markierung am Col de Bretolet hatten die ersten Fliegen den 3,6 km entfernten Paß La

Goleze schon in 10-15 Minuten erreicht. Die unter sehr ungünstigen Witterungsbedingungen am

Col du Glandon in 111 km Entfernung wiedergefangenen Tiere waren vermutlich drei Tage vorher

markiert worden (Aubert et al. 1969).

An verschiedenen Stellen der Schweiz und Österreichs ergaben kurzfristigere Versuche (z. B. Au-

bert & Jaccard 1981, Dethier & Goeldlin 1981, Gepp 1975) ähnliche Beobachtungen, so daß die Phä-

nologie der herbstlichen Schwebfliegenwanderungen im Alpenraum als gut bekannt betrachtet wer-

den kann.

Aus anderen Gebieten fehlen längerfristige Beobachtungen allerdings fast völlig.

Nachdem an der Station Randecker Maar seit Anfang der 1960er Jahre immer wieder auffälliger

Schwebfliegenzug beobachtet worden war (z. B. Gatter & Gatter 1973, Gatter 1975a), begann man

dort daraufhin mit der systematischen Untersuchung der Wanderungen mit Hilfe von Insektenreu-

sen. Erste Ergebnisse aus dem Jahr 1975 liegen bereits publiziert vor (Gatter 1976); sie lassen hinsicht-

lich der Artenzusammensetzung große Übereinstimmungen zwischen den Beobachtungen in den Al-

pen und denen im südwestdeutschen Mittelgebirgsraum erkennen. In einer Analyse dieses Artenspek-

trums wurden charakteristische Merkmale einer von Wanderarten geprägten Schwebfliegenfauna

deutlich (Schmid & Gatter 1988).

112

Erste systematische Beobachtungen zum Auftreten von Schwebfliegen auf einer Nordseeinsel ohne

bodenständige Syrphidenfauna (Schmid 1987) deuten darauf hin, daß auch im norddeutschen Tiefland

regelmäßige spätsommerliche Syrphidenwanderungen stattfinden, an denen dieselben Arten wie im

Hoch- und Mittelgebirgsraum teilhaben.

Die am Randecker Maar gewonnenen und hier dargestellten Daten zum Wanderverhalten der

Schwebtliegen dürften also grundsätzliche Gültigkeit wenigstens für den mitteleuropäischen Raum

haben.

2. Material und Methode

2.1 Lage der Station Randecker Maar

Die Forschungsstation liegt im Südwesten der Bundesrepublik Deutschland (Abb. 1) am Nordrand der Schwäbi-

schen Alb, einem sich vonSW nach NE erstreckenden Mittelgebirge, das steil über seinem nördlichen Vorland auf-

ragt. Im Bereich der Station liegt das Vorland in ungefähr 400 m NN, der Albtrauf in ca. 800 m NN (Abb. 2a, 2b).

Der nahezu runde Kessel des Randecker Maars ist nach Norden geöffnet und unterbricht den Albtrauf mit einer

tiefen Kerbe. Diesen Einschnitt nutzen in breiter Front aus Nordost heranziehende Vögel und Insekten, um hier

mit geringerem Energieaufwand die Höhe der Albhochfläche zu gewinnen. Dadurch kommt es am Randecker Maar

zu starken horizontalen und vertikalen Verdichtungen des Zuggeschehens (Gatter 1978, 1981a).

Die Beobachtungsstation selbst (48.35 N, 9.31 E) befindet sich südlich des Maars am Rand einer kleinen Kuppe

in 772 müber NN (Abb. 2a). Sie bietet freie Sicht nach allen Himmelsrichtungen außer nach SE, wo die Kuppe von

Gebüschgruppen mit einigen niedrigeren Bäumen bewachsen ist.

Abb. 1: Die Lage der Station „Randecker Maar“ in Südwest-Deutschland.

Fig. 1: Position of the Migration Research Station “Randecker Maar” in SW-Germany.

2.2 Fang der Schwebfliegen

Die wandernden Schwebfliegen werden mit Hilfe einer trichterförmigen, nach NNE geöffneten Insektenreuse

gefangen. Die Reusenöffnung hatte in den Jahren 1975-1977 eine effektive Größe von 5 m? (2,5 m Höhe =2 m

Höhe über der Wiesenvegetation; 2,5 m Breite), in den Jahren 1978-1982 und 1984— 1987 eine effektive Größe von

10 m? (5 m Breite). Die Wände und das zum Fangbeutel hin leicht ansteigende Dach bestehen aus Fliegengitter. Den

nach SSW gerichteten inneren Abschluß bildet ein 20 cm breites, mit Plastikfolie bezogenes Fenster. Im oberen

Winkel der Reusenkonstruktion befindet sich eine 10X20 cm große Öffnung. Über diese Öffnung wird ein durch-

sichtiger Plastiksack gestülpt. Die aus NNE ankommenden Insekten fliegen in der Reuse, von den schräg stehenden

Wänden geleitet, bis zum Fenster, steigen an ihm empor und gelangen so in den Plastikbeutel. Dieser wurde in den

Jahren 1975-1977 von Sonnenaufgang bis Sonnenuntergang, ab 1978 von Sonnenaufgang bis 16 Uhr MEZ stünd-

lich gewechselt.

Am Arbeitsplatz in der wenige Meter entfernten Station werden die Insekten betäubt und bestimmt, wobei eine

Vergleichsammlung zur Verfügung steht. Nicht immer kann so eine sichere Artdiagnose gestellt werden, so daß

Zweifelsfälle zur späteren Nachbestimmung gesammelt werden. Das Gros der Fänglinge überlebt die Betäubung

und wird wieder in die Freiheit entlassen.

In den Jahren 1977 und 1979— 1982 wurde als Kontrolle zusätzlich eine baugleiche Reuse mit einer nach $ gerich-

teten Öffnung betrieben. In dieser Reuse gefangene Schwebfliegen wurden im Jahr 1979 nur pauschal erfaßt, in den

anderen Jahren artlich getrennt ausgezählt.

Tab. 1 und Abb. 27a geben einen Überblick über Reusengröße, Zeitraum der Erfassung und Anzahl gefangener

und bestimmter Schwebfliegen im Untersuchungszeitraum.

Tabelle 1: Fangzeitraum, Reusengröße und Zahl der gefangenen Schwebfliegen am Randecker Maar 1975-1987.

Tab. 1: Period of investigation, size of trap opening, and number of trapped hoverflies at Randecker Maar

1973— 1987.

Jahr Zeitraum Reusengröße Südflieger Nordflieger

(S-Reuse) (N-Reuse)

1975 2642-3009. 5 m? 4568

1976 6. 30488} 5 m? 1214

1977 DIN 9% 5m? 1519 299

1978 1602 12210: 10 m? 6462

1979 16 2731: 10, 10 m? 8910 1417

1980 TEICHE 10 m? 8869 2091

1981 227 12510! 10 m? 11511 3708

1982 1307 3.10, 10 m? 5323 2300

1983

1984 182 10. 10) 10 m? 7190

1985 29. 764: 30. 10 m? 13215

1986 13: il: 10 m? 11971

1987 19,723 9° 10 m? 9297

12 Jahre 90049 9815

2.3 Registrierung der Wetterdaten

Jede Stunde werden folgende Wetterdaten erfaßt:

— Windrichtung (N, NE, E, SE, S, SW, W, NW) und Windstärke (in Bft.); Messung mit Hilfe eines auf der Höhe

der Kuppe stehenden und über die Baumschicht hinausragenden Windmeßgerätes;

— Bedeckungsgrad des Himmels in Achteln;

— Temperatur;

— Sicht;

— Besondere Vorkommnisse wie Niederschlag, Gewitter, Nebel.

2.4 Klimadaten der Station Randecker Maar

An der in geringer Entfernung und auf ungefähr gleicher Höhe (764 m) liegenden Wetterstation Schopfloch be-

trägt die mittlere Jahrestemperatur 6,9°C, der mittlere Jahresniederschlag 1065 mm. Während der Beobachtungs-

periode liegen die Mittelwerte bei 15,7°C/120 mm im Juli, 15,2°C/108 mm im August, 12,1°C/93 mm im Septem-

ber, 7,2°C/72 mm im Oktober und 2,6°C/70 mm im November.

Im Albvorland (Kirchheim, 315 m) beträgt die mittlere Jahrestemperatur 9,0°C bei einem Niederschlag von

752 mm. Der Nordrand der Schopflocher Berghalbinsel, auf der die Station liegt, gehört zu den nebelärmsten

Punkten des mittleren Württemberg. Die Nebel des Albvorlandes steigen am nördlichen Steilabfall selten über

600 m NN hoch.

2.5 Abkürzungen

RM: Randecker Maar

CB: Col de Bretolet

S-Reuse: nach Norden geöffnete Reuse, die nach Süden fliegende Insekten fängt

N-Reuse: nach Süden geöffnete Reuse, die nach Norden fliegende Insekten fängt

©": Männchen

9: Weibchen

3. Ergebnisse

3.1 Schwebfliegenarten am Randecker Maar

Eine vollständige Artenliste der am Randecker Maar beobachteten Schwebfliegen liegt bereits vor

(Schmid & Gatter 1988). Hier werden deshalb nur die Arten berücksichtigt, von denen genug Material

(>50 Ex.) vorhanden ist, um gut begründbare Aussagen über ihre Phänologie zu treffen (Tab. 2).

Nach der Ernährungsweise der Larven lassen sich bei Schwebfliegen verschiedene Gruppen unter-

scheiden, die hier nacheinander abgehandelt werden:

— Schwebfliegen mit zoophagen (meist blattlausfressenden) Larven;

Abb. 2a: Das nach Norden geöffnete Randecker Maar in der Aufsicht. Aus Norden und Nordosten trifft der Breit-

frontzug auf den 400 m hoch über dem Vorland aufragenden Steilabfall der Schwäbischen Alb. Vögel und Insekten

ziehen bevorzugt durch den Einschnitt des Maars. Der Punkt zeigt die Lage der Station (aus Gatter 1981a).

Fig. 2a: “Randecker Maar” in topview. Migrating birds and insects use the cleft in the 400m steep slope of the

Schwäbische Alb. The spot shows the site of the station.

- 800m

— 400m

Abb. 2b: Am Steilabfall der Schwäbischen Alb (im Schnitt dargestellt) kommt es zu einer vertikalen Verdichtung

des Zuggeschehens. Der Punkt zeigt die Lage der Station (aus Gatter 1981 a).

Fig. 2b: Vertical concentration of migrants passing the Randecker Maar. The spot indicates the station site.

Tabelle 2: Zwölfjährige Fangsummen einzelner Arten in der S-Reuse des Randecker Maars. Zum Vergleich sind die

am Col de Bretolet in einem ebenfalls zwölfjährigen Untersuchungszeitraum (1962-1973) gewonnenen Fang-

summen aufgeführt (nach Aubert et al. 1976).

Tab. 2: Total amount of hoverflies trapped in the S-trap (trap with opening to the north) at Randecker Maar during

12 years compared with the data of Col de Bretolet (1962-1973).

vo oNONvpruvuND

. Episyrphus balteatus

. Melanostoma mellinum

. Platycheirus clypeatus

. Sphaerophorıa scripta

. Metasyrphus corollae

. Syrphus spec.

. Platycheirus albimanus

. Scaeva pyrastri

. Erıstalis tenax

. Platycheirus manicatus

. Helophilus pendulus

. Rhingia campestris

. Cheilosia pagana

. Sphaerophoria menthastri Gruppe

. Platycheirus peltatus

. Parasyrphus lineolus

. Meliscaeva cinctella

. Helophilus trivittatus

. Scaeva selenitica

. Neoascia podagrica

. Erıstalis arbustorum

. Syritta pipiens

. Erıstalis interrupta

. Erıstalis pertinax

. Melanostoma scalare

Sonstige

Randecker Maar

29546

15937

13450

11856

8426

1138

885

757

)

724

700

448

233

272

270

247

180

120

389

90049

32,81%

15,48%

14,94%

13,17%

9,36%

5,72%

1,26%

0,98%

0,84%

0,82%

0,80%

0,78%

0,50%

0,33%

0,30%

0,27%

0,20%

0,19%

0,13%

0,10%

0,09%

0,08%

0,06%

0,43%

100,00%

Col de Bretolet

1264568

31.373

146210

160685

67121

3658

11168

635 294

1476

4189

356

127

180

45668

2396539

52,77%

1,32%

0,00%

6,10%

6,71%

2,80%

0,15%

0,47%

26,51%

0,06%

0,17%

0,01%

0,00%

0,01%

0,00%

0,24%

0,04%

0,08%

0,45%

0,00%

0,19%

0,00%

0,01%

0,00%

1,91%

100,00%

ET —g

— Schwebfliegen mit terrestrisch lebenden, saprophagen Larven;

— Schwebfliegen mit aquatisch lebenden, sapro-/microphagen Larven;

— Schwebfliegen mit phytophagen Larven.

Innerhalb dieser Gruppen sind die Arten systematisch angeordnet.

Die Artabschnitte sind folgendermaßen gegliedert:

— Verbreitung (nach Angaben von Knutson et al. 1975, Sack 1932, Thompson et al. 1976, Violovitsh

1983, Wirth et al. 1965);

— Angaben zur Biologie: Flugzeit, Generationenfolge, Überwinterung, Häufigkeit;

— Hinweise auf Wanderverhalten;

— Phänologie am Randecker Maar;

— Status: In diesem Abschnitt wird versucht, die Art einer der bekannten Migrationsformen zuzu-

ordnen, wobei wir die als Arbeitshypothese vorgeschlagene Gliederung von Gatter (1981a: 19— 27,

1981c) zugrundelegen (vgl. Abb. 3).

Wander insekten

Dispersionsprozesse

Migrationsformen

Anemomigration

Verdriftung

Migration

(Richtungsorientiert)

(Navigierend)

Saisonale Bagzabten

Migration

Saisonale

Migration

Dismigration

Zerstreuungswanderung

Expansive

Dismigration

mit geringer

Rückwanderung)

Saisonale

Migration

mit ausgeprägter

Rückwanderung

monovoltine Arten monovoltine Arten

ohne Diapause \ ohne Diapause

B monovoltine Arten mit Diapause

polyvoltine Arten mit Diapause

polyvoltine Arten ohne Diapause

Saisonale Horizontalwanderungen

Saisonale Vertikalwanderungen

C I et!

Aufsuchen von Diapausequartieren

Habitatwechsel

Abb. 3: Die Migrationsformen der Insekten. Die links mit Buchstaben bezeichneten Bereiche beziehen sich auf ob-

ligatorische Wanderer und bedeuten A) WIE, B) WER und C) WESHALB gewandert wird (aus Gatter 1981 a).

Fig. 3: Insect migration categories. Letters on the left mean A)HOW, B) WHO and C) WHY migration takes place.

177

Die Begriffsbestimmungen sollten dabei nicht als starre, restriktive Einteilung verstanden werden

(Baker 1978). Als Folge der großen biologischen Mannigfaltigkeit der Insektenwanderungen bestehen

zahlreiche Übergänge zwischen einzelnen Migrationsformen (vgl. 4.5.4):

1: Anemomigration (Verdriftung): Passive Verfrachtung flugbereiter Insekten durch den Wind

ohne Richtungsorientierung.

1. Dismigration (Zerstreuungswanderung): Endogen (dispersal) oder exogen (spacing) ausgelö-

ste aktive Zerstreuungswanderungen ohne bzw. mit nur geringer Richtungsorientierung

durch Selektion günstiger Winde.

Innerhalb der Dismigrationen können zwei verschiedene Formen unterschieden werden:

II.1 _Expansive Dismigration (Expansive Zerstreuungswanderung): Aktive Zerstreuungswande-

rung nicht oder nur indirekt richtungsorientiert wandernder Arten, die sich bei der Migration

vom Entwicklungsort entfernen. Sie und ihre Nachkommen kehren nicht zurück.

11.2 _Saisonale Dismigration (Saisonale Zerstreuungswanderung): Aktive saisonale Wanderung

nicht oder nur indirekt richtungsorientiert wandernder Arten in Sommer- bzw. Winterareale

oder Diapausequartiere und zurück. Durch saisonal unterschiedliche Winde (ohne Richtungs-

orientierung) oder Selektion günstiger Winde (indirekte Richtungsorientierung) können sai-

sonal unterschiedliche Hauptstoßrichtungen entstehen.

III. _ Migration (richtungsorientiert) (Richtungsorientierte Wanderung): Wanderungen von Indivi-

duen oder Populationen von Arten mit endogen programmierter Migrationsrichtung (kom-

paßorientiert).

III.1 Expansive Migration (Expansive richtungsorientierte Wanderung): Richtungsorientiert wan-

dernde Arten, die sich auch ohne Einfluß exogener Faktoren vom Entwicklungsort weg in eine

Vorzugsrichtung entfernen. Sie und ihre Nachkommen kehren nicht zurück.

III.2 Saisonale Migration (Richtungsorientierte Saisonwanderung): Richtungsorientierte Wande-

rungen in Sommer- oder Winterareale bzw. Diapausequartiere (Sommer- und Winterruhe)

und zurück, die von Individuen oder Populationen einer Art alljährlich ausgeführt werden.

Unter den Schwebfliegen fehlen Beispiele für reine Anemomigration und für Expansive Migration.

Ein Beispiel für eine Expansive Dismigration ist Volucella zonaria (vgl. 3.2.3).

Zahlreiche Syrphidenarten führen Saisonale Dismigrationen oder Saisonale Migrationen durch. Saisonale Zer-

streuungswanderungen stehen oft in sehr engem Zusammenhang mit dem Entwicklungszyklus einer Art. Sie die-

nen dabei neben dem Aufsuchen von Winter- oder Sommerarealen und von Diapausequartieren auch dem Auf-

suchen von Orten, die der Nachfolgegeneration günstige Entwicklungsbedingungen bieten. Treten z. B. Nah-

rungsressourcen, von denen die Larven einer Art abhängig sind, nicht gleichmäßig in Raum und Zeit auf (z. B.

Blattläuse), können sie durch saisonal dismigrierende Arten schneller und effektiver erschlossen werden.

Dismigrationen können von ganzen oder nur von Teilpopulationen durchgeführt werden; bei Migrationen sind

in vielen Fällen eher ganze Populationen beteiligt.

Saisonal unterschiedliche Winde oder Selektion günstiger Winde können saisonal unterschiedliche Hauptstoß-

richtungen entstehen lassen (bei Dismigrationen) oder solche unterstützen (bei Migrationen).

3.1.1 Schwebfliegen mit zoophagen Larven

Platycheirus albimanus (Fabricius 1781) (= cyaneus Müller 1764)

Verbreitung: holarktisch, orientalisch (Nepal?, Indien?, Philippinen)

Biologie: Platycheirus albimanus ist eine weit verbreitete und häufige Schwebfliegenart, die in vie-

len Lebensräumen angetroffen werden kann, wenn sie auch feuchtere Standorte mit dichtem Pflan-

zenwuchs bevorzugt. Sie ist in Mitteleuropa von (Ende März) Mitte April bis Mitte Oktober (Anfang

November) anzutreffen und bildet in Süddeutschland (Schmid 1986) wie in Großbritannien (Stubbs

& Falk 1983) drei Generationen. Die Larven sind meist in der Streuschicht und nur relativ selten an

Blattlauskolonien anzutreffen. Vermutlich leben sie dort zoophag mit weitem Beutespektrum (Rothe-

ray & Gilbert 1989). P. albimanus überwintert als diapausierende Larve.

Anzahl pro Pent. j

Anz. pro Tag (#+) 832 ZmMm3SmUus

1975 - 1987

200 |

Aoa- Ir &: 0: 6. 0 - 94 |

5% 25. 7 J

25% 2.8.

50% 13. 8

H 75% 24.8

95% 18. 9

Lerzabr10r

Anzahl pro Pent x

Anzahl pro Pent De 79 Anz. pro Tag (+) g3lbr1ı1m3amı ja

3 7 AOeENAN EIS V

inz. pro Tag (+) | 1975 - 1987

50 1975 - 1987 100 v Y v

25+[ Vu NV a Sa |r ar: 16.7 0 = 51 -

Er. 19. 6 (E 18 | 5425. 7 12 J 615

5 12 J= 218 Er mm

Ber a |

ul. 6.8.75 2 16. 7.76 3

40 80 z

50.1 50% 8.8. 25.7.76 27 on Zar Al nl 15. 8.77 1]

JS 6RLOR 28. 7.78 1 | 95% 19. 9 26. 7.78 7

2 9.0. AN. Tee) 1 | Le. 6.10. | 4.10.79 1

Le. 16.9. 1. 8.80 17 | 7

Bor |! 6.8.81 Hi 7

15 + 29. 7.82 4 7

14. 8.87 22 8

49 + 1

10 a

1 al a Ulm h

b Jun Jul Aug Sep Okt

Abb. 4a: Nach Süden gerichtete Migration von Platycheirus albimanus in den 12 Jahren 1975-1982 und

| 1984— 1987 in Tages- und Pentadensummen (1.—5. Jan., 6.— 10. Jan. etc.). Auf der Ordinate werden Tagessummen

(untere Zahl, mit + markiert) und Pentadensummen (obere Zahl) angegeben. An welchem Tag 5, 25, 50, 75 und

| 95% der Gesamtsumme durchgezogen sind, kann links oben abgelesen werden. Die dreieckigen Pfeile über dem

| Diagramm kennzeichnen die Lage der 25 %-,50 %- und 75 %-Werte. Rechts oben kann die Zahl aller im 12jährigen

\ Untersuchungszeitraum gefangener Tiere (n) und der Jahresdurchschnitt (©) abgelesen werden. Darunter ist für je-

des Jahr die maximale Tagessumme aufgeführt.

Fig. 4a: Migration of Platycheirus albimanus in southerly direction over 12 years (1975-1982; 1984-1987) with

the sum of days and the sum of 5-day periods (1.-5. Jan., 6.— 10. Jan. ....). Sum of days (lower number, marked

| with +) and sum of 5-day periods (upper number) are shown on the ordinate. On the left upper side is shown on

| which day 5, 25, 50, 75 or 95% of total have passed. The triangles show the position of 25%, 50% and 75 % values.

| Above on the right the total number of trapped P. albimanus (n), the annual average (2) and of each year the day

with most specimens are shown.

| Abb. 4b: Nach Süden gerichtete Migration von Platycheirus albimanus-O'C..

ı Fig. 4b: Southbound migration of O'O° of Platycheirus albimanns.

Abb. 4c: Nach Süden gerichtete Migration von Platycheirus albimanus-Q2.

Fig. 4c: Southbound migration of PP of Platycheirus albimanus.

3JDIM3NMUS

vahre 1975 — 1987

albIimamus W albimsanus M

vahre 1975 —- 1987 vaehre 1975 - 1987

25%= 164

* 50%= 328

* 75%= 492

Max= 134

*N = 656

N = 596

r T 1 —

6 8 10 12 14 16 18 20 Unr 4

Abb. 4 d: Tageszeitliche Verteilung von Platycheirus albimanus nach Fängen aus der S-Reuse in den Jahren

1975-1982 und 1984— 1987.

Die Stunde, die den 50 %-Wert enthält, ist kreuzschraffiert, die Stunden, die die 25 %- und 75 %-Werte enthalten,

sind eng schraffiert. Die Strichmarkierungen über diesen Säulen bezeichnen die aus Stundenwerten errechnete Lage

des 25 %-, 50 %- und 75 %-Wertes genauer. Die zugrundeliegenden, kumulierten Individuenzahlen sind rechts an-

gegeben (* 25% etc.).

Nur in den Jahren 1975-1977 wurde von Sonnenaufgang bis Sonnenuntergang beobachtet (vgl. Abb. 8b). In den

anderen Jahren endete die tägliche Beobachtungszeit um 16 Uhr. Die Zugbewegungen am Spätnachmittag wurden

aus den dreijährigen Beobachtungen nach 16 Uhr hochgerechnert. Sie sind als weiße Säulen dargestellt.

N gibt die Zahl der tatsächlich gefangenen Schwebfliegen an, *N (ebenso wie alle anderen mit * versehenen Zahlen)

die durch die Hochrechnung der spätnachmittäglichen Zugbewegungen modifizierte Zahl.

Zur Veranschaulichung der Zahlenverhältnisse dient der „Max“-Wert, der für die längste Säule die Zahl der in dieser

Stunde gefangenen Individuen nennt.

Fig. 4d: Diurnal activity pattern of southbound migrating Platycheirus albimanus (1975—1982, 1984— 1987). 50 %

of the total numbers have passed between the early morning and the hour marked by cross hatching. The hour at

which 25% and 75% have passed respectively, are closely hatched. The small lines on top of the vertical bars indi-

cate exactly the position of 25%, 50% and 75 %.

Only in 1975-1977 did observations last from sunrise till sunset. In other years they ended at 4 p.m. The catches

later in the afternoon (white bars) are calculated from the observations in 1975— 1977. N is the real number of caught

albimanus, *N (and other numbers with *) is a fictitious number arrived at by extrapolization of the afternoon cat-

ches from 1975-1977 over the whole period of investigation. “Max” means the number of specimens indicated by

the longest bar.

Wanderungen: Obwohl die Art am Col de Bretolet in nicht geringer Zahl gefangen wurde, lassen

Aubert et al. (1976) offen, ob es sich dabei um eine Wanderart handelt („consideree comme migra-

trice“). P. albimanus erscheint am CB während der ganzen Beobachtungsperiode regelmäßig, ohne

daf® Maxima erkennbar wären. Ein weiterer Hinweis auf Wanderungen in den Alpen findet sich nur

bei Prell (1925), der während einer Insektenmigration einige Exemplare sammelte. Aus Norddeutsch-

land liegen Nachweise nicht bodenständiger Individuen von der Insel Scharhörn (Schmid 1987) und

von Feuerschiffen vor (Heydemann 1967). Unter Belegexemplaren wandernder Schwebfliegen aus

Südengland war auch ein albimanus-Q (Parmenter 1960).

Phänologie am Randecker Maar: Hier erscheint die Art in den meisten Jahren in — gemessen an

ihrer Häufigkeit — sehr bescheidenem Ausmaß in den Reusen (Tab. 2, Abb. 4a). Auch wenn man da-

von ausgeht, daß die Spätsommergeneration weniger zahlenstark ist als die Frühlings- und die Som-

mergeneration (Schmid 1986), kann man das als Hinweis darauf deuten, daß wohl keine regelmäßigen

weitreichenden Wanderungen der ganzen oder erheblicher Teile der Population stattfinden. In der S-

Reuse wurden erheblich mehr albimanus gefangen als in der N-Reuse (3,3:1; n = 740). Die Analyse

der Windrichtungen an Hauptflugtagen zeigt, daß albimanus zu einem erheblichen Teil bei Gegen-

winden ausSW, S oder SE (59,1 %) in die S-Reuse fliegt, während bei Rückenwinden aus NW, N oder

NE weniger Tiere gefangen werden (12,9%) (Abb. 26h).

Während die Flugzeit der G'O’Ende August schon fast vorüber ist, werden @ Qdurch den ganzen

September hindurch gefangen (Abb. 4b, 4c). Insgesamt treten OO’ wesentlich spärlicher auf als Q2

(@X07210.—1:258:n 1833)

P. albimanns ist eine Art, die hauptsächlich um die Mittagszeit gefangen wird (Abb. 4d). Knapp die

Hälfte gerät zwischen 11 und 14 Uhr in die S-Reuse.

Status: Saisonaler Dismigrant.

Hauptsächlich Q9 führen im Zusammenhang mit der Suche nach Eiablageplätzen Wanderungen

aus.

Platycheirus clypeatus © (Ph.: U. Schmid)

Platycheirus clypeatus (Meigen 1822)

Verbreitung: holarktisch

Biologie: Platycheirus clypeatus ist eine meist häufige, eurytope Art, die sich zu einem großen Anteil

von Pollen blühender Gräser ernährt. Die Flugzeit beginnt (Anfang April) Anfang Mai und endet An-

fang (Ende) Oktober. Innerhalb der Flugzeit werden mehrere Generationen gebildet. Die Larve über-

wintert. Auch c/ypeatus-Larven werden nur selten an Blattlauskolonien gesehen. Meist werden sie in

der Streuschicht gefunden, wo sie vermutlich als unspezialisierte Prädatoren von Bodenarthropoden

leben (Rotheray & Gilbert 1989).

Wanderungen: Zum Wanderverhalten dieser Art gibt es bisher nur wenige Beobachtungen, die

überwiegend aus dem Küstenbereich stammen. Bei starkem Südwind (6 Bft.) und Temperaturen bis

zu 24°C erschienen Tausende von Tieren mit dem Wind auf der 15 km nördlich der niedersächsischen

Küste liegenden und nur 14 ha großen Düneninsel Scharhörn (Schmid 1987). Lucas (in litt.,

24.1.1988) fand im Spülsaum der holländischen Küste bei ’s Gravenzande am 29.8.1987 neben den

Saisonwanderern Syrphus torvus, Metasyrphus corollae und Episyrphus balteatus auch 2 Exemplare

von clypeatus. Auf dem Feuerschiff „Fehmarn-Belt“ wurden innerhalb drei Wochen im Juli/August

20 Exemplare gefangen (Heydemann 1967). Dagegen gerieten am CB im Verlauf von zwölf Jahren le-

diglich 25 clypeatus in die Reuse (Aubert et al. 1976).

Phänologie am Randecker Maar: P. clypeatus ist nach Episyrphus balteatus und Melanostoma mel-

linum die dritthäufigste Art in der S-Reuse (Tab. 2). Sie zeigt ein sehr deutliches Maximum im August

(Abb. 5a) mit Median am 10.8. Die Verteilungskurve ist nahezu symmetrisch. Aus den Diagrammen

einzelner Jahre (Abb. 5b) wird deutlich, daß sie im Einzelfall zwar erheblich vom Summendiagramm

Anzahl pro Pent

Anz. pro Tag (+) elypbestös

3200 1975 - 1987

1600 +[ EN N ’ I

Er. 19.6 () 1120 |

5% 21.7 12 J)= 13450 |

25% 3.8 75 "

2880 5 1

1440 ER we 17 76 181

75% 18. 8 ee 3:

95% 4.9 = EN

Le. 6.10 15. 8.80 98 |

es 1.8. ee]

1280 + Re 51

8.54 172

| fe 12.8.85 175

9.8.86 333

2240 |

1120 + | 8.87 266

[0]

Abb. 5a: Nach Süden gerichtete Migration von Platycheirus clypeatus (vgl. Legende zu Abb. 4a).

Fig. 5 a: Southbound migration of Platycheirus clypeatus.

er u EEE ns: Er Re

abweichen können, insbesondere, wenn nur wenige Tiere gefangen wurden. Das Grundmuster mit

dem ausgeprägten Maximum Mitte August ist aber fast immer deutlich erkennbar.

Das Geschlechterverhältnis ist ausgeglichen (CO: = 1:1,1;n = 10968). Im jahreszeitlichen

Auftreten der beiden Geschlechter liegt der Unterschied vor allem im weitgehenden Fehlen der GC

im September. Deshalb liegt der Median der 0°C (10. 8.) etwas früher als der der P9 (15. 8.) (Abb. 5c,

5d).

Anzahl pro Pent

Anz. pro Tag (+)

800

400 +

560

280 +

320

160 +

40 +

1980

An. Bee 72

ID SREad. 7,

25% 3.8

| 50% 8.8

r 75% 18. 8

1984

Enmocbe,;

I SR729. 7;

25% 1.8

50% 5.8

I 75% 13. 8

95% 28.8

la. Be)

u ——

Anzahl pro Pent

Anz. pro Tag (+)

800

400 +

550

280 +

320

160 +

40 +

L 75% 19

1985

Er. 29

5% 1

25% 9

50% 13

ommmm \n

95% 31

Le. 4.10

Tossa Toms Tamat

Jun

800 1982

400+[

= | 8

560 5%. 20.7

280+| 25% 1.8

50% 15. 8

320 75% 3.9

Acor | R

so Le. 27.9

40 +

Jun

800 1984

400

Ds Er

560 5%

320 | 7%

35%

80 Le

40+|

- T

Jun

bj}

Abb. 5 b: Nach Süden gerichtete Migration von Platycheirus clypeatus in den Jahren 1980 (oben) bis 1987 (unten)

in Tages- und Pentadensummen (1.-5. Jan., 6.-10. Jan. etc.). Auf der Ordinate werden Tagessummen (untere

Zahl, mit + markiert) und Pentadensummen (obere Zahl) angegeben. An welchem Tag 5, 25, 50, 75 und 95% der

Gesamtsumme durchgezogen sind, kann links oben abgelesen werden. Die dreieckigen Pfeile über dem Diagramm

kennzeichnen die Lage der 25 %-, 50 %- und 75 %-Werte.

Rechts oben wird die Jahressumme (n) und ihr Abweichen vom 12jährigen Mittelwert (in %) ausgewiesen. Die dar-

unter aufgeführten vier stärksten Zugtagc erleichtern eine Orientierung ım Diagramm.

Fig. 5b: Southbound migration of Platycheirus clypeatus in the years 1980 (above) to 1987 (below) with the sum of

days and the sum of 5-day periods (see legend of Fig. 4a). On the upper right side the total number of specimens

in each year (n), its deviation from the 12-year-average (in %) and the four days of the year with most migrating

specimens are shown.

Das tageszeitliche Fangmuster von clypeatus zeigt Abb. 5e: Die Art weist demnach beı nahezu

gleichmäßiger Verteilung einen typischen Höhepunkt am frühen Nachmittag auf. (Die nach Fängen

von 1975-77 extrapolierten Spätnachmittagsdaten nach 16 Uhr sind im Diagramm mit Sicherheit un-

terrepräsentiert.) Dieses Muster ist durchgängig zu beobachten (Abb. 5h). Ein geschlechtsspezifischer

Unterschied besteht in der längeren Aktivitätsphase der OO’ gegenüber einem deutlicheren nachmit-

täglichen Aktivitätsmaximum der PQ (Abb. 5f und 5g).

Anzahl pro Pent

Anz. pro Tag (+) clypegtus W

1300 _ 1975 - 1987 TER ©

SE ERROLE 0=- an

5% 29.7 12 J= 5646

285% 7.8

alaiv [Ess lan & : z in 4

ea0+| mes. 8 25.8.7 29

95x 11729 6.8.79 69

ua Naraelıl 26. B.80 33

1120 | 1.0.01 89|

560 + POREIMBE 41

2.8.84 37

ale has 73

980 9.8.86 129

490 + 21. 8.87 118 4

Anzah! pro Pent |

Anz. pro Tag (#) erivBbezazlSs

1200 Kassel re sao |

SEE en ee © f) 443 420 + h 4

' K 12 U= 5322 |

5% 28 |

| 25% 3.8 6.8.5 2 |

o 50% 10. 8 m 700 | |

ln = u 3 15 = ) 350 + 12] ’

| 35% 31.8 | I ] 2.8.79 49

2 ] 1.8.80 67 |

eco | | 1.8.81 142 |

40 20. 7.82 40

2.8.84 135

| 12.8.8565 105

- | | 10. 8.86 221

Sc | ) 22.8.87 166 -

300 +

c ! um Jul ver. Se ee [6 | Jun Jul Aug Sep Okt

Abb. 5c: Nach Süden gerichtete Migration von Platycheirus clypeatus-C'O (vgl. Legende zu Abb. 4a).

Fig. 5c: Southbound migration of OO’ of Platycheirus clypeatus.

Abb. 5d: Nach Süden gerichtete Migration von Platycheirus clypeatus-? (vgl. Legende zu Abb. 4a).

Fig. 5d: Southbound migration of PP of Platycheirus chypeatus.

eIYPE3aLUS ce/ypeatus M

Jeanne 1.975 — 71987 vehre 1975 —- 1987

e f

* 25%= 3427 * 25%= 1399

S * 50%= 5854 * 50%= 2797

* 75%= 10281 * 75%= 4196

f Max= 1726 Max= 692

— * N = 13707 * N = 5594

RSS Je N = 12560 N = 4979

a 6 B 10 12 14 16 fi 18 m ; Unr a uhr

Abb. 5e: Tageszeitliche Verteilung von Platycheirus clypeatus (vgl. Legende zu Abb. 4d).

Fig. 5e: Diurnal activity patterns of Platycheirus clypeatus migrating southward.

Abb. 5f: Tageszeitliche Verteilung von Platycheirus clypeatus-I’T (vgl. Legende zu Abb. 4d).

Fig. 5f: Diurnal activity patterns of OO’ of Platycheirus clypeatus migrating southward.

Eine Analyse der Windrichtungen an den jeweils vier individuenreichsten Tagen aller Jahre

(Abb. 26i) ergibt, daß 29,2% der Fliegen bei Rückenwinden aus NW, N oder NE in die S-Reuse flo-

gen, 55,8% bei Gegenwinden aus SW, S oder SE (vgl. 3.4).

celypeatus

vahre 1975 — 1987 vanre 1975 —- 1977

4 f en 8 10 12 14 16 Ten ai 4.6 8 10 12 14 46 4B 20 Unr

Max= 39

30.9. \ N= 193

K NS

Max= 151

N= 871

cl/ypestus W

Max= 469 ei I Max= 18

16.- >16h= 2.3 %

N= 3070 31. 8. Y N= 88

[S Ih

vehre 19753 - 1987

Max= 880

*N= 7253 de

N= 6471 15.8

Max= 29

>416h= 12.1 %

Nas rag] N” 340

Zur rn — a EEE ra SE nn EB

EEE nn ET an

20 Unr 4 6 8 10 ı2 14 16 1B 20 Uhr 4 6 8 10 12 44 16 418 20 Uhr

Abb. 5g: Tageszeitliche Verteilung von Platycheirus clypeatus-QQ (vgl. Legende zu Abb. 4d).

Fig. 5g: Diurnal activity patterns of QP of Platycheirus clypeatus migrating southward.

Abb. 5h: Tageszeitliche Verteilung von Platycheirus clypeatus im Jahresverlauf nach Fängen aus der S-Reuse in den

Jahren 1975— 1977 (rechts; Erfassung von der Morgen- bis zur Abenddämmerung) und 1975— 1987 (links; Erfas-

sung bis 16.00 Uhr).

Die Einzeldiagramme zeigen die tageszeitliche Verteilung der Fänge in Halbmonatswerten von Juli bis November.

Die Stunde, die den 50 %-Wert enthält, ist kreuzschraffiert, die Stunden, die die 25 %- und 75 %-Werte enthalten,

sind eng schraffiert. Die Strichmarkierungen über diesen Säulen bezeichnen die aus Stundenwerten errechnete Lage

des 25 %-,50 %- und 75 %-Wertes genauer. Die Höhe der Maximalwerte der Einzeldiagramme ist vereinheitlicht.

Die Zahl der im betreffenden Zeitraum gefangenen Individuen (N) und der höchste Stundenwert (Max) ist jeweils

rechts angegeben. Die Tageszeit ist unten aufgetragen (Abszisse). Die flankierenden Begrenzungslinien zeigen den

Zeitpunkt des Sonnenauf- und Sonnenuntergangs für Stuttgart (Astronomisches Recheninstitut Heidelberg).

In der rechten Spalte ist mit >16 Uhr die Zahl der nach 16 Uhr gefangenen Fliegen angegeben. Auf dieser Basis wur-

den für die linke Spalte die “N-Werte hochgerechnet und die 25 %-,50 %- und 75 %-Werte errechnet. Damit sind

Schraffur und Markierungen beider Darstellungen rechnerisch und optisch vergleichbar.

Fig. 5h: Diurnal activity patterns of Platycheirus clypeatus migrating southward throughout the year in 19751977

(on the right; observations from sunrise to sunset) and in 1975-1987 (on the left; observations till 4 p.m.) (see le-

gend of Fig. 4d). Each diagram shows the diurnal pattern of a fortnight between July and November. On the right,

the total number of specimens (N), the number of specimens in the most frequented hour (Max) and after 4 p.m.

(>16h) are shown. *N is a fictitious number arrived at by extrapolization of afternoon catches from 1975— 1977

over the whole period of investigation. In all diagrams the 25 %, 50 %, and 75 % values are calculated from *N.

The side lines of the columns show the time of sunrise (left) and sunset (right).

72)

Das weitgehend ausgeglichene Verhältnis der Fangzahlen zwischen S-Reuse und N-Reuse (1,7:1;

n = 5964) (Abb. 27c) ist ein Hinweis darauf, daß Platycheirus clypeatus zumindest im bodennahen Be-

reich ohne ausgeprägte Richtungspräferenz wandert.

Status: Windabhängiger Saisonaler Dismigrant.

Die Beobachtungen aus dem Küstenbereich und vom RM zeigen, daß c/ypeatus regelmäßig und oft

in großer Zahl wandert. Das Fehlen von Wanderbeobachtungen der Art in den Alpen läßt darauf

schließen, daß sie nicht ın der Lage ist, größere vertikale Distanzen zu überwinden.

Platycheirus manicatus (Meigen 1822)

Verbreitung: Europa, Altaı

Biologie: Die eurytope Art P. manicatus fliegt von (Mitte) Ende April bis Ende September (Mitte

Oktober) und bildet in Mittel- und Nordeuropa vermutlich zwei Generationen, wobei die Frühjahrs-

generation wesentlich zahlenstärker als die Spätsommergeneration ist. P. manicatus ist weit verbreitet

und nicht selten. Im Larvenstadium ist manicatus vermutlich nicht obligatorisch, sondern fakultativ

aphidophag (Chandler 1968a). Die Larve überwintert in Diapause.

Anzahl pro Pent

Anz. pro Tag (+) MSNIZEcEsgteus

140 1975 - 1987 a : .

70 + [ Er ee () 61

r ou

| 5% 26. 7 | ] ; =

25% 6.8 | 0.9.7

120 | ie] 0.. 9.78 8

E05 1a. || 1.77% N

| 75% 25. 8 | ar BUY, |

95% 12.9 DE Ei

| Le. 6.10 | 6.8.79 1

100 L | | 26. 8.80 35 |

50+| | | 15. 8.81 16

| 8 11

| 8 18

80 |

20+| | A

30 +

40 |

———— © ill

a Jun Jul Aug Sep J Okt

Abb. 6a: Nach Süden gerichtete Migration von Platycheirus manicatus (vgl. Legende zu Abb. 4a).

Fig. 6a: Southbound migration of Platycheirus manicatus.

Wanderungen: In den Alpen ist der Status von manicatus unklar. Aubert et al. (1976) lassen offen,

ob die Art wandert („consideree comme migratrice“). Das Fangdiagramm weist ein sehr deutliches

Maximum in der letzten August- und der ersten Septemberdekade auf. Ein weiterer Hinweis auf Wan-

derung in den Alpen findet sich bei Burmann (1978). Funde von manicatus auf Feuerschiffen und In-

seln (Heydemann 1967, Schmid 1987) geben Hinweise darauf, daß die Art Wandertendenzen auf-

weist. Auch in Südengland wurde aus wandernden Schwebfliegen ein ©’ von manicatus gefangen (Par-

menter 1960).

Phänologie am Randecker Maar: Die Unterschiede zwischen der Anzahl der Fänge in beiden Reu-

sen (Abb. 27d) sind bei dieser Art sehr gering (S-Reuse :N-Reuse = 1,5:1;n = 826), so daß zweifelhaft

erscheint, ob manicatus im Herbst gerichtete Sidwanderungen durchführt. Die Fangzahlen sind aller-

dings angesichts der Spärlichkeit der Spätsommergeneration von P. manicatus recht hoch (Tab. 2).

MIMICIEUS Mamicatcus M

vahre 1975 —- 1987 veahre 1975 - 1987

* 25%= 186

vehre 1975 — 1987

Abb. 6b: Tageszeitliche Verteilung von Platycheirus manicatus (vgl. Legende zu Abb. 4d).

Fig. 6b: Diurnal activity patterns of Platycheirus manicatus migrating southward.

Abb. 6c: Tageszeitliche Verteilung von Platycheirus manicatus-J'J (vgl. Legende zu Abb. 4d).

Fig. 6c: Diurnal activity patterns of OO’ of Platycheirus manicatus migrating southward.

Abb. 6d: Tageszeitliche Verteilung von Platycheirus manicatus-?Q (vgl. Legende zu Abb. 4d).

Fig. 6d: Diurnal activity patterns of @ of Platycheirus manicatus migrating southward.

Am Randecker Maar wird P. manicatus überwiegend im August gefangen (Abb. 6a). Die Phänolo-

gie dieser Artam RM zeigt große Ähnlichkeit mit der am CB. Der einzige Unterschied liegt in der zeit-

lichen Versetzung des Maximums: Mitte August am RM, Ende August/Anfang September am CB.

Das Gros der O'CJ erscheint etwas vor dem der 2 (Median der CC’: 13.8.; Median der PP: 17.8.).

QQ werden etwas häufiger gefangen (J'O':Q 9 = 1:1,2;n = 550).

Während der wärmeren Vor- und Nachmittagsstunden gerät P. manicatus ziemlich gleichmäßig ın

die S-Reuse (Abb. 6b). Die Flugaktivität der O’C° ist am Vormittag deutlich höher als die der P

(Abb. 6c, 6d).

An starken Flugtagen gerät ungefähr die Hälfte (52,5%) der Fliegen bei Gegenwind aus SW, S oder

SE in die S-Reuse, ungefähr ein Viertel (24,2%) bei Rückenwinden aus NW, N oder NE (Abb. 26k).

Status: Saisonaler Dismigrant (?).

Die gleichartige Phänologie der Art am RM und CB mit der zeitlichen Medianverschiebung zwi-

schen RM (früher) und CB (später) lassen großräumige Wanderungen vermuten, die über den Rah-

men einer Dismigration hinausgehen (vgl. 4.1). Die geringen Unterschiede zwischen beiden Reusen

stellen allerdings solche weitreichenden richtungsorientierten Wanderungen der Gesamtpopulation

in Frage. Auch verhält sich manicatus gegenüber unterschiedlichen Windrichtungen wie der typische

Dismigrant P. chypeatus (Abb. 26i, 26k; vgl. 3.4). Vermutlich haben die Ähnlichkeiten der Phänologie

am RM und CB regionale Ursachen und spiegeln nur die Flugzeit der Herbstgeneration am jeweiligen

Ort wider. Um den Status dieser Art endgültig zu klären, bedarf es weiterer Beobachtungen.

27,

Platycheirus peltatus (Meigen 1822)

Verbreitung: holarktisch

Biologie: Die Art ist weit verbreitet und zahlreich. Sie fliegt zwischen (Ende März) Mitte Mai und

Mitte (Ende) Oktober mit Maxima im Mai/Juni und August/September. Die Larve ist vermutlich fa-

kultativ aphidophag (Chandler 1968). P. peltatus überwintert als Larve ın Diapause.

Wanderungen: Die einzigen Hinweise auf mögliche Wanderungen stammen aus dem Küstenbe-

reich, wo einige Individuen auf Feuerschiffen und einer Insel gefangen wurden (Heydemann 1967,

Schmid 1987). In den Alpen trat P. peltatus nicht als Wanderer in Erscheinung (Tab. 2; Aubert et al.

1976).

Anzahl pro Pent.

Anz. pro Tag (#) BWedIeEnee)S

1975 - 1987

22 Aue] R A| v 2.

ans la BD = 22

BABEITEBr 12 J 272

25% 14. B. 2.9.5 N

so | 50% 17.8. |

25 + 25.68.77 6

LOrSUTERE 152387779 16

si Es: 29. 8.80 10

Eer230r,9 16. 8.81 20

so | 18. 8.82 g |

40 +

Zoe N

zians5

30 +

40 |

20 +

30 |

15 +

peltatus

20 A Jahre 1973 71987:

De b

= 2 * 25%= 64

= * 50%= 128

* 75%= 192

Max= 39

*N = 256

N = 256

r "a ame = | uf

a sr Sn AUS Sei elSıs > unr

Abb. 7a: Nach Süden gerichtete Migration von Platycheirus peltatus (vgl. Legende zu Abb. 4a).

Fig. 7a: Southbound migration of Platycheirus peltatus.

Abb. 7b: Tageszeitliche Verteilung von Platycheirus peltatus (vgl. Legende zu Abb. 4d).

Fig. 7b: Diurnal activity patterns of Platycheirus peltatus migrating southward.

Phänologie am Randecker Maar: P. peltatus wird überwiegend im August gefangen (Abb. 7a). CC

sind dabei wesentlich seltener als 9 (I’T':Q9 = 1:5,8;n = 203). In der S-Reuse werden mehr pel-

tatus gefangen als in der N-Reuse (3,0:1;n = 262) (Abb. 27e).

Das tageszeitliche Muster zeigt, von den frühen Morgenstunden abgesehen, eine gleichmäßig hohe

Aktivität (Abb. 7b).

Status: Saisonaler Dismigrant.

An den bodennahen Wanderungen nehmen überwiegend @ Q, wohl auf der Suche nach günstigen

Eiablageplätzen, teil.

Melanostoma mellinum (Linnaeus 1758)

Verbreitung: holarktisch, neotropisch? (Argentinien?)

Biologie: Melanostoma mellinum ist eine eurytope, meist sehr häufige Art, die in Mitteleuropa von

(Anfang) Mitte April bis Mitte Oktober (Mitte November) zu sehen ist. Innerhalb dieser Zeit sind oft

zwei Maxima im Frühsommer und im Spätsommer zu erkennen. Die größte Häufigkeit erreicht mel-

linum im Spätsommer. Melanostoma-Larven sind an Blattlauskolonien relativ selten. Häufiger kön-

nen sie in der Streuschicht gefunden werden (Rotheray & Gilbert 1989). Inwieweit dort auch pflanz-

liches Material aufgenommen wird, ist umstritten. Rotheray (1983) und Gilbert (1986) vermuteten,

Anzahl pro Pent.

Anz. pro Tag (+) me221mwum

3600 1975 - 1987 N:

SoSe a 7. D= 1161

BE. 7 12 U = 13937

s 25% 8.8. 15. 8.75 64

Pr GEIU SEISEE 16. 7.76 3

75% 23. 8. 17. 8.77 34

95% 11.9. 15. 8.78 436

Le. 13.10. 2.8.79 75

2sso | 15. 8.80 72

279 105% 7). 8.81 4177

26. 8.82 86

2.8.84 16

14. 8.85 244

BBeen 15. 8.86 109

1.9.87 209

mEell1MUM

vahre 1975 —- 1977 vehre 1975 — 1987

25%= 193 * 25%= 3759

B 10 12 14 16 18 20 Uhr

Abb. 8a: Nach Süden gerichtete Migration von Melanostoma mellinum (vgl. Legende zu Abb. 4a).

Fig. 8a: Southbound migration of Melanostoma mellinum.

Abb. 8b: Tageszeitliche Verteilung von Melanostoma mellinum in den Jahren 1975-1977 (vgl. Legende zu

Abb. 4d).

Fig. 8b: Diurnal activity patterns of Melanostoma mellinum migrating southward in 1975— 1977.

Abb. 8c: Tageszeitliche Verteilung von Melanostoma mellinum in den Jahren 1975-1987 (vgl. Legende zu

Abb. 4d).

Fig. $c: Diurnal activity patterns of Melanostoma mellinum migrating southward ın 1975-1987.

mellinum M mellinum W

vahre 1975 — 1977 vehre 1975 — 1977

h D\ S =

6 8 10 12 14 16 18 20 Uhr 4 6 8 10 12 14 16 18 20 Uhr

Abb. 8d: Tageszeitliche Verteilung von Melanostoma mellinum-J'C (vgl. Legende zu Abb. 4d).

Fig. 8d: Diurnal activity patterns of O’C’ of Melanostoma mellinum migrating southward.

Abb. 8e: Tageszeitliche Verteilung von Melanostoma mellinum-QQ (vgl. Legende zu Abb. 4d).

Fig. 8e: Diurnal activity patterns of PP of Melanostoma mellinum migrating southward.

daß mellinum-Larven auch verrottete Blätter aufnehmen. In einer jüngeren Arbeit (Rotheray & Gil-

bert 1989) vertreten sie dagegen die Auffassung, daß die Art rein zoophag sei und in der Streuschicht,

als Alternative zur bevorzugten Blattlausbeute, Bodenarthropoden nachstelle. Nach Bastian (1986)

gehört bei Nahrungsmangel neben Schmetterlingsraupen auch pflanzliches Gewebe zum Nahrungs-

spektrum von mellinum. Die Art überwintert im Larvenstadium.

Wanderungen: Am Col de Bretolet wurde M. mellinum so zahlreich in den Reusen gefangen, daß

Aubert et al. (1976) die Art als Wanderer bezeichnen (Tab. 2). Mellinum kommt dort während der

ganzen Beobachtungsperiode vor. Zwischen Mitte Juli und Mitte September ist sie immer häufig,

ohne daß Maxima erkennbar wären. Prell (1925) fing an einem Paß in den Vierwaldstätter Alpen wäh-

rend einer Insektenwanderung auch Belege von mellinum.

Aus dem Flachland fehlen entsprechende Beobachtungen. Angesichts der überall sehr großen Häu-

figkeit dieser Art sind Beobachtungen an Stellen, die nur durch Wanderung zu erreichen sind, wie

z. B. Nordseeinseln ohne bodenständige Populationen und Feuerschiffe, ziemlich spärlich (Heyde-

mann 1967, Lempke 1962, Schmid 1987).

Phänologie am Randecker Maar: Melanostoma mellinum ist nach Episyrphus balteatus die zweit-

häufigste in der S-Reuse gefangene Art (Tab. 2). Sie wird hauptsächlich im August gefangen und weist

in diesem Monat einen starken Höhepunkt auf. Der Median liegt am 15.8. (Abb. 8a). GC erscheinen

in wesentlich geringerer Zahl als PP (CT: PQ = 1:3,6; n = 11974). Die Phänologie beider Ge-

schlechter unterscheidet sich im wesentlichen nur dadurch, daß die GC (25% am 5.8., Median am

15.8., 75% am 20.8.) etwas früher auftreten als die PP (25% am 12.8., Median am 16.8., 75% am

26.8.).

Für M. mellinum ist eine ausgesprochen lange tägliche Aktivität typisch (Abb. 8b, 8c). Der im Dia-

gramm erkennbare Gipfel ın den frühen Morgenstunden geht auf das Konto der ?2 (Abb. 8e); die

IT zeigen dagegen einen gleichmäßigen Aktivitätsanstieg bis Mittag (Abb. 8d). In der S-Reuse wur-

den an den vier individuenstärksten Tagen aller Jahre 64,6 % der Tiere bei Gegenwinden ausSW,S und

SE gefangen, 22,3 % bei Rückenwinden aus NW, N oder NE (Abb. 261). Auch diese kleine und wenig

flugtüchtig scheinende Art kann also durchaus gegen schwache Winde anfliegen. Sie wandert vermut-

lich bei Rückenwinden zu einem erheblichen Teil oberhalb des Fangbereiches der Reusen.

Die Unterschiede in den Fangzahlen zwischen S-Reuse und N-Reuse sind nicht allzu hoch (2,4:1;

= 6678) (Abb. 27f).

Status: Saisonaler Dismigrant.

Auch bei dieser Art dominieren PP. Die meisten dieser PP haben fast kugelförmig angeschwol-

lene Hinterleiber. Ihre Wanderungen dürften hauptsächlich dazu dienen, günstige Eiablageplätze zu

finden (vgl. 4.4.4). Das weitgehende Fehlen dieser sehr häufigen Art an isolierten Beobachtungsstatio-

nen (z. B. Feuerschiffe) deutet darauf hin, daß ihre Wanderungen nur über kurze Distanzen führen.

Da mellinum selbst im Bereich der alpınen Matten in großer Zahl beobachtet werden kann, können

auch viele der am CB gefangenen Tiere aus der näheren Umgebung stammen.

Melanostoma scalare (Fabricius 1794)

Verbreitung: paläarktisch, orientalisch

Biologie: Melanostoma scalare ist, wenngleich immer noch häufig, so doch weniger zahlreich als

mellinum. Die Art ist besonders an waldigen Standorten zu finden. Die Flugzeit stimmt mit der von

mellinum überein. Auch scalare weist gewöhnlich zwei Maxima auf. Besonders häufig ist die Art im

Frühjahr. Die Larve überwintert. Ihre Ernährungsweise dürfte der von M. mellinum entsprechen.

Wanderungen: Am Col de Bretolet wurden fast keine scalare gefangen (Tab. 2). Auch weitere Hin-

weise auf Wanderungen im Alpenraum fehlen. Aus Norddeutschland liegen einige wenige Funde

nicht ansässiger Tiere von Feuerschiffen (Heydemann 1967, Lempke 1962) und von der Insel Schar-

hörn vor (Schmid 1987).

Phänologie am Randecker Maar: Die häufige Art wurde auch hier so selten in den Reusen gefangen

(Tab. 2), daß regelmäßige Wanderungen oder gelegentliches Verdriften in größerem Ausmaß ausge-

schlossen erscheint.

Status: Saisonaler Dismigrant mit sehr geringer Wanderdisposition.

Anzahl pro Pent

Anz. pro Tag (#) v2

1000 1975 - 1987 NER:

50o0o+[ ]

"7 2% 0.6

I Tl

rare

—

voomomsmmmmm m n

Dom mmmssusnsmn

ouLLnD-OoSDBHoNJau

o [nt [e]

[e]

[2]

Abb. 9a: Nach Süden gerichtete Migration von Syrphus vitripennis und Syrphus ribesü (vgl. Legende zu Abb. 4a).

Fig. 9a: Southbound migration of Syrphus vitripennis und Syrphus ribesii.

Anzahl pro Pent.

Anz. pro Tag (+) VIREN ENTER SEN

300 _ 1975 - 1987 n Al, !

an Ei: nie

5% 30.7 =

SSR dr 14. 8.75 4

wenn. 23. 8.76 44

er re Sr ;

ee 15. 8.78 g

Le. 10.10 14. 8.79 28

240 17. 9.80 5

seo 3.8.81 17

3.7.82 8

21. 8.84 37

210 59.8585 38

| | 20 3

Anzahl pro Pent .:

Anz. pro Tag (+) NEST 31 W |

240 1975 - 1987 186 | |

120+[ mlz Bo |

200 150 ! |

100+|[ 75+| |

160 ıB 120 | |

een so+| |

120 so | ll: I |

za: 45 + { 53

eu 80 if) |

40 + +

a |

20 +

TIDESII W

le lee — EEE

25%= 444

50%= 888

75%= 1332

Max= 253

N = 1776

N = 1549

* *

4 6 B 10 12 14 16 18 20 Unr

Abb. 9b: Nach Süden gerichtete Migration von Syrphus ribesü-QQ (vgl. Legende zu Abb. 4a).

Fig. 9b: Southbound migration of PP of Syrphus ribesii.

Abb. 9c: Nach Süden gerichtete Migration von Syrphus vitripennis-QQ (vgl. Legende zu Abb. 4a).

Fig. 9c: Southbound migration of PP of Syrphus vitripennis.

Abb. 9d: Tageszeitliche Verteilung von Syrphus ribesü-Q 9 (vgl. Legende zu Abb. 4d).

Fig. 9d: Diurnal activity patterns of QQ of Syrphus ribesii migrating southward.

VIETIDENMIS W

vahnre 1975 — 1987

* 25%= 274

* 50%= 548

* 75%= 821

Max= 177

* N = 1095

N = 1022

Uhr

Abb. 9e: Tageszeitliche Verteilung von Syrphus vitripennis-QQ (vgl. Legende zu Abb. 4d).

Fig. 9e: Diurnal activity patterns of PP of Syyrphus vitripennis migrating southward.

Syrphus ribesii (Linnaeus 1758)

Syrphus torvus Osten Sacken 1875

Syrphus vitripennis Meigen 1822

In den meisten Jahren wurde nur aufgrund der bei Sack (1932) genannten Kennzeichen zwischen

den Syrphus-Arten unterschieden. Die OO’ des Artenpaares rıbesi/vitripennis sind damit aber nicht

zuverlässig zu unterscheiden. Die drei häufigeren Syrphus-Arten werden deshalb hier gemeinsam ab-

gehandelt. Syrphus rıbesii und S. vitripennis dürften am RM in ungefähr gleich großer Häufigkeit vor-

kommen, Syrphus torvus ist hier wesentlich seltener.

Verbreitung: rıbesu: holarktisch; torvus: holarktisch, orıentalisch (Indien, Nepal, Thailand, For-

mosa); vitripennis: holarktisch, orientalisch (Formosa).

Biologie: Alle drei Arten sind weit verbreitet und gehören ın vielen Jahren zu den häufigsten

Schwebfliegenarten. Sie sind eurytop, wobei torvus seinen Schwerpunkt ın Waldbiotopen hat. Sie

überwintern als Diapause-Larven. Der Lebenszyklus von ribesiz wurde von Schneider (1948) nach

Daten aus der Schweiz genauer dargestellt: demnach können Imagines zwischen Mitte April (in Bel-

gien Mitte März; Verlinden & Decleer 1987) und Mitte November gesehen werden. In dieser Zeit wer-

den maximal vier Generationen gebildet. Meist treten jedoch nur zwei bis drei Generationen auf, weil

ein Teil der Altlarven — im Frühjahr weniger, im Herbst immer mehr — in Diapause geht. Insgesamt

ergibt sich so eine lange Flugzeit mıt Maxima ım Frühjahr und im Frühherbst.

Für S. torvus (Flugzeit [Mitte März] Anfang April bis Mitte Oktober) und $. vitripennis (Flugzeit

[Ende März] Mitte April bis Mitte Oktober [Ende November) sind ähnliche Verhältnisse zu erwar-

ten.

Syrphus-Larven sind aphidophag mit einem weiten Beutespektrum (Läska & Stary 1980) und so-

wohl auf Bäumen als auch an Büschen und Kräutern zu finden (Dixon 1960, Rotheray & Gilbert

1989).

Wanderungen: Gatter (1980) beobachtete im Mai 1979 ım Himalaya nordwärts gerichtete Dipte-

renwanderungen über einen 5400 m hohen Paß. Unter den 15 gesammelten Belegexemplaren gehör-

ten 6 zur Gattung Syrphus (4 S. rıbesii). Zahlreiche Beobachtungen von sommerlichen Südwanderun-

gen liegen von allen drei Arten sowohl aus dem Gebirge (Alpen: Aubert et al. 1976, Burmann 1978,

Gepp 1975, Jeekel & Overbeek 1968; Pyrenäen: Snow & Ross 1952, Williams et al. 1956) als auch von

Feuerschiffen und von den Küsten vor (Heydemann 1967, Lempke 1962, Mackworth-Praed 1929,

Overgaard Nielsen 1968, Schmid 1987). Syrphus ribesüi ist sogar von Spitzbergen nachgewiesen (Elton

1925). Ein ganzer Schwarm von ribesii wurde am 5.8.1957 nachts 200 km vom dänischen Festland

entfernt von einem hell erleuchteten Fischereiboot angezogen. „Die Menge der einfallenden Fliegen

war so groß, daß sie eine heftige Belästigung der auf dem Deck arbeitenden Mannschaft... darstellten“

(Weidner 1958). Auch in Lichtfallen auf Helgoland war ribesii mit Abstand die häufigste Schwebflie-

genart (Gatter 1981a: 67).

Alle drei Arten wurden von Aubert et al. (1976) als Wanderer eingestuft.

Phänologie am Randecker Maar: Die Syrphus-Arten werden hauptsächlich im August gefangen

(Abb. 9a). Die deutliche Zweigipfeligkeit des Diagramms kommt durch die unterschiedlichen Zugzei- |

ten der Syrphus-Arten zustande. Der erste Gipfel wird hauptsächlich von ribesii, der zweite von vitri-

pennis gebildet. Dies zeigt ein Vergleich der Diagramme der (in allen Jahren sicher bestimmten) O9

von ribesi und vitripennis (Abb. 9b, 9c) deutlich. Daß ribesii (Median am 15.8.) etwas früher fliegt als

vitripennis (Median am 21.8.) deckt sich mit den bei Aubert et al. (1976) gezeigten Diagrammen, die

für ribesii nach einer Zugperiode ohne deutliche Maxima eine starke Abnahme der Zahlen im Septem-

ber zeigen, während vitripennis im September das Maximum aufweist.

Syrphus rıbesiui O (Ph. U. Schmid).

Von vitripennis stammen auch Fernfunde in den Alpen markierter Tiere über eine Distanz von

111 km (Aubert et al. 1969). Während in den Alpen vitripennis (43832 Ex.) wesentlich häufiger ist als

ribesüi (4047), dominiert in den Fängen am Randecker Maar die letzte Art (1582 ausgezählte OP) über

die erste (1046 ausgezählte PP).

Das tageszeitliche Aktivitätsmuster beider Arten unterscheidet sich nur leicht (Abb. 9d, 9e): gegen-

über ribesii hat vitripennis einen etwas späteren Aktivitätshöhepunkt. Ein Vergleich zwischen N-

Reuse und S-Reuse zeigt, daß die Syrphus-Arten überwiegend in der S-Reuse gefangen werden (Abb.

278) (S-Reuse: N-Reuse = 3,6:1). Nach einer Auswertung der Windrichtungen an Hauptflugtagen

fliegen 76,2% der Tiere bei Gegenwind aus SW, S und SE in die S-Reuse und nur 7,5% bei Rücken-

wind aus NW, N oder NE (Abb. 26a).

Status: Alle drei Arten sind Saisonale Migranten.

Metasyrphus corollae (Fabricius 1794)

= Eupeodes corollae

Verbreitung: holarktisch, paläotropisch

Biologie: Metasyrphus corollae ist eine eurytope und meist häufige Art. Sie fliegt in Mitteleuropa ab

(ausnahmsweise Ende März) Mitte Mai, bildet mehrere Generationen und ist bis in den Spätherbst

hinein zu sehen. Selbst aus dem im Dezember existieren Beobachtungen (Schneider 1958). In Belgien

ist corollae im Frühsommer mäßig zahlreich, erreicht dann Ende Juli sehr schnell große Abundanzen,

die erst Anfang September wieder schnell nachlassen (Verlinden & Decleer 1987). M. corollae über-

wintert als Puparium (Scott 1939). Die spärlichen mitteleuropäischen Nachweise im zeitigen Frühjahr

lassen vermuten, daß die Zahl der Überwinterer hier nicht sehr groß ist. Im nördlichen Mitteleuropa

und Nordeuropa scheinen überhaupt keine Überwinterungsmöglichkeiten zu bestehen. Hier wan-

dern im Frühjahr befruchtete Weibchen von Süden her ein und bauen neue Populationen auf (Torp

1984). Dementsprechend wird corollae hier erst spät beobachtet. In den Niederlanden, in Dänemark

und in Südnorwegen erscheint die Art erst Mitte Mai (van der Goot 1981, Nielsen 1971, Torp 1984).

Die unspezialisiert aphidophagen Larven von corollae leben hauptsächlich an Büschen und in der

Krautschicht (Dixon 1960, Läska & Stary 1980, Rotheray & Gilbert 1989).

Wanderungen: Die Nordwanderung im Frühjahr ist nicht durch direkte Beobachtungen belegt,

wird aber durch das späte Erscheinen der Art im nördlichen Mitteleuropa deutlich. Dagegen existieren

zahlreiche Daten zu sommerlichen und herbstlichen Wanderungsbewegungen in südliche Richtungen

aus dem Gebirge wie auch von der englischen, schwedischen, dänischen und deutschen Küste (z. B.

Aubert et al. 1976, Aubert & Goeldlin 1981, Gepp 1975, Heydemann 1967, Jeekel & Overbeek 1968,

Johnson 1960, Johnson 1969, Lempke 1962, Overgaard Nielsen 1968, Schmid 1987, Svensson & Jan-

Metasyrphus corollae © (Ph. U. Schmid).

Anzahl pro Pent

Anz. pro Tag (+) er oOmMoO ae

2400 1975 - 1987 |

1200+[ INN B

Er. 45.7 | od ‚02

| 7 | 12 U 8426

Sn | ee s e 4.8.75 190

1080+| 75415. 8 5= A 2 %

95% 2.9 2.8.78 4

Le. 10.10 17. 8.79 234

1920 29. 8.80 23

960 + a a

Abb. 10a: Nach Süden gerichtete Migration von Metasyrphus corollae (vgl. Legende zu Abb. 4a) (eingefügte Zeich-

nung aus van der Goot 1981).

Fig. 10a: Southbound migration of Metasyrphus corollae.

zon 1984, Williams et al. 1956). Sowohl im Flachland als auch im Gebirge ist corollae oft einer der häu-

figsten Wanderer. Am Col de Bretolet stellte die Art nach Episyrphus balteatus (52,8%) und Eristalis

tenax (26,5 %) mit 6,7 % die meisten Fänglinge (Tab. 2) und wurde von Aubert et al. (1976) als Wan-

derer eingestuft. Das Durchzugsmaximum liegt im August. Über den Wiederfang markierter Exem-

plare berichten Aubert & Goeldlin (1981).

Phänologie am Randecker Maar: M. corollae wird in großer Zahl gefangen (Tab. 2). Die Art er-

scheint mit einem deutlichen Schwerpunkt in der ersten Augusthälfte (Abb. 10a). SC’ und P9 sind

ungefähr gleich stark vertreten (CO: P = 1:1,1;n = 7545). Die Diagramme der beiden Geschlech-

ter unterscheiden sich nur dadurch, daß PP in der zweiten Augusthälfte und im September etwas

zahlreicher sind (Abb. 10b, 10c).

M. corollae zeichnet sich durch eine lange tageszeitliche Aktivitätsperiode mit einem leichten

Schwerpunkt in den Vormittagsstunden aus (Abb. 10d). Bei QQ ist dieser Vormittags-Schwerpunkt

wesentlich deutlicher ausgeprägt als bei 0°C’ (Abb. 10e, 10f). Das Aktivitätsmuster ändert sich im Jah-

resverlauf durch die mit fortschreitender Jahreszeit zu beobachtende allmähliche Einengung der im

Sommer sehr langen Phase hoher Flugintensität (Abb. 10g). Dieses Aktivitätsmuster unterscheidet

sich grundsätzlich von dem von Grosser (1979) mit Hilfe von Gelbschalenfängen ermittelten. Hier lag

das sehr deutliche Maximum in den ersten drei Morgenstunden (5-8 Uhr). In dieser Zeit sucht corol-

lae also intensiv nach Nahrung; erst in den späteren Morgenstunden setzt dann die Migration ein.

An den Hauptflugtagen fängt sich corollae hauptsächlich bei Gegenwind in der S-Reuse (Abb. 26b):

75,7% aller Fliegen geraten bei SW-, S- oder SE-Wind in diese Reuse, nur 9,2% bei Rückenwind aus

NW,N oder NE.

Anzahl pro Pent. Anzahl pro Pent.

Anz. pro Tag (#) ceomNmoI2ZIlIse MM Anz. pro Tag (+) ZEINFEDN NEE W

ee 1975 - 1987

1200 1975 198 au yes “Tr vv 7

Sol 4.7 0 = 303 | Er. 15. 7. B= 35

9,7 12 4= 3644 BE. 7 12 J)= 3904

25% 1. B 4.8.75 A ae e5% 3.8. 4.8.75 79

1080 [50% 4.8 16.7.6 58-1 So | E78 Won €

SO AB Bo 2, 8.7 2 75% 18. 8 Me Be 3

95% 24. 8 2.8.78 12 95% 6.9 15. 8.78 35

Le. 10.10 zul 21. 8.79 85 Le. 9.10 21. 8.79 70

sso | 1.8.80 8 | 960 | 29. 8.80 19

480 + 7.8.81 234 A) =: 3.8.81 270

4.8.82 23 4.8.82 14

3.8.84 220 3. 8.84 61

au0 a a! m

ae Mo 2% 47 Zoe Bil 112

9.8.87 22 22. 8.87 25

720

360 +

800

300 +

a8o

2430 +

s60 |

150 +

240

120 +

120

80 +

Abb. 10b: Nach Süden gerichtete Migration von Metasyrphus corollae-J’C (vgl. Legende zu Abb. 4a).

Fig. 10b: Southbound migration of CO’ of Metasyrphus corollae.

Abb. 10c: Nach Süden gerichtete Migration von Metasyrphus corollae-QP (vgl. Legende zu Abb. 4a).

Fig. 10c: Southbound migration of QQ of Metasyrphus corollae.

Abb. 10d: Tageszeitliche Verteilung von Metasyrphus corollae (vgl. Legende zu Abb. 4d).

Fig. 10d: Diurnal activity patterns of Metasyrphus corollae migrating southward.

Die Schwankungen der jährlichen Fangzahlen sind hoch (Abb. 27h). In den vier Jahren, in denen

beide Reusen betrieben wurden, fingen sich in der S-Reuse dreimal soviel corollae wie in der N-Reuse

(S-Reuse :N-Reuse = 3,0:1;n = 3182).

Status: Saisonaler Migrant.

Svensson & Janzon (1984) beobachteten Anfang August 1981 in S-Schweden eine Schwebfliegen-

wanderung, die fast ausschließlich von M. corollae bestritten wurde. Sie halten solche Wanderungen

für unregelmäßig und exogen induziert. Demnach ermöglichte eine hohe Blattlausdichte im Jahr 1980

den Aufbau hoher Schwebfliegenpopulationen; im folgenden Jahr 1981 sollen niedrige Blattlauspopu-

lationen die Abwanderung der Schwebfliegen erzwungen haben. Das ist mit Sicherheit für M. corollae

nicht gültig, denn diese Art kann in Nordeuropa nicht überwintern (s. o.). Ihre Populationen sind dort

coroillge M

JVanre 1973 — 1977

corollae

Pe Vanre 1975 - 1987 vahre 1975 - 1977

50%= 253

75%= 379

Max= 65

>16n= 32.7 X

N = 505

4 6 8 10 12 14 16 18 Gen 4 6 8 10 12 14 16 18 20 Uhr

6 8 12 14 16 18 20 Unr

25%= 1101

50%=- 2202

75%- 3303

Max= 482

N = 4403

N = 3319

unr Max= 5

Ne 20

corollge W

wahre 1975 - 1977 Max= 16

Ne 61

25%- 103 Max= 68

50%= 205

75%- 308 N= 416

Max= 56

>16h= 21.5 %

N = 410

1 Max= 226 Max= 5

Unr 16.- >16h= 17.6 %

N= 1363 31.8 Ne 34

vanre 1975 - 1987

Max= 633 Max= B0

"N= 5848 1.- Ä >16h= 30.9 X

N= 4468 15. 8 a 4 N= 599

IS * 25%= 1106

E Sos= = Max= 182 Jul Max= 37

= = ERS "N= 1837 16.- N >16h= 22.8 %

> N= 1496 31.7 NI|N= 328

2 N R = INDANOR N

N = 3641 mas

\ 4 6 8 10 i2 14 16 18 20 Uhr 4 6 8 10 12 14 16 18 20 Uhr

—— ı

14 16 18 20 Uhr 9

Abb. 10e: Tageszeitliche Verteilung von Metasyrphus corollae-J'J (vgl. Legende zu Abb. 4d).

Fig. 10e: Diurnal activity patterns of O’O’ of Metasyrphus corollae migrating southward.

Abb. 10f: Tageszeitliche Verteilung von Metasyrphus corollae-Q 9 (vgl. Legende zu Abb. 4d).

Fig. 10f: Diurnal activity patterns of PP of Metasyrphus corollae migrating southward.

Abb 10g: Tageszeitliche Verteilung von Metasyrphus corollae ım Jahresverlauf (vgl. Legende zu Abb. 5h).

Fig. 10g: Diurnal activity patterns of Metasyrphus corollae migrating southward throughout the year.

nicht von der Blattlausdichte des Vorjahrs abhängig, sondern von der Zahl der Einwanderer und der

Höhe der Blattlauspopulation im selben Jahr.

Sowohl die Einwanderung in Nordeuropa ım Frühjahr als auch die spätsommerlichen und herbst-

lichen Südwanderungen von corollae sind nicht exogen induziert, sondern integraler, endogen fixier-

ter Bestandteil des Lebenszyklus dieser Art.

Scaeva pyrastri (Linnaeus 1758)

Verbreitung: holarktisch

Biologie: $. pyrastri ist eine polyvoltine Art, die im zentralen Europa als Imago überwintern kann

(Schneider 1947, 1948, 1958, Dusek & Läska 1974). Zahlreiche Winter- und Frühlingsbeobachtungen

liegen aus den Schweizer Alpen vor (Schneider |. c.). Im Mittelgebirgsraum glückt die Überwinterung

wohl nur sehr selten (z. B. 19 am 8.3.1975; Kormann 1977. Je ein Februar- und Märznachweis aus

Belgien; Verlinden & Decleer 1987). Gewöhnlich sieht man die Art erst ab Mitte Mai. Aus den Nie-

derlanden gibt es einen April-Nachweis (20.4.1983, 1). Aus dem Mai liegen von dort insgesamt

Nachweise von 10 (20.5.) und 15 (ab 4.5.) vor. Im Juni wurden 49 9 beobachtet. Erst ab An-

Scaeva pyrastrı © (Ph. W. Gatter).

fang Juli waren beide Geschlechter zahlreich (van der Goot 1981, 1983, 1986a). Ähnlich ist die Situa-

tion in Belgien (Verlinden & Decleer 1987): wenige Frühjahrsbeobachtungen ausschließlich weiblı-

cher Tiere; erste O'C’ Anfang Mai; bis Mitte Juli mäßig häufig; Ende Juli bis Ende August sehr auffäl-

liges Maximum; Anfang September schnelles Absinken der Zahlen. In Dänemark erscheint pyrastrı ab

Ende Mai (Torp 1984), in Sidnorwegen ab Ende Juni (Nielsen 1971), ın Irland ım Juni (Speight et al.

1975).

Es handelt sich bei $. pyrastri also um eine Art, bei der befruchtete PP in Zentraleuropa überwin-

tern können. Das nördliche Mitteleuropa wird im Frühjahr durch von Süden her einwandernde Tiere

besiedelt. In welchem Ausmaß pyrastri im zentralen und alpinen Europa überwintert, ist unklar. Ob

das Herkunftsgebiet der Masse der mitteleuropäischen Einwanderer hier liegt, scheint zumindest

zweifelhaft. Wahrscheinlicher ist, daß — wie auch bei Episyrphus balteatus (S. 44) vermutet — die Me-

diterraneis Haupt-Überwinterungsgebiet ist.

Die Larven von pyrastri sind aphidophag; sie wurden an zahlreichen Blattlausarten v. a. in der

Krautschicht gefunden (Dixon 1960, Läska & Stary 1980, Rotheray & Gilbert 1989).

Wanderungen: Direkte Beobachtungen von Nordwanderungen im Frühjahr fehlen. Dagegen gibt

es zahlreiche Berichte von im Sommer und Herbst in südliche Richtungen wandernden Tieren aus den

Alpen, den Pyrenäen, von der Nord- und Ostseeküste und aus Süd-England (z. B. Aubert et al. 1976,

Gepp 1975, Heydemann 1967, Jeekel & Overbeek 1968, Johnson 1969, Lempke 1962, Mackworth-

Praed 1929, Overgaard Nielsen 1968, Schmid 1987, Walker 1864, Williams et al. 1956). Auch am Mit-

telgebirgsrand bei Halle (Saale) deutet eine Beobachtung auf Wanderungen von pyrastri hin (Grosser

& Klapperstück 1976; dort allerdings durch „synchronisiertes Schlüpfen vieler Tiere“ erklärt). Gele-

Anzahl pro Pent.

Anz. pro Tag (+) DERSEENT A

300 & 1975 - 1987 et

[®] -

a a a 58

5% 29. 7 1ie)= 885

2 8, Br Kae &

e70 50% 7.8. : sa

135+| 754 14.8 a :

95% 30.8 an 3

ler 3710 Bone 37

240 . 8.80 za

120 .8.81 70

ee 4

8.84 11

210 8.85 51

105+ Bi =

180

15:

75+[

eo |!

Bor:

so |

AUS) ==

80 Ik

Sie) Z-

45 +1,

Isla Inmwlgengl, Tmzalı T Tal: T

Okt

a Jun

Abb. 11a: Nach Süden gerichtete Migration von Scaeva pyrastri (vgl. Legende zu Abb. 4a) (eingefügte Zeichnung

aus van der Goot 1981).

Fig. 11a: Southbound migration of Scaeva pyrastri.

gentlich kann es zu gewaltigen Massierungen kommen (Rogers 1864 und Symes 1864 [zitiert in John-

son 1969], Williams 1961). Am Col de Bretolet ist pyrastri während der ganzen Beobachtungszeit an-

wesend, ohne ein ausgeprägtes Maximum aufzuweisen. Von Aubert et al. (1976) wird die Art, die hier

mit 11168 Ex. 0,5% der Fänge ausmacht (Tab. 2), als vermutlicher Wanderer geführt („espece consi-

deree comme migratrice“).

Phänologie am Randecker Maar: Hier steht pyrastri mit 885 gefangenen Exemplaren an achter Stelle

(Tab. 2). Die Art wurde überwiegend im August gefangen.” In der ersten Monatshälfte ist ein ausge-

* Im Jahr 1989 wanderte die Art am Randecker Maar erstmals in sehr großer Zahl. Am 16. Juli zogen hier Tausende

nach Süden.

PIF3SETI

vehre 1975 — 1977

e5x= 7

50%= 15

75%= 22

* 50%= 541

Abb. 11b: Tageszeitliche Verteilung von Scaeva pyrastri (vgl. Legende zu Abb. 4d).

Fig. 11b: Diurnal activity patterns of Scaeva pyrastri migrating southward.

prägter Gipfel ausgebildet (Abb. 11a). Q Q erscheinen häufiger als "CO (CC:QQ9 = 1:1,7;n = 654).

Das tageszeitliche Diagramm (Abb. 11b) zeigt eine nahezu symmetrische Verteilung. Der hohe

Wert zwischen 17 und 18 Uhr entstand durch Hochrechnung aus den Beobachtungen der Jahre

1975-1977, als die Beobachtungen bis Sonnenuntergang durchgeführt wurden (vgl. Legende zu

Abb. 4d). In diesen drei Jahren wurden insgesamt lediglich 29 pyrastri gefangen, davon 6 Expl. in die-

ser Stunde — der „Abendgipfel“ existiert also in der Realität nicht.

Zwischen den beiden Reusen bestehen erhebliche Unterschiede (Abb. 271): S-Reuse:N-Reuse =

6,0:1 (n = 341). Dabei fangen sich weitaus die meisten Tiere bei Gegenwinden aus SW, S und SE in

der S-Reuse (71,8%). Bei Rückenwinden aus NW, N und NE geraten nur 17,7% in die Reuse

(Abb. 26c).

Status: Saisonaler Migrant

Scaeva selenitica (Meigen 1822)

Verbreitung: paläarktisch, orıentalisch

Biologie: Wie bei Scaeva pyrastri überwintern auch bei dieser weniger häufigen Art befruchtete 2 9

(Schneider 1947, 1958). Überwinterungsmöglichkeiten finden die Imagines wieder im zentraleuropäi-

schen und alpinen Raum und - so ist jedenfalls zu vermuten — in weiter südlich gelegenen Regionen.

In den süddeutschen Mittelgebirgen kann die Art wenigstens gelegentlich ebenfalls erfolgreich über-

wintern (z. B. Kormann 1977), ebenso in den Niederlanden (van der Goot 1981), in Norddeutschland

(Schmid 1987) und in Süd-Norwegen (Nielsen 1971). Fehlende Frühjahrsdaten aus Dänemark (Torp

1984) und Großbritannien (Stubbs & Falk 1983) zeigen, daß Überwinterungen im nördlichen Mittel-

europa eher selten sind. Auch fehlen im Norden die im südlichen Mitteleuropa üblichen (z. B. Schmid

1986, Schneider 1. c.) späten Herbstbeobachtungen, die auf regelmäßige Überwinterungsversuche hin-

weisen könnten (späteste Daten aus Süd-Norwegen 16.8. [Nielsen I. c.], aus Dänemark Anfang Sep-

tember [Torpl. c.], aus Schleswig 22.8. [Claußen 1980]). Zum mindesten im nördlichen Mitteleuropa

sind für den Aufbau der Sommerpopulationen also im wesentlichen Zuwanderer aus Süden verant-

wortlich. — Die blattlausfressenden Larven von S. selenitica wurden auf Fichten und Kiefern gefun-

den (Dixon 1960, Kula 1980), was die leichte Präferenz der Art für Nadelwälder erklären könnte.

Wanderungen: Am Col de Bretolet wurde selenitica recht zahlreich gefangen (Tab. 2) und von Au-

bert et al. (1976) als vermutliche Wanderart eingestuft („espece consider&e migratrice“). Die Vertei-

lung zeigt von Anfang bis zum Ende der Erfassungszeit ein nahezu gleichmäßiges Vorkommen ohne

Anzahl pro Pent EN

Anz. pro Tag (#) SS NE NEED

ze. 1975 - 1987

SZ | jEn. 49.37 S

| 5% 23.7 | i

£ | 25% 28.7 | 29.

24 |50% 4.8 |

IS 75% 40..18

95% 9.9 |

ER AA10

bennenvmenum> .n

1 L

[e:)

[e,]

Abb. 12: Nach Süden gerichtete Migration von Scaeva selenitica (vgl. Legende zu Abb. 4a).

Fig. 12: Southbound migration of Scaeva selenitica.

Maxima oder Minima. Auch an einer in Tirol beobachteten Schwebfliegenwanderung hatte die Artteil

(Jeekel & Overbeek 1968). Grey et al. (1953) sahen auch ın den Pyrenäen wandernde selenitica.

Aus Norddeutschland liegen Funde nicht bodenständiger Individuen von der Nordseeinsel Schar-

hörn vor (Schmid 1987). Weitere Hinweise auf Wanderungen sind Beobachtungen von Malec (1986),

daß die Art im Herbst bis in Stadtgärten komme (Umgebung von Kassel) und eigene Beobachtungen

aus Innenstädten (SW-Deutschland).

Phänologie am Randecker Maar: In den meisten Jahren werden nur wenige Individuen gefangen

(Abb. 27k). Dies dürfte einerseits auf die relative Seltenheit dieser Art zurückgehen, andererseits dar-

auf, daß diese großen und flugtüchtigen Schwebfliegen die Reuse oft als Hindernis erkennen und um-

fliegen (Gatter 1975). Selenitica erscheint überwiegend ım Juli und der ersten Augusthälfte (Abb. 12).

Dabei stellen QQ das Gros der Wanderer (FO:PY = 1:2,9;n = 149). In der S-Reuse werden deut-

lich mehr Tiere gefangen als in der N-Reuse (S-Reuse :N-Reuse = 5,4:1;n = 54).

Status: Saisonaler Migrant.

Parasyrphus lineolus (Zetterstedt 1843)

Verbreitung: holarktisch

Biologie: Diese Art kann zwischen Ende April und Anfang Oktober beobachtet werden. Am zahl-

reichsten ist sie im Hochsommer. Sie ist vor allem ın Wäldern und an Waldrändern zu finden. In den

Nordalpen ist lineolus nach eigenen Beobachtungen bis über die Baumgrenze verbreitet. Parasyrphus-

Larven leben überwiegend auf Bäumen (Rotheray & Gilbert 1989). Die aphidophagen Larven von

lineolus wurden in Fichtenwäldern gefunden (Kula 1980).

Wanderungen: Der einzige Hinweis auf herbstliche Wanderungen stammt aus den Alpen: Aubert

et al. (1976) fingen insgesamt 5746 Fx. (Tab. 2) und bezeichneten die Art, die zwischen Mitte Juli und

Mitte September in fast gleichmäßigen Zahlen gefangen wird, als Wanderer.

Phänologie am Randecker Maar: P. lineolus fiel lediglich in zwei Jahren auf: 1980 wurden 265 Ex.

in der S-Reuse und 16 Ex. in der N-Reuse, 1982 5 Ex. in der S-Reuse gefangen. Es ist möglich, daß

die Art in einigen anderen Jahren ebenfalls mit wenigen Stücken vorkam, aber übersehen wurde. In

den zwei Jahren, in denen lineolus festgestellt wurde, erschien die Art während eines eng begrenzten

Zeitraums Ende Juli (Abb. 13a). Ihr Aktivitätsmaximum hat sie während der Mittagsstunden

(Abb. 13b). PP überwiegen deutlich (I: PP = 1:3,0;n = 242).

Status: Saisonaler Dismigrant (Saisonaler Migrant ?).

Wahrscheinlich ist, daß die im allgemeinen eher spärlich vorkommende Art nur im Jahr 1980 hohe

Populationen aufbauen konnte und in der Folge als Wanderer in Erscheinung trat. Die Abundanzen

Anzahl pro Pent.

Anz. pro Tag (+) Zzimeolus

75 =

246) 188 1987 u

105+| &.80.7 i ge ee

5% 23.7. el

25% 25.7. 5.7.90 7

180 | 50% 28.7. 1.8.8 24

90+| 75% 1.8.

95% 3.8

IS: GE).

/1ImMeolus

vanre 1975 — 1987

Finn

a Jun |

Abb. 13a: Nach Süden gerichtete Migration von Parasyrphus lineolus (vgl. Legende zu Abb. 4a) (eingefügte Zeich-

nung aus van der Goot 1981).

Fig. 13a: Southbound migration of Parasyrphus lineolus.

Abb. 13b: Tageszeitliche Verteilung von Parasyrphus lineolus (vgl. Legende zu Abb. 4d).

Fig. 13b: Diurnal activity patterns of Parasyrphus lineolus migrating southward.

einer Schwebfliegenart können regional sehr unterschiedlich sein. Möglicherweise sind solche räumlıi-

chen Disparitäten für die ungewöhnlich starke Konzentration auf die S-Reuse verantwortlich.

Um den Status von P. lineolus zu klären, sind weitere Beobachtungen notwendig.

Meliscaeva cinctella (Zetterstedt 1843)

Verbreitung: holarktisch, orientalisch

Biologie: M. cinctella fliegt in vielen Lebensräumen, bevorzugt aber Wälder und waldnahe Gebiete.

Sie kann von (Mitte) Ende April bis Mitte Oktober beobachtet werden. Die größte Häufigkeit wird

im Spätsommer erreicht. Die aphıdophagen Larven von cinctella wurden auf Laub- und Nadelbäumen

gefunden (Eiche, Apfelbaum, Holunder, Kiefer, Fichte; Rotheray & Gilbert 1989). Wie die Art über-

wintert, scheint noch unklar zu sein. Die nahe verwandte Meliscaeva auricollis (vgl. 3.2.1) überwintert

als Imago (Schneider 1948).

Wanderungen: Herbstliche Wanderungen wurden bisher nur am Col de Bretolet festgestellt (Au-

bert et al. 1976). Obwohl die Art dort nicht sehr zahlreich erscheint (Tab. 2), wurde sie als Wanderer

eingestuft. Sie weist ein sehr deutliches Maximum in der ersten Septemberhälfte auf.

Phänologie am Randecker Maar: M. cinctella ist in den meisten Jahren eher spärlich (Tab. 2). Nur

im Jahr 1980 war sie mit 172 gefangenen Exemplaren zahlreicher. Starke Bestandsschwankungen sind

bei dieser Art, wie bei anderen aphidophagen Schwebfliegen, auch andernorts beobachtet worden

(z. B. Malec 1986).

M. cinctella tritt hauptsächlich im August auf (Abb. 14). Im September werden nur noch wenige

Tiere gefangen. In der S-Reuse wurden wesentlich mehr cinctella gefangen (S-Reuse:N-Reuse =

3,5:1;n = 258). Q2 dominieren (O:P9 = 1:3,8; n = 67).

Status: Saisonaler Migrant (?).

Anzahl pro Pent 4

Anz. pro Tag (+) e2necetells

1975 1987

ERAhRET, D = 20

5% 30

05% 7

ee Le

75% 18

935% 2

LEm2A

oommm.

TER = Z—_

Aug Sep

Abb. 14: Nach Süden gerichtete Migration von Meliscaeva cinctella (vgl. Legende zu Abb. 4a).

Fig. 14: Southbound migration of Meliscaeva cinctella.

Phänologie, Geschlechterverhältnis und das häufige Vorkommen nur in einem Jahr (1980) ent-

spricht weitgehend den Beobachtungen bei der vermutlich saisonal dismigrierenden Art Parasyrphus

lineolus (S. 42). Der am RM und CB fast identische Verlauf der Fangkurven mit der für Saisonwande-

rer typischen Zeitversetzung von ca. 2—3 Wochen (vgl. 4.1) macht aber sehr wahrscheinlich, daß cinc-

tella ein Saisonaler Migrant und kein Dismigrant ist.

Episyrphus balteatus (De Geer 1776)

Verbreitung: paläarktisch, orientalisch, australisch

Biologie: Die eurytope Art E. balteatus ist eine der häufigsten Schwebfliegen. Sie kann in allen Mo-

naten beobachtet werden. Dusek & Läska (1974) wiesen nach, daß) weder Larven noch Puparien den

Winter ın Zentraleuropa überleben. Nur Imagines können hier überwintern. (Aufenthaltsorte über-

winternder @Q sind allerdings bis heute noch nicht gefunden worden.) An den ersten Frühjahrsblü-

hern sind in den Alpen (Goeldlin 1974) wie in den Mittelgebirgen (z. B. Malec 1986, Schmid 1986), im

nordwesteuropäischen Flachland (van der Goot 1981) und in England (Stubbs & Falk 1983) regelmä-

Rig auffallend dunkel gefärbte, befruchtete balteatus-Q Q zu sehen. Zum Aufbau der Ovarien ist, wie

Schneider (1948) nachwies, Pollenfraß nötig.

Nach den ersten Frühjahrsbeobachtungen folgt häufig eine Zeit, in der man kaum Imagines begeg-

net. Im Frühsommer ist balteatus dann zahlreich. Überaus häufig, oft geradezu massenhaft tritt die

Art dann im Hochsommer (ab Juli) bis in den Herbst hinein auf.

Die Larven leben auf den verschiedensten Pflanzen der Baum- und Krautschicht. Sie sınd auffällig

polyphag (Läska & Stary 1980). Neben ihrer Hauptnahrung, vielen verschiedenen Blattlausarten, er-

nähren sie sich auch von Käferlarven und Artgenossen (Goeldlin 1974). Die Larven entwickeln sich

stets ohne Diapause. So könnten, wenn zwischen einer Eiablage und der nächsten ca. 1!/2 Monate ver-

streichen, ungefähr 4—5 Generationen im Jahr gebildet werden, wie Schneider (1948) nach Laborver-

suchen herausfand. Allerdings werden im Freiland in der Regel nicht so viele Generationen gebildet.

Abb. 15a: Nach Süden gerichtete Migration von Episyrphus balteatus (vgl. Legende zu Abb. 4a); eingeschaltet ist

das Zugmuster am Col de Bretolet (aus Aubert et al. 1976).

Fig. 15a: Southbound migration of Episyrphus balteatus; the insertion shows the phenology at Col de Bretolet.

Anzahl pro Pent.

Anz.

8000

4000

7eo0®8

3600

8400

3200

5600

2800

4800

2400

4000

2000

3200

1600

2400

1200

1600

800

400

pro Tag (#)

1975 - 1987

er. 19, 8.

5% 2 dd,

2a . d,

SR dead.

Br 7% 7a

5% Sl, O,

Le, A4,.ı10,

Jun

SI eBeradeus

(e0) I] (00) (60) (co) (a0) (a0) {oo) (09) III (eo)

» 48

„ 48

> HU

. 80

. al

. 84

‚&8

. 86

Episyrphus balteatus 9 (Ph. W. Gatter).

Rotheray (1989) gibt für Großbritannien lediglich eine oder zwei Generationen pro Jahr an, Krüger

(1926) für Norddeutschland „mindestens zwei Generationen“.

Wanderungen: Episyrphus balteatus ist einer der auffälligsten Wanderer unter den Schwebfliegen.

Viele Einzelveröffentlichungen über Schwebfliegenwanderungen gehen auf Massenzüge dieser Art

zurück. Fast alle diese Beobachtungen betreffen spätsommerliche Bewegungen in südliche Richtun-

gen.

Direkte Beobachtungen von Nordwanderungen sind dagegen äußerst spärlich. Einen Hinweis auf

nordgerichtete Bewegungen über einen Alpenpaß Ende Mai verdanken wir L. Verlinden (in litt.,

27.6.1989): bei einem Aufenthalt im südtiroler Hochpustertal im Jahr 1985 waren bis zum 22.5. nur

dunkel gefärbte PP zu sehen (s. 0.). Am 23.5. (ca. 5°C, wolkig, nur gelegentlich sonnig, stärkerer

NW-Wind) waren in der Nähe des Kreuzbergpasses (Alpe Nemes, 1850 m NN) erstmals auch CC

zu beobachten, die gemeinsam mit QQ im Windschatten von Weidenbüschen schwebten. Von Zeit zu

Zeit, offensichtlich vor allem dann, wenn die Sonne kurzzeitig erschien, flogen kleine Gruppen gegen

den Wind, also in nördliche Richtung, ab. Von S$ her erschienen immer wieder neue Gruppen. Inner-

halb kurzer Zeit zogen so mehrere hundert balteatus durch. Schlechtes Wetter unterbrach diese Beob-

achtung wenig später. Das Beobachtungsgebiet liegt im Norden des Talsystems der Piave, das ziem-

lich direkt nach $ ins italienische Alpenvorland geöffnet ist.

An einem 3600 m hohen Hıimalayapaß in Nepal war im März Nordwanderung von Dipteren zu be-

obachten. Unter den Belegexemplaren waren auch zwei balteatus-QQ (Westmacott & Williams

1954). Ebenfalls im Himalaya sah Gatter (1980) Dipteren im Mai über einen 5400 m hohen Paß nach

Norden wandern. Auch an diesen Wanderungen hatte balteatus teil. Weitere Hinweise auf Nordwan-

derungen lassen sich aus faunistischen Daten schließen: im nördlichen Europa, wo Überwinterungen

der Art nie festgestellt wurden (Torp 1984), liegen die frühesten Beobachtungen am 8.6. in Schleswig

(Claußen 1980), am 4.6. in SE-Jütland (Torp 1981) und am 18.6. in S-Norwegen (Nielsen 1971). Bal-

teatus wandert also im Laufe des Juni dort ein.

Anzahl pro Pent

Anz. pro Tag (+)

4000 1975 - 1987 Pu

2000 + En. N). G- [) 1175 |

5% 26 -

05% 3

3600 | 50% 7

1800 + | 76% 17

95% 29.

Le 240) | 21

3200 1

1500 + | 3),

mmmmn

Anzahl pro Pent

Anz. pro Tag (+)

2800 1975 - 1987

oo R. B= 1085

| 12 J= 13031

oo onmommmnAao

[e}

Abb. 15b: Nach Süden gerichtete Migration von Episyrphus balteatus-J'O (vgl. Legende zu Abb. 4a).

Fig. 15b: Migration of OO of Episyrphus balteatus in southerly direction.

Abb. 15c: Nach Süden gerichtete Migration von Episyrphus balteatus-QQ (vgl. Legende zu Abb. 4a).

Fig. 15c: Southbound migration of PP of Episyrphus balteatus.

Aber auch die mitteleuropäischen Populationen werden wohl alljährlich in großem Ausmaß von

Zuwanderern gestärkt. Verlinden & Decleer (1987) vermuten, daß der Frühsommerbestand in Belgien

von ansässigen balteatus aufgebaut wird, während die gewaltigen balteatus-Massen, die im Hochsom-

mer zu sehen sind, Folge von Einwanderungen aus dem Süden sind. Die Einwanderungen finden da-

bei allerdings nicht im Hochsommer statt — sämtliche beobachteten Wanderungen in dieser Zeit füh-

ren in südliche Richtungen — sondern im Frühjahr. Für das Populationsmaximum sind also nicht die

Einwanderer direkt, sondern deren Nachfolgegeneration verantwortlich.

Südwandernde balteatus wurden dagegen schon an zahlreichen Plätzen beobachtet. Einige dieser

Beobachtungen sollen hier kurz wiedergegeben werden, ohne Vollständigkeit anzustreben.

England:

— In Südengland erreichten zahlreiche balteatus gemeinsam mit anderen Arten Anfang August 1928 die Isle of

Wight über See aus nordöstlicher Richtung (Flugrichtung also SW) (Mackworth-Praed 1929). Am selben Ort

hatte schon Walker (1864) Mitte August große balteatus-Schwärme beobachtet. Anfang August 1960 wurde eine

aus SE kommende, z. T. von balteatus bestrittene Schwebfliegenwanderung in Sussex bemerkt (Spreadbury

1960, Parmenter 1960).

Nördliches Mitteleuropa:

— Feuerschiffe: Von 1388 während drei Wochen im Juli/August 1963 mit Gelbschalen gefangenen Schwebfliegen

gehörten 507 (255 30,252 Q 9) zu balteatus. Nach Metasyrphus corollae war balteatus damit die häufigste Art.

Anzahl pro Pent Anzahl pro Pent

JERZIEN EN ZHeKENSE

Anz. pro Tag (+] Anz. pro Tag (#)

5 1975 1979

1 00 “ir vw, [|

SH ee Er. 16.7 n = 2654

: + 7%

En 530.7 5% 30.7

1050 | -

S25+ 1 2% 3.8 25% 14. 8 21.8.= 434

50% 4.8 50% 17.8 14.8.- 25

600 | 75 8 75% 21.8 17.8.=- 1984

300 5X 16.8 9% 2.9 15. 8.= 198

Sc Le, 16. 9 )

= |

Jun Ju Aug Sep Se Jul Aug Sep Okt

He 1976

1500 _ Ei - 1500 Re 1980 = ß 4

so + =, 2

=> Sep. Aalı, D Se | Er. 17.7 n= 1323

: | -47%

| ax 16, 7 il 1050 | 5%.26.7 |

Hl os ı7. 7 in ses+ [25% 31,7 1.8.= 160

50% 18.7 16.7.= 47| I Ei 3. B.= 440

60 L7% 24.7 20.7.= 20 SSe L 75% 16.8 AUCH 97

300+| go 9.8 ne ke See Er | 26.7.= 8

Sc BasPSEaB | 150 [1 193.9

Er ı 1 7 ]

F ae San Lee

Jun Jul Aug Sep Okt Jun Jul Aug 1 Sep Okt

50 +

1050

525 +

600

300 +

so 1978 E n , el 8 vw 7

"| Er. 46.7 R BEE SS ER n= 14587 |

+37% | a -36% |

| 29 |

+ 4 I}

3 |

8600 B |

300 + 3 |

50 |

2 h

[1]

10]

[eo]

d; Jun Jul Aug Sep Okt d> 5

Anzahl pro Pent

Anz. pro Tag (+) 7

1500

750 +

1050

Sees

800

300 +

150

[e)

300 +

Jun ya) Aug | Sep Okt

oommm

d Jun Jul Aug Sep Okt

baltealcus balteatus M

vahre 1975 - 1977 vshre 1975 —- 1977

25%= 145

Max= 81

>16h= 12.4 %

Ne= 9

* 25%= 8300 * 25%= 3824

* 50%= 7649

* 75%= 11473

Max= 2929

* N = 15298

N = 13606

unr

baltestus W

VIREN VID ZEIT TE

* 25%= 3555

* 50%= 7109

* 75%= 10664

Max= 2307

* N = 14218

Dane

4 6 8 10 12 14 16 4B 20 Unr

Abb. 15e: Tageszeitliche Verteilung von Episyrphus balteatus (vgl. Legende zu Abb. 4d).

Fig. 15e: Diurnal activity patterns of Episyrphus balteatus migrating southward.

Abb. 15f: Tageszeitliche Verteilung von Episyrphus balteatus-O'J (vgl. Legende zu Abb. 4d).

Fig. 15f: Diurnal activity patterns of O'C of Episyrphus balteatus migrating southward.

Abb. 15g: Tageszeitliche Verteilung von Episyrphus balteatus-PP (vgl. Legende zu Abb. 4d).

Fig. 15g: Diurnal activity patterns of PP of Episyrphus balteatus migrating southward.

504 Expl. wurden dabei auf den zwei in der Ostsee liegenden Feuerschiffen „Kiel“ und „Fehmarn-Belt“ (Kü-

stenabstand 17-25 km) gefangen, nur 3 Expl. auf dem 30 km vor der Küste liegenden Nordsee-Feuerschiff „EI-

be I“ (Heydemann 1967). Dieser Unterschied dürfte darauf zurückgehen, daß balteatus, von Skandinavien

kommend, in sehr großer Zahl über die Ostsee wandert, während die offene Deutsche Bucht gewöhnlich wohl

nur selten in großer Zahl überflogen wird.

Auf dem zwischen England und den Niederlanden in ca. 70 km Küstenabstand liegenden Feuerschiff „Noord

Hinder“ konnte balteatus ebenfalls an einigen Tagen vor allem Ende Juli/Anfang August in größerer Anzahl ge-

fangen werden (Lempke 1962).

— Insel Scharhörn: Bei Planbeobachtungen im Jahr 1984 auf der 15 km vor der Küste im inneren Winkel der Deut-

schen Bucht gelegenen Insel, die keine bodenständige Schwebfliegenfauna aufweist, war balteatus zwischen

a rn ee

Abb. 15d: Nach Süden gerichtete Migration von Episyrphus balteatus in den Jahren 1975-1987 (vgl. Legende zu

Abb. 5b).

Fig. 15d: Southbound migration of Episyrphus balteatus in 1975— 1987.

B3altegstus balteatus M balteatus W

anre 1975 - 1987 Vanre 1975 - 1977 vanre 1975 - 1987 vanre 1975 - 1987

ı0 ı2 14 16

AI RreBe:

18 20 Unr 4 6 8 10 ı2 14 16 18 20 Unr 4 6 8 10 12 14 16 18 20 Unr 4

ee) era ee ee 6 8 10 ı2 14 16 18 20 Unr

NOV

Max= 9 Max= 27

y m

15.10 N= 29 | 15.10. N= 105

N NS DIN .

Max= 35

be NENNNN a 30.9 NR N N= 128

Max= 2

>16h= 18.2 % oz

N= 11 15. 9

Max= 101 Max= 129

N= 485 15.9 N \ N= 628

Max= 31

Max= 706 Max= 601

"Ne 7175 16 >46h= 11.4 % 16.- 16 “N= 3110

N= 6441 31.8 N= 158 31. 8 N= 3186 31.8 } IS N= 2837

DR NONE

Max= 3302 Max= 226 x] Max= 1821 Max= 1297

"N= 19229 1.- >16h- 18.5 % 1.- | "N= 8572 1.- - "N= 7588

N= 16229 15.8 Ne 1282 15.8 N N= 7913 15 IN N= 6931

JUL Max= 10 JUL Max= 38 JUL Max= 287

16.- Sl >16he 17.0 % 16.- "N= 2261 16.- 4 "N= 2232

31.7. [r ZNOAN Ne 711 31. 7. [RN = SON N= 1959 31.7 N= 1865

r 1 j r En | tr r Talstal

4 6 B 10 ı2 14 16 18 20 Unr 4 6 8 10 12 14 16 18 20 Uhr 4 6 8 10 12 14 16 :B 20 Unr

i k

Abb. 15h: Tageszeitliche Verteilung von Episyrphus balteatus im Jahresverlauf (vgl. Legende zu Abb. 5h).

Fig. 15h: Diurnal activity patterns of Episyrphus balteatus migrating southward throughout the year.

Abb. 151, 15k: Tageszeitliche Verteilung von Episyrphus balteatus-J'O' (151, links) und-QQ (15k, rechts) im Jah-

resverlauf (vgl. Legende zu Abb. 5h).

Fig. 151, 15k: Diurnal activity patterns of JO (151) and PP (15k) of Episyrphus balteatus migrating southward

throughout the year.

Ende Juli und Ende August an vielen Tagen oft zu Tausenden zu sehen. Gelegentlich konnten auch über dem

Watt nach $ fliegende Tiere beobachtet werden. Massenwanderungen fanden immer an warmen Tagen mit ab-

landigen (also aus NE, E, SE oder $ wehenden), schwachen oder mäßigen Winden statt (Schmid 1987). Während

desselben Jahres auf der Insel Mellum aus Farbschalenfängen gewonnene Daten (Barkemeyer 1988) zeigen die-

selben Häufigkeitsmaxima und dokumentieren damit den überregionalen Charakter der auf Scharhörn beob-

achteten Schwebfliegenwanderungen.

— Nördlicher Mittelgebirgsrand: Auf eine südwestwärts gerichtete Massenwanderung von E. balteatus bei Biele-

feld vom 7.—10. August weist Eitschberger (1973) hin. Meineke (1979) beobachtete am südwestlichen Rand des

Harzes Anfang September starken, ebenfalls nach SW gerichteten Zug.

Alpen:

— Am Col de Bretolet in den Walliser Alpen ist balteatus mit Abstand die häufigste Art (Aubert et al. 1976). Sie

stellt über die Hälfte aller Fänge (Tab. 2). Der Hauptdurchzugsmonat ist der August; der Zug setzt sich im Sep-

tember aber mit beachtlicher Stärke fort und reicht bis zum Ende der Beobachtungszeit im Oktober (Abb. 15a).

Dabei werden fast nur Weibchen gefangen. Eine Stichprobe ergab 93,4% @Q und nur 6,6% OO’ (15 Fangtage

zwischen 7.8. und 27.10.,n = 1569; Aubert 1962) (vgl. 4.2).

Von balteatus liegen auch einige Fernfunde markierter Individuen vor (Aubert et al. 1969, Aubert & Goeldlin

1981): die maximale Flugstrecke eines Individuums lag dabei bei 111 km (Col de Bretolet bis Col du Glandon).

Durch Fang- und Markierungsexperimente an mehreren in südwestlicher Richtung aufeinanderfolgenden Al-

penpässen konnte der Zug von balteatus (und anderen Arten) über ca. 160 km verfolgt werden. Dabei wird auch

das tief eingeschnittene Rhönetal gequert.

— An mehreren anderen Stellen der schweizer und österreichischen Alpen wurden südgerichtete Schwebfliegen-

migrationen beobachtet, bei denen gewöhnlich Episyrphus balteatus, im Herbst auch Eristalis tenax, die Haupt-

rolle spielte (z. B. Burmann 1978, Dethier & Goeldlin 1981, Gepp 1975, Harz 1965, Huss 1975, Jeekel & Over-

beek 1968, Prell 1925). Häufig gelangen diese Beobachtungen in Pässen. An „starken Zugtagen“ werden nach

Dethier & Goeldlin (1981) aber auch die höchsten Berggipfel überflogen. Zu auffälligen Massierungen an Pässen

kommt es insbesondere bei starkem Föhn; dann wandern die Schwebtliegen in großer Bodennähe gegen den

starken Wind und stauen sich an den Pässen (vgl. 3.4). Nach Burmann (1978) und Jeekel & Overbeek (1968) do-

minieren wie am CB Q9 stark. Reusenfänge im schweizer Jura zeigen keine Unterschiede gegenüber den Ver-

hältnissen in den Alpen (Aubert & Jaccard 1981).

Pyrenäen:

Am Paß Port de Gavarnie, auf der französisch-spanischen Grenze in über 2000 m (7500 ft.) gelegen, beobachte-

ten Lack & Lack (1951) am 13.10.1950 eine SW-Wanderung von Schwebfliegen. Die Belegexemplare gehörten zu

Episyrphus balteatus. Am selben Ort gelang Snow & Ross (1952) am 20.9.1951 folgende Beobachtung: bei Wind-

stille war zwar Schmetterlingswanderung nach SW, aber keine Schwebfliegenmigration festzustellen. Am Nach-

mittag begann ein allmählich stärker werdender Südwind zu wehen. Gleichzeitig setzte ein ebenfalls stärker wer-

dender Schwebfliegenzug ein. Die Fliegen zogen dabei in großer Bodennähe unterhalb 60 cm (vgl. 3.4). Am folgen-

den Tag hielten Wind und Schwebfliegenwanderungen an. Balteatus gehörte zu den häufigsten Arten. Williams et

al. (1956) beobachteten am 9. und 10. Oktober ebenfalls an diesem Paß Schwebfliegenwanderungen, an denen ne-

ben Eristalis tenax v. a. Episyrphus balteatus teilnahm. Auch Grey et al. (1953) sahen in den nördlichen Pyrenäen

Ende September/Anfang Oktober Schwebfliegenzug, der überwiegend von den Arten E. balteatus und Eristalis

tenax bestritten wurde.

Phänologie am Randecker Maar: Mit knapp 33 % der Fänge ist E. balteatus auch am Maar die häu-

figste Schwebfliegenart (Tab. 2). Sie wandert hauptsächlich im August (Abb. 15a) mitSchwerpunkt ın

der ersten Monatshälfte. Der Median liegt in der Hauptzugzeit am 7. August. Der steile Anstieg der

Fangzahlen Ende Juli geht z. T. darauf zurück, daß die Station den Betrieb in einigen Jahren erst in der

letzten Julidekade aufnahm. Das schnelle Abflauen des Fanges Ende August liegt jedoch voll in der

Beobachtungsperiode. Im September und Oktober erscheint balteatus mit 5% der Gesamtsummen

nur noch spärlich.

Beide Geschlechter sind am Maar ungefähr gleich stark vertreten (TO':PP = 1:0,92; n = 27135).

Die SC sind etwas stärker auf den Beginn der Zugzeit konzentriert (Abb. 15b, 15c).

Die Fangergebnisse weisen von Jahr zu Jahr starke Schwankungen auf (Abb. 271). Trotz dieser ho-

hen Variabilität lassen die jährlichen Durchzugsdiagramme (Abb. 15d) das Grundmuster der zusam-

mengefaßten Daten (Abb. 15a) fast immer erkennen. Nur in seltenen Fällen liegt der Medianwert au-

ßerhalb der ersten Augusthältte.

Das tageszeitliche Muster zeigt einen für balteatus typischen, ausgeprägten Vormittagsgipfel mit

einem Aktivitätsmaximum zwischen 8 und 11 Uhr (Abb. 15e). Bei OO’ ist diese einseitige Verteilung

sogar noch etwas extremer als bei 9 (Abb. 15f, 15g). Die Form der Aktivitätskurve bleibt im Jahres-

verlauf im wesentlichen erhalten (Abb. 15h). Nur der Medianwert, der im Juli noch vor zehn Uhr

liegt, verschiebt sich bis Mitte September in das Zeitintervall 10-11 Uhr und bis Mitte Oktober in das

Intervall 11-12 Uhr. Auch aus diesen Darstellungen wird ersichtlich, daß der Aktivitätsgipfel der

JO stets deutlich vor dem der QQ liegt (Abb. 151, 15k).

Mit Hilfe von Gelbschalenfängen ermittelte Grosser (1979) für balteatus ein sehr deutliches Akuvi-

tätsmaximum in den ersten zwei Stunden nach Sonnenaufgang (5-7 Uhr). In dieser Zeit widmet sich

balteatus also hauptsächlich der Nahrungsaufnahme. Erst nach dieser frühmorgendlichen Phase des

Nahrungserwerbs setzt die Migration ein.

In der N-Reuse wird balteatus nur sehr spärlich gefangen (Abb. 271): S-Reuse : N-Reuse = 7,4:1

(n = 8902). Es dürfte sich dabei um Tiere handeln, die in der Umgebung Nahrungsflüge unternehmen.

In die S-Reuse geraten die Fliegen vor allem bei Gegenwind, fast nie bei Rückenwind (Abb. 26d;

vgl. 3.4):79% werden bei Winden ausSW, S oder SE gefangen, nur 0,6 % bei Winden aus NW, N oder

INIE®

Status: Saisonaler Migrant.

Im südwestdeutschen Mittelgebirge zeigen die sehr starken quantitativen Schwankungen der im

Frühjahr beobachteten Q® (Schmid 1986 und i. Dr.), daß die Überwinterung hier mit großem klima-

tischem Risiko behaftet ist. In den Alpen dürften die Verhältnisse nicht wesentlich anders sein. Erst

im südlichen (mediterranen) Europa oder Nordafrika sind Bedingungen zu erwarten, die eine unge-

fährdetere Überwinterung gestatten. Hierhin dürften die spätsommerlichen Wanderungen von bal-

teatus führen. Allerdings sind die Lebensverhältnisse im Mittelmeergebiet im Sommer eher ungünstig.

Die dann weitgehend vertrocknete Krautflora steht als Blattlausnahrung nicht zur Verfügung. Die re-

lativ wenigen Blüten der sommerlichen Mittelmeerflora sind für die meist kurzrüsseligen Schwebflie-

gen überdies oft nicht leicht zugänglich. Wesentlich günstigere Verhältnisse findet balteatus im Som-

mer dagegen im humiden Mitteleuropa, wo für Larven ein großes Blattlaus- und für Imagines ein

reichhaltiges Blütenangebot zur Verfügung steht. Im Frühjahr wandert die Art deshalb nach Mittel-

europa ein.

Das hier skizzierte Bild des biologischen Hintergrundes der saisonalen Wanderungen von balteatus

ist aus wenigen Mosaiksteinen zusammengesetzt und hat deshalb noch weitgehend hypothetischen

Charakter.

Zur Überprüfung der Hypothese sind selbst auf den ersten Blick wenig aussagekräftige Beobach-

tungen wertvoll. So sah z. B. Gatter (unveröff.) im August in Tälern der nördlichen Appenninen und

der Pyrenäen auf ca. 1000-1800 m NN praktisch keine Syrphiden der bekannten und häufigen Wan-

derarten. Sie hatten diese von „einheimischen Populationen“ im Sommer freien Gebiete auf ihrer

herbstlichen Südwanderung vermutlich noch nicht erreicht.

Sphaerophoria scripta (Linnaeus 1758)

Verbreitung: paläarktisch, orıentalisch (Nordindien, Nepal)

Biologie: $. scripta ist eine sehr häufige und eurytope Art, die zwischen (Mitte März) Mitte April

und Mitte (Ende) Oktober beobachtet wird. Sıe bildet ın dieser Zeit mehrere Generationen (in Mittel-

deutschland nach Grosser & Klapperstück [1976] drei Generationen). Am zahlreichsten kommt die

Art im Hoch- und Spätsommer vor. In Belgien entspricht die Häufigkeitskurve von scripta weitge-

hend der von Episyrphus balteatus mit mäßig hohen Zahlen im Frühsommer und einem Maximum,

das Mitte Juli sehr abrupt beginnt und Anfang September fast ebenso abrupt endet (Verlinden &

Decleer 1987).

Scripta-Larven leben überwiegend in der Krautschicht, wo sie sich von Blattläusen ernähren (Ban-

kowska 1964). Nach Dusek & Läska (1974) überwintert scripta als Larve. Beweise für die von Ban-

kowska (1964) vermutete Überwinterung weiblicher Imagines fehlen.

Wanderungen: Nordgerichtete Frühjahrswanderungen von Dipteren, an denen auch S. scripta teil-

nahm, beobachtete Gatter (1980) an einem in 5400 m Höhe gelegenen Himalayapaß im Mai. Im Som-

mer und Herbst ist scripta ein auffälliger und zahlreicher Süd-Wanderer. Aus den Alpen liegen u. a.

Beobachtungen von Aubert et al. (1976), Gepp (1975) und Jeekel & Overbeek (1968) vor. Das Phäno-

logie-Diagramm bei Aubert et al. zeigt, nach steilem Anstieg ab Mitte Juli, einen ausgeprägten Gipfel

in der ersten Augusthälfte und ein rasches Abnehmen der Fangzahlen in der zweiten Augusthälfte.

Zwei am Krinnenpaß in der Schweiz markierte Individuen wurden am ca. 50 km südwestlich liegen-

den Col de Bretolet wiedergefangen (Aubert & Goeldlin 1981).

In Norddeutschland wurden nicht bodenständige Tiere dieser Art in einigen Exemplaren auf Feu-

erschiffen (Heydemann 1967, Lempke 1962) und in großer Zahl im Juli und August auf der Insel

Sphaerophoria scripta @ (Ph. W. Gatter).

Anzahl pro Pent.

Anz. pro Tag (#) Deren

3000 1975 - 1987

1500+[ N. ei

Eee BD = 988

5% 28.7 12 J= 11856

2544785

2700 40.8979 193

en 8. IT

73% ei. B. 2. 8,9% 45

95% 4.9. 248.78 62

ber 28210% el 8. 79 150

2400 29. 8.80 65

1200 + 7. 8.81 69

1982.82 69

3. 8.84 234

147. 8.285 332

2100

nenn 15. 8.86 245

222887 147

ı800 |

900 +

1500 |

750 +

2200 |

8600 +

900

450 +

600

300 +|

acer |.

150 +

Jul Aug Sep Okt

a Jun

Abb. 16a: Nach Süden gerichtete Migration von Sphaerophoria scripta (vgl. Legende zu Abb. 4a) (eingefügte

Zeichnung aus van der Goot 1981).

Fig. 16a: Southbound migration of Sphaerophoria scripta.

Scharhörn (Schmid 1987) gefunden. Hinweise auf Wanderungen in S-England liegen ebenfalls vor

(Parmenter 1960).

Phänologie am Randecker Maar: $. scripta ist wie am Col de Bretolet eine der häufigsten Arten

(Tab. 2). Sie wird auch am RM hauptsächlich im August gefangen (Abb. 16a). Im September erschei-

nen nur noch wenige Individuen. P treten am Maar fast doppelt so häufig auf wie FO (CO: PP

= 1:1,7; n = 11204). Im jahreszeitlichen Muster ihres Vorkommens besteht kein Unterschied

(Abb. 16b, 16c).

Auch das tageszeitliche Aktivitätsmuster der beiden Geschlechter ähnelt sich stark. Insgesamt ist

die Verteilung nahezu symmetrisch mit einem Maximum in der Mittagszeit (Abb. 16d).

Anzahl pro Pent. Anzahl pro Pent. 4 j

Anz. pro Tag (+) SERrRZABES WM Anz. pro Tag (+) sense A

1000 1975 - 1987 1800 1975 - 1987

vv I vo

SIE 7. am RO ]

Us= 4136

5% 29. 7. = 5% 28.

25% 4.8. 4.8.75 108 25% 5.

S00 | 50% 14. 8. Ä 1520 | 50% 14.

2450 + Ava / 076 19 BI0

75% 20.8. 20.,8.77. 22 75% 21.

95% 37.9.. 15, 8.78 18 95% 5.

Le. 4.10. | 21.8.79 63 to, 0,

soo | 29. 8.80 24 | 1440 |

400 + | 7.8.8 21 720 +

rm 15. 8.82 25

| 3.8.84 114

14. 8.85 90

700 | 1260

350+| 17 ..8..186 83 | 8s30+|

| 22. 8.87 51

I oso |

540 +

900

720

360 +

540

270 +

360 |

180 +

180 |

90 +

C Jun

Abb. 16b: Nach Süden gerichtete Migration von Sphaerophoria scripta-J'O (vgl. Legende zu Abb. 4a).

Fig. 16b: Southbound migration of Q’C’ of Sphaerophoria seripta.

Abb. 16c: Nach Süden gerichtete Migration von Sphaerophoria scripta-? Q (vgl. Legende zu Abb. 4a) (eingefügte

Zeichnung aus van der Goot 1981).

Fig. 16c: Southbound migration of PP of Sphaerophoria scripta.

Das Verhältnis zwischen S-Reuse und N-Reuse liegt bei 3,5:1 (n = 2696). Eine Auswertung der

Windrichtungen an den Hauptflugtagen zeigt, daß scripta überwiegend bei Gegenwind aus SW, S und

SE (66,9%) und nur selten bei Rückenwind aus NW, N und NE (17,7%) ın die S-Reuse fliegt (Abb.

27m).

Status: Saisonaler Migrant.

„Sphaerophoria menthastri-Gruppe“

Unter diesem Namen werden mehrere Arten zusammengefaßt, die zu Beginn der Untersuchungen

mit den Bestimmungsschlüsseln von Sack (1932) nicht unterschieden werden konnten. Auch heute

sind oft nur die G'C° dieser Artengruppe genau bestimmbar. Überwiegend werden aber Weibchen ge-

fangen. Die in der ersten Hälfte unseres Jahrhunderts sämtlich als Sphaerophoria menthastri (Linnaeus

1758) bezeichneten Tiere dieser Gruppe erweisen sich als zahlreichen verschiedenen Arten zugehörig.

Erst jüngst beschrieb Goeldlin (1989) vier neue Arten aus der westlichen Paläarktis.

Die vorliegenden Belegexemplare lassen vermuten, daß S. menthastri (Linnaeus 1758) — nach

Goeldlin (1989) ist der gültige Name $. interrupta (Fabricius 1805) — am Randecker Maar die häufig-

ste Art der Gruppe ist. Außerdem wurden $. fatarum Goeldlin 1989 — in Schmid & Gatter (1988) un-

ter dem Namen abbreviata Zetterstedt 1859 aufgeführt — und S$. taeniata (Meigen 1822) nachgewie-

sen. Insgesamt sind Fliegen dieser Gruppe aber eher selten. Sie werden hauptsächlich im August ge-

GERNE.

Jahre 1979 — 1977

25%= 485

50%= 970

75%= 1454

Max= 335

>16h= 5.8 %

N = 1939

Unr

Varıze 1970,19

* 25%= 3045

* 50%= 6090

* 75%= 9135

Max= 2027

*N = 12180

N = 415415

Unr

seripta M SCrIpPt3 W

vehre 1975 - 1977 VEanrEe 1333 1977:

vahre 1975 — 1987

* 25%= 1794

* 50%= 3589

* 75%= 5383

Max= 1232

*N = 7178

N = 6876

Abb. 16d: Tageszeitliche Verteilung von Sphaerophoria scripta (vgl. Legende zu Abb. 4d).

Fig. 16d: Diurnal activity patterns of Sphaerophoria scripta migrating southward.

fangen. Ihre geringe Zahl läßt keinen Schluß auf mögliches Wanderverhalten zu. Auch von anderen

Gebieten fehlen Hinweise auf zahlenstärkere Wanderungen weitgehend. Goeldlin (1989) weist darauf

hin, daß die in Speziation befindliche „menthastri-Gruppe“ auch intraspezifisch sehr varıabel ist,

während die wandernde Art Sphaerophoria scripta im gesamten riesigen Verbreitungsgebiet bemer-

kenswert wenig variabel ist — ein weiterer Hinweis auf die sehr geringe Mobilität der „menthastri-

Gruppe“ (vgl. auch 4.5).

3.1.2 Schwebfliegen mit terrestrischen saprophagen Larven

Rhingia campestris Meigen 1822

Verbreitung: paläarktisch

Biologie: Die Larven von Rhingia campestris entwickeln sich in Kuhdung (Krüger 1926, Coe 1942).

Die Imagines sind zwischen (Mitte März) Mitte April und Mitte Oktober (Anfang November) zu se-

hen. Sie fliegen in zwei deutlich ausgeprägten Generationen vor allem im Frühling und im Spätsom-

mer. Vermutlich überwintern diapausierende Larven (Coe 1942).

Wanderungen: Hinweise auf Wanderungen von R. campestris existieren nicht. Die am Col de Bre-

tolet hauptsächlich zwischen Anfang August und Mitte September gefangenen Tiere (Tab. 2) entstam-

men wohl ansässigen Populationen (Aubert et al. 1976).

Anzahl pro Pent.

Anz. pro Tag (#) SEINWDSSENZS

1975 - 1987

Eh Rn

Fasz. D = 58

a. T. a 700

Er e5% 1.8. 16. 8.75 5

L h 8. 77 |

Br : 5.8.76 ?

: Ad. El. 27 ?

Je 20. 9.7 6

g. 3.8.79 3

BO ||: 20. 8.80 100 |

so+| 14.8.8 4

12.8.8 2

28. 7.85 4

10 11.08.87 35

SO +

AO: |

40 IL

e0O +

a Jum Jul | Aug sep , Okt

Abb. 17a: Fänge in der S-Reuse von Rhingia campestris (vgl. Legende zu Abb. 4a) (eingefügte Zeichnung aus van

der Goot 1981).

Fig. 17a: Trapped individuals with northerly opening trap (S-trap) of Rhingia campestris.

C3MPESEFTIS

JENE NIT LGETE

Abb. 17b: Tageszeitliche Verteilung von Rhingia campestris (vgl. Legende zu Abb. 4d).

Fig. 17b: Diurnal activity patterns of Rhingia campestris.

Phänologie am Randecker Maar: Auch im Bereich der kleinen, gebüschbestandenen Kuppe, auf der

die Station Randecker Maar liegt, lebt eine Population von campestris, von deren Sommergeneration

sich regelmäßig einige Fliegen in den Reusen fangen (Tab. 2 und Abb. 17a). Da die 0°C’ oft stunden-

lang schwebend an Gebüschrändern und über der Reuse in der Luft stehen, während die PQ auf Blü-

ten der Krautschicht Nahrung suchen, sind 0'O’ in den Reusenfängen leicht überrepräsentiert

(CO: DD = 1m 307).

Abb. 17b zeigt im Tagesverlauf bis zum frühen Nachmittag eine weitgehend ausgeglichene Vertei-

lung. Das Flugaktivitätsmaximum liegt am späteren Nachmittag. Es unterscheidet sich damit grund-

sätzlich vom Aktivitätsmuster der Wanderarten mit ihrem Aktivitätsmaximum am späten Vormittag

oder um die Mittagszeit (Gatter 1981b).

Das starke Ungleichgewicht zwischen den beiden Reusen (S-Reuse : N-Reuse = 5,3:1;n = 498) läßt

sich dadurch erklären, daß die S-Reuse auf der Kuppe in einer Schneise zwischen Büschen steht — dort

schweben die campestris-O'O' bevorzugt und dort sind in der Krautschicht auch im Spätsommer zahl-

reiche Blüten zu finden, an denen die nahrungssuchenden @Q zu sehen sind — während sich die

N-Reuse direkt auf eine offene, im Spätsommer nahezu blütenlose Wirtschaftswiese öffnet.

Status: Bivoltine Art ohne Migrationsneigung.

Neoascia podagrica (Fabricius 1775)

Verbreitung: paläarktisch

Biologie: Diese kleine Schwebfliege besiedelt viele Lebensräume und ist sehr häufig. Ihre Larven le-

ben saprophag von pflanzlichen und tierischen Abfällen. Die Flugzeit beginnt (Ende März) Mitte

April und endet Mitte Oktober. Nach Hartley (1961) überwintern die Larven. Nielsen (1972) vermu-

tet allerdings, daß Aprilbeobachtungen i in S-Norwegen auf überwinternde Imagines zurückgehen.

Wanderungen: Der einzige Hinweis auf nicht bodenständige Individuen liegt von der Nordseeinsel

Scharhörn vor, wo an einem Tag 7 Tiere gefangen wurden (Schmid 1987).

Phänologie am Randecker Maar: Diese häufige Art wird regelmäßig, wenn auch nicht sehr zahl-

reich, in den Reusen gefangen (Tab. 2, Abb. 18). Vermutlich werden die sehr zarten Fliegen vom

Wind dorthin verdriftet. Dabei geraten wesentlich mehr Fliegen in die S-Reuse (S-Reuse :N-Reuse =

4,6:1;n = 124). Der Grund dürfte derselbe sein wie bei Rhingia campestris: N. podagrica lebt über-

Anzahl pro Pent.

Anz. pro Tag (#) DBoalgeogräee

1975 - 1987

IM V V v

Sp, 29. 2% D = 10

5% 3.8 BuJ= 1

25% 15.8. ae 5

e4 50% 2.9. ;

ler BR 9,8. i

95% 28.9. :

Le. 11.10. 3

18 8 7

9g9+ 5

12 IL

8 +

6 Io

Jun | Slam: Aug SIEIB Okt

Abb. 18: Fänge in der S-Reuse von Neoascia podagrica (vgl. Legende zu Abb. 4a) (eingefügte Zeichnung aus van

der Goot 1981).

Fig. 18: Trapped individuals with northerly opening trap (S-trap) of Neoascia podagrica.

]

wiegend im Bereich der einschürigen Wiese der Gebüschkuppe und wird deshalb überwiegend in der

S-Reuse gefangen.

Status: Polyvoltine Art mit geringer saıisonaler Dismigrationsneigung.

Syritta pipiens (Linnaeus 1758)

Vorkommen: holarktisch, orientalisch (Nepal)

Biologie: S. pipiens gehört zu den häufigsten Schwebfliegenarten. Sie ist zwischen (Ende März)

Mitte April und Mitte Oktober (Anfang November) in nahezu allen Lebensräumen zu beobachten.

Am häufigsten ist die Art im Hochsommer. Die Larven ernähren sich von pflanzlichen und tierischen

Abfällen wie Kompost, Dung usw. (Hartley 1961, Krüger 1926). Die Larven überwintern (Hartley

1961).

Wanderungen: Die einzigen Hinweise auf Wanderungen sind einige Funde (6 Expl.) auf Feuer-

schiffen (Heydemann 1967) und Beobachtungen (18 Expl.) auf der Nordseeinsel Scharhörn (Schmid

1987). Aus den Alpen fehlen Wanderbeobachtungen (vgl. Tab. 2).

Phänologie am Randecker Maar: Von dieser Art werden — verglichen mit ihrer est — nur

sehr wenige Tiere in den Reusen gefangen (Tab. 2). Sie entstammen mit großer Wahrscheinlichkeit der

ansässigen Population. In der Umgebung der Station, vor allem im 200 m südlich gelegenen Schopf-

locher Moor, ist die Art oft extrem häufig.

Status: Polyvoltine Art mit sehr geringer saisonaler Dismigrationsneigung.

3.1.3 Schwebfliegen mit aquatischen sapro-/microphagen Larven

Eristalis arbustorum (Linnaeus 1758)

Verbreitung: paläarktisch, orientalisch (Nordindien); nearktisch verschleppt

Biologie: Die eurytopen Imagines der häufigen E. arbustorum können zwischen (Anfang März)

Mitte April und Ende Oktober beobachtet werden, dürften also in mehreren Generationen auftreten.

Im Spätsommer ist die Art am häufigsten. Ansonsten können meist keine weiteren deutlichen Maxima

beobachtet werden. Die Larven werden gewöhnlich in stark eutrophierten Gewässern (Hartley 1961),

aber auch in Haufen faulender Pflanzen (Krüger 1926) gefunden. Die Larve überwintert (Hartley

1961, Krüger 1926).

Wanderungen: Am Col de Bretolet wird die Art, obwohl nicht allzu zahlreich gefangen (Tab. 2),

\ als Wanderer eingestuft (Aubert et al. 1976). Sie kommt dort fast während der ganzen Fangperiode

Anzahl pro Pent

Binz. pro Tag (#) =Iı

19757 1987

Erw o3 er, D = 7

52.29. 7 Aal STDUSTOTUM

Eu Al, Re E

=) 50% 20. B.

|| 7er. © b

95% 17.9.

kerEPaRag

10 25%= 21

5 + 50%= 42

75%= 63

Fo Max= 17

} | N = 84

| BRIUIRET > zu +

a Jum Jul 4 6 8 410 14 16 18 20 un

Abb. 19a: Nach Süden gerichtete Migration von Eristalis arbustorum (vgl. Legende zu Abb. 4a).

Fig. 19a: Southbound migration of Eristalis arbustorum.

Abb. 19b: Tageszeitliche Verteilung von Eristalis arbustorum (vgl. Legende zu Abb. 4d).

Fig. 19b: Diurnal activity patterns of Eristalis arbustorum migrating southward.

3%)

ohne ausgesprochene Maxima vor. Bei einer in Tirol beobachteten Schwebfliegenwanderung war

arbustorum ebenfalls unter den gesammelten Belegexemplaren (Jeekel & Overbeek 1968).

Aus Norddeutschland liegen einige Funde von Feuerschiffen (Heydemann 1967) und Beobachtun-

gen auf der Nordseeinsel Scharhörn vor (Schmid 1987). Dort erscheint die Art zwar regelmäßig, aber,

gemessen an ihrer Häufigkeit, in nur geringer Zahl.

Phänologie am Randecker Maar: Auch hier wird arbustorum nur in geringer Zahl hauptsächlich im

August in den Reusen gefangen (Tab. 2, Abb. 19a). Das tageszeitliche Aktivitätsmuster zeigt

Abb. 19b. Obwohl das Material sehr gering ist, ist der Unterschied zwischen den beiden Reusen deut-

lich (S-Reuse: N-Reuse = 5:1;n = 30.

Status: Saisonaler Migrant mit geringer Migrationsneigung.

Verglichen mit der Häufigkeit von arbustorum ist die Zahl der Wanderer klein. Für die alljährliche

Populationsentwicklung in Mitteleuropa dürften Migranten keine wesentliche Bedeutung haben.

Eristalis interrupta (Poda 1761) (= E. nemorum auct.)

Verbreitung: holarktisch

Biologie: Die Entwicklung dieser Art dürfte weitgehend der von arbustorum gleichen (Hartley

1961). Auch E. interrupta ıst eine eurytope, allerdings offene Habitate eher meidende, und häufige

Art; sie kann von (Mitte April) Anfang Mai bis Anfang (Mitte) Oktober gesehen werden. Ihre größte

Häufigkeit erreicht sie im Hochsommer.

Wanderungen: Von interrupta liegen keine auf Wanderungen hinweisende Beobachtungen vor.

Phänologie am Randecker Maar: Die Art wird — verglichen mit ihrer Häufigkeit — nur selten ge-

fangen (Tab. 2). Es dürfte sich dabei um Tiere der näheren Umgebung handeln.

Status: Der Status dieser Art entspricht möglicherweise dem von E. arbustorum. Die Wandernei-

gung von interrupta ist allerdings äußerst gering.

Eristalis pertinax (Scopoli 1763)

Verbreitung: westpaläarktisch

Biologie: Die sehr häufige, eurytope pertinax kann von (Ende Februar) Mitte April bis Ende Okto-

ber (Anfang November) beobachtet werden. Sie ist schon im Mai sehr zahlreich anzutreffen und ist

im Sommer oft eine der häufigsten Schwebfliegenarten. Die Larven leben in eutrophen Gewässern.

Die Art überwintert im Larvenstadium (Hartley 1961).

Wanderungen: Am Col de Bretolet trat die Art kaum in Erscheinung (Aubert et al. 1976; Tab. 2).

Die wenigen Fänge konzentrieren sich auf den August. Dethier & Goeldlin (1981) bezeichnen perti-

nax als Wanderart. Aus Norddeutschland liegt ein Fund von einem Feuerschiff vor (Heydemann

1967). Auf der Nordseeinsel Scharhörn konnte pertinax zwar nicht häufig, aber doch regelmäßig

nachgewiesen werden. Von hier stammen auch Beobachtungen nach SW fliegender Tiere (Schmid

1987).

Phänologie am Randecker Maar: Verglichen mit der Häufigkeit der Art wird pertinax in den Reusen

sehr selten gefangen (Tab. 2). Allerdings sagt das wenig über den tatsächlichen „Flugverkehr“ aus. Für

pertinax könnte hier dasselbe gelten wie für tenax: die meisten Fliegen erkennen die Reusen als Hin-

dernis und um- oder überfliegen sie gezielt. Einen Hinweis auf gerichtete Wanderungen von pertinax

geben die Zahlenverhältnisse zwischen den Reusen: auf 34 in der S-Reuse gefangene Ex. kam nur 1 Ex.

aus der N-Reuse.

Status: Saisonaler Migrant mit geringer Wanderneigung (vgl. E. arbustorum).

Eristalis tenax (Linnaeus 1758)

Verbreitung: kosmopolitisch

Biologie: PP von E. tenax überwintern selbst im nördlichen Europa noch regelmäßig (z. B. Torp

1984, Nielsen 1972). Sie sind im frühesten Frühjahr schon auf Blüten zu sehen. Im späten Frühjahr ist

tenax ausgesprochen selten. Erst im Frühsommer wird die Art wieder regelmäßig angetroffen. Im

Hoch- und vor allem ım Spätsommer und Herbst ist tenax eine der häufigsten Schwebfliegenarten. Sie

kann bis in den November hinein gesehen werden. Ihre Rattenschwanz-Larven leben in eutrophen

Gewässern. Selbst Extremhabitate werden besiedelt (Jauche, sich zersetzendes Aas etc.).

Wanderungen: Eristalis tenax gehört zu den auffälligsten Wanderern unter den Schwebfliegen. Gat-

ter (1980) beobachtete im Frühjahr (Mai) nordwärts gerichteten Schwebfliegenzug über einen 5400 m

hohen Himalaya-Paß. Unter den Belegexemplaren waren auch 3 Eristalis tenax. Beobachtungen

herbstlicher SW-Wanderungen liegen aus den Pyrenäen (Snow & Ross 1952, Williams et al. 1956), den

Alpen (z. B. Aubert etal. 1976, Eimer 1880, Prell 1925), den südwestdeutschen Mittelgebirgen (Gatter

1975a, 1976, Gatter & Gatter 1973) und aus dem mitteleuropäischen Küstenbereich vor (Lempke

1962, Schmid 1987). Williams (1961: 102) erwähnt regelmäßige Herbstwanderungen von Eristalis ent-

lang der Küste von New Jersey/USA in südliche Richtung. Möglicherweise handelt es sich auch hier

um E. tenax.

Erıstalis tenax ist, vor allem wenn es sich um Beobachtungen im Frühherbst handelt, oft die häufig-

ste und auffälligste Wanderart. Am Col de Bretolet ist tenax nach Episyrphus balteatus die am häufig-

sten gefangene Art (Tab. 2, Abb. 20a). Die Durchzugszahlen steigern sich von geringen Tagessum-

men im Juli bis zum sehr ausgeprägten Maximum im September und der ersten Oktoberdekade. Bis

Ende Oktober nehmen die Zahlen wieder sehr stark ab.

Eristalıs tenax © (Ph. W. Gatter).

Anzahl pro Pent.

Anz.

200

100

180

160

140

120

100

=[6)

ei®

pro Tag (+)

1975 - 1987

ER /ER

SR 10 MER

25% 28.8

50%. 11,29%

95% 37210

ler, 4152.10

Jun

rtengax

vooov0o0o00o 0

v v v

el RMV. 2 =

= 7.9 12 J 398

EO2SSIEEer 4.8.75 1

SOBE 50x 24.7.7 |

45 + 6. Wi

75% 19.9. 20. 8.78 2

9% 4.10. 17. 9.79 14

Le. 15.10. 19. 9.80 23

80 25. 9.81 4

205% 20. 9.82 3

Anzahl pro Pent. 4. 9.84 2

Anz. pro Tag (+) samaIE VU 19. 8.85 4

1975 - 1987 70 26. 8.86 19

N 7 35+| 20. 9.87 7-

Er. 44. 7. B = 18

3.8. 12 y 220

25% 30. 8. 19. 9.80 18 er

50% 11.9. 12.9.8 34 FH

9. 30 +

| 75x 19.9. 20. 9.82 2

95% 1.10. 29. 8.84 4

Le. 6.10. 19. 9.85 4

so | 15. 9.86 14 | 50

25 + 20. 9.87 2 e5+

40 40

eo + 20 +

30 30

As5+[T 15+

20 20

10+| 10 +

10 10

er 5+

KMITTUTTF T, T T: = -

b Jun Jul Aug Sep Okt C

Anzahl pro Pent.

Anz. pro Tag (+) tens3ax W

100 1975 - 1987

Abb. 20b: Nach Süden gerichtete Migration von Eristalis tenax-J'C' (vgl. Legende zu Abb. 4a).

Fig. 20b: Southbound migration of JO of Eristalis tenax.

Abb. 20c: Nach Süden gerichtete Migration von Eristalıs tenax-?Q (vgl. Legende zu Abb. 4a).

Fig. 20c: Southbound migration of QQ of Eristalis tenax.

Als große Insekten können fliegende Eristalis-Arten auch im Feld gut angesprochen und über kurze

Strecken mit dem Fernglas verfolgt werden. Wie Stichproben ergaben, handelte es sich am Randecker

Maar dabei überwiegend um E. tenax. Richtungsmessungen ergaben, daß die Fliegen fast ausschließ-

lich nach SW zogen (Gatter 1981a). Dabei entwickeln sie beachtliche Zuggeschwindigkeiten von ca.

25 km/h beı Windstille (Gatter 1981a mit weiteren Einzelheiten).

In den Alpen wurden am Col de Bretolet markierte tenax an einem 3 km südwestlich gelegenen Paß

innerhalb von 10-15 Minuten wiedergefangen (Aubert et al. 1969).

Phänologie am Randecker Maar: Hier steht tenax in der Fangstatistik mit 757 Expl. erst an neunter

Stelle (Tab. 2). Frühere Beobachtungen (Gatter & Gatter 1973, Gatter 1975a) belegen allerdings, daß

diese geringen Fangzahlen keineswegs Folgen eines nur sehr schwachen Zuggeschehens sind. Gatter

(1976) konnte beobachten, daß tenax die Reusen gewöhnlich als Hindernisse erkennt und um- oder

überfliegt. Trotzdem dürften auch die relativ wenigen Fänge zuverlässig Aufschluß über die Phänolo-

gie von tenax geben. Das auf den Fängen basierende Diagramm (Abb. 20a) dieser Art mit seinem typi-

schen Schwerpunkt erst im September entspricht im wesentlichen den durch Sichtbeobachtungen ge-

wonnenen Diagrammen bei Gatter (1975a). Auch am Col de Bretolet ist tenax die Art mit dem späte-

Abb. 20a: Nach Süden gerichtete Migration von Eristalis tenax (vgl. Legende zu Abb. 4a); eingeschaltet ist das

Zugmuster am Col de Bretolet (aus Aubert et al. 1976).

Fig. 20a: Southbound migration of Eristalis tenax; the insertion shows the phenology at Col de Bretolet.

FEN3IX

Wahre 1GFa EI GEZ

4.6 8 10 12 14 16 18 20 Uhr

L DIESE 1}

Ma 150 1

NEE 15. 9 N= 225

Uhr

Max= 32

> 16.-

CENGX M 31.8 Ne 443

Max= 15

Ike

15.8 N= 62

25%= 53

50%= 105

75%=- 158

Max= 50 JUL

N = 240

Unr e 4 6 8 10 12 414 16 18 20 Uhr

Abb. 20d: Tageszeitliche Verteilung von Eristalis tenax (vgl. Legende zu Abb. 4d).

Fig. 20d: Diurnal activity patterns of Eristalis tenax migrating southward.

Abb. 20e: Tageszeitliche Verteilung von Eristalis tenax im Jahresverlauf (vgl. Legende zu Abb. 5h).

Fig. 20e: Diurnal activity patterns of Eristalis tenax migrating southward throughout the year.

sten Maximum (vgl. Abb. 20a). PP werden am RM etwas häufiger gefangen als CC’ (O0: 29 =

1:1,3;n = 701). Die Diagramme für die beiden Geschlechter weichen nicht wesentlich vom Summen-

diagramm ab (Abb. 20b, 20c).

Auch das tageszeitliche Muster der Reusenfänge (Abb. 20d) bestätigt die durch Sichtbeobachtungen

(Gatter 1975a) gewonnenen Daten, nach denen das Gros in der Zeit von ungefähr 10-16 Uhr zieht,

während nach 16 Uhr nur noch wenige ziehende tenax beobachtet werden können. Mit fortschreiten-

der Jahreszeit verlagert sich der Aktivitätsschwerpunkt vom späten Vormittag auf den frühen Nach-

mittag (Abb. 20e). Das Aktivitätsmuster der beiden Geschlechter unterscheidet sich nicht wesentlich

voneinander.

Eristalis tenax wird Jahr für Jahr ın stark wechselnder Anzahl gefangen (Abb. 27n). Die weitaus

meisten Tiere geraten dabei in die S-Reuse (S-Reuse : N-Reuse = 46,5:1;n = 285). Gewöhnlich wer-

den sie dort bei Gegenwinden aus SW, S oder SE gefangen (75,8%, n = 317), nur gelegentlich bei Rük-

kenwind aus NW, N oder NE (7,5%) (Abb. 26f).

Status: Saisonaler Migrant.

Die Zahl der im mittleren und nördlichen Europa überwinternden Q 9 ist, im Vergleich zu der gro-

ßen Häufigkeit der Art, eher gering. Verlinden & Decleer (1987) vermuten, daß ein großer Teil der

Fliegen im Herbst nach Süden zieht. Die großen Zahlen (Hochrechnungen für das Randecker Maar

bei Gatter & Gatter 1973) zeigen, daß tatsächlich enorm viele Fliegen an der herbstlichen Wanderung

teilhaben.

Direkte Beobachtungen von Frühjahrswanderungen fehlen, jedoch lassen die sehr steil ansteigen-

den Zahlen im Sommer vermuten, daß Zuwanderung aus dem südlichen Europa für den Aufbau der

Sommerpopulationen mit verantwortlich ist. Dabei sind allerdings nicht die Immigranten selber, son-

dern deren Nachkommen für das Soemmermaximum verantwortlich.

Helophilus pendulus (Linnaeus 1758)

Verbreitung: paläarktisch

Biologie: H. pendulus ist eine eurytope und oft häufige Art. Die Larven wurden in eutrophen Ge-

wässern, aber auch in sehr nassem Dung und sich zersetzendem pflanzlichen Material gefunden. Die

Larven überwintern (Hartley 1961). Die Imagines können zwischen (Ende März) Mitte April und

Ende Oktober, gelegentlich auch noch im November und Dezember (Verlinden & Decleer 1987) ge-

sehen werden. Gewöhnlich wird im Juni ein erstes Maximum gebildet. Ihre größte Häufigkeit erreicht

die Art aber im Hochsommer (August).

Wanderungen: Am Col de Bretolet erscheint H. pendulus zwar nicht in sehr großer Zahl (Tab. 2),

aber regelmäßig, so daß Aubert et al. (1976) die Art als Wanderer einstufen. Das Maximum wird hier

Mitte September erreicht. Aus Norddeutschland existieren Beobachtungen nicht-ansässiger Tiere von

Feuerschiffen (Heydemann 1967) und von der Insel Scharhörn (Schmid 1987). Ein Mai-Nachweis von

dort stammt möglicherweise von einem Tier, das sich auf Nordwanderung befand. Regelmäßig wurde

die Art erst ab Anfang Juli gesehen.

Phänologie am Randecker Maar: H. pendulus wird in der S-Reuse nur wenig seltener gefangen als

Eristalıs tenax (Tab. 2). Der Zughöhepunkt liegt aber früher als bei dieser Art, nämlich an der Monats-

wende August/September (Abb. 21a). H. pendulus ist von den späten Vormittagsstunden bis in den

Nachmittag hinein ohne auffälliges Maximum flugaktiv (Abb. 21b). Der „Abendgipfel“ kommt durch

Volucella pellucens, eine monovoltine Art ohne saisonale Wanderungen (Ph.: W. Gatter).

Anzahl pro Pent

Anz. pro Tag (#) pendulaga

Es 1975 - 1987

e -

Se oe7 ar

5% 31.7

25% 21

Se EL E 16. 7.76 2

5% 10.9 15 78 6

9% 21.9 2 79 37

| te. 12.10

144 15 81 7£

72+ 4 82 3

venoamomomnu MW

pendaulg

Elan ee) EEE

Abb. 21a: Nach Süden gerichtete Migration von Helophilus pendulus (vgl. Legende zu Abb. 4a).

Fig. 21a: Southbound migration of Helophilus pendulus.

Abb. 21b: Tageszeitliche Verteilung von Helophilus pendulus (vgl. Legende zu Abb. 4d).

Fig. 21b: Diurnal activity patterns of Helophilus pendulus migrating southward.

die Extrapolation der Beobachtungen von 1975-77 (n = 14) zustande und bezieht sich nur auf zwei

gefangene Tiere, ist also zu vernachlässigen. Abb. 27 o zeigt die jährlichen Fangsummen und das Ver-

hältnis der Fangzahlen zwischen beiden Reusen: S-Reuse : N-Reuse = 6,5: 1 (n = 195).

Status: Saisonaler Migrant.

Helophilus trivittatus (Fabricius 1805)

Vorkommen: paläarktisch

Biologie: Die Larven von trivittatus und ihre Habitatansprüche sind nach unserem Wissen noch un-

beschrieben. Sie dürften aber denen von pendulus ähneln. Auch die Imagines von trivittatus sind eben-

sowenig an bestimmte Habitate gebunden wie die von pendulus. In vielen Gebieten ist trivittatus,

wenn auch nicht so häufig wie pendulus, in großer Zahl zwischen (Mitte April) Mitte Maı und Anfang

(Mitte) Oktober anzutreffen. Frühjahrsnachweise vor Mitte Mai sind ausgesprochen spärlich. Die

größte Häufigkeit wird meist im August und September erreicht.

Wanderungen: Aus den Alpen ist trivittatus vom CB als Wanderer mit ausgeprägtem Maximum in

der ersten Septemberhälfte bekannt (Tab. 2; Aubert et al. 1976). Williams et al. (1956) fingen 1 Expl.

unter migrierenden Syrphiden in den Pyrenäen. Auch aus Norddeutschland gibt es Wanderbeobach-

tungen von der Insel Scharhörn. Hier wurden die meisten Fliegen Ende August gesehen (Schmid

1987). Im Golf von Biskaya wurde 1 Expl. von trivittatus auf offener See gefangen (3.7.1957, Weidner

1958). Hinweise auf Wanderungen sind auch Herbstdaten aus Gebieten ohne Sommerbeobachtungen

(z. B. Schmid 1986).

Anzahl pro Pent.

Anz. pro Tag (+) ErRZAVZECECSES

1975 - 1987

V V V

Er. 18.7. ae 15

= 18

5% 26. 7 0

25% 13. 8. 1.8.7 >

36 50% 21.8.

a : u 24. 8.76 2

7% 3.9 15.08.78 7

95% 20.9. >3.8.79 6

leo 1oR 29. 8.80 4

30 | 15. 8.84 4

15 + oeae2 2

31. 8.84 3

13.8.85 2

15. 8.86 3

12 + 18. 9.87 5

6 +

N) BR [II }

Jun Jul Aug sep Okt

Abb. 22: Nach Süden gerichtete Migration von Helophilus trivittatus (vgl. Legende zu Abb. 4a) (eingefügte Zeich-

nung aus van der Goot 1981).

Fig. 22: Southbound migration of Helophilus trivittatus.

Phänologie am Randecker Maar: H. trivittatus erscheint zwar nur in geringer Zahl, aber regelmäßig

am RM (Tab. 2, Abb. 27p). Die Art wird dabei fast ausschließlich in der S-Reuse (S-Reuse :N-Reuse

= 33,5:1;n = 69) bei Gegenwinden aus SW, S oder SE (78,5 %) gefangen (Abb. 26g). Bei Rückenwin-

den aus NW, N oder NE geraten nur 6,8% der Fliegen in die S-Reuse. Trivittatus wandert überwie-

gend im August und September (Abb. 22). Das Maximum liegt am RM Ende August, während es am

CB erst in der ersten Septemberhältte erreicht wird.

Status: Saisonaler Migrant.

3.1.4 Schwebfliegen mit phytophagen Larven

Cheilosia pagana (Meigen 1822)

Verbreitung: paläarktisch

Biologie: Die sehr artenreiche Gattung Cheilosia ist in den Reusenfängen stark unterrepräsentiert

(vgl. Schmid & Gatter 1988). Lediglich die eurytope und häufige C. pagana, deren Larven phytophag

in Wurzeln des Wiesenkerbels Anthriscus sylvestris minieren (Stubbs 1980), wird zahlreich gefangen.

C. pagana bildet mehrere Generationen. Ihre Flugzeit beginnt gelegentlich schon Mitte März (Kor-

mann 1977), meist jedoch im April und dauert bis Mitte Oktober. Dabei sind oft Maxima ım Frühjahr

Anzahl pro Pent

nz. pro Tag (+) esgarna

1975 1987

Ep2 MER Z 1 D= 37 |

ala 7

| 25% 27.7 3.8.75 5

7 7 4

12. 8.77 1 PLIGaNgG

vahre 1975 - 1977

144 50% A

75% 14

95% 12

| Le. 30

wo omm

Dom.

* 25%= 125

* 50%= 250

* 75%= 376

Max= 82

*N = 501

N = 437

Uhr

Abb. 23a: Fänge in der S-Reuse von Cheilosia pagana (vgl. Legende zu Abb. 4a).

Fig. 23a: Trapped individuals with northerly opening trap (S-trap) of Cheilosia pagana.

Abb. 23b: Tageszeitliche Verteilung von Cheilosia pagana (vgl. Legende zu Abb. 4d).

Fig. 23b: Diurnal activity patterns of Cheilosia pagana.

und im Hochsommer deutlich. In welchem Entwicklungsstadium pagana überwintert, wird bei

Stubbs (1980) nicht deutlich. Zahlreiche Cheilosia-Arten überwintern als Puparium (z. B. Rotheray

1988, Smith 1979).

Wanderungen: Beobachtungen von Wanderungen fehlen bei sämtlichen Cheilosia-Arten. Am Col

de Bretolet sind zwar zahlreiche Arten nachgewiesen (Aubert et al. 1976), aber selbst im Bereich der

alpinen Matten lebende Cheilosien wurden nur in sehr geringer Zahl gefangen — Hinweis auf die im

allgemeinen sehr geringe Mobilität der Tiere. C. pagana ist nur mit 10 Expl. vertreten.

Phänologie am Randecker Maar: Auch die hohe Zahl von pagana-Fängen in den Reusen (Tab. 2)

dürfte nicht Ausdruck einer Wanderbewegung sein, sondern auf eine große ansässige Population zu-

rückgehen. Das an Doldenblütlern reiche Busch- und Wiesengelände der Stationsumgebung ist für

pagana ein ıdealer Lebensraum. Das Diagramm (Abb. 23a) spiegelt die lokale Häufigkeit der Art ge-

gen Ende des Hochsommer-Maximums wider.

Das Aktivitätsmaximum dieser Art liegt um die Mittagsstunden (Abb. 23b).

Der Unterschied zwischen den Reusen (S-Reuse:N-Reuse = 3,1:1;n = 232; Abb. 27q) geht ver-

mutlich auf die kleinräumigen Unterschiede im vor der Reusenöffnung gelegenen Lebensraum zurück

(vgl. Rhingia campestris, Neoascia podagrica).

Status: Polyvoltine Art ohne oder mit sehr geringer saisonaler Dismigrationsneigung.

3.2 Weitere Wanderarten in Europa

Nicht alle in Europa bisher wandernd beobachteten Syrphidenarten konnten am RM in einer zur

Klärung ihres Status hinreichend großen Zahl beobachtet werden. Deshalb soll hier zunächst ein

Überblick über diese Arten gegeben werden.

Die meisten Beobachtungen stammen vom Col de Bretolet. Hier konnten insgesamt 186 Schweb-

fliegenarten nachgewiesen werden (Aubert etal. 1976), von denen 30 als wandernd oder zumindest als

wanderverdächtig eingestuft wurden. In welche Kategorie diese Arten am CB eingestuft wurden, ist

jeweils sowohl übersetzt als auch im Wortlaut zitiert.

3.2.1 Schwebfliegen mit zoophagen Larven

Platycheirus scutatus (Meigen 1822)

Verbreitung: holarktisch

CB: n = 186, während der ganzen Beobachtungsperiode verteilt, vielleicht Wanderart („peut-etre

migratrice“); RM: n = 16, nur in zwei Jahren beobachtet.

Die Art ist im allgemeinen zahlreich. Sie kommt in Mitteleuropa von (Anfang April) Anfang Mai

bis Mitte Oktober (Anfang November) vor und bildet ı. a. zwei Generationen (Krüger 1926) die als

deutliche Maxima erkennbar sind (Verlinden & Decleer 1987). Angesichts der Spärlichkeit der Fänge

ist regelmäßiges Wandern in größerem Ausmaß ausgeschlossen. Gelegentliche Dismigrationen kön-

nen vorkommen (vgl. Schmid 1987). Die im Larvenstadium an Blattlauskolonien der Krautschicht le-

bende, obligatorisch aphidophage (Rotheray & Gilbert 1989) Art kann als Saisonaler Dismigrant mit

geringer Migrationsneigung betrachtet werden.

Xanthandrus comtus (Harrıs 1780)

Verbreitung: paläarktisch, orientalisch (Formosa)

CB: n = 441, Herbstwanderart („Migrateur automnal“) mit Maximum Mitte Oktober und Vor-

kommen 7.7.—21.10. Von den 441 Expl. allein 231 Expl. in einem Jahr; RM: n = 17.

Die in Mitteleuropa zwischen Mitte Mai und Mitte Oktober mit einem deutlichen Schwerpunkt im

Spätsommer vorkommende Art ist meist recht selten. Gelegentliches häufigeres Vorkommen könnte

mit Gradationen der Nahrungstiere zusammenhängen: comtus-Larven fressen v. a. Schmetterlings-

raupen, insbesondere Gespinstmotten (Yponomeutidae) und Wickler (Tortricidae), aber auch Kie-

fernprozessionsspinner (Thaumetopoea pinivora Tr.) (weitere Nahrungstiere in der Zusammenfas-

sung von Torp [1984]). Snow & Ross (1952) fingen comtus am 21.9. unter wandernden Syrphiden in

den Pyrenäen. Weitere Hinweise auf Wanderungen fehlen. Der Status der Art erscheint weiterhin un-

geklärt.

Metasyrphus lapponicus (Zetterstedt 1838)

= Enpeodes lapponicus

Verbreitung: holarktisch

CB: n = 10235, während der ganzen Beobachtungsperiode anwesend, Maximum variiert von Jahr

zu Jahr („Migrateur“); RM:n = 4.

M. lapponicus gehört zu den Arten, die in Mitteleuropa als Imago überwintern (Schneider 1958,

Goeldlin 1974, Dusek & Läska 1986) und kann deshalb schon früh im Jahr beobachtet werden (März).

Auch im Herbst ist die Art lange aktiv (November). Die Flugzeitangaben aus dem nördlichen Mittel-

europa und Nordeuropa (Belgien: Anfang April bis Ende August [Verlinden & Decleer 1987], Nie-

derlande: Ende Mai bis Anfang August [van der Goot 1981], Südnorwegen: Ende Mai bis Mitte Au-

gust [Nielsen 1971], Schleswig: Mitte Mai bis Mitte August [Claußen 1981], Dänemark: Ende Mai bis

Anfang September [Torp 1984]) zeigen, daß hier keine Überwinterung stattfindet. Die fehlenden

Spätherbstdaten lassen vermuten, daß eine Überwinterung auch nicht versucht wird. Die Art muß in

diese Gebiete also alljährlich einwandern. — M. lapponicus ist in Mitteleuropa in den meisten Jahren

eine eher seltene, waldbewohnende Art; nur gelegentlich ist sie häufiger anzutreffen. In den Alpen ist

lapponicus dagegen stets zahlreich. Möglicherweise ist das der Hauptgrund für die geringen Zahlen am

RM und den stärkeren Durchzug am CB mit Jahressummen zwischen 82 und 3765 Exemplaren.

Ein Hinweis auf Wandertendenzen dieser Art im Mittelgebirgsbereich ist z. B. die Bemerkung von

Malec (1986) aus dem Kasseler Raum, daß lapponicus im Herbst „mobiler“ zu sein scheine und dann

selbst in Großstadtgärten anzutreffen sei. Im Sommer 1989 war lapponicus nach einem sehr blattlaus-

reichen Frühling auch um Tübingen (SW-Deutschland) ungewöhnlich zahlreich und immer wieder in

Gärten und selbst in Häusern zu finden (Schmid unveröff.). Möglicherweise beziehen sich auch Hin-

weise auf Wanderungen von „Syrphus arcuatus“ in Südengland bei Walker (1864) auf lapponicus —

allerdings ist die Art in Großbritannien selten (Stubbs & Falk 1983).

Aufgrund ihres Lebenszyklus und der Flugzeiten in Mittel- und Nordeuropa ist zu erwarten, daß

M. lapponicus zu den Saisonalen Migranten gehört.

Metasyrphus latifasciatus (Macquart 1829)

= Eupeodes latifasciatus

Verbreitung: holarktisch

CB: n = 4909, während der ganzen Beobachtungsperiode anwesend, undeutliches Maximum An-

fang September („Migrateur“); RM: n = 20.

Der Lebenszyklus dieser Art entspricht weitgehend dem von M. corollae; allerdings scheint noch

nicht bekannt zu sein, in welcher Form die Überwinterung stattfindet. Typisch für latifasciatus sind

die extremen Häufigkeitsunterschiede von Jahr zu Jahr (Stubbs & Falk 1983, Verlinden & Decleer

1987). Am RM wurde die Art in vielen Jahren gar nicht beobachtet. M. latifasciatus ist eurytop mit

einer Vorliebe für offene und feuchte Habitate. Sie fliegt ab Anfang April bis Anfang Oktober und bil-

det mehrere Generationen. Um den Status der Art zu klären, bedarf es weiterer Beobachtungen.

Metasyrphus luniger (Meigen 1822)

= Eupeodes luniger

Verbreitung: paläarktisch, nordorientalisch (Assam)

CB: n = 12002, während der ganzen Beobachtungsperiode anwesend, Maximum variiert („Migra-

teur“),; RM:n =7.

Wie M. corollae ist auch luniger eine polyvoltine Art, die als Puparium überwintert (Scott 1939, Du-

sek & Läska 1974). Möglicherweise können gelegentlich auch Imagines überwintern (Gauss 1961,

Nielsen 1971), der Normalfall ist dies aber sicher nicht (Dusek & Läska 1986). Die Larven von /uniger

leben von zahlreichen Blattlausarten (Läska & Stary 1980) überwiegend in der Krautschicht (Dixon

1960, Rotheray & Gilbert 1989). M. luniger kann gewöhnlich zwischen Mitte April und Oktober be-

obachtet werden, gelegentlich auch noch später (9 QQ am 17. 12., [Schneider 1958]). Die früher oft als

häufig und weit verbreitet bezeichnete Art ist heute vielerorts ausgesprochen selten. Torp (1984)

spricht von einem sicheren Rückgang der Art in Dänemark. Dies dürfte für die wenigen Nachweise

der Art am RM mit verantwortlich sein. Der mit corollae übereinstimmende Lebenszyklus macht

wahrscheinlich, daß auch M. luniger zu den Saisonalen Migranten gehört, wenn auch Wanderdaten

nur spärlich vorliegen (z. B. Johnson 1960, Schmid 1987).

Dasysyrphus albostriatus (Fallen 1817)

Verbreitung: paläarktisch

CB: n = 548, Maximum Mitte September („Migrateur“); RM: n = 13.

D. albostriatus hat einen unter mitteleuropäischen Schwebfliegen ungewöhnlichen Lebenszyklus:

die Art ist bivoltin mit Hauptflugzeit im Frühjahr und Herbst und obligatorischer Larvendiapause

(Schneider 1948). Die zweigeteilte Flugzeit ermöglicht albostriatus die Ausbeutung des Frühjahrs-

wie des Herbstmaximums der Blattlauspopulationen (Dusek & Läska 1986). Die Art ist hauptsächlich

in Wäldern und an Waldrändern weit verbreitet, meist aber nur in geringer Anzahl anzutreffen. Dasy-

syrphus-Larven leben aphidophag überwiegend auf Laub- und Nadelbäumen (Rotheray & Gilbert

1989).

Am CB erscheint albostriatus vereinzelt ab Mitte Juli. Bis Anfang September steigern sich die Fang-

zahlen kontinuierlich, dann nehmen sie bis Anfang Oktober wieder ab. Hier spiegelt sich die Flugzeit

der Herbstgeneration getreulich wider. Vermutlich wandert D. albostriatus allenfalls in geringem

Ausmaß und ist als Saisonaler Dismigrant mit geringer Migrationsneigung zu betrachten.

Melangyna cincta (Fallen 1817)

Verbreitung: Westliche Paläarktıs, Nearktis

CB: n = 1075, deutliches Maximum Ende August/Anfang September, bis Mitte September sehr

schnelle Abnahme der Zahlen („Migrateur“); RM: n = 16.

Von Mitte April bis Mitte September kann diese Art vor allem ın Wäldern und an Waldrändern an-

getroffen werden. Ihre Larven wurden an Blattläusen auf Buchen (Fagus sylvatica) gefunden (Läska

& Stary 1980). Innerhalb ihrer Flugzeit ıst ein ausgeprägtes Maximum ım Frühjahr (Mai bis Anfang

Juni) ausgebildet. Im Sommer kann ein zweiter, schwächerer Höhepunkt ausgebildet sein. Dusek &

Läska (1962, 1986) halten cincta für eine univoltine Art; die Sommerdaten stammen demnach von ver-

früht geschlüpften und nicht erfolgreich fortpflanzungsfähigen Individuen. M. cincta kann am besten

mit Malaisefallen erfaßt werden (Schmid 1986, Verlinden & Decleer 1987); Angaben über die Häufig-

keit im Vergleich zu anderen Arten lassen sich somit nach Kescherfängen schlecht machen. Insgesamt

scheint cincta nicht selten zu sein, aber sehr starke Populationsschwankungen aufzuweisen (z. B. Ma-

lec 1986). — Der Status der Art erscheint unklar. Weitere Beobachtungen sind notwendig.

Parasyrphus annulatus (Zetterstedt 1838)

Verbreitung: paläarktisch

CB:n = 1786, Maximum zu Beginn der Beobachtungen im Juli („Migrateur probable“); RM: 3 Ex.

P. annulatus kann zwar gelegentlich bis in den Herbst hinein angetroffen werden, kommt aber nur

im Frühjahr in so großer Anzahl vor, daß sie lokal — so z. B. im Tübinger Raum (Schmid 1986), am

Alpennordrand (Schmid unveröff.) oder in der Westschweiz (Goeldlin 1974) — sehr häufig sein kann.

Möglicherweise ist die im Larvenstadium an Blattläusen in Fichtenwäldern (Kula 1980) gefundene Art

univoltin (z. B. Niederlande: Beobachtungen nur Mitte Mai bis Anfang Juli [van der Goot 1981]) mit

der Tendenz der Ausbildung einer zweiten Generation (vgl. Melangyna cincta). Überwintern dürfte

die diapausierende Larve. Von Arten mit einem solchen Entwicklungstyp sind keine gerichteten Sai-

sonwanderungen bekannt. P. annulatus führt Saisonale Dismigrationen im Zusammenhang mit der

Suche nach geeigneten Eiablageplätzen durch. Dafür spricht auch, daß in einem als Malaisefalle arbei-

tenden Gebäude am Rand von Tübingen ganz überwiegend PP gefangen wurden (Schmid 1986 und

unveröff.).

Parasyrphus punctulatus (Verrall 1873)

Verbreitung: Europa, Altai

Auch die QQ dieser ebenfalls nur in einer Frühjahrsgeneration auftretenden Art führen wie P. an-

nulatus saisonale Dismigrationen aus (vgl. Schmid 1986). Auch punctulatus ıst eine aphidophage Art,

die überwiegend, wenn auch nicht ausschließlich an Nadelbäumen lebt. Rotheray & Gilbert (1989) ge-

ben Funde von Abies, Pinus, Acer psendoplatanus und Rosa an. Bei Tübingen wurden Mitte Mai 1988

zahlreiche QQ bei der Eiablage an verlausten Fichtentrieben (Picea abies) beobachtet (Schmid unver-

öff.).

Parasyrphus vittiger (Zetterstedt 1843)

Verbreitung: paläarktisch

CB:n = 1568, undeutliches Maximum Ende August („Migrateur“); RM: 4 Ex.

Auch diese im allgemeinen eher seltene, im Larvenstadium an Blattläusen auf Tanne, Kiefer und

Johannisbeere (Ribes nigrum) lebende Art (Goeldlin 1974, Rotheray & Gilbert 1989) hat eine relativ

lange Flugzeit (Mitte April bis Mitte Oktober). Im Tübinger Raum existiert ein ausgeprägtes Früh-

jahrsmaximum (Schmid 1986). Hinweise darauf fehlen andernorts, so daß ungeklärt ist, wie viele Ge-

nerationen vittiger normalerweise ausbildet. In den Alpen kommt vittiger zahlreich bis über die

Waldgrenze vor (Goeldlin 1974, eigene Beobachtungen). Der Status dieser Art dürfte dem von P. an-

nulatus entsprechen: Saisonale Dismigration.

Didea alneti (Fallen 1817)

Verbreitung: holarktisch

CB: n = 817, ab Mitte Juli bis Ende September mit einem wenig ausgeprägten Maximum Ende Au-

gust („considere comme migrateur“); RM:n = 2.

D. alneti fliegt zwischen Mitte Mai und Mitte Oktober ohne deutliche Höhepunkte, dürfte also po-

lyvoltin sein. Diese aphidophage Art scheint nirgends zahlreich vorzukommen. Wanderhinweise ab-

seits des CB fehlen. Der Status der Art ist unklar.

Didea fasciata Macquart 1834

Verbreitung: holarktisch, orientalisch (Formosa)

CB: n = 3171, ausgeprägtes Maximum Ende Juli/Mitte August („Migrateur“); RM:n = 11.

Die Flugzeit dieser in Wäldern, aber auch in Gärten anzutreffenden, aphidophagen Art beginnt

Ende April und endet Ende Oktober. Maxima liegen in Dänemark Mitte Juni und Anfang August

(Torp 1984), in Belgien im August. Verlinden & Decleer (1987) vermuten, daß dieses Maximum durch

Wanderer verursacht wird. Direkte Hinweise auf Wanderungen gibt es außer vom CB nur noch aus

der Steiermark (Gepp 1975) und aus den Pyrenäen (1 Ex. am 10.10.; Williams et al. 1956). Der Status

von D. fasciata bedarf weiterer Klärung.

Meliscaeva auricollis (Meigen 1822)

Verbreitung: Westliche Paläarktis

CB: n = 3127, Maximum Ende September („Migrateur“); RM:n = 11.

Wie bei der sehr nahe verwandten Art Episyrphus balteatus können die PP, wie Schneider (1948)

im Alpenraum nachwies, als Imagines überwintern. Goeldlin (1974) nennt für die Westschweiz eine

Flugzeit von Februar bis November. Nördlich der Alpen scheint die Überwinterung nur in Ausnah-

mefällen zu glücken. Hier erscheint auricollis nur in Ausnahmefällen schon im Februar (Großbritan-

nien [Stubbs & Falk 1983]) oder im März (ein Fund am 26.3. in Belgien [Verlinden & Decleer 1987]).

Normalerweise fliegt die Art erst ab Mitte April, im Norden sogar erst ab Mai. Torp (1984) vermutet,

daß die dänische Population alljährlich von Einwanderern aufgebaut wird. Wandernde Exemplare

wurden auch in Südengland (Mackworth-Praed 1929) und auf der Insel Scharhörn festgestellt (Schmid

1987).

Die Larven von auricollis sind aphidophag und bewohnen Bäume, Büsche und Kräuter. Auch ın

Kolonien von Blattflöhen (Psyllidae) konnten sie gefunden werden (Dixon 1960, Laska & Stary 1980,

Rotheray & Gilbert 1989). Meliscaeva auricollis ist zwar sehr weit verbreitet, kommt aber meist nur

vereinzelt vor. Das erklärt ihre Seltenheit am RM. Der fast völlig mit dem von Episyrphus balteatus

übereinstimmende Lebenszyklus dieser Art und ihre Phänologie in Mitteleuropa machen sehr wahr-

scheinlich, daß auch sie zu den Saisonalen Migranten gehört.

3.2.2 Schwebfliegen mit terrestrischen saprophagen Larven

Xylota segnis (Linnaeus 1758)

Verbreitung: Westliche Paläarktis, nearktisch

CB: n = 130, einzelne Exemplare zwischen dem Beginn der Beobachtungsperiode und Anfang Ok-

tober; die Art könnte Wanderer sein? („pourrait etre migrateur?“) [dagegen Aubert mdl. an Gatter:

die Art wandert nicht]; RM: n = 47.

Diese häufigste und am weitesten verbreitete Xylota-Art kann zwischen (Ende März) Anfang Mai

und Mitte Oktober (Anfang November) beobachtet werden. Ihre Larven leben in vermoderndem

Holz, in nassem Sägemehl oder auch in Sılage (Hartley 1961). Sie überwintern und verpuppen sich im

Frühjahr. In SW-Deutschland werden vermutlich drei Generationen gebildet (Schmid 1986). Die O'C°

dieser Art verhalten sich territorial. Vor allem legereife PP zeigen dagegen ein ausgeprägtes saisonales

Dismigrationsverhalten, wohl auf der Suche nach Eiablageplätzen (Schmid |. c.). Weiterführende

Wanderungen finden nicht statt.

Xylota florum (Fabricius 1805)

Verbreitung: paläarktisch

CB:n = 24;RM:n=0.

Ein dem von X. segnis entsprechendes, vor allem von PQ ausgeführtes, saisonales Dismigrations-

verhalten zeigt auch die nahe verwandte X. florum. Diese Art wird deshalb ebenfalls sehr zahlreich in

Einflugfallen gefangen (vgl. Löhr 1987, Schmid 1986).

Xylota sylvarum (Linnaeus 1758)

Verbreitung: paläarktisch

CB:n=7;RM:n=15.

Auch sylvarum wird nicht selten in Einflugfallen gefangen (Löhr 1987, Schmid 1986). Hier sind

ebenfalls fast ausschließlich PQ an der nur über kurze Distanzen reichenden Saisonalen Dismigration

beteiligt, während die Q’C’ ein ausgesprochenes Territorialverhalten an den Tag legen (Schmid 1986).

3.2.3 Schwebfliegen mit sapro-/zoophagen Larven

Volucella zonaria (Poda 1761)

Verbreitung: südliche Paläarktıis

Volucella-Larven leben in Nestern von Hummeln und Wespen. Dort ernähren sie sich einerseits

von Abfällen und toten Tieren, andererseits zumindest gelegentlich auch von den Larven ihrer Wirte

(vgl. Torp 1984).

Volucella zonaria ist eine im Mittelmeergebiet sehr zahlreich vorkommende Art. Im Hochsommer

(meist August) wandern immer wieder einzelne dieser großen und sehr auffälligen Schwebfliegen in

Mitteleuropa ein. Gelegentlich kommt es zu zahlenstärkeren Einflügen (z. B. van der Goot 1986b).

Fast alle dieser Einwanderer sind weiblich. Infolge solcher expansiver Dismigrationen kann es zu

Arealerweiterungen kommen: V. zonaria etablierte sich in den 1940er Jahren in Südengland und ist

seither dort heimisch. Bis 1960 war die Art dort relativ häufig. In den letzten Jahren wurde sie wieder

seltener (Gilbert 1986). Diese Beobachtungen decken sich mit Arealausweitungen und -verlusten me-

diterraner Vogelarten in Mitteleuropa. Im Zusammenhang mit einer längeren Klimaphase mit hohen

Sommertemperaturen und geringen Niederschlägen in der Mitte des 20. Jahrhunderts dehnten solche

Arten ihr Verbreitungsgebiet weit in den mitteleuropäischen Raum hinein aus. Die in den 1970er Jah-

ren einsetzende, für diese Arten ungünstige Klimaphase mit feuchteren und kühleren Sommern führte

wieder zu starken Arealverlusten. Sowohl die Ansiedlung von Volucella zonaria ın Südengland wie

ihre zunehmende Seltenheit in den letzten Jahren dürften auf dieselben klimatischen Ursachen zu-

rückgehen.

Status: Expansiver Dismigrant.

Die Neigung zu expansiven Dismigrationen ermöglicht Volucella zonaria eine schnelle Reaktion

auf wechselnde Umweltbedingungen.

3.3 Übersicht über die Wanderschwebfliegen Europas

Tabelle 3: Übersicht über die Wanderschwebfliegen Europas.

Tab. 3: Migratory hoverfly species of Europe.

Larvennahrung Überwinterung Lebenszyklus #

Saisonale Migranten

Syrphus ribesu Blattläuse Larve 3

Syrphus torvus Blattläuse Larve 3

Syrphus vitripennis Blattläuse Larve 3

Metasyrphus corollae Blattläuse Puparium 2

Metasyrphus lapponicus Blattläuse Imago 1

Metasyrphus lunıiger Blattläuse Puparıum 2

Scaeva pyrastri Blattläuse Imago 1

Scaeva selenitica Blattläuse Imago 1

Melscaeva auricollis Blattläuse Imago 1

Melıiscaeva cinctella Blattläuse (Imgao?) (?1)

Episyrphus balteatus Blattläuse Imago 1

Sphaerophoria scripta Blattläuse Larve 3

Eristalis tenax aquat. sapro-/microphag Imago

Helophilus pendulus aquat. sapro-/microphag Larve

Helophilus trivittatus aquat. sapro-/microphag (Larve?)

Saisonale Migranten mit geringer Wanderneigung

Eristalis arbustorum aquat. sapro-/microphag Larve

Eristalis interrupta” aquat. sapro-/microphag Larve

Eristalis pertinax aquat. sapro-/microphag Larve

Saisonale Dismigranten

Platycheirus albimanus Zoophag in der Streu- Larve 3

schicht, Blattläuse

Platycheirus clypeatus Zoophag in der Streu- Larve 3

schicht, Blattläuse

Platycheirus manicatus” Blattläuse (fakultatıv) Larve 3

Platycheirus peltatus Blattläuse (fakultatıv) Larve 3

Melanostoma mellinum Blattläuse, zoophag Larve 3

in der Streuschicht

Parasyrphus annulatus Blattläuse Larve (?4)

Parasyrphus lineolus“ Blattläuse Larve (?3)

Parasyrphus punctulatus Blattläuse Larve (?7)

Parasyrphus vittiger Blattläuse Larve ?

Xylota florum terrestrisch saprophag Larve

Xylota segnis terrestrisch saprophag Larve

Xylota sylvarum terrestrisch saprophag Larve

Saisonale Dismigranten mit geringer Wanderneigung

Melanostoma scalare Blattläuse, zoophag Larve 3

in der Streuschicht

Platycheirus scutatus Blattläuse Larve 3

Dasysyrphus albostriatus Blattläuse Larve 6

Neoascıa podagrica terrestrisch saprophag Larve

Syrıtta pipiens terrestrisch saprophag Larve

Expansıve Dismigranten

Volucella zonarıia sapro- und zoophag in (Larve?)

Wespen- und

Hummelnestern

Status ungeklärt

Xanthandrus comtus Schmetterlingsraupen, Larve (25)

Blattläuse

Metasyrphus latıfasciatus Blattläuse ? 23)

Melangyna cincta Blattläuse Larve 4

Didea alneti Blattläuse Larve (?3)

Didea fasciata Blattläuse Larve (?3)

*: Status nicht ganz sicher

Aufbauend auf Schneider (1948, 1969) geben Dusek & Läska (1986) einen Überblick über die

Lebenszyklen der aphidophagen Syrphiden. Sie unterschieden (vereinfacht) folgende Typen

(der letztgenannte Typ 7 ist vorerst nur bei einer Art verifiziert (Dusek & Läska 1986), kommt

aber möglicherweise noch bei anderen Frühjahrsarten vor):

mehrere Generationen im Jahr, Überwinterung als Imago.

mehrere Generationen, Überwinterung als Puparium.

mehrere Generationen, Überwinterung als Larve.

eine Generation, Überwinterung als Larve, Flugzeit: Frühjahr.

eine Generation, Überwinterung als Larve, Flugzeit: Herbst.

zwei Generationen, Überwinterung als Larve, Flugzeit: Frühjahr und Herbst.

eine Generation im Frühjahr, Verpuppung im Herbst, Überwinterung als Puparium.

Bei Arten, deren Lebenszyklus noch nicht genauer beschrieben wurde, wird der aus der Phänologie

der Imagines geschlossene wahrscheinliche Zyklus in Klammern angegeben.

3.4

Wanderaktivität in Abhängigkeit von den Windverhältnissen

Unter den Wetterfaktoren, die Insektenwanderungen am stärksten beeinflussen, kommt dem Wind

eine herausragende Bedeutung zu (Gatter 1975, 1977a, 1977b). Die Beziehungen zwischen Schweb-

fliegenwanderungen und Windverhältnissen sollen deshalb ausführlicher dargestellt werden.

Im folgenden wird zunächst das Verhalten des Saisonwanderers Episyrphus balteatus und des Dis-

migranten Platycheirus clypeatus genauer analysiert.

Dazu werden zuerst Daten aus dem Jahr 1986 miteinander verglichen. In diesem Jahr waren beide

Arten annähernd gleich häufig. Anschließend werden die Verhältnisse im Jahr 1981 dargestellt. Aus

diesem Jahr liegen Daten aus beiden Reusen vor.

3.4.1 Episyrphus balteatus 1986

Abb. 24a zeigt, daß balteatus ganz überwiegend bei Gegenwind gefangen wird. Das größte Kontin-

gent bilden bei S- oder SW-Winden mit Stärken von 2-4 Bft. erfaßte Tiere. Dieses scheinbar paradoxe

Phänomen ließ sich am Randecker Maar durch Beobachtungen klären. Demnach nutzen die Fliegen

fördernde Rückenwinde aus, indem sie in größere Höhen aufsteigen. Den widrigen Einfluß von Ge-

genwinden versuchen sie dagegen dadurch zu verringern, daß sie bodennah ziehen. Bei mäßigen Ge-

genwinden geraten deshalb die weitaus meisten Tiere in die S-Reuse.

Schwebfliegen wie Episyrphus balteatus zeigen also ein Verhalten, wie es auch bei migrierenden

Schmetterlingen (Gatter 1981a) oder ziehenden Vögeln beobachtet werden kann.

Die Strategie des Unterfliegens widriger Gegenwinde wird am RM an vielen Tagen, an denen der

Wind dreht, deutlich (vgl. Gatter 1981b). Bei Gegenwinden fängt sich balteatus in großer Zahl in der

Reuse; sobald der Wind dreht, lassen die Fangzahlen schlagartig nach (Tab. 4). Stichprobenhafte Be-

obachtungen mit dem Fernglas zeigen dann häufig hohen Zug (vgl. 3.4.6.1).

3.4.2 Platycheirus clypeatus 1986

Der Saisonale Dismigrant P. clypeatus zeigt ein anderes Verhalten als der Saisonmigrant Episyrphus

balteatus. Ein großer Teil gelangt mit schwachen Rücken- oder Seitenwinden (1-2 Bft.) in die

S-Reuse (Abb. 24b). Die Art scheint demnach keine dem Verhalten von balteatus entsprechende Stra-

tegie der Nutzung günstiger Luftströmungen durch Aufsteigen in höhere Luftschichten zu haben.

Die Art wurde 1986 gewöhnlich bei etwas schwächeren Winden gefangen als E. balteatus

(Abb. 25a).

Abb. 24: (Seite 77) Windrichtung und Windstärke bei Wanderungen von Episyrphus balteatus und Platycheirus

chypeatus in ausgewählten Jahren.

Für einzelne Jahre wurden für sämtliche Individuen der jeweiligen Art Windrichtung und -stärke zum Zeitpunkt

des Fanges dargestellt. Die Diagramme zeigen, wie viele Individuen (in Prozent) bei welcher Windrichtung und

-stärke in der S-Reuse oder N-Reuse gefangen wurden. Die Zahlen geben den Anteil (in Prozent) am Gesamtfang

an. (Ablesebeispiel: 24a; 41,7 % aller im Jahr 1986 in der S-Reuse gefangenen balteatus wurden bei S-Wind gefan-

gen, die meisten davon bei Windstärken von 3 und 4 Bft.).

Fig. 24: Wind direction and wind force during migration of Episyrphus balteatus and Platycheirus chypeatus in selec-

ted years.

In each year for all specimens, wind direction and force at the time of trapping are shown. The diagrams indicate

how many individuals (in %) were collected in different wind conditions. (Example: Fig. 24a; 41,7 % of all catches

of E. balteatus in 1986 in the trap opening to the north are collected with a southerly wind, most of them at wind

forces of 3 and 4 Bft.). S-trap: trap with opening to the north (for south-flying insects); N-trap: trap with opening

to the south (for north-flying insects).

a) E. balteatus, S-Reuse/S-trap, 1986, n = 2937 b) P. clypeatus, S-Reuse/S-trap, 1986, n = 3485

c) E. balteatus, S-Reuse/S-trap, 1981, n = 4244 d) P. clypeatus, S-Reuse/S-trap, 1981, n = 1528

e) E. balteatus, N-Reuse/N-trap, 1981, n = 355 f) P. clypeatus, N-Reuse/N-trap, 1981, n=987

Tabelle 4: Stündliche Fangsummen (S-Reuse) von Episyrphus balteatus an drei ausgewählten Tagen mit drehendem

Wind.

Tab. 4: Catches per hour of Episyrphus balteatus in the S-trap (trap with opening to the north) at three selected days

with shifting winds.

1.8.1982

Uhrzeit 5 6 7 8 9 10 11 12 13 14 15 16 17 18

Wind SWFSWE SWR FSW. W W N N N N N N N N

Stärke l 1 1 1 1 1 1 3 ö) 2 3 2 3 2

balt. 3 22 130 10 30 42 6 — 1 2 1 _ _ _

22.8.1984

Uhrzeit 5 6 7 8 9 10 11 12 13 14 15

Wind S SE SE S SE S S N N N N

Stärke l 2. 1 2 2 1 1 1 1 2 1

balt. 3 3 4 70 84 126 43 22 4 7 8

14.8.1985

Uhrzeit 5 6 7 8 9 10 11 12 13 14 15 16

Wind S S S S SW NW N N S S SE SE

Stärke 2 2 2 2 1 2 2 1 2 1 2 2

balt. A DD 75554 169 37 10 9 25 12 12 16

3.4.3 Episyrphus balteatus 1981

Abb. 24c gleicht im wesentlichen Abb. 24a: ein großer Teil der Fliegen wird in der S-Reuse bei Ge-

genwinden mäßiger Stärke gefangen. Auch in die N-Reuse geraten die meisten Tiere gegen den Wind

(Abb. 24e). Auch hier handelt es sich also keineswegs um einen Verdriftungseffekt. Eher ist denkbar,

daß sich im Augenblick nicht wandernde balteatus gegen den Wind orientieren, um einer Verdriftung

entgegenzuwirken; zum anderen muß berücksichtigt werden, daß hochsteigende Syrphiden sich ge-

gen den Wind wenden, um danach ihre Wanderung mit dem Wind fortzusetzen (vgl. 3.4.6.1; Gatter

1981a).

In die N-Reuse fliegt balteatus überwiegend bei ganz schwachen Winden, während sich die Art in

der S-Reuse in größerer Zahl bei etwas stärkeren Winden fängt (Abb. 25a, 25b).

3.4.4 Platycheirus clypeatus 1981

Abb. 24d bestätigt die an Abb. 24b gewonnenen Ergebnisse: c/ypeatus wird überwiegend bei Rük-

kenwinden aus NW, N oder NE (59,8%) und nur in bedeutend geringerer Zahl bei Gegenwinden aus

SW, S oder SE (24,4%) in der S-Reuse gefangen. Auch in der N-Reuse fangen sich mehr c/ypeatus mit

dem Wind (47,0%) als gegen ihn (38,8 %) (Abb. 24f). Damit wird deutlich, daß c/ypeatus im bodenna-

hen Bereich in erheblichem Ausmaß mit dem Wind wandert.

Dabei könnte die Möglichkeit bestehen, für eine bevorzugte Migrationsrichtung günstige Winde zu

wählen und sich von ihnen „gezielt“ verdriften zu lassen. In die N-Reuse flogen nur 47,0% der dort

gefangenen c/ypeatus mit Rückenwinden aus SW, S oder SE, während 59,8% mit Rückenwinden aus

NW, N oder NE in die S-Reuse gerieten (Abb. 24d, 24f). Nicht zuletzt angesichts der Tatsache, daß

SW die häufigste Windrichtung ist, könnte dieses Zahlenverhältnis Ausdruck einer gezielten Auswahl

in südliche Richtungen führender Windrichtungen sein. Dafür könnte auch die Relation der Fangzah-

len mit der, allerdings nur wenig auffälligen, Überzahl der S-Flieger sprechen (S-Reuse: N-Reuse =

1,7:1;n = 5964). Allerdings fand die einzige andernorts beobachtete zahlenreiche Wanderung von

P. clypeatus bei starkem Südwind statt, so daß die Insekten nach Norden verdriftet wurden (Schmid

1987). Die Frage, ob die Dismigrationen von c/ypeatus zu einer gleichmäßigen Streuung im Raum füh-

ren oder durch Wahl bestimmter Windrichtungen gerichtete Komponenten enthalten (vgl. 4.5.4;

Abb. 28), muß vorläufig offen bleiben (siehe dazu Gatter 1981 c).

Die Flugaktivität von P. clypeatus beschränkt sich am RM weitgehend auf relativ windarme Zeiten

mit Windstärken zwischen 1 und 3 Bft. Zwischen den beiden Reusen ist kein Unterschied zu erkennen

(Abb. 25a, 25c).

3.4.5 Das tageszeitliche Windangebot und seine Auswirkungen auf die Wanderaktivität von

Episyrphus balteatus und Platycheirus clypeatus

Insekten wandern bevorzugt bei Hochdruck-(= Strahlungs-)wetter. Bei solchen Wetterlagen ist

am Randecker Maar (und nicht nur dort) eine typische tageszeitliche Drehung der dann im allgemei-

nen nur schwachen bis mäßigen Winde festzustellen (Gatter 1981a, 1981b mit zahlreichen Einzelhei-

ten). Wie auch ın Tab. 4 (3.4.1) deutlich zu erkennen ist, herrschen in den Morgenstunden Winde aus

südlichen Richtungen vor. Um die Mittagszeit dreht der Wind über West und bläst nun aus nördlichen

Richtungen.

balteatus clypeatus 2 Bft.

}

E ] Int

s 4.0 0 2 090 0 10 2% 30 4 50%

6 Bft.

3 b

OT OT HTM S 020050

6 Bft.

3 c

50 40 30 20 10 0 %0 20 30 0 5 40 30 20 0 20 30 40 50%

Abb. 25: Windstärken bei Flugbewegungen von Episyrphus balteatus und Platycheirus clypeatus.

Fig. 25: Wind forces during migration of Episyrphus balteatus and Platycheirus clypeatus.

a) Beide Arten/both species, 1986, S-Reuse/S-trap, n (balteatus) = 2937, n (clypeatus) = 3485

b) E. balteatus, 1981, S-Reuse/S-trap (n= 4244) und N-Reuse/N-trap (n=355)

c) P. clypeatus, 1981, S-Reuse/S-trap (n = 1528) und N-Reuse/N-trap (n = 987)

Für Episyrphus balteatus bedeutet das, daß die Art vormittags niedrig und nachmittags in höheren

Luftschichten zieht. Damit wird deutlich, daß der bei dieser Art besonders ausgeprägte Vormittags-

gipfel (Abb. 15e, 15h) in der täglichen Verteilung nicht auf eine weitgehende Beschränkung der Zug-

aktivität auf die Vormittagsstunden zurückgehen muß, sondern vermutlich nur eine Folge der tägli-

chen Winddrehung ist: vormittags herrscht gewöhnlich Gegenwind und die niedrig ziehenden Fliegen

geraten in die Reuse, nachmittags überfliegen die mit Rückenwind ziehenden Insekten die Fangein-

richtung. Diese Strategie, mit den wechselnden Windrichtungen die Zughöhe zu ändern, ist wahr-

scheinlich einer der bedeutenden Grundpfeiler zur Ermöglichung transkontinentaler Insektenwande-

rungen (Gatter 1981a: 40ff, 1981 bb, Gatter & Gatter 1990).

Anders sind die Verhältnisse bei Platycheirus clypeatus. Diese Art weist im tageszeitlichen Verlauf

einen ausgeprägten Nachmittagsgipfel auf (Abb. 5e, 5h). Auch hierfür könnte das tageszeitlich wech-

selnde Windangebot verantwortlich sein. Während der Vormittagsstunden, an denen meist schwache

Winde aus südlichen Richtungen wehen, werden weniger clypeatus gefangen. Nachmittags fördern

leichte nördliche Winde die Südwanderung dieses Dismigranten und steigern damit auch die Fangzah-

len.

N N N

W E W E W E W E

S a S b c

S S q

N N

W E W E W E

e S f S g

„N N

W E W E W E W E

S ——

— S l

S— S ee

h k

Abb. 26: Windrichtungen an Hauptzugtagen. Für alle Individuen einer Art (= 100 %) sind die Windrichtungen an

den jeweils vier individuenstärksten Tagen jeden Jahres dargestellt. An der Länge der Linien ist ablesbar, wieviele

Individuen der Art bei welcher Windrichtung in der S-Reuse gefangen wurden. Der Maßstab rechts unten ent-

spricht 10 %.

Fig. 26: Wind direction at the four main days of migration in each year. The length of each of the eight lines is cor-

related with the number of individuals of the species which were caught in the S-trap (trap with opening to the

north) with the wind blowing from the direction to which the line points. Scale below on the right: 10 %.

a) Syrphus spec., n= 2013 g) Helophilus trivittatus, n= 102

b) Metasyrphus corollae, n = 3869 h) Platycheirus albimanus, n=418

c) Scaeva pyrastri, n= 209 i) Platycheirus clypeatus, n= 4612

d) Episyrphus balteatus, n= 13666 k) Platycheirus manıcatus, n= 322

e) Sphaeropharia scripta, n = 4420 l) Melanostoma mellinum, n = 4342

f) Eristalis tenax, n= 317

3.4.6 Zusammenfassung: Windabhängige Wanderstrategien Saisonaler Migranten und

Saisonaler Dismigranten

3.4.6.1 Saisonale Migranten

Die exemplarisch am Beispiel Episyrphus balteatus dargestellte Strategie der Nutzung günstiger und

Vermeidung ungünstiger Winde läßt sich auch bei anderen Saisonalen Migranten beobachten. Wie

balteatus wurden auch Syrphus spec., Metasyrphus corollae, Scaeva pyrastri, Sphaerophoria scripta,

Eristalis tenax, Helophilus pendulus und H. trivittatus zu einem sehr deutlich überwiegenden Teil bei

Gegenwinden gefangen (Abb. 26-9).

Diese am RM gewonnenen Ergebnisse lassen sich auf andere Orte und Situationen übertragen. Ty-

pisch ist z. B. die Bemerkung bei Snow & Ross (1952), die in den Pyrenäen südwärts gerichteten In-

sektenzug beobachteten: „Syrphids were notnnoticed until the wind got up. From then on they crossed

in increasing numbers, flying at 0 to 2 feet.“ Nachdem vorher Windstille geherrscht hatte, hatte ein

langsam immer stärker werdender Südwind eingesetzt. Die Schwebfliegenwanderungen verlagerten

sich deshalb in Bodennähe. Auch in den Alpen werden Massenwanderungen immer bei (oft föhnartig

starken) Gegenwinden auffällig (z. B. Burmann 1978, Gepp 1975, Harz 1965, Jeekel & Overbeek

1968, Prell 1925). Nur dann fliegen die Syrphiden in Massen in Bodennähe und stauen sich an Pässen.

Eindrucksvoll ist z. B. die Schilderung von einer Migration über den Weit-Riß-Paß (2350 m) bei Prell

(1925), die sich v. a. auf Eristalıis tenax, aber auch auf Episyrphus balteatus bezieht: „Ununterbrochen

kamen in geradem Flug aus der Richtung des Melchtales Insekten herangezogen, welche deutlich ge-

gen den von Süden her wehenden Föhn ankämpfend über den Grat hinwegzukommen versuchten.

Handhoch bis meterhoch über die Felskante hinwegstreichend, die höchsten etwa mannshoch flie-

gend, kamen die Tiere scharenweise heran. Deutlich konnte man sehen, daß manche sich bemühten,

den Windschatten hinter dem Grat auszunutzen und dann unter Anspannung aller Kraft den Grat

selbst zu überwinden. ... Nicht ein einziges Tier wurde während der ganzen Zeit in entgegengesetzter

Richtung fliegen gesehen.“

Nicht nur im Gebirge, sondern auch an Küsten wurden große Schwebfliegenkonzentrationen v. a.

bei Gegenwinden beobachtet (z. B. Mackworth-Praed 1929, Svensson & Janzon 1984).

Wie Syrphiden die in größerer Höhe stärkeren Rückenwind nutzen, hat Gatter (1981a: 51) anschau-

lich geschildert: „Große Arten der Gattung Eristalis, Scaeva usw. können mit einem normalen Fern-

glas etwa 200 m weit verfolgt werden. Dabei zeigt sich, daß das Hochsteigen bei Rückenwinden auch

hier durch Drehung gegen den Wind geschieht. Die Fliegen, die mit Rückenwinden im bodennahen

Bereich Geschwindigkeiten von 40 bis 56 Stundenkilometern erreichen können, drehen sich in

einem weiten Bogen gegen den Wind, werden dabei ca. 10 bis 30 m hochgerissen und fliegen dann mit

dem Wind weiter. Am 20.9.1980, einem sehr heiffen Sommertag mit günstigem Nordostwind um

15 km/h, verließen die wenigen ziehenden Tiere, die in Bodennähe flogen, fast alle auf diese Weise die

tiefere untere Zone. In 1200 bis 1500 m über Grund (ca. 2000 m NN) herrschte dagegen reges Zugge-

schehen wohl eben dieser Arten nach Südwest. Bei einer Eigengeschwindigkeit von 25 km/h und einer

Windgeschwindigkeit von 17 km aus Nordost zogen die Fliegen mit ca. 40 km/h... Das Hochsteigen

hat hier wohl auch den Zweck, eine drohende Überhitzung zu vermeiden. In 2000 m lag die Nachmit-

tagstemperatur an diesem Tag bei 12,8° (...). Die Schwebfliegen fanden hier stoffwechselphysiolo-

gisch günstigere Bedingungen zur Migration vor als im 28° warmen Tiefland.“

Insektenzug in großer Höhe ist natürlich nicht leicht zu beobachten. Gute Indikatoren sind in sol-

chen Höhen jagende Vogelschwärme (vgl. Gatter 1981a: 53f., Spreadbury 1960). Glick (1939, 1942,

1960) und Glick & Noble (1961) (zitiert in Johnson 1969) fingen mit Hilfe von Flugzeugen Schweb-

fliegen in Höhen bis zu 5000 Fuß.

3.4.6.2 Saisonale Dismigranten

Bei den Dismigranten wie Platycheirus clypeatus, P. albimanus, P. manicatus oder Melanostoma

mellinum ist die Streuung der Windrichtungen beim Fang in der S-Reuse größer als bei den Saisonalen

Migranten (Abb. 26h-1). Häufiger als diese geraten die Dismigranten bei Rückenwinden in die

S-Reuse — ein Hinweis darauf, daß die oben für Saisonwanderer geschilderte, windabhängige Wahl

1000

clypeatus

2000 100

albimanus

75 80 SEeeBst-

2000

1000

2000

10.000

3000

manıcatus

100 1000

mellinum

50 peltatus ]

75 80 - - 85 - 75 80 - 85

3000

500 Syrphus 500 corollae ri Dypasiki

75 80 - 85 75 80 = 85 75 80 - 85

1000

2000

1000

50 selenitica | balteatus 500 scripta

75 80 = 85 75 80 = 85 75 80 S 85

1000

S m

50 pendulus 20

trivittatus

75 80 = 85 75 80 - 85

50 pagana

75 80 SM=3852=

Abb. 27: Fangsummen in der S-Reuse (unten) und der N-Reuse (oben) in den Jahren 1975— 1987. Der Pfeil gibt den

Zeitpunkt der Verdoppelung der Größe der S-Reuse an (1978). Zur besseren Vergleichbarkeit sind die verdoppelten

Fangzahlen der Jahre 1975-1977 gebrochen dazugezeichnet. Jahre ohne Erfassung der jeweiligen Art sind in der

Zeitspalte mit — gekennzeichnet.

Fig. 27: Numbers of Syrphidae caught in the S-trap (trap with opening to the north; below) and the N-trap (opening

to the south; above) in 1975— 1987. The arrow indicates the date when the S-trap size was doubled (1978). For better

comparison, the diagram shows double numbers of 1975— 1977 in broken lines. Years without identification of the

respective species are marked with — .

a) Syrphidae insgesamt/total g) Syrphus spec. n) Erıstalis tenax

b) Platycheirus albimanus h) Metasyrphus corollae o) Helophilus pendulus

c) Platycheirus clypeatus i) Scaeva pyrastri p) Helophilus trivittatus

d) Platycheirus manicatus k) Scaeva selenitica q) Cheilosia pagana

e) Platycheirus peltatus l) Episyrphus balteatus

f) Melanostoma mellinum m) Sphaerophoria scripta

der Zughöhen von Dismigranten nicht im selben Ausmaß durchgeführt wird. Dismigranten wandern

zwar hauptsächlich mit dem Wind. Die trotz allem hohen Zahlen der bei Gegenwinden gefangenen

Fliegen lassen aber ebenso wie die bei allen Arten zu beobachtenden zahlenstärkeren Fänge in der

S-Reuse vermuten, daß auch sie im Herbst eine Tendenz haben, in südliche Richtungen zu wandern

und dabei auch Gegenwindbewegungen durchführen können (vgl. 4.5.4). Diese Frage bedarf weiterer

Klärung.

3.5 Populationsdynamik

Tab. 1 und Abb. 27a zeigen die relativ starken Schwankungen der jährlichen Fangzahlen am RM.

Noch extremer sind die Häufigkeitsunterschiede von Jahr zu Jahr bei einzelnen Arten (Abb. 27b-q).

Ähnlich stark wie am RM sind diese Schwankungen am CB (Aubert et al. 1976). Sie sind auch überall

im Freiland zu beobachten.

Die Zahl der Jahr für Jahr in den Reusen gefangenen Insekten wird natürlich von sehr vielen Fakto-

ren beeinflußt. Wichtig ist mit Sicherheit der konkrete Witterungsverlauf während einer Fangperiode.

Allerdings ist das Wetter nicht direkt und allein für die schwankenden Fangzahlen verantwortlich.

Wie der unterschiedliche Verlauf der Kurven einzelner Arten zeigt (s. u.), spiegeln die Diagramme

auch die tatsächliche Häufigkeit der einzelnen Arten in den betreffenden Jahren wider.

Ausschlaggebend für den Populationsaufbau bei aphidophagen Formen — und dazu gehören fast

alle Saisonalen Migranten unter den Schwebfliegen (vgl. Tab. 3) — ist in erster Linie das Blattlausange-

bot im Frühjahr und Frühsommer. Die Diagramme für die aphidophagen Saisonwanderer Syrphus

spec., Metasyrphus corollae, Scaeva pyrastri, S. selenitica, Episyrphus balteatus und Sphaerophoria

scripta zeigen allerdings keinen einheitlichen Verlauf: im Jahr 1980 erschienen z. B. nur wenige baltea-

tus, corollae, pyrastri und scripta, während Syrphus spec. und Scaeva selenitica ausgesprochene Ma-

xima aufwiesen. Auch die Saisonalen Dismigranten Platycheirus albimanus, P. manicatus und P. pel-

tatus zeigen ausgeprägte Höhepunkte. Alle genannten Arten sind im Larvenstadium Blattlausfresser

mit weitem Beutespektrum, die auch weitgehend dieselben Habitate besiedeln (vgl. 3.1; Rotheray &

Gilbert 1989). Die Zahlen lassen vermuten, daß sich ihre Populationen gegenseitig über (Nahrungs-)

Konkurrenz beeinflussen. Nachgewiesen ist, daß Syrphus ribesii Blattlauskolonien effektiver nutztals

Melanostoma scalare (Rotheray 1983) und daß die Mortalität der Larven von Metasyrphus corollae bei

der Anwesenheit von rıbesii-Larven steigt (Benestad Hägvar 1972).

Auch aquatische Formen können starke Schwankungen aufweisen (Abb. 27n—p). Ihre Diagramme

lassen jedoch keine gegenseitige Beeinflussung erkennen.

4. Diskussion

4.1 Vergleich der Phänologie der Schwebfliegenwanderungen am Randecker Maar

und am Col de Bretolet

Bei zahlreichen Arten ist, wie in den Artabschnitten jeweils kurz erwähnt (vgl. 3.1), der Zughöhe-

punkt am Col de Bretolet deutlich später als am Randecker Maar. Besonders klar ist dieser Unter-

schied bei Episyrphus balteatus (Abb. 15a): An beiden Stationen steigen die Fangzahlen in der zweiten

Julihälfte stark an. Während der Durchzug am RM aber nach einem ausgeprägten Höhepunkt Anfang

August in der zweiten Monatshälfte so stark abflaut, daß im September nur noch wenige balteatus (ca.

5% der Gesamtsumme) gezählt werden, ist am CB nach dem ebenfalls in der ersten Augusthälfte er-

reichten Maximum bis Mitte Oktober starker Durchzug festzustellen. Der Median, am RM der 7.8.,

ist also am CB weit nach hinten verschoben. Auch im Geschlechterverhältnis besteht ein gravierender

Unterschied: am RM sind JO’ und 29 gleich stark vertreten, am CB werden fast ausschließlich PP

gefangen (vgl. 4.2).

Der Col de Bretolet liegt ca. 350 km südwestlich des Randecker Maars. Für den Brachpieper, einen

Zugvogel, dessen Winterquartiere südlich der Sahara liegen, errechnete Gatter (1970) eine Verschie-

bung des Herbstzugsmedians von zehn Tagen zwischen Kirchheim (am RM) und dem Col de Bretolet

und damit eine tägliche Vorrückstrecke von ca. 35 km. Eine solche Tagesetappe ist auch für Schweb-

fliegen mit Sicherheit gut zu bewältigen: trotz ungünstiger Wetterbedingungen fingen Aubert et al.

(1969) am Col du Glandon Schwebfliegen, die sie drei Tage zuvor am 111 km entfernten Col de Bre-

tolet markiert hatten. Die Medianverschiebung zwischen RM und CB dürfte bei Syrphiden genau wie

beim Brachpieper eine Folge der Lagebeziehungen der beiden Stationen sein: die in Zugrichtung nach-

geschaltete Beobachtungsstation am CB wird von der Hauptmasse der Durchzügler später erreicht.

Auch das Ende der Zugperiode wird von der Lage natürlich beeinflußt. Beim Brachpieper wie bei Epi-

syrphus balteatus endet die Zugzeit am CB wesentlich später als am RM.

Ähnlich deutlich wie bei balteatus ist die Verschiebung der Maxima und Verlängerung der Flugzeit

bei den Saisonalen Migranten Metasyrphus corollae (S. 35), Meliscaeva cinctella (S. 43), Eristalis tenax

(S. 61) und Helophilus trivittatus (S. 66) und bei dem Saisonalen Dismigranten (?) Platycheirus mani-

catus (S. 26).

Mehrere Arten zeigen zwar eine Verlängerung der Flugzeit, aber keine Verschiebung des Maxi-

mums (bzw. sind am CB keine deutlichen Maxima ausgeprägt), so die Saisonalen Migranten Syrphus

spec. (S. 33), Scaeva pyrastri (S. 38) und S. selenitica (S. 41) und der Saisonale Dismigrant Melano-

stoma mellinum (S. 29) sowie der Saisonale Dismigrant (?) Parasyrphus lineolus (S. 42).

Weitgehende Entsprechung der Phänologie beider Orte weist der Saisonale Migrant Sphaerophoria

scripta (S. 52) auf; das könnte auf kürzere Migrationsentfernungen schließen lassen.

4.2 Veränderungen der Zahlenverhältnisse der Geschlechter im Verlauf der Migration

Balteatus-O'O haben eine wesentlich geringere Lebenserwartung als O9. „Am 49. Gesamtlebens-

tag (d. h. ca. 4 Wochen nach dem Schlupftermin) waren bereits 50% der O’C' tot... Die Hälfte der

QQP wurde 60 Tage alt. Dieses Alter erreichten nur ca. 25% der O’O', während von den Q9 25% (13

von 52 Individuen) ein Alter von 80 Tagen und 9 Individuen sogar ein Alter von über 100 Tagen er-

langten. Eine deutlich erhöhte Sterblichkeit führte bei den O’C’ schon ab dem 55. Gesamtlebenstag zu

einer raschen Abnahme der Zahl der Überlebenden. Die durchschnittliche Lebensdauer ... war für die

QQP mit 39,5 Tagen ebenfalls deutlich höher als die der G'C’ mit 23,5 Tagen“ (Geusen-Pfister [1987]

nach Versuchen unter Gewächshausbedingungen). Dadurch erklärt sich die Beteiligung beider Ge-

schlechter von balteatus an den Fängen im RM (wie übrigens auch an den norddeutschen Feuerschif-

Tabelle 5: Geschlechter-Verhältnis von Wanderschwebfliegen an Feuerschiffen der Nord- und Ostsee (nach Hey-

demann 1967), am Randecker Maar und am Col de Bretolet (nach Aubert 1962).

Tab. 5: Sex ratios of migratory hoverflies at lightships in the North Sea and the Baltic Sea, at Randecker Maar and

at Col de Bretolet.

Art Feuerschiffe Randecker Maar Col de Bretolet

SL... @ SELRERU) SCH TERT)

P. albimanus 1:0,96 (165) 1:2,82 (833) 1:20,25 (85)

Pemanicasıs 1:0,44 (13) 1:1,18 (550) 1:0,11 (40)

M. mellinum 1:0,10 (34) 1:3,58 (11524) 1:2,34 (1535)

S. ribesi 1:2,00 (36) 1:8,00 (9)

S. torvus 1:1,86 (4030) 16 292 (16)

S. vitripennis 1:1635 (7) 68 29 (68)

M. corollae” SZ (526) 1811,07 (7545) 3072. (142)

M. lunıger 1:16,00 (17)

M. auricollıis 1:61,00 (62)

E. balteatus 1E0,99 (507) 1E0,92 (27135) 1:12,02 (1569)

S. scripta 1:1,71 (11204) 1:3,98 (209)

E. tenax 1:1,32 (511) 1611634 (1230)

= In Südschweden Anfang August 1:1,6 (n=4338) (Svensson & Janzon 1984).

fen [Heydemann 1967]) und das weitgehende Fehlen der Q'O’ am CB. Der CB mit seinem wesentlich

später liegenden Durchzugsmedian wird von O’O’ kaum mehr erreicht. Die meisten J’O’ sterben vor-

her. — (Leider liegen vom CB nur wenige Stichproben zum Geschlechterverhältnis vor [Aubert 1962],

ohne daß eine genaue zeitliche Zuordnung der Daten möglich ist. Zu erwarten wäre, daß ein zu Beginn

der Zugzeit weitgehend ausgeglichenes Verhältnis sich schnell zugunsten der 2 verschiebt. Am RM

wird jedenfalls deutlich, daß balteatus-J'O’ ım September praktisch nicht mehr vertreten sind [vgl.

Abb. 15b].)

Die Verschiebung des Geschlechter-Verhältnisses ist wie die Medianverschiebung zwischen RM

und CB ein Beleg dafür, daß sich eine Zugwelle wenigstens über das südliche Nord- und über Mittel-

europa hinweg bewegt.

Für einige häufigere Arten liegen Daten zum Geschlechter-Verhältnis von mehreren Orten vor

(Tab. 5).

Für die Saisonalen Wanderer Syrphus spec., Metasyrphus corollae, Episyrphus balteatus und Sphae-

rophoria scripta zeigt sich zwischen RM und CB durchgehend eine Verschiebung des Verhältnisses

zugunsten der Weibchen. Für alle diese Arten ist, wie oben für balteatus geschildert, anzunehmen, daß

im Verlauf einer großräumigen Migration die O’CO’ absterben. Das gilt auch für E. tenax, selbst wenn

hier als Folge der sehr späten Flugzeit das Geschlechterverhältnis zwischen RM und CB keine Unter-

schiede aufweist. Auch bei dieser Art dominieren zu Beginn der Zugzeit am CB im August und An-

fang September die O’CO’, später die PP (Aubert 1962); wie bei den anderen Saisonalen Migranten

werden also auch hier die S’C’ im Herbst seltener. Die Daten für die am RM seltenen Saisonwanderer

Meliscaeva auricollis und Metasyrphus luniger zeigen mit ihrem extremen Q Q-Überschuß am CB, daß

die Verhältnisse denen von E. balteatus wohl weitgehend entsprechen.

Während die aus Farbschalenfängen von norddeutschen Feuerschiffen (Juli/August) gewonnenen

Angaben im wesentlichen den Daten vom RM entsprechen, liefen sich die am CB beobachteten Zah-

lenverhältnisse auch andernorts in den Alpen bestätigen. Jeekel & Overbeek (1968) beobachteten am

9.8.1967 in Tirol für die Saisonalen Migranten Episyrphus balteatus, Sphaerophorıa scripta, Syrphus

torvus und $. vitripennis eine starke Dominanz der PP. Auch die von Burmann (1978) ebenfalls aus

Nordtirol mitgeteilten Daten lassen ein starkes Überwiegen der Q 9 bei Episyrphus balteatus, Syrphus

ribesüi, Metasyrphus corollae und Sphaerophoria scripta erkennen.

Demnach sind die oben für Episyrphus balteatus dargestellten Verhältnisse wohl für die ganze

Gruppe der Saisonalen Migranten unter den Schwebfliegen gültig.

Unter der Gruppe der Dismigranten (Platycheirus albimanus, P. manicatus und Melanostoma mel-

linum) sind die Verhältnisse weit weniger klar. Die Daten lassen aufgrund des geringen Umfanges der

meisten von anderen Orten vorliegenden Stichproben auch kaum weitergehende Deutungen zu. Mit

Sicherheit besteht aber der für die Saisonalen Migranten typische, auf weitreichende Wanderungen

hinweisende Unterschied zwischen dem Geschlechterverhältnis an den einzelnen Beobachtungssta-

tionen nicht. Der häufig zu beobachtende P Q-Überschuß geht auf die höhere Dismigrationsbereit-

schaft der PP zurück.

4.3 Biologie der Wanderschwebfliegen

Aus Tab. 3 (vgl. 3.3) wird zweierlei deutlich: (1) alle Saisonmigranten sind entweder aphidophag

oder aquatisch sapro-/microphag (vgl. Schmid & Gatter 1988). Unter den anderen trophischen Grup-

pen findet sich keine Art, die saisonale Migrationen durchführt und (2) sämtliche Schwebfliegenarten,

die nicht als Larve, sondern als Imago überwintern, gehören zur Gruppe der Saisonalen Migranten,

ebenso die als Puparium überwinternden polyvoltinen Arten. Diese beiden Ergebnisse werden im fol-

genden diskutiert.

4.3.1 Arten mit phytophagen und terrestrischen saprophagen Larven

Für eine Art wie beispielsweise Cheilosia fasciata Schiner & Egger 1853, deren Larven als Nah-

rungsspezialisten in Blättern des Bärlauchs (Allium ursinum) minieren, ist der Vorteil, nicht in größe-

rem Ausmaß zu wandern, deutlich: die Nahrungsquelle steht mit Sicherheit im nächsten Jahr am sel-

ben Ort wieder zur Verfügung. Ähnlich spezialisiert sind, soweit bekannt, sehr viele phytophage

Schwebfliegen z. B. der Gattungen Cheilosia (z. B. Smith 1979, Rotheray 1988), Merodon oder Eume-

rus. Die Überwinterung findet geschützt in der Erde statt. Damit wird plausibel, warum es unter den

phytophagen Schwebfliegen keine Wanderarten gibt.

Ähnliches dürfte für die Gruppe der terrestrischen saprophagen Syrphiden gelten. Auch sie beuten

eine Nahrungsressource aus, die im nächsten Jahr mit großer Wahrscheinlichkeit in der Nähe wieder

verfügbar ist. Mit Sicherheit bewegen sich die meisten dieser Arten vom Geburtsort weg; die Distan-

zen sind aber fast immer sehr gering. Lediglich einige Arten der Gattung Xylota führen Saisonale Dis-

migrationen aus, die etwas weiter führen (Tab. 3).

4.3.2 Arten mit zoophagen Larven

Aphidophage Schwebfliegen stehen dagegen vor dem Problem, daß ihre Larven eine Nahrungs-

quelle benötigen, die in Raum und Zeit oft unvorhersagbar auftritt. Hier helfen in den Lebenszyklus

integrierte Dismigrations- oder Migrationsphasen bei der Auffindung und Ausnutzung solcher Res-

sourcen.

Sehr viele aphidophage Schwebfliegen sind monovoltine Frühjahrsarten, die zudem oft mehr oder

weniger spezialisiert sind. Ihre Flugzeit deckt sich mit dem Frühjahrsmaximum der Blattlausentwick-

lung (vgl. Dusek & Läska 1986). Für einige dieser Arten (Parasyrphus) konnte oben nachgewiesen

werden, daß sie Dismigrationen durchführen. Es ist wahrscheinlich, daß sehr viel mehr dieser Arten

dismigrieren. Durch ihre frühe Flugzeit entgehen sie aber der Erfassung sowohl am RM wie am CB.

Saisonale Dismigranten sind, neben diesen monovoltinen Frühjahrsarten, vor allem Arten der in-

nerhalb der Zoophagen phylogenetisch ursprünglichen Gattungen (Rotheray & Gilbert 1989) Platy-

cheirus und Melanostoma. Bei beiden Gruppen stehen Wanderungen nicht im Zusammenhang mit

dem Aufsuchen von Winterquartieren, sondern dienen in erster Linie dem Auffinden von Eiablage-

plätzen. Bei den monovoltinen Frühjahrsarten erleichtert das Frühjahrsmaximum der Blattläuse diese

Suche, bei Platycheirus und Melanostoma ıhre weite ökologische Potenz. Fast alle Arten dieser beiden

Gattungen ziehen zwar Blattläuse als Nahrung vor, sind aber auch sehr oft in der Bodenstreu zu fin-

den, wo sie Jagd auf andere Arthropoden machen. Sie entgehen damit während des Soemmerminimums

der Blattläuse der Konkurrenz durch die aphidophagen, polyvoltinen Saisonwanderer. Rotheray

(1983) konnte zeigen, daß Syrphus ribesii Blattlauskolonien effektiver nutzen kann als Melanostoma

scalare.

Die Saisonalen Migranten unter den Aphidophagen sind alle polyvoltin und, was die Nahrung und

das Habitat (vgl. Chandler 1968b, Läska & Stary 1980) betrifft, zwar spezialisierter als die meisten Pla-

tycheirus- und Melanostoma-Arten, aber gewöhnlich wesentlich weniger spezialisiert als die unıvolti-

nen Arten. Sie stellen während des sommerlichen Minimums der Blattlausentwicklung die Haupt-

masse der Schwebfliegen (vgl. Dusek & Läska 1986). Ihre ausgeprägte Migrationsneigung kann (ge-

meinsam mit ihrem weiten Nahrungs- und Habitatspektrum) einerseits als Anpassung an die Not-

wendigkeit verstanden werden, die in dieser Jahreszeit nur spärlich vorhandenen Blattläuse aufzufin-

den, andererseits steht sie, zum mindesten bei den Formen, die den Winter als Puparien oder Imagines

überdauern, in engem Zusammenhang mit der Überwinterung. Für Episyrphus balteatus, die am be-

sten bekannte Art, wurde der adaptive Wert der spätsommerlichen und herbstlichen Wanderung ın

südliche Gebiete bereits diskutiert (S. 52). Es ist anzunehmen, daß das dort Gesagte auch auf die ande-

ren Arten übertragen werden kann. Demnach ist einerseits im nördlichen Europa Überwinterung

nicht oder nur sehr eingeschränkt möglich; auch in Mitteleuropa ist das Überwinterungsrisiko für

diese Arten beträchtlich, weshalb große Teile der Populationen im Herbst nach Süden ziehen. Ande-

rerseits finden die Larven (möglicherweise auch die Imagines) im Frühjahr und Sommer in Mittel-

europa bessere Entwicklungs- und Lebensbedingungen als in Südeuropa vor. Es bleibt zu prüfen, ob

die Wanderungen auch für die den Winter als diapausierende Larven überstehenden Saisonwanderer

Syrphus spec. und Sphaerophoria scripta einen ähnlichen adaptiven Wert besitzen.

Zusammenfassend (und vereinfachend) lassen sich also unter den zoophagen Schwebfliegen drei

Migrationsstrategien erkennen:

— Unspezialisierte, zoophage, polyvoltine Arten, die Saisonale Dismigrationen ausführen, um geeig-

nete Eiablageplätze aufzufinden: Arten der Gattungen Platycheirus und Melanostoma.

— Etwas stärker spezialisierte, aphidophage, polyvoltine Arten, die Saisonale Migrationen durchfüh-

ren, um einerseits Nahrungs- und Eiablageplätze, andererseits günstige Überwinterungsplätze zu

finden: Arten siehe Tab. 3.

— Spezialisierte, monovoltine Arten, die Dismigrationen durchführen, um geeignete Eiablageplätze

aufzufinden: Arten der Gattung Parasyrphus und vermutlich noch viele weitere Species.

4.3.3 Arten mit aquatischen sapro-/microphagen Larven

Larvenhabitate dieser Gruppe sind Gewässer — vom offenen Teich bis zur wasserdurchtränkten

mulmigen Baumhöhlung. Vor allem offene Kleingewässer können wetterabhängig leicht verschwin-

den; für manche Arten ist das Angebot an Larvenhabitaten somit vielfachen Schwankungen ausge-

setzt.

Unter den Arten der aquatischen Gruppe sind verschiedene Strategien erkennbar. Sehr viele Species

wandern nicht, so z. B. die häufige Myathropa florea (Linnaeus 1758), deren Larven in Kleinstgewäs-

sern zwischen Baumwurzeln o.ä. leben. Manche Arten wandern nur zu einem mehr oder weniger gro-

ßen Teil, so Eristalis arbustorum und E. pertinax. Sie verfolgen also eine „Doppelstrategie“, bei der

der Schwerpunkt bei der Überwinterung von Larven im Heimatgebiet ist. Bei Helophilus pendulus

und /. trivittatus ist die Wanderneigung dagegen sehr viel ausgeprägter. Für pendulus lassen die zahl-

reichen Beobachtungen im frühen Frühjahr vermuten, daß ein beträchtlicher Anteil als Larve in Mit-

teleuropa überwintert; bei trivittatus sind Frühjahrsbeobachtungen so spärlich, daß angenommen

werden muß, daß der Aufbau mitteleuropäischer Populationen überwiegend auf Immigranten zu-

rückgeht.

Die einzige Art dieser Gruppe, die als Imago überwintert, ist Eristalis tenax. Für das Überwinte-

rungsrisiko in Mitteleuropa dürfte dasselbe gelten wie bei den aphidophagen, als Imagines überwin-

ternden Schwebfliegen (4.3.2). Im Mittelmeergebiet überwinternde Tiere haben vermutlich wesent-

lich größere Überlebenschancen. Dagegen bietet im Sommer das humide Mitteleuropa den wasser-

lebenden tenax-Larven mit Sicherheit mehr und günstigere Entwicklungschancen als das trockene

Südeuropa. — Vermutlich spielen beim Aufbau der Sommerpopulationen in Mittel- und Nordeuropa

hier überwinternde tenax-QP gegenüber den Immigranten in den meisten Jahren nur eine unterge-

ordnete Rolle.

4.4 Zur Bedeutung der Wanderschwebfliegen bei der biologischen Schädlingsbekämpfung

Unter den aphidophagen Insekten kommen Marienkäfern (Coccinellidae), Florfliegen und Blatt-

lauslöwen (Chrysopidae und Hemerobiidae) und Schwebfliegen besondere Bedeutung zu (Rotheray

1989). Nach Versuchen von Sundby (1966) hat dabei die Larve der Schwebfliege Syrphus ribesii eine

größere Fraßleistung als die des Marienkäfers Coccinella septempunctata L. und die der Florfliege

Chrysoperla carnea St. Die hohe Nachkommenzahl der Florfliege läßt diese (im Labor) als den beiden

anderen Arten weit überlegenen Blattlausvertilger erscheinen. Außerhalb des Labors ist es von vielen

Faktoren abhängig (beteiligte Arten, Luftfeuchtigkeit, Temperatur etc.; vgl. Bastian 1986), welche der

drei Insektengruppen als effektivster Prädator auftritt.

Unter den aphidophagen Schwebftliegen gibt es weitgehend spezialisierte Arten wie z. B. Neocne-

modon vitripennis (Meigen 1822), eine hauptsächlich von Tannenstammläusen (v. a. Dreyfusia piceae

Ratzeburg) lebende Art, die einen mit den Populationsmaxima der Beutetiere synchronisierten bivol-

tinen Lebenszyklus aufweist (Delucchi et al. 1957). Zahlreiche univoltine Frühjahrsarten unter den

Schwebfliegen nutzen das Frühjahrs- und Frühsommermaximum der Laus-Populationen. Viele die-

ser Arten sind hauptsächlich in Wäldern anzutreffen (Arten der Gattungen Dasysyrphus, Parasyrphus,

Epistrophe).

Die häufigsten aphidophagen Schwebtliegenlarven in einem von Bastıan (1984) untersuchten Koni-

ferenjungwuchs waren allerdings keine solchen „Waldarten“, sondern typische Wanderschwebflie-

gen. Dazu gehörten die Saisonalen Migranten Syrphus rıbesü, S. torvus, S. vitripennis, Episyrphus bal-

teatus und Sphaerophoria scripta und der Saisonale Dismigrant Melanostoma mellinum. Bastian (1984)

konnte zeigen, daß unter den Blattlausprädatoren Schwebtliegen in einem Jahr mit witterungsbedingt

stark verzögertem Aufbau der Läusepopulationen am raschesten auf diese veränderte Situation rea-

gierten.

Voraussetzung für diese Flexibilität ist einerseits die hohe Mobilität der Wanderschwebfliegen, an-

dererseits ihr polyvoltiner Lebenszyklus und ihre geringe Spezialisierung auf bestimmte Beutetiere.

Die polyvoltinen Arten sind nicht an eng begrenzte Fortpflanzungszeiten gebunden; sie nutzen Blatt-

lauskolonien auch im Sommer, wenn die Populationshöhe und Fortpflanzungsrate der Läuse einen

Tiefpunkt erreicht hat und damit auch der Einfluß von Prädatoren höher wird. Die Merkmalskombi-

nation mobil/polyvoltin/wenig spezialisiert macht gerade Wanderschwebtliegen zu forst- und land-

wirtschaftlich bedeutenden Blattlausvertilgern. Will man sie ökonomisch im Rahmen der biologi-

schen Schädlingsbekämpfung nutzen, ist eine sehr genaue Kenntnis ihrer individuellen Lebenszyklen

unerläßlich. Gerade dem Phänomen der Migrationen muß hierbei wesentlich mehr Aufmerksamkeit

als bisher geschenkt werden.

4.5 Allgemeine Aspekte der Schwebfliegenwanderungen

4.5.1 Verbreitung

Gatter (1981a) wies darauf hin, daß Wanderinsekten große Verbreitungsgebiete haben. Dies trifft

auch auf die Wanderschwebfliegen zu.

Von den 18 als Saisonale Migranten eingestuften Arten (Tab. 3) sind neun wenigstens holarktisch

verbreitet. Unter diesen Arten sind viele auch im nordorientalischen Bereich zu finden; eine Art (Eri-

stalis tenax) ist sogar kosmopolitisch. Sieben der 18 Arten sind wenigstens paläarktisch verbreitet.

Auch von diesen kommen die meisten auch noch nordorientalisch vor, das Areal von Metasyrphus co-

rollae reicht bis in die Paläotropis, das von Episyrphus balteatus schließt die australische Region mit

ein. Lediglich zwei der 18 Arten haben ein relativ kleines Verbreitungsgebiet: Meliscaeva aurıcollis

und die nur in geringer Zahl wandernde Eristalis pertinax scheinen auf die westliche Paläarktis be-

schränkt zu sein.

Auch unter den 17 Dismigrantenarten sind acht wenigstens holarktisch und sieben wenigstens pa-

läarktisch verbreitet. Zwei Arten (Platycheirus manicatus, Parasyrphus punctulatus) kommen nach

heutiger Kenntnis nur in der westlichen Paläarktıis vor.

Demgegenüber haben z. B. viele Arten der im Larvenstadium phytophagen, sehr artenreichen Gat-

tung Cheilosia, von denen für keine Species weitreichende Wanderungen nachgewiesen sind, recht be-

schränkte Verbreitungsgebiete.

4.5.2 Diapause und Migration

„Unter Diapause verstehen wir eine unter natürlichen Bedingungen mehrere Wochen, Monate oder

sogar Jahre andauernde Depression im Wachstum embryonaler Gewebe oder der Ovarien, für welche

die herrschende Temperatur und andere Außenfaktoren nicht allein verantwortlich gemacht werden

können“ (Schneider 1948), also eine endogen fixierte Ruheperiode (Sommerruhe, Winterruhe etc.) der

Larve oder der Imago.

Auf den engen Zusammenhang zwischen der Diapause von Imagines und Migrationen machte

zuerst Johnson (1969) aufmerksam. Beispiele aus dem Bereich der Schmetterlinge werden unten aus-

führlich dargestellt (4.5.4). Unter den Käfern (Coleoptera) ist der Marienkäfer Coccinella septempunc-

tata L. die eindrucksvollste Wanderart in Europa. Die Jungkäfer schlüpfen meist zwischen Mitte Juli

und Anfang August. In Gradationsjahren — die fast immer mit denen der aphidophagen Syrphiden

zusammenfallen — klettern die Jungkäfer an Halmen hoch und starten an geeigneten Tagen, manch-

mal zu Millionen, zur Migration (Gatter & Gatter 1973, Hodec 1973). Bereits wenige Tage nach der

kurzen Migrationsphase suchen die meisten Tiere Sommer- und Winterruhequartiere auf, die bis zum

Frühjahr nicht mehr verlassen werden. Diesen Prädiapauseflügen (Hagen 1962, Williams 1961) folgen

Nachdiapauseflüge ım Frühling.

Während bei den zoophagen, sich meist von Blattläusen ernährenden Marienkäfern (vgl. 4.4) Pol-

lennahrung nur sehr untergeordnete Bedeutung hat, können Syrphiden beim Vor- und Nachdiapau-

seflug unentwegt Pollennahrung zu sich nehmen. Dies könnte die Evolution der Langstreckenmigra-

tion ım Sinne Johnsons (1969) begünstigt haben.

Wie oben gezeigt wurde (Tab. 3; 4.3), führen auch sämtliche als diapausierende Imagines überwin-

ternden Schwebfliegenarten richtungsorientierte Saisonwanderungen aus, die im Herbst dem geziel-

ten Aufsuchen günstiger Überwinterungsareale dienen, während die Nachdiapausewanderungen im

Frühjahr in Gebiete führen, die gute Entwicklungsbedingungen gewährleisten. Direkte Hinweise auf

diese Frühjahrswanderungen sind zwar spärlich (3.1; Gatter 1980, Westmacott & Williams 1954), was

aber mit Sicherheit einerseits an der Unauffälligkeit der Tiere und ihrer im Frühjahr gegenüber dem

Herbst erheblich niedrigeren Zahl hängt, andererseits aber auch daran, daß diesem Phänomen bisher

kaum Aufmerksamkeit geschenkt wurde.

4.5.3 Generationenfolge und Migration

Gatter (1981a) untersuchte die mitteleuropäischen Schmetterlinge und stellte erstmals fest, daß es

neben den Arten mit Diapause im Imaginalstadium (4.5.2) die Arten mit mehreren Generationen sind,

die ausgeprägt migrieren. Bei den Schwebfliegen bestätigt sich dieses Ergebnis: sämtliche Saisonalen

Migranten bilden in der Tat mehrere Jahresgenerationen aus (3.1, 3.2, Tab. 3).

Andererseits wandern nicht alle Arten, die in mehreren Jahresgenerationen auftreten. Unter den

polyvoltinen Syrphiden haben sich, in Anpassung an die ökologischen Bedürfnisse einzelner Arten

und Artengruppen, verschiedene Strategien herausgebildet, von denen Migration nur eine ist (vgl.

4.3). Die ausgeprägtesten Wanderer sind polyvoltine Arten, die zudem eine Winterruhe (Diapause)

haben und vor und nach dieser Winterruhe wandern. Die sich im Jahresverlauf ergebenden Habitatän-

derungen und die unterschiedlichen Ansprüche während verschiedener Lebensphasen bringen die

Notwendigkeit zum Ortswechsel mit sich (Gatter 1981c, Gatter & Gatter 1990). Die weitere Erfor-

schung dieser vitalen Wanderer wird noch manches aufregende Ergebnis bringen.

4.5.4 Die Stellung der Schwebtliegen innerhalb der Wanderinsekten

Der Monarch Danans plexippus L. (Lepidoptera) gilt als das klassische Wanderinsekt schlechthin.

Die Herbstgeneration dieses in der Alten und Neuen Welt beheimateten Schmetterlings wandert in

Amerika aus Kanada und den Vereinigten Staaten im Extremfall bis in das mexikanische Winterquar-

tier, um dort gesellig auf bewaldeten Berggipfeln zu überwintern. Nach der Überwinterung wandern

die Falter nordwärts. Im Verlauf ihrer Wanderung (und nicht erst am Endpunkt) legen die @Q Eier

ab. Die Nachkommen der Überwinterungsgeneration erreichen dann wieder die Gebiete des nördli-

chen Amerika (Urquhart 1960). — Der Monarch wandert kompaßorientiert und erreicht seine Über-

winterungsplätze navigierend.

In Europa kennen wir innerhalb der Schmetterlinge ähnliche Fälle. Geselliges Überwintern ist aller-

dings nicht bekannt; so waren — wenn man von den methodisch unglücklichen Versuchen bei Roer

(z. B. 1961) absieht (Baker 1978) — bisher keine Massenkennzeichnungen möglich, die die Verfolgung

der Zugwege einzelner Individuen ermöglicht hätten. Mit Distelfalter Vanessa cardni L. (Harz 1975,

Baker 1972), Admiral Vanessa atalanta L. (Baker 1972, Gatter 1981a), Gammaeule Autographa

gamma L. (Baker 1978, Gatter 1981a) und anderen Arten haben wir Schmetterlinge, deren Migra-

tionszyklus dem des Monarchen Danaus plexippus ähnlich ist. Bei Distelfalter (Baker 1978), Winden-

schwärmer Agrıins convolvul L. (Gatter & Gatter 1990) und weiteren Arten können sich bei diesen

Wanderungen Verbindungen bis in den nordafrikanischen und afrotropischen Bereich ergeben. Ver-

gleichen wir diese Lebenszyklen und Flugleistungen mit denen von Episyrphus balteatus (S. 44), so

finden wir zahlreiche Parallelen: Die Herbstgeneration zieht wochenlang, möglicherweise über Mo-

nate hinweg zu bisher unbekannten Überwinterungsgebieten. In Mitteleuropa, den Alpen und den

Pyrenäen führt die Wanderung kompaßorientiert in südwestliche Richtung. Die ?Q offensichtlich

derselben Generation wandern im Frühjahr wieder nach Norden. Ihre Nachkommen erreichen

Nordeuropa, wo sie nicht überwintern können. Die Sommergeneration wandert wiederum nach

Süden.

Hinsichtlich ihrer physischen Leistungsfähigkeit sind Schwebfliegen vielen Wanderfaltern überle-

gen. Bei der Überwindung von Gebirgspässen ist z. B. der Kleine Kohlweißling Pieris rapae L. als Sai-

sonaler Migrant unseren Wanderschwebfliegen derselben Kategorie weder in der Vitalität noch in der

Fluggeschwindigkeit gewachsen. Ein Vergleich der Relation von Körpergröße/-gewicht zur Flügel-

fläche zwischen Episyrphus balteatus und Danaus plexippus macht wahrscheinlich, daß die Fluglei-

stungen dieser Schwebtliegenart denen des Monarchen kaum nachstehen. Ihre Flugleistungen und

-geschwindigkeiten werden wohl nur von wenigen Libellen (Odonata) und Schmetterlingen (Lepi-

doptera) übertroffen.

Auch unter den wenigen Wanderinsekten, die Spitzbergen über das Nordmeer erreichen (Lokki et

al. 1978), ist eine Schwebfliegenart (Elton 1925). Zusammen mit einer Reihe anderer Syrphidenarten

gehört balteatus mit zu den leistungsfähigsten Wanderinsekten im paläarktischen Raum. Vergleichen

wir quantitatives Ausmaß und Regelmäßigkeiten dieser Schwebfliegenwanderungen mit dem der

Wanderungen der Schmetterlinge, Libellen und anderen Wanderinsekten, so stellen die Schwebflie-

gen das wohl eindrucksvollste Beispiel für Insektenwanderungen in Europa.

Neben diesen alljährlichen Migrationen über weite Strecken bei den physiologisch starken Fliegern

haben wir ein weites Feld von verschiedenen Formen der Dismigration (3.1.), wobei alle hier abgehan-

delten Migrationsformen und die Begriffsbestimmungen (3.1., Gatter 1981a, 1981c) nicht als starre,

restriktive Einteilung verstanden werden sollten (Baker 1978). Schon in den Abschnitten 3.1 und 3.2

wird deutlich, daß es zahlreiche Übergänge zwischen Migrationsformen gibt; trotzdem ist es notwen-

dig, mit der begrifflichen Kennzeichnung bestimmter biologischer Abläufe Abgrenzungen zu anderen

zu schaffen — schon um dadurch zu weiterer Forschungsaktivität anzuregen.

Bei den Dismigranten unter den Schwebfliegen kommen dem Migrationsvorgang die unterschied-

lichsten Bedeutungen zu. Der Verteilung der Individuen im Raum dienen die einfachen Formen der

Anemomigration oder Verdriftung und der Expansiven Dismigration. Durchmischung der Popula-

tionen, das Auffinden neuer Habitate und die Möglichkeit der Arealerweiterung sind einige der Ziele

dieser emigrationsähnlichen Formen der Wanderung. Im Ergebnis sind sie vielfältig: in der durch-

schnittlichen Emigrationsdistanz einer Generation unterscheiden sich die Wanderungen der winzigen

und sehr flugschwachen Neoascıa podagrıca von den Einflügen der großen, flugphysiologisch kräfti-

gen Volucella zonaria zwar sehr, biologisch ist es aber derselbe Vorgang.

Bei allen am Randecker Maar wandernden Dismigranten werden @ Q häufiger in den Reusen gefan-

gen als OO (3.1., Tab. 5). Oft sind erstere erheblich weniger flugtüchtig. Besonders deutlich ist dies

z. B. bei Melanostoma mellinum (S. 29). Hier dienen die Migrationen der 9 wohl in erster Linie dem

Aufsuchen günstiger Eiablageplätze, während die Q’CO’ den aktiveren Teil des Migrationsprozesses

übernehmen. Das würde den Dispersionsprozessen entsprechen, die wir von Spinnerarten (Bomby-

ces, Lepidoptera) kennen. Auch hier kommt den in vielen Fällen flugagileren G’CO’ der aktivere Teil des

Dismigrationsgeschehens zu, während sich die schwerfälligen PP nur wenig von ihrem Schlüpfort

entfernen (Schwerdtfeger 1977). Als bemerkenswerten Fall konnte Mikkola (1968) Einflüge der süd-

ostrussischen Rasse des Schwammspinners Lymantria dispar L. vom zentralen Eurasien nach Finn-

land nachweisen. Im Extremfall wandern selbst Männchen von Arten, deren 2 Q flügellos sind, große

Strecken. Im 19. Jahrhundert erreichte der Große Frostspanner Erannis defoliaria Cl. Helgoland in

ungeheuerer Zahl (Gätke 1900).

Abb. 28: Beispiel einer Saisonalen Dismigration unter dem Einfluß vorherrschender Windrichtungen: Wander-

insekten, die sich beispielsweise im Mittelmeerraum (1) entwickelt haben, dismigrieren ım Frühjahr. Nach Süden

wandernde Tiere kommen um (gestrichelte Linien). Nach Norden dismigrierende Tiere erzeugen eine zweite Ge-

neration in Mitteleuropa (2). Die mitteleuropäischen Tiere wandern im Herbst wieder mit der Vorzugswindrich-

tung. Nach Norden wandernde Tiere kommen um, nach Süden wandernde errreichen das Herkunftsgebiet (aus

Gatter 1981).

Fig. 28: Examples for seasonal dismigration under the influence of predominant wind directions: Migratory insects

which developed in the Mediterranean area (1) dismigrate in spring. Insects migrating southward will die (broken

lines). Insects migrating northward produce a second generation in Central Europe (2). In autumn, they will mi-

grate again with predominant wind directions. Insects migrating northward will die, insects migrating south will be

able to return to the Mediterranean area (see Gatter 1981a).

Neben diesen Formen der Zerstreuungswanderung gibt es bei den polyvoltinen Arten sicher alle

Übergänge zu höheren Migrationsformen. Bei flugschwachen Arten, die den bodennahen Bereich sel-

ten verlassen, würde die biologische Bedeutung dieser Dismigrationen der Fxpansiven Migration ent-

sprechen. Sobald sich aber wenigstens kleine Teile einer Population in größere Höhen begeben, setzen

sie sich den Windströmungen aus und wandern mit ihnen (Abb. 28). Seit den Untersuchungen von

Glick (z. B. 1939, 1942, 1960) wissen wir, daß sich fast alle Insektengruppen und selbst Milben und

Spinnen in großer Höhe mit saisonalen Vorzugswindrichtungen verdriften lassen. Ist diese Form der

Migration erfolgreich, so wird daraus die „Produktion von Migranten durch Selektion“ im Sinne

Johnsons (1969). Es gibt zahlreiche Hinweise, daß viele Insektenarten Strategien entwickelt haben,

aus dem Windangebot bestimmte, für sie vorteilhafte Winde zu selektieren (Baker 1978, Gatter 1977b,

1981a; vgl. Abb. 28).

Ebenso ist belegt, daß Dipteren außerhalb Europas so extreme Wanderungen ausführen, wie wir sie

für Episyrphus balteatus und andere Arten annehmen. Johnson (1969) nennt einen Fall, der vermuten

läßt, daß sich selbst Stubenfliegen Musca domestica L. (Muscidae) über große Entfernungen in Höhen

von 300-1500 m mit dem Wind verfrachten lassen. Für die amerikanische Fliege Chochlomyıa homi-

nivorax Cog. (Calliphoridae) sind individuelle Zugentfernungen von PP über 290 km nachgewiesen.

Sie sucht im Herbst nahrungsreiche Gebiete einige hundert oder tausend Kilometer weiter südlich auf

und wandert im Frühjahr zurück nach Norden. In beiden Zugzeiten wird angenommen, daß Selektion

die Fliegen fördert, die vorteilhafte Winde nutzen. Extrementfernungen zwischen Winteraufenthalt

und den nördlichsten Sommerfunden liegen 2400 km auseinander (Baker 1978). Die australische

„bushfly“ Musca vetustissima Walk. (Muscidae) wandert saisonal und legt dabei z. T. Entfernungen

von 1500 km in offenbar einer Generation zurück (Hughes & Nicholas 1974).

Unterschiedliche Lebenszyklen, kurze Lebensdauer einzelner und Langlebigkeit anderer Syrphi-

denarten haben unterschiedliche Anpassungen an die Jahreszeiten und an die vorhandene Nahrung

notwendig gemacht. Die unterschiedlichen Habitatansprüche von Larven und Imagines verschiede-

ner Generationen zu verschiedenen Jahreszeiten wie auch die Ansprüche an den Überwinterungsort

haben bei den Schwebtliegen zu einer breiten Palette von Anpassungen geführt, auf die mit Wande-

rung reagiert wird. Baker (1969, 1978) konnte nachweisen, daß zu- und abnehmende Tageslängen

bzw. Temperaturen während der Larvenzeit bzw. Überwinterung steuernd auf die Zugrichtung ein-

wirken, die ein Insekt später einschlägt. Diesbezügliche Ergebnisse für Dipteren stehen bisher völlig

aus.

5. Zusammenfassung

Im Zeitraum 1975— 1987 wurden an der Station Randecker Maar/Schwäbische Alb (SW-Deutschland; 48.35 N,

9.31 E; 772 m über NN) die Wanderungen der Schwebftliegen erforscht. Zwölf Jahre lang wurde hier zwischen

Mitte/Ende Juli und Anfang Oktober (Tab. 1) eine nach Norden geöffnete Reuse betrieben, die nach Süden flie-

gende Insekten erfaßte (S-Reuse). Während fünf Jahren wurden zusätzlich mit einer baugleichen, nach Süden geöff-

neten Reuse Insekten gefangen, die nach Norden flogen (N-Reuse). Während dieser Zeit wurden 90049 Syrphiden

in der S-Reuse und 9815 Syrphiden in der N-Reuse gefangen und determiniert (Tab. 1). Die Leerung der Reusen

erfolgte stündlich.

Mit Hilfe dieses Materials wird die Phänologie der am Randecker Maar im Untersuchungszeitraum mit über

50 Exemplaren festgestellten Arten beschrieben (3.1).

Untersucht und für jede Art dargestellt werden jeweils folgende phänologischen Aspekte:

— Jahreszeitliches Vorkommen der Art;

— Jahreszeitliches Vorkommen der Geschlechter;

— Tageszeitabhängige Aktivität der Art;

— Tageszeitabhängige Aktivität der Geschlechter;

— Veränderungen der tageszeitlichen Aktivität ım Jahresverlauf;

— Unterschiede zwischen S-Reuse und N-Reuse;

— Geschlechterverhältnis;

— Einflug in die S-Reuse in Abhängigkeit von der Windrichtung.

Zur Einordnung dieser Ergebnisse in den Zusammenhang enthalten die Artabschnitte kurze Abrisse der Biologie

der Art und eine Zusammenfassung der bisherigen Wanderbeobachtungen. Wenn möglich, wird die Art abschlie-

ßend einer der bekannten Migrationsformen zugeordnet (Definitionen s. 3.1).

Um einen Überblick über die Wanderarten in Europa zu erhalten, werden darüber hinaus auch Arten dargestellt,

die am Randecker Maar nur in geringer Zahl in Erscheinung treten, andernorts aber als Wanderer beschrieben wur-

den (3.2).

Daraus ergibt sich eine Darstellung und Bewertung des Status sämtlicher als Wanderarten bekannten

Schwebfliegen Europas (Tab. 3).

Dabei wurden typische Merkmale der Phänologie Saisonaler Migranten (Arten s. Tab. 3), die richtungsorien-

tierte Wanderungen in Winter- und Sommerareale ausführen, deutlich:

1. Der Vergleich der Fangergebnisse beider Reusen und direkte Beobachtungen zeigen, daß die spätsommerli-

chen und herbstlichen Flugbewegungen gerichtet in südliche und südwestliche Richtungen führen.

2. Einige direkte Beobachtungen machen deutlich, daß Saisonale Migranten im Frühjahr in nördliche Richtungen

wandern.

3. Zwischen dem Vorkommen am Randecker Maar und dem am 350 km südwestlich gelegenen Col de Bretolet

bestehen charakteristische Unterschiede. Die meisten Saisonmigranten weisen am Col de Bretolet ein später liegen-

des Maximum auf. Wie bei Zugvögeln dürfte das zeitversetzte Auftreten auf die unterschiedliche Lage der Stationen

zurückgehen; demnach wird die in Zugrichtung nachgeschaltete Station am Bretolet später erreicht (4.1).

4. Während die meisten Saisonwanderer am Randecker Maar und an weiter nördlich liegenden Stellen ein weitge-

hend ausgeglichenes Geschlechterverhältnis aufweisen, dominieren in den Alpen die @ 9 sehr stark. Für Episyrphus

balteatus konnte nachgewiesen werden, daß die P sehr viel langlebiger sind. Die wesentlich kurzlebigeren ’C’

nehmen anfangs noch an der Migration teil (Randecker Maar), sterben aber in ihrem Verlauf ab (Alpen) (4.2;

Tab. 5). Gegen Ende der Zugzeiten erscheinen auch am Randecker Maar fast nur noch 9. Der Durchzugs-Me-

dian liegt deshalb bei PQ stets später als bei OO’.

5. Saisonale Migranten wandern nur bei Gegenwinden in bodennahen Schichten, um den widrigen Einfluß des

Windes zu verringern. Dagegen nutzen sie den fördernden Einfluß von Rückenwinden, indem sie in größere Höhen

aufsteigen. Dementsprechend werden Südwanderer fast ausschließlich bei Winden aus südlichen Richtungen gefan-

gen (3.4).

6. Sowohl das jahreszeitliche als auch das tageszeitliche Muster des Auftretens der häufigen Saisonmigranten

zeigt von Jahr zu Jahr, von den hohen Populationsschwankungen abgesehen, relativ geringe Unterschiede. Das

spricht gegen eine exogene Auslösung und für eine genetische Fixierung der Migrationsphasen.

Die Gruppe der Dismigranten (Arten s. Tab. 3), die aktive Zerstreuungswanderungen ausführen, ist weniger

eindeutig charakterisiert. Nicht für alle Arten gelten sämtliche der angeführten Merkmale der Phänologie der Dis-

migranten — eine Folge unterschiedlicher Funktionen, Ausmaße und Reichweiten dieser Zerstreuungswanderun-

gen (vgl. 4.5.2).

1. Die Unterschiede zwischen den Fangsummen in den beiden Reusen sind oft nicht sehr hoch; eine eindeutige

Vorzugsrichtung ist damit nicht erkennbar. Allerdings lassen die bei manchen Dismigranten höheren Unterschiede

vermuten, daß sie gezielt günstige Windrichtungen nutzen, um in Vorzugsrichtungen zu wandern.

2. Zwischen dem jahreszeitlichen Auftreten am Randecker Maar und in den Alpen (Col de Bretolet) besteht meist

kein Unterschied (4.1).

3. Das Geschlechterverhältnis zwischen Randecker Maar und Col de Bretolet zeigt keine gleichsinnigen Verän-

derungen, die auf einen Zusammenhang schließen lassen (4.2.; Tab. 5).

4. Am Randecker Maar dominieren bei allen Dismigranten die Weibchen, deren Flugaktivität im bodennahen Be-

reich im Zusammenhang mit dem Aufsuchen günstiger Eiablageplätze größer ist als die der GC’.

5. Saisonale Dismigranten verfügen nicht über die weitreichende Strategie der Meidung widriger und Nutzung

günstiger Winde der Saisonwanderer. Bei ihnen finden auch Mitwindwanderungen wenigstens zum Teil in Boden-

nähe statt. Im Gegensatz zu reinen Driftinsekten geraten Dismigranten bei allen Windrichtungen in die Reuse; Mit-

windbewegungen überwiegen aber leicht (3.4).

Für verschiedene ökologische Gruppen unter den Schwebfliegen sind Saisonale Migrationen integraler Be-

standteil des Lebenszyklus (4.3, 4.5):

1. Saisonwanderungen sind nur von Arten mit zoophagen (aphidophagen) Larven und Arten mit aquatischen

sapro-/microphagen Larven bekannt.

2. Zu den Saisonwanderern gehören insbesondere die Arten, die als Imago oder Puparium überwintern (vgl.

Tab. 3). Diese Arten überstehen den Winter im nördlichen Bereich ihres Verbreitungsgebietes nicht oder nur in re-

lativ geringer und von den Witterungsbedingungen des Winters stark abhängiger Anzahl. Sie wandern deshalb in

jedem Frühjahr dort ein; im Herbst finden Rückwanderungen in südliche Richtungen statt. — Da auch einige Ar-

ten, die im Larvenstadium überwintern, solche Wanderungen durchführen, kann vermutet werden, daß auch für sie

das Überwinterungsrisiko im südlichen Europa geringer ist.

3. Der adaptive Wert der Sidwanderungen im Sommer/Herbst besteht in einer Vergrößerung der Überlebens-

chancen im Winter. Der adaptive Wert der Nordwanderungen im Frühjahr kann für die aphidophagen Arten darın

gesehen werden, daß in Mitteleuropa dann optimale Entwicklungs-(Blattlausmaximum) und Ernährungsbedingun-

gen (Reichtum an „Fliegenblumen“) für Larven und Imagines bestehen, für die aquatischen Formen darin, daß im

humiden Bereich die als Larvenhabitate benötigten Kleingewässer in wesentlich höherer Dichte und mit geringerem

Austrocknungsrisiko anzutreffen sind als im sommerlich-ariden Südeuropa.

Dismigrationen dienen im wesentlichen dazu, Stellen mit für die Larven günstigen Lebensbedingungen leichter

zu erschließen. An solchen Wanderungen haben wieder in erster Linie Arten teil, deren Larven zoophag sind (4.3),

d. h. auf eine in Raum und Zeit nicht vorhersagbar auftretende Ressource angewiesen sind (Arten s. Tab. 3).

Aphidophage Schwebfliegen haben unterschiedliche Lebenszyklen, die zum Teil mit dem Frühjahrs- und/oder

dem Herbstmaximum der Blattlauspopulationen korreliert sind (mono- und bivoltine Arten; vgl. Tab. 3). Um sich

während des Soemmerminimums der Blattläuse erfolgreich fortpflanzen zu können, müssen polyvoltine Syrphiden-

arten in ihren Lebenszyklus Wanderphasen integrieren. Darüber hinaus sind sie hinsichtlich des Habitats und der

Larvenbeute wenig spezialisiert. Zwei Gruppen konkurrieren dabei um die Larvennahrung: Die Gruppe der Saiso-

nalen Migranten und die der Saisonalen Dismigranten (vgl. Tab. 3). Während erstere in der direkten Nahrungskon-

kurrenz überlegen sind, können letztere ein wesentlich weiteres Nahrungsspektrum nutzen (4.3).

Bei den Schwebfliegenarten, deren Larven terrestrisch saprophag oder phytophag sind, sind keine weitreichen-

den Wanderungen nötig, um die in Raum und Zeit vorhersagbar auftretende Larvennahrung zu erschließen. Wäh-

rend unter den Saprophagen einige Arten über kurze Strecken dismigrieren (Tab. 3), sind von den gewöhnlich stark

spezialisierten Phytophagen keine Wanderungen bekannt (4.3).

Ein Vergleich mit anderen Wanderinsekten zeigt, daß das Wanderverhalten der Schwebtliegen weitgehende

Ähnlichkeiten mit dem anderer bekannter Wanderinsekten v. a. unter den Schmetterlingen (Lepidoptera) aufweist

(4,5).

Enge Zusammenhänge bestehen zwischen Diapause und Migration (als Imagines diapausierende Arten migrie-

ren; 4.5.2), zwischen Generationenzahl und Migration (v. a. polyvoltine Arten migrieren; 4.5.3) und zwischen Ver-

breitungsgebiet einer Art und ihrem Wanderverhalten (Wanderer haben riesige Verbreitungsgebiete; 4.5.1).

Die Leistungsfähigkeit einzelner Schwebfliegenarten und Ausmaß und Regelmäßigkeit ihrer Wanderungen ma-

chen die Migrationen der Schwebfliegen zum eindrucksvollsten Beispiel für Insektenwanderungen ın Europa.

Darüber hinaus gehört der weit überwiegende Teil der land- und forstwirtschaftlich wichtigen Blattlausvertilger

unter den Syrphiden in die Gruppe der aphidophagen Wanderschwebfliegen. Bei allen Versuchen, einzelne dieser

Schwebfliegenarten im Rahmen der biologischen Schädlingsbekämpfung zu nutzen, ist eine genaue Kenntnis des

individuellen Lebenszyklus von entscheidender Bedeutung. Dabei wurde in der bisherigen Forschung die Bedeu-

tung des Phänomens der Wanderungen weit unterschätzt.

6. Dank

Seit dem Bestehen der Station am Randecker Maar haben sich zahlreiche Mitarbeiter große Verdienste erworben

beim Aufbau und der Unterhaltung von Station und Fanganlagen, durch technische Hilfe im Bereich der Datener-

fassung und -auswertung und bei der Betreuung der Mitarbeiter der Station. Dafür danken wir R. Braun, E. Blo-

cher, B. Dauner, Dorothea Gatter, Martin Gatter, R. Gardner, H. Grau, Herr und Frau Dr. Höß, Dr. Hohloch,

Barbara Hoschek, E. Kotzke, V. Kirchner, R. Laih, Dr. H. Löhrl, Prof. Dr. H. Mattes, S. Maskow, E. Miller,

W. Neuhäuser, Frau P. Schmid, K. Sıll, J. Schweier, Chr. und I. Schwotzer, H. J. Ullrich, H. Graf Westarp (f) und

I. Gräfin Westarp (f).

An den Beobachtungen und dem Auswerten der Reusenfänge und der Betreuung der Reuse beteiligten sich fol-

gende Mitarbeiter, bei denen wir uns herzlich bedanken:

F. Alkemeyer, G. Arnold, S. Baumung, M. Behrndt, B., M. und W. Beissmann, Renate Benath, Heide Berger,

Anke Borchert, $. Bräger, R. Braun, Jutta Brüggemann, H. Buchmann, K. Burbach, A. Burnhauser, B. Dauner,

W. Dickore, T. Dolich, Ina Donat, W. Dornberger, M. Drösler, Dr. H. Ebenhöh, K. Einig, F. Engele, Dr. R. Er-

tel, D. Fausel, J. Fehling, H. Feil, K. H. Fiala, Ursula Firsching, M. Flade, U. Friedlin, D. Frühling, R. Gardner,

C. und P. Gatter, M. Göpfert, Christiane Guth, A. Hachenbersg, $. Häfele, V. Hennig, Ute Hermann, Eva Hesse,

C. Höller, R. Hömke, W. Hörnle, T. Hübner, B. Hündorf, K. Hund (f), D. Ikemayer, D. Ilg, ©. Jahn, R. Jahr-

aus, $. Jansen, W. Kantner, M. Kasparek, Ulrike Kay, Britta Kiesewetter, Dr. F. Kipp, C. Kleemann, Dr.

G. Klump, A. Koch, H.M. Koch, K.H. Kolb, Karın Krämer, ©. Krösche (f), Jutta Krusenbaum, V. Laske, Caro-

line Liepert, van Lin, F. Llimona, K. Lohrmann, K. Loos, K. H. Loske, Ehepaar Luce, E. Ludwig, J. Lüttmann,

M. Luy, U. Maier, D. v. Mallinkrodt, J. Maser, S. Maskow, Prof. Dr. H. Mattes, M. Mayer, Dr. T. Meineke, Mör-

gelin, Renate Mogck, W. Müller (}), Barbara Nebel, Dr. M. Neub, G. Neumann, ©. Neumann, Ursula Nigmann,

Silvia Nikulski, R. Oppermann, K. Penski, P. Pfeilsticker, Pfitzenmaier, R. Pliefke, Dr. H. Prahl, A. Prinzing,

Prof. Dr. R. Prinzinger, E. und B. Raddatz, D. Räpple, M. Ralston, B. Ratzke, K. Reiner, W. Remm, W. Riedel,

Dr. Ch. Rieger, J. Rinn, R. Rochau, M. und $. Rösler, Rotter, Andrea Ruf, H. Schlüter, I. Schmoll, W. Schreck,

I. Schröder, R. Schütt, Petra Schwartz, R. Seibold, F. Seifert, Dr. B. Seitz, K. Siedle, K. Sievers, Petra Sperlbaum,

T. Stadtlander, H. Stäbler, K. Steiof, D. Streng, H. Strunck, J. H. Stuke, M. Stumpf, J. Trittler, T. Volpers, Chr.

Wagner, H. Wahl, F. Weber, P. Weber, V. Weiß, Frau Winchenbach, Ursula Witte, Wolf.

Für technische Unterstützung bei der Datenauswertung danken wir den Firmen Hewlett Packard, IBM und

MBB.

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100

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Axel Hausmann

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Zur Dynamik von Nachtfalter-Artenspektren:

TURNOVER UND DISPERSIONSVERHALTEN ALS ELEMENTE

VON VERBREITUNGSSTRATEGIEN

von Axel Hausmann

Inhalt

I. ERFASSENDER TEIL: DAS ARTENSPEKTRUM

k; Einleitung ee hanorne non ie ee ae se ee een 4

2% Untersuchungsgebietar zur 2... 2. eat ass anna ep nehe ehe nee he nee een 5

Dal GLOBLaUmE NE a a ee: 5

2.2. ° Botanische Charakteristik der Fallenstandorte......................2z02220200s0eneneeennenn 6

2.3 KlimadatenundJWetter..............000 222020. RT en 11

3 Melhode N ee ee 11

3.1 Bann esscaohedeeasee en eere Te ee en ee E 11

3.1.1. Bebend„Lichtlangtees a ass rpalee eBensris re Re 11

Balz. AndereMethodikente. mn. ae nenege. eeeen r 13

3.2. Bangmodust en m m a see ds re a N: 14

RE TIERE REN REN RE 14

Bu2.2 Raum Re as a ee Eee 14

3.2.3 N 15

3.2.4. Zusätzliche, Experimenten... essen ern 16

33. PNUSWELLUNSEAR TE A a I ee en ee 16

3.4. Markierung a ee ee ee 17

35. Materialen ve N er Re een en ale ses enge ehren San ne rapie ee 19

4 INTEEHUSLE en Be ee ee 19

4.1. Vorbemerkungene ne a nee een es ee 19

4.2. Magtalter a ee ee ee TR, 20

4.3. Nachtfalten m nen ee RE N 21

4.4 Baunistischer Hinweise nt her nn RL nenn EN RL NEAR IR EINE DANKEN TOR RREU EBEN ET RORRR, 43

4.4.1. Arealenweiterungen.n. einen een miese 43

4.4.2. TEA VENIUSLEN N erste ae ensure res ee 43

4.4.3. Artensmit»lokalem; Vorkommen...................0.0.0..000.ensssneeseneeaan sangen 44

5% Das: Phänomen Turnovet.......r.:-.%.3.8---..H ee ne ee 46

ST @ Arten-Zeit-Beziehung ..... Aa IR ARENA N nennen 46

3.2; Fosrmeln...24.... 2 er RE ET RT ee 47

6. Interpretation des apparenten Turnovers .................040@0440404snnssnnnnennnnnennnnnnnnnnnn 49

6.1. Berechnung dersAusfauschraten 2:12. I ER RAN ee nnd anne none ne 49

6.1.1. Vorbemerkungen... 00 dee a 49

6.1.2. Liste der -Turnoverwerle.......tscsscseseineeae res RE 49

617214: Garten ("SINE 49

6.1.2.2. Zusammenstellung der in den letzten 3 Jahren festgestellten Turnoverwerte 51

6.1.3. Folgerungen‘. 2... 52

6.2. Turnover durch Kolonisationsversuche biotopfremder Arten.............u.zeusrsne0. 53

6.3. Abhängigkeit des Turnovers vom Flächeninhalt des abgedeckten Areals ....... 56

6.3.1: Vergleich: der Werte... 56

6.32. Rückschlüsse für faunistische Ansätze ...............z.0:0...0..000000000000000 nenne nee nennen 56

6.4. Turnover. Konstanz und. Fluktuation............5...sse0:0.0aane ame renen E 56

6.4.1. Tagfalter (unterer Inn: REICHHOLF, 1986).........uuuuesseeeeeeeeneesssssssssennnnnnnennnn 56

6.4.2. Nachtfalter im Gitschtal (WIESER, 1987)............uuunneseeeeesnaennennneessnnnnnnnennneenn 57

6.4.3. Nachtfalter in Oberschleißheim (Garten)..................uu4u0000000000@seneeeenennneeneeenenn 59

6.4.4. Nachtfalter in Oberschleißheim (Offenland: "HM" und "HO".)...................... 61

6.4.5. Eulenfalter (Noctuidae) in Südböhmen und England ........................22220022000000 64

1: Rückschlüsse aus den Häufigkeitsverteilungen .....................-2...02s0200002020en en 65

71: Artenzahl' und 'Stichprobengröße...........csc.cmccoseeeencnneen nenn anne een nano 65

12: Vergleich mit den "log-series" WILLIAMs (1964).............u.uussss0nnneneenneneen 67

1:3. Zum Problem der Erfassung der seltenen Arten.................z2s2s2s0s0snenenenen nenn 67

7.3.1: Artenzahl’ und’ Fangrhyihmus........ +... arena nennen ge ee 67

3.2: PRESTONS vet line" ee ee ne nen ana ae ee 69

II. EXPERIMENTELLER TEIL: DISPERSIONSVERHALTEN UND TRIVIAL MOVEMENT

8. Die Experimente: ....-...cc essen sine genen een en 72

8.1. Vorstellung der Markierungsexperimente ...........u.---40s444snennnennnnnnnn nennen nennen T2

SIT Allgemeines: .....uum.ens0sceseeneueneaanee nenne nennen ernennen ae 2

8-1.2- "Fern" -Wiederläange .....nsseasenanuessnnssennensen sten nnenen armen ee ae 72

8:13. Verringerte Fallendistanzen ........s.<u0040.00000040sS00neRann en ine ee nee 2.

8.1.4. Versetzexperiment ..........uu0 Herne ana snne nam eneen ernennen an san nee 12

8.1.5. Rückschlüsse aus Ortswiederfängen ...................u.susuensnossennnnonensnnnsnonnnnnnnennnne 23

8.2. “"Fern"-Wiederfänge ..........uusscssesssenncsnennunendennennee denne een ee ee 75

82:1. ÜDErsIcht nenn 75

8.2.2. DiskUSSIOR '.......4:0.00. 00.400 nase ons ernennen gerne ee 76

8.3: Verringerte Fallendistanzen .........üsosaneeuesnensssseuuenseneeneanennensn seen ag ara ne 77

823]: ÜBersicht"uber ‚das Material... {hl

8.3.2: Weitere Ergebnisse, aufgezeigt an ausgewählten Arten .............-ussussnsenennen 80

8.4. Versetzexperiment nen ER, 82

8.4.1 Übersicht#über" das Material... 0... 82

8.4.2 Ergebnisse Hin. N IE OL I Eee 83

8.4.3 Vergleich der sich ergebenden Muster _..........cesssessesssensnnnennnsnnesnnnnnnenennnn nenn 97

8.4.4 Hinweise für faunistische Arbeitsansätze .............. wagen 99

8.5. Rückschlüsse aus Ortswiederfängen ............uscccesensessesenennensensnnnsnnnnnnnnnnnnneenenn 99

8.5.1. Die Problematik atirta. lat a aa ein 99

8.5.2 Beispiel:JAmathes) triangulumn.. 222.....204. 2.8. er. rel alsan. 101

8.5.3 Beispiel:#Calospilosasylvatant an..n:n.. este Na a Vin erieden: 102

9. Übersicht über die Reaktion der Arten ......cnnesnneesenenennnnennnnenennenneennnnnnnn 103

9.1. Vorbemerkungen #r..2.. 2 HH IHN kuss .eteben seen. eh 103

I Attenlusten a ee ea Renee aa ade Nee 105

9E3% AUSWETLUNgHHA ME ee een ee ee ne ee a ea 195

9.3.1. T=K -Kontinuums Me an a a a ee 195

9.3.2 Elugdistanzen 2 an se ae seen 195

93.3. Ökotypenn a a a en 196

9.3.4 Arten jextremersJahreszeiten ru... nennen ee 197

9.33. Broterandrie tu. m nn 2 Nr sad I a a ea se aser en 197

9.3.6. Bi-2und Bolyvoltinismussı.. an. 197

9.3.7 Einflüsse der Witterung auf die Mobilität ...........eessesceneeeeeneenseenenneeeneeneennn 197

10. Ergebnisse zur Biologie von Nachtfalterpopulationen ..............ccucusesneeeneeen- 199

10.1. Wanderfalter ee 0a... 00:00 Me a ee no, 199

10.2. Direkter Einfluß>von=lichtquellen ı..... ra... nn 199

10.3. Eebensdauer. 2 nr ee RE. en er een can 200

10.4 Weibchen Raten 42. He Eat ee 200

10.5. Bi cund ;Polyvoltinismusu.e. 123... sa ee 201

10.6. Ortstreue/Habitattreue 1... 2. Mater een.) 201

10.7. Geländestrukturen rm. EN ts 201

10.8 Bopulationsdiehtener rs a el en 202

10.9. Nektarquellen®e....... 2. ee es een ande Nennen sen. Are 202

10.10 Witterung a ne are Biesiee 202

10.11 Dispersionsaktivität, trivial movement, Flugdistanzen ...............cccenneenne. 202

10.12 SPTALE RITA ee en en lei ee. ats 203

10.13 TrittsteınKolonisationens. eu... N es a kale 205

10.14 Wirtspflanzen, Spektren) 4.9 9 et ee ee 205

11. Zusammenhänge zwischen Artenspektren-Dynamik und Artendynamik ........ 205

11.1. Untersuchungsgebiet 8%... users 205

11.2. Fluktuation von Eulenfaltern in Südböhmen ...............uuus2cccenasseeenseeeenneeeneneen 206

12. Diskussionws a. ern ea are 207

ZUSAMMENFASSUNG HR. ER EMEEE ERILEETATRE NR en 208

N N 210

BMTERASTURS 3 37.00: 20 a rn ee anne nce denne sn ee ge 211

ANHANG: SUMMARY me nasse ee ee rasen 216

ANHANG: -ARTENREGISITER FE U ans a era 219

I. ERFASSENDER TEIL: DAS ARTENSPEKTRUM

1. EINLEITUNG

Die Dynamik von Arten, von Populationen einer Art und von Artenspektren eines defi-

nierten Areals sind wichtige Gesichtspunkte für ökologische Beurteilungen einer Art

oder eines Biotops und somit für den Naturschutz bedeutsam.

Bei vielen Tiergruppen sind wir heute, was die in den Artenspektren stattfindenden

Austauschprozesse betrifft, auf einem einigermaßen befriedigenden Kenntnisstand ange-

langt, wie bei den Vögeln (z.B. DIAMOND, 1969; JONES & DIAMOND, 1976; u.s.w.),

oder den sessilen marinen Organismen (SCHOENER & SCHOENER, 1981).

Die Tagfalter sind bezüglich der Artendynamik (z.B. Dispersionsverhalten) ebenfalls

recht gut untersucht, hier vermittelt vor allem die amerikanische (z.B. EHRLICH,

1961; EHRLICH & DAVIDSON, 1961; SCOTT, 1975; WATT et al., 1977, SCHRIER et

al. 1976) und die englische (z.B. DOWDESWELL, FISHER & FORD, 1940 und 1949)

Literatur ein einigermaßen solides Basiswissen, bestehend aus vielen Einzelinformatio-

nen. Zu interessanten Ergebnissen kamen SMOLIS & GERKEN (1986) in ihrer Arbeit

über die intrapopulare Mobilität von Widderchen (Zygaenidae).

Was jedoch auf der Ebene der Artenspektren die Austauschprozesse betrifft, die diese

Artendynamik durch ständiges lokales Verschwinden und Wiederbesiedelung hervorruft,

ist vergleichsweise wenig publiziert worden (z.B. REICHHOLF, 1986). Einen Versuch,

den einheimischen Tagfalterarten verschiedene Verbreitungsstrategien zuzuordnen, un-

ternahm WEIDEMANN (1986a; 1986b).

Bei den Nachtfaltern ergeben die bisher publizierten Arbeiten bezüglich der genann-

ten Thematik erst ein sehr bruchstückhaftes Bild. Erst UTSCHICK (1989) versucht bei

einer allerdings recht kleinen Stichprobe aus Lichtfallenfängen am Inn eine Biotop-

beurteilung über Artenumsatz-Raten. Ohne detailliertere Diskussion reißt auch MA-

LICKY (1974b) durch die Ermittlung einer Artenübereinstimmung von Jahr zu Jahr die

Problematik an.

Während beispielsweise bei den Vögeln selbstverständlich zwischen Brutvögeln und

Durchzüglern unterschieden wird, fehlt eine solche Differenzierung - abgesehen von

einigen wenigen Wanderfalterarten - in Publikationen von Nachtfalter-Artenspektren in

fast allen Fällen. Dies liegt in erster Linie an den durch Nachtaktivität und fehlendes

Revierverhalten verständlicherweise erschwerten Erfassungsbedingungen.

Nachdem in einer vorbereitenden Arbeit bei der Noctuiden-Unterfamilie Amphipyrinae

in der Methodik des Lebend-Lichtfallenfangs ein Jahr-zu-Jahr-Artenaustausch (Turn-

over) von ca. 35 % festgestellt wurde, aber offenbleiben mußte, wie stark die Methodik

dieses Ergebnis beeinflußte, war nun die Idee, ein ganzes Fangstellen-Netz zu errich-

ten. Mehrere Fallen sollten bei geringeren Distanzen parallel in den selben Nächten

betrieben werden, mit dem Ziel, anhand ausgewählter Arten Zusammenhänge zwi-

schen der Dynamik der Arten in Raum und Zeit und Austauschprozessen in der Zu-

sammensetzung der Artenspektren von definierten Arealen zu erarbeiten.

Hauptgegenstand der Untersuchung sollen die nachtaktiven Macroheteroceren-Familien

in der Methodik des Lebend-Lichtfallenfangs sein. Aus Markierungexperimenten resul-

tieren weitere Hinweise, vor allem auf die Dispersionsdynamik der Arten.

In fast allen Fällen waren bisher ähnliche Untersuchungen auf die großräumige Dyna-

mik der Wanderfalter gerichtet, jedoch können sich Erkenntnisse gerade über die Aus-

tauschprozesse, die sich in Distanzen von unter 5 km abspielen, als nützliche Hilfsmit-

tel erweisen, beispielsweise für Programme im Zuge von Biotopvernetzungsmaßnahmen.

2. UNTERSUCHUNGSGEBIET

2.1. GROSSRAUM

Das Gemeindegebiet Oberschleißheim (31 km?, 475-488 m ü. NN) liegt an einer inte-

ressanten Stelle im Nordwesten der Münchner Schotterebene (vergleiche hierzu die

Übersichtskarte, Abb. 1).

Abb. 1: Gemeindegebiet Oberschleißheim. 1 = Kiefern-Eichenwaldgürtel; 2 = Flughafen-

gebiet (Magerrasen); 3 = Dachauer Moos (entwässert und größtenteils landwirt-

schaftlich genutzt). Abkürzungen der Fundorte siehe Text (nach HAUSMANN, 1988,

verändert).

Types of habitat in the municipality of Oberschleissheim near Munich. 1 = Pine-Oak

forest belt, 2 = air strip (low fertility pasture), 3 = fen area of Dachau (drained and

mostly changed to arable land), Abbreviations cf text.

Das eigentliche Siedlungsgebiet (48°15 N/11°34 E) ist im Nordosten und Osten von

einem relativ trockenen Kiefern-Eichenwald ("Vaccino-vitis- idaea-Quercetum"), im

Süden von einem Magerrasengebiet, das einem Trespen-Halbtrockenrasen (Mesobrome-

tum erecti) nahekommt, und im Westen vom ehemaligen Niedermoor "Dachauer Moos",

das jetzt weitgehend landwirtschaftlich genutzt ist, begrenzt (siehe auch HAUSMANN,

A., 1988; HAUSMANN, S., 1982, 1984, 1987).

Das Gelände ist, von unbedeutenden Bodenwellen abgesehen, eben. Im ganzen Gebiet

bildet ein aus tonigen und sandigen Mergeln bestehender tertiärer Flinz eine wasser-

stauende Schicht, die von ca. 15 Meter mächtigen quartären und postglazialen Schot-

tern bedeckt ist (vergleiche HAUSMANN 1984 und 1987, Bayerisches Geologisches

Landesamt briefl.).

2.2. BOTANISCHE CHARAKTERISTIK DER FALLENSTANDORTE

Da praktisch alle Biotope in irgendeiner Weise vom Menschen beeinflußt oder sogar

geschaffen wurden, ist die im folgenden teilweise getroffene Einteilung in Pflanzen-

gesellschaften oft nur tendenziell zu verstehen.

Siedlung

Wie es für Wohngebiete typisch ist, zeichnet sich auch der Siedlungsbereich Ober-

schleißheim durch eine uneinheitliche, reichhaltige Staffelung der Vegetationselemente

aus. Standort und Artenzusammensetzung sind fast ausschließlich vom Menschen nach

ästhetischen Gesichtspunkten bestimmt.

- Siedlung Süd ("SiS"): Der Fangplatz befindet sich auf einem knapp 2 ha großen

ungenutzten Geländestreifen, der auf einer Seite vom Würmkanal, auf den ande-

ren Seiten von Gärten begrenzt ist. Die Krautschicht besteht fast nur aus Bren-

nessel-Giersch-Saum (Urtico dioicae-Aegopodietum), der stark von Brombeere

(Rubus fruticosus) durchsetzt ist. Im artenreichen Baumbestand dominieren Ge-

wöhnliche Esche (Fraxinus excelsior), Schwarzer Holunder (Sambucus nigra),

Hybrid-Pappel (Populus euramericana) und Weiden (Salix). Der Beschattungsgrad

beträgt ca. 70 %.

- Siedlung Mitte ("SiM"): Die nähere Umgebung der stark befahrenen Straßenkreu-

zung ist von Gärten beherrscht. Botanisch interessanter ist v.a. die Begleitflora

des Schloßkanals in 50 (fragmentarisch) bis 100 m Entfernung: Charakteristisch

sind Rohrschwingel-Rasen (Dactylo-Festucetum arundinaceae), Blaubinsen-Roßmin-

zen-Gesellschaft (Junco inflexi-Menthetum longifoliae) sowie Weiden und

Schneeball (Viburnum).

- Siedlung Nord ("SiN”): Es handelt sich hier um den Garten des Verfassers, der

Fangplatz befindet sich auf der Südseite des Hauses. Das Siedlungsgebiet ent-

stand 1952 unter partieller Rodung des Bergliwaldes, der Waldcharakter blieb

jedoch bestimmend. Unter den Bäumen dominieren 15-20 m hohe Kiefern (Pinus

sylvestris), Stieleiche (Quercus robur), Gemeine Fichte (Picea abies), Hängebirke

(Betula pendula), Winterlinde (Tilia cordata) und Hainbuche (Carpinus betulus).

Der Beschattungsgrad beträgt ca. 60 %. Im Garten wurde der größte Teil der

Wiesenfläche (ca. 200 m?) in den Jahren seit 1985 abgemagert, was zu einer

sehr artenreichen Krautschicht führte. In manchen anderen Gärten sind Ehren-

preis-Erdrauch-Gesellschaften (Veronico-Fumarietum officinalis) zu finden.

- Wald Süd ("WaS"): Dieser Standort befindet sich nur 30 m vom letztgenannten

entfernt, nämlich auf der gegenüberliegenden Seite des Hauses. Hier erstreckt

sich ein zwischen den Gärten liegender 20-40 m breiter Waldstreifen, der mit

dem 150 m entfernten Berglwaldgebiet in direkter Verbindung steht. Es dominiert

die Gemeine Fichte bei einem Beschattungsgrad von ca. 90 %. Erwähnenswert sind

ein Bergweidenröschen-Stinkstorchschnabel-Saum (Epilobio montani-Geranietum

robertiani) sowie ein Schlehen-Weißdorngebüsch (Pruno spinosae-Crataegetum) in

je 20-30 m Entfernung (Lageskizze siehe 8.1.).

- Wald Mitte ("WaM”): Der Fangplatz befand sich auf einer 0,5 ha großen baumlo-

sen Fläche im Berglwald (ca. 150jähriger Kiefern-Eichenwald mit starkem Laub-

holzunterwuchs), die in der Vergangenheit als Wildacker genutzt wurde, im

Untersuchungsjahr jedoch von Pionierpflanzen wie Acker-Kratzdistel (Cirsium

arvense), Gewöhnlichem Pastinak (Pastinaca sativa), Gemeinem Beifuß (Artemisia

vulgaris) aber auch von Tüpfel-Johanniskraut (Hypericum perforatum) und Gän-

se-Fingerkraut (Potentilla anserina) dominiert war. Die Flächenränder bestehen

aus Schlehen-Weißdorn-Gebüsch (Pruno spinosae-Crataegetum) mit viel (ange-

pflanztem) Liguster (Ligustrum vulgare).

- Wald Nord ("WaN"): Der Standort befindet sich im relativ naturnah gestalteten

Wasserwerksgelände am nordöstlichen Rand des Berglwaldes. Vor ca. 10 Jahren

bestand die Fläche (wie heute noch in der näheren Umgebung) aus einem ziemlich

abrupt in das angrenzende Ackerland übergehenden forstwirtschaftlich genutzten

Hochwald. 50 m südöstlich bzw. 100 m westlich von "WaN" liegen die Fangplätze

"WNo" und "WNw" (siehe Lageskizze).

Die ziemlich abgemagerten arten- und blütenreichen Wiesenflächen sind an man-

chen Stellen noch von Pionierpflanzen dominiert (Ackerkratzdistel, Gewöhnlicher

Pastinak), sie kommen an anderen Stellen jedoch dem Mittelklee-Odermennig-

Saum (Trifolio medii-Agrimonietum eupatoriae) nahe, häufig ist weiterhin die bun-

te Kronwicke (Coronilla varia). Diese Magerrasenflächen sind mit ähnlichen Bio-

topen in der Gegend der Fröttmaninger Heide und des Mallertshofer Holzes

("M") im Osten über Trittstein-Lebensräume im Abstand von 100-300 m vernetzt.

Am Standort "WNo" befindet sich ein ca. 20-30 m? großes Teichröhricht (Scir-

po-Phragmitetum).

Die von jungen Bäumen durchsetzten Gebüschflächen sind äußerst artenreich und

haben manchmal, vor allem an "WNw" den Charakter eines Schlehen-Weißdorn-

Gebüsches (Pruno spinosae-Crataegetum).

Im Süden schließt ein Kiefern-Hochwald (Unterwuchs nur Him- und Brombeere)

an, in der Waldfläche im Südwesten sind dagegen Wald-Kiefer und Gemeine

Fichte zu ungefähr gleichen Teilen vertreten. In der Nähe des Fangplatzes WNw

ist eine ca. 100 m? große Buchenenklave (Fagus sylvatica) eingestreut und im

Norden umgrenzt das Gelände schließlich ein kleines Fichtenwäldchen.

Der Flächeninhalt des teilweise verbuschten Magerrasens im Wasserwerksgelände

beträgt ca. 2,3 ha. Wenn man bedenkt, daß auch Tiere, die sich im toten Winkel

der Falle A (WaN) befanden (bereits relativ stark verbuschte Wiesenfläche), schon

durch geringe Ortsveränderungen in den Einzugsbereich der Falle gelangten, dann

wird deutlich, daß durch die 3 Standorte fast die gesamte Fläche abgedeckt wurde.

Wald (>10m : : Gebüsch (1-5m

hoch) Ba hoch)

Abb. 2: Das Wasserwerksgelände Oberschleißheim mit den Fangplätzen "WaN" (=A),

"WNw" (=B) und "WNo"” (=C).

The localities "WaN" (A), "WNw" (B) and "WNo"” (C); dark grey = wood (trees

over 10 m), grey = shrubs (1-5 m).

- Das Mallertshofer Holz im Nordosten ("M"), stellt einen Ausläufer des Kiefern-

Eichenwaldgürtels dar. Für die vorliegende Untersuchung wichtige Teilflächen

dieses Gürtels sind der Berglwald ("B") im Norden, das Schweitzerholz ("S") im

Osten und das Korbinianiholz ("K") im Süden.

Halbtrockenrasen

Der Charakter des Trespen-Halbtrockenrasens (Mesobrometum erecti) ist auf dem

Flughafengelände (ca. 3,5 km?) im Süden des Siedlungsgebietes nicht eindeutig ausge-

prägt. Durch die extensive, nicht allzuhäufige Beweidung durch Schafe werden Pflan-

zenarten, die von den Schafen gemieden werden (z.B. Gemeine Schafgarbe, die an den

meisten Stellen sehr häufig ist), bevorzugt. Eine Entwicklung zu einem Enzian-Zwen-

kenrasen (Gentiano-Koelerietum pyramidatae) hat jedoch bisher nicht stattgefunden.

Abb. 2b: Luftbild des Wasserwerksgeländes Oberschleißheim (Freigabe Reg. v. Obb. GS

300/91 75/82.

Abb. 2c: Blick vom Fangplatz "WNw" nach "WaN" im Wasserwerksgelände Ober-

schleißheim.

- Halbtrockenrasen (Nord-)Ost ("HO"): Der Fangplatz befindet sich an der Stelle,

wo das Schweitzerholz und der Schloßpark (hier Stiel-Eiche als dominante Art)

aneinandergrenzen in einer kleinen Einbuchtung des Halbtrockenrasen-Gebietes.

Der Waldcharakter ist hier noch deutlich ausgeprägt. Durch die Randlage kommt

es zu einem großen Arten- und Strukturreichtum der Vegetation.

- Halbtrockenrasen Mitte ("HM"): Der Standort ist durch reinen Offenlandcharakter

bestimmt, zur Ausprägung des Trespen-Halbtrockenrasens siehe obige Bemerkun-

gen. Der Rand des Flughafengebiets ist in allen Richtungen ungefähr 800-1000 m

entfernt.

In ca. 150-300 m Entfernung befindet sich eine ca. 5 ha große ruderalartige Flä-

che ("R") mit 5-10 m hohen Weiden, Rainfarn-Beifuß-Gestrüpp (Tanaceto vulga-

ris-Artemisietum vulgaris) sowie Großer Brennessel (Urtica dioica), Schilfrohr

(Phragmites communis) und Land-Reitgras (Calamagrostis epigejos) als weitere

häufige Pflanzenarten.

Das im Südosten liegende Drittel des Flughafengebiets wird teilweise intensiv

landwirtschaftlich genutzt.

Am Flughafen-Nordrand befindet sich in ca. 800 m Entfernung ein neuentstande-

ner Teich mit Teichröhricht (Scirpo-Phragmitetum), Natternkopf-Steinklee-Gesell-

schaften (Echio vulgaris-Melilotetum) und einer Reihe von Pionierpflanzen in der

näheren Umgebung.

- Halbtrockenrasen West ("HW"): Am Westrand des Halbtrockenrasengebiets wurde

an einem offen strukturierten Standort gefangen, in dessen unmittelbaren Umge-

bung vereinzelt 1-2 m hohes Gebüsch und junge Bäumchen verschiedenster Arten

angepflanzt wurden.

In ca. 150 m Entfernung befindet sich der Würmkanal mit kleineren auwaldarti-

gen Baumbeständen. Die Sicht zur Lichtquelle war von dort durch ein 10 m ho-

hes Gebäude versperrt.

Dachauer Moos

- Würmauen ("Au"): Eine auwaldartige Begleitflora, durchsetzt von jahrhundertealten

Baumveteranen (siehe HAUSMANN, 1987) stellt zusammen mit einem an manchen

Stellen ausgeprägten Brennessel-Giersch-Saum (Urtico dioicae-Aegopodietum) ty-

pische Aspekte der Vegetation dieses Lebensraumes, der an der Fangstelle von

Ackerland begrenzt ist, dar.

Weiher im Torfeinfang ("We"): Der Torfeinfang ("T") ist ein Relikt ehemals aus-

gedehnterer "minerotropher Erlenbrücher" (Alnetalia glutinosae), die von den Be-

ständen der Moorbirke (Betula pubescens) beherrscht werden. Am Fangplatz sind

weiterhin Brennessel-Giersch-Saum (Urtico dioicae-Aegopodietum) und Teichröh-

richt (Scirpo-Phragmitetum) wichtige Pflanzenassoziationen.

Birket: Moorbirkenwald ("Mb"): Das Birket ist ein dem Torfeinfang recht ähnli-

cher Standort. Die Größe des relativ isolierten Wäldchens beträgt 16,9 ha.

Franzosenhölzl ("F"): In seiner Vegetation entspricht dieses ebenfalls isoliert

liegende Moorbirkenwäldchen den beiden vorhergehend beschriebenen Standorten.

Moorversuchsgut ("Mo"): Der Fallenstandort befindet sich auf einer kleinen Wirt-

schaftswiese, die an dieser Stelle von Brennessel-Giersch-Saum (Urtico dioicae-

Aegopodietum) begrenzt wird. In ca. 70 m Entfernung befinden sich 2 Entwässe-

rungskanäle, deren Vegetation als "Moorgebüsch"” (Frangulo-Salicion auritae)

bezeichnet werden könnte und von Weiden (Salix) dominiert wird. Einen ähnlichen

Charakter besitzt ein anderer Entwässerungsgraben ("E”) zwischen dem Birket

und dem Franzosenhölzl, von dem einige Beobachtungen stammen.

2.3. KLIMADATEN UND WETTER

Im langjährigen Mittel treten im Siedlungsgebiet Oberschleißheim Niederschlags-Jahres-

summen von 814 mm auf, in Badersfeld (500 m vom Birket entfernt) sind es 844 mm.

Das Temperaturmittel beträgt für Schleißheim 7,5 °C (Deutscher Wetterdienst/Wetter-

amt München, briefl.).

Insgesamt betrachtet dürften beide Fangjahre als durchschnittlich bis gut einzustufen

sein, was die Individuen- und Artenausbeute betrifft. Vergleichsweise katastrophal

wirkte sich jedoch die naßkalte Witterung von April bis Juni 1987 auf die Nachtfalter-

aktivitäten aus.

3. METHODE

3.1. FANG

3.1.1. Lebend-Lichtfang

Bei der repräsentativen Erfassung von Nachtfalter-Zönosen stellt die nächtliche Le-

bensweise einen Unsicherheitsfaktor dar. Durch moderne Methoden, vor allem durch

verbesserte Produkte der Beleuchtungsindustrie (z.B. UV-Röhren) konnten seit der

Jahrhundertwende gerade in der wichtigsten Methode, dem Lichtfang, entscheidende

Fortschritte erzielt werden.

Die "klassische" Methode besteht darin, an einem angestrahlten Tuch in Anwesenheit

des Beobachters die anfliegenden Tiere zu registrieren.

Moderne Lebend-Lichtfallen bieten dagegen bei Fragestellungen wie im vorliegenden

Fall eine Reihe von entscheidenden Vorteilen:

- Die mit UV-Röhren bestückten Fallen verbrauchen wenig Strom und sind daher vom

Stromnetz ohne aufwendige Apparatur (Aggregat) potentiell unabhängig.

- Der Beobachter wird nicht der schädlichen Strahlung ausgesetzt.

- Das Fangergebnis ist in gewissen Grenzen quantitativ auswertbar ("relative Häufig-

keiten"). Nach RETZBANYAI-RESER (1974) muß "die moderne faunistische For-

schung unbedingt auch quantitativ sein".

- Ein Parallelfang gleichzeitig mit mehreren Fallen ist durch einen Beobachter möglich.

- Die Methode ist konstant, so gut wie unabhängig vom Beobachter und in gewissen

Grenzen standardisierbar.

Einige Nachteile werden noch erörtert werden, diese können jedoch durch entsprechen-

de Maßnahmen auf ein Minimum reduziert werden.

Für die Parallelfänge wurden drei identisch gebaute Lichtfallen vom Minnesota-Typ

verwendet. Sie waren nach dem in REICHHOLF (1984) beschriebenen Prinzip konstru-

iertt und mit 18-W Schwarzlichtröhren (Maximum der relativen Leuchtstärke bei

360-370 nm) bestückt. Die Unversehrtheit der gefangenen Falter wurde durch Ver-

schlupfmöglichkeiten (Eierschachteln) und Regenschutz sowie durch einige in Kapitel

3.2. erwähnte Maßnahmen gewährleistet.

Abb. 2d: Lebendlichtfallen.

Abb. 2e: Die Lebendlichtfalle am Fangplatz "WaN” bei Sonnenaufgang. Die Auswer-

tung des Fangs an dieser Stelle war zum Zeitpunkt der Aufnahme bereits abgeschlossen.

Im Juli 1987 und von Juni bis August 1988 wurden zu Ergänzungsfängen teilweise noch

zwei weitere Fallen eingesetzt.

Eine der drei "Hauptfallen” wird von einer Autobatterie gespeist, die beiden anderen

arbeiten mit Netzstrom. Um die Konstanz der Methode zu gewährleisten, wurde gete-

stet, ob die Fangergebnisse der verschiedenen Fallen gleich waren. Hierzu wurden sie

im Flughafengebiet, wo an allen drei Standorten Netzstrom zu Verfügung stand, im

"Rotationsprinzip" jeweils durchgewechselt und die Verhältnisse der Individuen- bzw.

der Artenzahlen zu den beiden anderen Fallen ermittelt. Das Ergebnis zeigte eine

Drosselung der Individuen- und Artenausbeute bei der batteriebetriebenen Falle auf ca.

60 %. Dieses Manko wurde im Flughafengebiet und im Wasserwerk durch das erwähnte

Rotieren-Lassen ausgeglichen, an den Standorten SiS, WaM, Au, We und Mb ist bei

Vergleichen ein entsprechender Korrekturfaktor zu veranschlagen.

3.1.2. Andere Methodiken

Mit Lichtfang allein kann man kein 100 prozentiges Nachtfalter-Artenspektrum erfas-

sen. Dies hat seinen Grund darin, daß es nachtaktive Arten gibt, die Lichtquellen nicht

oder nur mit äußerst geringer Affinität anfliegen. Hierher gehören beispielsweise viele

Arten der Gattungen Cucullia und Catocala (Noctuidae). Manche Arten reagieren zwar

auf das Licht, fliegen die Lichtquelle jedoch nicht direkt an, so daß sie, wie manche

Flechtenbären (Nolidae), in Lichtfallen relativ selten erfaßt werden. Andere sind obli-

gatorische Tag- oder Dämmerungsflieger geworden, wie die Eulenfalter (Noctuidae)

Ectypa glyphica und Callistege mi, die Spanner (Geometridae) Archiearis parthenias

und Odezia atrafa sowie die meisten Sackträger (Psychidae) und Glasflügler (Aegerii-

dae). Ca. 5 % des Artenspektrums der Macroheteroceren dürfte in die bisher bespro-

chenen Kategorien fallen.

Andere Arten fliegen tagsüber und mehr oder wenig häufig auch nachts, z.B. die Wan-

derfalter Autographa gamma, Chloridea viriplaca und Macroglossum stellatarum (letz-

terer am Licht nur selten), aber auch Diacrisia sannio, Eustrotia olivana, Jaspidia de-

ceptoria, Phytometra viridaria, Hypena proboscidalis, Epirrhoe tristata, E. alternata,

Chiasmia clathrata, Ematurga atomaria, Siona lineata und einige weitere. Es handelt

sich hierbei ebenfalls um ca. 5 % des Artenspektrums.

Die Affinität zum Licht ist von Art zu Art, bei vielen Arten auch geschlechtsdifferen-

ziert verschieden, was beim Versuch einer Quantifizierung der Ergebnisse zu berück-

sichtigen ist.

Was kann also getan werden, um für faunistische Fragestellungen die Erfassung des

Artenspektrums zu optimieren?

Bezüglich der Tagflieger sind regelmäßige Tagexkursionen erforderlich. Diese wurden

im Untersuchungsgebiet in den letzten 15 Jahren jeweils 50-100mal pro Jahr bei einem

Zeitaufwand von 1-3 Stunden in den verschiedenen Biotopen durchgeführt. Zusätzlich

war bei der Installation der Lichtfang-Apparatur natürlich auch die Umgebung Ziel der

Aufmerksamkeit. In der Dämmerung schwärmende Falter wurden desöfteren mit dem

Käscher gefangen. Dies bringt vor allem im Frühsommer an blütenreichen Feldhecken

und Waldrändern oft erstaunliche Ausbeuten an Spannerarten (Geometridae).

Bei den Exkursionen wurden auch Präimaginalstadien gesammelt. Cucullia verbasci ist

beispielsweise im Raupenstadium in Oberschleißheim stets häufig, am Licht konnten

bisher jedoch noch keine Imagines beobachtet werden. Ähnlich wird dies auch bei vie-

len Psychiden und einigen anderen Taxa die Hauptmethodik sein, Artnachweise zu "lan-

den".

Köderfang erfolgte nur einige Male (ca. 20), meist mit schlechtem Erfolg (siehe UR-

BAHN, 1973). Hierbei war keine einzige Art zu "erbeuten”, die nicht auch irgendwann

ans Licht gekommen wäre.

3.2. FANGMODUS

Die letztgenannten Methodiken sollten Zusatzinformationen zur Vervollständigung der

einzelnen Artenspektren sammeln, um zu testen, ob in bestimmten Fällen apparente

Austauschprozesse von Jahr zu Jahr (turnover) in den Lichtfallen-Artenspektren nur

methodisch-bedingter Natur waren.

Um mit Hilfe der Lebend-Lichtfänge, die die Hauptgrundlage für die vorliegende Ar-

beit darstellen, in bezug auf die Fragestellungen verwertbare Informationen zu erhal-

ten, ist ein bestimmtes Muster in der angewendeten Methodik vonnöten. Dieses läßt

sich nach folgenden Kriterien gliedern:

1. Ökologie (Abdeckung verschiedener Habitate, Habitattypen)

2. Raum ("Fangstellennetz", verschiedene Distanzen)

3. Zeit (Fangnacht-Abstände)

4. Zusätzliche Experimente (Markierung)

3.2.1. Ökologie

Das Gebiet von Oberschleißheim wurde - generalisierend - in folgende vier "Haupt-

Biotoptypen” eingeteilt:

Siedlungsgebiet / Wald / Offenland (Halbtrockenrasen) / landwirtschaftlich genutztes Da-

chauer Moos (siehe "Untersuchungsgebiet”, Kapitel 2).

Es sollte, wo möglich, das Verbreitungszentrum der Arten ermittelt werden, um sie

in Einzelfällen an anderen Standorten als zugeflogene Gastarten einstufen zu können.

Hierzu war natürlich eine besondere Berücksichtigung der Larvalökologie nötig, um ei-

ne zahlenmäßig geringe aber bodenständige Anwesenheit einer Art aufgrund suboptima-

ler Bedingungen von solchen Dispersionsereignissen abtrennen zu können.

3.2.2. Raum

Es wurde jeweils mit drei identisch gebauten Lebend-Lichtfallen in derselben Nacht

parallel gefangen. Die drei parallel kombinierten Fangplätze blieben pro Jahr konstant

dieselben und lagen im gleichen Haupt-Biotoptyp.

So wurden 1987 jeweils drei Standorte im Flughafengebiet, im Berglwald und im Sied-

lungsbereich miteinander kombiniert. Im Birket (Mb) wurde zusätzlich gefangen, um

Verbreitungsschwerpunkte eventueller Zuflieger zu ermitteln, der Fang erfolgte hier

nicht über das Jahr hinweg parallel zu anderen Fangplätzen. Die Fallenabstände betru-

gen bei den Parallelfängen 0,5-2,0 km. Die Maximal- (Mb/HO) und Minimalabstände

(SIN/WaS) lagen bei 4,9 km bzw. 30 m.

1988 sollte durch eine Beibehaltung der Methode im Flughafengebiet ein Vergleich der

Jahre 1987/1988 ermöglicht werden. Am Nordrand des Waldes wurden die Fallendi-

stanzen um ca. eine Zehnerpotenz verringert (siehe Untersuchungsgebiet, Kapitel 2). Im

Siedlungsbereich erfolgte der Fang nur im Garten des Verfassers (SiN, WaS), meist

kombiniert mit den beiden anderen Standorten im Torfeinfang (We) und am Würmkanal

(Au).

Die durch dieses Netz "abgedeckte Fläche” (jedoch ohne vollständige Erfassung aller

Biotope!) beträgt ca. 6 km?, wenn man einmal die Standorte im Dachauer Moos unbe-

rücksichtigt läßt. In einzelnen Fällen sind nicht erfaßte Häufigkeits-Peaks oder sogar

zusätzliche Arten in dazwischenliegenden Biotopen außerhalb der Reichweite der Licht-

quellen zu erwarten (wohl v.a. im Bereich des Schloßparks).

Teilweise wurden auch Fangergebnisse aus den über 1000 Fängen der anderen Jahre

(1974-1986, 1989) berücksichtigt, von besonderem Interesse sind hierbei die an HO und

HM 1986 durchgeführten Parallelfänge.

3.2.3. Zeit

BETTMANN (1985a; 1985b; 1986) stützte seine Auswertungen auf Markierungs/Wie-

derfang-Versuche bei teilweise täglichem Lichtfang an einem definierten Standort, was

von verschiedener Seite kritisiert wurde (z.B. RETZBAYAI-RESER, 1980).

Um nicht zu stark in das Dispersionsgeschehen einzugreifen und den Tieren die Mög-

lichkeit zu geben, wegzufliegen, ohne in der darauffolgenden Nacht gleich wieder im

Bann der Lichtquelle gefangengehalten zu werden (bzw. nach anderen Theorien: "vom

Licht geblendet zu werden”, siehe SCHACHT & WITT, 1986), wurde ein Fang an der-

selben Stelle in zwei aufeinanderfolgenden Nächten vermieden. Eine Ausnahme wurde

1988 im Garten (WaS) gemacht (siehe II. Teil, "Vorstellung der Markierungsexperi-

mente", Kapitel 8.1.).

Die Anzahl der Fänge pro Monat verteilte sich nur im Garten und Wasserwerk 1988

im zweitägigen Rhythmus (vereinzelt Intervalle von 3 Tagen) gleichmäßig über das

Jahr, sie richtete sich ansonsten nach der schwerpunktmäßigen Verbreitung der Arten

auf die in den Markierungsexperimenten gezielt wurde. So wurde z.B. im Juni verstärkt

am Flughafen gefangen, wo in diesem Monat Pachetra sagittigera, Leucania comma,

Apamea sublustris und viele andere typische Falter offener, trockener Wiesen häufig

sind.

Die 1438 Fangnächte in den Jahren 1987 und 1988 verteilten sich folgendermaßen auf

die verschiedenen Monate:

300 N (Fangnächte)

F] 1987 BE 1988 > = 1438

Abb. 3: Verteilung der 1438 Lichtfänge auf die verschiedenen Monate der beiden Er-

fassungsjahre 1987 und 1988.

Distribution of the 1438 light-trap-captures over the months from January to De-

zember 1987 and 1988.

3.2.4. Zusätzliche Experimente

In einigen zusätzlichen Experimenten sollte anhand markierter Nachtfalter versucht

werden, weitere Aufschlüsse über Dispersionsverhalten und Austauschprozesse in Nacht-

falterpopulationen zu erhalten. Hierbei wurde jeweils mit besonderem Fangmodus gear-

beitet.

3.3. AUSWERTUNG

Die jeweils von Sonnenuntergang bis Sonnenaufgang betriebenen Fallen wurden noch

vor der Morgendämmerung aufgesucht, verschlossen, an einen kühlen, schattigen Ort

gebracht und an den heißesten Tagen im Hochsommer bisweilen auch mit etwas Eis

gekühlt, da sich in Spitzenflugnächten die Auswertung bis in den Vormittag hinein er-

streckte, obwohl sie bereits gegen 3.°° am Morgen begonnen wurde. Die frühe Aus-

wertung ist ratsam, um Verluste durch Vögel zu vermeiden und eine Schädigung der

Falter durch unnötig lange Gefangenschaft und Austrocknung zu verhindern. Zur Be-

schleunigung des Vorgangs halfen in der Hauptflugzeit desöfteren mein Vater Stefan,

meine Frau Silvia und meine Schwester Susanne Hausmann durch die Übernahme der

Schreibarbeiten.

Die in der Umgebung um die Falle herum sitzenden Falter wurden getrennt notiert, bei

den Auswertungen jedoch zum Fangergebnis gerechnet. Beim Öffnen der Falle wurde

mit größter Sorgfalt gearbeitet, so daß nur einzelne Tiere vor der Registrierung fliehen

konnten.

Die Artbestimmung erfolgte im Freiland nach den Bestimmungswerken KOCHs (1984)

und FORSTER & WOHLFAHRT (1955-1981). Bei markierten sowie bei offensichtlich

biotopfremden Tieren wurde auch das Geschlecht ermittelt.

Im Zuge einer fotographischen Dokumentation des Fangergebnisses entstanden in den

letzten 10 Jahren über 10.000 Fotos; in besonderen Fällen erschien es ratsam, Beleg-

exemplare in die Zoologische Staatssammlung München zu geben.

Ca. 80-90 % der im Untersuchungsgebiet vorkommenden Arten konnten an der Fang-

stelle determiniert werden. Bei einigen (artenmäßig vergleichsweise wenigen) proble-

matischen Arten mußten zusätzliche Differenzialmerkmale wie Fühlerbau oder (v.a.) die

Struktur der Genitalapparaturen untersucht werden. Letzteres wurde bei den Gattungen

Mesapamea, Amphipoea (1987) und vielen Arten der Gattung Eupithecia obligatorisch

durchgeführt, bei einer Reihe weiterer Gattungen wie z.B. Oligia, Hydraecia oder Opo-

rinia geschah dies in größerem Ausmaß. So wurden in den beiden Untersuchungsjahren

zur Absicherung der Determination über 1000 Genitalpräparate angefertigt, davon über

die Hälfte aus der Gattung Eupithecia.

Das Freilassen der Tiere erfolgte in einem Radius von 5-10 m um die Falle herum an

möglichst vielen verschiedenen Stellen in dichter und hoher Vegetation, auch dies, um

Verluste durch Vögel zu vermeiden.

3.4. MARKIERUNG

Zur Markierung der Falter wurden im Handel erhältliche Fingernagellack-Farben ver-

wendet, bei der Auswahl der 20 verschiedenen Farben wurde darauf geachtet, daß sie

auch nach dem Trocknen noch gut unterscheidbar waren; gleichzeitig sollten sie nicht

zu auffällig sein, um eine höhere Mortalität durch Vögel und andere Feinde zu verhin-

dern.

Von den beiden bei der Erstmarkierung aufgetragenen Punkten codierte der eine den

Fangplatz, der andere das Fangdatum.

Hierzu wurde der Thorax des Falters vorsichtig von Haaren und Schuppen freigepin-

selt, danach wurde die Farbe in Punkten von ca. 1 mm Durchmesser mit einem vorher

präparierten Pinselchen auf den Thorax aufgetragen, wobei darauf zu achten war, mit

dem Lack nicht in die Tergit-Fugen zu geraten, z.T. wurde zusätzlich ein "Sicher-

heitspunkt" auf dem Flügel angebracht. Einige kleinere Arten, vor allem Spanner (Geo-

metridae) konnten aus Platzgründen nur auf den Flügeln markiert werden.

Bei Wiederfängen wurde die den Tag codierende Farbe im Wurzelfeld des Flügels an-

gebracht, ab dem Zweitwiederfang waren die Tiere dann meist individuell erkennbar

und bedurften keiner weiteren Kennzeichnung.

Es zeigte sich, daß diese etwas kompliziert klingende Methodik bei einiger Übung eine

gute und rasche Markierung von Nachtfaltern zuließ: Von 23.492 markierten Faltern

mußten nur 17 wegen vermutlicher Schäden oder Behinderungen der Flugfähigkeit aus

der Wertung genommen werden.

Drei Fehlerquellen seien hier noch kurz besprochen:

- Obwohl die Farbe schnell trocknet, könnte es durch ein Abwischen beider Punkte zu

fehlerhaften Ergebnissen kommen. Der Anteil solcher Exemplare mit "Glatze”

betrug jedoch in den verwendeten Lichtfallen je nach Witterung ca. 2-5 %. Dieser

Prozentsatz war bei Arten, die nicht markiert wurden, genauso hoch. Eine bevor-

zugte "Glatzenbildung” bei markierten Tieren durch Abwischen der Farbe fand also

nicht statt. Es ist also nur eine sehr geringfügige Unterschätzung der Wieder-

fang-Quoten zu veranschlagen.

Abb. 3b: Mamestra w-latinum (d, HW, 16.6.87) mit zwei verschiedenfarbigen Markie-

rungspunkten auf dem Thorax.

- Verluste bzw. Beeinträchtigungen durch den Fangvorgang sind vermutlich zu ver-

nachlässigen: Einige Falter wurden 8-10mal gefangen und waren dennoch am Ende

in noch erstaunlich gutem Zustand.

- Verluste bzw. Beeinträchtigungen durch den Markierungsvorgang sind vermutlich

ebenfalls zu vernachlässigen: Ein erster Hinweis darauf ist die eben erwähnte Be-

obachtung. Weiterhin konnte dreimal beobachtet werden, wie markierte Falter eine

Kopula eingingen. Schließlich wurden einige markierte Falter (je 3 Noctuiden- und

Geometridenarten sowie eine Arctiide) in Gefangenschaft bei Fütterung gehalten und

die Überlebensdauern von 1-2 Wochen, in einem Fall von 16 Tagen zeigen, daß

diese Freilandfalter, die vor dem Anfliegen der Lichtfalle vielleicht schon einige

Tage alt waren, eine normale Lebenserwartung hatten. Ein 9 legte dabei seinen Ei-

vorrat ab.

3.5. MATERIAL

Folgende Werte stellen die Eckdaten für das 1987 und 1988 mit der Methodik des Le-

bend-Lichtfangs gesammelte Material dar:

49.072 Individuen (Macroheterocera)

462 Macroheteroceren-Arten (zusammen mit den Tagbeobachtungen und den Mel-

dungen aus den anderen Jahren: 514 Arten)

23.434 markierte Individuen (1989 384 weitere)

1290 Wiederfänge (1989 57 weitere)

Aus Tabelle 1 wird die Verteilung der gefangenen Individuen auf die verschiedenen

Monate des Jahres ersichtlich, in der Hauptflugzeit von Mitte Juni (11.6.) bis Ende

August flogen 81,6 % aller in den Lichtfallen ausgezählten Nachtfalter (Macrohetero-

cera).

Tab. 1: Verteilung der Individuen-Monatssummen in den Erhebungsjahren 1987 und 1988.

Distribution of the individuals over the months from January to Dezember 1987 and 1988.

Monat | Jan Feb März Apr Mai Juni Juli Aug Sep Okt Nov Dez

ee] - 1 309 2181 1836 9615 18.039 14.258 1764 643 458 6

4. ARTENLISTE

4.1. VORBEMERKUNGEN

Die Unterscheidung zwischen "Macro-" und "Microlepidoptera" entstand aus mehr äs-

thetischen Beweggründen der Sammler, die die kleineren "Motten" verschmähten. Diese

Einteilung entbehrt jedoch einer Grundlage in der Systematik. Einige üblicherweise zu

den "Großschmetterlingen" gezählte Familien, z.B. Hepialidae oder Psychidae müßten

bei genauerer Betrachtung der Verwandtschaftsverhältnisse ins System nahe bei

Schmetterlingfamilien gestellt werden, die zu den "Microlepidoptera" gehören. Dennoch

soll hier auf diese gängige Einteilung zurückgegriffen werden.

Hauptgegenstand der Untersuchung sind, wie erwähnt, die "Macroheterocera", worunter

alle "Großschmetterlinge"” mit Ausnahme der Tagfalter ("Rhopalocera") verstanden

werden. Die überwiegend tagaktiven Widderchen (Zygaenidae), Sackträger (Psychidae) und

Glasflügler (Aegeriidae) werden in Lichtfallen so gut wie nie erfaßt und werden daher

bei vielen der folgenden Auswertungen ausgeklammert. Die so eingegrenzte Gruppe von

Schmetterlingen bezeichnet MEINECKE (1984) als "überwiegend nachtaktive Groß-

schmetterlingsfamilien".

Die in der vorliegenden Untersuchung verwendete Nomenklatur und Systematik richtet

sich aus Praktikabilitätsgründen nach FORSTER & WOHLFAHRT (1955; 1960; 1971;

1981), da dieses Werk allgemein gebräuchlich ist. Neuere Erkenntnisse der Taxonomie

(z.B. LERAUT, 1980) sollen dadurch jedoch nicht in Frage gestellt werden. Lediglich

bei den Psychiden wurde einer moderneren Systematik gefolgt und beispielsweise die

früher zu den "Microlepidoptera" gerechnete Narycia monilifera mitaufgenommen. |

4.2. TAGFALTER (RHOPALOCERA)

Tab. 2: Im Untersuchungsgebiet 1974-1989 festgestellte Tagfalterarten (Legende/Häufızä

siehe Nachtfalterliste).

Species of Butterflies, recorded 1974-1989 in Oberschleißheim (Southern Bavaria). |

Häufigkeit ||

Nr. Art Häufigkeit "Ne. Art

1 Papilio machaon L. n-h 31 Clossiana selene Schiff. sh

2 Pieris brassicae L. h 32 Clossiana euphrosyne L. n-h

3 Pieris rapae L. h 33 Clossiana dia L. n-h

4 Pieris napi L. h-sh 34 Issoria lathonia h

>) Anthocharis cardamines L. h 35 Thecla betulae L. n

6 Gonepteryx rhamni L. sh 36 Nordmannia ilicis Esp. n

7 Colias hyale L. h 37 Strymonidia w-album Knoch 2

8 Colias australis Vrty. v 38 Strymonidia pruni L. 1

9 Erebia medusa Schiff. h 39 Callophrys rubi L. n-h

10 Melanargia galathea L. sh 40 Lycaena phlaeas L. n

11 Aphantopus hyperantus L. sh 41 Cupido minimus Fuessly n

12 Pararge aegeria L. n-h 42 Celastrina argiolus L. 1

13 _Lasiommata maera Il. h 43 Lycaeides idas L. h (Iokal)

14 _Lopinga achine Scop. n-h 44 Plebejus argus L. 2

15 Maniola jurtina L. sh 45 Aricia agestis Schiff. n

16 _Coenonympha glycerion Bkh. n 46 Cyaniris semiargus Rott. n

17 _Coenonympha arcania L. h-sh 47 Polyommatus icarus Rott. sh

18 _Coenonympha pamphilus L. sh 48 Lysandra bellargus Rott. n

19 Apatura iris L. n 49 _Lysandra coridon Poda h

20 Limenitis camilla L. n 50 Erymnis tages L. n

21 Vanessa atalanta L. h 51 Pyrgus malvae L. h

22 Vanessa cardui L. n-sh 52 Pyrgus alveus Hbn. 2

23 Aglais urticae L. sh 53 Spialia sertorius Hffmgg. z

24 Inachis io L. sh 54 Carterocephalus palaemon

25 Nymphalis polychloros L. 1 Pallas h-sh

26 Polygonia c-album L. h 55 Adopaea lineola O. n-h

27 Araschnia levana L. sh 56 Adopaea sylvestris Poda n

28 Mesoacidalia aglaja L. n 37 Qehiaden yenahu. Er ch

m Atsyaoispaphiacl, ah 58 Hesperia comma L n

30 Brenthis ino Rott. 1 ö

Aus Gründen der Vollständigkeit wird diese Großschmetterlingsgruppe, die eigentlich

nicht Gegenstand der Untersuchung ist, hier kurz aufgeführt. Bei einigen ökologischen

Mustern kann das Aufzeigen von Parallelen zu den Tagfaltern angebracht sein.

Die Tagfalterhäufigkeiten beziehen sich auf die addierten Jahressummen des ganzen

Untersuchungsgebietes und sind daher in dieser Form sehr verallgemeinernd dargestellt.

4.3. NACHTFALTER (MACROHETEROCERA)

In der Nachtfalterliste kam es ab dem 28.5.1988 am Fangplatz "WaS" aus zwei Grün-

den zu einer bisweilen starken Überhöhung der Werte: Durch (methodisch bedingte)

Wiederfänge sind durchschnittlich 18 % bei den betreffenden Arten zu veranschlagen. In

besonders krassen Fällen (»50%) wurden die Werte besonders ("mÜ”) gekennzeichnet.

Eine Überhöhung im Vergleich mit dem Fangplatz SiN erfolgte zusätzlich durch häufi-

geren Fang, das Ausmaß wird zumindest bei den in größerem Umfang markierten Arten

im II. Teil (Kapitel 9.2.) aus den Markierungstabellen ersichtlich. So ergibt sich bei-

spielsweise bei Mythimna impura aus einem SiN/WaS-Verhältnis von 4/32 nach der

Bereinigung ein Wert von 4/11, bei Lymantria monacha statt 2/ll nur 2/6 und bei

Arctia caja statt 2/13 sogar ein umgekehrtes Verhältnis von 2/1, bedingt durch ein an

WaS 10mal gefangenes d.

Einige besonders gekennzeichnete Nachweise stammen aus einer Bestandserhebung 1988

im Korbinianiholz (KOLBECK, in Vorber.).

Die Angaben zur Ökologie sind nur als eine approximative und vorläufige Charakteri-

sierung der Verhältnisse im Untersuchungsgebiet zu verstehen. Es wurden hierbei aus-

gehend von den Beschreibungen BERGMANNs (1951-1954), KOCHs (1984), GERST-

BERGER & STIESYs (1983) und REJMANEK & SPITZERs (1982) nach einer wegen der

Frage der geographischen Vergleichbarkeit sehr kritischen Betrachtung der Literaturan-

gaben die im Untersuchungsgebiet gemachten Beobachtungen interpretiert. Die Über-

tragbarkeit dieser Aussagen auf andere Gebiete kann, wenn überhaupt, jeweils erst

nach einer Überprüfung konstatiert werden, da oft ein und dieselbe Art in verschiede-

nen Höhenstufen, in verschiedenen Breitenlagen u.s.w. unterschiedliche ökologische An-

passungen zeigt.

Eine Schwierigkeit stelli auch der Umstand dar, daß die Einnischung verschiedener Ar-

ten oft von ganz unterschiedlichen Faktoren abhängt: Manchmal richten sie sich nach

mikroklimatischen Verhältnissen (z.B. die starke Sonneneinstrahlung bei polyphagen

Arten, die sowohl auf Heiden wie auch in Mooren vorkommen), oder sie ist fast nur

durch die Raupenfutterpflanze bestimmt (viele monophage Arten).

Viele Arten sind recht euryök. Aus diesem Grunde erfolgte in den meisten Fällen die

Angabe mehrerer möglicher Lebensräume, die jedoch als Anhaltspunkt stets auf ein

Hauptvorkommen "präzisiert" wurde.

In der Artenliste (Tab. 3) werden folgende Abkürzungen und Symbole verwendet:

Legende/Fangplätze:

SiS = Siedlung Süd

SiM = Siedlung Mitte

SiN = Siedlung Nord (Garten vor dem

Haus)

WaS = Wald Süd (Waldstreifen hinter

dem Haus)

WaM= Wald Mitte

WaN = Wald/Wasserwerk Nord

WNw= Wald/Wasserwerk Nordwest

WNo = Wald/Wasserwerk Nordost

HO = Halbtrockenrasen (Nord-) Ost

(Waldrand)

HM = Halbtrockenrasen Mitte (Offenland)

HW = Halbtrockenrasen West

Au = Auwald am Würmkanal

We = kleiner Weiher mit Moorbirken-

wäldchen (Torfeinfang)

Mb = Moorbirkenwald (Birket) im Inne-

ren

vor 1987:

Gar = Garten (1974-1986)

Mo = Moos zwischen zwei Entwässe-

rungsgräben, 250 m vom Birket

entfernt (1985)

HO = siehe oben (1986)

HM = siehe oben (1986)

weitere Fundorte:

Schw.holz = Schweitzerholz

Regattastr. = Ruderregattastrecke

Entw.graben = Entwässerungsgraben

Legende/Ökologie:

Ub = Ubiquisten

Wf = Wanderfalter

Wf(RR) = Wf nach RETZBANYAI-RESER

(siehe Literaturverzeichnis)

mGr = Arten der mesophilen Grasland-

schaften

Agr = Arten der unter

stehenden Flächen

Agrarnutzung

Geb = mesophile Arten der Hecken, Ge-

büsche und der Waldränder

Geb/R = Geb, jedoch bevorzugt an rude-

ralartigen Standorten

W = mesophile Arten der Wälder

WL = Laub- und Mischwaldarten

WN = Nadel- und Mischwaldarten

Hy = Hygrophile Arten im weiteren Sinn

Xe = Xerothermophile Arten im weite-

ren Sinn

Legende /Häufigkeiten:

1-3 = unter 4 Individuen

V = vereinzelt

n = nicht selten

h = häufig

sh = sehr häufig

sonstige Abkürzungen:

E = Fund von Eiern/Eigelegen

R = Raupenfund

P = Fund von Puppen bzw. Kokons

dd‘ = AmLicht nur dd

909 = AmLicht nur 99

Ex. = Exemplar

T = Imagines tag- bzw. dämmerungs-

aktiv, daher am Licht nicht

optimal erfaßbar

Däm = (Abend-)Dämmerungsflieger / in

der Dämmerung gekäschert

zus. = zusätzlichle, er, es)

Gen.prp. = Genitalpräparat (nur in wich-

tigen Fällen angegeben)

am L.n. opt. = am Licht nicht optimal

F.F.w.r. = Fänge in der Flugzeit wenig

repräsentativ

mÜ = methodisch bedingte Überhöhung

der Werte

In der Nachtfalterliste bezieht sich je-

weils die erste Zeile der ersten vier

Spalten auf das Erhebungsjahr 1987, die

zweite Zeile auf 1988.

Nr. Art

Nolkdae

69 Celama

confusalis

H.-S. 1847

Lymanträdae

60 Dasychira

selenitica

ESP. 1789

61 Dasychira

udibunda

. 1768

62 Orgyia

recens

HBN. 1819

63 Lymantria

monacha

L. 1768

64 Euproctis

chrysorrhoea

L. 1768

65 Porthesia

similis

FUESSLY 1776

Arotiidae

66 Cyboslia

mesomella

L. 1768

67 Miltochrista

miniata

FORSTER 177

68 Lithosia

quadra

L. 1768

69 Eilema

depressa

ESP. 1787

70 Eilema

lutarella

L. 1768

71 Eilema

complana

L. 1768

72 Eilema

lurideola

ZINCKEN 1817

73 Systropha

sororcula

HUFN. 1766

74 Atolmis

rubricollis

L. 1768

76 Phragmatobia

fulıginosa

L. 1768

76 Spilarctia

lubricipeda

L. 1768

77 Spilosoma

menthastri

ESP. 1786

78 Diacrisia

sannio

L. 1768

79 Arctia

caja

L. 1768

Endrosidae

80 Pelosia

muscerda

HUFN. 1766

Notodontidae

81 Harpyia

CL. 1769

82 Cerura

vinula

L. 1768

(R)

[>78]

WALD

ın

174

162

[+17]

107

109

Gar

n-h

VOR

1987

Mo HO HM

- 16 1

- 29 -_

6 -_ -

6651

2 - —

- 4 -

- 64 1

A ©

- 6 =

- 60 1

m 17 -

37 32 23

20 a

4 20 4

ı 08

67 3 8

en, 1 >

- 58

UKO-

LO-

GIE

Von)

eed)

Vsed)

Geb)

(m6r,

Geb

(wı)

Geb/R

FLUG-

ZEIT

E4-A6

M6-E6

A7-M8

A7-A9

M7-E7

A7-M8

A6-E7

E6-AB

A7-AB

E6-A9I

E7-AB

E6-E8

M7-M8

M6-A7

A6-M7

AB-EB,

A7-E8

E6-M7

A6B-M7

E6S-A7,

AB-A9

M7-M9

A7-M7

E4-M7

BEMER-

KUNGEN

1989 ein zus

Ex. im Korbi-

nianiholz

am 29.56.89 im

Mallertshofer

Holz ein 9

dd, auch T,

am L.n. opt.,

Birket 1083 E

S-Bahnhof

1987 1 Ex.

Im Mallertsho-

ter Holz 1989

9 Ex., auch T

® 1986 I R

1. Generation

selten

Birket 1983 h

2. Generation

1988 h, auch T

°ml.

R im Schw.-

holz h

SIEDLUNG WALD VOR UKO-

Nr. Art ı Garten | Wasserwerk 1987 LO- FLUG- BEMER-

ZEIT KUNGEN

SiS SIM} SIN Gar Mo HO HM | GIE

83 Stauropus = u nh- - - - - - - 6 ia = gan 4 A6-M7 *S-Bahnhof

fagı - 2 Ara 1 3 - x WAREN | Geb) 1 zus. Ex

L. 1768 1986

84 Hybocampa - - - - - - 1 - - - 2 - == Ws M6-A6 1989 im Fran-

Ba) = = S 1 48, < u - = Geb) zanenhölz| =

. 1776 an WaS je

u frisches \

85 Gluphisia = 1 = = - = = = m { 2 - Il 2 SR M6-A7 1989 m. zanı

crenata = = = = = = = = zosenhölzl +

EsP. 1786 eb] an We je ein

86 Drymonia - - 1 4 — = 8 = 1 - v - 19 = L M6-E6

trimacula - 3 - 1 1 6 - - = = Geb)

ESP. 1786

87 Drymonia - - 2 1 7 1 16 - - - h -" 104 - vi E4-A6 °FEFwe,

ruflcornis - 1 1 2 - Ur - IR | Geb) im Kapuziner-

HUFN. 1766 holz 1989 h

88 Peridea — = 3 6 6 3 21 = 3 18 n-h - 656 - ws E4-M6 ®am Rand des

anceps 3 3 2m- 743 60 1 1 Du = Geb) Birkets

GOEZE 1781

89 Pheosia = = = - = 1 1 1 7 12 L) Ze 22T H E6-EB,

tremula = - = 1 = = 6 4 10712 m. M7-M8

CL. 1769 eb/R)

90 Pheosia 1 2 2 = - 1 1 2 2 43 n-h26b 6 4 H A6B-A7,

noma - 14 1 - 1 2 = 1 2 2 {w M7-M8

1972; eb

91 Notodonta = = = _ = = = = = - 1 a u = 7: E7

hoebe - - - - - - - - - = Geb)

IEBERT 1790

92 Notodonta — 1 2 1 = 2 = 2 1 1 v-h- 8.772 m A6-A7, SiN 1987 R

dromedarius 1 2 3 1 6 - - 1 3 WE WL) M7-M8

L. 1767

93 Notodonta = 2 = 2 1 2 1 10 6 7 3 A612 Geb/R AB-E6

ziezac 1 1 I u 1 aa (Ub) A7-ME

L. 1768

94 Leucodonta = = - 2 1 2 1 = = 4 1 - 2. }= 7: M6-A7 1989 an SiN

bicoloria = 2 - 1 = 6 = = - 1 Geb, M6 Io

SCHIFF. 1776 Hy

96 Odontosia - - - 1 - - - = = - - =e2 .}=- wL E4-A6 .FFwer

carmelita = - = = = - = = - -

EsP. 1799

96 Lophopteryx = - 2 4 2 1 6 = 4 9 ten = 117 7- Mr 6-M8

camelina 2 13 6 1 7. 12 1 4 17 6 Geb) 2 Gene-

L. 1768 rationen)

97 Lophopteryx - - 1 - = 1 9 = - 1 1 - 91 L M6-A8® Regattastr

AR. - 3 - 1 1 - - 1 1 - Weed) 1983 IR

EsP. 1786 = a

98 Ptilophora = = z = E S = = = - - ld no ws EIO-MI1

umigera - 2 - - 1 7 = 7 Geb)

CHIFF. 1776

99 Pterostoma 1 - 2 - - = = 2 4 | -v 6 1 4 | Geb/R Bar ® am Rand

alpina - 2 2 2 Zu ve 10 2 (Ub) 2 Gene- des Birkets

e 1769 rationen)

100 Phalera — = = 10 3 1 1 1 2 28 n-h 77720 °2 Geb E6-E7

bucephala - 6 4 1 1 6 1 2 7“ (wu)

L. 1768

101 Clostera 1 2 - = = 2 1 3 6 3 = 7 2 4 | Geb/R E4-Eß6,

curtula - - - 1 2 - - 3 - 6 (Hy) M7-AB

L. 1768

102 Clostera - - _ - - - - = 1 2 - 1 = cz "% AB, 1989 3 zus

anachoreta u - = = = = = = = = eb/R) E7-AB Ex. am Fran-

F. 1787 zosenhölzl

103 Clostera > = = = 1 1 1 3 4 3 - 2 - 6 Geb/R E4-M6, HM 1986 I R

igra -. - 1 1 - - - 2 = 03 (Hy) A7-E8 ®S-Bahnhof

FN. 1766 1988 1 Ex

Zygaenidae

104 Lictoria = = = = = = = = = = = = = - = - - - r M6-M7 T, 1989 im

achilleae = = = _ = = = = = = = = = = Xe Mallertsho-

4 Esp 170 Geb) a ie Holz h

105 Thermophila = = = = — = = = ° = = = = = = = . = =

meiion ee Br | Fe ee oen,

SP. 1793 r

106 Zzygaena z = = - + . + + . . . - + - - ne Geb E6-M8 T

IDESndlee = ie = = + + + + . . . Zueer > Imer,

. 1768

107 Huebneriana Se a de = = = = U Sl - - 0. > r M7® T, °1989 im

lonicerae = = = = = = = = = = = - = - Xe Mallertshofer

SCHEV. 1777 5 eb) Holz 1 Ex.

108 Burgeffia =/#= "1 > = = = zer . ° Z =. .= ar Na)e ie ne E7-M8 T *®200 m

ephlaltes - .- - - - - _ z + + = u mGr) vom Garten

L. 1767 entfernt

Cochiidiidae

109 Apoda 2 1 6 10 27, 44 22 21 24 che 219223 ur M6-E7

limacodes 2 10 2 2 2 17 2 8 1 2 WL

HUFN. 1766 Geb]

110 Heterogenea = 0-0 = = za pl ZIP) IR Sun = = - SE Wi: E6 am L. n. opt

ne a TO DO ee 1087 3. 1988

SCHIFF. 1776 40 Kokons

Nr. Art

Sphinsklae

111 Mimas

tiliae

L. 1768

112 Laothoe

Bepul

. 1768

113 Smerinthus

ocellata

L. 1768

114 Herse

convolvuli

L. 1768

116 Sphinx

ligustri

L. 1768

116 Hyloicus

Inastri

. 1768

117 Dellephila

elpenor

L. 1768

118 Deilephila

orcellus

. 1768

119 Macroglossum

stellatarum

L. 1768

120 Hemaris

tuciformis

L. 1768

Ihyatirkdae

121 Habrosyne

yritoides

FN. 1766

122 Thyatira

atis

L. 1768

123 Tethea

fluctuosa

HBn. [1803]

1796

124 Tethea

duplaris

L. 1761

125 Tethea

SCHIFF. 1776

126 Tethea

ocularis

L. 1767

127 Kohrploca

lavicornis

L. 1768

Drepanidae

128 Drepana

alcataria

L. 1768

129 Drepana

lacertinarla

L. 1768

130 Drepana

binaria

HUFN. 1767

131 Drepana

cultraria

F. 1776

132 Cilix

laucata

COP. 1763

Saturnidae

133 Eudia

avonia

. 1768

Von

- 8 -

3 9

2 =

v-n 1 6

f-v - -

3 a

v-h- +

1° - =

v-n 12 11

1 SS U

n 10 1

1 - =

n mel-

n IEE16

1 23

n-h 9 126

one 3

1 a

UOKO-

LO-

GIE

Med)

bebr)

Geb/R

Ub/Wt

Agr)

Geb

N .

Geb)

nA b/R,

mGr)

Ub/Wt

Xe)

Geb

(w)

Fed)

Geb

Ver ®)

Hr.

Geb]

Geb

(wı)

Med)

Geb®®

(wı)

Von)

ir,

Veen)

Geb®®

FLUG- BEMER-

ZEIT KUNGEN

AB-E6 T: SIN 1987

1 Ex.

6-A8 "Mo" 1983

vermutl. 3 Ex.

Gener.)

M6-M7 "Mo" 1983

2 Ex

°m.d.

A7-A9

ES-M7 SiN 1977 R

AB-M8 SIiN 1988 R

“wt (RR)

M6-M7 WaN 1982 1

Ex. ruhend (T}

M6-M7,

M8 Ne

A6-M7 T, 1983 ein

AY-AIO Ex. am Licht

° 1974

E6-AB

E6-M7, HO 1988 R

M7-E8

A7-E7 ® auch Moore

und Heiden

M6-M8

A6-M8

E65

E3-E4 ®FEFwer

se auch Moore

und Heiden

2 Gene-

rationen)

6-M8

2 Gene-

rationen)

AB-M6, 3. Generation

M7-EB, nur partiell

AB5-E5,

A7-E8

E4-A6

am L:99, dd

nur T, Po dd

im Mallertshofer

Holz, ®F.F.wer.

@® auch Moo-

re und Heiden

Nr. Art

Lasiooampidae

134 Malacosoma

neustria

L. 1768

136 Poecilocampa

opuli

. 1768

136 Pachygastria

ariloll

SCHIFF. 1776

137 Macrothylatla

rubi

L. 1768

138 Philudoria

otatoria

. 1768

139 Cosmotriche

lunigera

ESP. 1784

140 Dendrolimus

ini

. 1768

Psyohidae

141 Sterrhopteryx

fusca

HAw. 1809

142 Psyche

crassiorella

BRUAND 1849

143 Psyche

casta

PALLAS 1767

144 Proutia

betulina

Z. 1839

146 Bacotia

seplum

SPR. 1846

146 Talaeporlia

tubulosa

RETZ. 1783

147 Dahlica

triquetrella

HBN. 1812

148 Narycia

monilitera

GEOFFR. 1786

Asgerikdae

149 Chamaespheclia

empiformis

EsP. 1783

Cossklao

160 Cossus

cossus

L. 1768

151 Zeuzera

prsa

. 1761

Heplalidao

162 Hepialus

umull

L. 1768

153 Hepialus

sylvina

L. 1761

164 Hepialus

ecta

L. 1768

{R)

WALD

er:

3 361

3 = —

eier

36 ire2

R - = -

Eike Dee

RR al IR}

Ua)

1 =

1 1

3 = 4

2 6

1 3 ng

zei

a 1

8 12

1,2

{R)® (R)

-a

23, 2

(R)

Gar

v({R)

h(R)

1987

Mo HO HM

- (R)

UKO-

LO- FLUG- BEMER-

AR E6-E7 Im Moos als

eb) R mehrfach

7 MIO-MII *®FF.wer.

Geb)

r AB-M9 IgG » am

6) L.{n) HH

viele R (1988

mer M6-M7 amLl. va. 99,

Geb/R) auch T, im

Bergliwald als

R regelmäßig

1989 wieder

Hs) A7-EB

Web) T

WN A6-M7

Hm E6

Geb

(w)

Fed)

Geb

(Hy)

Geb)

Geo)

up)

Geb)

Ten)

Geb M7

(Hy)

Geb A7-M8

(wı)

H A6-AB

{ms P

rc A8-M9

Geb,

Geb A6-A7

(Hy)

1 cd im Garten

1989 1 Ex. im

Mallertshofer

Holz

T, d4,

® am Licht

T, dd

T, dd, * auch

im Schw.holz,

®® im Ruderal

T,dd

T, dd, im

Mallertshofer

Holz 1989 1

Sack

IhonSgenese)

dd «+ e-

lügen 1989

1 Sack im Mal-

lertshoter Holz

T, Bergiwald

1982 1 Ex

® Entw.graben

1 Ex., dort 1983

Ex, amtL.n

opt.

®Däm. 10 Ex,

®®Däm. ein zus.

Ex., Bergiwald

1982/86 T2Ex

WALD

Nr. Art Wasserwerk

ı Garten |

SiS SIM, SIN |WaS} WaM WaN WNw WNo

Noctuldae

Noctuinae

155 Euxoa - - = = e S & = ei =

obelisca - 1 > > E 2 1 1 2. Er

SCHIFF. 1776

156 Euxoa - - 1 o - > a 5 2 a

tritici - - - 1 - = - 1

L. 1761

157 Euxoa - 1 _ o = S = = 5 u

nigricans - - - _ 5 a = 1 1 je

L. 1761

Euxoa _ - - - = 2 - 2 2 -

aquilina - = 16 = 6 = = 2 -

SCHIFF. 1776

Scotia > 3 2 1 = 2 3 1 4 -

segetum = = = = = < 1 1

SCHIFF. 1776

Scotia 3 28 32

clavis 27

HUFN. 1766

161 Scotia 2 38 36 29 9 62 66 32

u malionis 32 84 39 4 29 62 45

. 1768

Scotia 4 94 8 29 3

ipsilon 3 46

HUFN. 17686

Ochropleura 2 41 23 21 3 64 40 39 119 24

lecta 124 1656 18 28

. 1761

158

159

160 48

176 14 20 7 33 32

162

[e,}

163

164 Eugnorisma - - - - - - = - = =

depuncta - - - - - - - = > en

L. 1761

166 Rhyacia - - - - - - - _ > E

ucipeta - - 1 - - - - — = =

SCHIFF. 1776

166 Rhyacia - - - - - - - - = =

simulans - - _ - = - & 1 3 &

HUFN. 1766

Noctua 9 38 18 80 4 26 45 26

ronuba 48 | 1011 76 94 42 12355135395 35 4

. 1768

Noctua - 7 5 6 _ 2 3 2

comes 5

Hen. [1813]

167

168

169 Noctua - 1

fimbriata

SCHREBER 17689

-n

170 Noctua 1 1

janthina 2 16 1 - 2

SCHIFF. 1776

171 Spaelotis - - - - - - - = = =

ravida = - 1 - 1 = = 1 & en

SCHIFF. 1776

172 Graphiphora = - - - - - - > _ w

augur 1 2 - - - = = = = a

F. 1776

173 Paradiarsia = - - = - 1 - = 1 7

unicea - - - - - 1 e = e =

BN. [1803]

Diarsia = = 2 g 3 2

mendica 1 23 4 ı0 4 g - = 1 =

F. 1776

Diarsia - 1 7 18 4 6 6 = - 1

brunnea 6 120 ih 9 8 3 = - 3 2

SCHIFF. 1776

174

176

176 Diarsia _ — _ 1

rubi = 3 4 = 1 3 S 6 1 1

VIEWEG 1790

Amathes - 82 9 16 6 30

c-nigrum 48 | 336 180 32

L. 1768

Amathes 8 6 12 23 3 44 25

ditrapezium 19 127 60 31 43 28

SCHIFF. 1776

Amathes 8 12 10

triangulum 69

HUFN. 1766

Amathes 1 2 2 10 - 22 31 1 6 8

baja 1

SCHIFF. 1776

181 Amathes 2 1 1 =

sexstrigata 1 22 141 62

HAWw. 1809

177 21 96 4

208 138 72 176 27 9

178

32 26

179

641 86 42 80 46 12 re ]

180

106 16 46 38

173 138 63 82 bzı

VOR

1987

Gar Mo HO HM

h-sh - 106 21

h-sh 9 66 16

h-sh 16 20 14

h 272398

VchEalZzse2

VE]

3 =I B2ı05=

v-hla= 362 5-

1 3 U =

n-sh 22° 13 16

h 126 62 11

OKO-

LO-

GIE

FLUG- BEMER-

ZEIT KUNGEN

M8-E8

M7-M9

A8-M9

M7-M8 Im Garten nur

1983

A6-E7, ewt (RR)

E8-MIO

A6-E7

MB5-E7, 2. Generation

E7-E8 nur sehr partiell

.wf AR)

A6-A11

A5B-M7,

M7-E9

A8 ®auch Heiden

und Moore

M8 ®wt (RR)

EI “wt (RR)

A6-AI11

A8-E9®® ®Wf (RR)

®®1 Ex. bereits

M7-M9 ®Ww# (RR)

M7-E9I ewt (RR)

A7-M7, ®Wf (RR)

M9-E9

M6-M7

E6-M7 WaN 1989

1 Ex.

E6S-M7

M6-M8

MB-M6,

A7-A9

M65-A7, °"Mo” am

E7-MIO 6.6.83 11 yo

swf (RR

M6-M8

A6-AB

M7-M9

E7-A9

VOR

BR De FLUG- BEMER-

Mo HO HM K EN

182 Amathes bb 47,3 7 a 33 1827 2 h 7 93762 r M8-E9 ®| frisches E

xanthographa 4 36 32639 E17. 18 2107 226 6 1 = b, MIO® ? am 16.10.88 x

SCHIFF. 1776 Ub

183 Phalaena = = = = = = = = = = 1 1 Sn Geb M7-Ag® *®WaS 1989

typica - 3 Je ee u- - 1 (Hy) schon M6

L. 1768

184 Eurols - 1 - - 1 - - - - - 1 a r A7-EB

occulta = 2 = = = = = = = = W,

L. 1768 eb)

185 Anaplectoides u - _ 3 5 B 5 2 = 567727 7Geb A6-E7

rasina - 2211 3,4 - - 1 3 3 (mGr)

CHIFF. 1776

186 Cerastis = 1 1 1 2 48 6 6 2 = v | Gr E3-E6®® *®FFwre,

rubricosa - 2 39 26 26 10721 4 = = Geb) ®® am Ende der

SCHIFF. 1776 Beuel! frische

tücke

187 Cerastis - - - 1 - 8 1 = - z = -® 2 - I Geb M4-E5 ®FF.wır.

leucographa 1 - 1) =) 3 1 - - - -

SCHIFF. 1776

188 Mesogona — - - - - - - 4 = = = Ze We >) Geb/R EBS-E9

oxalina - - - - - - 1 - - 10 (Hy)

Hen. [1803]

Hadeninae

189 Discestra = ehe = = 3 1 = il = 3 ER TZE2E I EUHBAWIZGEBZAG 2. Generation

trifolü - 2 34 = ee! - > Agr) M7-Ad zahlreicher

HUFN. 1766

190 Polia = 1 - = 1 14 3 1 1 = v 3 310 Diet M6-AB

bombycina - 2 zit 42 10 AA = Geb)

HUFN. 1766

191 Polia ne} Bm er 1:03) = 3222, |5Geb A6-E7

nebulosa = 12 3 2 7 x = = 2,

HUFN. 1766

192 Pachetra - - - - - - 6 35 44 = = = 3 24 m M5-E6 WaN 1989

sagittigera - . - - - 3, Hr S15 -.- Xe) 2 Ex.

HUFN. 1766

193 Sideridis = - = - = - = 1 4 = = => Kal = M5-A7

albicolon - - - - - - - 1 == mGr)

Hen. [1813]

1809

194 Heliophobus = = - - - - - 4 4 = - - =. 2 e A6-M7

reticulata - - - - - 1 4 3 - .- mGr)

GOEZE 1781

196 Mamestra - 2 - 1 - 1 4 1 v = ia r2 gr® Ba ®wt (RR)

brassicae - 30 1 1 6 AA 6 - = mGr) (2 Gene-

L. 1768 rationen)

196 Mamestra 1 10 g 8 3 13 2 1 = - hr 28, ai A6-AB

ersicariae ıo| a7 De DER sr 5 (eb,

. 1761 na

197 Mamestra Ze] En SEN Te 1 3 - =. 2 | Geb E6-M7, *2. Generation

contigua = 3 Si u 2 2m - - (mGr) A8-A9® nur partiell

SCHIFF. 1776

198 Mamesitra = - - 1 - 65 1 7515 - -v - 1% r M5-A7

w-latinum = 3 = = = _ 27210 = = ee

HUFN. 1766

199 Mamestra - 2 4 U - 2 2 1 3 - - - M6- 198 E7

thalassina 2| 35 an © a ea 2 een En user

HUFN. 1766

200 Mamestra - 14 4 2 1 12 8 19 30 1 veh. (au aeg U E4A-E6, 2 vorzeitig ge-

suasa 6 58 49 10 658 8 920 167 7) = mGr, M7-M9 schlüpfte Ex. A7

SCHIFF. 1776 Agr)

201 Mamesitra = 7 & 6 1 1 = = = - - 6-AB = i I-

oleracea 3 | ee on N ee Dh 2 0 a

L. 1768

202 Mamestra sa - = JS, = ae =, y®, = = r A6-M7 *nur vor 1982

isi - - - - 1 = 2 = - - ur: r,

. 1768 Geb)

203 Hadena Zen. < == = Pr Ze = ev = - m a E5S-E6, *HM 1986 zu-

rivularis 5 2 ei BE CE == 1 - =» Agr) E7-A9 sätzlich I R

F. 1776

204 Hadena Sn = = = 5 = = 6 - 1 - .-..- Gr M6-M7 1989 wieder

lepida - - - - - - 1 3 - = Xe) 1 Ex. im Garten

EsP. 1790

2056 Hadena = = = 1 = = = = 1 - t-v - u r M6-M7

compta 1 1 - - - - = 1 - - Tee)

SCHIFF. 1776

206 Hadena = z 7 - - = 7 = 2 - 1 1 NE mGr E6

confusa Z = = = = 7 = = - =

HUFN. 1766

207 Hadena = = 1 = - = 1 = 2 - 1 - 1 _ r M6-AB, *1 Ex. 1987

bicruris - 1 - - Z = = = - .- Ted) A9*®

HUFN. 1766

Nr. Art

208 Lasionycta

209 Eriopygodes

imbecilla

F. 1794

210 Cerapteryx

raminis

. 1768

211 Tholera

cespitis

SCHIFF. 1776

212 Tholera

decimalis

PoDA 1761

213 Panolis

flammea

SCHIFF. 1776

214 Orthosia

cruda

SCHIFF. 1776

216 Orthosia

Bepul

TROM 1783

216 Orthosia

racilis

CHIFF. 1776

217 Orthosia

stabilis

SCHIFF. 1776

218 Orthosia

Incerta

HUFN. 1766

219 Orthosia

munda

SCHIFF. 1776

220 Orthosia

gothica

L. 1776

221 Mythimna

turca

L. 1761

222 Mythimna

conigera

SCHIFF. 1776

223 Mythimna

ferrago

F. 1787

224 Mythimna

albipuncta

SCHIFF. 1776

226 Mythimna

vitellina

Hen. [1808]

226 Mythimna

udorina

CHIFF. 1776

227 Mythimna

impura

Hen. [1808]

228 Mythimna

allens

. 1768

229 Mythimna

I-album

L. 1767

230 Leucania

comma

L. 1761

Amphipyrinae

231 Amphipyra

Buyamigea

. 1768

232 Amphipyra

berbera

RUNGS 1949

233 Amphipyra

tragopoginis

CL. 1769

E20}

-a

-._

aan

-oa

Gar

= Sud

n-h 1 27 130

h eg?!

nh -®6 2

2 -ı _e _e

h-she_u2232

hzsh2 2213556:

Vo Dee u

sh -® 29 20

Fi dd A

2 a

Dh 3

v en

1 -_ = =

n® Isa1bzEg

Vones2 02507017,

1° =. -_ -

1 EEE HATGT,

h 3 14 -

1 - - -

h 9 14

FLUG- BEMER-

ZEIT KUNGEN

Geb M6° °im Garten im

(mGr) Juli” 1983

Hy E6 1989 ein Ex

im Franzosen-

hölzl

u M8

Geb,

mGr A9-M9

mGr E8-M9

Sn M3-A6 ®F.F.wer.

Geb)

„: M3-M5 *F.F.wer.

Geb)

Geb M3-M4 ®FFwer.

(Hy)

ur A4-M5E *F.F.wır.

Hy)

U M3-M6 1987 noch am

WL 22.6. 1 Ex

eb] ®FF.wrr.

SR A3-M6 1982 noch am

WL 22.6. I Ex.

Geb] SFFwı

“RE M3-E4 Birket 1983 1 R,

Geb) HO 1987 2 R

®F.F.wır.

ur A3-E6 ®FEF.wır.

Geb)

H M6-E7

(Gen,

mGr

"5 5 E6-E7

Me E6-M8 ®Wf (RR)

Geb)

EN A6-M9

mGr) (2 Gene-

rationen)

e/Wt E7

mGr)

Hy E6-E7

"ll,

mGr)

Ub)

Hy)

mGr

Weed)

ee)

M6-AB®®, ®früher z.T

A8-E9 fehlbestimmt

2. Gen. e®A6 einzeln

nur part. Entwgr. "87 1Ex

A6-M7°®®, ®früher z.T

A8-AIO fehlbestimmt

°®E7 einzeln

M6 ®1977

M6-M7 1989 WaN

1cd

A7-MIO im Juli nur

einzel

®w#t (RR)

AI ®W#(RR), Kor-

binlaniholz 1988

2 Ex. Gen.prp.:

KOLBECK

E7-MIO | Ex. am 11.7.

1988)

236

238

239

240

241

242

243

244

245

246

247

248

249

250

261

262

264

266

256

267

2568

269

260

Art

Rusina

ferruginea

EsP. 1786

Talpophila

matura

HUFN. 1766

Euplexia

lucipara

L. 1761

Phlogophora

meticulosa

L. 1768

Ipimorpha

retusa

L. 1761

Ipimorpha

subtusa

SCHIFF. 1776

Enargia

aleacea

SP. 1788

Enargia

ipsilon

CHIFF. 1776

Cosmia

affinis

L. 1767

Cosmia

trapezina

L. 1768

Cosmia

yralina

CHIFF. 1776

Auchmis

comma

SCHIFF. 1776

Actinotia

Bann

L. 1769

Actinotia

hyperici

SCHIFF. 1776

Apamea

monoglypha

HUFN. 1766

Apamea

lithoxylea

SCHIFF. 1776

Apamea

sublustris

Esp. 1788

Apamea

crenata

HUFN. 1766

Apamea

characterea

HBn. [1803]

Apamesa

lateritia

HUFN. 1766

Apamea

remissa

HBn. [1809]

Apamea

unanimis

HBn. [1813]

Apamea

anceps

SCHIFF. 1776

Apamea

sordens

HUFN. 1766

Apamea

scolopacina

EsP. 1788

Apamea

ophiogramma

ESP. 1793

Oligia

strigilis

L. 1768

SIEDLUNG

ı Garten |

SiS SM) SiN

WaS| WaM WaN WNw WNo

WALD

Wasserwerk

1 68 43

5 ©

IN3:

Su =>

3 1 3

5 A v4

EL a

al 2

Een

Ei O8

6 9 8

23 13 24

21°

VOR

1987

Gar Mo HO HM

(EVA IE Fe]

- Me

= Sn

1 an =.

- 1 - -

h BIR217 2

ch 92 74. =

v =. al

v Er

v - 9 37

3 a |

v=h#3 2 =

= are HE

h 2 3

OKO-

io: FLUG- BEMER-

ZEIT KUNGEN

GIE

” E6-E7

Es:

mGr

mGr M7-M8

Geb E6-E7

en E6 1. Generation

mGr) M8-All nur 1 Ex. 1986,

1988 1 Ex. am

S-Bahnhof.

Geb/R ET7-A9 Regattastr.

Hy) 1983 2 R

H E7-M8

(Geb/R)

Geb® E7-AB ®v.a. Moore

und Heiden

H M7-E7

(Geb)

Geb Korbinianiholz

(wL) 19 Ex

KOLBECK

“s A7-M9 ®Birket-Rand

Geb) 1988 3 Ex

Geb E6-AB

(wı)

Geb A7-E8

(Xe)

> E6,

mG([, AB-M8

Geb

Xe M8

A - M6-A9

eb)

g\ A6-AB

mGr)

mGr A6-A7

mer E6-E7

Geb)

Geb A6-E7

(mGr)

mGr AB

Gr E6-E7

Hy)

H E5-E6

(mGr)

gr E6-E7

mGr)

gr E6S-M7

mGfr,

Geb M7-E8

(wıL

mGr)

Hy E7-M8

Geb E6-E7

(mGr)

auch T

®wt (RR)

WaN 1989 ein

Ex.

® "Mo"

1983 3 Ex.

261 Oligia

versicolor

BKH. 1792

262 Oligia

atruncula

SCHIFF. 1776

263 Miana

furuncula

SCHIFF. 1776

264 Mesapamea

secalis

L. 1768

265 Mesapamea

secalella

REMM 1983

266 Photedes

minima

HAw. 1809

267 Photedes

extrema

Hen. [1809]

268 Photedes

ftluxa

Hen. [1809]

268b siehe Addenda

269 Luperina

testacea

SCHIFF. 1776

270 Amphipoea

oculea

L. 1761

271 Amphipoea

ucosa

FRR. 1830

272 Amphipoea

lucens

FRR. 1846

273 Hydraecia

micacea

ESP. 1789

274 Gortyna

flavago

SCHIFF. 1776

275 Celaena

leucostigma

Hen. [1808]

276 Nonagria

typhae

HNBG. 1784

277 Nonagria

nexa

HBn. [1808]

278 Rhizedra

lutosa

Hen. [1803]

279 Meristis {

trigrammica

HUFN. 1766

280 Hoplodrina

alsines

BRAHM 1791

281 Hoplodrina

blanda

SCHIFF. 1776

282 Hoplodrina

ambigua

SCHIFF. 1776

283 Atypha

ulmonarls

SP. 1790

284 Spodoptera

exigua

Hen. [1808]

286 Caradrina

morpheus

HUFN. 1766

286 Paradrina

clavipalpis

ScoP. 1763

287 Eremodrina

ilva

ONZ. 1837

288 Agrotis

venustula

Hen. 1790

-n

18 6

116

87 29

14 =

U 2

3 1

4 =

34 8

7 3

=, 6

4 6

33311

6 1

4 =

23 26

8°

vu NR

BTROK-

KENRASEN

-n

"DACH.

MOOS’

n-h

v-h

VOR

1987

Mo HO HM

V..@ =

2 00

a1 -

3) - =)

= 3 fe)

289 =

716 16

6u17

Slee

SEE 2

- 26 34

67 48 18

I 2078

eb)

Teen,

mGr

Geb

(mGr)

Teer)

eye

Xe/Wt

"Imgı.

Geb,

r/Wt

Agr)

Ab)

FLUG- BEMER-

ZEIT KUNGEN

M6-E7 1988 1 Ex. A6

A6-A8

M7-E8

M7-E8 alle Stücke

Gen.Prp.

E6-E8® alle Stücke

Gen.Prp

®E8 nur 2 Ex

E6-AB

E6-A7 1989 WaN Io,

Mallertshofer

Holz do

A7-M8 1988 1 Ex. E6

AB-M9

M7-M8

A7-M8

A8 Gen.Prp

M7-E9 viele GenPrp,,

noch keine

H. ultima

M9-AIO

M8 ®wt (RR)

M8-MI1O

E9-MIO

M6-A7 1987 1 Ex. M7

M6-AB 1987 auch M8

und EB

M6-M8

E6-E6,

E7-E9

M7-E7 * feuchtwar-

me Standorte

AB8-E8

A6-A8® °1989 ab M6

E7-AB

A6-M7 S-Bahnhof 1988

2 Ex., ®zT.

Zuflug aus den

Alpen ("wtf

A6-E7 ®1 zus. Ex.

Art

Cucullinae

289

290

291

292

293

294

296

297

298

299

300

301

302

303

304

306

307

308

309

310

312

313

314

315

Cucullia

lucifuga

SCHIFF. 1776

Cucullia

umbratica

L. 1768

Cucullia

verbascl

L. 1768

Cucullla L

scrophulariae

SCHIFF. 1776

Calophasia

lunula

HUFN. 1766

Brachionycha

sphinx

HUFN. 1766

Lithophane

socia

HUFN. 1766

Lithophane

ornitopus

HUFN. 1766

Lithophane

turcifera

HUFN. 1766

Xylena

vetusta

Hen. [1813]

Allophyes

oxyacanthae

L. 1768

Griposia

aprilina

L. 1768

Blepharita

satura

SCHIFF. 1776

Blepharita

adusta

EsP. 1790

Antitype

chi

L. 1768

Eupsilia

transversa

HUFN. 1766

Conistra

vaccinli

L. 1761

Conistra

rubiginosa

ScoPr. 1763

Agrochola

circellaris

HUFN. 1766

Agrochola

macilenta

HBn. [1809]

Agrochola

nitida

SCHIFF. 1776

Agrochola

helvola

L. 1768

Agrochola

litura

L. 1761

Agrochola

Iychnidis

CHIFF. 1776

Agrochola

lota

CL. 1769

Parastichtis

suspecta

Hen. [1817]

1814

Cirrhia

aurago

SCHIFF. 1776

SIEDLUNG

ı Garten |

= 96

1, =W5

= a

"sei

202

WALD

- 01

3 6

6 00

-n

{R)

VOR

1987

Gar Mo HO HM

Vlenzeise-

v{r) (R) (R) (R)

1 - - -

3 Su er

IV. = 3, =

(SV 2

1 = oe

2 - - -

vh = 0-0 -

v-h- 1 -

Geb

(mGr)

Geb

(Hy)

ed)

Tess)

FLUG- BEMER-

ZEIT KUNGEN

E6 am L.n. opt.

E6-AB am L. n. opt

am L. n. opt

®Ruderal und

nördl. Hügel

®®Birket-Rand

E6-A7 am L.n. opt.

im Schw holz

1987/88 eini-

ge R

M6,

E7-AB

A10-All 1983 im Bergl-

wald 2 R

E9-Win-

ter-E6

M3-A5 Überwinterer

M9-MIO Überwinterer

M3-M5 Überwinterer,

1989 I d am

Franzosenhölzl

M9-EIO

E9-AIO

EB-E9

M8-M9

E9-Win- *am nördl

ter-AB Hügel I R,

eos lluR

M9-Win-

ter-M5

E1O-Win- *alle 1986

ter-M4

E9-EN

M9-AI11

A9-M9

E9-EIO

E8-A1O

M9-EIO

E9-MIO ANGE

1983 I R

M7-AB

Nr. Art

316 Cirrhia

togata

ESP. 1788

317 Cirrhia

icteritia

HUFN. 1766

8 Cirrhia

ilvago

CHIFF. 1776

319 Cirhia

ocellaris

BKH. 1792

320 Cirrhia

citrago

L. 1768

Melicleptriinae

321 Chloridea

viriplaca

HUFN. 1766

322 Chloridea

eltigera

CHIFF. 1776

323 Pyrrhia

324 Panemeria

tenebrata

ScoP. 1763

3256 Axylia

utris

. 1761

Bryophilinag

326 Euthales

algae

F. 1776

327 Bryoleuca

raptricula

SCHIFF. 1776

Apatelinae

328 Panthea

coenobita

EsP. 1786

329 Daseochaeta

alpium

OSBECK 1778

330 Colocaslia

coryli

L. 1768

331 Subacronicta

megacephala

SCHIFF. 1776

332 Acronicta

aceris

L. 1768

333 Acronicta

leporina

L. 1768

334 Apatele

336 Apatele

. 1768

336 Hyboma

strigosa

SCHIFF. 1776

337 Pharetra

auricoma

SCHIFF. 1776

338 Pharetra

rumicis

L. 1768

339 Craniophora

ligustri

SCHIFF. 1776

SIEDLUNG

nes

2er

Se)

1 - -

am 2

E20

ı®

en) 2 a

[Ay

WALD

Dn-

-n

w—

3“

VOR

1987

Mo HO HM

3 - =

1 - -

- 1 -

(bye ur?

1 1 1

1 En -

5.9 =

. 9 -

Au =

= 2 >

2 - -

= 1 &

T®® m -

= 1 -

2. |

3 29

FLUG-

er) M9-MI1O

Geb/R EB8-A1O

(Hy)

Geb M9-E9

Geb M9

(Hy)

Geb E8-A1O

r/Wf EB-A7,

Geb/R, E7-AB

E8-M9

E6-AB

Teen) "°

en. EB-E7

Geb E7-&9

Xe E7-M8

Geb M6-E6

(wn)

Ms A6-E6

Geb)

WwL E4-M6,

M7-M8

H 6-M8B

(Geb) 2 Gene-

rationen)

Geb 6-M8

2 Gene-

rationen)

Geb E5-E7

Geb M6B-A7

Geb MO:

MB (2 Ex.)

Geb A6-E7

(wL)

Geb/R E4-M6,

(Agr) A7-M8

AR AB-M6,

eb/R) M7-E8

Geb einzeln:

(WL) AB-A6

2ER

BEMER-

ZEIT KUNGEN

auch T, 1981 in

Kiesgrube 500m

von WaN ent-

ternt IC T

T, 1989 3 Ex

in im Mal-

ertshofer Holz

°1i Ex. am

S-Bahnhof

“nur 1983

"FFwr.,

2. Genera-

tion 1986

bis 1988 n

SiN 1988 1 R

9891

Was)

als R verbrei-

tet, ® S-Bahn-

hof 1 zus. Ex.,

®®Birket-Rand

n Klesgruben

(Ber I-, Schw.-

olz) als Rh

VOR

Nr Art n . 1987 |

Gar Mo HO

Jaspidiinae

340 Jaspidia El = a lo0= 8 - | - - 1 = mGr EB-A7®® T häufiger, vo,

eceptoria - - mer 3 35 062007 it . = Geb) W un u

ScoP. 1763 Ruderal *®

“22.787 1 Ex

341 Jaspidia 3 716 47 14 286 99 3 2 12 h-shr 3872 un M6-AB8 auch T, zB

ygarga 22 127 361 417 494 102 2 10 4 8 Geb, Berglwald sh

FN. 1766 Ww)

342 Eustrotia = 0-0 => See Sr = ,2 EV, E7

uncula = = = = - - = - = =

CL. 1769

343 Eustrotia 1 = - - - 1 8 1 1 = 1 eg, mar A6-M7 T sh (M): HM

olivana - 1 - 1 3 9 -. - Sl Geb, Ruderal ®),

SCHIFF. 1776 Hy egattastr,

Schw.holz

Nycteolinaeg

344 Nycteola Sr= - 1 1 - - - - - 1 - = | WL MA-A6, Überwinterer

revayana - 3 - =. - - - =u = E7 (2 Ex)

ScOP. 1763

Beninae

345 Earias = = = = = 3 = = 1 1 = - = = Geb M6-E7

chlorana - - 3 a =. = - -.- (Hy)

L. 1761

346 Bena - - 1 3 1 6 8 - - - t-h - Be Ws M6-E7

rasinana 6 I 21 8 6 6 - 1 1 - Geb)

1768

347 Pseudoips = = = 1 = = = = = = 3 = om. = WL E6-E7 WaN 1989 2

bicolorana = = - = 5 = = = ZT 7 Ex

FUESSLY 1776|

Plusiinae

348 Chrysaspidia - - - = = _ = = = = = 1 EZ lEhy; E6

utnami - - - - - - - - =

ROTE 1873

349 Autographa 7 360131 81 21 74 69 70 127 2 sh 21 13 16 Up/Wt 6-A1l auch T, zB

gamma 30 65 28 6 23 17.12 47 1 1 Agr) 2 Gene- Entw.gräben,

L. 1768 rationen) BapaMBeL

Schw holz sh (M)

350 Autographa - 1 - - - - 1 - 1 - 1 mu = au A6-A8 .wi?

ulchrina = = = = = = = = - = Geb)

AW. 1809

361 Autographa - - - 1 1 - - - - - - - 0-0 - Geb*® M8-E8 “wtf (RR)

bractea - - = - - 0 => - - = (W, Hy)

SCHIFF. 1776

352 Macdunoughia - 0-3 1 - 2 - - 4 (R) Inh 2 - 14 Agr/Wtf M7-AlIO *auch hier

confusa - - - - - - = - = 4A (Xe (2 Gene- 1987 I R

STEPHENS mGr) rationen)

1860

363 Plusia 1 2 = 1 2 4 1 1 1 1 n-hı "Wi 22 Ub A6-E7 *1988 1 Ex.

chrysitis 3 12 3 7 3 6 4 7 - - . M8-M9® der 2. Gen.

L. 1768 schon AB

364 Plusia 1 = 1 3 - 1 1 - 8 -*!in 41 3 Ub E6-M7, ® Entw.graben

tutti - 13 - 2 2 6 - 2 - 1 AB-E8 1 Ex.

KOSTR. 1961

365 Plusia = = — = = = = = = = 1 = == H AB

chryson - - == = - 0 = - - = (Geb)

ESP. 1789

356 Polychrysia = = = = = E = = = = =) = = = Geb A7

moneta - - - - - - - - -.-

F. 1787

357 Chrysoptera - - = 1 - - = = = - 1 - - - Geb E6-E7

c-aureum 1 - - - - - - - 1 5 (wu)

KNOCH 1781

368 Abrostola - - = = — 2 1 = = — 3 Pe Geb 6-M8 Genprp.

triplasia = 4 _ = 2 1 5 1 1 - 2 Gene-

L. 1768 rationen)

369 Abrostola - - 1 = - 1 - - - = = =. er erlläGeb A6-E7 Gen.prp.

asclepiadis - 6 2 3 = - - - =. - (Xe)

SCHIFF. 1776

360 Abrostola - - - - 1 - 1 1 - - 2 - => Geb M6-E9 Gen.prp.

trigemina - 2 - = 4 2 - 1 - -

WERNEBG

1864

Catocalinae

361 Astiodes - - - = - - - - = - - = SZ WwL® E8 am L. n. opt

sponsa - 1 - - 0. - = - = “wi (RR)

L. 1767

362 Catocala - - 1 1 - - = 1 = - 1 Ge Ber Se Geb AB-E9I am L. n. opt

nupta - - - - - - - 1 - .- (wı)

L. 1767

363 Callistege - + - - — . . * . . . - - - - er er r M5-E6 T, auch

mi = + - - - + * + + + - . = - Agr, Regattastr.,

CL. 1769 eb) Schw.holz

m nn mn rn

VOR UOKO-

w An 1987 LO- Zei KUNGEN

Gar Mo HO HM | GIE

364 Ectypa . ® - = . . ° . « « . Q Ö > = ’ 0,0 Gr A5S-M7 T, auch

piyphica . + - - + . . + + ’ + * . = gen. E7 (1 Ex.) Be attastr,,

. 1768 Agr chw.holz

iderin

365 Scoliopteryx 1 = - 6 = 3 = - 1 1° tv 1 us Geb E6-E7 am Köder

libatrix - - 1 SE 1 - - > pc (Hy) Es-Win- besser,

L. 1768 ter-A6 ®zus. I R

e®zus. I P

astinum - - 1 2 2 - 1 Se Imer. M6

R. 1826 e

367 Parascotia - - -. 2 = - - - = = Vo Be = “ E6-A9 “S- Bann

fuliginaria - 3 - - - - = - Zu = Geb) 1987 1 Ex

L. 1761

368 Phytometra SE ul - (= S |< ge} Ds AB-A6, auch T, Mal-

viridaria = = = 1 - - - 1 = = W A8-M8 Ruelaıeı Holz

CL. 1768 Xeb) 1989 3 Ex,

® auch 1 MöBıe

369 Rivula - - 1 2 3 4 13 - 1 67 Au. =, Se Geb 6-M9 auc

sericealis 3 20 8 14 22 35 - 6 118 (mGr) (2 Gene- Däm

ScoP. 1763 rationen)

Hypeninae

370 Laspeyria 1 1 6 26 2° 23 3 - 1 - n-h 164 1 Geb M6-M8 *WaM 1987

tlexula 12 71 40 34 37 2 1 1 S26 (w) zus. IR

SCHIFF. 1776

371 Colobochyla - — = = _ = - Ze = = Eee EGeb/RI AG v.a. T,

sallcalis - - 1 Ss - See Zi = (Hy) ®im Ruderal

SCHIFF. 1776 mehrere Ex

372 Herminia - — 2 1 1 1 6 = = - - = = Geb E6S-M7

barbalis - 6 - - 3 4 - 1 Ser - (wıL)

CL. 1769

373 Zanclognatha == 3 6 = 3 1 = 1 = 3 Zee Geb M6-M8® geopip

tarsıpennalis 2 48 3a 324 - = - - > (mGr) “in 8 frische

TR. 1836 Ex. einer 2.

Generation

374 Zanclognatha 8 = 6 16 326 = 1 42 n-h 11 SZ Geb A6-E7® 1987 1 Ex

tarsicrinalis 1 14 1327032 14 - - 6 18 (wı) am 128.,

KNOCH 1782 n Birket 1986

375 Zanclognatha = = - 4 — - - - - = 2 en, > Geb A6-E7

risealis - 8 1 - 2 1 - - 1 2 (wı)

CHIFF. 1776

376 Trisateles - 1 < 6 2 2 2 - - 1 Vene - SE E5S-E7

emortualis 2 1 1 - - 1 - 1 1 2 Geb)

SCHIFF. 1776

377 Hypena 62 6 14 - 2 2 2 1 10°| n-h 24 1 3 fe Ei auch T: (U

roboscidalis 13 68 2 6 7 3 - - 26 16 Geb/R) (2 en. Biken h

. 1768 rationen) 2. G

378 Hypena = - - = = = = = = - - Ze ee Geb A6° 1989 1 d im

obsealis = = = = = = - = SE (Hy) Franzosen-

TR. 1829 hadlzl,

berwinterer

Geomatrkdae

Archiearinae

379 Archiearis. - = = = = = 2 Z = "7 - _ 70h Geb E3-E4 T, Birket und

arthenias - - = = = = - - SEAT (WL, Regattastr

.. 1761 Hy) stets h

Oenochrominag

380 Alsophila 2° 4 67 ASE SZ, SEEE19 = h-sh -" -e -° L A3-E4 [2207

aescularia 8 9 7 = p2 2 - 1 29 1 Geb) °F Ewır.

SCHIFF. 1776

381 Odezia - = = = = = = = (T)| - - =. - mon A6-M7 T: Birket und

L. 1768 WaM 1983

1 Ex.

Geometrinag

382 Geometra - = 3 1 3 - 2 = 1 8 -v 10 91 [L E6-M8

Raptloneria - 1 - - 1 - - - > 2 Geb,

. 1768 Hy

383 Comibaena - S - = = = = = = - - - 4 - | WL E6

ustulata = - = - = = = = - -

FN. 1767

384 Hemithea 1 Ss u 2 2 = = 2 vn - 4 - | Geb M6-M7 Raupenfunde im

aestivarla 1 1 1 3 6 1 - - 2 3 (wu) Ort, WaM, HO

Hen. [1799]

1796

386 Thalera - = > - > = 1 3 4 - - - -.- e A7-M8

fimbrialis - - So, 0.000 1 1 6 Ir = mGr)

ScoP. 1763

386 Hemistola - = = = - = = = = - - - oo Geb E7

chrysoprasaria - = 1 - 1 = - — - -

ESP. 1794

387 lodis ee. = = = 1 - = 2°| 2 - (T:3) - | Geb M65-A7 Däm, amL

lactearia 2 2 - 1 2 6 - - Sun (wı) n. opt.,

L. 1768 °Däm 6 zus.

Ex., 1983 5 Ex

WALD VOR

Mr. Art i 1987

Mo HO HM

Sterrhinag

388 Sterrha - - - - - - - = = = = = en -

serpentata - - - = = = 5 = = a

HUFN. 1767

389 Sterrha = = = = = 4 - - 2 = > Sa. u

muricata - = 1

HUFN. 1767

390 Sterrha - _ 6 12 - 3 = = - 5 vh = 6 -

biselata 4 16 13 9 8 6 = 1 2.58

HUFN. 1767

391 Sterrha - - - - - - - - - a = = ih 2

inquinata - - 1 - = = < = & =

ScOP. 1763

392 Sterrha = = = = - - - - 3 Be

seriata 1 - - - - - - _ = =

SCHRANK 1802

393 Sterrha - .- 2 4 - 3 - - - = = 1 Pe Dr:

dimidiata 1

HUFN. 1767

394 Sterrha = = - - - - - 1 - - 1 Fr

emarginata - - - - - - = = = =

L. 1768

396 Sterrha 1 341 22 8.2 I) 2 9 6 h IE I5) ==

BEReate 24 84 9 20 14 9 - 7 2 6

. 1768

396 Ercppkoca - - - - - - - - - 4 t-n 1 1 -

my punctata 1 2 4 1 - 3 = = = 6°

HUFN. 1767

397 Cyclophora - - 6 19 3 2 29 - 3 6 h-sh - 44 -

unctaria 40 | 220 20 19 16 66 - 9 7

1768

398 Cyclophora - - - - 1 - - - - - 22 - - +

linearia 2 = = = = 1 = E - -

Hen. [1799]

1796

399 Calothysanis - - 3 3 3 3 9 (n

griseata 4 19 10 16 8 10 = 2 4 1

PETERSEN 1902

400 Scopula = wa) - - - 1 3 ı0 - - -#. mM 83

immorata - - - 1 he 1 2 47 - -

L. 1768

401 Scopula = = = 4 Sse1D 6 = = 8 3° = 87

nigropunctata 1 1 Skler ir! 3 - 1 - 3

HUFN. 1767

402 Scopula - - - - - - - 6 2 - 1 a1 -

ornata - - 1 - - _ 2 3

ScoP. 1763

403 Scopula u > - - - - - - 6 = - =4 14 2

rubiginata - - 2 — 1 1 1 g - -

HUFN. 1767

404 Scopula = = — - - - - - - = 1 Se

immutata - - - - - - - - - =

L. 1768

405 Scopula = = 1 = = - - - - > 1 — (im =

lactata - - 2 2 = = - 2 = =

HAw. 1809

Larentiinag

406 Scotopteryx 1 -

chenopodiata

L. 1768

407 Anaitis - - - - = 1 = = = = = = =I2-

raeformata 1 - - - - =: en = = =

Bn. [1826]

408 Anaitis 3157210 10 =

efformata 6 39

GN. 1867

409 Acasis - - - - 1 - - - = = 2) u ae

viretata - - - - 1 - - Ss = —

Hen. [1799]

1796

12 143 13 17 37, it SE E6

119 218 26 = 150 2 =

{=}

n=hı = 79281

-a

410 Nothopteryx Zu „= 1 = = = 1 = 1 = vn -+ -ı 0

olycommata - 2 1 - - - - = en =

CHIFF. 1776

411 Nothopteryx = - - - - 1 1 - - 1 = El u

carpinata - - - 1 1 3 - W 4%

BKH. 1794

412 Lobophora - - - - - = = = = = = 2.30

helterata 1 - - - - - - = = 1

HUFN. 1767

413 Pterapheraptery 1 4 1 - = 8 1

sexalata - = 7

RETZ. 1783

414 Operophthera {R) - 93 3 2 - 1 ı ({R -| h-sh -" 6 -

Duansia 66 106 8 I, - 18 = 3 22 8

. 1768

A7-M8

Mo Ex)

1 E5)

E6-A7,

AB-A9I

M6-E6,

AB-E8

E6-E6

M7-A9

AB8-EB

M6-A7,

E7-AIO

AS-A6

E3-M4

M4-M6

AB-M6

M5-AB

EIO-A12

T 1989 im

Mallertshofer

Holz h

®auch Moore

Heiden, WaN

1978 +1986 2 E

Entw.graben

1987

®v.a. an Ge-

bäuden

va. an Ge-

bäuden, am L

nicht opt. 198

S-Bahnhof I E

Entw.graben

1987 4 Ex,

5- el 198

x“.

2. Generation

weniger zahl-

reich

Däm, Birket

1983 2 Ex,

®°Dam h

3. Generation |

nur sehr par-

tiell

1989 WaN I 9

auch T, Birker

1981/83 3 Ex

un raben

87 v, HM '86

WNo T 2 Ex, |

2. Gen. stärke

®im bewaldet

Teil, Schw. hol/f

1987 TI Ex, |

auch T- HW

1987 1 Ex

auch T,

2. Generation

störker

auch T

®Birket-Rend |

8 Ex, auch T:|

WaN, HO + HM]

Ruderal h-sh |

Genpr

auch Piv)

S-Bahnho

1988 1 Ex

1989 Dere

en SN I

°FFwer.

“FFwer.

®EFwer.

dd, ®FFwı

Nr. Art

Oporinia

dilutata

SCHIFF. 1776

416 Oporinia

autumnata

BKH. 1794

417 Triphosa

dubitata

L. 1768

418 Calocalpe

cervinalis

ScoP. 1763

419 Philereme

vetulata

SCHIFF. 1776

420 Philereme

transvorsata

HUFN. 1767

421 Lygris

runata

. 1768

422 Lygris

testata

L. 1761

423 Lygris

Bopulate

. 1768

424 Lygris

mellinata

F. 1787

425 Lygris

Pryabele

CHIFF. 1776

426 Cidaria

fulvata

FORSTER 177

427 Plemyria

rubiginata

SCHIFF. 1776

428 Thera

variata

SCHIFF. 1776

429 Thera

obeliscata

HBn. 1787

430 Thera

[nipersie

. 1768

431 Thera

firmata

Hen. [1822]

432 Chloroclysta

siterata

HUFN. 1767

433 Dystroma

truncata

HUFN. 1767

434 Dystroma

citrata

L. 1761

435 Xanthorhoe

tluctuata

L. 1768

436 Xanthorhoe

montanata

SCHIFF. 1776

437 Xanthorhoe

spadicearia

SCHIFF. 1776

438 Xanthorhoe

terrugata

CL. 1769

439 Xanthorhoe

biriviata

BKH. 1794

440 Xanthorhoe

designata

HUFN. 1767

441 Ochyria

uadrifasciata

L. 1769

(T:1)

97,87

201

-n

7°

21°

Gar

1-v

n-h

VOR

1987

Mo HO HM

1°

UKO-

LO-

GIE

Med)

wı

Geb]

Geb®®

Geb

(wı)

Geb

(Ub)

Geb®

Geb)

Gev)

Geb*®

Geb

(mGr)

Geb

Geo,

Web)

Weed)

Geb

(wN)

ev)

Geb

(wı)

Geb

(w. Hy)

May)

Geb

Ub)

(Geb)

Me b,

May)

God)

FLUG- BEMER-

ZEIT KUNGEN

AIO-Ali Genprp.

A1IO-MII Gen.prp.

M7-Win- ®am Sender-

ter-E6 gebäude,

“va. an Ge-

bäuden

E3-A6 °EF.we.

E6-E7 “Birket 1987 T

Ex., Garten

erst ab 1986

E6-MB8 ®v.a. an Ge-

bäuden

M6-A9 ®v.a. Gärten

1988 3 Ex. im

Korbinianiholz

KOLBECK

Was 1989 1

M6-A7 ®v.a. Gärten

E6-M8

M6-E7

E6-E7®® ®T ein zus.

Ex., °°1987

1 Ex. M8

M6-A7, Gen.prp., keine

MB8-E9, T. britannica

M10 (2 Ex.)

E6-E7, ® regelmäßig

E7-AB® frische Ex

Jedoch 1. Gen

stärker

E9-EIO S-Bahnhof

1987 6 Ex

M8°-MIO *AB einzeln

M8-Win-

ter-M6

E6-M7 ®Birket 1983

8-EIO 1 Ex, M-EIO

A8 2 Ex.) z.T. frische Ex,,

Gen.Pr

AB-E9I “vor 1987 4

fotogr. Belege,

sonst Überse-

hen, Gen.Prp

AB-E6,

M7-E9,

MIO (1 Ex.)

A6-M7 T, *am L.n.

np! Birket

A 87 T) 600

Ab-EG6, GenPrp., auch

A7-E8 T: Birket-Rand

1987 3 Ex., 1988

1 Ex.

A6B-M6° °1987 bis E6,

A7-M9®® *®in 9 z.T. frisch

Genprp., auch

T: Birket-Rand n

M4-ABb, ®zus. IR,

E6-E7 “oT 2 zus. Ex

E6, SiN E6 1989 1

M/-M8 frisches 9

M6-E8® °1987 1 Ex. A9,

Entw.graben

1987 1 Ex

VOR

SEREE 1987 FLUG- BEMER-

ZEIT KUNGEN

Gar Mo HO HM

442 Nycterosea Ze 1 = = = = = - - - - - - | Ub/Wt Ai

obstipata - en E = = n - - -

F. 1794

443 Calostigia _ = 1 1 1 1 _ - - - 2 = Ic w M7-AB

olivata 1 8 = 1 = = - - — Se

SCHIFF. 1776

444 Calostigia = 2 77 N 2 - 4 - 3 > 5 Geb A6-E7, ® 1 Ex. 1989

ectinataria 2 22 6 7, 2 7 - > (mGr) E8®

NOCH 1781

445 Lampropteryx 1 - = 3 = 1 2 - 4 - f-V = -' = are M5-A7, Birket 1983

ocellata 1 21 1 4 16 30 = 7) 1 1 Geb) E7-A9 1 Ex.

L. 1768

446 Lampropteryx - - = - = 2 1 - - = EV == Geb A6B-M6

suffumata 1 1 2 1 4 2 = - - > (w)

SCHIFF. 1776

447 Coenotephria = - 6 18 4 7 7 1 7 2 h = 3 5 Geb AS-A7,

berberata 13 119 6 1 6 12 - 2 1 - (w) M7-A9

SCHIFF. 1776

448 Euphyia | 2 7 3 4 6 2 1 r in -- =. - | Geb A6B-AB

cuculata 4 6 10° 107 79 Pe] 4 =4 er (mGr,

HUFN. 1767 Ww

449 Euphyia - - - 1 - - - - - - 1 = =] = Geb E6-E7 WaN 1989

molluginata - 2 - 1 - - - - - > (mGr) 1 Ex

Hen. [1813]

1796

450 Euphyia (IT) 1 8 7 4 3 og 16 - n-h - 20 1 er A6-E9I auch T: HO +

bilineata - 48 a. 1 - VE) 2 - ee (1-2 Ge- HM-Ruderal

L. 1768 mGr neratio- stets h

nen

451 Diactinia - - - - - _ - = - - - = 2-7 =,ldH E6-M7 We 1989 1 Ex

capitata - 1 = = = > = - - 2 (Geb,

H.-S. 1839 W)

452 Diactinia - - - - 2 = z = = 202173 = 1 - M AB-A6, auch T/Däm:

silaceata 2 12 - 2 1 = = = 2 = Hy) A7-M8 ®Birket 2 Ex

SCHIFF. 1776

4563 Electrophaes = = = - = = - - = 2 th »-# 31 € L E65-A7

corylata 1 4 Be 1 | = = Geb)

THNBG. 1792

454 Mesoleuca = Def =# (MI = u 13 av ZI u M6-E8 auch T/Däm:

SRicHRte - 6 - - - = - - 8 Hy) Be % N

. 1768 stets h,

Schw.holz 1 Ex

455 Melanthia - - - - - - - - = = 2 -4 =] | Geb A6-A7,

rocellata - 3 - 1 1 - => - - .- (Xe) E7-M8

CHIFF. 1776

456 Epirrhoe 4 W02 - 1 - 711 3 - 18 1 I-V SE BI E mer A6-M7, auch T, va

tristata 2 6 86 28 33 ar 27760 I (hi Geb) M7-E8 WaN und HW,

L. 1768 un graben

457 Epirrhoe 456 1 12 2.426 10 1 31 14 h si 37% U 4-A9® auch Tü [)

ALERT 27 | 196 116 656 109 31 4 69 20 28 (T:6) (Geb) 2-3 h —_

LLER, O.F Genera- *In 9 zT. fri-

e 1764 tionen sche Ex

4568 Perizoma 1 9 8 6 1 9 12 - 21 9 h sh 2 1 U M6-EB

alchemillata 26 61 n 7 6 12 #38 621 [Geb)

L. 1768

469 Perizoma - - - - - - = = > = = = te e M8

bifaciata - 1 - = - = = = = = "Ins ,

HAw. 1809 eb

460 Perizoma = = - - - - - - - = 2 = =) = H A6-E6

blandiata - - - - = - = 4 = - (Geb)

SCHIFF. 1776

461 Perizoma - - - - - 1 - - - 1 - - - | Geb A6-M7

flavofasciata = - - - 2 - - - = = (mGr)

THNBG. 1792

462 Hydriomena 2 = 3 5 „ 2 - 1 6 1 sh 2 - | Geb E6-A8

furcata 1 3 Se ! 2 2 3 16 (w,

THN8G. 1784 Hy)

463 Hydriomena - - - 3 - 3 n = 2 35 n -- 4: - | Geb A5-M7 Birket 1983

coerulata 1 3 13 a7 6:5. = - - 3 (Hy) 3 Ex.

641777

464 Anticlea - - = - = 1 - = = - = - - .- Geb E3-E4A 1989 WaS Ic

badiata - - 1 3 1 - - - - -

SCHIFF. 1776

465 Pelurga - Sle = = Ver VUEZT -0e=T ES Gr E7-E8

comitata - 1 = = _ 1 4 2 = = Agr,

L. 1768 ar,

466 Hydrelia - - - 1 - 1 - - - 63 - af <7=%H E6-AB WaN 1989

testaceata - - - | - = - 3 10 (Geb) 1 Ex., im Fran-

DENOYAN zosenhölzl sh

467 Hydrelia - 1 1 2 6 3 - - 16 - =1 31 > L M6-AB8

flammeolaria 2 31 3 7 3 8 - 1 2 3 Geb,

HUFN. 1767 Hy

468 Euchoeca - - 2 2 - 9 10 - 1 30 - =3537 € L M6-A7, 1. Gen. 1987

nebulata 1 3 RE Er 21 - - 6 18 Hy. A7-E8 bis M7,

Scor. 1763 eb) 2. Gen. zahl-

reicher

469

470

471

472

473

474

475

476

477

478

479

480

481

482

483

484

485

486

487

488

489

490

491

492

493

494

495

496

Asthena

albulata

HUFN. 1767

Asthena

anseraria

H.-S. 1866

Eupithecia

tenuiata

Hen. [1813]

1796

Eupithecia

inturbata

Hen. [1817]

1814

Eupithecia

Iumbeolata

AW. 1809

Eupithecia

ini

ETZ. 1783

Eupithecia

bilunulata

ZETT. [1839]

Eupithecia

linarlata

SCHIFF. 1776

Eupithecia

exiguata

Hen. [1813]

1796

Eupithecia

valerianata

Han. [1813]

1796

Eupithecia

venosata

F. 1787

Eupithecia

egenaria

H.-S. 1848

Eupithecia

extraversaria

H.-S. 1862

Eupithecia

centauresata

SCHIFF. 1776

Eupithecia

selinata

H.-S. 1861

Eupithecia

trisignaria

H.-S. 1848

Eupithecia

intricata

ZETT. [1839]

Eupithecia

satyrata

Hen. [1813]

1796

Eupithecia

tripunctaria

H.-S. 1862

Eupithecia

absinthiata

CL. 1769

Eupithecia

assimilata

DOUBLEDAY

1866

Eupithecia

vulgata

HAw. 1809

entfällt

Eupithecla

castigata

Hen. [1813]

1796

Eupithecia

icterata

VILL. 1789

Eupithecia

succenturlata

L. 1768

Eupithecia

subumbrata

SCHIFF. 1776

Eupithecia

millefoliata

ROÖSSLER 1866

SIEDLUNG

ı Garten

SiS SIMı SiN

-n

113

WALD

N Wasserwerk

WaS! WaM WaN WNw WNo

KENRASEN

HO! HM HW

-n

@anS

5°

Gar

3®

Ev)

Geb

(w)

Geb/R

Geb

(mGr)

mGr

Geb/R)

Geb)"

FLUG-

BEMER-

ZEIT KUNGEN

ES-A7

A6-A7

E7-E8

A7-AB

ES-M6

AB-M6

A7-E8

AS5-M6

E6-M7

M6-E7

M6-M7

A7-E7

M5-A6,

E7-E8B

E6-E7

AB-M6,

AT7-E7

AB-E6

E6-E6

M6-A6,

M7-E8

A8-E8®

MB-AB6,

E7-E8

M5-A7

AB-E7

M7-E8

A7-E8

E6S-M7

A7-AB

im Garten

WaS) auch

989

Gen.prp.

Genprp.

Gen.prp

Gen.pr

Wa IR)

1 Ex.

“alle 1983,

Nachzucht,

WasS 1989 1

Gen.prp.

Gen.prp

Gen.prp.

Gen.prp

Gen.pr

® zus. ? 1 Ex

®® nur 1986

Gen.pr

® zus R

®® nur 1986

Gen.prp.

® nur 1d86,

2. Generation

aurker

en.prp.

°1987 1 Ex. E6

Gen.pr

eva. Gärten,

2. Generation

stärker

Genprp.

Gen.prp

°T 2 weitere

Ex.

Gen.prp

Gen.prp.

Gen.prp

Gen.prp.

SIEDLUNG VOR FLUG- BEMER-

N. Art | Garten ! 1987 ZEIT KUNGEN

SiS SM, SIN Gar Mo HO HM

497 Eupithecia = = - = = = = = = = n/ M6-E7

Talnuosarla = 4 - = = = = = - = Ru

Ev. 1848

498 Eupithecie 1 2 1 - = - = [ze Hz 9 WN AB-E6 Genprp

indigata 1 6 = = 1 = — = = = FF.wer.

Hen. [1813] Birket 1983

1796 1 Ex.

498 a siehe Addenda

499 Eupithecia - - - 1 - 1 = 3° a 2 r AB-EB Gen.prp.

roten - 1 - - - - - 1 = = ee) “nur Bas

HUFN. 1767

500 Eupithecia 2 2 - 1 - 2 3 2 = E4-A6 Gen.prp

"irgaureate 7 | 28 Das er | (Gen. E7-AQ"

DOUBLEDAY Ww

1861

601 Eupithecia - = - = = = = = a Io WL 1989 1 Ex. im

abbreviata = = _ = = = = = = = Korbinianiholz

STEPH. 1831 KOLBECK

602 Eupithecia = = = = = = 5 = = =, WL A6B-M6 1989 an SN »

dodoneata = = — = = = = TEE WaS 3 09

GN. 1867 Genprp.

503 Eupithecia - - - - - - Z i-h -_-.I- Geb E7-E8B Gen.prp.

sobrinata = = = = = = = 1 = =

Hen. [1817]

604 Eupithecla - 1 - 2 = = = = => =. = WN A6-M7 Gen.prp.

lariciata = 1 - = = = = = = =

FRR. 1842

505 Eupithecia 6 ul 3 3 - 1 1 Veh m 2. ui A5-A7 Gen.prp

tantillaria 6 19 1 6 052 a = = Geb)

Bsp. 1840

606 Eupithecia 2 1 - - - - = tn a 51 ur M4-M6 en.prp

lanceata 2 6 4 2 4 - - - Ze Geb) °FFwıI

Hen. [1826]

1816

607 Chloroclystis 6 U 6 14 1 2 8 h > u = U E4-M6 Gen prp

Tal 12 32 10 14 12 7 - 4 4 4 R& b, E6-MB'

HAW. 1809 Hy

508 Calliclystis 1 1 - - - - = = =. == Geb M6-AB Genprp, im |

chioerata - - I ap = Garten (Was) |

MABILLE 1870 u wieder

509 Calliclystis 17 7 - 2 - 6 4 h = 82 U M6-E7® Genprp

ar 17 29 2 - 6 4 - 9 > 1 Ioer. °1983:

L. 1768 Agr E6-E6

510 Horisme 1 - 1 - = = = = u Geb E6-E7

tersata 1 8 1 1 2) - - - == (w)

SCHIFF. 1776

Boarminnae

511 Arichanna - - - = - = Re E = Er

melanaria - - - 2 = = = = u = Hy A7

L. 1768

612 Abraxas - - - - - - - - = - = Geb® M7 “va. Gärten,

rossulariata - = - = E = = = = 1 We 1989 1

. 1768

613 es = A ae 2 Zur sr „ai2® - - 3 - | Geb M6-EB aut T (zahl-]

SCoP. 1763 1 (wı) reich

614 Lomaspilis 6 2 48 g SeEEB 3*| n 146 20 18 U A5B-MB®® ® h T: Birk

Damen 4 77, 6I 31 44 6 SI ln 16 71 [6 b, abe #

B Hy HM-Ruderal v, |

616 Ligdia 2008 3 1 1 4 hıı A A) VE E6

adustata 3|ı a 6, 6 Br 2 le 2 er |

SCHIFF. 1776

BIiombanıa = 1 = Se: ee| vn - 1 - L AB-A7 *Däm: Birket

bimaculata - 9 - 2 6 2 = = ı u Geb, 1981/87 je 1 Ex

F. 1776 Hy |

617 Bapta 7 4 10 4 - 3 9 n=h) = 97 = L AB-M7, *T ein weitere

temerata 8| 2 15° 26 12 7 1 .2|14 24 Ub, MB (IEx) Ex.

SCHIFF. 1776 Geb)

518 Cabera 9|ı7 27 Bi, 2 3|nh - 1 -| Geb 6-E8 _Genprp, auch

re 37 17.17 36 6 — 1 6 16 (wı) am Flug- T: nd (986/87 |

i zeitende Birket je 1 Ex |

2.T. frisch) s |

519 Cabera 4 6 37 1 3 = 49 -n 9 71 ur 6-E8 Genprp., auch |

exanthemata 3 16 26 21 42 10 6 3 130 Geb) (2 Gene- T/Däm:. v.a. im’)

ScoP. 1763 rationen) Dach. Moos h

620 Plagodis 1 1 1 Des e ER 4 a D |

abraria 3 5 eo 1 = = De v-h 10 Geb M6-E6 ed, ellorai

L. 1767 |

621 Ellopia - - - - - 1 - f-v _ - 1 - “a M6-M7 sensu FORSTER

asciaria - - - - - - - _ - .- Geb) & WCOHLFAHR

L. 1768 i

622 Elopa - = = == = ZelEvenS Saee on A6-A7 _sensu FORSTER)

rasinaria - 1 - :) = 2 a = - = Geb) & WOHLFAHRTI|

CHIFF. 1776 |

623 Campaea - - 1 - = = - vn - = [ke A6-E7,

margaritata 1 16 - - 3 1 - 2 1 - Geb) A9-M9

L. 1767

624

626

627

628

629

630

531

632

633

634

636

637

538

639

640

641

642

543

644

545

646

647

648

649

660

Ennomos

autumnaria

WRNBG. 1869

Deuteronomos

alniaria

L. 1768

Deuteronomos

fuscantaria

HAw. 1809

Deuteronomos

erosaria

SCHIFF. 1776

Selenia

bilunaria

ESP. 1796

Selenia

tetralunaria

HUFN. 1767

Apeira

syringaria

L. 1768

Gonodontis

bidentata

CL. 1769

Colotois

ennaria

. 1761

Crocallis

elinguaria

L. 1768

Angerona

runarla

. 1768

Ourapteryx

sambucaria

L. 1768

Opisthograptis

luteolata

L. 1768

Epione

repandaria

HUFN. 1767

Cepphis

advenaria

HBnN. 1790

Lozogramma

chlorosata

ScoPr. 1763

Macaria

notata

L. 1768

Macaria

alternaria

HBn. [1809]

1796

Macaria

signaria

HBn. [1809]

1796

Macaria

liturata

CL. 1769

Chiasmia

clathrata

L. 1768

Itame

wauaria

L. 1768

Itame

fulvarla

VILL. 1789

Theria

rupicaprarla

SCHIFF. 1776

Erannis

bajaria

SCHIFF. 1776

Erannis R

leucophaearia

SCHIFF. 1776

Erannis

aurantiarla

HBn. [1799]

1796

ı®

ı Garten

SIS SIM| SIN |WaS| WaM WaN WNw WNo

aan

-n

[Ay

]

boenı

6 1

1 =

8 32

1 7

SasR2

1 7

1 4

10 34

1 s

1 1

1 3

16°

UOKO-

LO- FLUG- BEMER-

ZEIT KUNG

GIE an

Geb E8-EIO

(wı)

Geb M8-A9

(wı)

Geb E7-M9

ws M7-M9

Geb)

1. MA-MS5,

Geb) A7-A8

wL M4-E6,

Geb) M7-M8

Geb A7-M8

(wı)

Geb AB-M6

(wı)

Geb E9-A11

(wı)

Geb A7-M8 ®zus. je I R

(wı)

Geb E6-E7 WaS 19891

(wı)

vs E6-E7

Geb)

Geb Ab-E6°® 21987 1 Ex. M7,

(wıL) auch Däm

Geb/R 6-E9

(Hy) 2 Gene-

rationen)

E6-E6 ®Birket 1983

“ M6

Hy)

Geb)

WwL A6-AB,

(Geb/R) AB-M8

Me A6-AB

Geb)

AS5-E7

AB8-M8

Sin M6-E7,

Geb) E7-M8

ac: AB6-E6,

ep: A7-E8

Agr

Geb*® E6-E7

Geb/R E6-M7

(Hy)

Geb E2-E3

Geb M10-M11

(Xe)

Ws A3-A4

Geb)

M10-M11

Veen)

2 Ex, auch T:

Birket stets h

2. Generation

nur partiell

2. Generation

nur partiell

auch T: Birket n

“alsRv

2. Generation

m.o.w. partiell

DICH am

aN|w, oJ, HO

HM-Ruderal, HW '

Birket 1983 1 Ex

®v.a. in Gärten

1989 WaS und

WaN je 1 Ex

Is, *FF. wer,

im Garten nur

1983 + 1989

II, ®F.F.w.r.

dd, ®F.F.w.r.

WALD

FLUG- BEMER-

Nr. Art ı Garten | Wasserwerk

H ) ZEIT KUNGEN

561 Erannis A3-M4®® dd, ®FFwr,

marginaria ®*1986 1 Ex

E17 4

6562 Erannis EIO-EN dd, ®FFwr

defoliaria

CL. 1769

6563 Phigalia AI-A4 dd, ®FFwr

edarla

. 1787

6564 Apocheima

hispidiaria

SCHIFF. 1776

E3 dd, ®F.Fwır

555 Lycia A3-MB *FFwı

hirtaria Gen.prp

CL. 1769

556 Biston - 2 - - - - - 1 1 - a Zi] Geb M3-E4 °EFwı

strataria 1 - - 2 - - - - - .- (wı)

HUFN. 1767

557 Biston 1 - 2 2 - 6 9 2 12 16 v eg #2 WE M6-E7 ® Entw.graben

betularia 4 3 19 6 - 8 == 112 34 Geb) 1983 8 Ex

L. 1768

558 Peribatodes 1 20 38 43 3 6 3 - 7 = h = F877= 7 AT7-A9

rhomboldaria 31 | 374 26 9 16 22 - 6 3 Geb)

SCHIFF. 1776

659 Peribatodes - = 1 8 1 12 2 — 3 1 n-h - 10 - 2 M7-E8B SIN + WaS

secundarla 4 73 40 69 12 n 1 9 1 - Geb) öfters als R

EsP. 1797

560 Cleora - - = = = 1 1 = = - - - 1 1 Gr A5B-M6 Mallertshofer

cinctaria - 1 - - - - - - - => Geb) Holz 1989 h,

SCHIFF. 1776 WaN 1989 2 «a

661 Deileptenia - - 4 18 14 7 16 2 1 8 3a Zu d- Y E6-MB®* "nur 1986,

ribeata 3 128 VERRLSIE iz 21 1 - 21 8 Geb) **1987 bis EB

CL. 1769 j

5662 Alcis 7 Aw 36 19 16 4 29 = 3 9 h-sh - 39 - " A6-AB, *3 Ex. 1988

repandata 18 | 728 72 169 132 69 1 4 n 9 Geb) M9-MIO®

L. 1768

663 Boarmia - = 2 = = 4 = = 21 1 In 7 E- WL M6-E7

roboraria 6 - - = 3 E 1 #12

SCHIFF. 1776

5664 Serraca - 1 4 15 1 2 31 1 4 16°| vn - 7 - Ss A5B-M7, *Birket 1983

unctinalis 6 39 8 3 10 26 - 8 566 21 Geb) E7*®® 3 Ex, **1988

COP. 1763 1 frisches Ex

565 Ectropis a 622137210, 9 - 1 16 h-sh - 8 2|U M3-E5°, S-Bahnhof 1988

bistortata 13 115 Br szel 14 - 2 13 9 W, E6-MB, 2.Ex., °2.T. bis

GOEZE 1781 eb) M9-MIO®® a sehr

partie

566 Ectropis - - - 2 - 1 - - - - (SE m: E6-A7

extersaria - 1 - = - — = = Pr Geb)

Hen. [1799]

1796

667 Aethalura — = = - 1 2 - = = 8 3 - 1 — LE M4-E6

unctulata - - 1 - 2 1 = = - 6 ir.

CHIFF. 1776 eb)

668 Ematurga N - - = (Mn 2 (n) 3 | - = 1m) Gr 6-M8 *T h-sh ("HM’>

atomaria - - De 3 1 6 a "ke 2 Gene- Ruderal),

L. 1768 eb) rationen) am L.n. opt

669 Bupalus - = 7 3 2 8 14 = 2 = n-h + = 26. = "s A6b-A7® ®1987 I Ex. M7,

iniaria 1 1 3 237 E19 icu == 2 - = Geb) Schw.holz 1988

1768 1Ex. T

570 Sons = EM = - - - \ I I - - - - 1 | mGr ES-A7 ee

neata = = = = = = nuderre=

ScoPr. 1763 BannStacke h

am L. n. op

Addenda:

268b Photedes - - - - - - - - - - - =. Se FSlcHy AB-A9 Wan 1989 2 dc

ygmina = = = = = = = = = =

AW. 1809

498b Eupithecia - = = = - - - - - - - - - - Geb A8 1989 im Mal-

impinellata = - - - - - - - = (Xe lertshofer Holz

BN mGr) 4 Ex

Artenzahlen: Gesamt: Tagfalter 68 Arten

Bombyces + Sphinges 96 Arten

Noctuidae 226 Arten

Geometridae 193 Arten

Summe 672 Arten

SIEDLUNG WALD HALBTROK- "DACH. VOR

Jahr ı Garten | Wasserwerk KENRASEN MOOS” 1987 2

SiS SIM} SN WaS!WaM WaN WNw WNo HO} HM HW Au We Mb Gar Mo HO HM

1987 116 136 217 240 171 260 267 160 266 181 431

1988 222 304 277 263 283 261 132 260 183 184 446

367 144 270 162 > 2 614

Die Artenzahlen spiegeln konstante, vom Jahr weitgehend unabhängige Muster wieder,

wie dies aus dem Beispiel des Flughafengebiets (HO, HM, HW) schön zu ersehen ist.

In gewissen Grenzen gilt das auch für die Individuenausbeuten, das Verhältnis HO/

HM/HW zeigt beispielsweise recht konstante Werte von 3,3/1/2,9.

4.4. FAUNISTISCHE HINWEISE

Auf eine eingehende Erläuterung der faunistischen Besonderheiten, so interessant dies

auch wäre, soll im Rahmen dieser Arbeit verzichtet werden. Da dies teilweise bereits

an anderer Stelle geschah, sei hier nur auf HAUSMANN (1988) verwiesen.

Einige für die Thematik relevante Informationen aus der Faunistik werden jedoch im

folgenden kurz aufgelistet. Es handelt sich hierbei um Arten, bei denen Arealerweite-

rungen festzustellen sind, solche mit Arealverlusten und schließlich Arten mit stark

lokaler Verbreitung. Die Angaben beziehen sich auf Prozesse bzw. Befunde dieses

Jahrhunderts in Südbayern.

Als Basis wird die von OSTHELDER (1925-1933) vor ca. 60 Jahren publizierte relativ

vollständige Fauna Südbayerns und der angrenzenden nördlichen Kalkalpen verwendet.

Die Hauptreferenz stellen die in der Folgezeit durch WOLFSBERGER (1945-1949;

1950: 1953/1954; 1954/1955; 1958; 1960; 1974 u.a.) veröffentlichten Ergänzungen dar.

4.4.1. Arealerweiterungen

Bei den Arealerweiterungen wie in den unten aufgeführten Fällen sind Ortsveränderun-

gen von Schmetterlingen (99) zu postulieren, die entweder in Vorstößen über viele Ki-

lometer hinweg (mindestens 10-50) oder in einer mehr oder weniger stetigen Weise

über mittlere Distanzen von 3-10 Kilometer erfolgten.

Derartige Ortsveränderungen entsprechen jedoch (meist) nicht dem "trivial movement"

(sensu SOUTHWOOD, 1978) der betreffenden Art. Vielmehr kommt es oft im Zuge

solcher Arealerweiterungen zu erfolgreichen Kolonisationen von geeigneten Habitaten,

wo die Art dann vergleichsweise orts- und habitattreu bleibt.

Folgende Arten sind zu dieser Gruppe zu rechnen:

Bombyces und Sphinges: Gluphisia crenata

Noctuidae: Euxoa tritici, E. aquilina, Noctua janthina, N. comes, Apamea scolopacina,

Hoplodrina ambigua, Eremodrina gilva, Calophasia lunula, Euthales algae, Bryoleuca

raptricula

Geometridae: Lygris mellinata, Thera juniperata, Eupithecia intricata, Eupithecia mille-

foliata, Eupithecia sinuosaria

Die Liste erhebt keinen Anspruch auf Vollständigkeit, bespielsweise könnten auch

Agrotis venustula, Zanclognatha tarsipennalis und Eupithecia virgaureata hierher gehö-

ren, u.U. wurden sie jedoch in vergangenen Zeiten auch nur übersehen bzw. verwech-

selt.

Wanderfalter, die neuerdings in Südbayern häufiger nachgewiesen werden, wie z.B.

Mythimna albipuncta interessieren uns in bezug auf die Thematik hier weniger.

4.4.2. Arealverluste

Arealverluste sind schwieriger zu belegen als Arealgewinne.

Wenn man den Blick jedoch nicht auf so große Areale, wie dies Südbayern ist, richtet,

sondern Arealverluste auf lokalerem Niveau untersucht, werden sich - vor allem kor-

reliert mit Zerstörungen von Lebensräumen - viele Beispiele finden lassen.

In einer vorbereitenden Arbeit konnte in diesem Sinne bei den hygrophilen Arten der

Noctuiden-Unterfamilie Amphipyrinae ein überdurchschnittlicher Rückgang in den Ar-

ten- und Individuenzahlen belegt werden. So sind beispielsweise die auf Schilf ange-

wiesenen Eulen Rhizedra lutosa und Mythimna pudorina im Untersuchungsgebiet nur

vereinzelt zu finden, während sie vor 60 Jahren von OSTHELDER (1925-1933) aus

Schleißheim noch als "sehr häufig” gemeldet wurden. Andere "Röhricht-Eulen” wie

Mythimna straminea TR., Leucania obsoleta HBn., Archanara geminipuncta HAw. unc

Archanara algae EsP. sind im Verlauf der letzten 100 Jahre aus dem Gebiet Ober-

schleißheim wohl ganz verschwunden. Diese Befunde sind mit hoher Wahrscheinlichkeit

auf die Zerstörung der Niedermoorflächen im Dachauer Moos zurückzuführen.

Eine parallele Entwicklung bei den Tagfaltern, nämlich die Extinktion von Populationen

vieler hygrophiler Arten v.a. im Dachauer Moos wurde in HAUSMANN (1988) belegt.

4.4.3. Arten mit lokalem Vorkommen

Das Verbreitungsmuster lokal verbreiteter Arten kann im Zusammenhang mit Speziali-

sierungen auf bestimmte Raupenfutterpflanzen wertvolle Informationen zur Beurteilung

von Verbreitungsstrategien liefern.

Auch die folgende Liste ist sicher nicht vollständig, sie spiegelt einige aus den Publi-

kationen OSTHELDERs (l.c.) und WOLFSBERGERs (l.c.) entnommene Verhältnisse

wieder, die für Südbayern bzw. im besonderen für die untere Hochebene des Faunenge-

bietes zutreffen. Eine Reihe von Zusatzinformationen stammt auch aus Gesprächen mit

verschiedenen Wissenschaftlern.

Bombyces und Sphinges: Dasychira selenitica, Eilema lutarella, Pelosia muscerda, Hy-

bocampa milhauseri, Lophopteryx cuculla, Heterogenea asella, Tethea ocularis,

Tethea fluctuosa, Cilix glaucata, Cosmotriche lunigera, Bacotia sepium, Narycia

monilifera

Noctuidae: Euxoa obelisca, Eugnorisma depuncta, Rhyacia lucipeta*, Rhyacia simulans*,

Sideridis albicolon, Eriopygodes imbecilla (Glazialrelikt), Cerapteryx graminis, Or-

thosia populi, Mythimna pudorina, Amphipyra berbera*, Talpophila matura, Ipimorpha

subtusa, Cosmia affinis, Apamea lateritia, Apamea unanimis, Photedes extrema, P.

fluxa (vielleicht ein Arealerweiterer), Amphipoea lucens, Nonagria typhae, Nonagria

nexa, Atypha pulmonaris, Cucullia lucifuga, Parastichtis suspecta, Cirrhia aurago,

Cirrhia gilvago, Cirrhia ocellaris, Hyboma strigosa (Arealerweiterer?), Chrysaspidia

putnami, Chrysoptera c-aureum, Parascotia fuliginaria, Colobochyla salicalis, Zanc-

lognatha tarsipennalis, Trisateles emortualis

Geometridae: Comibaena pustulata, Hemithea aestivaria, Hemistola chrysoprasaria,

Sterrha muricata, Sterrha inquinata, Sterrha dimidiata, Sterrha emarginata, Scopula

rubiginata, Anaitis praeformata, Acasis viretata, Lygris testata, Thera firmata, Xan-

thorhoe designata, Calostigia olivata, Euphyia molluginata, Perizoma bifaciata, Peri-

zoma flavofasciata, Anticlea badiata, Pelurga comitata, Hydrelia testaceata, Asthena

anseraria, Eupithecia inturbata, Eupithecia valerianata, Eupithecia venosata, Eupithe-

cia egenaria, Eupithecia extraversaria, Eupithecia selinata, Eupithecia abbreviata,

Eupithecia dodoneata (neu für Südbayern! HAUSMANN in Vorber.), Eupithecia laricia-

ta, Calliclystis chloerata, Arichanna melanaria, Abraxas grossulariata, Ellopia fasciaria,

Abb. 3c und 3d: Sterrha muricata (oben) und Abraxas grossulariata (unten), zwei in

Südbayern nur lokal verbreitete Spanner (Geometridae).

Deuteronomos alniaria, Lozogramma chlorosata, Itame fulvaria, Iheria rupicap-

raria, Erannis bajaria, Erannis leucophaearia, Apocheima hispidaria, Boarmia robo-

raria, Ectropis extersaria

Interessant sind auch die extremen Häufigkeiten von sonst weniger beobachteten Arten

wie Scotia clavis, Amathes sexstrigata, Peribatodes rhomboidaria und einiger anderer.

Die mit * gekennzeichneten Arten sind als wanderverdächtige Arten wie Chloridea

peltigera oder Nycterosea obstipata in bezug auf die Thematik dieser Arbeit natürlich dif-

ferenziert zu betrachten. Es handelt sich hierbei wohl nicht um stabile lokale Populationen.

5. DAS PHANOMEN TURNOVER

5.1. ARTEN-ZEIT-BEZIEHUNG

Jedem, der sich faunistisch mit der Erstellung von Nachtfalter-Artenspektren befaßt,

begegnet schon bald eine Reihe von Gesetzmäßigkeiten, wie z.B. das Phänomen, daß

man im ersten Erhebungsjahr bei einer "punktuellen"” Erfassung von Imagines auch

unter größten Anstrengungen (Licht-, Köder-, Pheromonfang u.s.w.) nie ein vollständi-

ges Artenspektrum eines definierten Standortes erhält. Es werden in den Folgejahren

immer weitere Arten hinzukommen.

So erwähnt URBAHN (1973) neben langfristig beobachteten Häufigkeitsfluktuationen

auch das Phänomen, daß manche Arten in Zeiträumen von einigen Jahren oder gar

Jahrzehnten tatsächlich verschwinden. Sie werden "gewissermaßen ersetzt durch Neuan-

kömmlinge. Dies alles ist bekannt und immer so gewesen ..." (URBAHN, .c.).

Wenn man sich als Beispiel einmal die im Garten (SiN) in den letzten 5 Erhebungs-

jahren nachgewiesenen Eulenfalter (Noctuidae) betrachtet, ergibt sich folgendes Bild (der

Kurvenverlauf wurde durch die Mittelwerte gelegt):

200

150

100

t (Jahre)

1 2 3 4 5 6

Abb 4: Zunahme der Artenzahl (n) der Familie Noctuidae im Lauf der letzten 5 Erhe-

bungsjahre im Garten des Verfassers (SiN).

Increase of species number (n) in the family Noctuidae during the last 5 trapping-

years in the garden of the author.

Der Kurvenverlauf zeigt eine Annäherung an eine Gesamt-Artenzahl von 170 Arten.

Nimmt man diesen Wert mit 100 % an, so ergibt sich im 1. Jahr ein Prozentsatz von

58,5 %, im 2. Jahr 72,6 %, im 3. Jahr 82,2 %, im 4. Jahr 89,4 % und im 5. Jahr 93,5 %.

Das bedeutet, daß bei derartigen punktuellen Methodiken Erfassungsintervalle von min-

destens 4 Jahren erforderlich sind, um ein einigermaßen "komplettes" Artenspektrum zu

erhalten.

In einem durchschnittlichen ersten Jahr fehlen also ca. 70 Arten. Es bleibt jedoch un-

klar, ob es sich um eine tatsächliche Abwesenheit handelt, bzw. wie hoch der Prozent-

satz der Arten ist, die lediglich nicht erfaßt wurden.

Bei einer flächendeckenderen Methodik ergeben sich vollständigere Werte für das erste

Jahr und eine schnellere Annäherung an die maximale Artenzahl (siehe 6.3.1.).

Zur Abschätzung der Fehlerquellen ist zu berücksichtigen, daß sich die Abbildung 4

ausschließlich auf die durchgeführten Lichtfallenfänge stützt. Eine Reihe von Arten (ca.

10), die ein offensichtlich nicht optimales Anflugverhalten ans Licht zeigen, wurde

daher ausgeklammert.

Inwieweit einige weitere Arten auf diese Weise methodisch bedingte "Zuwachsvorgänge"

vorgetäuscht haben könnten, muß an dieser Stelle ebenso offenbleiben wie die Frage,

wie groß der Anteil der Arten ist, die durch ihre Seltenheit in der vorliegenden Me-

thodik unter der "Erfassungsschwelle" geblieben sind. Da es sich bei Lichtfallenfängen

um eine Probeentnahme handelt, können sehr seltene Arten in einer Stichprobe auftau-

chen, in einer anderen jedoch fehlen und so zu einer Unterschätzung der Artenzahlen

in den ersten Erhebungsjahren führen. Weitere Aufschlüsse werden uns hierzu die

Häufigkeitsverteilungen (Kapitel 7) geben.

Fehlerquellen sind also theoretisch alle Faktoren, die Arten unerkannt lassen, die im

Einzugsbereich der Lichtfalle anwesend sind, wie z.B. auch die Witterung, der Fang-

rhythmus, Bestimmungsprobleme (siehe 3.3.), Fallenbeschaffenheit, Lichtqualität u.s.w..

Prognosen, die mittels Arten-Zeit-Beziehungen getroffen werden, können jedoch zu gu-

ten Ergebnissen führen: Die in HAUSMANN (1988) abgebildete Kurve postulierte für

das Jahr 1988 einen Artenzuwachs von 30 Arten, real waren es dann 28 neue Nach-

weise.

Bei der Betrachtung der Arten-Zeit-Kurven (z.B. Abb. 4) werden Artenumsätze von

Jahr zu Jahr ersichtlich: Im zweiten Jahr kommen im Garten, verglichen mit dem Aus-

gangsjahr, durchschnittlich 24 Noctuiden-Arten hinzu. Bei gleichbleibender Artensumme

bedeutet das auch 24 fehlende Arten. Diesen Artenaustausch bezeichnet man als "Turn-

over".

5.2. FORMELN

Die dynamischen Prozesse, die sich auf der Ebene der Artenspektren abspielen, finden

erst seit der richtungsweisenden Publikation MAC ARTHUR & WILSONSs (1967) in ver-

stärktem Maße Beachtung. Jene hatten anhand von kleinen Inseln sehr anschaulich ge-

zeigt, daß sich der Artenbestand in einem dynamischen Gleichgewicht befindet, das von

der Größe der Insel ("Auffangtrichter" für Immigranten) und den Barrieren gegen die

Verbreitung (Abstand zum Festland) abhängig ist. Dieses Modell ist in gewissen Gren-

| zen auch für "Habitat-Inseln auf dem Festland, z.B. ... einem zurückgebliebenen

Sumpf" anwendbar (MAC ARTHUR & WILSON, 1.c.). Man könnte also auch im Unter-

suchungsgebiet die Artenspektren relativ isolierter Standorte, wie beispielsweise das

Wasserwerk oder die verbliebenen Moorbirkenwäldchen (Birket, Torfeinfang, Franzo-

senhölzl) als in einem Gleichgewicht befindlich betrachten, das von einem ständigen

Artenaustausch geprägt ist (Immigration und Extinktion von Populationen). An weniger

verinselten Lebensräumen finden vermutlich auf lokalem Niveau ähnliche Prozesse

statt, die sich jedoch schneller wieder ausgleichen können.

Zur Berechnung der Arten-Austauschraten (Turnover, is) von Jahr zu Jahr verwen-

det man zweckmäßigerweise die Formel für den absoluten Turnover, dem prozentualen

Anteil der ausgetauschten Arten am Ausgangsartenspektrum:

(X, ? X,) 100 % S; = Ausgangsartenzahl

S, X] Zahl der neu hinzugekommenen Arten

X, = Zahl der fehlenden Arten

Wenn beispielsweise im Garten (SiN) 1987 90 Noctuidenarten, 1988 dagegen 89 festge-

stellt wurden, so spiegelt die Stabilität der Artenzahl keineswegs stabile Verhältnisse

in der Artenzusammensetzung wieder: 21 Arten kamen hinzu, 22 konnten nicht mehr

nachgewiesen werden. In die o.g. Formel eingesetzt ergibt sich folgender Artenumsatz:

(21 + 22) 100

Für unser eingangs erwähntes Beispiel der Austauschrate von Jahr I nach Jahr 2 in der

Arten-Zeit-Beziehung (Abb. 4) errechnet sich ein Turnover von 48,3 %.

Nach DIAMOND (1969) ist die Höhe des Turnovers von der Länge des "Census Inter-

valls" zwischen zwei Aufnahmen abhängig. Zur Berechnung des Relativen Turnovers

(T,) dient folgende Formel:

(T + E) 100 I = Zahl der hinzukommenden Arten

Tea less) E = Zahl der fehlenden Arten

S, = Gesamtartenzahl (1. Aufnahme)

S,= Gesamtartenzahl (2. Aufnahme)

Zeitspanne (Jahre) zwischen S, und S,

(Census Intervall)

Bei längeren Zeitspannen kommt in zunehmendem Maße ein "in-and-out-effect" (das

Wiederauftauchen ausgelöschter Populationen und umgekehrt) zum Tragen, wodurch die

Summe der Austauschereignisse unterschätzt wird (DIAMOND & MAY, 1976).

In den folgenden Berechnungen wird unter "Turnover” stets der absolute Turnover

verstanden.

Es erscheint wichtig, den "Pseudoturnover” (NILSSON & NILSSON, 1983), verursacht

durch Stichprobenfehler (siehe 5.1.) vom tatsächlichen Austauschgeschehen abzutrennen.

Bei punktbezogenen Erhebungen sind nun einige verschiedene Reaktionsmuster der Arten

denkbar: Bei mobilen, mehr oder weniger ubiquitären Arten hängt das Ausmaß von

verursachten Turnoverereignissen von der Abundanz ab, häufige Arten (z.B. die häufig-

sten Wanderfalter) werden stets nachgewiesen, seltene Arten verurachen Turnover;

hier wäre als Beispiel der Wanderfalter Nycterosea obstipata zu nennen.

Bei relativ ortstreuen Arten kommt es nun neben dem Ausmaß der Dispersionsaktivität

auf die Entfernung zum typischen Habitat an: Wird die Art im typischen Lebensraum

erfaßt, kommt es zu wenigen Turnoverereignissen, ist das Habitat sehr weit entfernt,

wird ein Zuflug selten erfolgen. Auch hier ist - in langen Zeiträumen betrachtet - der

Turnover vergleichsweise klein. Bei "mittleren" Entfernungen kommt es dagegen in

einer starken Weise zu einem Wechsel von Auftauchen und Verschwinden. Eine Ermitt-

lung solcher Distanzen kann für Fragen bezüglich der Vernetzungen von Biotopen von

Interesse sein.

6. INTERPRETATION DES APPARENTEN TURNOVERS

6.1. BERECHNUNG DER AUSTAUSCHRATEN

6.1.1. Vorbemerkungen

Um den Turnover über mehrere Jahre hinweg auf die Konstanz des Wertes hin zu te-

sten, werden in einem ersten Schritt die Austauschvorgänge im Garten des Verfassers

(SiN) etwas genauer beleuchtet. Für die Tests der Standortabhängigkeit und der Ab-

hängigkeit von verschiedenen Artengruppen (nach Kriterien der Systematik, der Jahres-

zeiten u.s.w.) scheinen sich v.a. die Werte, die sich aus den Fängen der letzten 3

Jahre errechnen, durch die konstant gehaltene Meihode zu eignen.

Für die Berechnungen wird die Formel für den absoluten Turnover verwendet.

Einige Arten mußten wegen ihres nicht optimalen Anflugverhaltens ausgeklammert

werden.

6.1.2 Liste der Turnover-Werte

6.1.2.1. Garten (SiN; Tab. 4):

In den 5 Erhebungsjahren wurde mit einigermaßen vergleichbarer Methode Lichtfang

betrieben: Standort und Bau der Falle waren praktisch konstant, die Anzahl der Fang-

nächte in der Hauptflugzeit (10.6.-31.8.) betrug 25, 39, 38, 30 bzw. 35. Mit einer Aus-

nahme (1985) wurde stets über 100mal pro Jahr gefangen.

Eine Verfälschung der Austauschraten 1983 > 1985 findet durch die Addition zweier

Jahresturnover und durch den "in-and-out-effect" statt: Es handelt sich um zwei ge-

genläufige Komponenten, die sich vielleicht mehr oder weniger egalisieren (vergleiche

DIAMOND & MAY, 1976).

"BOMBYCES UND SPHINGES" (ohne Zygaenidae, Sesiidae, Psychidae und einige

nicht optimal erfaßbare Arten, z.B. Macroglossum stellatarum)

für TO verwen- Immigrations- Extinktions- Arten

Re Turnover Ä

dete Arten 2 ereignisse ereignisse addiert

species-2, used accumulated

for turnover species-,

nur vor

1983 1 = - = (1)

1983 33 = = 2 33

1985 24 6 15 63,6 % 39

1986 29 12 Z 79,2 % 47

1987 30 9 8 58,6 % 49

1988 Z7 6 9 50,0 % 51

62,8 %

NOCTUIDAE (ohne einige nicht optimal erfaßbare Arten, z. B. Ectypa glyphica)

für TO verwen- Immigrations- Extinktions- Arten 2

IH R.33 Turnover Ä

dete Arten I ereignisse ereignisse addiert

nur vor

1983 1 = = = (1)

1983 111 = = - 111

1985 104 19 26 40,5% 130

1986 100 PER 31 55,8 % 148

1987 90 22 32 54,0% 154

1988 89 21 22 47,8 % 159

49,5 %

GEOMETRIDAE (ohne einige nicht optimal erfaßbare Arten, z.B. Ematurga atomaria)

für TO verwen- Immigrations- Extinktions- Arten &

sc Fi Turnover 3

dete Arten I ereignisse ereignisse addiert

nur vor

1983 2 = = = (2)

1983 97 = = z 97

1985 56* Se 46* 52,6*% 102*

1986 100 528 8* 107,1*% 129

1987 93 20 27 47,0% 141

1988 102 23 14 39,8% 146

((61,6 %))*

Die mit * gennzeichneten Werte sind verfälscht, da bei den Geometriden 1985 die

Artbestimmung nicht in allen Fällen mit der nötigen Genauigkeit durchgeführt wurde.

EEE

Der vom Jahr 1983 auf das Jahr 1986 berechnete Turnover beträgt 56,7 %, eine

Verfälschung dieses Austausches durch das Mehrjahresintervall und durch stattfin-

denden in-and-out-effect ist zu veranschlagen.

Der durchschnittliche Geometriden-Turnover im Garten würde dann 47,8 % (Mittel

aus drei Werten) betragen.

MACROHE TEROCERA &

für TO verwen- Immigrations- Extinktions- Arten 2

RN SR Turnover 2

dete Arten 2 ereignisse ereignisse addiert

nur vor

1983 4 = = = (4)

1983 241 = = = 241

1985 183* Zu 86* 47,1*% 271

1986 229 89* 45% 13.2°% 324

1987 213 51 67 51,5 % 347

1988 218 50 45 44,6 % 356**

(654,1 %))*

* Siehe Bemerkungen zur Familie Geometridae! Ein besserer Durchschnittswert er-

rechnet sich wohl aus den beiden letzten Werten mit 48 %.

*#*Dje bisher im Garten (vorne) festgestellte Gesamtartensumme unter Einschluß der

Arten, die für die Turnoverberechnungen nicht berücksichtigt wurden, beträgt 366.

Nimmt man auch die 30 m entfernt an WaS sowie die bis Juni 1989 neu festgestell-

ten Arten hinzu, so kommt man auf 414 Arten.

6.1.2.2. Zusammenstellung der in den letzten 3 Jahren festgestellten Turnoverwerte (Tab. 5)

"BOMBYCES UND SPHINGES"

SiN WaN HO HM HW Mittel

1986/87 58,6 % = 29,8 % 38,7% zu 153%

1987/88 500% 38,6% 455% 70,4 % 31,1% 3

Mittel 54,3% 38,6 % 37,6 % 54,6 % 31,1%

NOCTUIDAE

SiN WaN HO HM HW Mittel

1986/87 54,0% = 35,3% 61,0 % > } 46.0 %

1987/88 47,8 % 33,9% 40,7% 50,0 % 45,3 % :

Mittel 50,9 % 33,9 % 38,0 % 553% 45,3%

GEOME TRIDAE

SiN WaN HO HM HW Mittel

1986/8 47,0% = 48,4% 102,6 % = 578 %

1987/88 39,8 % 47,3% 420% 882% 47,1% A

Mittel 434% 473% 45,2 % 95,4 % 471%

MACROHE TEROCERA %&

SiN WaN HO HM HW Mittel

1986/87 51,5 % = 39,1% 66,9 % > hr

1987/88 44,6 % 39,8 % 42,0 % 62,9 % 43,4 % f

Mittel 48,1% 39,8 % 40,6 % 64,9 % 43,4%

6.1.3. Folgerungen

Der Turnover scheint an einem definierten Standort von Jahr zu Jahr konstant zu

bleiben, deutlichere Schwankungen wurden bisher nur bei den Bombyces und Sphinges

beobachtet (z.B. HO und HM), was aber vermutlich an der größeren Störanfälligkeit

aufgrund der geringen Stichprobengröße liegt.

Wie groß der Einfluß des Witterungsverlaufs eines Jahres auf den Turnover ist, läßt

sich anhand des Materials noch nicht abschätzen, da im Zeitraum der Untersuchun-

gen keine genügend krassen Unterschiede auftraten.

Das Ausmaß der Artenaustauschprozesse ist stark vom jeweiligen Standort (Biotop-

beschaffenheit) abhängig:

Der hohe Turnover im Offenland (HM) erklärt sich durch das vergleichsweise starke

Auftreten von Gastarten aus mehr oder weniger entfernten Habitaten (siehe 6.2.).

Am reich strukturierten Waldrand (HO) liegen zwei unterschiedliche Lebensraumty-

pen in unmittelbarer Nähe des Fallenstandortes. Zufliegende Nachtfalterarten spielen

hier eine geringere Rolle, der Turnover ist daher niedriger.

Der im Siedlungsgebiet (SiN) relativ große Artenaustausch deutet vermutlich eine

Abhängigkeit des Turnovers vom Sukzessionsstadium bzw. dem Ausmaß der Störungen

und Eingriffe in das Ökosystem durch den Menschen an. SPITZER & LEPS (1988)

wiesen eine Korrelation von Sukzessionsstadium des Lebensraumes und der Höhe der

Abundanz-Schwankungen (Fluktuation) der Arten nach. Inwieweit nun Turnover im

Artenspektrum und Fluktuationen in den Häufigkeiten der Arten miteinander verknüpft

sind, wird in einem folgenden Kapitel getestet (6.4.).

Die Frage, ob es sich nicht nur um eine Abhängigkeit von der Stichprobengröße han-

delt (diese war an den Standorten HM und SiN am kleinsten), bleibt primär jedoch

offen und wird gesondert untersucht (z.B. Kapitel 7).

Der Turnover ist abhängig von der gerade betrachteten Artengruppe. Besonders deut-

lich wird dies bei den Geometriden an HM, bei denen die ausgetauschten Arten bis-

weilen das Ausgangsartenspektrum an Zahl übertreffen. Da diese Familie vorwiegend

aus Bewohnern der Wälder und deren Ränder besteht, zeigt sich, daß die Abhängigkeit

von der Artengruppe im Grunde eine Abhängigkeit vom Ökotyp der betrachteten

Arten und den jeweiligen Distanzen und Flächengrößen der benachbarten geeigneten

Lebensräume ist.

Der Gesamtturnover für die Spanner (Geometridae) fällt wegen des Offenlandwerts

(HM) relativ hoch aus. In manchen Habitaten, z.B. im Garten (SiN) liegt er jedoch

niedriger als der Artenumsatz bei den Noctuiden.

Die Geometriden-Unterfamilie Boarmiinae fällt an Waldrand-Standorten mit einem

niedrigen Turnover von 36,9 % (HO, Mittelwert aus zwei Jahren) bzw. 37,1 % (WaN)

aus dem Rahmen. Im Garten lag er sogar noch darunter (siehe 6.4.3.). Im österrei-

chischen Gitschtal (aus WIESER, 1987) war Ähnliches festzustellen, hier war der

Effekt bei 44,0 % für die Geometriden und 23,4 % für die Boarmiinen noch stärker.

Bemerkenswert ist auch das gleichmäßige Niveau der Geometriden-Austauschraten an

allen Standorten, wenn man das Offenland (HM) außer Acht läßt: Es ergibt sich ein

Wert von 45,3 % bei einem Variationskoeffizienten von nur 7,7 %.

- Die Größenordnung des Turnover liegt in den meisten Biotopen zwischen 35 und 55

Prozent/Jahr, die Ergebnisse sind bei definierter Methodik reproduzierbar. Der Arten-

austausch von durchschnittlich 36,7 %, der sich aus den Zahlenkolonnen WIESERs

(1.c.) errechnet, liegt in diesem Rahmen und ähnelt stark den im Wasserwerk (WaN)

festgestellten Verhältnissen.

6.2. TURNOVER DURCH KOLONISATIONSVERSUCHE BIOTOPFREMDER ARTEN

Eine indirekte Methode, Artenspektren ökologisch zu gewichten, ist die Ermittlung des

"Spanneranteils”. In dieser Familie ist der Anteil an Wald- und Waldrandbewohnern

zwar deutlich höher, als z.B. bei den Eulenfaltern (Noctuidae), doch geben so ermit-

telte Werte nur einen sehr unvollständigen ersten Anhaltspunkt über die Präsenz

dieses Ökotyps.

An den Fangstellen ergibt sich folgendes Bild (Angaben in %):

Tab. 6: Spanneranteile (%) an den verschiedenen Standorten 1987 und 1988.

Percentages of the Geometridae at the various trapping-sites 1987 and 1988.

SIEDLUNG WALD HALBTROK- "DACH.

| Garten | Wasserwerk KENRASEN MOOS”

SiS SIM|SiN WaS|WaM WaN WNw WNo HO| HM HW Au We Mb

187 4 31 #3 4 40 3% TEEN Al

1988 46 43 ae 40 00 3 ae

Im Offenland (HM), aber auch an Randbiotopen mit leichtem Offenlandcharakter (SiM,

HW) ist der Spanneranteil niedriger. Sogar die etwas polarisierten Wasserwerks- Werte

sind mit der Lage der Standorte korreliert, da WNw dem Waldrand am nächsten liegt.

Genauere Informationen ergeben sich allerdings, wenn die Beurteilung auf der Basis

der in der Artenliste (4.3.) angegebenen ökologischen Charakteristik der Arten erfolgt.

Wertet man die Haupt-Ökotypen des Gesamtartenspektrums aus, so kommt man zu

folgender Zusammenstellung:

Tab. 7: Anteile der verschiedenen Ökotypen am Gesamtartenspektrum (Macroheteroce-

ra) im Untersuchungsgebiet.

Ecotype-percentages of the species spectrum (Macroheterocera).

UÜb: 50 Arten (9,8 %) Geb: 174 Arten (34,3 %)

mGr: 80 Arten (15,8 %) W: 120 Arten (23,6 %)

Agr: 17 Arten (3,3 %) Hy: 44 Arten (8,7 %)

Xe: 23 Arten (4,5 %)

Im Offenland (HM) zeigen sich bei einer derartigen Einteilung erwartungsgemäß deut-

lich erhöhte Werte bei den Ubiquisten (20,4 %) und den mesophilen Arten des Gras-

landes (23,1 %), leicht erhöht sind sie bei den Arten des Ackerlandes (5,4 %) und den

Xerothermophilen (5,0 %), während die Arten der Wälder, der Waldränder, der Ge-

büschformationen und Hecken deutlich unter den Gesamtwerten liegen (Geb: 24,9 %;

w: 17,2 %).

Man könnte also (verallgemeinernd) von ca. 46 % "biotopfremden” Arten sprechen,

wenn man einmal alle Offenlandarten ohne die hygrophile Fauna als biotoptypisch be-

zeichnet.

Eine verfeinerte Methode der Betrachtung wird zeigen, daß der Prozentsatz der "Über-

fremdung” dieses Standortes noch höher ist: Hierzu wurde unter Berücksichtigung der

Raupenfutterpflanzen getestet, welche Arten mindestens aus dem 150-300 m entfernten

Ruderalgelände (siehe 2.2 und 9.2.) und welche mindestens vom Flughafenrand

(800-1000 m) herbeigeflogen sein müssen.

Vom Ruderal stammen - so beurteilt - 88 Arten (39,8 %) und vom Flughafenrand 36

Arten (16,3 %). Bei diesem Anteil der Gastarten von 56 % handelt es sich wie gesagt

um Mindestangaben!

Tab. 8 zeigt, daß solche biotopfremde Arten verstärkt am Turnovergeschehen beteiligt

sind, daß also der apparente Turnover, der in den mit der vorliegenden Methodik erar-

beiteten Artenspektren zu beobachten ist, durchaus mit tatsächlich in der Natur statt-

findenden Prozessen verknüpft und keineswegs nur ein methodischer Artefakt ist.

Tab. 8: Artenzahlen und Turnover bei bodenständigen und zugeflogenen Arten im Of-

fenland (HM) 1986-1988.

Species number and turnover of within habitat propagating and the visiting species in

the study areas of open habitats.

Arten-2_ Turnoverereignisse

St 1986-1988 pro Art und Jahr

atus

2 ber events of turnover per

BROGIES NINE species and year

potentiell bodenständig 97 0,29

species potentially propagating in site

mindestens aus 150-300 m 88 0,57

at least from 150-300 m distances

mindestens aus 800-1000 m 36 0,54

at least from 800-1000 m distances

Wenn man eine grobe Abschätzung des Ausmaßes von Kolonisationsversuchen ins Of-

fenland hinein wagt, kann man von ca. 150 Wald-, Waldrand- bzw. Gebüscharten

ausgehen, die an den Rändern des Flughafengebiets vorkommen dürften, aber im Rude-

ral, wo Weiden die einzigen Bäume darstellen, keine Lebensgrundlage haben. Für ca.

1/4 dieser Arten konnte also im 3-Jahres-Zeitraum die Bewältigung der Distanz von

ca. 1 km nachgewiesen werden. Da sich darunter vergleichsweise wenige 99 befanden,

wäre ein hypothetisches, an dieser Stelle neugeschaffener Biotop in dieser Zeitspanne

nur von recht wenigen Arten dieser Gruppe besiedelt worden. Möglicherweise könnten

jedoch von den Futterpflanzen ausgehende Geruchsstoffe 99 bevorzugt anlocken.

Es zeichnet sich aber ab, daß man bei einer Anlage von solch isoliert liegenden Bio-

topen (z.B. auch im Ackerland) mindestens 10 Jahre, wenn nicht länger, warten muß,

um auf natürlichem Wege zu einem einigermaßen biotoptypischen Artenspektrum zu

gelangen. Ähnlich langsame Besiedelungsprozesse vermutet HEYDEMANN (1980) für

die Arten der Feuchtbiotope. Erfolgskontrollen nach 1-5 Jahren, wie sie oft praktiziert

werden, sollten in solchen Fällen daher differenziert betrachtet werden.

Im Ruderal kann man den Artenbestand, beurteilt nach Fläche und Reichhaltigkeit der

Vegetation (Krautschicht sehr artenreich, wenige Gebüsch- und Baumarten) auf ca.

250-350 Arten schätzen. Abzüglich der auch an HM möglicherweise bodenständigen

Arten ergeben sich 150-250 Arten, von denen in 150-300 m Entfernung ca. 1/3 bis die

Hälfte in der Zeitspanne von 3 Jahren beobachtet werden konnte. Zu berücksichtigen

ist hier die im Vergleich zu den umliegenden Wäldern geringere Ausgangsfläche (nur

ca. 5 ha).

An den anderen Standorten ist eine derartige Beurteilung schwieriger, es können nur

Einzelinformationen gegeben werden, wie im Fall der im Garten nachgewiesenen Nacht-

falter, die durch die Larvalökologie an Pflanzen der Gattungen Salix, Populus und Al-

nus gebunden sind. Keine dieser Pflanzen kommt in einem Radius von 200-300 m um

den Fangplatz herum vor.

In 5 Erhebungsjahren (siehe 6.1.2.1. und 9.2.) traten 17 so spezialisierte Arten auf, die

durchschnittlich 0,51 Turnoverereignisse pro Art und Jahr verursachten. Auch zwei

weitere außerhalb dieses Intervalls gefangenen und auf Weiden spezialisierte Arten

wurden nur in einem Jahr (? Austauschereignisse) beobachtet.

Für den Rest des Artenspektrums (339 Arten) errechnen sich 0,31 Turnoverereignisse

pro Art und Jahr. Diese Ergebnisse ähneln den an HM (Ruderal) gemachten Beobach-

tungen stark.

6.3. ABHÄNGIGKEIT DES TURNOVERS VOM FLÄCHENINHALT DES ABGEDECKTEN

AREALS

6.3.1. Vergleich der Werte

Tab. 9: Vergleich der Austauschraten in der Gesamtfläche mit denen eines Einzel-

standortes.

Comparison of turnover-rates in the total area with those of a single trap.

| GESAMTFLÄCHE l

Jahr | Standorte für Turnover ver- | GARTEN (SiN)

| wendete Arten-2 Turnover | Turnover

| TOTAL AREA |

year | localities species-2, used | GARDEN

| for turnover |

1986 4 351 = =

1987 10 416 24,8 % 31,5 %

1988 10 428 15,4 % 44,6 %

Eine Unschärfe dieses Vergleichs ergibt sich aus zwei Gründen: Erstens wurde durch

das Fallennetz nicht eine einheitliche Fläche abgedeckt, es war von Jahr zu Jahr ein

Ausweichen von Populationen in nicht fangabgedeckte, zwischen den Fallen liegende Bio-

tope möglich.

Zweitens sind einige "Austauschereignisse” auf der Gesamtfläche auf den Wechsel von

Standorten und die damit verbundene Veränderung der Biotopbeschaffenheit zurückzu-

führen. Daher sind vor allem die Jahre 1986/1987 wenig vergleichbar.

Nach einer Bereinigung dieser Unsicherheitsfaktoren würde der Gesamtflächen- Turnover

noch niedriger liegen, vermutlich bei 5-10 %!

Bei einer größeren Flächenabdeckung durch mehrere Fallen verringert sich also der

Turnover in einer drastischen Weise. Es findet eine Art Abpufferung des sich meist

lokal abwickelnden Artenumsatzes statt.

6.3.2. Rückschlüsse für faunistische Ansätze

Aus dem genannten Phänomen leitet sich ein erhöhter Wirkungsgrad der Methode ab:

Anstelle der normalerweise bei einer punktuellen Nachtfaltererhebung pro Jahr nach-

zuweisenden 50-65 % des Artenspektrums (siehe Arten-Zeit-Kurve, 5.1.) wurden 1987

und 1988 jeweils ca. 440 von 550 im Gebiet zu erwartenden Nachtfalterarten beo-

bachtet (siehe HAUSMANN 1988). Das bedeutet im ersten Jahr einen Anteil von 80 %,

im Zweijahresintervall 1987/1988 (473 Arten) sogar 86 %.

6.4. TURNOVER, KONSTANZ UND FLUKTUATION

6.4.1. Tagfalter (Unterer Inn: REICHHOLF, 1986)

Die Auswertung von umfangreichem Tagfalter-Material aus 10jährigen Linientaxierun-

gen am unteren Inn ergab den interessanten Befund, daß bei recht stabilen Artenzahlen

pro Jahr ein "außerordentlich hoher Artenumsatz ..., der im Mittel 36 % beträgt"

(REICHHOLF, 1.c.) festzustellen war. "»Stabilität« paart sich hier also mit starker

»Dynamik«" (REICHHOLF, 1.c.), was sich auch in der Konstanz des Auftretens nieder-

schlug: Diese verteilte sich keineswegs gleichmäßig über das Artenspektrum, sondern

es überwogen die ganz unregelmäßig sowie die sehr regelmäßig auftretenden Arten,

während Tagfalterarten mit mittlerer Konstanz unterrepräsentiert waren.

Im Tageslicht spielen bei der angewandten Methode der Linientaxierung - anders als

bei Lichtfallenfängen - die Möglichkeit einer unvollständigen Erfassung des Artenspek-

trums eine sehr untergeordnete Rolle. Und dennoch kam es in punkto Turnover zu

einem recht ähnlichen Ergebnis wie bei den Nachtfaltern in der vorliegenden Arbeit.

Inwieweit sich die Befunde auch im Hinblick auf die Konstanz des Auftretens entspre-

chen, wird im folgenden geprüft werden.

6.4.2. Nachtfalter im Gitschtal (WIESER, 1987)

Auf der Suche nach geeignetem Vergleichsmaterial stößt man bei den Nachtfaltern

schnell auf einen eklatanten Mangel an quantifizierten Artenlisten. Eine solche Quanti-

fizierung wird oft mit dem Argument, die Mengenangaben haben nichts mit den tat-

sächlichen Abundanzen in der Natur zu tun, unterlassen, sie unterbleibt bisweilen

jedoch wohl aus Angst vor Kritik. Auch wenn derartige Zahlenkolonnen natürlich nicht

direkte Rückschlüsse auf Populationsgrößen erlauben, so bieten dennoch die relativen

"Lichtfallen-Häufigkeiten” in bestimmten Fragestellungen wertvolle Informationen.

So kann die Nachtfalterfauna des Gitschtales (WIESER, 1987), die in Lichtfallenfängen

der Jahre 1983-1986 erstellt wurde, gut für die Berechnung von Konstanz, Fluktuation

und Turnover verwendet werden.

‚Das Problem, daß ein Zeitraum von 10 Jahren "als ein Mindestmaß einer verläßlichen

Beobachtung populationsdynamischer Vorgänge" gelten kann (VARGA & UHERKOVICH,

1974), kennzeichnet die folgenden Berechnungen als vorläufig. Statistische Fehlerquellen

werden hierbei zumindest teilweise durch die Betrachtung relativ großer Artengruppen

kompensiert.

Der mittlere Turnover, errechnet aus drei Austauschraten, beträgt im Gitschtal 36,7 %.

Tab. 10: Konstanz, Turnover und Fluktuation im Gitschtal (Österreich) 1983-1986, er-

rechnet aus WIESER (1987).

Constancy, turnover and fluctuation in the "Gitschtal” (Austria) from 1983 to 1986,

calculated from WIESER (1987).

KONSTANZ ARTEN TURNOVER- TURNOVEREREIG- VARIATIONS-

EREIGNISSE NISSE/ ART x JAHR KOEFFIZIENT

constancy species events of events of turnover coefficient of

turnover per species and year variation

20% 102 137 0,45 200,0 %

50% 59 110 0,62 129,1 %,

75% 74 115 0,52 91,9%

100 % 222 = 0,00 58,7%

GESAMT 457 362 0,26 104,7 %

total

n (Arten)

300 eh

V@%

200

100

25 50 IB, 100 y4 25 50 73 100

KONSTANZ

Abb. 5: Konstanz des Auftretens von Abb. 6: Variationskoeffizient (V, mit Staı

Nachtfaltern im Gitschtal. dardabweichung) in Abhängigkeit von d

Constancy of moths in the "Gitschtal" Konstanz bei Nachtfaltern im Gitschta

(Austria). Coefficient of variation (V, with Stan

dard deviation) and constancy in the

Gitschtal.

Die Abhängigkeit von der jeweiligen Artengruppe wird in Tabelle 11 ersichtlich.

Tab 11: Mittlerer Variationskoeffizient und Turnover verschiedener Artengruppen im

Gitschtal (errechnet aus WIESER, 1.c.).

Coefficient of variation and turnover of some systematic groups in the "Gitschtal".

ARTENGRUPPE VARIATIONSKOEFFIZIENT TURNOVER

group of species coefficient of variation turnover

Bombyces und Sphinges 103,0 % 32.2.%

Noctuidae 102,0 % 332%

Geometridae 108,9 % 44,0 %

(Boarmiinae 80,9 % 23,4%)

Die Arten mit mittlerer Konstanz sind wie bei den Tagfaltern am unteren Inn unterre-

präsentiert; es sind hier bei einer größeren Stichprobe (knapp 20.000 Individuen) und

einem kürzeren Erfassungsintervall im Vergleich mit REICHHOLF (1986) die regelmä-

Big auftretenden Arten (rechter Teil der Kurve) etwas stärker vertreten. Dennoch wird

ein nicht unbedeutender Teil des Artenspektrums sehr unregelmäßig nachgewiesen.

Konstanz und Fluktuation sind zwei streng negativ miteinander korrelierte Parameter.

Die meisten Turnoverereignisse eines definierten Artenspektrums werden von den sehr

unregelmäßig auftauchenden Arten verursacht. Pro Art und Jahr gerechnet sind es

allerdings die Arten mittlerer Konstanz, die die größten Austauschraten aufweisen. Das

häufige Auftauchen und Verschwinden solcher Arten kann durch die Position der Licht-

falle in einer bestimmten Distanz zum typischen Habitat bedingt sein: Wird im Rand-

bereich der potentiellen Dispersionsaktivität einer Art geleuchtet, werden verstärkt

derartige Effekte auftreten (siehe Schlußbemerkungen zu 5.2.).

6.4.3. Nachtfalter in Oberschleißheim (Garten)

Im 5-Jahresintervall 1983/1985/1986/1987/1988 wurden nur ausgewählte Arten aus den

Bombyces, Sphinges und aus der Familie Noctuidae berücksichtigt. In bezug auf die

Geometridae gilt die in 6.1.2.1. angesprochene Problematik. Bei einigen weiteren Arten

erschien die quantitative Vergleichbarkeit fraglich, sie wurden ebenfalls ausgeklammert.

Variationskoeffizienten aus 5-Jahresintervallen, wie sie in Tabelle 12 angegeben wer-

den, können allerdings nicht direkt mit 4-Jahreswerten (WIESER, 1987 oder Tabelle

13) verglichen werden. Zur Abschätzung der Korrelation der verschiedenen Parameter

liefern die folgenden Ergebnisse jedoch genügend Einzelinformationen.

Tab 12: Konstanz, Turnover und Fluktuation bei Nachtfaltern im Garten des Verfassers

(SiN) in 5 Erhebungsjahren.

Constancy, turnover and fluctuation of Macroheterocera in the garden of the author

in 5 trapping years.

KONSTANZ ARTEN-2 TURNOVER- TURNOVEREREIG- VARIATIONS-

EREIGNISSE NISSE / ART x JAHR KOEFFIZIENT

constancy species-2 events of events of turnover coefficient of

turnover per species and year variation

20 % 46 1122 0,39 223,6 %

40% 29 60 0,52 142,1 %

60 % 24 51 0,53 113,9 %

80 % 27 43 0,40 92,8%

100 % 39 = 0,00 69,5 %

GESAMT 165 226 0,34 135,3 %

total

n (Arten)

20 40 60 80

100

Abb. 7: Konstanz des Auftretens von

Nachtfaltern im Garten.

Constancy of moths in the

garden (Southen Bavaria).

200

100

% 20 40 60 80

KONSTANZ

Abb. 8: Variationskoeffizient (V, mit Stan-

dardabweichung) in Abhängigkeit von der

Konstanz bei Nachtfaltern im Garten.

Coefficient of variation (V, with Stan-

dard deviation) and constancy in the

garden.

Die Werte und Kurven ähneln im Prinzip den unter 6.4.1. und 6.4.2. gefundenen Ver-

hältnissen.

Die konstanten Arten (100 %) zeigen eine etwas stärkere Fluktuationsdynamik als die

des Gitschtales. Dies könnte auf größere anthropogene Störeinflüsse in Siedlungsgebie-

ten (siehe 6.4.5.) zurückzuführen sein. Jedenfalls entspricht dieser Befund auch der im

Gesamtbild festgestellten größeren Artenumsatz-Dynamik.

Nach verschiedenen systematischen Einheiten aufgegliedert, lassen sich folgende inte-

ressante Einzelwerte feststellen:

Tab. 13: Gegenüberstellung des mittleren Variationskoeffizienten und des

mittleren

Turnovers verschiedener systematischer Gruppen in unterschiedlichen Erfassungszeit-

räumen im Garten.

Mean of coefficient of variation and mean of turnover of various taxa in the gar-

den.

MITTLERER MITTLERER

VARIATIONSKOEFFIZIENT TURNOVER

mean of coefficient mean of

of variation turnover

(5 Erhebungsjahre)

5 years

Bombyces + Sphinges 138,9 % 62,8 %

Noctuidae 133,4 % 49,5 %

Tab. 13 (Fortsetzung)

MITTLERER MITTLERER

VARIA TIONSKOEFFIZIENT TURNOVER

mean of coefficient mean of

of variation turnover

(4 Erhebungsjahre: 1983, 1986, 1987, 1988)

4 years

Bombyces + Sphinges B32% 62,5 % (n=47)

Noctuidae 121.1 % 49,2 % (n=122)

Geometridae 112,2 % 47,8 % (n=131)

- Unterfamilie Boarmiinae 96,7% 28,9 % (n=42)

- Gattung Eupithecia 136,8 % 78,5 % (n=24)

Die gemittelten Variationskoeffizienten und der mittlere Artenumsatz einer definierten

Artengruppe sind also offensichtlich miteinander korreliert. Bei stärkeren Fluktuationen

kommt es statistisch häufiger zu Artenaustausch-Ereignissen.

Trotz der relativ hohen Turnoverraten und Fluktuationen bei den Bombyces und Sphin-

ges ist bei einzelnen Arten die Konstanz der Populationen bemerkenswert, wie z.B. bei

Spilosoma menthastri (100 % Konstanz, V = 25,0 %) oder Spilarctia lubricipeda (100 %

Konstanz, V = 43,9 %). Dieser Einzelbefund deckt sich gut mit den Ergebnissen in

REICHHOLF (1974).

Die in der Zeitspanne von 4 Jahren festgestellten Werte für die Spanner (Geometri-

dae) und die Unterfamilie Boarmiinae stimmen erstaunlich gut mit dem Material aus

dem Gitschtal (WIESER, 1987) überein. Die Boarmien, deren Raupen vorwiegend auf

Bäume angewiesen sind, sind durch die geringen Fluktuationen und den niedrigen Arte-

numsatz als Arten gekennzeichnet, die stabile, relativ ortsfeste Populationen bilden.

Dies steht im Einklang mit den extrem niedrigen Arten-Umsatzraten der Futterpflanzen.

Die erhöhte Dynamik der an SiN nachgewiesenen Eupithecienarten könnte z.T. im Zu-

sammenhang mit instabileren Ressourcen stehen, zu berücksichtigen ist jedoch stets die

Standortabhängigkeit solcher Phänomene: Am Fangplatz WaS beispielsweise scheinen

sich die populationsdynamischen Prozesse bei den Arten der Gattung Eupithecia auf

einem viel niedrigeren Niveau abzuspielen. Andersherum unterliegen die Boarmien im

Offenland (HM) einem relativ hohem Artenaustausch. Eine hohe Ortstreue der Eupithe-

cien konstatieren auch VARGA & UHERKOVICH (1974).

6.4.4. Nachtfalter in Oberschleißheim (Offenland: "HM" und "HO"”)

Im Flughafengebiet (HM und HO) stehen leider nur Ergebnisse aus drei Erhebungsjah-

ren (1986, 1987, 1988) zur Verfügung, wodurch Randeffekte, die zu einer Verzerrung

der Ergebnisse führen können, stärker in Erscheinung treten. Es wird sich aber zeigen,

daß sich dies durch die Fülle des Materials (über 200 Arten an HM) wohl zum großen

Teil gegenseitig ausgleicht und aufschlußreiche Folgerungen zuläßt.

Besonders wertvoll ist eine genauere Analyse der Verhältnisse im reinen Offenland

(HM), da hier die Abgrenzung der Habitate in einer schärferen Weise erfolgen kann

(vergleiche 6.2.):

Tab. 14: Konstanz des Auftretens verschiedener Macroheteroceren-Taxa im Offenland (HM).

Constancy of some Macroheterocera-Taxa in the open grassland (HM).

KONSTANZ" BOMESCEs } meer Re 0 00008

33,3 % 13 44 2 e

66,6 % 15 25 21 6l

Arten-2 42 117 62 “=

species-Z

Den tatsächlich festgestellten Kurvenverlauf der Konstanz, der ganz im Einklang mit

dem bereits diskutierten Befund einer Unterrepräsentation der Arten mittlerer Kon-

stanz steht, kann man sich nun in einem vereinfachten Modell als aus zwei Komponen-

ten zusammengesetzt vorstellen. Hierzu soll von einer hypothetischen scharfen Eintei-

lung in "biotoptypische”, und "biotopfremde"” Arten ausgegangen werden. Folgende

Einzelverteilungen bezüglich der Regelmäßigkeit des Auftretens sind zu erwarten:

Su beobachtet

S\ „ biotoptypisch

50 Rn

8 biotopfremd

Abb. 9: Beobachtete und erwartete Konstanz des Auftretens, aufgeschlüsselt nach der

Bodenständigkeit der Arten.

Observed and expected constancy of species occurrence, separated into the catego-

ries ‘'habitat specific‘ (dotted line) and ‘origin off site‘ (of capture) (broken line).

The solid line gives the observed distribution of constancy .

Je weiter "das Heimathabitat" von der Falle entfernt ist, umso weiter sollte sich in

Abb. 9 die Kurve der Gastarten nach links verschieben. Je großflächiger und unge-

störter der typische Lebensraum um die Falle herum ist, umso steiler wird der Kur-

venverlauf der potentiell bodenständigen Arten sein.

Die Verteilung der Familie der Spanner (Geometridae) in den verschiedenen Konstanz-

klassen entspricht ziemlich genau der postulierten Kurve für biotopfremde Arten. Lenkt

man das Augenmerk darauf, daß der Wald für HM einen solchen entfernten Biotop

darstellt und die Spanner vorwiegend Wald-, bzw. Waldrandarten sind, so scheint

schon dies ein erster Anhaltspunkt für eine Bestätigung der Hypothese zu sein.

Da jedoch in den allerwenigsten Fällen ein solches 2-Biotope-Modell vorliegen wird,

sondern Überlagerungen von mehreren, verschieden weit entfernten Zuwanderungs-

quellen, so werden, falls Beurteilungen überhaupt möglich sind, differenziertere Tests

vonnöten sein.

Hierzu wurde das an HM nachgewiesene Artenspektrum über eine genauere Betrach-

tung der in der Literatur angegebenen Raupenfutterpflanzen, z.T. durch eigene Beo-

bachtungen präzisiert, in drei verschiedene Kategorien eingeteilt: Es können potentiell

bodenständige Arten von solchen abgetrennt werden, die entweder aus mindestens

150-300 m oder aus über 800 m Entfernung stammen müssen (siehe 6.2.). Es handelt

sich also um Mindestangaben. Andere Kleinstbiotope spielen in der näheren Umgebung

auf dem Flughafen eine nur sehr untergeordnete, zu vernachlässigende Rolle.

50 bodenständig

aus mind.

150-300 m

u... aus mind.

O 800-1000 m

33,3 66,7 100 Konstanz (%)

Abb. 10: Beobachtete Konstanz des Auftretens bei Nachtfaltern verschiedener "Hei-

mat-Habitate" im Offenland (HM).

Constancy of occurrence in the light-trap-captures in relation to distance of the

species‘ breeding sites (solid line = highly probable in site propagation, broken line =

distance 150 to 300 metres, dotted line = distance 800 to 1000 metres).

Abb. 10 bestätigt nun die eingangs aufgestellten Überlegungen:

- Der rechte Teil der Kurve (also die regelmäßig auftretenden Arten) besteht fast aus-

schließlich aus den potentiell bodenständigen Arten, während der linke Teil überwie-

gend von den zugeflogenen Gästen beherrscht wird. Das jedoch bedeutet, daß die in

Lichtfallenausbeuten festgestellte Konstanz des Auftretens als Parameter der Popu-

lationsdynamik durchaus mit real in der Natur ablaufenden Prozessen verknüpft ist.

Es handelt sich mit höchster Wahrscheinlichkeit nicht nur um den unregelmäßigen

Nachweis von ständig anwesenden Arten, sondern tatsächlich um ein unregelmäßiges

Auftreten. Dies wiederum bekräftigt den Aussagewert des in Lichtfallen-Artenspek-

tren auftretenden Turnovers!

- Mit zunehmender Entfernung der Ausgangs-Habitate verschiebt sich die Teilkurve der

biotopfremden Arten weiter nach links, wie dies im Falle der vom Flughafenrand

(800-1000 m) stammenden Nachtfalter geschah. Das Auftreten einer Art wird also

mit zunehmender Entfernung vom Habitat unregelmäßiger.

Schon im Gesamtbild (siehe 6.2.) fallen die in verstärktem Maße fehlenden Waldarten

auf. Eine Vielzahl von Arten dieses Lebensraumtyps, die in ca. 1 km Entfernung (z.B.

an HO) nachgewiesen werden konnte, ist also mit 0 % Konstanz zu veranschlagen.

An anderen Standorten ist eine Beurteilung schwieriger, da sich die Effekte einer Viel-

zahl mehr oder weniger kleiner Lebensräume verschiedenster Entfernungen überlagern

und das im reinen Offenland so klar ausgeprägte theoretische Modell im Gesamtbild

verwischen.

Am Flughafenrand (HO) ergibt sich beispielsweise folgendes Bild:

Tab. 15: Konstanz des Auftretens verschiedener Macroheteroceren-Taxa am Flugha-

fenrand (HO).

Constancy of some Macroheterocera-Taxa at the edge of a wood (HO).

KONSTANZ BOMBYCES +

onslancy SPHINGES NOCTUIDAE GEOMETRIDAE 2

33,3% 14 32 29 75

66,6 % 12 30 35 711

100 % 3 76 60 167

Arten-2 37 138 124 319

species-2

Die regelmäßig auftretenden Arten sind hier stärker vertreten, was dadurch erklärt

wird, daß an diesem Waldrand-Standort sowohl Waldarten wie auch Offenlandarten als

"biotoptypisch" angesprochen werden können. Diese auf den Standort bezogene verrin-

gerte Artenspektrendynamik schlägt sich in ebenfalls deutlich niedrigeren Artenum-

satz-Raten nieder als im Offenland.

6.4.5. Eulenfalter (Noctuidae) in Südböhmen und England

Umfangreiche, auf sehr großen Stichproben beruhende Auswertungen von Lichtfallenfän-

gen in Südböhmen und England ergaben wertvolle Informationen v.a. in bezug auf die

Fluktuationsdynamik. Die im folgenden kurz skizzierten Ergebnisse stützen sich auf die

Publikationen von REJMANEK & SPITZER (1982), SPITZER, REIJIMANEK & SOLDAN

(1984), GLAZIER (1986), GASTON (1988) und SPITZER & LEPS (1988). Einleitend

muß darauf hingewiesen werden, daß in diesen Arbeiten nur Arten mit einer gewissen

Mindestkonstanz des Auftretens bzw. einer Mindestabundanz berücksichtigt wurden.

Dies ist bei Vergleichen stets zu bedenken.

Nach SPITZER, REJMANEK & SOLDAN (1984) sind Variationskoeffizient und poten-

tielle jährliche Wachstumsrate der Population (PGR) miteinander stark positiv korre-

liert. Die Arten mit den höchsten festgestellten Variationskoeffizienten sind r-Strate-

gen, ihre Raupen sind polyphag.

Wie wir im vorigen Abschnitt (6.4.4.) gesehen haben wurden durch die bevorzugte

Verwendung konstant auftretender Arten bei diesen Berechnungen v.a. die potentiell

bodenständigen Arten erfaßt und statistisch verwertet. Außerhalb ihrer Habitate können

auch tendenziell von K-Strategen beherrschte Gruppen mitunter höhere Variationskoef-

fizienten aufweisen. Am Standort SiN trifft dies vielleicht bei den Eupithecien (6.4.3.)

zu. Hier waren bei den Berechnungen auch die unregelmäßig auftretenden Arten hinzu-

gezogen worden. Die Boarmien reagierten als K-Strategen mit niedrigen Fluktuationen

jedoch ganz im Sinne der o.g. Autoren.

Die Absolutwerte der in REIJMANEK & SPITZER (1982) publizierten Variationskoef-

fizienten für Arten ab einer Konstanz von 25 % (3 von 12 Jahren) liegen bei durch-

schnittlich 94,6 %. Sie sind mit Werten aus unterschiedlichen Erfassungszeiträumen

jedoch nicht direkt vergleichbar. Die relativen Abweichungen vieler Arten von diesem

Mittelwert entsprechen zumeist den Ergebnissen im Gitschtal und aus dem Untersu-

chungsgebiet.

Das Ausmaß der Abundanzschwankungen ist von der jeweils untersuchten Artengruppe

abhängig (REJMANEK & SPITZER, 1.c.): Wanderfalter und schädliche Arten, vor allem

die der Landwirtschaft, zeigen hohe Variationskoeffizienten, hygro- und mesophile

Arten des Graslandes sowie die Bewohner von "Baum- und Gebüschformationen” unter-

liegen dagegen geringeren Fluktuationen. Dies deckt sich mit den Beobachtungen im

Untersuchungsgebiet.

Die positive Korrelation von Variationskoeffizient und PGR belegen auch SPITZER &

LEPS (1988). Dieselben Autoren fügen hinzu, daB in einem Klimax-Ökosystem die

Häufigkeiten der Arten konstanter sind als in Ökosystemen früher Sukzessionsstadien

mit größeren Störeinflüssen. Für diesen Vergleich hatten sie die Variationskoeffizienten

der jeweils gleichen Arten herangezogen. Ephemere Lebensräume werden darüber hin-

aus von einer größeren Artenzahl an r-Strategen besiedelt. Dies könnte eine Erklärung

dafür sein, daß im Siedlungsgebiet von Oberschleißheim (anthropogene Störeinflüsse

relativ groß) etwas höhere Fluktuationen festzustellen waren, als beispielsweise im

Gitschtal. Diese größeren Häufigkeitsschwankungen führen statistisch häufiger zu einem

lokalen Verschwinden von Arten und damit zu einem höheren Turnover als an anderen

Standorten.

Populationen lokal vorkommender Arten, die als K-Strategen relativ stabile Ressourcen

und Habitate benutzen, besitzen niedrige Variationskoeffizienten (GLAZIER, 1986;

GASTON, 1988).

7. RUCKSCHLUSSE AUS DEN HAUFIGKEITSVERTEILUNGEN

7.1. ARTENZAHL UND STICHPROBENGRÖSSE

In einer ersten Betrachtung könnte man die höheren Umsatzraten im Garten (SiN) und

im Offenland (HM) auf die relativ geringen Individuenzahlen (ca. 1700 bzw. 1000 In-

dividuen pro Jahr) an diesen Standorten zurückführen und behaupten, daß hier durch ge-

ringere Erfassungsgenauigkeit ein größerer Einfluß methodisch bedingter Fehler vorliegt.

Die Abhängigkeit der Artenzahl von der Menge der gefangenen Individuen (ein Maß

der Diversität) - bezogen auf Jahressummen jeweils eines Standorts - geht aus Abbil-

dung 11 hervor.

Gesamtartenzahl,

Bodlren Yorller mach" Tue, Wie ae sk Bett .

EN,

,* x

LERSR

200: ,

NOEMORO

(Individuen)

5000 10000

Abb. 11: Artensumme pro Jahr in Relation zur Individuenzahl; mit Kreis sind die Fang-

plätze We und Mb gekennzeichnet, mit x die übrigen Fangplatz-Jahresergebnisse.

Total species per year and number of individuals; o = two localities in wet woods

("We" and "Mb"): x = the results of the other localities.

Große Individuenzahlen pro Jahr bedeuten bei definierten Einzugsbereichen der Falle

aber auch biologisch reichhaltigere Habitate, die von Natur aus mehr Arten beinhalten.

Andersherum ausgedrückt ist die Stichprobengröße im Garten und im Offenland relativ

gesehen mit der der anderen Standorte vergleichbar, da die Proben aus einem kleineren

Individuen-Pool entnommen wurden.

Relativ stark aus dem Rahmen fallen die zwei mit Kreis gekennzeichneten Werte im

Dachauer Moos (We, Mb). Sie liegen vor allem wegen der überaus großen Häufigkeit

des Spanners Calospilos sylvata unter dem Niveau der anderen Fangplätze. Diese nied-

rigere Diversität könnte entsprechend den Ergebnissen MADERs (1980) in Zusammen-

hang mit der Störung des Lebensraumes durch die Zurückführung des ehemaligen

Niedermoors auf wenige kleinflächige Reliktstandorte und durch Einflüsse seitens der

Landwirtschaft von den Rändern dieser Restflächen her stehen. Deren Moorcharakter

ist seit vielen Jahren durch den stark gesunkenen Grundwasserspiegel verlorengegangen.

Die Stichprobenentnahme erfolgte an den drei Standorten im Dachauer Moos jedoch

mit geringerer Frequenz als an den anderen Stellen. Diese Problematik soll in Kapitel

7.3. besser beleuchtet werden.

Zur Abschätzung von Fehlerquellen bezüglich der Erfassungsgenauigkeit sollte man

vielleicht eher einzelne Standorte im Lauf der Jahre unter Berücksichtigung der Domi-

nanzstrukturen heranziehen.

7.2. VERGLEICH MIT DEN "log-series” WILLIAMS (1964)

Teilt man das 1987 und 1988 im Untersuchungsgebiet erarbeitete Artenspektrum in der

Weise ein, wie dies WILLIAMS (1964) in seinen Auswertungen des Fangergebnisses der

Lichtfalle in Rothamsted tat, so ergibt sich bei einer Zuteilung in die einzelnen geo-

metrischen X 3-Klassen das in Abb. 12 gezeigte Bild.

Artenzahl

number of species

100

I m m VW v VI VI VI x 3-Klassen

Abb. 12: Häufigkeitsverteilung der in den Lichtfallenfängen 1987 und 1988 nachgewie-

senen Arten in X 3-Klassen.

I = 1 Individuum IV = 14-40 Individuen VII = 365-1093 Individuen

2-4 Individuen V = 41-121 Individuen VIIIl= > 1093 Individuen

II = 5-13 Individuen VI = 122-364 Individuen

Abundance structure (in 3 geometric classes) of species of Macroheterocera,

caught by light-trap-captures 1987 and 1988.

Die Verteilung ähnelt am meisten derjenigen eines 8 Jahre dauernden, täglich erfolgten

Lichtfanges in Rothamsted bei einer Ausbeute von knapp 33.000 Individuen.

In bezug auf die Problematik der Erfassungsgenauigkeit interessieren uns nun vor allem

die selteneren Arten der ersten beiden Klassen an Einzelstandorten, sind es doch ge-

rade diese seltenen Arten, die in verstärktem Maße am Turnovergeschehen beteiligt

sind (SCHOENER, 1983).

17.3. ZUM PROBLEM DER ERFASSUNG DER SELTENEN ARTEN

7.3.1. Artenzahl und Fangrhythmus

Aus der Überlegung heraus, daß sich die beobachtete Artenzahl bei einer Optimierung

der Methode durch steigende Anzahl von Fangnächten pro Jahr an einen maximalen

Wert annähern sollte, soll nun die Abhängigkeit vom Fangrhythmus näher beleuchtet

werden. Durch ein auf die Fänge 1988 im Garten (WaS) gelegtes Raster wird getestet,

wieviele Arten nachgewiesen worden wären, wenn nur in jeder zweiten, vierten u.s.w.

Nacht geleuchtet worden wäre. Die Grundlage bilden die Ergebnisse aus den Fängen in

einem mittleren Rhythmus von 1,15 Tagen von Mai bis August bzw. von 1,45 Tagen

von April bis Oktober.

Das Ergebnis zeigt Abb. 13:

300 ee

200

100

Fangnächte (%)

50 100

Abb. 13: (Hypothetische) Artensumme n in Relation zur Anzahl der Fangnächte pro

Jahr (in % der 182 Gesamtfangnächte) am Standort WaS 1988.

Correlation of species number (n) and number of captures per year (in %X of the to-

tal of 182 nights) 1988 in the garden (Southern Bavaria).

Die Kurve zeigt eine Annäherung an einen Wert von 325 Arten im Jahr 1988. Wenn

man bedenkt, daß bei langjährigen Beobachtungen im Garten bisher 411 Arten nachge-

wiesen werden konnten, ist dies ein weiteres Argument dafür, daß in den Ergebnissen

der Einzeljahre tatsächlich Arten fehlen und das Artenspektrum somit einem mehr oder

weniger großen Artenumsatz von Jahr zu Jahr unterliegt.

Bei fast kontinuierlichem Betrieb der Lichtfalle scheinen etwas über 90 % des Arten-

spektrums nachgewiesen werden zu können. Für jede 2. Fangnacht errechnet sich ein

theoretischer Wert von 262,5 Arten (81 %); im Jahr 1987 waren bei einem (vergleich-

baren) durchschnittlichen Fangnachtabstand von 2,3 Tagen (Mai-August) bzw. 2,6

Tagen (April-Oktober) real 240 Nachtfalterarten beobachtet worden. Durch täglichen

Fang werden viele Individuen länger an der Lichtquelle festgehalten, als es einem

natürlichen Aufenthalt im Einzugsbereich der Falle ohne LichteinflußB entsprechen

würde. Dadurch kommt es zu einer leichten Überschätzung der theoretisch ermittelten

Artenzahlen für die Hälfte der Fangnächte. Dies fällt aber offensichtlich ab einer

gewissen Fülle von Einzelinformationen () 10.000 gefangene Individuen) nicht mehr so

stark ins Gewicht. Die Frage bleibt offen, inwieweit die Jahre 1987 und 1988 von der

Witterung her zu vergleichen sind.

In einem weiteren Test der Abhängigkeit der Artenzahl von der Fanghäufigkeit zeigte

die Familie Noctuidae im Jahr 1986 ähnliche Ergebnisse, obwohl hier fangfreie Nächte

zwischen den Stichprobenentnahmen lagen. Die Berechnungen stützen sich auf das

Fangergebnis im Garten am Fangplatz SiN. Die Gesamtartenzahl nähert sich an einen

Wert von ca. 125 Arten an. Bei Fangnachtabständen von durchschnittlich 2,3 Tagen

(Mai-August) und 2,0 Tagen (April-Oktober) waren demnach 79 % in der Probeentnah-

me enthalten, in der Hälfte dieser Fangnächte waren es noch 66 %. Im Jahr 1988

(WaS, alle Nachtfalterarten) betrugen die entsprechenden Werte 81 % bzw. 68 %!

7.3.2. PRESTONs "veil-line”-Theorie

Bei der Betrachtung der Häufigkeitsverteilung des am Flughafenrand (HO) für das Jahr

1988 nachgewiesenen Nachtfalter-Artenspektrums zeigt sich eine Dominanz der Arten,

die in 2-4 Individuen festgestellt wurden:

50 n

x 3-Klassen

I I III IV V VI VI

Abb. 14: Häufigkeitsverteilung der am Fiughafenrand (HO) 1988 nachgewiesenen Mac-

roheteroceren in x 3-Klassen (Legende siehe Abb. 12).

Abundance structure of Macroheterocera (in x 3-—-classes), captured 1988 at the

edge of a wood ("HO").

PRESTON (1948) unterstellte für derartige Häufigkeitsstrukturen eine lognormale Ver-

teilung (vergleiche auch MAY, 1980), bei der wie im vorliegenden Fall durch eine

begrenzte Stichprobenentnahme ein Teil der Kurve "abgeschnitten" wurde. Er nannte

diesen "Schnitt" bei den Individuen, die nur in einem Exemplar festgestellt wurden

"veil line”. Die Arten links davon waren in der Stichprobe nicht enthalten, hätten aber

in einer anderen auftauchen können (WILLIAMS, 1964).

Trägt man nun die aufsummierten prozentualen Anteile der Klassen I-VI an der Ge-

samtartenzahl gegen den Logarithmus der Klassenobergrenze auf (Abb. 15), so erhält

man unter der Annahme eines Nullwertes von 0 Arten einen etwas asymmetrischen

Kurvenverlauf. Dieser kann nun nach WILLIAMS (l.c.) auf der Basis größtmöglicher

Symmetrie durch eine Nullwert-Hypothese korrigiert werden (Abb. 15).

98% 7

log [upper

limit of class)

N (m) I) M MM

Abb. 15: Aufsummierte prozentuale Anteile der x 3-Klassen an der Gesamtartenzahl mit

und ohne Annahme eines Nullwertes am Fangplatz HO 1988 (vgl. WILLIAMS, 1964).

Accumulated percentage of the x 3-classes of the total species on the assumption

of no zero value and on the assumption of a zero value of 35 (see WILLIAMS, 1964).

Führt man diese Operation bei den Artenspektren aller drei Erhebungsjahre an HO

durch, so kommt man bei einer durchschnittlichen Gesamtartenzahl von 256 Arten auf

einen (hypothetischen) mittleren Nullwert von ca. 50 Arten.

Aus der Gesamt-Häufigkeitsstruktur der an diesem Standort 1986-1988 gefangenen

Nachtfalter (338 Arten) ergibt sich ein Nullwert von 40 Arten.

Diese Angaben könnte man benutzen, um die Arten-Zeit-Kurve (siehe 5.1.) zu korri-

gieren. Im ersten Jahr würde unter Annahme der oben genannten Nullwerte dann

durchschnittlich knapp 3/4 des Artenspektrums nachgewiesen werden können (statt ca.

60 %). Der Turnover würde sich von ca. 40 % auf ungefähr 30 % korrigieren.

Mit einer anderen Vorgehensweise waren wir schon an anderer Stelle (7.3.1.) zu einem

fast identischen Ergebnis gekommen: Bei einer Optimierung der Fangintensität wären im

Garten nach zwei voneinander unabhängigen Berechnungen (WaS, SiN) ungefähr 3/4

der hochgerechneten langjährigen Gesamtartenzahl in einem Einzeljahr nachzuweisen

gewesen (statt ca. 60 %).

Hervorzuheben ist allerdings der hypothetische Charakter dieser Aussagen, da offen-

bleibt, inwieweit sich ein solcher theoretisch ermittelter Nullwert mit der tatsächlich

unter der Erfassungsschwelle gebliebenen Artenzahl deckt!

Zu einer derartigen Beurteilung wären direkte Nachweise von methodisch bedingten

Artefakten nötig. Am Flughafenrand (HO) konnten beispielsweise im Sommer 1986

Raupen von Eupithecia tripunctaria und 1988 Raupen von Thyatira batis gefunden wer-

den. Die jeweiligen l. Generationen mußten also mit mindestens einem 9 im Gebiet

vertreten gewesen sein, obwohl in den betreffenden Jahren am Licht kein Imago be-

obachtet wurde. Der von Thyatira batis verursachte Turnover kann daher als methodisch

bedingt charakterisiert werden, bei Eupithecia tripunctaria ergibt sich jedoch ein zu-

sätzliches Turnoverereignis.

Derartige Einzelinformationen sind, gemessen an der Größe des Artenspektrums, nur

Mangelware und werden auch bei einer Optimierung anderer Methoden nie zu einem

umfassenden Bild des Anteils methodischer Störeinflüsse im apparenten Turnover füh-

ren.

Wertvolle Zusatzinformationen kann dagegen eine Betrachtung der Dynamik auf dem

Art-Niveau liefern. Wenn sich zeigen sollte, daß ein großer Teil des Artenspektrums

einer ausgeprägten Populationsdynamik unterliegt und die jeweilige Besiedelung eines

Habitats stark von einem Fließgleichgewicht abhängt, das durch ständige "Trittstein-

Sprünge", d.h. durch ein Neu-Aufsuchen, Bodenständig-Werden (mehr oder weniger

kurzzeitig) und bei Störungen durch ein Wieder-Verschwinden seitens der Arten ent-

steht, so sollte dies ein weiterer Hinweis darauf sein, daß der apparente Turnover eine

tatsächlich stattfindende Dynamik wiederspiegelt. Der apparente Artenumsatz mag nu-

merisch nicht ganz exakt sein, er stellt aber vielleicht eine brauchbare Annäherung

der Verhältnisse dar.

Hierzu sollen im II. Teil vor allem die Dispersionsaktivitäten der Arten in Zusammen-

hang mit Verbreitungsstrategien betrachtet werden.

II. EXPERIMENTELLER TEIL: DISPERSIONSVERHALTEN UND.

TRIVIAL MOVEMENT

8. DIE EXPERIMENTE

8.1. VORSTELLUNG DER MARKIERUNGSEXPERIMENTE

8.1.1. Allgemeines

Unter "Dispersionsdynamik" versteht man nach SCHWERDTFEGER (1978) "den durch

Ortswechsel ihrer Glieder sowie durch Dichteänderung bewirkten Wandel in der örtli-

chen Verteilung der Population".

Die Auftrennung der Nachtfalter-Flugaktivitäten in Dispersions- und Migrationsverhal-

ten einerseits und "trivial flights” andererseits soll im Sinne JOHNSONs (1969) ver-

standen werden (siehe Kapitel 12).

Zur Erforschung dieses Fluggeschehens sind drei Wege zu unterscheiden (REINHARDT

& DROBNIEWSKI, 1979):

- die Erfassung offensichtlich zugeflogener Arten,

- das Markieren und Freilassen von Faltern und

- zusätzliche experimentelle Arbeiten.

Insgesamt wurden im Untersuchungsgebiet über 23.800 Falter aus 131 Arten markiert.

Die Artauswahl richtete sich nach folgenden Kriterien:

- Abdeckung der Bandbreite verschiedener Taxa (Familien, Unterfamilien)

- Unterschiedliche Ökotypen

- Unterschiedliche Flugzeiten (auch Arten, die unter extremen Bedingungen fliegen)

- Unterschiedliche Körpergrößen

- Unterschiedliche Abundanz (insgesamt gesehen, wurden jedoch häufigere Arten bevor-

zugt)

- Repräsentanz von uni- und bivoltinen Arten

- Wanderfalter als "Nullprobe" geringer Ortstreue

8.1.2. "Fern"-Wiederfänge

Durch das Betreiben eines ganzen Standortnetzes (siehe 2.2., Abbildung 1) sollte gete-

stet werden, ob auf Distanzen von 400 m (SiS>SiM) bis 3,25 km (FW>WaN) Disper-

sionsaktivitäten markierter Nachtfalter direkt durch Wiederfang an einer anderen Stelle

nachgewiesen werden können.

Zur Technik der Markierungen siehe Kapitel 3.4..

8.1.3. Verringerte Fallendistanzen

Nachdem 1987 über die oben angeführten Distanzen bei 7617 markierten Faltern nur 2

Ortswechsler nachgewiesen werden konnten, wurde im Wasserwerk ein Parallelfang-

Versuch mit Fallenentfernungen gestartet, die ungefähr eine Zehnerpotenz unter denen

des Jahres 1987 lagen (siehe 2.2., Abbildung 2). Der Fang erfolgte mit einer Frequenz

von ca. 2-3 Tagen über das ganze Jahr.

8.1.4. Versetzexperiment

Um abschätzen zu können, welchen Anteil am Anflugverhalten die direkte Lichtattraktion

hat, und inwieweit Dispersionsaktivitäten eine Rolle spielen, wurde 1988 im Waldstreifen

hinter dem Garten (Fangplatzz WaS) ein Versetzexperiment durchgeführt: Die

Wiederfang-Wahrscheinlichkeiten von Tieren, die an der Falle freigelassen wurden,

geben im Vergleich mit solchen, die in 30 m/60 m/90 m und 120 m Entfernung freige-

lassen wurden, Aufschlüsse über die Anteile solcher Einflüsse.

Es wurde hierzu an WaS täglich geleuchtet, um "Verlustkurven" besser dokumen-

tieren zu können. Aus diesem Grund wurden auch die Wiederfänge in fast allen Fällen

bei "O0 m" (=Radius von ca. 7-8 m um die Falle herum, siehe 3.3.) ausgesetzt. Für das

Versetzexperiment wurden einige genügend häufige Arten ausgewählt.

Im Torfeinfang erfolgten 1988 einige Versuche an der Spannerart Calospilos sylvata,

um eine relativ ortsfeste Population mit hoher Populationsdichte auf deren Antworten

auf die Versuchsbedingungen zu testen.

Die Versetzdistanzen wurden im Garten laufend von Nacht zu Nacht vergrößert, so

daß in der jeweiligen 6. Nacht wieder bei 0 m begonnen werden konnte. In anderen

Fällen (z.B. bei Alcis repandata und Peribatodes rhomboidaria) wurde stets die Hälfte

der Exemplare bei 0 m und die andere Hälfte bei der jeweiligen Versetzdistanz frei-

gelassen, um durch eine Bereinigung mit Hilfe des mittleren "Null-Wertes” die Ergeb-

nisse aus den verschiedenen Entfernungen unabhängiger gegen Störeinflüsse zu machen

(z.B. Witterung). Die errechneten bereinigten Ergebnisse entsprachen jedoch in allen

Fällen denen, die ohne diese Maßnahme zustandegekommen wären.

Abb. 16: Garten des Verfassers mit den Fangplätzen SiN (A) und WaS (B); markierte

Tiere wurden in 30m (1), 60 m (2), 90 m (3) und 120 m (4) Entfernung freigelassen;

Flächen mit überwiegendem Waldcharakter (Kronenschluß) sind mit Raster gekenn-

zeichnet.

Garden of the author (SiN=A; WaS=B): marked specimens were released at distan-

ces of 30 m (1), 60 m (2), 90 m (3) and 120 m (4); reticulate: wood character.

Abb. 16b: Nähere Umgebung des Fangplatzes Was.

Im Grunde genommen stellen auch die 30 m südlich des Standorts WaS an SiN mar-

kierten Falter "versetzte" Exemplare dar, deren Anflug an WaS in den für SiN fang-

freien Nächten getestet wurde. Durch Vergleich mit den im Waldstreifen bei 30 m

freigelassenen Tieren ergeben sich Hinweise auf die Einflüsse der schlechteren Ein-

sehbarkeit der Lichtquelle sowie der vielleicht als Barriere wirkenden Häuser- und

Garagenzeile (2-10 m hoch).

Die Einsehbarkeit der Lichtquelle war bei keiner der vier Freilaß-Punkte gegeben, bei

30 m war diese jedoch nur durch einen Busch verstellt.

8.1.5. Rückschlüsse aus Ortswiederfängen

Die mit einigen Problemen behafteten Interpretationen von Ortswiederfängen bei Nacht-

faltern können bei Berücksichtigung der Fehlerquellen (siehe 8.5.) in vielen Fällen

Aufschlüsse über populationsdynamische Vorgänge liefern.

Nach BETTMANN (1986) benötigt man eine Stichprobe von mindestens 100 markierten

Faltern, um einigermaßen repräsentative Ergebnisse zu erhalten. Dies war im Untersu-

chungsgebiet bei 50 Arten der Fall.

8.2. "FERN-" WIEDERFÄNGE

8.2.1. Übersicht

Tab. 16: Wiederfänge über Mindestdistanzen (Luftlinie) ab 1 km 1987 und 1988 im

Untersuchungsgebiet.

Recaptures over distances of at least 1 km 1987 and 1988 in the study area.

JAHR ART GESCHLECHT STRECKE DISTANZ (km) INTERVALL (Tage)

year species Sex route distance (km) interval (days)

1987 Scotia clavis [6] HW>SIiS 1,0 3

Alcis repandata 6) SiIS>HW 1,0 3

1988 Scotia clavis 6) WaN>HO 1,9 2

Scotia clavis [oj HM>HO 1,0 2

Ochropleura plecta cd HO>HM 1,0 2

Noctua pronuba [6] WaS>WNo 1,4 2

Noctua pronuba q WaS> WNw 1,3 2

Amathes triangulum WaS> WNw 1,3 4

Rusina ferruginea q HM>HO 1,0 6

Meristis trigrammica SIN>WaN 1,4 3

Hoplodrina alsines cd HW>WNo 3,25 5

Peribatodes rhomboidaria WNo>HO 1,9 1

Alle Angaben wurden genanen Prüfungen unterzogen, auch die beiden Wiederfänge nach

1 Tag können als gesichert gelten. Eine Verschleppung mit der Falle durch ungenügen-

de Achtsamkeit beim Absammeln von markierten Faltern, die dort Unterschlupf such-

ten, konnte in den genannten Fällen durch Markierung der Fallen ausgeschlossen wer-

den: Die Falle des Erstfangs war am Wiederfundort nicht eingesetzt worden. Die

Auswertung erfolgte auch in genügendem Abstand zum Fahrzeug, und eine Tasche mit

den nötigen Utensilien wurde nur zur Auswertung, nicht jedoch beim Aufstellen der

Falle mitgenommen; auch dadurch können Fehlerquellen bezüglich einer Verschleppung

von Faltern ausgeschlossen werden.

In vier Fällen blieben diesbezüglich jedoch Zweifel übrig, die betroffenen Wiederfänge

wurden daher nicht berücksichtigt.

8.2.2. Diskussion

Bei der Betrachtung der 13 Wiederfänge, die über Distanzen von mindestens 1 km er-

folgten, zeigt sich, daß es sich hier vor allem um Eulenfalter (Noctuidae) handelt: Sie

stellen 64,1 % aller markierten Individuen, aber 84,6 % der Wiederfänge über größere

Distanzen. Auch die beiden Spanner (Geometridae) sind große Arten mit relativ star-

kem Thorax. Die Dispersionsaktivitäten über Distanzen von 1-4 km scheinen also in

Korrelation zur mechanischen Flugfähigkeit zu stehen, was jedoch nicht umgekehrt

bedeuten muß, daß für jede flugkräftige Art solche Entfernungen zum normalen

Dispersal gehören.

SCOTT (1975) stellte in seiner Untersuchung von Tagfalter-Flugaktivitäten fest, daß

die meisten Arten die Distanz von mehreren Kilometern nicht bewältigen. Der weiteste

nachgewiesene Flug fand über eine Strecke von 2940 m statt. Die Aussagen entspre-

chen dem im Untersuchungsgebiet für die Nachtfalter Gefundenen.

Bei allen Wiederfängen handelte es sich um Männchen. Die Weibchen-Rate bei den

markierten Faltern lag im Bereich von durchschnittlich 27 %. Es handelt sich also auch

hier um eine Überrepräsentierung.

Ubiquisten (r-Strategen) sind in der oben aufgeführten Liste verstärkt vertreten: Fallen

9,9 % der im Untersuchungsgebiet festgestellten Gesamtartenzahl in diese Kategorie, so

waren 56 % der Arten, bei denen größere Flugdistanzen beobachtet wurden, Ubiquisten.

Die Arten des mesophilen Graslandes sind mit 22 % vertreten (15,8 % im Gesamt-Ar-

tenspektrum). Stark unterrepräsentiert sind dagegen die Arten der Wälder, der Wald-

ränder und der Gebüschformationen mit 22 % statt 57,9 %. Diese können also als we-

niger expansiv gelten.

Der Befund, daß die als Wanderfalter bekannte Noctua pronuba in relativ kurzen Zeit-

spannen jeweils von Süd nach Nord flog, ist vermutlich auf ein solches Migrationsver-

halten zurückzuführen.

1987 ereigneten sich auffallend wenig Fern-Wiederfänge, nämlich 0,026 % der markier-

ten Falter im Gegensatz zu 0,070 % 1988. Vielleicht spielte hier die etwas feuchtere

Witterung (v.a. von April bis Juni) im Jahr 1987 eine Rolle.

Geländestrukturen spielen zumindest in Einzelfällen eine Rolle bei der Ausbreitung:

Beide Ortswechsel 1987 erfolgten entlang der auwaldartigen Kanalbegleitflora des

Würmkanals. Langgestreckte, schmale Strukturen sind auch nach WATT et al. (1977)

ein begünstigender Faktor für Dispersionsaktivitäten. Da auch die 1988 von Peribato-

des rhomboidaria zurückgelegte Strecke entlang der Luftlinie fast ausschließlich aus

Waldrandstrukturen besteht, sind beide Geometriden-Wiederfänge über größere Entfer-

nungen hinweg potentiell durch Geländestrukturen begünstigt worden.

Bei den Fern-Wiederfängen ist kein signifikanter Zusammenhang zwischen Entfernung

und Zeitspanne erkennbar. Die betreffenden Arten scheinen also nicht die Summe vie-

ler Tagesdistanzen zur Bewältigung der beobachteten Entfernung zu benötigen.

Aus den Tatsachen, daß bei einem derartigen Umfang der Stichprobe () 23.800) Wie-

derfänge über Distanzen von 1-4 km nicht einmal im Promillebereich und ausschließ-

lich von Männchen nachzuweisen waren, ist abzuschätzen, daß hypothetisch neugeschaf-

fene Biotope in einer entsprechenden Entfernung zu stabilen Nachtfalterpopulationen

des gleichen Lebensraumtyps nur langsam und unvollständig besiedelt werden können.

Eine Vernetzung von Biotopen sollte daher mit Abständen von unter 1 km erfolgen.

Diese Befunde stehen im Einklang mit den von SCOTT (1972) und WATT et al. (1977)

publizierten Ergebnissen, die bei nordamerikanischen Tagfaltern schon in einer Entfer-

nung von wenigen 100 m nur wenig Austauschereignisse bzw. eine weitgehende Isolie-

rung von Populationen feststellten.

8.3. VERRINGERTE FALLENDISTANZEN

8.3.1. Übersicht über das Material

Zur Beschreibung des Experiments und des Geländes siehe 8.1.2. und 2.2..

Im Jahr 1988 wurden im Wasserwerksgelände Oberschleißheim 5335 der 11.392 gefan-

genen Nachtfalter (Macroheterocera) markiert. Es erfolgten 159 Wiederfänge. Zu

Vergleichszwecken können auch die 1280 im Jahr 1987 markierten Nachtfalter (17

Rückfänge) herangezogen werden.

Aus Abb. 17 werden die 154 im Wasserwerk 1988 nachgewiesenen ÖOrtswiederfänge

bzw. Dispersionsaktivitäten ersichtlich. Fünf Rückfänge beziehen sich auf zugeflogene

Stücke anderer Fangplätze (siehe 8.2.).

w I Ss

Abb. 17: Ortswiederfänge und Ortswechsler 1988 im Wasserwerk (siehe 2.2.).

Recaptures at the same site and changes of site 1988 in the pump station (see 2.2.).

Es zeigt sich ein deutlicher Abfall der Austauschraten um ca. die Hälfte schon bei ei-

ner Verdoppelung der Distanz von 50 m (WaN/WNo) auf 100 m (WaN/WNw). Die

Distanz scheint jedoch nicht der einzige die Verbreitung begrenzende Faktor zu sein,

da beispielsweise auf der Strecke WNo-WNw (120 m) ein größerer Individuenaustausch

stattfand als zwischen WaN und WNw. Hier spielen offensichtlich Geländestrukturen

eine Rolle. Hin- und Rückflug zeigen fast identische Werte: Es liegen, insgesamt ge-

sehen, keine konstanten, gerichteten Prozesse vor.

Die durchschnittliche Wiederfang-Quote lag mit 2,9 % deutlich über dem 1987 an WaN

festgestellten Wert von 1,3 %. 1988 wurden durch die bessere Abdeckung des Geländes

(3 Fallen) auch die in der Nähe verweilenden Falter miterfaßt, die bei einem Einzel-

standort außerhalb der Fallenreichweite geblieben wären.

Dementsprechend verteilten sich auch die Anteile der 1988 festgestellten Wiederfang-

Quoten: An WaN entfielen von den 3,0 % beobachteten Wiederfängen 1,8 % auf orts-

treue Falter, der Rest stammte von den Nachbarstandorten.

Andersherum betrachtet konnte das Wiederfangergebnis durch die Erhöhung der Fallen-

zahl von 1,8 % um 1,2 %, die an den anderen Fallen wiedergefunden wurden, auf eben-

falls 3,0 % gesteigert werden.

Am Standort WNw war die geringste Stabilität nachzuweisen, hier lag die Ortswieder-

fang-Quote mit 0,7 % deutlich unter derjenigen der beiden anderen Fangplätze (WaN

und WNo jeweils 1,8 %).

Diese für alle markierten Arten etwas pauschale Skizzierung der Verhältnisse wird in

den Tabellen 17 und 18 etwas weiter differenziert: In die Liste wurden die Arten mit

aufgenommen, von denen mindestens 100 Individuen markiert wurden und/oder zumin-

dest 5 Wiederfänge vorlagen.

Tab. 17: Die wichtigsten Wiederfänge an den drei Standorten im Wasserwerk 1988,

geordnet nach der Wiederfang-Quote.

The most important recaptures of the three capture-sites at the pump station 1988,

according to their recapture-rate.

ART MARKIERT WIEDERFÄNGE WIEDERFANG-

QUOTE (%)

species marked recaptures percentage of

recaptures

Chiasmia clatrata (I+1I) 36 8 239

Diacrisia sannio 50 6 12,0

Cerastis rubricosa 83 7. 8,4

Mythimna impura 200 15 743

Spilosoma menthastri 125 9 1,2

Scotopteryx chenopodiata 326 19 5,8

Orthosia gothica 208 10 4,8

Amathes ditrapezium 124 6 4,8

Alcis repandata 294 9 3.1

Amathes sexstrigata 332 9 27

Amathes triangulum 184 5 27

Amathes xanthographa 106 2 |)

Apamea anceps 110 2 1,8

Rusina ferruginea 121 2 27

Eilema depressa 336 3 0,9

Ochropleura plecta 380 2 0,5

Amathes c-nigrum 361 1 0,2

Noctua pronuba 185 = 0,0

Es ergibt sich eine mehr oder weniger kontinuierliche Verteilung, deren Gradient ab-

gesehen von einigen Ausnahmen in etwa dem r-K-Kontinuum entsprechen dürfte. So

finden sich in der oberen Hälfte der Tabelle signifikant mehr Spanner (Geometridae)

und Spinner (Bombyces), während in der unteren Hälfte mehr Eulenfalter (Noctuidae)

vertreten sind. Aufgrund unterschiedlicher Raupenfutterpflanzen sind - wie schon im

ersten Teil angeklungen ist - bei den Noctuiden mehr r-Strategen zu finden als in den

anderen Gruppen. Dementsprechend befinden sich auch die in der Literatur als Wan-

derfalter bezeichneten Noctua pronuba und Amathes c-nigrum am Ende der Tabelle.

Zur Problematik der Interpretation von Ortswiederfängen sind einige Aspekte, die in

8.5. näher beleuchtet sind, zu beachten. In der Nähe herumvagabundierende Individu-

en dürften jedoch zu einem Großteil durch die Erhöhung der Fallenzahl erfaßt worden

sein.

Tab. 18: Die wichtigsten Wiederfänge im Wasserwerk 1988, geordnet nach der mittleren

Verweildauer.

The most important recaptures at the pump station 1988, according to their mean of

residence time.

ART MITTLERE MAXIMAL BEOBACH- MITTLERE

VERWEILDAUER TETE VERWEILDAUER FLUGDISTANZ

(Tage) (Tage) (Meter)

species mean of residence observed maximum of mean of flight

time (days) residence time (days) distance (m

Cerastis rubricosa 6,33 10 63

Diacrisia sannio 5,33 16 28

Orthosia gothica 022. 13 59

Chiasmia clatrata 5,00 11 40

Alcis repandata 4,14 7 (3 Exemplare) 13

Mythimna impura 3,92 9 (HO 1987 27 Tage) 37

Scotopteryx chenopodiata 3,83 15 26

Spilosoma menthastri 3,75 8 24

Amathes triangulum 3,20 7 (sonst 3 Tage) 54

Amathes sexstrigata 2,63 5 39

Amathes ditrapezium 2,33 3 33

Stichproben zu klein

Apamea anceps 3,50 4 25

Eilema depressa 2,66 3 17

Rusina ferruginea 2,50 3 120

Amathes xanthographa 2,00 2 60

Ochropleura plecta 2,00 2 50

Amathes c-nigrum 2,00 2 0

Noctua pronuba er > =

Die Frühlingsarten (O. gothica, C. rubricosa) fallen durch verlängerte Verweildauern

auf, ein vermutlich witterungsbedingtes Phänomen. Die eigentiichen Flugnächte liegen

hier zwischen Phasen der Unbeweglichkeit bei nicht ganz optimalen Flugbedingungen.

Bei den Arten des "Sommerblocks” erfolgen dagegen offenbar Ortsveränderungen auch

unter suboptimalen Voraussetzungen.

Unter der Annahme einer vergleichbaren Mortalität sind "residence time” und "disper-

sal" zwei negativ korrelierte Parameter (WATT et al., 1977).

Ein signifikanter Anstieg der mittleren Flugdistanzen von oben nach unten (zunehmend

starke r-Strategen) in der Tabelle ist nicht zu belegen, da die Flugstrecken der r-

Strategen in anderen Größenordnungsbereichen liegen. Bei den beobachteten Flugdistan-

zen handelt es sich vermutlich um trivial movement; auch die am meisten ortsgebun-

denen der untersuchten Arten können Strecken von ca. 100 m bewältigen.

Nach systematischen Gruppen aufgeschlüsselt ergibt sich für die Eulenfalter (Noctui-

dae) eine mittlere Flugdistanz von 51 m. Bei den Spannern (Geometridae) sind es wie

bei den Spinnern (Bombyces) 26 m, wenn man nur die Arten mit über 5 Wiederfängen

berücksichtigt. Die geringen Artenzahlen bei den Spannern (3) und Spinnern (2) stellen

für das Ergebnis jedoch noch Unsicherheitsfaktoren dar. Wenn man die Anzahl der

Wiederfänge (51) betrachtet, so stellen diese beiden Gruppen 50 % des Materials, was

genügen sollte.

8.3.2. Weitere Ergebnisse, aufgezeigt an ausgewählten Arten

Diacrisia sannio:

Bei Streifzügen durch magere Wiesen entdeckt man oft aufgeschreckte Exemplare

dieser Art. Diese fliegen dann normalerweise 5-10 m, höchstens 20 m weit und las-

sen sich dann wieder im Gras nieder.

An WNw war kein Ortswiederfang festzustellen, an den anderen Standorten je zwei.

Der westliche Fangplatz ist auch durch die geringeren Ausbeuten nur als suboptima-

ler Lebensraum für D. sannio charakterisiert. An ungünstigeren Stellen finden ver-

mehrt Austauschprozesse statt.

Ochropleura plecta:

Bei beiden Wiederfängen handelte es sich um 99, die vielleicht beim Kopulations-

oder Eiablagegeschehen eine größere Ortsfestigkeit zeigen als die dd.

Amathes ditrapezium:

Ähnlich dem Bärenspinner Diacrisia sannioe An WNw schlägt sich die vermutlich

durch die Habitatrand-Lage bedingte höhere Dynamik in den fehlenden Ortswieder-

fängen nieder. 5 der 6 Wiederfänge belegen ein Verbleiben in einem 50 m-Radius

des Verbreitungszentrums (WaN/WNOo), allerdings bei kurzen Verweildauern.

Amathes sexstrigata:

Ähnlich dem Bärenspinner Diacrisia sannio fehlten an WNw Ortswiederfänge, die

Ausbeuten waren hier geringer. Dies mag mit erhöhten Austauschprozessen und

Dispersionsaktivitäten an den Rändern von (Teil-)Populationen erklärt werden.

8 der 9 Wiederfänge, nämlich die Ortswiederfänge an WaN und WNo sowie die

Wechsler zwischen den beiden Standorten, zeugten von einem Verbleiben in einem

50 m-Bereich innerhalb des Verbreitungszentrums.

Cerastis rubricosa:

Trotz der hohen Wiederfang-Quoten und Verweildauern zeigt sich innerhalb des

Wasserwerkgeländes eine Dynamik auf relativ hohem Niveau: Es überwiegen hier die

Ortswechsler, und bei einem d' wurde sogar ein Ortswechsel von WaN> WNw> WNo

(220 m) über 8 Tage hinweg mitverfolgt.

Orthosia gothica:

Die hohe Wiederfang-Quote und Verweildauer sollte nicht vorschnell zu einer Cha-

rakterisierung dieser Art als K-Strategen führen. Auffallend ist schon die Verteilung

der Wiederfänge (siehe auch 9.2.): Ähnlich der vorhergehenden Art überwiegen auch

hier die Ortswechsler, Ortswiederfänge erfolgten nur an WaN (3). Bei einigermaßen

vergleichbaren Zahlen der markierten Individuen steht den 8 Wiederfängen an WaN

kein einziger an WNw gegenüber; an letztgenanntem Standort scheint die Dynamik

am höchsten zu liegen.

Da WNw auch am weitesten von der nächstgelegenen Weide, der wohl wichtigsten

Nektarquelle von O. gothica, entfernt liegt, ist hierin ein Zusammenhang zu vermu-

ten: An Standorten mit blühenden Weiden kommt es zu einer starken Reduktion des

trivial movement.

Bei einem J konnte ein Ortswechsel von WNw>WNo>WaN (170 m, 4 Tage) mit-

verfolgt werden.

Mythimna impura:

Es ist das gleiche Phänomen festzustellen wie bei Diacrisia sannio, Amathes ditrape-

zium und A. sexstrigata. Das Verbreitungszentrum ist hier WNo, dort erfolgten wie

auch an WaN drei Wiederfänge am selben Ort; an WNw fehlte ein solcher.

Im Gegensatz zu den drei oben genannten Bewohnern von tendenziell trockeneren

Wiesen ist M. impura eher an Schilf und Seggen gebunden. Dies ist eine denkbare

Erklärung für den Sachverhalt, daB in das dynamische Geschehen der Fangplatz

WNw stärker miteinbezogen wurde: Dies war in 6 der 15 Wiederfänge der Fall.

Zwei dd flogen von WNo nach WNw und wieder zurück (240 m). Dies ereignete

sich in einer Zeitspanne von 4 bzw. 7 Tagen.

M. impura ist ein schönes Beispiel dafür, daß offensichtlich ortstreuere Arten in Be-

reichen von ca. 100 m durchaus eine ausgeprägte Dynamik im Sinne eines trivial

movement besitzen.

Scotopteryx chenopodiata:

Diese Art reagierte nun etwas verschieden von den bisher besprochenen Mustern: An

allen Standorten erfolgten proportional ungefähr gleiche Anteile an Ortswiederfängen.

Diese lagen insgesamt mit 13 von 19 auf einem hohen Niveau. Man kann also auch

bei so klein gewählten Distanzen von geringen Austauschraten ausgehen!

Die 99 scheinen bei einer mittleren Flugdistanz von 17 m ortsfester zu sein als die

dc (30 m). Die mittlere Verweildauer der 99 liegt jedoch genau im Schnitt.

Chiasmia clathrata:

Wie Mythimna impura: Das Verbreitungszentrum liegt an WNo (tagsüber wie in der

Nacht!). 6 der 8 Wiederfänge spielten sich an/zwischen WaN und WNo ab. Am

westlichen Standort erfolgte kein Ortswiederfang, an dieser Habitatrand-Lage ist also

eine höhere Dynamik, verursacht durch geringere Bodenständigkeit zu beobachten.

Der hohe Anteil der Ortswiederfänge an den anderen beiden Fangplätzen (5 von 7)

kennzeichnet C. clathrata ebenfalls als vergleichsweise ortstreue Art.

Die Wiederfang-Quote liegt in der ersten Generation mit 26,3 % deutlich über den

17,6 % der zweiten Generation. Da ein J sehr früh in der 2. Generation zweimal

wiedergefangen wurde, kann man ab Mitte Juli von einer spürbar erhöhten Disper-

sionsaktivität ausgehen.

Alcis repandata:

Bei dieser Art wurde nur ein Ortswechsler bei 9 Wiederfängen festgestellt! Diese

eigentlich auf hohe Ortstreue hinweisende Beobachtung steht etwas im Gegensatz zu

den oft nur kurzen Verweildauern von 2 Tagen. Berücksichtigt man auch die niedri-

geren Fang- und Wiederfangzahlen an WaN, so ist zu vermuten, daß viele Exempla-

re von den Waldrändern herbeigeflogen waren und an WNo und WNw, die diesen

Rändern näher liegen, vielleicht ein mehrfaches Hin- und Herfliegen erfolgte.

Die 99 sind im Wiederfang (11 %) gegenüber den Erstfängen (34 %) unterrepräsentiert.

8.4. VERSETZEXPERIMENT

8.4.1. Übersicht über das Material

Die Zahl der 1988 im Garten (WaS) im Rahmen des Versetzexperiments (ausgewählte

Arten) markierten Individuen betrug 2465, es wurden 475 Wiederfänge verzeichnet.

Im Torfeinfang (We) wurden 1988 629 Spanner der Art Calospilos sylvata markiert,

von denen 46 rückgefangen werden konnten.

Da - wie bereits erwähnt - auch Ortswechsler zwischen SiN und WaS (30 m) als

"versetzte" Nachtfalter interpretiert werden können (wenn nicht eine präferenzielle

Rückkehr vorliegt, vergleiche KELLER, MATTONI & SEIGER, 1966), ist eine Betrach-

tung der zwischen diesen beiden Fangplätzen stattfindenden Austauschraten interessant:

- 1987 waren die Ortswiederfang-Quoten an SiN und WaS (bei fangfreien Nächten da-

zwischen) mit 1,5 % bzw. 1,2 % ungefähr gleich. In 30 m Entfernung waren bei einer

Richtung SiN> WaS nach einem Intervall von 1 Tag weitere 0,4 %, insgesamt weitere

1,0 % wiederzufangen. Umgekehrt betrugen die Werte 0,6 % und 1,0 %.

- 1988 zeigte sich an SiN bei gleicher Methode eine entsprechende Ortswiederfang-

Quote von 1,2 %, der nach WaS abwandernde Prozentsatz fiel mit 1,5 % etwas höher

aus, da dort über einen Großteil der Flugzeit hinweg täglich geleuchtet wurde und

die Erfassung somit vollständiger erfolgte. Durch diesen veränderten Rhythmus stie-

gen auch die Ortswiederfänge an WaS drastisch auf 21,5 %, während der Anteil der

von WaS nach SiN fliegenden Tiere mit 0,9 % ungefähr gleich blieb.

Die Berechnungen wurden durch Abzug der im eigentlichen Versetzexperiment ge-

sammelten Daten bereinigt.

Bei einem Vergleich mit dem Wasserwerk zeigt sich, daß der Individuen-Austausch

zwischen den Fallen beispielsweise im Vergleich mit dem Fallenpaar WaN/WNo bei

einer etwas kleineren Distanz ungefähr proportional höher ausfiel: Dividiert man die

Summe der ausgetauschten Individuen durch die Summe der an beiden Standorten

markierten Falter ergeben sich für den Garten (1987) 1.0 %, für das Wasserwerk

(1988, WaN/WNo) 0,8 % ausgetauschter Nachtfalter. Es handelt sich hierbei wohl-

gemerkt um keine Absolutzahlen, die realen Austauschraten liegen sicherlich höher.

Die Barriere der Häuserzeile scheint demnach zusammen mit den anderen Störein-

flüssen (Straßenbeleuchtung u.s.w.) kein Hindernis darzustellen, das sich wesentlich

von der Geländestruktur im Wasserwerk, nämlich einer relativ naturnahen halbver-

buschten Fläche unterscheidet.

Diese Aussage muß aber, da sie über eine gemeinsame Betrachtung aller markierten

Arten getroffen wurde, nicht für alle Arten gleichermaßen zutreffen.

8.4.2. Ergebnisse

Scotia clavis:

Tab. 19-21: Ausbeute, Markierungen und Wiederfänge im Garten 1988 bei Scotia clavis.

Numbers of individuals, marked specimens and recaptures of Scotia clavis in the

garden 1988.

Ausbeute (Individuen) Markierungen Wiederfänge/Mehrfachwiederfänge

SIiN WaS zZ SIiN WasS 2 W.L.1 2.2.0832.4: 51.6.7228 =

dc 25 158 183 23 119 142 26: x bed == NZ

99 20.202022 D1822 20 EEE ET a)

_ 27 178 205 25 137 162 28 ben Prise 40

Par. 23] 62

In Tabelle 19 ist eine Erhöhung der Werte durch die hier mitgezählten Wiederfänge zu

veranschlagen, eine Bereinigung der Angaben wird durch Tab 2i möglich.

Unter "Par." ist in der Tabelle 19 das aus den Parallelfang-Ergebnissen stammende di-

rekt vergleichbare Individuenverhältnis aufgeführt. Gelegentlich auftretende Differenzen

zu den in der Artenliste (4. Kapitel) angegebenen Zahlen entstehen durch entkommene

Exemplare, deren Artbestimmung erfolgte, nicht aber die Geschlechtsbestimmung.

Die Weibchen-Rate ist an WaS fast doppelt so hoch wie an SiN. Im Wiederfang sind

die 99 mit 5 % etwas unterrepräsentiert.

Tab. 22: Zwischen Fang und Wiederfang festgestellte Intervalle bei Scotia clavis 1988

im Garten.

Intervals (in days) between two catches of Scotia clavis 1988 in the garden.

lee DEE Re So an as

Wiederfänge SSCRE SE a ee EN OBEN 40

Das Intervall zwischen Fang und Wiederfang beträgt im Mittel 1,13 Tage.

Die Tabelle 22 offenbart bereits eine hohe Dynamik von Scotia clavis an diesem Stand-

ort, die kurzen Verweildauern, wie sie aus den Abbildungen 18 und 19 ersichtlich

sind, können nicht allein auf Mortalität zurückzuführen sein.

4 Aufsummierte Verweilzeiten (Annahme: Stän-

dige Anwesenheit, ‚nur nicht immer gefangen)

Wiederfä h in Relati

20 = ie ee lese x Tagen in Relation

1 2 3 4 5 (Tage)

Abb. 18: Apparente Verweilzeiten von Abb. 19: Aufsummierte Verweilzeiten und real

Scotia clavis 1988 im Garten. nach x Tagen beobachtete Wiederfänge.

Apparent residence times of Scotia Accumulated residence times and really

clavis 1988 in the garden. after the time x recaptured specimens.

Die durchschnittliche Verweilzeit, errechnet aus den Wiederfängen, betrug nur 1,57

Tage. Nicht berücksichtigt bleiben die Individuen, von denen keine Wiederfänge vorlie-

gen. Es handelt sich daher um einen relativen Wert.

Auch die Verlustkurven in Abb. 19 zeugen von einem schnellen Verschwinden aus dem

Einzugsbereich der Falle.

Tab. 23: Reaktion von Scotia clavis auf ein Versetzen in verschiedene Entfernungen.

Recaptures of specimens of Scotia clavis, released at different distances from the

light source.

0 m 30m 60 m 90 m 120 m

freigelassen* 95 22 20 25 12 *Summe aus markierten Faltern

released und einigen freigelassenen Wie-

Wiederfänge 25 4 3 3 1 derfängen.

recaptures

y2 26 18 15 12 8

Das Versetzexperiment zeigt einen kontinuierlichen Abfall der Wiederfang-Wahr-

scheinlichkeit mit zunehmender Versetzdistanz. Dies ist vermutlich auf ein zickzackar-

tiges, mehr oder weniger ungerichtetes Umherschweifen in der weiteren Umgebung zu-

rückzuführen (siehe Peribatodes rhomboidaria). Ausgeprägte und zielgerichtete Dynami-

ken würden zu einem stärkeren Abfall der Werte bei 90 und 120 m führen (siehe Noc-

tua pronuba).

Eine Verlängerung der Rückkehrdauer mit zunehmender Versetzdistanz war nicht zu

beobachten. Distanzen bis 120 m scheinen also im Bereich des trivial movement einer

Nacht zu liegen.

Scotia exclamationis:

Die Stichprobengröße bei Scotia exclamationis 1988 im Garten ist relativ klein (98

markierte, 14 wiedergefangene Falter). Der starke Abfall von 11 Erst- auf 3 Zweit-

wiederfänge entspricht ungefähr den Verhältnissen bei $. clavis.

Die Weibchen-Rate liegt auch hier an WaS deutlich höher als an SiN. Im Wiederfang

sind die 99 jedoch mit 64 % deutlich überrepräsentiert! Im Gegensatz zu S. clavis

scheinen bei S. exclamationis die dd‘ mobiler zu sein. Die 99 stellen auch alle beo-

bachteten Zweitwiederfänge.

Tab. 24: Zwischen Fang und Wiederfang festgestellte Intervalle bei Scotia exclamatio-

nis 1988 im Garten.

Intervals (in days) between two catches of Scotia exclamationis 1988 in the garden.

Intervall: (Tage) ne 1a»231.1 311.3 65.5 de Irni1O, AL =

Wiederfänge 12.7 1 = = _ = — 14

Das Intervall zwischen Fang und Wiederfang beträgt im Mittel 1,43 Tage.

Die mittlere Verweildauer liegt bei 1,73 Tagen, ein Unterschied zwischen den Ge-

schlechtern ist nicht ersichtlich.

Die Wiederfang-Quote der an der Falle freigelassenen Tiere entspricht mit 23 % in

etwa dem Wert von Scotia clavis (n=53). Von den in den verschiedenen Entfernungen

ausgesetzten Faltern (n=26) konnte nur ein d aus 60 m zurückgefangen werden. Die

Stichprobengröße ist hierbei noch zu klein, um nähere Rückschlüsse zu erlauben.

Noctua pronuba:

Als bekannter Wanderfalter sollte Noctua pronuba - wie es bei einer hochmobilen Art

zu erwarten ist - mit kurzen Verweildauern und sehr niedrigen Rückfang-Quoten bei

einem Versetzen um 90 bzw. 120 m reagieren. Letzteres deshalb, weil hier bei gerad-

linigen starken Flugaktivitäten Individuen, die nicht die Richtung der Lichtquelle ein-

schlagen (Winkel von ca. 330-340° = 92-94 % unter der Voraussetzung einer freien

Beweglichkeit ohne Barrieren) auf Nimmerwiedersehen verschwinden.

Bei einem Herumvagabundieren mit häufigeren Richtungswechseln kommt es dagegen zu

einer erhöhten Wiederfangwahrscheinlichkeit auch bei größeren Versetzdistanzen. Dies

ist jedoch bei manchen Arten auch durch eine "Kanalisierung” des Fluges durch die Bio-

topgrenzen im Waldstreifen denkbar. Die Unterschiede zu den Biotopstrukturen der

umliegenden Gärten sind jedoch nicht besonders gravierend.

Ein Vergleich mit einem ähnlichen Experiment an Spodoptera littoralis in Israel

(PLAUT, 1971), ebenfalls einem Wanderfalter, ist hier möglich.

Tab. 25-27: Ausbeute, Markierungen und Wiederfänge im Garten 1988 bei Noctua pronuba.

Numbers of individuals, marked specimens and recaptures of Noctua pronuba in the

garden 1988.

Ausbeute (Individuen) Markierungen Wiederfänge/Mehrfachwiederfänge

SIiN WaS = SIiN WaS 2 WiE.1152 73.2425, 007 BEE

dd 26 492 518 26 400 426 32,17, ,8,..3 2, 1,71 ze89

09 14 433 442 14 355 369 See re rene iR

>2 48 1011 1059 40 755 795 107.:36,.12. 37.217 Pie 7162

Par. 48 448

In Tabelle 25 ist eine Erhöhung der Werte durch die hier mitgezählten Wiederfänge zu

veranschlagen, eine Bereinigung der Angaben wird durch Tab 27 möglich.

Die Weibchen-Raten liegen an SiN (35 %) und WaS (47 %) auf ähnlich hohem Niveau

(NOVAK, 1974: 19 %), in den Wiederfängen zeigen sich die gleichen Verhältnisse.

Die proterandrische Phänologie kommt in Abbildung 20 gut zum Ausdruck. Sie steht im

Gegensatz zu dem von MEINEKE (1984) im südlichen Niedersachsen festgestellten Be-

fund. Nach NOVAK (l.c.) ist N. pronuba weder proterandrisch noch protogyn.

Abb. 20: Ansteigen der Weib-

chen-Rate (Proterandrie) im

Lauf der Flugzeit bei Noc-

tua pronuba im Garten

1988.

Increase of sex-ratio of

Noctua pronuba in the gar- (Stichprobe

den 1988.

A6 M6 E6 AT M7T ET A8 M8 ES AI MI

(Stichprobe

Hauptflugzeit lklen

Tab. 28: Zwischen Fang und Wiederfang festgestellte Intervalle bei Noctua pronuba

1988 im Garten.

Intervals (in days) between two catches of Noctua pronuba 1988 in the garden.

Intervall lTage) se. iv.ı2. 3 074.. Zee ea gm le 12 2

Wiederfänge 141 16 1 1 Zu 0 2 = | = = = 162

Das Intervall zwischen Fang und Wiederfang beträgt im Mittel 1,23 Tage.

Einige der nach einem 2-Tages-Intervall wiedergefangenen Tiere könnten sich in der

dazwischenliegenden Nacht in der Nähe der Falle versteckt haben und übersehen wor-

den sein. Die hohe Zahl der Wiederfänge täuscht also darüber hinweg, wie das Ergeb-

nis richtig zu interpretieren ist: Der Anteil an Wiederfängen, die man nicht auf ein

Festgehalten-Werden im Bann des Lichts zurückführen kann, liegt unter 1 %!

Aus Abbildung 21 errechnet sich eine mittlere Verweilzeit von 1,88 Tagen. Der Wert

der dd’ liegt mit 1,93 Tagen etwas über dem der 99 (1,82 Tage). Die genannten Zah-

len sind in Vergleichen (z.B. mit Scotia clavis) wohl zum großen Teil Ausdruck der

anziehenden Wirkung des Lichts, durch das der Aufenthalt in der Nähe der Lichtquelle

hinausgezögert wird. Noctua pronuba verweilt unter natürlichen Verhältnissen deutlich

kürzer in Flächenausschnitten, die dem Einzugsbereich der Lichtfalle entsprechen.

Aufsummierte Verweilzeiten (Annahme:

Ständige Anwesenheit, nur nicht gefangen)

Wiederfänge nach x Tagen in Relation

zum Erstfang

1 2 3 4 5 6 7 8 9 10 t (Tage)

Abb. 21: Apparente Verweilzeiten von Abb. 22: Aufsummierte Verweilzeiten und real

Noctua pronuba 1988 im Garten. nach x Tagen beobachtete Wiederfänge.

Apparent residence times of Noctua Accumulated residence times and really

pronuba 1988 in the garden. after the time x recaptured specimens.

Die weitgehende Übereinstimmung der beiden Kurven in Abbildung 22 ist auf die be-

sonders große Affinität zum Licht bei dieser Art zurückzuführen. Differenzen, die bei

anderen Arten durchaus auftreten können (siehe Alcis repandata), können zwei Ursa-

chen haben: Entweder ist die anziehende Wirkung des Lichtes nicht besonders groB und

viele in der näheren Umgebung verbleibende Falter werden in den Fangintervallen nicht

erfaßt, oder die Individuen bewegen sich in einer Art Zickzackflug und können so nach

mehreren Tagen der Abwesenheit wieder in den Einzugsbereich der Lichtquelle zurück-

kehren.

Wiederfang-Quote

w£f£.2

® W£3

0 m 30m 60 m 90 m 120 m Versetzdistanz 0m 30 m 60 m 90 m 120 m Versetzdistanz

Abb. 23: Wiederfang-Quoten von Noctua Abb. 24: Korrelation: Rückkehrdauer

pronuba im Versetzexperiment. und Versetzdistanz.

Probability of recapture of Noctua Correlation of distance of displace-

pronuba, released at different distances. ment and time, needed for return.

Schon in einer Entfernung von 30 m ist ein starker Rückgang der Wiederfang-Wahr-

scheinlichkeit festzustellen. Hier nimmt der direkte Lichteinfluß ab und von den ver-

setzten Individuen wird nur der Anteil in den "Lichttrichter" wiedereingefangen, der

zufällig die entsprechende Richtung einschlägt. Daher kommt es mit abnehmendem

Winkel zu einem leichten Abfall zwischen 30 und 120 m.

Zwischen der Rückkehrdauer und der Versetzdistanz ist keine eindeutige Korrelation

festzustellen, ein Befund, der für diesen Wanderfalter zu erwarten war, da es bei der

Bewältigung einer Distanz von 120 m zu keinen Schwierigkeiten kommen dürfte.

Äußerst ähnliche Ergebnisse liegen von einem anderen Wanderfalter (Spodoptera litto-

ralis) in einer ganz anderen Gegend (Israel) vor: PLAUT (1971) berichtet ebenfalls von

einem starken Rückgang der Wiederfang-Wahrscheinlichkeit schon zwischen der ersten

und der zweiten Nacht nach dem Markieren sowie zwischen Versetzdistanzen von 10-50 m.

Die Ergebnisse scheinen also reproduzierbar zu sein.

Amathes c-nigrum:

Bei dieser Art erfolgte das Versetzexperiment nur in einer Serie von 5 aufeinanderfol-

genden Nächten. Die zu kleine Stichprobengröße läßt daher noch keine statistisch un-

termauerten Rückschlüsse zu.

Die Verteilung der Erst- und Zweitwiederfänge (47/5) zeugt von einer starken Dyna-

mik. Beim Wanderfalter Noctua pronuba und bei Scotia clavis waren die Mehrfachwie-

derfänge stärker repräsentiert, was aber teilweise durch eine größere Affinität zu

Licht bedingt sein könnte.

|, Abgesehen von einem nach 4 Tagen zurückgefangenen J ereigneten sich alle Wieder-

fänge nach 1- oder 2-Tages-Intervallen.

ı Auch die mittlere Verweilzeit von nur 1,21 Tagen belegt die hohe Mobilität von Ama-

| thes c-nigrum.

Tab. 29: Reaktion von Amathes c-nigrum auf ein Versetzen in verschiedene Entfernungen.

Recaptures of specimens of Amathes c-nigrum, released at different distances from

the light source.

0 m 30m 60m 9m 120m

freigelassen 239 12 20 17 18

released

Wiederfänge 50 2. _ = A

recaptures

% 21 17 > N £

Ein vergleichbares 30 m-Experiment stellen die 38 an SiN markierten Tiere dar, von

denen keines wiedergefangen wurde. Auch umgekehrt konnte von den 237 an WasS frei-

gelassenen Faltern keiner an SiN nachgewiesen werden. Ähnlich wie bei Noctua pronu-

ba sinken auch bei dieser wanderverdächtigen Art die Wiederfang-Quoten drastisch ab,

sobald sich die Lichtquelle außer Sichtweite befindet.

Rusina ferruginea:

Bezüglich der Stichprobengröße gilt hier das gleiche wie für die vorige Art.

Bei Rusina ferruginea fehlen Zweitwiederfänge gänzlich, und abgesehen von einem 9,

das in 3 Tagen von WaS (0 m) nach SiN flog erfolgten alle Wiederfänge nach einem

Tag und sind durch die Methode bedingt. Die mittlere Verweilzeit liegt dementspre-

chend auf einem sehr niedrigen Niveau von 1,13 Tagen.

Tab. 30: Reaktion von Rusina ferruginea auf ein Versetzen in verschiedene Entfernungen.

Recaptures of specimens of Rusina ferruginea, released at different distances [rom

the light source.

0 m 30m 60 m 90m 120 m

freigelassen 49 10 8 9 f)

released

Wiederfänge 10 | - 3 —

recaptures

% 20 10 - 33 -

Es wäre natürlich Unsinn, Rusina ferruginea aufgrund der oben erwähnten kurzen Ver-

weilzeiten als Wanderfalter zu bezeichnen, sicherlich nicht falsch ist dagegen die Kon-

statierung einer hohen Dynamik und Dispersionsaktivität bei dieser Art.

Meristis trigrammica:

Tab. 31-33: Ausbeute, Markierungen und Wiederfänge im Garten 1988 bei Meristis tri-

grammica.

Numbers of individuals, marked specimens and recaptures of Meristis trigrammica in

the garden 1988.

Ausbeute (Individuen) Markierungen Wiederfänge/Mehrfachwiederfänge

SiN WaS Z SIiN WaS 2 Wf.1h,2, 3,4 5,6 7er su!

dd AU 120 147 22 66 88 33 12. 8, 4.10 12m 58

09 1 5 6 1 322.6 m ee et =

>2 28 125722153 23 71 94 332 na il. ann, 8

Par. Zu) 30

In Tabelle 31 ist eine Erhöhung der Werte durch die hier mitgezählten Wiederfänge zu

veranschlagen, eine Bereinigung der Angaben wird durch Tabelle 33 möglich.

Drei Erstwiederfänge und ein Drittwiederfang erfolgten am Fangplatz SiN.

Die Weibchen sind im Wiederfangergebnis unterrepräsentiert, was aber nicht unbedingt

auf höhere Mobilität hindeutet, sondern auch durch das schlechtere Anflugverhalten in

Verbindung mit den dadurch bedingten statistischen Störeinflussen erklärt werden kann.

Tab. 34: Zwischen Fang und Wiederfang festgestellte Intervalle bei Meristis trigram-

mica 1988 im Garten.

Intervals (in days) between two catches of Meristis trigrammica 1988 in the garden.

Intervall (Tage). 1.2.23, 22075 76. 7.28. 9,10 11.2 >

Wiederfänge 39 8 8 1 RR N 1 1 - - = 58

Das Intervall zwischen Fang und Wiederfang beträgt im Mittel 1,72 Tage.

Im Gegensatz zu den bisher besprochenen Arten zeigt sich hier ein deutlich erhöhter

Anteil der Wiederfänge mit dazwischenliegenden Intervallen, was als Hinweis auf ver-

stärktes Verbleiben in der näheren Umgebung zu deuten ist.

4 Aufsummierte Verweilzeiten (Annahme:

Ständige Anwesenheit, nur nicht gefangen)

Wiederfänge nach x Tagen in Relation

zum Erstfang

23a Se Tao | (le) 123456789 10111213 ! (Tase)

Abb. 25: Apparente Verweilzeiten von Abb. 26: Aufsummierte Verweilzeiten und real

Meristis trigrammica 1988 im Garten. nach x Tagen beobachtete Wiederfänge.

Apparent residence times of Meristis Accumulated residence times and really

trigrammica 1988 in the garden. after the time x recaptured specimens.

Sowohl die hohe durchschnittliche Verweildauer von 3,00 Tagen als auch die deutliche

Differenz der Kurven in Abbildung 26 (siehe Bemerkungen zu Noctua pronuba) ver-

deutlichen, daß Meristis trigrammica im Rahmen des trivial movement zumindest im

Bereich des Gartens relativ ortstreu bleibt.

Wiederfang-Quote

(x)

t (Tage)

| I

| I)

mo Versetzdistanz 0 m 30 m 60 m 90 m* Versetzdistanz

Abb. 27: Wiederfang-Quoten von Meristis Abb. 28: Korrelation: Rückkehrdauer

trigrammica im Versetzexperiment. und Versetzdistanz (* n=3)

Probability of recapture of Meristis Correlation of distance of displace-

trigrammica, released at different distances. ment and time, needed for return.

* only three recaptures

In der Entfernung von 120 m wurde nur ein J freigelassen, das nicht mehr zurück-

kehrte. Vom 30 m entfernten Fangplatz SiN, wo 23 Individuen markiert wurden, flogen

3 (13 %) nach WaS, was für eine Barrierewirkung der Häuserzeile spricht. Umgekehrt

waren es nur 5,6 %, z.I. wohl aber durch den 2tägigen Fangrhythmus an SiN bedingt.

Die Darstellung (Abb. 27) erfolgte unter Hinzunahme der Mehrfach-Wiederfänge, da

einige Wiederfänge auch an den Versetzdistanzen freigelassen wurden.

In kühlen Nächten unternehmen die Tiere keine weiteren Flüge: Zwischen dem 4.6. und

dem 7.6.1988 beispielsweise flogen bei naßkaltem Wetter nach zwei Tagen 100 %, nach

drei Tagen noch 50 % der am 4.6. markierten Falter die Falle an.

Hoplodrina alsines:

Tab. 35-37: Ausbeute, Markierungen und Wiederfänge im Garten 1988 bei Hoplodrina

alsines.

Numbers of individuals, marked specimens and recaptures of Hoplodrina alsines in

the garden 1988.

Ausbeute (Individuen) Markierungen Wiederfänge/Mehrfachwiederfänge

SIiN WaS = SIiN Was 2 wenn VyR Su Tanzes

dd 23 216 239 21 186 207 214 5 1 0-0= Ve

99 i 26,29 La 126 1 - = og

>» 24 242 266 2221179233 VER EE: TEE ve nn an

Par. 24 69

Ein Wiederfang stellt einen Ortswechsler (WaS>SiN) dar, sonst handelt es sich aus-

schließlich um an WaS gemachte Beobachtungen.

Tab. 38: Zwischen Fang und Wiederfang festgestellte Intervalle bei Hoplodrina alsines

1988 im Garten.

Intervals (in days) between two catches of Hoplodrina alsines 1988 in the garden.

Intervall (Tage) 1 200374 2556 70582097: 10. 11% 012 &

Wiederfänge 21.04 Denen 1 - = - = 3

Das Intervall zwischen Fang und Wiederfang beträgt im Mittel 1,81 Tage.

= Aufsummierte Verweilzeiten (Annahme:

Ständige Anwesenheit, nur nicht gefangen)

Wiederfänge nach x Tagen in Relation

20 zum Erstfang

LEE t (Tage)

2a > a8 8 789.

Abb. 29: Apparente Verweilzeiten von Abb. 30: Aufsummierte Verweilzeiten und real

Hoplodrina alsines 1988 im Garten. nach x Tagen beobachtete Wiederfänge.

Apparent residence times of Hoplo- Accumulated residence times and really

drina alsines 1988 in the garden. after the time x recaptured specimens.

Die durchschnittliche Verweildauer beträgt 2,24 Tage, das einzige rückgefangene

Weibchen flog die Falle nach 4 Tagen zum zweiten Mal an.

Die beobachteten Intervalle und Verlustkurven deuten auf eine Ortstreue hin, die nicht

ganz so ausgeprägt ist wie bei Meristis trigrammica, jedoch deutlich höher als bei-

spielsweise bei Noctua pronuba oder Amathes c-nigrum.

Tab. 39: Reaktion von Hoplodrina alsines auf ein Versetzen in verschiedene Entfernungen.

Recaptures of specimens of Hoplodrina alsines, released at different distances from

the light source.

0 m 30 m 60 m 90 m 120 m

freigelassen 135 15 29 24 377,

release

Wiederfänge 24 = 2 1 3

recaplures

% 18 - 7 4 8

Anders als bei Meristis trigrammica zeigt sich hier ähnlich wie bei Noctua pronuba ein

Knick der Kurve bei 30 m. Es ist keine Korrelation zwischen Rückkehrdauer und

Versetzdistanz erkennbar.

Xanthorhoe ferrugata:

Wegen des starken Anfluges im Jahr 1988 mußte bei dieser Art das Versetzexperiment

abgebrochen werden, das Arbeitspensum wäre sonst nicht mehr zu bewältigen gewesen.

So wurden nur 5 Individuen dieser Art bei 30 m, 3 bei 60 m und 2 bei 90 m freige-

lassen. Von diesen markierten Faltern konnte keiner wiedergefangen werden.

Calospilos sylvata:

Diese Spannerart wurde 1988 am Fangplatz "We" im Dachauer Moos näher untersucht.

Zur Hauptflugzeit wurde 5 Fangnächte lang im 3-Tage-Rhythmus gefangen. Auf 629

markierte Falter kamen hierbei 46 Wiederfänge, die sämtlich an der Fangstelle freige-

lassen wurden. Einige Falter wurden dagegen in je 100 m Entfernung an verschiedene

Stellen gebracht um den Einfluß von Geländestrukturen auf die Rückkehr- Wahrschein-

lichkeit zu testen. Hierzu wurden ein Waldweg, eine Weidenhecke und eine Wirt-

schaftswiese ausgewählt.

Calospilos sylvata verweilt sehr lange am selben Standort, wie aus Abbildung 31 er-

sichtlich wird:

Abb. 31: Apparente Verweilzeiten von

Calospilos sylvata im Dachauer

Moos 1988.

Apparent residence times of Calo-

spilos sylvata in the "Dachauer

Moos" 1988.

3 6 9 12 t (Tage)

Aus den Verweildauern der Wiederfänge errechnet sich ein Durchschnittswert von 4,6

Tagen.

Tab. 40: Reaktion von Calospilos sylvata auf ein Versetzen an verschiedene Stellen.

Recaptures of specimens of Calospilos sylvata, released at different places.

0 m 100 m Hecke 100 m Wald 100 m Wirtschaftswiese

hedge. wood meadow

freigelassen 342+ 46 147 90 50

release

Wiederfänge 35 6 S) =

recaptures

% 9 4 6 e:

Ein Ortswiederfang-Ergebnis von 9 % nach zwei fangfreien Nächten ist als sehr hoch

einzustufen!

Zwischen zwei definierten Punkten scheint ein Individuenaustausch dann am meisten

begünstigt zu sein, wenn die dazwischenliegende Strecke waldartig strukturiert ist.

Im Offenland (Wirtschaftswiese) ist jedoch vermutlich eine höhere Mortalität durch

Freßfeinde zu veranschlagen.

Abb. 31b: Calospilos sylvata d (We, 4.7.88)

Abb. 3lc

Peribatodes rhomboidaria (WaS, 26.7.88)

Peribatodes rhomboidaria:

Tab. 41-43: Ausbeute, Markierungen und Wiederfänge im Garten 1988 bei Peribatodes

rhomboidaria.

Numbers of individuals, marked specimens and recaptures of Peribatodes rhomboida-

ria in the garden 1988.

Ausbeute (Individuen) Markierungen Wiederfänge/Mehrfachwiederfänge

SIN Was 2 SIiN Was 2 Wr. 102 3,805 ne LTE

dd 23 3197342 18207 285 44 ER Te

99 a 2 5 Zi __ ET NE EEE

ee ae -

> 31 374 405

Par. 31 154

Die Weibchen-Rate ist an SiN (17,9 %) mehr als doppeit so hoch als an WaS (7,2 %).

Die 99 sind im Wiederfangergebnis unterrepräsentiert, was auch hier durch nicht opti-

males Anflugverhalten und damit verbundene Störungen der statistischen Vergleich-

barkeit zu erklären ist.

Abbildung 32 verdeutlicht das proterandrische Erscheinungsbild dieser Art:

Abb. 32: Ansteigen der Weib-

chen-Rate (Proterandrie) im Lauf

der Flugzeit bei Peribatodes

rhomboidaria im Garten 1988.

Increase of sex-ratio of Periba-

todes rhomboidaria in the garden

1988.

Dekade

Tab. 44: Zwischen Fang und Wiederfang festgestellte Intervalle bei Peribatodes rhom-

boidaria 1988 im Garten.

Intervals (in days) between two catches of Peribatodes rhomboidaria 1988 in the

garden.

Intervallölfage )» 1.212. 32540075, "er 89, = 10, SET? FR

Wiederfänge 35.211 6 = 1 = = = = = = = 53

Das Intervall zwischen Fang und Wiederfang beträgt im Mittel 1,51 Tage.

Aufsummierte Verweilzeiten (Annahme:

Ständige Anwesenheit, nur nicht gefangen)

Wiederfänge nach x Tagen in Relation

zum Erstfang

t (Tage)

Abb. 33: Apparente Verweilzeiten von Abb. 34: Aufsummierte Verweilzeiten und real

Peribatodes rhomboidaria 1988 im nach x Tagen beobachtete Wiederfänge.

Garten.

Apparent residence times of Periba- Accumulated residence times and really

todes rhomboidaria 1988 in the garden. after the time x recaptured specimens.

Die mittlere Verweildauer liegt mit 1,80 Tagen unter dem Wert von Noctua pronuba.

Dies verdeutlicht die Notwendigkeit einer vorsichtigen Interpretation solcher Einzelin-

formationen. Erst in Verbindung mit zusätzlichen Ergebnissen kann sich dann ein voll-

ständigeres Bild der Populationsdynamik ergeben. Im vorliegenden Fall könnte es durch

eine im Vergleich mit Noctua pronuba geringere Affinität ans Licht zu einem weniger

ausgeprägten "Festhaltephänomen" durch die direkte Lichtwirkung gekommen sein. Die

Vergleichbarkeit leidet auch unter den nicht exakt gleich verteilten Individuenzahlen an

den verschiedenen Versetzdistanzen. Da bei P. rhomboidaria (im Gegensatz zu N.

pronuba) überproportional viele Tiere direkt an der Falle freigesetzt wurden, könnte es

zu einer Überbetonung der 1-Tage-Wiederfänge und einer methodisch bedingten Ver-

ringerung der apparenten Verweildauern gekommen sein (siehe Alcis repandata).

Die Differenz der beiden Kurven in Abbildung 34 ist jedoch ein Hinweis auf weniger

dynamische Prozesse als bei den hochmobilen Arten wie Amathes c-nigrum oder Noc-

tua pronuba.

A ns 173

nach |

Ri Widrlng Qui

gesamt

t (Tage)

; / Versetz-

oe 30 m Fo 90 m 120 m Versetz- 0m 30m 60 m 90 m 120 m distanz

distanz

Abb. 35: Wiederfang-Quoten von Periba- Abb. 36: Korrelation: Rückkehrdauer

todes rhomboidaria im Versetzexperiment. und Versetzdistanz.

Probability of recapture of Peribatodes Correlation of distance of displace-

rhomboidaria, released at different distances. ment and time, needed for return.

Von den Wiederfängen (Distanz: 0 m) wurden am 1. Tag 11,8 %, und insgesamt 17,6 %

ein weiteres Mal gefangen. Dies entspricht dem Wert für die Erstfänge.

Bei einer gemäß den Ausführungen in 8.1.3. über die parallel laufenden 0 m-Werte

durchgeführten Bereinigung der Wiederfang-Quoten ergibt sich folgendes Bild, das sich

nicht grundlegend von den Ergebnissen in Abb. 35 unterscheidet:

Tab. 45: Versetzexperiment bei Peribatodes rhomboidaria, bereinigt über die parallel

laufenden 0 m-Werte.

Recapture-rates of Peribatodes rhomboidaria, released in different distances, after

correction by the means of the 0 m-terms.

Versetzdistanz

distance of displacement 0m 320m 60m 90m 120m

W.f.-Quote, 1.Tag (%) 1157 125 17,0 3.3 3,3

W.f.-Quote, 2 (%) 14,6 24,0 12,7 6,2 12,6

Im Prinzip gleiche Ergebnisse resultieren auch dann, wenn man nur die in der Haupt-

flugzeit dieser Art (1.8.-18.8.) gesammelten Daten berücksichtigt. Etwas aus dem Rah-

men fallen lediglich die in einer Entfernung von 120 m ausgesetzten Tiere mit Wie-

derfang-Anteilen von 7,4 % (1. Tag) und 18,5 % (Gesamt).

Aus der Beobachtung, daß am I. Tag nach dem Freilassen ein Gradient festzustellen

ist, der sich danach einigermaßen ausgleicht, könnte man schließen, daß bei zufällig in

verschiedene Richtungen startenden Exemplaren anfangs noch der Winkel zum Einzugs-

bereich der Lichtquelle bestimmend für die Wiederfang-Wahrscheinlichkeit ist. Später

könnte durch "zickzackartige” Richtungsänderungen umherschweifender Falter ein Wie-

dereinfangen in den Trichter der Lichtfallen-Reichweite erfolgen.

In diesem Sinne ist mit zunehmender Versetzdistanz eine Verlängerung der Rückkehr-

dauer feststellbar (siehe Abb. 36).

Ortswechsler von SiN nach WaS (30 m) waren mit insgesamt nur 4,3 % gegenüber den

versetzten Individuen (sogar dem 120 m-Wert) deutlich unterrepräsentiert. Hierbei mag

die Barrierewirkung der Häuserzeile eine besondere Rolle gespielt haben.

Ähnliches gilt für die Ortswechsler in umgekehrte Richtung (? Wiederfänge nach einem

Tag). Die aus 90 m (1) und 120 m (2) nach SiN geflogenen Stücke wurden jeweils

nach 2 Tagen wiedergefangen, was einen weiteren Hinweis auf eine Verlängerung der

Rückkehrdauern bei zunehmender Distanz (vergleiche Abb. 36) darstellt.

Alcis repandata:

Tab. 46-48: Ausbeute, Markierungen und Wiederfänge im Garten 1988 bei Alcis repandata.

Numbers of individuals, marked specimens and recaptures of Alcis repandata in the

garden 1988.

Ausbeute (Individuen) Markierungen Wiederfänge/Mehrfachwiederfänge

SIN WaS Du SiN Was 23 WR 2 3a 5 D=

dd 12 596 608 NES8B77234:8 de letters. eo 50

09 2 55 SU 253 55 Zu em, Ze 2

entkom- “

zen 4 77 81 _

ı 18 728 746

Par. 18 310

In der Weibchen-Rate liegen ähnliche Verhältnisse vor wie bei Peribatodes rhomboida-

ria, sie betrug an SiN 14,3 %, an WaS 8,9 %. Das nicht optimale Anflugverhalten

führte vermutlich zu der feststellbaren Unterrepräsentierung der o9 im Wiederfanger-

gebnis (siehe Bemerkungen zu Peribatodes rhomboidaria).

Ein proterandrisches Erscheinungsbild ist nicht erkennbar (Abbildung 37):

13 590 603 EN)

o9-Rate

(x)

Abb. 37: Verlauf der Weibchen-Rate in der

Flugzeit von Alcis repandata im Garten 1988.

Sex-ratio of Alcis repandata in the months

of the flight time. :

M6 E6 AT M7 Dekade

KawallIere) Ri Or

Wiederfänge 45 et Zen een 1 1 1 - 1 = = 58

Tab. 49: Zwischen Fang und Wiederfang festgestellte Intervalle bei Alcis repandata

1988 im Garten.

Intervals (in days) between capture and recapture of Alcis repandata 1988 in the garden.

Das Intervall zwischen Fang und Wiederfang beträgt im Mittel 1,76 Tage. Auffällig ist

auch die hohe Zahl von Intervallen über 2 Tagen (siehe Bemerkungen zu Noctua pronuba)).

Aufsummierte Verweilzeiten (Annahme:

Ständige Anwesenheit, nur nicht gefangen)

Wiederfänge nach x Tagen in Relation

zum Erstfang

t (Tage)

a 07, Be 9 Ne, TAN EETESON

Abb. 38: Apparente Verweilzeiten von Abb. 39: Aufsummierte Verweilzeiten und real

Alcis repandata 1988 im Garten. nach x Tagen beobachtete Wiederfänge.

Apparent residence times of Alcis Accumulated residence times and really

repandata 1988 in the garden. after the time x recaptured specimens.

Die durchschnittliche Verweildauer von 2,08 Tagen liegt zwar über dem Wert von

Noctua pronuba, jedoch nicht wesentlich. Dies ist, wie bereits unter Peribatodes rhom-

boidaria angedeutet wurde, durch eine Überrepräsentierung der bei 0 m freigelassenen

Tiere verursacht. Diese verblieben durchschnittlich nur 1,35 Tage im Fallenbereich,

bedingt durch einen hohen Anteil an Faltern, die sich eine Nacht lang an der Licht-

quelle festhalten ließen! Geht man von einer gleichmäßigen Verteilung auf die ver-

schiedenen Entfernungen aus, so ergibt sich ein theoretischer Wert von ungefähr 2,8

Tagen, also deutlich mehr als beim Wanderfalter Noctua pronuba.

Auch die relativ große Differenz der beiden Kurven in Abbildung 39 steht vermutlich

im Zusammenhang mit einer niedrigeren Mobilität als beispielsweise bei Noctua pronu-

ba oder Amathes c-nigrum.

[=] Nuederlang: gar De:

nach |

10 Se -Quote 4

gesamt

t Versetz-

oa ee 0 m 30 m 60 m 90 m 120 m distanz

Abb. 40: Wiederfang-Quoten von Alcis Abb. 41: Korrelation: Rückkehrdauer

repandata im Versetzexperiment. und Versetzdistanz.

Probability of recapture of Alcis Correlation of distance of displace-

repandata, released at different distances. ment and time, needed for return.

Die in Abbildung 41 für 60, 90 und 120 m angegebenen Werte sind wegen der geringen

Stichprobengröße (3, 2 und 2) noch wenig aussagekräftig.

Auch hier ergeben sich wie bei Peribatodes rhomboidaria für die Wiederfang-Wahr-

scheinlichkeiten der Mehrfachwiederfänge Prozentsätze, die dem Ergebnis bei 0 m ent-

sprechen. Die Bereinigung mittels der gemittelten 0 m-Werte sowie die ausschließliche

Berücksichtigung der Hauptflugzeit (17.6.-6.7.) führen im Prinzip zu den gleichen Aus-

sagen, wie sie den Abbildungen 40 und 41 zu entnehmen sind.

Das von Peribatodes rhomboidaria deutlich unterschiedene Ergebnis könnte auf vermin-

dert starke Ortsveränderungen im Zickzackmuster zurückzuführen sein. Vielleicht ver-

bleibt ein Teil der Falter, die ab einer Entfernung von 60 m ausgesetzt wurden, an

den entsprechenden Stellen. Wichtig ist jedoch, daß ein Muster, das dem des Wander-

falters Noctua pronuba ein wenig ähnelt - wenn auch ohne den starken Knick in der

Kurve bei 30 m - , nicht unbedingt entsprechend interpretiert werden kann. Die durch-

schnittlich längeren Intervalle zwischen Fang und Wiederfang deuten auf ausgeprägte

Unterschiede hin.

8.4.3. Vergleich der sich ergebenden Muster

Im Versetzexperiment waren in den Reaktionen z.T. deutliche Unterschiede zwischen

den bekannten Wanderfaltern und den Arten, von denen kein Migrationsverhalten be-

kannt ist, erkennbar: Noctua pronuba und Amathes c-nigrum kehren nach einem Ver-

setzen von 30 Metern und mehr nur mehr selten ans Licht zurück; der Prozentsatz

dürfte von dem Winkel vom Freilaß-Punkt zum Einzugsbereich der Lichtquelle abhängen.

Ganz ähnliche Ergebnisse erzielte PLAUT (1971) beim Wanderfalter Spodoptera litto-

ralis in einem ähnlichen Experiment.

Scotia clavis, Scotia exclamationis und Rusina ferruginea reagieren in etwa entspre-

chend, bei Hoplodrina alsines allerdings lassen sich schon Anzeichen für eine nicht so

hohe Dynamik finden.

Stärker sind solche Hinweise bei Meristis trigrammica, Peribatodes rhomboidaria und

Alcis repandata. Diese Arten schweifen vermutlich mehr oder weniger ungerichtet in

der weiteren Umgebung herum und verursachen so Wiederfänge, die nicht durch die

direkte Lichteinwirkung erkiärt werden können.

Die einzige relativ ortstreue Art, die getestet wurde, ist der Spanner Calospilos sylvata.

Im Garten war bei vielen Arten (z.B. Peribatodes rhomboidaria) die Barrierewirkung

der Häuser- und Garagenzeile (siehe 8.1.) aufgefallen.

8.4.4. Hinweise für faunistische Arbeitsansätze

Bei Lebend-Lichtfallenfängen mit quantitativen Arbeitsansätzen erscheint es wichtig,

nicht kontinuierlich zu fangen, da die Häufigkeiten durch verschiedene Wiederfang-

Wahrscheinlichkeiten verzerrt werden.

Ein 2-Tage-Rhythmus ist als die günstigste Methodik anzusehen. Häufigkeitsverfäl-

schungen spielen hier praktisch keine Rolle mehr.

Der direkte Einfluß der Lichtquelle (Schwarzlichtröhre) sinkt offensichtlich schon in

Entfernungen von 30-50 m drastisch ab. Die Grenzwerte einer positiven Antwort sind

artspezifisch verschieden. Diese Befunde entsprechen den Beobachtungen vieler anderer

Forscher (z.B. GROTH, 1951; DANIEL, 1952; SCHADEWALD, 1955 und 1956; RETZ-

BANYAI-RESER, 1986), auch wenn bezüglich der Reichweite noch andersartige Mei-

nungen existieren (z.B. SCHEERPELTZ, 1968 oder URBAHN, 1973).

In einem Versetzexperiment ermittelte PLAUT (1971) für einen Wanderfalter die Fal-

lenreichweite von ca. 30 m, in Tunnelexperimenten kamen GRAHAM et al. (1961) auf

40 m (allerdings an einer Argon-Lampe) und schließlich STEWART et al. (1969) auf

maximal 60-135 m Entfernung für eine positive Antwort auf Lichtreize, ermittelt an

jeweils nur einzelnen Arten.

8.5. RÜCKSCHLÜSSE AUS ORTSWIEDERFÄNGEN

8.5.1. Die Problematik

BETTMANN (1985 a; 1985 b; 1986) versuchte, über Interpretationen von Ortswiederfän-

gen die verschiedenen Nachtfalter hinsichtlich ihrer Ortstreue zu charakterisieren.

Ähnliches publizierten REINHARDT & DROBNIEWSKI (1979) für einige Tagfalterarten.

Als Grundlage diente BETTMANN Material, das aus einer kontinuierlich arbeitenden

Lichtfalle stammte.

Tab. 50: Wiederfänge bei Nacht-Großschmetterlingen in Rheydt (aus BETTMANN, 1986).

Recaptures of moths in Northern Germany (from BETTMANN, 1986).

Markiert Wiederfänge nach x Tagen

released recaptures alter x days

222730475 6-7 „8,59.,.10:,.11,:12.13214.:15,.16.,>16

4877 13 Role ar 3 ne 21, 2 ren a el

Abgesehen von der auffällig niedrigeren Wiederfang-Quote von durchschnittlich 3,5 %

gegenüber 18,8 % im Garten ab Mai 1988 (siehe Tab. 51) ergibt sich eine ähnliches

Verteilungsmuster wie im Untersuchungsgebiet. Berücksichtigt man die Tatsache, daß

BETTMANN die Wiederfänge nach x Tagen in Relation zum Erstfang ermittelte und.

nicht die apparenten Verweilzeiten, dann zeigt sich ein leichter Mangel an Wiederfän-

gen über mehrere Tage hinweg.

Tab. 51: Fang-Wiederfang-Intervalle (Interv.) und Apparente Verweildauern (Verw.) bei

täglichem Betrieb der Falle (WaS ab Mai 1988: 1 und 2) sowie Apparente Verweil-

dauern aus Fängen mit fangfreien Nächten dazwischen (3).

Capture-recapture-intervals (1) and apparent residence times (2) in a garden in Sou-

thern Bavaria (daily catches); apparent residence times (3) in the study area (with

census intervals of at least one night between the catches)

Markiert Fang-Wiederfang-Intervall/Apparente Verweildauer (Tage)

released Capture-recapture-interval/Apparent residence time (days

23456 7 89 1107 111 122703 SITES

1 WaS 4713 7290890 3115107 duilanl4 624 \elol iX imnliEieezEe

1988

Interv.

2 WaS 4444431 482123 76 ie ER Rt] 11 1 1 1

1988

Verw.

3 Gesamt- 18.721 ya Ve 1 3 LA SE a 3 Fa 1 a Br ä er ZN | 1 1 1

fläche

198 1458 * Ortswechsler SiN/WaS

Wenn wir nun die in 8.4. dargestellten Ergebnisse betrachten, wird schnell deutlich,

welche Probleme Beurteilungen mit sich bringen, die sich ausschließlich auf Material

aus kontinuierlichen Fängen stützen:

Von Noctua pronuba beispielsweise wurden über 40 % der Falter am nächsten Tag

wiedergefangen (bei BETTMANN nur 1,2 %). Dieser Prozentsatz sinkt schon dann, wenn

man jeweils nur eine fangfreie Nacht zwischen den Probeentnahmen läßt, je nach

Standort auf Werte um oder unter 1 %.

Das bedeutet, daß eine Nacht (mit einigermaßen günstigen Flugbedingungen) genügt, um

den Tieren ausreichend Gelegenheit zu bieten, sich in einem den natürlichen Gegeben-

heiten entsprechendem Maß zu vermischen. Bei einigen wenigen Arten, z.B. Calo-

spilos sylvata, scheint dies schon durch Tagaktivitäten zu erfolgen.

Dies erklärt auch die Überrepräsentierung der 1-Tages-Wiederfänge, wie sie auch

BETTMANN (l.c.) selbst hervorhebt. Es handelt sich hierbei zum Großteil um Tiere,

die durch die direkte Lichtwirkung "gefangengehalten” wurden.

Ein weiteres bereits angesprochenes Problem ist die unterschiedliche Affinität der Ar-

ten und Geschlechter ans Licht: Bei einigen Weibchen beispielsweise (z.B. Hoplodrina

alsines) wird nur ein kleiner Teil der tatsächlich anwesenden Individuen nachgewiesen.

Diese geringeren Stichproben-Ausschnitte haben - methodisch bedingt - aber eine klei-

nere Wiederfangwahrscheinlichkeit zur Folge.

100

REZBANYAI-RESER (1986) erläutert in seiner Antwort auf die Veröffentlichungen

BETTMANNs die Problematik in sieben Unterpunkten genauer und kommt zum Schluß,

daß "die Ortstreue von Nachtgroßfaltern mit der Kombination der Markierungs- und

Lichtfangmethode an einem einzigen Standort nicht ausreichend erforscht werden kann."

Die offenbleibenden Fragen der Mortalität, der Möglichkeit eines unerkannten Verblei-

bens in der weiteren Umgebung u.s.w. "könnte man eventuell beantworten, wenn im

weiteren Umkreis eines Markierungs-Standortes zahlreiche Lichtfallen oder Lichtfang-

Beobachtungs-Stationen regelmäßig verteilt, aufgestellt würden. Wenn die an einem

bestimmten Tag markierten Nachtgroßfalter aus der Gruppe der Nichtwanderer auch

weiterhin am Leben bleiben und ihre Aktivität unverändert ist, müssen sie auch in den

nächsten Tagen in der weiteren Umgebung durch Licht irgendwo angezogen werden.

Eine solche Untersuchung wäre sehr wünschenswert. Sie ist jedoch äußerst zeit- ar-

beits- und personalaufwendig ...” (REZBANYAI-RESER, l.c.).

Es ist sicherlich richtig, daß Rückschlüsse aus Ortswiederfängen an einem Einzelstand-

ort zu sehr verzerrten Ergebnissen führen können. Wenn man die in der von REZBA-

NYAI-RESER (l.c.) postulierten Methodik (Fangstellen-Netz) gesammelten Daten mit-

einschließt, können Beurteilungen aus Ortswiederfängen gute Zusatzinformationen lie-

fern.

8.5.2. Beispiel: Amathes triangulum

Insgesamt wurden bei 919 markierten Individuen 122 Wiederfänge registriert.

Im Garten (1988) verteilten sich Fänge und Wiederfänge folgendermaßen:

Tab. 51-53: Ausbeute, Markierungen und Wiederfänge im Garten 1988 bei Amathes tri-

angulum.

Numbers of individuals, marked specimens and recaptures of Amathes triangulum in

the garden 1988.

Ausbeute (Individuen) Markierungen Wiederfänge/Mehrfachwiederfänge

SiN WaS D SiN WasS 2 WEL 12 30 W4n SER SE 7Me RB OTEESE

dd 37 313 350 33 311 344 72: 14, 5.,222 02. 112°172127100

99 16 97 113 16 97 113 14,51 SZ ZZ 5

>x 53 410 463 49 408 457 86T 13A1I, 282272,2 El

Par. 5770, 200

Tab. 54: Zwischen Fang und Wiederfang festgestellte Intervalle bei Amathes triangu-

lum 1988 im Garten (nur Ortswiederfänge an WaS).

Intervals (in days) between two catches of Amathes triangulum 1988 in the garden.

Intervall ([age) ı 2 345 678 9 1 2 3 ZZ

Wiederfänge 84 13.3.4 1 2 en = = 1 1 109

Das Intervall zwischen Fang und Wiederfang beträgt im Mittel 1,62 Tage.

101

ni Aufsummierte Verweilzeiten (Annahme:

Ständige Anwesenheit, nur nicht gefangen)

Wiederfänge nach x Tagen in Relation

50 zum Erstfang

t (Tage) ie

age

5 10 15 20

1 5 10 15 20

Abb. 42: Apparente Verweilzeiten von Abb. 43: Aufsummierte Verweilzeiten und real

Amathes triangulum 1988 im Garten. nach x Tagen beobachtete Wiederfänge.

Apparent residence times of Amathes Accumulated residence times and really

triangulum 1988 in the garden. after the time x recaptured specimens.

Die durchschnittliche apparente Verweildauer beträgt 2,20 Tage. Die 99 (1,50 Tage,

n=14) scheinen wesentlich kürzer im Gebiet zu bleiben als die dd (2,63 Tage, n= 72).

Die Weibchen-Rate nimmt auch bei den Wiederfängen, stärker noch bei den Mehrfach-

wiederfängen ab.

Nimmt man als Bewertungsgrundlage nur die im Garten 1988 gesammelten Daten, so

ist die Ähnlichkeit mit dem bei Hoplodrina alsines (siehe 8.4.2.) festgestellten Muster

am größten. Es handelt sich demnach um eine Noctuide, deren Mobilität eine Zwi-

schenstellung zwischen den hochdynamischen Arten wie Noctua pronuba oder Amathes

c-nigrum und der relativ häufig in der weiteren Umgebung verbleibenden Meristis tri-

grammica einnimmt.

Diese Beurteilung deckt sich mit den Ergebnissen der anderen Standorte (siehe z.B.

8.3.1.).

8.5.3. Beispiel: Calospilos sylvata

Diese Art eignet sich durch die vergleichsweise hohe Ortstreue und durch das zahlrei-

che Auftreten im Dachauer Moos gut zu Berechnungen der Populationsgröße. Da nur

wenige Probeentnahmen bei relativ niedriger Fangfrequenz vorliegen, sind die Berech-

nungen als vorläufig anzusehen und bedürfen weiterer Untersuchungen zur Absicherung

der Ergebnisse.

Teilt man das Produkt der beiden Fangergebnisse zweier Probeentnahmen (in der ersten

Nacht nur die markierten Individuen) durch die Anzahl der in der zweiten Fangnacht

wiedergefangenen Individuen so erhält man einen ersten Anhaltspunkt über die absolute

Populationsgröße im Einzugsbereich der Lichtfalle.

So ergibt sich aus dem Fangergebnis vom 1.7.88 (159 markierte Falter) und den Beo-

bachtungen vom 4.7.88 (126 Individuen, davon 22 Rückfänge) eine Populationsgröße von

911 Individuen.

Am selben Standort ergab sich bei einem in zwei aufeinanderfolgenden Nächten 1989

durchgeführten Versuch mit ähnlichen Stichprobengrößen ein Wert von 685 Individuen.

102

Einen Überblick über die Problematik solcher Berechnungen geben unter anderen EHR-

LICH & DAVIDSON (1961), JOLLY (1965), ZINNERT (1966) und ROFF (1973 a und b).

9. UBERSICHT UBER DIE REAKTION DER ARTEN

9.1. VORBEMERKUNGEN

Aufbauend auf den Ergebnissen der Markierungsexperimente und genaueren Betrachtun-

gen der larvalökologischen Ansprüche wird versucht, das ganze im Untersuchungsgebiet

nachgewiesene Artenspektrum hinsichtlich Dispersionsverhalten, trivial movement und

Verbreitungsstrategien zu charakterisieren.

Nach einem Überblick über das 1987 und 1988 im Untersuchungsgebiet gesammelte

Material erfolgt eine Wertung der Wiederfangergebnisse Dies ist nötig, da nicht an

allen Standorten mit der gleichen Methodik vorgegangen wurde.

Bei einem Materialumfang von über 100 markierten Exemplaren werden Markierungs-

tabellen beigefügt. Hierin spiegelt "2 par." das aus den streng vergleichbaren Paral-

lelfängen stammende Fangergebnis wieder, während sich die unter "2 zus.” angegebe-

nen Individuensummen auf die zusätzlichen Fänge beziehen.

In einem zweiten Schritt wird die betreffende Art in das von SCOTT (1975) vorge-

schlagene System zur Beurteilung von (Tagfalter-)Flugdistanzen eingeteilt. Die vier

Gruppen sind folgendermaßen definiert:

1 = "Very small movements"” (gewöhnlich weniger als 100 m)

2 = "Larger movements” (manchmal einige 100 m)

3 = "Still larger movements” (manchmal 1 km)

4 = "Often move many kilometers (or can migrate many kilometers)"; Bewegungen

über viele Kilometer sind keine Seltenheit, hierher gehören auch die einheimi-

schen Wanderfalter.

Durch die Gruppeneinteilung wird ein Bereich abgesteckt, in dem sich das trivial mo-

vement sowie der größte Teil des Dispersionsverhaltens abspielt. Auf die Zuteilung in

eine 5. Gruppe (Wanderfalter über Distanzen von vielen Tausend Kilometern) soll hier

verzichtet werden.

Über eine genaue Betrachtung der Larvalansprüche können weitere Rückschlüsse gezo-

gen werden, diese sind im Abschnitt "Larvalökologie" festgehalten. Fehlende Nachweise

biotopfremder Tiere theoretisch auch durch vermindertes Anflugverhalten an solchen

Standorten bedingt sein. Dies erscheint jedoch angesichts der zahlreichen Beobachtungen

von Gastarten unwahrscheinlich. Außerdem ist oft eine Korrelation zwischen ausgepräg-

ten Häufigkeitsgradienten und stark lokal vorkommenden Raupenfutterpflanzen erkennbar.

Als Unsicherheitsfaktor ist hierbei zu berücksichtigen, daß in Einzelfällen die Literatur-

angaben zu Raupenfutterpflanzen nicht hinreichend präzise oder sogar falsch sind. Bei

einer Reihe von Arten flossen daher in die Beurteilung eigene Beobachtungen mit ein.

Als weitere Hinweise zur "Populationsbiologie” werden Angaben über Bivoltinismus und

sexuelle Koinzidenz gegeben, wenn möglich, erfolgen auch Schätzungen der Popula-

tionsgröße.

Der Versuch einer Einordnung in das r-K-Kontinuum ("r-Stratege”, "K-Stratege", "in-

termediärer Typ") bildet den Abschluß der jeweiligen Artcharakteristik. Das r-K-Kon-

tinuum stellt ein vereinfachtes Schema dar, ist aber, wie dies auch WEIDEMANN

103

(1986; 1986b) für die Tagfalter und SRITZER & LEPS (1988) für die Nachtfalter

betonen, eine plausible Art und Weise, Verbreitungsstrategien zu charakterisieren.

Als Beurteilungskriterien dienten

- die Ergebnisse aus den Fang/Wiederfangexperimenten

- Nachweise biotopfremder Tiere

- einige zusätzliche mehr oder weniger vage Hinweise, die mit der gebotenen Vorsicht

behandelt wurden: Schlechte Flugtauglichkeiten (z.B. flügellose 99), Ausmaß der

"Nacht-zu-Nacht-Fluktuationen” und Häufigkeitsgradienten. Vor allem der letzte

Parameter zeigt deutlich, wie vorsichtig man vorgehen muß, da auch hochmobile

Wanderfalter z.T. in solchen Häufigkeitsgradienten auftreten, was auf "Zugstraßen”

und nicht auf lokale Populationen zurückzuführen ist. Nach SCOTT (1975) sind solche

Gradienten jedoch in heterogenen Lebensräumen Hinweise auf Arten mit kurzen

Flugdistanzen. Arten, die mehr lokal verbreitet sind, benutzen relativ stabile Res-

sourcen und Habitate (GLAZIER, 1986), was ein Merkmal der K-Strategen darstellt.

Nicht berücksichtigt wurden hierbei Bi- bzw. Polyvoltinismus, welche nach MEINICKE

(1984) und GATTER (1981) für hochmobile Arten typisch sind. Dasselbe gilt für die

Fluktuationen von Generation zu Generation, deren Korrelation mit der Position im

r-K-Kontinuum SPITZER & LEPS (1988) bei den Noctuiden Südböhmens belegten.

So können in einem zweiten Schritt die entsprechenden Zusammenhänge vergleichend

getestet werden.

Abschließend wird die Art noch ergänzend hinsichtlich ihrer Verbreitung im Untersu-

chungsgebiet und ihrer beobachteten Habitat- bzw. Ortstreue charakterisiert. Hierzu

dienen folgende 6 Gruppen:

1. Gruppe: Wanderfalter oder "wanderverdächtig"” nach EITSCHBERGER & STEINI-

GER (1980).

2. Gruppe: Ubiquisten, deren Häufigkeit überall mehr oder weniger gleich ist.

3. Gruppe: Die Art ist mehr oder weniger überall anzutreffen, auch die Potenz zur

Verbreitung und die Mobilität sind hoch, es sind jedoch Verbreitungsschwerpunkte

festzustellen.

4. Gruppe: Die Art ist ebenfalls mehr oder weniger überall anzutreffen, sie ist je-

doch vergleichsweise ortstreu; es existieren Verbreitungsschwerpunkte.

5. Gruppe: Die Habitatbindung ist relativ stark ausgebildet, dennoch sind die Potenz

zur Verbreitung und die Mobilität (v.a. innerhalb des Habitats) groß.

6. Gruppe: Die Art ist in starkem Maße habitat- und ortstreu.

Es soll noch darauf hingewiesen werden, daß die Angaben die Verhältnisse im Unter-

suchungsgebiet wiederspiegeln; eine Übertragbarkeit auf andere Gebiete muß nicht un-

bedingt gegeben sein, beispielsweise wird die xerothermophile Actinotia hyperici im

Norden Münchens aufgrund des Mangels an geeigneten Biotopen zweckmäßigerweise

eine Strategie verfolgen, die eher in der K-Region zu suchen ist, während sie in Sü-

deuropa, wo die Gefahr, in biotopfremdes Gelände verschlagen zu werden, geringer

ist, als r-Stratege zu bezeichnen ist und manchmal in unvorstellbaren Mengen gefangen

wird. Es ist an distalen Bereichen des Vorkommens von Arten also bisweilen nicht nur

eine unterschiedliche ökologische Nischenbeanspruchung zu beobachten ("ökologische

Kompensation", siehe WEIDEMANN, 1986, p. 52), sondern auch eine Änderung der

Strategie.

104

9.2. ARTENLISTE

NOLIDAE

Celama confusalis (1 Individuum)

Distanzen: vermutlich 1-2

Verbreitungsstrategie: vermutlich K-Stratege,, 6. Gruppe

LYMANTRIDAE

Dasychira selenitica

Distanzen: 1-2

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Dasychira pudibunda

36 Individuen 11,1 % 9- Rate

33 markiert kein Wiederfang

Wiederfang-Quote: niedrig, Stichprobe noch zu klein

Distanzen: 3

en:

Larvalökol gie. im Raupenstadium an Rot-Buche, Birke, Hain-Buche und Eiche ge-

bunden ( Si 1984), die beiden HM-Stücke sind also zugeflogen (mindestens

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Orgyia recens (7 Individuen)

Distanzen: o9 1 (flugunfähig); I 3

Larvalökologie: im Raupenstadium an verschiedenen Laubhölzern (auch Salix spec.),

das HM-Stück stammt also zumindest aus dem Ruderal (150-300 in).

Verbreit strategie: intermediärer Typ, 3. Gruppe; die o9 sind durch den Verlust

der Flugfähigkeit gezwungenermaßen biotop- und ortstreu, sie legen den Eivorrat

meist auf dem Puppenkokon ab. Die dc‘ sorgen dagegen durch ihre große Mobilität

für eine gute Gendurchmischung. Bei der Kolonisation neuer Lebensräume spielen

wohl die lang behaarten Einsupchen. die leicht vom Wind verdriftet werden, eine

besondere Rolle. Schon durch kurzes Anpusten können 1-3 m zurückgelegt werden.

Lymantria monacha

137 Individuen 1,4 % 9- Rate

65 markiert 5 Wiederfänge

Wiederfeng-Quote: niedrig, alle Wiederfänge WaS 1988 nach 1-Tages-Intervallen,

‚also durch die direkte Lichtwirkung gefangengehalten.

Distanzen: 2-3

Larvalökologie: an Bäume gebunden (v.a. Nadelhölzer), die Strecke von 1 km (nach

HM) wird normalerweise nicht bewältigt, schon in 50 m Entfernung vom Nadel-

waldrand starker Rückgang der relativen Häufigkeit (siehe Wasserwerk

Verbreitungsstrategie: intermediärer Iyp; 9. GEUppe: ei Massenvermehrungen wohl

erhöhte Mobilität; SCHWERDTFEG (1978) berichtet von Massenflügen diese

rn mobei es sich seiner Meinung nach nur um einen passiven Transport (Wind)

andelte.

Euproctis chrysorrhoea (2 Individuen)

Distanzen: 2-3

Larvalökologie: im Raupenstadium an Laubholz gebunden, vor allem an Obstbäume

und Eiche. Das HM- emntaralaı) legte also wahrscheinlich 800-1000 m zurück.

Verbreitungsstrategie: vermutlich r-Stratege, 3. Gruppe

Porthesia similis

138 Individuen 13,8 % o- Rate

46 markiert kein Wiederfang

105

Wiederfang-Quote: wohl niedrig, die Probeentnahme erfolgte im Moos jedoch nur in

wöchentlichen Abständen

Distanzen: 2 ee Siedlungsbereich noch nie festgestellt!)

Verbreitungsstrategie: K-Stratege, 6. Gruppe

ARCTIDAE

Cybosia mesomella

167 Individuen 0% 9- Rate

49 markiert 1 Wiederfang

Wiederfang-Quote: ein Wiederfang (4) erfolgte nach 6 Tagen (WaN>WNo= 50 m)

Distanzen: 2-3 i

Verbreitungsstrategie: intermediärer Typ, 3. Gruppe

Miltochrista miniata (9 Individuen)

Distanzen: 1-2

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Lithosia quadra (17 Individuen)

Distanzen: 2-3

Larvalökologie: die Raupen dieser Art können bei Massenvermehrungen von Lyman-

u monacha als Mordraupe zur Einregulierung der Bestände dieses Schädlings

eitragen.

Verbreitungsstrategie: vermutlich ähnlich wie L. monacha intermediärer Typ, 5.

Gruppe; nach BAHN 11973) mit langfristigen Häufigkeitsschwankungen

Eilema depressa

siehe verringerte Fallendistanzen 8.3.

713 Individuen 55,1 % 9- Rate

575 markiert 4 Wiederfänge

Tab 55: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von Eile-

ma depressa

SIEDLUNG WALD HALBTROK- Dat WALD

l KENRASEN MOL 2 1988 Garten Wasserwerk

1987 | Garten |!

SIS SIMıSN WaS;ıWaM WaN HO HM HW eS WaN WNw WNo HO HM HW Au We

Z par. a So 3 2 6 42 1 1 2 136 Zpar. 7 13 173 162 81 107 2 Es - 1

zu = 0-8 Se 4 - - - 15 zus - 23 u. ER hi =

dd a 9 6 36 17 1 - 2 75 [0102 3 9 86 62 37 39 1 2 = =

99 -. - 6 - 6 26 28 _ 1 = 70 99 4 27 86 87 43 66 1 2 = =

Mark. 12 10 57 32 1 1 2 126 Mark. 6 27 146 124 66 76 1 3 = =

Wif. Ssshalrs il Asse Age - 1 we = = BE Ne nn >=. "| 2

Wiederfang-Quote: gering! An WaS trotz täglichem Fang 1988 kein Rückfang; WaS

1987 ein J nach 2 Tagen wiedergefangen

Distanzen: 3

Larvalökologie: die Raupen leben an Nadelholzflechten, die Strecke nach HM () 800 m)

wird regelmäßig bewältigt.

Populationsbiologie: deutlich proterandrisch!

Verbreitungsstrategie: r-Stratege, 3. Gruppe

Eilema lutarella (\ Individuum)

Distanzen: vermutlich 2 E.

Verbreitungsstrategie: vermutlich intermediärer Typ, 5. Gruppe

Eilema complana

52 Individuen 36,6 % o- Rate

40 markiert 1 Wiederfang

Wiederfang-Quote: wohl durchschnittlich, der Wiederfang (9, HO) erfolgte nach 3

agen.

Distanzen: 2-3

Verbreitungsstrategie: r-Stratege, 5. Gruppe

106

HALBTROK- ze |

ER RR

Eilema lurideola (9 Individuen)

Distanzen: vermutlich 2-3

Verbreitungsstrategie: vermutlich intermediärer Typ, 5. Gruppe

Systropha sororcula

153 Individuen

112 markiert

14,2 % 9 - Rate

kein Wiederfang

Tab 56: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von Sys-

tropha sororcula.

DLUNG WALD HALBTROK- "DA! WALD

SIEDE! l KENRASEN MOOS: 2 1988 Garten

1987 ı Garten ! Wasserwerk

SiS SIMı SN WaS,WaM Wa HO HM HW N WaS WaN WNw WNo HO HM HW Au We

2Ep8l> 3 nmel 1 3 5 30 - - 3 43 I par. 1 - 16 15 18 34. - - 1 2

Zus - - 4 (ya - = - - 22 % zus. - 1 - - - > = 2 a =

dd - - 2 1 2 12 21 - - - 38 dd 1 1 8 12 12 23 - - - 2

99 a = ee DE. 1 1099 - - 2.1 Ze een =

Mark ZU > 1 48 Mark 1 ) Je ee #25

W.f. - - - = = = = > = = — W.f. - - - - = = = = = 3

Wiederfang-Quote: sehr niedrig

Distanzen: 3; innerhalb des Habitats ist die Art beweglicher (siehe "Wiederfang-

Quote”) als außerhalb davon (siehe "Larvalökologie"

Larvalökologie: im Raupenstadium auf Laub- und Nadelholzflechten angewiesen, das

HM-Stück ist also eın Zuflieger, die Strecke von (mindestens) 800-1000 m über

benopiremdes Gebiet wird nur gelegentlich bewältigt.

a biologie: ‚proterandeisch: O am 11.6.88 ein "schwarmartiges" Auftreten

21 Ex.), 2 Tage davor und danach jedoch kein einziges Stück; solch starke Fluk-

tuationen von Nacht zu Nacht sind oft mit Ortswechsel-Ereignissen korreliert.

Verbreitungsstrategie: r-Stratege, 5. Gruppe

Atolmis rubricollis (85 Individuen)

Distanzen: 2

Larvalökologie: Larvalansprüche wie bei der vorhergenannten Art, im Offenland bis-

her jedoch noch nie beobachtet.

Verbreitungsstrategie: iniermediärer Typ, 5. Gruppe

Phragmatobia fuliginosa

382 Individuen 6,8 % o- Rate

360 markiert 4 Wiederfänge

Tab 57: Fangergebnisse,

ı br Geschlechterverteilung und Markierungsergebnisse von

Phragmatobia fuliginosa.

HALBTROK- "DA

KENRASEN MOOS £

SEDLUNG WALD HALBTROK- IDAEE D HALBTROK- Be:

1987 ı Garten ! KENRASEN M 2 1988 Garten Wasserwerk KENRASEN M Di

SiS SMıSN WaS,WaM Wa HO HM HW Mb SN WaS WaN WNw WNo HO HM HW Au We

Z par. 1 2 - 10 35 65 41 48 12 215 Z par. 1 9 3 1422 33 9222 5 4 150

dd 1 1 1 _ 8 33 69 43 45 10 211 dd 1 10 27 14 18 30 7.19 6 3 134

99 SoE Sn 220 2 2E a Mi 3 1 1399 = - a Wen ana ea

Mark. 1 1 1 - 10 35 70 43 43 11 215 Mark. 1 9 31 14 20 33 87721 6 3 145

Wit. ee Sa > Me) - 4 wi - - =, TE Eee =

Wiederfang te: vergleichsweise niedrig, alle Wiederfänge erfolgten schon 2 Tage

nach dem Erstfang, auch am Wasserwerk 1988 keine besonders starke Ortstreue

nachgewiesen.

Distanzen; 3; an SiS ist eine as Häufigkeitsabnahme gegenüber dem Flughafen-

gebiet (200-300 m Entfern ‚in dem die Art wohl allgemein häufig ist, festzu-

stellen. Vermutlich ist ein Großteil der in der Siedlung festgestellten Exemplare

zugeflogen. Am 29.7.88 wurde hier aus einem wohl durchziehenden "Schwarm" 8

Stücke gefangen, tags darauf war noch ein Tier festzustellen, dann keines mehr. _

ulationsbiologie: bivoltin; im Mai 1989 wurden 3 dd‘ tagsüber bei hoher Agilität

gekäschert. Ein Nachweis einer solchen Flugaktivität steht für die 2. Generation

noch aus.

Verbrei strategie: intermediärer Typ, 3. Srunz est lich können auch die

Be schnell laufenden Raupen iR an warmen Herbsttagen) 100 m und mehr zu-

rücklegen.

107

Spilarctia lubricipeda (116 Individuen)

Distanzen: 2-3

Larvalökologie: findet durch Polyphagie überall Lebensgrundlage

Verbreitungsstrategie: K-Stratege, 4. Gruppe; zur Konstanz der Populationsverhält-

nisse vergleiche REICHHOLF (1974).

Spilosoma menthastri

siehe verringerte Fallendistanzen (8.3.) und Versetzexperiment (8.4.)

434 Individuen 10,1 % o- Rate

421 markiert 23 Wiederfänge

Tab 58: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von Spi-

losoma menthastri.

m on 2 SEE 5

SIEDLUNG WALD HALBTROK- DA WALD HALBTROK- "DA

1987 ! Garten ! KENRASEN M 2% 1988 Garten Wasserwerk M

SiS SIMıSN WaSıWaM WN HO HM HW Mb SN WaS WaN WNw WNo HO HM HW Au We

Diherc It Bye 2u2s- 26 - 26).:3.1,46 Muo3,-ass Zpar.* 3° 65 Mi 28. 60. 30 WS

Eh Te ee) 7 me 32 u - aa izüs er 7 - - - a zw =

dd 3.2 4 6 4.2,,68, 20 340 18 66 6 2 0 3% 27 55 36 Bess

9 a Aal 23. 2.08 3 2099 1 3 3 1 _,6 3 ja Dee

Merk. 3 3 5 10°, 2: 68:,,23 37482 7231 173 Mark. , (3 "13 38.27” 60’ W a0 a nn

Wit. RL = ir) alu 2 - 5 wi. = am u u

Wiederfang te: hoch, bei langen Verweildauern: Die 1987 insgesamt sowie die

1988 an HOÖ und HW registrierten Rückfänge (6 d’d', 10) erfolgten nach einem In-

tervall von durchschnittlich 4,6 Tagen.

tanzen: 2

Larvalökologie: S. menthastri findet durch Polyphagie überall eine Lebensgrundlage.

Populationsbiologie: der a an den iederfängen entspricht in etwa der

-Rate der Erstfänge. Dennoch dürften die 99 - vor allem diejenigen mit vollem

ivorrat - ortstreuer als die dc’ sein. Ein nicht optimales Anflugverhalten der 99

könnte das Wiederfangergebnis verfälscht haben. Die Konstanz der Populationsver-

hältnisse entspricht den Befunden in REICHHOLF (1974).

Verbreitungsstrategie: K-Stratege, 4. Gruppe; die schnelllaufenden Raupen können

100 m und mehr zurücklegen.

Diacrisia sannio

siehe verringerte Fallendistanzen (8.3.)

167 Individuen 0% 9-Rate

90 markiert* 6 Wiederfänge* *1989 weitere 19 markiert, 2 Wiederfänge

Tab 59: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von Dia-

crisia sannio.

WALD HALBTROK- DACH

198 Garten Wasserwerk KENRASEN M 2

SN WaS WaN WNw WNo HO HM HW Au We

pa. - - 19 14 24 6 4 34 1 - 102

og - - 19 14 24 6 4 34 1 - 102

2) = = E = = zu ik =

Mark. = - Ian] 6 4 29 1 - 90

W.f. - = 2 1 3 = = - - - 6

Wiederfang-Quote: am Wasserwerk hoch, am Flughafen bei vergleichbarer Abun-

danz niedrig! Die Ortstreue muß also nicht eine artspezifische Konstante sein,

sondern kann auch vom Biotop oder von der Geländestruktur abhängen. Auffallend

sind bei dieser Art die langen Verweildauern.

Distanzen: 2-3; im Ort wurde diese Art noch nie beobachtet, die Strecke von 1 km

wird also, zumindest über biotopfremdes Gebiet, so gut wie nie bewältigt.

Populationsbiologie: tagsüber sind durchaus auch 99 zu beobachten. Bivoltın.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Arctia caja

149 Individuen 24,4 % o- Rate

49 markiert 9 Wiederfänge

108

Wiederfang-Quote: alle 9 Wiederfänge beziehen sich auf ein d (SiN 1988), das die

Falle in 10 aufeinanderfolgenden Nächten anflog.

istanzen: 2-3, die nur recht vereinzelt im Ort anzutreffenden Stücke werden meist

am Flugzeitende beobachtet (Zuflug?).

Populationsbiologie: man könnte sich die Population in einem dynamischen Fließ-

gleichgewicht befindlich vorstellen, das zu dem beobachteten Ost/West-Gradien-

ten in der Häufigkeit führt. !

Verbreitungsstrategie: r-Stratege, 3. Gruppe; die schnellaufenden Raupen können

vermutlich 100 m und mehr zurücklegen.

ENDROSIDAE

Pelosia muscerda (1 Individuum)

Distanzen: vermutlich 2(-3?); _

Verbreitungsstrategie: vermutlich K-Stratege, 5. Gruppe; der Lebensraum am Was-

serwerk wurde jesoen vor einigen Jahren neu besiedelt, höchstwahrscheinlich sogar

über viele km hinweg (nächstgelegenes bekannte Vorkommen: Isarauen bei Lands-

hut, ca. 40 km, vielleicht aber auch bei Ismaning: ca. 7 km).

NOTODONTIDAE

Harpyia furcula (3 Individuen)

Distanzen: vermutlich 2-3 R

Verbreitungsstrategie: vermutlich intermediärer Typ, 5. Gruppe

Cerura vinula

15 Individuen 20,0 % o- Rate

15 markiert kein Wiederfang

Wiederfang-Quote: Stichprobe zu klein

Distanzen: 3

Larvalökologie: Raupenfunde im Gebiet bisher nur an Salix spec.; die HM-Exempla-

re stammen wohl aus dem Ruderal, die nächstgelegenen Raupenfutterpflanzen lie-

gen in dieser Richt 200-300 m entfernt. Die SiN-Stücke sind ebenfalls zugeflo-

gen: Einzelstehende Weiden sind in einer Entfernung von ca. 200-300 m zu finden.

Verbreitungsstrategie: r-Stratege, 5. Gruppe

Stauropus fagi

21 Individuen 0% o- Rate

16 markiert kein Wiederfang

Wiederfang-Quote: Stichprobe zu klein

Distanzen: 2-3 r

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Hybocampa milhauseri (1 Individuum)

Distanzen: vermutlich 2

Larvalökologie: mOnophas an Eiche; das 9 im Franzosenhölzl 1989 flog mindestens

50 m weit zur Lichtfalle, sonst nur im typischen Habitat.

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Gluphisia crenata (5 Individuen)

Distanzen: 2-3

Larvalökologie: auf Populus spec. spezialisiert; von den Fundorten SiM und HW lie-

gen die nachstgelegenen Pappelvorkommen jeweis ca. 200 m entfernt.

Verbreitungsstrategie: K-Stratege, 5. Gruppe. Die Kolonisierung des Raums Ober-

schleißheim in den letzten Jahrzehnten erfolgte vermutlich von den Isarauen her,

die an der nächsten Stelle 7 km entfernt sind. Hierbei könnte die Begleitvegetation

des Schloßkanalsystems als Trittsteinle) eine besondere Rolle gespielt haben.

Drymonia trimacula

24 Individuen 0% o- Rate

23 markiert kein Wiederfang

Wirgerlang Quo wohl niedrig, Stichprobe noch zu klein

anzen:

Larvalökologie: oligophag (im Gegensatz zur folgenden Art)

109

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe; die Oligophagie erlaubt der. Art

eine größere Mobilität, die Gefahr, in ungeeignete Biotope zu gelangen ist geringer

als bei einer monophagen Art wie Drymonia ruficornis.

Drymonia ruficornis

56 Individuen 18,2 % o - Rate

55 markiert 2 Wiederfänge

Wiederf: te: relativ hoch; die beiden Wiederfänge (dd, HO) erfolgten nach

jeweils agen.

Distanzen: im Habitat 2-3, außerhalb 1-2

Larvalökologie: eine ee an Eiche SS nunlene Art, die sehr biotoptreu ist: Sie

wurde an den Fangplätzen HM und HW nie festgestellt, obwohl sie im KelesEs

Eichenwaldgürtel rund um den Flughafen herum allgemein verbreitet ist (1989

wurde auch südlich des Flughafengebiets eine beträchtliche Anzahl Eelongehhk Von

HO liegen. die OeHatzP1epenen Vorkommen ca. 800-1000 m, von HW 500-600 m

entfernt, einzelstehende Eichen befinden sich in letzterem Fall sogar noch näher.

Der normale Aktionsradius über biotopfremdes Gebiet von D. ruficornis sollte also

unter, diesen Werten liegen.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Peridea anceps

112 Individuen 5,3 % 9- Rate

110 markiert 5 Wiederfänge

Tab 60: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von Peri-

dea anceps

SIEDLUNG WALD HALBTROK- "DA WALD HALBTROK- "DA

1987 ‚| Garten _! MOOS 2 1988 Garten Wasserwerk Moosr

SiS SIMıSN WaS,WaM WaN HO HM HW Mb SN WaS WaN WNw WNo 2 HO HM HW Au We

Epe. - - - = -

EaesueBenh Hi unfpr tae Mz nd ue Ei Satin

oe ANERLZ Be EZ 1 3 de 3%: 09 2.0 ig Eee -

99 zwi - Ir taste = re - 4.09 = 1 zujente 1sıh= = SF:

Mark. - - 3 6 6 2 19 - 3 1 40 Mark 3 3 2 - 3 55 1 1 2 -

W.f. = pe yx Sur une Ina = - 1 wi. - - = 4 - > - >

Wiederfang-Quote: hoch, bezogen auf den Fangplatz HO 7,8 %! Bei den Wiederfän-

en handelt es sich um 5 dd‘, die nach einer durchschnittlichen Verweildauer von

‚8 Tagen wiedergefangen wurden.

Distanzen: 2-3

Larvalökologie: Wie die vorige Art monophag an Eichen. Im Prinzip gilt das für D.

ruficornis Gesagte, eine etwas größere Mobilität ist jedoch festzustellen. Die HW-

Stücke, vor allem das 9 stammen wohl von den einzelstehenden Bäumen der nä-

heren a Sue: 2-3 m hohe Eichen finden sich schon in ca. 100 m Entfernung.

Das HM-Stück ist jedoch aus mindestens 800-1000 m Entfernung bei windstillem

Wetter zugeflogen, was aber vielleicht schon die obere Grenze der Reichweite

über biotopfremdes Gebiet darstellt. Diese Art fällt auch durch einen etwas plum-

pen Flug auf. Im Franzosenhölzi wurden 1989 in 50 m Entfernung zu einer einzel-

stehenden Eiche 7 Exemplare, in 90 m Entfernung nur noch 2 Exemplare ge-

fangen.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Pheosia tremula

54 Individuen 0 % 9- Rate

43 markiert kein Wiederfang

Wiederfang-Quote: relativ niedrig, im Birket 1987, wo die meisten Markierungen er-

folgten, wurde allerdings nur in wöchentlichem Fangrhythmus gefangen.

Distanzen: 2-3 e

Larvalökologie: im Larvenstadium hauptsächlich an Pappeln und Weiden. Die an HM

gefangbnen Tiere stammen wohl aus den 150-300 m entfernten Weidenbeständen

es Ruderals. An den auch im Ort vorkommenden Birken kann P. tremula_offenbar

keine auf Dauer erfolgreichen Kolonisationsversuche durchführen: Das Exemplar

1986 im Garten war vermutlich ein Zuflieger.

Populationsbiologie: bivoltin

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

110

Pheosia gnoma

75 Individuen 0% o- Rate

73 markiert 2 Wiederfänge

Wiederfang-Quote: niedrig! Die beiden Falter (WaS 1988) waren durch die_direkte

‚Lichtwir ng festgehalten worden, ihre Rückfänge erfolgten nach jeweils 1 Tag.

Distanzen: 3- k

Larvalökologie: die Raupen fressen monophag an Birken. Umso erstaunlicher ist_das

fast regelmäßige Auftreten im Offenland {HM), wovon die nächstgelegenen Rau-

Beaiutierpflanzen 800-1000 m entfernt sind. Der Zuflug war hier im Jahr 1987 in

eiden Fällen von Westwinden begünstigt.

1988 wurden 10 der 12 WaS gefangenen Falter an 2 aufeinanderfolgenden Tagen

gefangen, was eher durch "schwarmartiges" Auftreten als durch eine eriolerciche

rut in der näheren Umgebung zu erklären ist: Ein solch synchrones Schlüpfen

sollte ein Zufall sein.

989 tauchte P. gnoma im Garten zum ersten Mal in der ersten Generation auf

frisches Stück). Es handelt sich vermutlich um eine erfolgreiche Kolonisation der

8 zugeflogenen Exemplare.

Populationsbiologie: bivoltin

Verbreitungsstrategie: r-Stratege, 3. Gruppe

Notodonta phoebe

Distanzen: vermutlich 2-3 3 ei

Verbreitungsstrategie: vermutlich intermediärer Typ.

Notodonta dromedarius

27 Individuen 3,8 % 9 - Rate

26 markiert kein Wiederfang

Wiederfang-Quote: relativ niedrig, Stichprobe jedoch noch zu klein.

Distanzen: 2-3

Larvalökologie: SHFopas an einigen Laubholzarten; die ut der HM-Stücke

liegt wahrscheinlich in den eidenbeständen des Ruderals (150-300 m), diese

Strecke scheint regelmäßig bewältigt zu werden.

Populationsbiologie: bivoltiin

Verbreitungsstrategie: intermediärer Typ, 3. Gruppe

Notodonta ziczac

46 Individuen 14,0 % o- Rate

43 markiert 1 Wiederfang

Wiederfang-Quote: durchschnittlich; ein 1987 an HM nach 2 Tagen rückgefangenes J

war in der dazwischenliegenden Nacht (günstige Flugnacht) wohl nicht am Fang-

latz verblieben, an dem die Art als ' jiopfremd, anzusprechen ist. Vermutlich

on das Stück die 150-300 m in das Ruderal (zurück) und dann erst wieder an die

ichtfalle, was einer Mindest-Gesamitstrecke von 450-900 m entspricht, unter der

Minimal-)Annahme, daß das J aus dem Ruderal stammt.

Distanzen: 5-4 A

Larvalökologie: Raupenfutterpflanzen sind Salix- und Populusarten. Die ne,

strecke der an HM festgestellten Tiere beträgt also 150-300 m (Ruderal ‚ die der ım

Garten Belangenen Stücke 200-300 m. Diese Entfernungen werden ohne Probleme und

in größeren Stückzahlen geflogen, im Falle des Gartens auch einmal von einem 9.

Populationsbiologie: bivoltin

Verbreitungsstrategie: r-Stratege, 3. Gruppe

Leucodonta bicoloria

17 Individuen 0 % o-Rate (1989 1 9 am Licht)

13 markiert 2 Wiederfänge

Wiederfang-Quote: relativ hoch, Stichprobe jedoch noch zu klein. Ein Wiederfang ist

methodisch durch den direkten Lichteinfluß begründet, es handelt sich um ein nach

einem Tag rückgefangenes Tier 1988 WaS. Ein anderes c' wurde 1988 an HO nach

2 Tagen wiedergefangen.

Distanzen: 2-3

Larvalökplogie: monophag an Birke (nur selten Eiche). Diese Art ist nach OSTHEL-

DER (1925-1933) sowie nach WOLFSBERGER (1974) in Schleißheim bzw. allge-

mein ein typischer Bewohner der Birkenmoore, L. bicoloria wich wahrscheinlich

erst in den letzten Jahrzehnten nach der Trockenlegung des Dachauer Mooses

verstärkt auf andere Waldgebiete aus. Trittsteine für eine solche Besiedlung sind

in ausreichendem Maße vorhanden.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

111

Odontosia carmelita (1 Individuum)

Distanzen: vermutlich 2-3 j J

Larvalökologie: Futterpflanzen nur Birke, Erle; bisher keine biotopfremden Tiere

Populationsbiologie: Art mit lokalem, inselartigem Vorkommen beı meist geringer Po-

pulationsdichte

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Lophopteryx camelina

93 Individuen 2,2 % o- Rate

78 markiert 2 Wiederfänge

Wisderfang-Quote: niedrig, beide rückgefangenen dd nach 1-Tages-Intervallen (1988,

Distanzen: 2-3

Larvalökologie: die Raupe ist auf abge angewiesen, das HM-co 1988 flog also

mindestens vom Ruderal her zu (150-300 m).

Populationsbiologie: bivoltin

Verbreitungsstrategie: intermediärer Typ, 3. Gruppe

Lophopteryx cuculla

19 Individuen 0 % 9 - Rate

16 markiert kein Wiederfang

Wiederfang-Quote: niedrig, Stichprobe jedoch noch zu klein

Distanzen: 2-3

Larvalökologie: oligophag an Ahornarten; an die HM-Falle flog 1986 bei windstillem

Wetter ein Exemplar, das mindestens 500 m zurückgelegt hatte: In dieser Entfer-

nung befinden sich einige junge Ahornbäumchen (Allecan flanzung).

Populationsbiologie: die Generationenfrage ist im Untersuchungsgebiet noch ungeklärt:

ie Daten bis 1988 deuten auf Monovoltinismus hin, wie ın KOCH (1984) be-

schrieben. Das Exemplar E5 1989 ist jedoch ein Hinweis auf "zwei voneinander

unabhängige Stämme” (FORSTER & WOHLFAHRT, 1960).

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Ptilophora plumigera

14 Individuen 0.%” = o- Rate

1 markiert 1 Wiederfang

Wiederfang-Quote: Stichprobe natürlich noch zu klein: I (WaS) nach 2 Tagen, in

‚der dazwischenliegenden Nacht erfolgte kein Lichtfang.

Distanzen: 2,

Larvalökologie: wie L. cuculla, jedoch keine biotopfremden Tiere festgestellt

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Pterostoma palpina

21 Individuen 0% o9-Rate

20 markiert kein Wiederfang

Wiederfang-Quote: niedrig, Stichprobe noch zu klein

Distanzen: 3

50-300 m) wird ähnlich wie bei Notodonta ziczac ohne Probleme und regelmä-

iE geflogen.

Populationsbiologie: bivoltin

Verbreitungsstrategie: r-Stratege, 3. Gruppe

Phalera bucephala

97 Individuen 4,2 % 9- Rate

95 markiert 4 Wiederfänge

Wiederfang te: durchschnittlich; 3 dd , die im Garten (WaS) nach einem 1-Ta-

es-Intervall wiedergefangen wurden, erklären sich durch das Gefangenschaft im

nziehungsbereich der täglich betriebenen Lichtfalle, 1 @ wurde im Birket 1987

nach 5 Tagen wiedergefunden.

Ein markiertes J ging interessanterweise am nächsten Tag in der Falle eine Ko-

pula mit einem unmarkierten 9 ein.

Distanzen: 3, £

Larvalökologie: Larvalansprüche sowie Bemerkungen wie bei Fer elome palpina

Verbreitungsstrategie: r-Stratege, 3. Gruppe; im Ort wurden (ähnlich wie bei Phrag-

matobia fuliginosa und Pheosia gnoma) 1988 sämtliche nachgewiesenen Stücke an

-4

O0 m} polyphag Laubbäume Kan Weide, Pappel); die Strecke Ruderal>HM

112

zwei aufeinanderfolgenden Tagen registriert. Ein solches Auftreten in "Schwärmen"”

im Siedlungsbereich steht den vermutlich geringeren Dispersionsaktivitäten in

feuchteren Wäldern (Dachauer Moos) gegenüber.

Clostera curtula

28 Individuen 4,3 % o9- Rate

23 markiert kein Wiederfang

Wiederfang-Quote: niedrig, Stichprobe noch zu klein

istanzen: 2-3, in der 2. Generation mit größerer Dispersionsaktivität

Larvalökologie: an Salix Ep und Populus spec.. Die nächstgelegenen Raupenfutter-

pflanzen liegen von SiM wie von HM ca. 200-300 m entiernt. Solche Distanzen

scheinen ohne Mühe und regelmäßig bewältigt zu werden. Im Garten wurde C.

curtula dagegen nur einmal 1989 beobachtet: Die Distanz von I km (von SiM

bzw. 1,3 km (Wasserwerk) liegt nicht im normalen Aktionsradius der Art und wir

nur ausnahmsweise geflogen. N 2

Biotopfremde Tiere treten vor allem in der 2. Generation auf!

Populationsbiologie: bivoltin

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Clostera anachoreta (3 Individuen)

Distanzen: vermutlich 2-3 :

Larvalökologie: wie C. curtula, biotopfremde Tiere wurden noch nicht beobachtet.

Populationsbiologie: bivoltin

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Clostera pigra

21 Individuen 4,8 % o- Rate

18 markiert kein Wiederfang

Wiederfang-Quote: wohl niedrig, Stichprobe noch zu klein

Distanzen: 2-3

Larvalökologie: wie C. curtula. Die Distanz von 150-300 m (Ruderal>HM) liegt

auch hier im Bereich der normalen Dispersion. Die Strecke von 1 km (Zuflug ın

den Garten) wird im Normalfall nicht bewältigt. Tas Exemplar am S-Bahnhof

28 uß mindestens 300 m zurückgelegt haben (nächstgelegene Raupenfutter-

anze).

Populationsbiologie: bivoltin

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

ZYGAENIDAE

Diese Familie soll hier ausgeklammert werden, da die Imagines tagaktiv und mit

Lichtfang nicht zu erfassen sind.

Die Dispersionsaktivitäten von 5 Arten (darunter Zygaena filipendulae und Huebneriana

lonicerae) sind in SMOLIS & GERKEN (1 56) usführlich abgehandelt, zu den ökologi-

schen Ansprüchen der Arten siehe BLAB (19 2).

COCHLIDODAE

Apoda limacodes (196 Individuen)

Distanzen: 3-4

Larvalökologie: als Raupe an Laubgehölzen, v.a. Eiche und Buche. Nach HM, wo A.

limacodes keine Lebensgrundlage hat, erfolgt regelmä iz ein Zuflug, der seinen

Ursprung in mindestens 150-300 m Entfernung (Ruderal at, vermutlich jedoch in

den umliegenden Wäldern, deren Mindestabstand 800-1000 m beträgt.

Verbreitungsstrategie: r-Stratege, 3. Gruppe

Heterogenea asella (1 Individuum, ca. 45 Kokons)

Distanzen: 1-

Larvalökologie: oligophag an Laubbäumen, v.a. Buche (Fagus sylvaticus); im Ver-

gleich mit der vorigen Art mit eingeschränktem Wirtspflanzenspektrum. Die Fund-

orte der Kokons konzentrieren sich auf ein relativ kleines Areal (ea. 50 ha) im

Nordosten des Berglwalds. An anderen Stellen des Berglwaldes wurde mit gleicher

Intensität ohne Erfolg gesucht. } ®

Die im Berglwald eingestreuten Buchenenklaven (je ca. 200 m2) liegen in Abstän-

den von durchschnittlich etwa 100 m, die offensichtlich noch bewältigt werden.

Verbreitungsstrategie: K-Stratege, 6. Gruppe. Der Unterschied in der Strategie

dieser Art zur vorigen erklärt sich nicht nur durch das eingeschränkte Wistepflanzen

spektrum, sondern auch durch die Kleinheit der wohl schlecht fliegenden Imagines.

113

SPHINGIDAE

Mimas tiliae

35 Individuen 3,1 % 9- Rate

32 markiert kein Wiederfang

Wiederfang-Quote: niedrig, Stichprobe noch zu klein

Distanzen: 3-4

Larvalökologie: oligophag an Laubbäumen, das HM-Stück (1986) flog aus mindestens

800-1000 m herbei.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Laothoe populi

23 Individuen 13,0 % 9- Rate

21 markiert kein Wiederfang

Wiederfang-Quote: niedrig, Stichprobe noch zu klein

Distanzen: 3_

Larvalökologie: ne rpil nern Arten der Gattungen Salix und Populus; die Distanz

von 150-300 m wird also häufig zurückgelegt: Zufliegende Exemplare wurden an

M, SiM und WaS beobachtet. An HM auch ein 9.

Populationsbiologie: vermutlich bivoltin

Verbreitungsstrategie: intermediärer Typ, 3. Gruppe

Smerinthus ocellata

21 Individuen 14,3 % 9- Rate

20 markiert 4 Wiederfänge

Wiederfang-Quote: niedrig! Alle Wiederfange beziehen sich auf ein einziges d', das

im Garten (WaS 1988) in 5 aufeinanderfolgenden Nächten gefangen wurde, das al-

so von der Lichtwirkung gefangengehalten wurde. Stichprobe noch zu klein

Distanzen: 3 {

Parratkuingie: oligophag an Laubgehölzen, v.a. aus den Gattungen Salix und Popu-

lus. Die 150-300 m zwischen HM und dem Ruderal liegen im Bereich der norma-

len Dispersionsaktivität dieser Art. Im Ort tritt diese Art Haie auch die vorige

unregelmäßig auf; dies läßt den Schluß zu, daß Distanzen von 2 km (HM>S

nur selten geflogen werden. Vermutlich gibt es jedoch noch en Vor-

kommen dieser Ärt, die Obergrenze der "normalen" Flugaktivität (trivial move-

ment) verringert sich dementsprechend.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Herse convolvuli (4 Individuen)

Distanzen: 4 f

Verbreitungsstrategie: r-Stratege, 1. Gruppe

Sphinx ligustri (4 Individuen)

Distanzen: 3-4

Larvalökologie: die Raupe lebt in erster Linie an verschiedenen Hecken und Büschen

wie Liguster, Flieder u.s.w.. Das HM-Stück stammt also wahrscheinlich nicht aus

dem Flughafengebiet und ist mindestens 800-1000 m geflogen.

Verbreitungsstrategie: intermediärer Typ, 3. Gruppe

Hyloicus pinastri

89 Individuen 9,0 % o- Rate

86 markiert 1 Wiederfang

Wiederfang-Quote: niedrig; ein an WaN markiertes d wurde an WNo nach 4 Tagen

wiedergefunden. Die in dieser Zeit tatsächlich zurückgelegte Strecke ist vermutlich

bedeutend größer als nur die Luftlinie von 50 m.

Distanzen: 2-4 .

Larvalökologie: auf Nadelbäume spezialisiert; die Distanz von ca. 1 km (Zuflug nach

H kann in günstigen Jahren auch in größeren Stückzahlen geflogen werden. Der

Einflug 1986 nach HM deckt sich mit einem ähnlichen Phänomen beim Eulenfalter

Panolis flammea. Die als Kiefernschädlinge bekannten Schmetterlinge haben in

diesem Jahr günstige Bestandsentwicklungen durchgemacht. Daraus resultiert of-

fensichtlich eine vergrößerte Dispersionsaktivität aufgrund hoher Populationsdichte.

9 wurde ein d am Franzosenhölzl gefangen, die nächstgelegene Fichte befindet

sich ca. 500 m entfernt.

Verbräimpatzätenje r-Stratege, 5. Gruppe (in normalen Jahren); nach

SCHWERDTFEGER (1978) unternimmt H. pinastri in seinem Verbreitungsgebiet

und aus ihm hinaus Wanderungen.

114

Abb. 43b: Leucodonta bicoloria (Notodontidae; Mb, 12.6.87)

Abb. 43c: Deilephila porcellus & (Sphingidae; Garten, 12.6.85)

115

Deilephila elpenor

10 Individuen 22,2 % o- Rate

9 markiert kein Wiederfang

Wiederfang-Quote: niedrig, Stichprobe noch zu klein

Distanzen: 3-4 (nach HEYDEMANN, 1981, "0,5 - 10 km und mehr")

Larvalökologie: das HM-Stück könnte aus dem Ruderal stammen (Epilobium-Vor-

kommen) und ist mindestens 150-300 m geflogen. Neue Epilobium-Standorte sind

meist bald mit Raupen besetzt.

Verbreitungsstrategie: r-Stratege, 3. Gruppe

Deilephila porcellus

32 Individuen 32,1 % o-Rate

28 markiert 3 Wiederfänge

Wiederfang-Quote: hoch; bezogen auf den Standort HW 1987 sogar 16,7 %. Die

Fang-Wiederfang-Intervalle betrugen bei den 3 dd durchschnittlich 3,0 Tage.

Distanzen: 2(-3?

Larvalökologie: an Arten der Gattungen Galium und Epilobium; die HM-Stücke flo-

gen mindestens 150-300 m (Ruderal); diese Distanz scheint D. DOrseln: mehr oder

weniger regelmäßig zurückzulegen. Ein Sich-Entfernen vom Verbreitungszentrum

um 1000 m nach SıS konnte 1987 nicht nachgewiesen werden.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Macroglossum stellatarum

Diese Art soll bei den weiteren Auswertungen ausgeklammert bleiben, da sie bedingt

durch die Tagaktivität in Lichtfallenfängen (mit Ausnahme eines Exemplars 1983) nicht

achzuweisen ist. Es handelt sich jesenfalle bekanntermaßen um einen Wanderfalter

r-Stratege, I. Gruppe). Distanzen:

Hemaris fuciformis

Auch H. fuciformis soll aus den o.g. Gründen ausgeklammert werden, die Art gehört

vermutlich einem intermediären Strategietyp der 5. Gruppe an. Distanzen: 2-3

THYA TIRIDAE

Habrosyne pyritoides (126 Individuen)

Distanzen: 2-3

Larvalökologie: auf Brom- und Himbeere spezialisiert; die HM-Exemplare stammen

vielleicht aus dem Ruderal (Brombeer-Vorkommen), sind also mindestens 150-300 m

geflogen, was offensichtlich regelmäßig geschieht.

Verbreitungsstrategie: intermediärer Typ, 3. Gruppe

Thyatira batis (41 Individuen)

Distanzen: 2

Larvalökologie: Raupenfutierpflanzen wie H. pyritoides, interessanterweise im Offen-

land (HM) noch nie nachgewiesen. Bemerkenswert ist auch der fehlende Nachweis

an y w, obwohl gleichzeitig in 100 bzw. 120 m Entfernung 11 Stücke beobachtet

wurden.

Populationsbiologie: bivoltin

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Tethea fluctuosa (6 Individuen)

Distanzen: 1-2

Larvalökologie: monophag an Birke. Bisher wurden keine biotopfremden Tiere festge-

stellt.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Tethea duplaris (96 Individuen)

Distanzen: 2-3 .

Larvalökologie: oligophag an Erle, Birke und ap bisher wurden nur wenige bio-

topfremde Tiere festgestellt: Lediglich die beiden HM-Stücke müssen 800-1000 m

zurückgelegt haben.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

116

Tethea or (7 Individuen)

Distanzen: 2

Larvalökologie: auf Salix und Populus spec. PpEzialieient, bisher wurden keine bio-

topfremden oder in den Siedlungsbereich einfliegenden Exemplare beobachtet.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Tethea ocularis

Distanzen: vermutlich 2-3

Larvalökologie: nur an Pappeln und Espe; das im Garten nachgewiesene Stück muß

mindestens 200 m geflogen sein.

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Polyploca flavicornis

32 Individuen 11,8 % 9- Rate

16 markiert 1 Wiederfang

Wiederfang-Quote: vergleichsweise hoch, Stichprobe noch zu klein. Es wurde ein d’

(WaN nach 5 Tagen am gleichen Standort wiedergefangen.

Distanzen: 2

Larvalökologie: monophag an Birke (Betula spec.). 1989 war im Franzosenhölzl schon

in einer Entfernung von 40 m, verglichen mit dem Fangergebnis am Waldrand ein

Abundanzabfall auf die Hälfte festzustellen. Im Offenland wurde diese Art bisher

un festgestellt. Die Distanz von 1 km liegt außerhalb der normalen Dispersions-

aktivität.

Verbreitungsstrategie: K-Stratege, 6. Gruppe.

DREPANIDAE

Drepana falcataria (77 Individuen)

Distanzen: 2-3

Larvalökologie: Birke und Erle, vorwiegend an Büschen; diese Aussage, entnommen aus

dem Werk KOCHs (1984) impliziert eine vermutlich erhöhte Dispersionsaktivität, da

im Verlauf des Wachsens der Futterpflanzen desöfteren ein Standortwechsel zu

Stellen auftreten sollte, wo Jungformen von Alnus bzw. Betulus spec. vorkommen.

So wurden auch an HM 2 Exemplare festgestellt, die mindestens 1 km weit geflo-

gen waren, eines davon war ein fertiles ®; das zu einer Kolonisation fähig gewe-

sen wäre. In beiden Fällen wurde der Ortswechsel von z.T. böigen Winden unter-

stützt. Beide Fälle traten in der 2. Generation auf. Der starke Haufigkeitsabfall von

WaN nach WNo auf einer Sirecke von nur 50 m spricht jedoch eher gegen eine

höhere Austauschraten zwischen diesen beiden Standorten.

Populationsbiologie:, bivoltin

Verbreitungsstrategie: intermediärer Typ, 4. Gruppe

Drepana lacertinaria (6 Individuen)

Distanzen: 1-2

Larvalökologie: ebenfalls nur an Birken und Erlen; bisher wurden keine biotopfrem-

den Tiere festgestellt (abgesehen von vier 1989 in 40 m Entfernung zum Franzo-

senhölzl gefangenen Exemplaren).

Populationsbiologie : bivoltin

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Drepana binaria

111 Individuen 46,2 % o- Rate

88 markiert kein Wiederfang

Wiederfang-Quote: niedrig!

Distanzen: 3,

Larvalökologie: als Raupe an Eiche, Rot-Buche und Erle gebunden. An HM wurden 3

Exemplare festgestellt, die mindestens 800-1000 m geflogen waren: 2 dd‘ bei

windstillem Wetter und I 9 in einer stürmischen Nacht. h R

Populationsbiologie: bivoltin (einzelne Tiere einer 3. Generation); biotopfremde Stük-

e wurden auch in der I Generation gefunden.

Verbreitungsstrategie: r-Stratege, 3. Gruppe. Auch die starken Häufigkeitsschwan-

kungen von en zu Jahr und von Generation zu Generation deuten auf einen r-

trategen hin.

117

Drepana cultraria (92 Individuen)

Distanzen: 3

Larvalökologie: Raupenfutterpflanze: Buche, vermutlich auch Eiche. Die 3_HM-

Stücke legten also mindestens 800-1000 m zurück, in einem Fall war der Zuflug

durch hälge Winde unterstützt, an den beiden anderen Tagen herrschte windstilles

Wetter. Am Wasserwerk ist ein Häufigkeitsabfall auf 40% in ca. 100 m Entfer-

nung zum Habitat (WNw: Buchenwaldrand) zu erkennen.

Populationsbiologie:, bivoltin, biotopfremde Tiere auch in der I. Generation.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Cilix glaucata (3 Individuen)

Distanzen: vermutlich 1-2

Larvalökologie: an allen Fundorten wachsen in unmittelbarer Umgebung die Raupen-

futterpflanzen.

Populationsbiologie: bivoltin

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe; der Falter zeichnet zieh

überdies durch einen vergleichsweise plumpen, mit den Spannern (Geometridae

vergleichbaren Flug aus.

SA TURNIDAE

Eudia pavonia (3 Individuen)

Distanzen: 2 (09); 3-4 (dd); 1989 wurden zwei Jod bei einem raschen (ca. 20

km/h). sera inigen Flug über mehrere 100 Meter beobachtet.

<) stammt wohl aus dem Ruderal (150-300 m),

Larvalökologie: das HM-Stück (fertiles

orkommen von Raupenfutterpflanzen dieser

dort findet sich das nächstgelegene

Art (z.B. Weiden).

Populationsbiologie: am Licht wurden nur 99 festgestellt. Die dc’ fliegen tagsüber.

Verbreitungsstrategie: K-Stratege, 5. Gruppe; durch die große Agilität der dd‘ wird

eine gute Gendurchmischung erreicht.

LASIOCAMPIDAE

Malacosoma neustria (14 Individuen)

Distanzen: 2-3

Larvaliäningie: verschiedene Laubbäume, dementsprechend im Offenland nie beo-

achtet

Verbreitungsstrategie: r-Stratege, 5. Gruppe

Poecilocampa populi

116 Individuen 13,8 % 9- Rate

98 markiert kein Wiederfang

Wiederfang-Quote: niedrig! An HO wurde z.B. nach 46 in einer Nacht markierten

Stücken bei der Probeentnahme 2 Tage später kein einziges markiertes Exemplar

‚mehr nachgewiesen.

Distanzen: 3

Larvalökologie: durch die Raupenfutterpflanze an Laubhölzer geb den. Das an HM

SEIAngene Stück stammt zumindest aus dem Ruderal (150-300 m). Die Strecke von

km ins Flughafeninnere wird jedoch von den am Flughafenrand zahlreich vor-

handenen Exemplaren dieser Art nicht regelmäßig geflogen.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Pachygastria trifolii

117 Individuen 30,7 % 9- Rate*

109 markiert kein Wiederfang

* Ein Problem bei dieser Art, scheint zu sein, daß nur die 29 ans Licht kommen,

die gefangenen cd’ erklären sich durch indirekte Anlockung durch die Sexualphero-

mone von bereits gefangenen AR in der Falle. Darüber hinaus ist eine hohe Mor-

Bea der 99, die oft unmittelbar nach der Eiablage entkräftet sterben, zu veran-

schlagen.

118

Abb. 43d: Eudia pavonia 9 (Saturnidae; WaN, 22.4.88). Die im Tageslicht fliegenden

c’d' sind mit der Methodik des Lichtfangs nicht nachzuweisen.

119

MT — pe

Tab 61: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Pachygastria trıfolii.

SIEDLUNG HALBTROK- 2a

NRASEN M

1988 Garten Wasserwerk

SiN W

WALD HALBTROK- =

1987 ' Garten ! KENRASEN M L

SiS SIMı SN WaS,WaM WaN HO HM HW Mb aS WaN WNw WNo HO HM HW Au We

I par. - 0-0 - - - - 4 1’ WTA6 - 61 2 par. 1 - 3 - - 87 1232 - -

dd - - - = - - 3 I 37 = 47 dd 1 - - - — 4 Br 222 - -

99 = Bei him - - - 1 4 8 - 13 99 - - 3 - = 4 % 9 - -

Mark. - - = = = = 4 1 43 - 58 Mark. 1 - 3 = - 8 1128 - -

Wit. 2m 0. SIE: Se - = Wi. = = Ben re se Ste

Wiederfang-Quote: sehr gering, trotz aller methodischer Bedenken dena.) Die Flug-

zeit fällt der Hauptfangperiode im Offenland zusammen, bei Ortstreue hätten

Wiederfänge stattfinden müssen.

Distanzen: sy Außerhalb des Habitats nur selten anzutreffen: Keine Nachweise in

400 m (SiS) und 800 m (SiM) Entfernung. Schon am Rand (HO) eine deutliche

bnahme der Abundanz. Das Ex. im Garten (SiN, allerdings ein 9). sowie die

Trittstein-)Besiedelung des Wasserwerks lassen erkennen, daß in Mehrjahresin-

ver aen ausnahmsweise einmal 1 km bewältigt wird (über biotopfremdes Ge-

ıet).

Larvalökologie: P. trifolii gehört zu den Arten, die gut mit der Schafbeweidung zu-

recht kommen: Die Raupen leben zwar oberirdisch, sie sind aber durch ihre auf-

fällige Färbung gut sichtbar und werden gemieden.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Macrothylatia rubi (11 Individuen)

Distanzen: 2-3 ,

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Philudoria potatoria (8 Individuen)

Distanzen: 3; die ST WaS 1988/89 sind wohl zugeflogen. Die Population am Was-

serwerk ist als positives Resultat eines Kolonisationsversuchs innerhalb der letzten

10 Jahre zu verstehen.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe.

Cosmotriche lunigera

Distanzen: vermutlich 2

Larvalökologie: an Nadelbäume gebunden; bisher keine biotopfremden Tiere

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Dendrolimus pini (3 Individuen)

Distanzen: vermutlich innerhalb des Habitats 3, außerhalb 2

Larvalökologie: die Raupe lebt an Kiefer, Fichte und Weiß-Tanne werden nur aus-

nahmsweise angenommen; bisher keine biotopfremden Tiere beobachtet

Populationsbiologie: eine große Belat ale Populations-Wachstumsrate führt zu den

efürchteten gelegentlichen assenvermehrungen (vergleiche z.B. ODUM &

ICHHOLF, 1980)

Verbreitungsstrategie: r-Stratege, 5. Gruppe

PSYCHIDAE

Diese interessante Familie soll in der weiteren Auswertung ausgeklammert werden, da

ihre Vertreter im Gebiet (mit Ausnahme von Sterrhopteryx fusca) mit Lichtfängen

nicht nachgewiesen werden können. Die Arten dieser Gruppe haben bis auf wenige

iälen) eine flügellose 89 so daß den oft tagaktiven dd‘ (meist hohe Dispersionsaktivi-

täten) eine besondere le für die Gendurch en der Populationen zukommt. ni

entfällt bei einigen parthenogenetischen Arten (im Gebiet z. B. Dahlica triquetrella).

Für die Verbreitungsstrategie spielen vermutlich die Larvalstadien eine größere Rolle:

Die Eier werden oft direkt am Sack des 9 abgelegt, der in vielen Fällen an erhöhten

Standorten befestigt wird. Die schlüpfenden Eiräupchen "seilen sich ab” und können

durch Winde oder durch Vögel, die ım Flug den Faden und damit das Räupchen mit-

reißen, verbreitet werden.

Weiterhin zeichnen sich bisweilen die erwachsenen Raupen durch größere Mobilität aus

Die Arten sind wohl als K-Strategen der 6. Gruppe (zumindest, was die 99 betrifft)

zu bezeichnen.

In einem ca. 1 ha großen Kiefernwäldchen in der Nähe des Mallertshofer Holzes fan-

den sich Säcke von Psyche casta, Talaeporia tubulosa, Bacotia sepium und Narycia mo-

nilifera. Der Biotop ist stark isoliert und in allen Richtungen von mindestens 500 m

intensiv en Ackerland umgeben (seit über 15 Jahren). Dennoch können die ge-

nannten Ärten offensichtlich stabile Populationen unterhalten.

120

AEGERIIDAE

Auch diese Familie soll in der weiteren Auswert wegen der geringen Erfaßbarkeit

durch die angewandte Methodik unberücksichtigt bleiben.

COSSIDAE

Cossus cossus (1 Individuum)

Distanzen: vermutlich 2-3 N

Larvalökologie: zur Larvalentwicklung werden 2-4 Jahre benötigt; dies erlaubt C.

cossus keine Strategie, die eine rasche Wachstumsrate der Populationen bein-

halten würde. Da befallene Bäume des öfteren absterben, muß in der Folgegene-

ration das Aufsuchen neuer Futterquellen vorprogrammiert sein, die Polyphagie

gewährleistet dies jedoch. Es werden ältere Bäume bevorzugt.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Zeuzera pyrina (2 Individuen)

Distanzen: vermutlich 2-3

Larvalökologie: zur Larvalentwicklung werden 2-3 Jahre benötigt, siehe Bemer-

kungen zu C. cossus. Z. pyrina zeigt eine von voriger Art etwas unterschiedene

Einnischung: Es werden jüngere Bäume bevorzugt.

Verbreitungsstrategie: K-Siratege, 6. Gruppe

HEPIALIDAE

Hepialus humuli (20 Individuen)

Distanzen: 2-3, im trockeneren Offenland (HM, auch HW) noch nie beobachtet

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Hepialus sylvina

197 Individuen 22,7 % o- Rate

166 markiert 3 Wiederfänge

Tab 62: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Hepialus sylvina.

SIEDLUNG WALD HALBTROK- Dat WALD HALBTROK- "DA

1987 ! Garten ! KENRASEN M 2% 1988 Garten Wasserwerk KENRASEN M L

SiS SIMı SN WaSı WaM Wan HO HM HW Mb SN WaS WaN WNw WNo HO HM HW Au We

Soparı br 8 2 6 102-217 - 54 XZpar. 20 15 11 3 9 230225539 2,0 1 1265

Suse ls 1 - - >. 1 5 - Sr22272us} - 15 - - - - = 15

dd a7, N ee 2 SR) - 34 dd 13 24 BET NA 230 re 99

99 Sa a 3 3 sag - 22.99 1 2 As ae) - = 17

Mark. 4 - 8 84.2206 9-1 - 52 Mark. 14 26 oo... 0 20.2 & Zen MA

Wk. - 0 - a -.- 0 - - =... Wi. - 3 = 0-0 = 20-0 = - .- 3

Wiederfang-Quote: niedrig: Die 3 wiedergefangenen JG beziehen sich auf 2 Rück-

funde am Standort WaS 1988 nach 1-Tages-Intervallen und nur auf einen "ech-

ten" Wiederfang am selben Ort nach 2 Tagen (in der dazwischenliegenden Nacht

‚erfolgte kein Lichtfang).

Distanzen: 3-4

Larvalökologie: stark polyphag i

Beralalisuepin lo e: proterandrisch; im Garten fällt ein erhöhter Jco‘-Anteil auf, die

„ st jedoch auch hier bodenständig (Beobachtung eines schlüpfenden Exem-

plars). }

An HM ist der Häufigkeitswechsel 1986/1987 bemerkenswert. Auch diese Un-

stetigkeit kann als Hinweis auf eine hohe Dynamik in der Verbreitungsstrategie

dieser Art gewertet werden.

Verbreitungsstrategie: r-Stratege, 2. Gruppe

Hepialus hecta (? Individuen, in der Dämmerung einige weitere)

Distanzen: 2-3

Populationsbiologie: am Würmkanal (Au) wurden am späten Abend ca. 10 Exempla-

re beim Ausschwärmen beobachtet. Der Flug diente offensichtlich zum Auffinden

der Geschlechtspartner. Es wurde eine Kopula registriert. Die Aktivitäten be-

schränkten sich auf den Waldrand, wohingegen im Waldesinneren und auf dem

freien Feld kein Exemplar gefunden werden konnte.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

121

NOCTUIDAE

NOCTUINAE

Euxoe tritici (5 Individuen)

Distanzen: 3; das Exemplar, im Ort war vielleicht vom Verbreitungszentrum (Halb-

trockenrasen) zugeflogen (ca. I km).

Verbreitungsstrategien: r-Stratege, 5. Gruppe

Euxoa obelisca (3 Individuen)

istanzen: 3; die en im Ort waren vielleicht vom Verbreitungszentrum

Halbtrockenrasen) zugeflogen er. 1 Kar

Verbreitungsstrategien: r-Stratege, 5. Gruppe

Euxoa nigricans (? Individuen)

Distanzen: vermutlich 3; Verhältnisse wohl wie bei den vorhergehenden Arten.

Verbreitungsstrategien: vermutlich r-Stratege, 5. Gruppe

Euxoa aquilina (30 Individuen)

Distanzen: 3. Von den Verbreityngszentren (Halbtrockenrasen) her zufliegende

Exemplare gelegentlich im Ort (ca. I km), ein relativ starker Vorstoß 1983.

Verbrei strategien: r-Stratege, 5. Gruppe

Scotia segetum (18 Individuen)

Distanzen: 3-4

Larvalökologie: in früheren Zeiten oft an Getreide schädlich, durch Pestizideinsatz

nahmen die Bestände drastisch ab. Halbtrockenrasen scheinen im Untersuchungs-

gebiet Refugialstandorte darzustellen.

Verbreitungsstrategien: r-Stratege, 1. Gruppe

Scotia clavis

siehe Fernwiederfänge (8.2.) und Versetzexperiment (8.4.)

582 Individuen 17,8 % o- Rate

546 markiert 61 Wiederfänge

Tab 63: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Scotia clavıs.

SIEDLUNG WALD HALBTROK- Dez WALD HALBTROK-

1987 ! Garten ! KENRASEN M L 1988 Garten Wasserwerk KENRASEN M

SiS SIM ı SN WaSı WaM WaN HO HM HW Mb SN WaS WaN WNw WNo 2 HO HM HW Au We

Bar NZ IITEIO ST ÜNERTG> Van» Diia* Mag 1 170° "Fpar.” 27..,62% | 20, 776 320

X zus. SER — 26 = 3 21 8 38 = 96 I zus. - 76 - - - - - - = -

dd 11923 27 3 7 47 21 48 = 196 dd 25 120 719 5 26 28 41 = =

99 1 877 13 2 * 8 - 1 67 99 2 18 6 1 2 6 4 4 1 =

Mark. 2 26 29 38 4 ll 65 20 53 1 239 Mark. 267 137 12 20 7 30 31 44 1 =

Wr& morıree BR aan are E 15 wi. 2.: „SB - ji ij ’= „35 n PRRR TE

Wiederfang-Quote: BREI bEE IE: Im Garten, HO und HW mit 3-10% Wieder-

fängen, die nicht auf Gefangenschaft durch die Lichtwirkung beruhen, relativ hoch.

Garten 1987 bei den 7 do‘ eine mittlere Verweildauer von 2,7 Tagen. HO und

HW bei den Ortswiederfängen (9 od, 1 Drittfang) eine mittlere Verweildauer von

5,0 Tagen, 3 Exemplare nach 14, 9 bzw. 8 Tagen! 3

Im Wasserwerk und HM niedrige Wiederfang-Quote, hier wohl erhöhte Disper-

sionsaktivitäten.

Distanzen: 2-4; im Moos tauchten bisher nur sehr wenige Stücke auf. In feuchte-

res Gelände hinein scheint die Mobilität dieser Art drastisch reduziert zu sein.

P Kuatiombiologie: proterandrisch;, abgesehen von den beiden im Versetzexperiment

ehandelten Stücken wurden keine 99 wiedergefangen! Die Ursache liegt viel-

leicht in einer höheren Mobilität der ‘99 (siehe Sc. exciamationis)

Verbreitungsstrategien: intermediärer Ip Er ZUpBE: im Vergleich zu $S. exclama-

tionis weniger expansiv, was als Erklärung für das in Bayern etwas lokalere

Vorkommen Ü) in Frage kommt. An anderen südbayerischen Standorten ist die Art

bisweilen recht selten.

122

Scotia exclamationis

siehe Versetzexperiment (8.4.)

679 Individuen 34,2 % o9- Rate

634 markiert 23 Wiederfänge

Tab 64: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Scotia exclamationis.

mm ss, ml m mV SV ——— ———

SIEDLUNG WALD HALBTROK- DACH, WALD HALBTROK- "DA\

1987 ı Garten ! KENRASEN M 3 1988 Garten Wasserwerk KENRASEN MOOS 2

SiS SIMıSN WaSıWaM WaN HO HM HW Mb SN WaS WaN WNw WNo 2 HO HM HW Au We

Zpar. 2 37 29 3 9 3 46 31 60 - 258 2 par. 31 33 393 .4 22 62 45 69 - 4 36

zus: - 16 14 21 Be 7, = 67 I zus. _ 38 - - - - - - e = 38

dd - 21 25 5 6 29 34 25 651 - 206 og 18 34 zn 733838 64: - 2 25

9 2 15 8 9 217 146 1 - 85 29 n 35 mn I gy - 2 134

Mark. 2 35 32 23 8 45 47 30 63 I 285 Mark. 29 69 39 4 29 62 45 68 - 4 349

W.f. ae gi 2 Er = 453 = 9:3 wi 1 13 = Belle zu ee ae 14

Wiederfang-Quote: im Vergleich mit der vorigen Art deutlich niedriger; alle Wie-

derfänge, soweit sie nicht im Versetzexperiment behandelt wurden, erfolgten nach

einem Intervall von 2-3 Tagen! Im Gegensatz zum Garten wurden an den anderen

Standorten keine 99 iedergefängen. Da sich die Mortalität wohl auf einem

ähnlichen Niveau bewegt wie bei den J’c' lWiehe Versetzexperiment), scheinen hier

die 99 mobiler als die Jc' zu sein!

Distanzen: 3-4; z.T. hohe Nacht-zu-Nacht-Fluktuationen: 8 der 9 WaM-Stücke

wurden in einer Nacht gefangen; ein solch schwarmartiges Auftreten läßt ver-

muten, daß die Art hier nur zugeflogen war.

Am Wasserwerk zeit sich im Gegensatz zu S. clavis eine Häufigkeitsverteilung

wie bei den hochmobilen Arten Ochropleura plecta und Amathes c-nigrum. Man

könnte darin entlang der Waldränder ziehende Exemplare vermuten.

Populationsbiologie: proterandrisch; die 99-Rate sinkt mit zunehmendem Offenland-

charakter und ist an HM und HW relatıv niedrig.

Verbreitungsstrategie: r-Stratege, 1. Gruppe

Scotia ipsilon

261 Individuen 44,1 % 9- Rate

215 markiert 11 Wiederfänge

Tab 65: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Scotia ipsilon.

SIEDLUNG WALD HALBTROK- DAR WALD

i 1 KENRASEN MOX 2 1988 Koauen Wasserwerk

1987 ! Garten !

SiS SIM,SN WaS,WaM WaN HO HM HW WaS WeaN WNw WNo 2 HO HM HW Au We

Z par 4 86 8 21 3 4 1 6 25 1 169 Z par. 3 15 7 4 5 2 7 10 2

& zus - 8 - 8 - 1 1 - - - 183 2 zus. - 19 - = = = = - -

ds 17 467776 11 1 2 6 3 11 - 86 dd 2 15 5 1 2 1 5 6 -

99 Ser26023 13 1 2 6 3 9 1 67 99 1 15 2 3 3 1 1 4 -

Mark. 4 0 8 24 2 3 12 6719 1 149 Mark. 3 30 17) 4 4 2 6 10 -

Wir. az ee _ wie Sa le Se Fre -

Wiederfang-Quote: sehr niedrig, alle Wiederfänge erfolgten nach 1-Tages-Interval-

len und sind somit kein Hinweis auf Ortstreue! Wie auch bei den anderen Wan-

derfaltern erwies sich die mark-recapture-Methode in einer solchen "Nullprobe"

als gut brauchbar.

Je ein d' und ein 9 konnten beim täglichen Fang WaS 1988 3 Tage lang festge-

halten werden.

Distanzen: 4 ö

Populationsbiologie: oft, jedoch nicht immer konnte Proterandrie beobachtet werden

9 ab September häufiger).

Verbreitungsstrategie: Se 1. Gruppe; die Art ist an den Inpise en "Wan-

derfalterstandorten” (SiM, HW, entlang der Hecken im Dachauer Moos) besonders

häufig, vermutlich handelt es sich um bevorzugte "Straßen", wie sie bei Autogra-

pha gamma direkt tagsüber) beobachtet wurden.

123

HALBTROK- "DA

EEE DöcH 5

Ochropleura plecta

siehe Fernwiederfänge (8.2.) und verringerte Fallendistanzen (8.3.)

1803 Individuen 49,8 % o- Rate

1578 markiert 59 Wiederfänge

Tab. 66: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

i9g7 SIEDLUNG WALD

Ochropleura plecta.

HALBTROK- "DA "0° SIEDLUNG WALD HALBIROK- "DACH

KENRASEN MOos 2 1987 SEDIME MAD 7 FRENRASEN MOOS

1.6 | arten | ı Garten |!

. Gen. sis SIMISIN WaS;ıWaM WaN HO HM HW Mb 2. Gen. sis SIMISN WaS;ıWaM WaN HO HM HW Mb

a a ie u u) Re 13 EN DE 5 3 45 26 32 12

2 zus ı St Su an 76 - 1) zizus) We 2 7 En 36 (oe = = f

de men auteiies.t refr2alie 6 A de en2ze] 2 - 2 5 10 3% 3 |

99 3 - M2 2, -E —- 343 4 150 - 26 9 m %3: 20. or 7 il

Mrk. 1 - 1 au U gm El Te 10 44 Mark. 1 37 19 13. 30 41, 275338101 oe a

Wi. Su ur Eli > - Eu we u 2 er =E Tu er <

W HALBTROK- mas 1988 WALD HALBTROK- Zac |

1988 Garten. Wasserwerk KENRASEN M I 50 Garten Wasserwerk KENRASEN M |

1. Gen. SN WaS WaN WNw WNo HO HM HW Au We . Gen. sn WaS WaN WNw WNo HO HM HW Au We |

Sonara A 13 Pe 29,u 78,280 3209 m 29 8 3 163 per. 100 188 233 49 96 180 40 126 25 10 10

zus, = 9 - - - ne = —z gr 72Tzus, 4 = 2220 - - - =. 20 Ir >

oc DNA. OA ZERO EN I 719 A 66. 162° 1377 1A, 371 Les Piz?

99 1 6 > O2 7 DE, 1 Aa 38'220 8630 a2 Warn 22m

Marktes #208 126.4 IB 26:0, 309 Ri 25 8 3067 Mark 82 3691 ?206.:'42° 727 168, a7 Tom ma Au

W.t. - 4 = = - = 1 - = = 5 W.t. 1 48 1 1 - 1 - - - -

Wiederfang-Quote: sehr niedrig bei sehr kurzen Verweildauern: Im Garten erfolg-

ten, abgesehen von 5 Wiederfängen nach 2 Tagen alle Rückfänge nach 1-Tages-

Intervallen. Die niedrige mittlere Verweildauer von 1,22 Tagen ist schon durch

festgehaltene Exemplare überhöht.

Die o-Rate liegt bei den Erstwiederfängen bei ca. 50 %. Bei den durch die Licht-

wirkung über mehrere Tage EN ehaltenen Stücken (Mehrfachwiederfänge) und bei

den "echten" Ortswiederfäangen (Fangnacht-Pausen dazwischen) zeigt sich ein stark

erhöhter 9-Anteil: 80 %. Auch der HO-Wiederfang 1988 nach 3 Tagen (das

längste festgestellte Intervall!) war ein 2

In beiden Generationen hohe Dispersionsaktivitäten!

Distanzen: 4

Larvalökologie: die Raupen finden wohl überall eine Lebensgrundlage.

ulationsbiologie: bivoltin, die o-Rate schwankt in Ort und Zeit bisweilen be-

rächtlich: Im Wasserwerk aN) waren am 10.8.88 noch 65 %, am 12.8.88 nur

noch 9 % 09 zu beobachten. we en war am 2.8.88) an zwei nur 50 m vonei-

nender entfernten Standorten aN, WNo) eine 9-Rate von 37,5, % bzw. 73 %

festzustellen. Dies und eine Reihe anderer Beobachtungen rechtfertigen die Hypo-

these von geschlechispolarisiorien Schwärmen.

Eine "Popu BVISSBEESEHNLEIE hnung” im Garten vom 1.8. auf den 2.8.88 (40 bzw.

47 Individuen, 6 iederfänge) ergäbe eine "Populationsgröße" von 313 Individuen

im Einzugsbereich der Lichtfalle, die hohe Mobilität dieser Art stört derartige

Berechnungen jedoch gewaltig.

Verbreitungsstraiegie: r-Stratege, 2. Gruppe; auch das besonders häufige Auftreten

in Wanderfalternächten und das Häufigkeitsmuster mit Peaks an denselben Stand-

orten wie bei den als Wanderfalter bekannten Arten sind bemerkenswert und

wohl als weiterer Hinweis auf die hohe Mobilität dieser Art zu verstehen. Es

(siche EITSCHBERGER & STEI-

handelt sich jedoch nicht um einen Wanderfalter

NIGER, 1980]

Eugnorisma depuncta

Distanzen: vermutlich 2

Larvalökologie: die Raupenfutterpflanzen sind überall verfügbar h h

Verbrei strategie: vermutlich K-Stratege, 5. Gruppe; einen Hinweis darauf gibt

das lokale Vorkommen bzw. die Seltenheit dieser Art.

Rhyacia lucipeta (1 Individuum)

Distanzen: 4, das Stück ist vermutlich aus den Alpen zugeflogen!

Verbreitungsstrategie: r-Stratege, 1. Gruppe

Rhyacia simulans (1 Individuum)

Distanzen: 4, wie Rh. lucipeta

Verbreitungsstrategie: r-Stratege, 1. Gruppe

124

ESTER EERFEHEERERSEEEEERSEEEEREE

Noctua pronuba

siehe Fernwiederfänge (8.2.), verringerte Fallendistanzen (8.3.) und Versetzexperiment (8.4.)

1675 Individuen 44,2 % 9 - Rate

1457 Markiert 167 Wiederfänge

Tab. 67: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Noctua pronuba.

SIEDLUNG WALD HALBTROK- Dans WALD HALBTROK- DACH

1987 ! Garten ! KENRASEN M % 1988 Garten Wasserwerk KENRASEN M ”

SiS SIMıSiN WaSıWaM WaN HO HM HW Mb SN WaS WaN WNw WNo HO HM HW Au We

Ip. 9 2ı 8 74 4 23 40 26 54 - 259 Xpar. 48 48 75 9 42 193 3 5 35 A 977

zus. - 17 8 Be. Bro 70 - 3 LTzus - 40 SIDENRE re rn Ss Aal

gs 4 28 1 5° 1 18 28 10 34 - 184 de 26 403 4 47 24 60 3 37 9 - 640

99 Se 210, BE Do - 75.09 14. 7360,,,.26, „as an 63, SE Bo, ES a E78

Mark. 9 34 2 MN 4 20 37 21 49 - 257 Mark. 40 755 64 87 34 13 1 84 12 - 1200

WA. 2 TAN EUR - 3 wi. - 162 = 24 081 = 0 = ae fe 1168

Wiederfang-Quote: sehr iedde bie

Q),

Garten erfolgten nach 2 | g) und 4 Tagen. Die vereinzelten Ortswie-

derfänge, die nicht auf ein Festgehalten-Werden durch die Lichtwir zurück-

zuführen sind, deuten auf Einzelexemplare hin, die ortstreuer werden (v.a. 99).

Distanzen: 4 RD

Populationsbiologie: die. Sommerruhe, die diese Art einlegt ist nach MEINECKE

1984) ähnlich wie bei den bivoltinen Arten ein Hinweis auf hohe Dispersionsakti-

vitäten. Proterandrisch.

Verbreitungsstrategie: r-Stratege, 1. Gruppe

j 0.8. Eöperimenie); die Wiederfänge 1987 im

Noctua comes (112 Individuen)

Distanzen: 4; im Verbreitungsmuster und in den Flugnächten relativ stark mit Noc-

tua pronuba korreliert.

Verbreitungsstrategie: r-Stratege, 3. Gruppe; GYULAI & VARGA (1974) berichten

von gerichteten Bewegungen innerhalb des Areals dieser Art.

Noctua fimbriata (43 Individuen)

Distanzen: 3-4

Larvalökologie: entsprechend der Polyphagie der Raupen sind auch die Imagines

mehr oder weniger überall zu finden.

Verbreitungsstrategie: r-Stratege, 1. Gruppe

Noctua janthina (35 Individuen)

Distanzen: 3-4

Larvalökologie: Siehe Bemerkungen zur vorigen Art

Verbreitungsstrategie: r-Stratege, 3. Gruppe

Spaelotis ravida (3 Individuen)

Distanzen: 4

Verbreitungsstrategie: r-Stratege, 3. Gruppe

Graphiphora augur (3 Individuen)

Distanzen: vermutlich 3

Verbreitungsstrategie: vermutlich r-Stratege, 3. Gruppe

Paradiarsia punicea (10 Individuen)

Distanzen: 2; schon 400 m nördlich des Birkets ("Moos 1985") konnte kein Exem-

plar mehr nachgewiesen werden.

Verbreitungsstrategie: K-Stratege, 5. Gruppe

Diarsia mendica (76 Individuen)

Distanzen: 2. ; ; ;

Larvalökologie: obwohl die Raupen "niedrigen Pflanzen” leben, ist D. mendica

ziemlich stark habitatgebunden Wald). ie polarisierten Zahlenverhältnisse

WaS>SiN sowie WNw> WaN bzw. > o zeigen, daß schon in 30-100 m Entfer-

nung vom Habitat deutliche Häufigkeitsverluste feststellbar sind.

Verbreitungsstrategie: K-Stratege, 5. Gruppe

125

Diarsia brunnea (204 Individuen)

Distanzen: 2

Larvalökologie: siehe Bemerkungen zu Diarsia mendica

Verbreitungsstrategie: K-Stratege, 5. Gruppe

Diarsia rubi (22 Individuen)

Distanzen: 2-3: an HM nie beobachtet, dagegen war D. rubi 1989 am Franzosen-

hölzl eher im Freien zu finden: Zehn Exemplaren am Waldrand standen neunzehn

in einer Entfern von 40 m gegenüber.

Populationsbiologie:, bivoltin

Verbreitungsstrategie: r-Stratege, 3. Gruppe

Amathes c-nigrum |

siehe verringerte Fallendistanzen (8.3.) und Versetzexperiment (8.4.)

1460 Individuen 32,9 % 9- Rate

1255 markiert 54 Wiederfänge

Tab. 68: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Amathes c-nıgrum.

SEDLUNG WALD HALBTROK- "DA SEDLUNG WALD HALBTROK- "DACH, |

> Garten ! KINRASEN Moos 2 1987 Garten ' KENRASEN MOOS )

1. Gen. sig SIMıSN WaS,WaM WaN HO HM HW Mb 2. Gen. sis SiM | SIN Was | WaM WeaN HO HM HW Mb

Euer, Jar = - ige

N a 11 hg NE Br FE En 1 0 0 0

dd - - = - 1 2 4 - 6 - 13 dd - 46 6 8 3 8 14 13 46 1 1

99 -.- > MED ZUR SR - 3.9 - 34 3 PL E 2 3

Mark. EEE - 1 =! 4 - 7 - 15 Mark. = 99 12 7 22: 20771 3 23

W.f. a Sun. > use - 1 W.f. Sechs ENT NG- ee =

WALD HALBTROK- "DA! W HALBTROK- "DA

1988 Garten Wasserwerk NRASEN MOOS = 1988 Garten Wasserwerk KENRASEN Moos )

1. Gen. sy WaS WaN WNw WNo HO HM HW

Erna ar Tach 2m SE ar, Ba 220, wa 30 198 128 147

Zen am f a AT ee ade 1 = mie - -

dd za BI UI: 959 4.025 VER NEO dd 24 139° ‚97 ‘9 122 87°’52 ma „ee

99 hrs! Brishatsätlatjeisıvatltt 63H 29 58 12 N3_ 50,74 a2.) 20.01 aa

Mark. 13 haadomtal. ar rain: Ana nn 70% Mark. 361260 147 26 164 12 3 sb u

Wit. - 2 Sue ee 2 f. - 50 ri 7

Wiederfang te: ist als niearig

dazwischenlie

3 Tagen und

arten überwiegen sehr stark die Rückfänge nach 1-Tages-Intervallen, aus

dem Rahmen fällt lediglich ein 9 nach 4

Au We

acht

einzustufen! Nur

See fangfreier bei über 1000

o 1988 nach 2 Tagen (jeweils ein c).

aN WNw WNo HO HM

2 "reguläre"

Wie derkänse (mit

arkierungen: HW 1987 nach

HW Au We

Tagen. Die mittlere Verweilzeit betrug

hier 1988 nur 1,21

Die o-Rate in den

‚gestellten.

Distanzen: 4: in der ersten Generation wohl nur 3-4: In Jahren mit relativ ungün-

stiger Bestandsentwicklung (z.B. 1987) zeigt sich ein Fehlen in der 1. Generation

an suboptimalen Standorten, nämlich Sied ‚ HM und im Dachauer Moos. Die

Distanzen von 1 km HW>HM) und 1,4 km [WaN>WaS) scheinen, zumindest bei

solchen Konstellationen für die Individuen der 1. Generation nicht im Rahmen der

normalen Dispersionsaktivitäten zu Kesen. Das Häufigkeitsmuster HW)HO+

WaN))>HM, Siedlung, Moos und Wald (WaM, WNw) scheint in der 1. Generation

von Jahr zu Jahr konstant zu sein.

In der zweiten Generation kommt es dann zu einem Einflug an solche "subopti-

male” Standorte, besonders deulich war dies 1987 an SiM mitzuverfolgen. Jedoch

werden offenbar nicht alle Standorte gleichermaßen überschwemmt, was die signi-

fikante Häufigkeitsdifferenz zwischen SiM und SiS bei einer Distanz von nur 400

m veranschaulicht. Es werden "Zugstraßen" bevorzugt. Das im Netz der Fangstel-

u use IE Häufigkeitsmuster entspricht dem charakteristischen Wanderfal-

terbıld.

Larvalökologie: die Art findet wohl überall eine Lebensgrundlage

Populationsbiologie: bivoltin; die zum Teil sehr starken Nacht-zu-Nacht-Fluktuatio-

nen deuten auf einen Flug in Trupps oder zumindest auf besondere flugstimulie-

rende Faktoren hin. Diese Faktoren sollten eine gewisse Spezifität aufweisen, da

sich in mehreren Fällen "Häufigkeitstäler" zwischen 2 Aktivitätsmaxima genau in

solchen Nächten befanden, in denen andere Arten recht gut flogen und öfters sehr

gute Lichtfallen-Ausbeuten erzielt wurden.

Jape:

iederfängen entspricht ungefähr der in den Erstfängen fest-

126

Verbreitungsstrategie: r-Stratege, 1. Gruppe; Amathes c-nigrum kann jedoch im

Gegensatz zu manchen anderen Arten aus der Gruppe der typischen Wanderfalter

unseren Winter gut überstehen. Die starken Fluktuationen von Generation zu Ge-

neration sind für eine r-Strategie typisch.

Amathes ditrapezium

UNG WALD HALBTROK- "DA WALD HALBTROK- "DA

1987 m arten ! KENRASEN Moos 2 1988 Garten Wasserwerk KENRASEN Moos L

SiS SMıSN WaSıWaM Wa HO HM HW Mb SN WaS WaN WNw WNo HO HM HW Au We

Z par. e 6 9 16 339 18 6 3 24 132 Z par. 19 63 59 31 41 28 4 4 32 24 294

Dezusupe 002723 6 - 4 7 2 - 24 X zus. - 62 - - - - - - - - 62

dd uU. .Ö 11 - 21 7 4 3 14 88 ‘gs 16 77 34 172230 24 1 4 20 17 240

99 1 - 4 8 3 14 6 4 2 4 46 99 2 26 23 13 10 2 3 - 6 6 9

Mark. 8257210 19 I) &s 23 8 3 16 130 Mark. 17 102 57 29 38 26 4 4 26 23 326

W.f. - - - 1 > 1 - - - - 2 W.f. 1 12 1 1 4 - - - - 1 20

Wiederfang-Quote: a endortsbhängie: Im Wasserwerk relativ hoch, sonst eher nied-

rig. Die 1987 rückgefangenen Tiere waren 2 Jod‘ nach 2 Tagen. 1988 errechnete

sich für die Wiederfänge im Garten eine mittlere Verweildauer von 1,64 Tagen,

wobei es sich bei nur 2 Ausnahmen um Intervalle von | das handelte: 1 c nach

: Tagen as ein 9, an WaS markiert und nach 5 Tagen 30 m entfernt an SiN

estgestellt.

Die ER sind im Garten in den Wiederfängen deutlich überrepräsentiert! ß

Ein Festhalten durch die direkte Lichteinwirkung bei täglichem Fang findet in weit

eringerem Ausmaß statt als bei folgender Art. Die Proportionen Fang/Wieder-

ang/Mehrfachfänge liegen ähnlich wie bei Noctua pronuba, nur etwas AI

‚Die Fluchtdisposition ist deutlich größer als bei der ähnlichen Art A. triangulum

Distanzen: 2-3 (-4?), im Wasserwerk ortstreuer, sonst relativ mobil.

Populationsbiologie:, proterandrisch;

Verbreitungsstrategie: intermediärer Typ, 2. Gruppe

Amathes triangulum

siehe_Fernwiederfänge ae verringerte Fallendistanzen (8.3.) und Rückschlüsse aus

den Ortswiederfängen (8.5.

996 Individuen 24,0 % o9- Rate

919 markiert 122 Wiederfänge

Tab. 70: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Amathes triangulum.

SIEDLUNG WALD HALBTROK- "DA‘ WALD HALBTROK- "DA

1987 ! Garten ! KENRASEN Moos L 1988 Garten Wasserwerk KENRASEN MOSE L

SiS SIMıSN WaSı WaM WaN HO HM HW Mb SN WaS WaN WNw WNo 2 HO HM HW Au We

X par. 87312258 31 353 37 5 11 6 174 X par. 57 200 84 42 79 44 21212) 17 9 546

zug? 16 - 8 1a 4 - 45 % zus. - 23 - - - - - - - - 231

[oje] 46 6 33 238 38 5 6 3 141 og 37 313 63 34 656 42 1 9 14 8 567

99 4 4 4 8 1 14 13 - 6 1 54 99 16 97 27 6 17 1 1 2 3 1 170

Mark. sr1010 41 3.62! 50 6 ii 4 194 Mark. 49 408 279736572072 43 2210 17 9 725

TEE a N a ee ee ER Wi A ar 425 en Dar ann

siehe verringerte Fallendistanzen (8.3.)

512 Individuen 29,5 % 9- Rate

456 markiert 22 Wiederfänge

Tab. 69: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Amathes ditrapezium. !

Wiederfang-Quote: durchschnittlich; an HO wurde 1988 1 cd’ nach 4 Tagen rück-

gefangen. Die 99 sind im Gegensatz zur vorhergehenden Art in den Wiederfän-

‚gen unterrepräsentiert und zeigen eine kürzere Verweildauer.

Distanzen: 2-4 |

Populationsbiologie: proterandrisch; 1988 wurde an HO nach der Flugzeit der 99 am

8.7. noch ein Trupp von 18 geflogenen Jo‘ gefangen. An diesem Standort fallen

besonders starke Nacht-zu-Nacht-Fluktuationen auf, die wohl auf eine erhöhte

Dispersionsaktivität hinweisen.

Verbreitungsstrategie: r-Stratege, 2. Gruppe;

Amathes baja (240 Individuen)

form auffiel, wurde 3 Tage später wiedergefangen (in der Falle). Die gering-

Wiederfang-Quote: ein S, das an HO 1987 mit einer "Ua der Falle)“ Flügeiver-

fügige Abnormalität hatte auf das Flugverhalten höchstwa

rscheinlich keinen Einfluß.

127

Distanzen: 2-3; die Distanz von ca. | km von den Flughafenrändern zum Standort

HM (Ofteniand wird nur ausnahmsweise bewältigt, wie der starke Häufigkeits-

abfall HO/HM zeigt.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Amathes sexstrigata

siehe verringerte Fallendistanzen (8.3.)

1122 Individuen 26,5 % o- Rate

603 markiert 18 Wiederfänge

Tab. 71: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Amathes sexstrigata.

WALD HALBTROK- DACH

Wasserwerk KENRASEN M 2

1988 Garten

SN WaS WaN WNw WNo HO HM HW Au We

> para 7 B: 13921624172. Miss: 62% 82 6 - 666

zu en EM - 2 - ao 10

dc ) 10 93 47 94 107 48 60 a MEETS

99 - a "2 Mammea ea 2 18 ie

Mark. a a ee ae 52 77 603

Wf. - 6 ya AN) ae En 18

Wiederfänge am Flughafen erfolgten nach 2,

und 5 Tagen. Im Garten, einem offensichtlich eher ungeeigneten Lebensraum für

diese Art, kam es 1988 zu einem verstärkten Zu- bzw. Durchflug. Dies fand auch

seinen Niederschlag darin, daß sich die Rückfänge ausschließlich nach 1-Tages-In-

tervallen ereigneten. Die Tiere waren also von der direkten a estge-

halten worden. Die 99 sind im Wiederfang deutlich unterrepräsentiert (nur I 9

WaS nach 1 Tag).

Miederiang:Chaute, durchschnittlich; die

Distanzen: 2-3

Eopelnt biologie: deutlich proterandrisch' Im Wasserwerk (WaN) stieg in allen 3

ahren (1987-89) die Populationsdichte zwischen dem 4. und dem 10. August stetig

auf einen Maximalwert an, der einen Fang von 30-50 Individuen pro Nacht be-

wirkte. Relativ pünktlich am 20. August erfolgte dann jeweils ein ziemlich abrup-

ter Zusammenbruch der PORU Alıen: Die Lebensdauer übersteigt offensichtlich in

der Regel einen Wert von 10 Tagen nicht wesentlich. N

Merbreilmganizn ige: intermediärer Typ, 5. Gruppe; diese Art kommt in anderen

Gebieten nur sehr kokal und meist nicht häufig vor. Dies erlaubt der Art wohl

keine r-Strategie. Bei stärkeren Vermehrungen, wie 1987 und 1988 im Untersu-

chungsgebiet kann es jedoch zu Abwanderungen auch von fertilen 99 über Strek-

ken von mindestens 1 kommen.

Amathes xanthographa

1987

Z par.

% zus.

[e 102

Mark.

W.f.

siehe verringerte Fallendistanzen (8.3.)

393 Individuen 24,8 % 9- Rate

315 markiert 8 Wiederfänge

Tab. 72: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Amathes xanthographa.

SIEDLUNG WALD HALBTROK- Dacı

KENRASEN M

HALBTROK- Da

| Garten _! KENRASEN M 2 1988 Garten Wasserwerk

SIS SIMıSN WaSıWaM WaN HO HM HW Mb SN WaS WaN WNw WNo 2 HO HM HW Au We

DEsE 2 5 7.40 32...182227 2 141 Z par. 4 10 18 26 1

= a Wr EM ZUR TS - oa ae me Ss, = Set

- Ya | 1 4 24 23. 107218 1 81 dd 2 22 17 27 40 15 14 22 - -

5.71 2 4 3 13 6 5 7/ 1 47 99 1 4 2 6 15 2 1 1 - -

BÄAL5E 5 TE: 29 14 20 2 125 Mark 3 26 18,733 55 171 19023 - -

Er ehe = 2 wi. = 4 en Zu Ser Bade

Wiederfang te: relativ niedrig; die längste nachgewiesene Verweildauer beträgt

pur 2 Tage! 1988 erfolgten alle Wiederfänge im Garten nach Intervallen von 1 Tag.

istanzen:

Populationsbiologie: deutlich proterandrisch, die dd schwerpunktmäßig im August, die

28 im September. erre 3 }

Verbreitungsstrategie: r-Stratege, 3. Gruppe; A. xanthographa ist im Vergleich mit

A. sexstrigata dıe Art mit der größeren Dispersionsaktivität.

Phalaena typica (5 Individuen)

Distanzen: vermutlich 2, bisher keine biotopfremden Tiere festgestellt

Verbreitungsstrategie: vermutlich intermediärer Typ, 5. Gruppe

128

Eurois occulta (4 Individuen)

Distanzen: 2-3 & fe

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Anaplectoides prasina (51 Individuen)

Distanzen: 2-3 2

eralöKalogie oligophag, bevorzugt findet man die Falter jedoch an Stellen, wo

Him- und Brombeeren wachsen. Im reinen Offenland (Flughafen) ist A. prasina

bestimmt nicht podens and . Die beiden HM-Stücke (1986) sind daher vermutlich

aus dem Ruderal (ca. 150-300 m Entfern zugeflogen.

Die Distanzen von 1 km (Abstand von zum nächsten sicheren bodenständi-

en Vorkommen) bzw. 1,3 km, (Wasserwerk>Garten) werden nicht regelmäßig

Bewalker Die 11 WaS-Stücke sind vermutlich zugeflogen (mindestens 50 m

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Cerastis rubricosa

siehe verringerte Fallendistanzen (8.3.)

171 Individuen 4,8 % o- Rate

100 markiert 7 Wiederfänge

Tab. 73: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Cerastis rubricosa.

WALD HALBTROK- se:

198 Garten Wasserwerk KENRASEN M 2

SN WaS WaN WNw WNo HO HM HW Au We

2 par. = 1 38 24 25 Os] 4 S - 103

zus. - 1 Su SP che Ze 1

[0202 = 2 34 24 24 10 1 4 = = 99

99 - — 4 = 1 = - = = = 6

Mark. - 2 37 22 24 10 1 4 - > 100

Wi. Su 2) 4 en, Erg 7

Wiederfang-Quote: hoch, bei langen Verweildauern, die siedcch - zumindest teil-

weise - durch kühlere Witterung bedingt sein dürften. Vielleicht suchen die Ima-

gines ähnlich wie die Arten der Gattung Orthosia nach dem Schlüpfen Nektar-

quellen und werden an Standorten mit blühenden Weiden "seßhaft". Ein "Einflug"

in den Siedlungsbereich und in das Gebiet des Dachauer Mooses kommt jedoch in

nur geringem Maß vor, wodurch im Gegensatz zu den Orthosien ein deutliches

Häufigkeitsgefälle entsteht.

Distanzen: ? >»

Larvalökologie: polyphag an niedrigen Pflanzen

Populationsbiologie: ın allen Jahren tauchten bisher im Juni frische Exemplare auf,

es handelt sich jedoch wohl nicht um eine zweite Generation.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Cerastis leucographa (23 Individuen)

Distanzen: ?; die Distanz von ca. 1 km ins Offenland hinein (HM, auch HW) wird

anscheinend nur selten bewältigt, an den genannten Stellen keine Nachweise

Verbreitungsstrategie: K-Stratege, 6. Gruppe; die gesammelten Daten deuten auf ein

ähnliches Verhalten wie bei C. rubricosa hin, C. leucographa scheint lediglich

etwas stärker habitatgebunden zu sein.

Mesogona oxalina (15 Individuen)

Distanzen: 2, :

Eeralessioge: die Raupe lebt SUBEnEE an verschiedenen Laubgehölzen, v.a. Wei-

den und Pappeln; die HM-Stücke stammen daher sehr wahrscheinlich aus dem

150-300 m entfernten Ruderal. Diese Distanz wird mit erstaunlicher Konstanz

bewältigt, 1 km jedoch so gut wie nie: We>Mo, HM>HO und HM>HW,; an den

letztgenannten Standorten konnte die Art nicht nachgewiesen werden.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

HADENINAE

Discestra trifolii (28 Individuen)

Distanzen: 4 \

Verbreitungsstrategie: r-Stratege, 1. Gruppe; in dieses Bild passen auch die Häufig-

keitsschwankungen von Jahr zu Jahr, das den anderen Wanderfaltern entsprechende

Häufigkeitsmuster innerhalb des Fangstellen-Netzes sowie das gleichzeitige Auf-

treten mit diesen anderen Arten in besonderen "Wanderfalternächten".

129

Polia bombycina

70 Individuen 19,0 % 9 - Rate

62 markiert 2 Wiederfänge

EEE ee: relativ hoch; es handelt sich um zwei dd, einen Ortswieder-

Ians an o nach 3 Tagen und einen Ortswechsler von WaN>WNo (50 m) nach

agen.

Distanzen: 2-3

Larvalökologie: die HM-Stücke könnten zu einem Verbreitungszentrum im Ruderal

mit Weide als bevorzugter Futterpflanze gehören, eine eindeutige Aussage ist

wegen der Oligophagie dieser Art nicht möglich.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Polia nebulosa (38 Individuen)

Distanzen: 2-3

Larvalökologie: etwas mehr an bebuschtes Gelände gebunden als vorige Art, bio-

topfremde Tiere wurden bisher nicht beobachtet.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Pachetra sagittigera

114 Individuen 9,9 % o-Rate

110 markiert 2 Wiederfänge

Tab. 74: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Pachetra sagıttigera.

SIEDLUNG WALD HALBTROK- "DA! WALD HALBTROK- "DA

1987 ' Garten ' KENRASEN Moos 2 1988 Garten Wasserwerk NRASEN MOosı

SIS SIMıSN WaSıWaM WaN HO HM HW Mb SN WaS WaN WNw WNo 2 HO HM HW Au We

Ipaı. - - - - - - 6 36 42 - 82 Xpar. -

dd u Pre 49327436 - 72 dd “

KL u ae *= I 2, 4 = 7...%99 2

Mark. - = u - = = 6 34 40 - 79 Mark. -

W.f. - - - = = = - - 2 - 2 WE

Wiederfang-Quote: durchschnittlich, es handelt sich um zwei Ortswiederfänge (do)

an HW nach jeweils 2 Tagen.

Distanzen: im Habitat 3, außerhalb 2

Larvalökologie: pokyplias an niedrigen Pflanzen und Gräsern! Dennoch ist eine

starke Habitatbin (Halbtrockenrasen) im Untersuchungsgebiet festzustellen:

ußerhalb der drei Flughafen-Standorte wurde diese Art bisher nur 1989 an er

20) beobachtet. Schon in einer 30 m tiefen Einbuchtung in den Wald (Ho ist

_ je weise konstant - ein starker Häufigkeitsabfall festzustellen. Im Flughafen-

gebiet ist P. an wohl allgemein verbreitet, in 300-400 m Entfernung (SiS)

war 1987 kein Stück nachzuweisen.

Populationsbiologie:, proterandrisch

Verbreitungsstrategie: K-Stratege, 5. Gruppe

Sideridis albicolon (6 Individuen)

Distanzen: im Habitat 2-3, außerhalb 1-2

Larvalökologie: Auch hier wäre durch die (angebliche) Polyphagie der Raupen die

Lebensgrundlage in einem weitaus größeren Areal gegeben als re be-

setzt wird. $. albicolon ist streng habitatgebunden (Steppehheidechar kun und

war schon an HO nicht mehr nachzuweisen.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Heliophobus reticulata (16 Individuen)

Distanzen: 2

Larvalökologie: oligophag an einigen Carophyllaceen, wie die beiden vorigen Arten

strenger habitatgebunden, als dies von den weiter verbreiteten Raupenfutterpflan-

zen her "nötig wäre”.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Mamestra brassicae (63 Individuen)

Distanzen: 3-4

Populationsbiologie: bivoltin

Verbreitungsstrategie: r-Stratege, 1. Gruppe

130

Mamestra persicariae (146 Individuen)

Distanzen: 3-4; die deutliche Häufigkeitsdifferenz zwischen WaN und WNw (nur 100

m voneinander entfernt) weist dennoch auf Barrieren gegen die Verbreitung hin: In

Wäldern ist M. persicariae seltener anzutreffen.

Verbreitungsstrategie: r-Stratege, 3. Gruppe

Mamestra contigua (20 Individuen)

Distanzen: 3

Larvalökologie: M. contigua-Raupen sind relativ polyphag; vermutlich bilden jedoch

für die an HM nachgewiesenen Stücke die eiden des Ruderals (150-300 m

entfernt) die Lebensgrundlage. 1988 befand sich darunter auch ein fertiles 9. In-

teressanterweise konnte in 1 km Entfernung (HO) über 3 Jahre hinweg kein

Exemplar festgestellt werden.

Populationsbiologie:, bivoltin

Verbreitungsstrategie: r-Stratege, 3. Gruppe

Mamestra w-latinum

43 Individuen 24,4 % o9- Rate

41 markiert 2 Wiederfänge

Wiederfang-Quote: durchschnittlich: ein 9 wurde an WaS 1988 am nächsten Tag

wiedergefangen, war also nur vom Licht festgehalten worden, dazu kommt ein

"echter" Wiederfang 1987 HW nach 2 Tagen ()

Distanzen: 3; unter den im Untersuchungsgebiet festgestellten Mamestren wohl die

Art mit der größten Habitattreue (trockene Wiesen). Die im Garten vereinzelt zu

beobachtenden Stücke sind vielleicht zugeflögen, auch die (Trittstein-)Besiedlung

des Wasserwerks läßt vermuten, daB M. w-latinum-oo9 bisweilen Distanzen von

1 km zurücklegen. }

Populationsbiologie: bivoltin, proterandrisch

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Mamestra thalassina (77 Individuen)

Distanzen: 2-3

Larvalökologie: M. thalassina ist zwar eine polyphage Art, bevorzugte Futterpflan-

zen sind jedoch Laubgehölze und -gebüs N ım reinen Offenland ist diese Art

wohl nicht bodenständig. Das HM-Stück (6 ist vielleicht vom Ruderal her zuge-

flogen (200-300 m).

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Mamestra suasa

371 Individuen 32,2 % o- Rate

93 markiert 3 Wiederfänge

Wiederfang-Quote: sehr niedrig! Alle drei Wiederfänge (dT‘9) erfolgten an WaS

1988 nach 1 Tag und stellen somit keinen Hinweis auf ein natürliches Verbleiben

in der Umgebung dar. Hier waren 31 Individuen markiert worden.

Distanzen: 4 _ h

Populationsbiologie: bivoltin; in der Siedlung wurde die erste Generation von M.

suasa nur sehr vereinzelt beobachtet, es gilt analog das bei Amathes c-nigrum

Gesagte, vor allem für den Fangplatz SiM. Die Weibchen-Rate der 2. Generation

liegt höher als die der ersten.

Au alle ist die besondere Korrelation der Anflugaktivität mit den sogenannten

Wanderfalternächten. Dies ist ein weiterer Hinweis darauf, daß auch einige andere

hochmobile Arten "mitfliegen".

Verbreitungsstrategie: r-Stratege, 2. Gruppe

Mamestra oleracea (88 Individuen)

Distanzen: ad

Larvalökologie: Wenn die Angabe in KOCH (1984) stimmt, daß die Raupen (nur)

Garten und Feldgewächse fressen, müßte das HM-Exemplar (d) 1986 zumindest

vom Gut Hochmutting her zugeflogen sein (ca. 500-600 m). Interessant ist auch

der starke Häufigkeitsuntersched WaN-WNw, der für Barrieren gegen die freie

Beweglichkeit dieser Art spricht.

Verbreitungsstrategie: r-Straiege, 3. Gruppe

Mamestra pisi (4 Individuen)

Distanzen: 3 (-4?)

Larvalökologie: die HM-c’S 1988 entwickelten sich vermutlich 150-300 m entfernt

an den Weiden des Ruderals zum Falter.

Verbreitungsstrategie: r-Stratege, 5. Gruppe

131

Hadena rivularis (10 Individuen)

Distanzen: 3

n die Raupe lebt an verschiedenen Nelkengewächsen, die in der Nä-

he aller Fundorte vorkamen.

Populationsbiologie: bivoltin

Verbreit ae ei r-Stratege, 3. Gruppe; ähnlich wie wohl bei allen Vertretern

dieser Gruppe der "Nelkeneulen” wird auch hier ein vorprogrammiertes Umher-

streifen der 99 zur Aufsuche der nicht immer standorttreuen Futterpflanzen nötig

sein.

Hadena lepida (10 Individuen)

Distanzen: 3

Larvalökologie: wie vorige Art; bei dem frischen J, das 1986 im Garten gefangen

wurde, könnte es sich um einen Nachkommen eines für die Dauer eines Jahres

geglückten Kolonisationsversuches handeln: Seit 1985/1986 wächst im Garten ei

sich von Jahr zu Jahr ausbreitender Bestand von Roten Lichtnelken (Siene dien.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Hadena compta (5 Individuen)

Distanzen: 2-3

Larvalökologie: die Grundlage für die Konstanz des Auftretens im Garten in den

letzten Jahren liegt wohl in den Beständen der Roten Lichtnelke.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe; nach LATTIN (1967) unternahm

H. compta in diesem Jahrhundert Arealausweitungen.

Hadena confusa

Distanzen: vermutlich 2-3

Verbreitungsstrategie: vermutlich intermediärer Typ, 5. Gruppe

Hadena bicruris (5 Individuen)

Distanzen: 2-3

Larvalökologie: im Garten ist gleichzeitig mit dem Erscheinen der Bestände der

Roten Lichtnelke seit 1986 auch regelmäßig H. bicruris nachgewiesen worden. Die

allen: erfolgte erstaunlich rasch, wenn man bedenkt, daß vorher nie ein

zugeflogenes) Exemplar festgestellt wurde.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Lasionycta nana (1 Individuum)

Distanzen: vermutlich 2-3 £

Verbreitungsstrategie: vermutlich intermediärer Typ, 5. Gruppe

Eriopygodes imbecilla (1 Individuum)

Distanzen: 1-2

Larvalökologie: Die Raupe lebt an "niedrigen Pflanzen und Gräsern". Trotz dieser

breitgefächerten Lebensgrundlage handelt es sich in Südbayern um ein sehr lokal

vorkommendes Glazialrelikt.

Verbreitungsstrategie: K-Stratege, 6, Gruppe; ein neu geschaffenes_(hypothetisches)

Feuchtgebiet im Siedlungsbereich (3,8 km entfernt) wäre in der Zeit der. bisheri-

gen Lichtfallenerhebungen im Garten (ca. 10 Jahre) höchstwahrscheinlich noch

nicht besiedelt worden, auch wenn es noch so naturnah gestaltet worden wäre.

Cerapteryx graminis (1 Individuum)

Distanzen: vermutlich 2-3

Larvalökologie: an Graswurzeln, gern an Sauergräsern, die Futterquelle wäre also

reichlich vorhanden; dennoch ist diese Art in der unteren Hochebene Südbayerns

eine recht lokale Erscheinung.

Verbreitungsstrategie: vermutlich intermediärer Typ, 5. Gruppe

Tholera cespitis (1 Individuum)

Distanzen: vermutlich 2-3

Larvalökologie: an Gräsern, vorwiegend an den Wurzeln; dadurch spielen Flächen

wie das Flughafengebiet, das extensiv bewirtschaftet wird (Schafweide), eine be-

Seutende „ER le für diese Art, die wohl durch den Verbiß nicht besonders ge-

stört wird.

Verbreitungsstrategie: vermutlich intermediärer Typ, 5. Gruppe

132

Tholera decimalis

282 Individuen 6,3 % o- Rate

183 markiert 4 Wiederfänge

Tab. 75: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Tholera decimalis, 1988 wurden nur wenige Stücke markiert.

SIEDLUNG WALD HALBTROK- "DA

1987 ' Garten ! KENRASEN Moos L

SiS SIMıSN WaSıWaM WaN HO HM HW Mb

par. - 3 227767 7,5377,667740 1 173

2 zus pr - 1 Se = - 1

dd 3 9 2 8.80 1 164

99 = 0.9 = = 1 23 4 = 10

Mark. Ne 2 3 2 6 53 66 40 1 173

Wit. au ao = = 1 1 Ai = = 4

Wiederfang-Quote: durchschnittlich; es handelt sich um 4 dc, das WaN-Stück

wurde nach 4 Tagen, die drei Flughafen-Exemplare nach je 2 Tagen wiederge-

fangen. Die mittlere Verweilzeit errechnet sich also zu relatıv niedrigen 2,5 Tagen.

Für eine Offenlandart liegen diese Wiederfang-Daten jedoch vergleichsweise hoch.

Distanzen: SS 3-4, 09 2-3

Larvalökologie: siehe Bemerkungen zu, voriger Art; auch die Raupen dieser Art

bevorzugen eher trockene Wiesen, sie wurde beispielsweise 700 m vom Garten

entfernt in einer Wegwarten-Gesellschaft (denken intybi) gefunden. Unter

Umständen ist ‚T. decimalis im eigentlichen Ortsbereich nicht bodenständig.

Populationsbiologie: die schwerfällig wirkenden und fliegenden 99 sind vermutlich

ortstreuer als die dd; dieses verminderte Flugvermögen kann auch für die ge-

ringe o-Rate in der Lichtialle verantwortlich sein.

Die Tatsache, daß im Ort noch nie ein 99 gefunden wurde, mag als weiterer

Hinweis darauf gewertet werden, daß es sich bei den Siedlungs-Stücken nur um

zufliegende cd handelt. n

en intermediärer Typ (nach SPITZER et al., 1984 nahe einer

K-Strategie), 3. Gru Bei die starke Vermehrung 1986 im Flughafengebiet (eine

Fangnacht mit über 180 Exemplaren an HM) war von einem ebenfalls starken

Flug im Garten begleitet, was wohl zumindest teilweise auf abwandernde Indivi-

duen zurückzuführen ist.

Panolis flammea

85 Individuen 16,7 % o- Rate

83 markiert kein Wiederfang

Wiederfang-Quote: sehr niedrig

Distanzen: 4

Larvalökologie: an Nadelhölzer gebunden (v.a. Kiefer), die HM-Stücke sind daher

mit Sicherheit mindestens 1 km von den Flughafenrändern her zugeflogen. Diese

Distanz scheint für P. flammea keinerlei Verbreitungshemmnis darzustellen, sie

wird zwar nicht Tepe ann bisweilen jedoch in größeren Anzahlen und auch von

fertilen 09 (1987 I 9, 1986 war keine Geschlechtsbestimmung durchgeführt wor-

den) geflogen! Die festgestellte Expansion 1986 fällt mit einer besonders guten

Bestandsentwicklung vieler Kiefernarten in diesem Jahr zusammen.

Auch bei kleineren Fallendistanzen (Wasserwerk) sind im 100 m-Bereich keine

Häufigkeitsgradienten festzustellen. .

Derbreisunganiralegie. r-Stratege, 3. Gruppe; im Gegensatz dazu steht die Beobach-

tung SCHWERDTFEGERS (1978) einer relativ langsamen Wiederbesiedlung eines

künstlich "entleerten" Waldabschnitts.

Orthosia cruda

175 Individuen 33,1 % 9 - Rate

173 markiert 2 Wiederfänge

Tab. 76: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Orthosia cruda.

SIEDLUNG WALD HALBTROK- "DA =

KENRASEN Moos 2 KENRASEN Mass

WALD

198 Garten Wasserwerk

1987 ! Garten !

SiS SIMıSIN WaSıWaM WN HO HM HW Mb SIN WaS WeN WNw WNo HO HM HW Au We

Zpa. - 8 14 ER I ee I) = 78 I par. 10 1 Ss er], 2 26

DEzus un A ee REN WE - 5 5 - SE

ds a am BIN Mei ES; Ba - 53 ES re Sa TE ee.

99 2.808 SIRURSMR DM ABIT NE GIER - son. ar Re

Mark - 8 14 30 2 weg 000g - N REN LE REN En

133

Wiederfang: Oiyote: niedrig! Nur im ern: etwas höher, 1987 wurde ein 9

nach 2 Tagen, 1988 ebenfalls ein 9 nach 9 (!) Tagen wiedergefangen. Die lan-

e Verweildauer sowie die deutliche Überrepräsentierung im iederfangergebnis

Tasten erkennen, daß die o9 im Vergleich zu den d’d etwas ortstreuer sind.

Distanzen: oo 2 (-3?), dd 3-4 ei i

Larvalökologie: an verschiedenen Laubbäumen, die nach HM zugeflogenen Stücke

Yan fertile 99!) stammen mindestens aus dem 150-300 m entfernten Ruderal,

teilweise wohl auch von den Flughafenrändern () I km).

Populationsbiologie: proterandrisch

Verbreitungsstrategie: intermediärer Typ, 3. Gruppe

Orthosia populi

37 Individuen 14,3 % o- Rate

34 markiert kein Wiederfang

Wiederfang-Quote: niedrig, Stichprobe noch zu klein; im Moos wurden am Stand-

ort We 1988 31 Individuen markiert, die fehlenden Rückfänge erklären sich durch

den im Frühling an diesem Ort nur wöchentlich betriebenen Lichtfang. Hier könnte

die Art dennoch relativ ortsfest sein.

Distanzen: 2-3 #

Larvalökologie: an Espe und Schwarzpappel, die nächstgelegenen Futterpflanzen

liegen von HM ca. 1 km, vom Garten ca. 500 m entfernt. Diese Distanzen liegen

nicht im normalen Bereich der Dispersionsaktivität von O. populi (siehe Häufig-

keitsdifferenz zu "We”), werden jedoch ohne größere An, me bewältigt. Im Ort

1988 auch ein fertiles 9! Die Distanz von 2 km (We>Au), ausgehend vom Ver-

breitungszentrum konnte 1988 offensichtlich nicht zurückgelegt werden.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Orthosia gracilis

31 Individuen 60,0 % 9 - Rate

30 markiert kein Wiederfang

Wiederfang-Quote: niedrig, Stichprobe noch zu klein.

Distanzen: 2-3

Larvalökologie: Durch die Raupenfutterpflanzen eher an feuchte ls gebun-

den, nimmt jedoch auch Schafgarbe (Achillea millefolium) und Beifuß (Artemisia

spec.).

Verbreitungsstrategie: intermediärer Typ, 4. Gruppe

Orthosia stabilis

192 Individuen 30,5, % .9;> Rate

187 markiert 4 Wiederfänge

Tab. 77: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Orthosia stabiılis.

SIEDLUNG WALD HALBIROK- DACH _— HALBTROK- "DA

G Dass 5 ENRASEN MOOSE

WALD

1987 „Garten | 1988 Garten Wasserwerk

SIS SIM!SIN WaS!WaMWaN HO HM HW Mb SiN WaS WaN WNw WNo HO HM HW Au We

Zpa. 3 7 322 10113 SIGRSINE- 3 BAER > par A15u12 air, 2 5 ee!

2 An "aaa ELLE. Pur, E Yansz 10: | Van re Fer ea es

ds 1 4 20 a = 54 09 Eiuhi- en EI U 018 En:

99 ai Be u, 3 36

Mk: 4. 2 231 6 7 2

Mark, 37 3 Be We 3 89 we r r SEA Mn 7 62.5

f. ee eh Kalle = 1

VWiederinugrtiunte: durchschnittlich, die mittlere Verweildauer betrug 3,0 Tage; aus

es Rahmen fällt das 9 im Wasserwerk mit einem Rückfang-Intervall von 5

agen.

‚Die 09 sind im Wiederfangergebnis mit 50 % vertreten.

Distanzen: 3-4; die Häufigkeitsdifferenz WaN/WNw spricht dennoch für Hemmnis-

se in der Beweglichkeit dieser Art.

Larvalökologie: unter den von KOCH (1984) aufgezählten Futterpflanzen befindet

sich keine, die am HM-Ruderal vorkäme. Die mit hoher Konstanz zufliegenden

Stücke (auch 99) hatten also mindestens 800-1000 m vom Flughafenrand her zu-

rückgelegt. Das Ruderal wird offensichtlich nur zur Nektaraufnahme der Imagines

aufgesucht. \

Populationsbiologie: proterandrisch

Verbreitungsstrategie: r-Stratege, 2. Gruppe

134

Orthosia incerta

269 Individuen 25,1 % o- Rate

265 markiert 8 Wiederfänge

Tab. 78: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Orthosia incerta.

SEDLUNG WALD HALBTROK- "DA WALD HALBTROK- "DA

1987ER NGaztene! KENRASEN MOOS" 2 1988 Garten Wasserwerk KENRASEN Moosr 2%

SiS SIMı SN WaSıWaM WaN HO HM HW Mb SN WaS WaN WNw WNo 2 HO HM HW Au We

Z par. 4 29 48 8 6 22 15 8 10 4 153 X par. 5 5 2 1 ORE22 16 1 5 1 15 107

a Re nn an Sa ee mache U & Yale age 1 10% re

[o1°7 4 20 41 12 a 11 6 7 3 122 99 1 = 6 4 8 3 & 2 E 4 28

Mark. 4 29 48 14 5 24 15 8 10 4 161 Wi. = = 3 = 1 z & pi = = a

Wit. See Frag - 0-0 = - 4

DE naz note relativ hoch (vor allem im Wasserwerk), bei einer recht langen

mittleren Verweildauer von 4,25 Tagen! oo-Rate im Wiege fans erschuts USERS:

Beide o-Rückfänge en schon nach einem 2-Tage-Intervall.

Im Wasserwerk 1988 (verringerte Fallendistanzen) betrug die Wiederfang-Quote

sogar 7 % bei einer mittleren Verweildauer von 4 Tagen.

Gemessen am Wiederfangergebnis handelt es sich hier um die ortstreueste der im

‚Gebiet beobachteten Orthosien. __ _ f

Distanzen: 2-3; der deutliche Häufigkeitsgradient WNw>WaN spricht auch für

Barrieren gegen die freie Beweglichkeit dieser Art. '

Larvalökologie: im Vergleich zu den anderen Orthosien relativ Bolyph g, die HM-

Stücke stammen aber wohl zumindest aus dem Ruderal (200- 0 m Diese Di-

stanz scheint auch von 99 ohne Probleme bewältigt zu werden.

Populationsbiologie:, proterandrisch

Verbreitungsstrategie: intermediärer Typ, 4. Gruppe

Orthosia munda

26 Individuen 23,1 % o- Rate

26 markiert kein Wiederfang

Wiederfang-Quote: niedrig, Stichprobe noch zu klein

Distanzen: 2-3

Larvelbkalosis: das 1987 an HM festgestellte J stammte mindestens aus dem Ru-

deral (150-300 m entfernt).

Populationsbiologie: proterandrisch

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Orthosia gothica

siehe verringerte Fallendistanzen (8.3.)

835 Individuen 32,3 % o- Rate

814 markiert 15 Wiederfänge

Tab. 79: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Orhosia gothica.

SIEDLUNG WALD HALBTROK- "DA WALD HALBTROK- "DA

1987 ' Garten ! Moos X 1988 Garten Wasserwerk KENRASEN MOOS ®

—____ Beh) VESmTEN le) ilelmin 2 ln Sn Was TE WENMNWEWNOL HOSEN SEN SRauWe

X par. 30 25 118 37 10 121 SEE 13 421 2 par. 39 37 86 60 7 37 7 16 8 21 381

% zus Simianise 19 Seo - .- - = 32 X zus. au 1 = > - EHE = = 1

00, 200,168 181,62.00.46 5, 92,.00.5128. 4.7, 2120.u1,9.....204 5.4 11.224.130, 15,66% -36,4,47,,4-29] 0668 snas 5,116) 266

99 15 73 8 5 61 94 6 4 143 09 17 ER 300 230 Zar DE ses

Mark. 30 25 116 62° 137 131 a ei 13 445 Mark. 39 35 si 58 69 36 DEN 8 21 369

WDR = NEL Bean SE SE were ISIN ee 0

Wiederfang-Quote: standortabhängig: Im Wasserwerk hoch, sonst recht niedrig. Die

99 sind im Wiederfangergebnis mit 2/15=13,3 % etwas unterrepräsentiert; deren

nenreildauern liegen mit jeweils 2 Tagen auch unter denen der dd (durchschnitt-

ich 5,5 Tage).

135

Im Garten wurde 1988 ein J, das an SiN markiert wurde nach 4 Tagen 30 m

entfernt an WaS wiedergefunden. In diesem Intervall lagen 3 gute Flugnächte für

O. gothica. Bisweilen scheint jedoch kühle Witterung für ein verlängertes Ver-

Bleiheii im Gebiet verantwortlich zu sein.

Distanzen: 3; für die 99 ist vielleicht eine höhere Dispersionsaktivität zu veran-

schlagen.

Larvalökologie: unter den Arten der Gattung Orthosia vergleichsweise polyphag;

jedoch an HM wohl nicht bodenständig, die o9-Rate liegt hier im gleichen Be-

reich wie im Gesamtergebnis.

Populationsbiologie: deutlich proterandrisch

Verbreitungsstrategie: intermediärer Typ, 2. Gruppe; wie auch bei einigen anderen

Orthosien scheint an Standorten mit großem Nektarangebot für die Imagines (blü-

hende Weiden) die Ortstreue erhöht zu sein. An den anderen Stellen (z.B. im

Garten) werden verstärkt solche Individuen erfaßt, die sich auf "Suchflügen"”

befinden.

Die Dispersionsaktivitäten der o9 liegen offensichtlich auf etwas höherem Ni-

veau als bei den dd‘. Da es für die 99 wohl keine Schwierigkeiten bereitet, die

mehr oder weniger ubiquitär verbreiteten nen ten flanzen zur Eiablage zu

suchen, scheint es sich um Suchflüge nach Nektarquellen zu handeln, um eine

optimale Nährstoffversorgung für das Eiablagegeschehen zu gewährleisten; in den

noch etwas kühlen Nächten der Flugzeit er Art dürfte überdies auch der

Wärmeverlust eine bedeutendere Rolle spielen als bei anderen Arten.

Mythimna turca (133 Individuen)

Distanzen: 2; das HM-Stück (4) stammt vielleicht aus dem Ruderal (150-300 m

entfernt), sonst konnten bisher keine biotopfremden Tiere festgestellt werden. Der

Häufigkeitsgradient SiN> WaS (30 a

persionsaktivität der an WaS gefangenen Tiere sprechen.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Mythimna conigera (71 Individuen)

Distanzen: 3

Larvalökologie: die Raupe lebt an "Gräsern und niedrigen Pflanzen”, die Art ist, im

Gebiet an trockenere Wiesen gebunden. Die 3 im Garten festgestellten Tiere (das

Stück 1987 war ein d) sind wahrscheinlich von den Verbreitungszentren (Flugha-

fen, Wasserwerk) zugeflogen, haben demnach 1-1,3 km zurückgelegt. Betrachtet

man auch das stark polarisierte Häufigkeitsverhältnis WaN/ WNw. so scheinen

Distanzen von ca. | km schon eher an der Obergrenze der normalen Dispersions-

aktivität zu liegen.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Mythimna ferrago (63 Individuen)

Distanzen: 3 (-4?) _

Verbreitungsstrategie: r-Stratege, 3. Gruppe

Mythimna albipuncta (23 Individuen)

Distanzen: 4

Verbreitungsstrategie: r-Stratege, 1. Gruppe; auch das Haufigkeitsmuster innerhalb '

des betriebenen Fallennetzes entspricht dem der anderen typischen Wanderfal-

terarten.

Mythimna vitellina (1 Individuum)

Distanzen: 4 5

Verbreitungsstrategie: r-Stratege, 1. Gruppe

Mythimna pudorina (5 Individuen)

Distanzen: 2-3 # R

Larvalökologie: die Raupe ist an "Schilf und nach der Überwinterung an Moorgrä-

ser" gebunden. An HM wurden 2 höchstwahrscheinlich aus dem Ruderal (150-300

m entfernt) stammende do‘ gefangen. Das HO-c flog zumindest vom mit Schilf

bestandenen Schloßkanal her zu 1200 m), die Lebensgrundlage der Garten-Exem-

lare (Jo liegt unter Umständen in der Begleitvegetation von Gartenteichen

[mindestens 200 m), das WNo-Stück en entwickelte sich direkt am

orst

angplatz an einem 1 m? großen Schi wenn es nicht vom nächsten

Standort der Futterpflanze (ca. 1 Ki) her zuflog. Der Schilfbestand befand sich -

erst seit 2 Jahren an dieser Stelle, eine Kolonisation über (mindestens) 1 km kann

also im Einzelfall recht schnell erfolgen.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

136

könnte ebenfalls für eine geringe Dis- |

Mythimna impura

siehe verringerte Fallendistanzen (8.3.)

439 Individuen 20,6 % o- Rate

424 Markiert 25 Wiederfänge

Tab. 80: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Mythimna impura.

SIEDLUNG WALD HALBTROK- In WALD HALBTROK- DAS

1987 | Garten ! KENRASEN M 2% 1988 Garten Wasserwerk KENRASEN M

SiS SIM ı SN WaSı WaM WaN HO HM HW Mb SN WaS WaN WNw WNo 2 HO HM HW Au We

2 par. SEr2EEE6) U 1a 70 21 6 6 2 136 2 par. 4 1 VER37 98 19 gl 5 3

2a 2. oo 1 zog 89. & = A an —< 17 SER FEN Re =. 00,.°8 =

dd 3 2.8 5 95552 205 8 1 112 dd 3 2i ET el, Ze 18 9 8 3 3

99 Se San 30 21 mEEb ae 1 38 09 1 7.07 19:.,.,06%, Na nel po

Mark. b2=336 STRBI2E73 25 9 6 1 147 Mark. 4 28 75 35 90 19 9 10 4 3

Wif. =. = Ei Doz - 6 wi. - 004 Salem Tach San 00m

Wiederfang-Quote: sehr hoch bei langen Verweildauern, ein x (1987) sogar

erst nach 27 Tagen. Im Garten nur kurze en 1-2 Tage), hier und

vielleicht auch am Flughafen (außerhalb des Habitats) etwas mobiler. Die sind

mit 1/25= 4 % im Wiederfangergebnis deutlich unterrepräsentiert, der einzige

‚9-Wiederfang erfolgte auch WaS 1988 nach 1 Tag, also irregulär.

Distanzen: im Habitat 2, außerhalb 2-3; oo vielleicht 3

Keuslikolegie. Raupenfutterpflanze vermutlich nicht "Gräser" (KOCH, 1984) allge-

mein, sondern mit starker Vorliebe feuchtigkeitsliebende Pflanzen wie Carex spec.

oder Lee Die HM-Ex. wären dann vom Ruderal her (150-300 m

gen, die HW-Stücke vom Würmkanal (ca. 100 m).

Populationsbiologie: deutlich proterandrisch.

(vgl. folgende Art), dies stellt wohl

Verbreitungsstrategie: K-Siratege, 4. Crüppe

eine Anpassung an die Bedürfnisse hinsichtlich der Futterpflanzen dar, welche ja

ewässern oder anderen

nicht gleichmäßig über das Gebiet verbreitet sind, an

feuchten Stellen jedoch über Jahre hinweg stabile Bestände bilden können. Eine

Emigration aus solchen Stellen stellt dann stets eine Gefahr dar, nicht mehr zur

Besip anzuns zu gelangen, bzw. die zur Eiablage optimalen Pflanzen nicht mehr

zu finden. -

zugeflo-

Mythimna pallens

148 Individuen

139 Markiert

Tab. 81: Fangergebnisse,

Mythimna pallens.

31,2 % o- Rate

1 Wiederfang

Geschlechterverteilung und Markierungsergebnisse von

SIEDLUNG WALD HALBTROK- "DA WALD HALBTROK- "DACH.

1987 ' Garten ! KENRASEN MOOS 9 NRASE OOS

SiS SIMıSN WaS,WaM WaN HO HM HW Mb 2 Ei u es = HM ı a We

»z 5 1 3 2 1 1 8 3

a ee nen

[0107 1 1 2 1 1 9 3 6 7 1 32 [oj 07 3 5 8 8 11 7 7, - 1

99 oo 9 °8 = - 3 = — 2 1 fe) 99 - 1 3 3 9 3 See12: = -

Mark. 1 4 [2 1 1 12 <) 6 9 2 41 Mark. 3 6 11 11 20 10 8 28 - 1

Wi. NE Auer, Spaniel = EA WÄE Rt RR TEREIEU SERIEN. >

Wiederfang-Quote: sehr niedrig! Der Wiederfang WaS (c) fand nach 1 Tag statt,

das Tier war also von der direkten Lichtwirkung gefangengehalten worden.

Distanzen: 3-4

Larvalökologie: im Gegensatz zur vorhergehenden Art werden Pflanzen der Familie

Poaceae bevorzugt.

Populationsbiologie: bivoltine Art (M. impura scheint dagegen im Gebiet nur univol-

in zu sein).

Merbreitungsstrategie: r-Stratege, 2. Gruppe; völlig anders als bei der habituell

eigentlich sehr ähnlichen M. impura zeigt die Verbreitung von M. pallene im Ge-

biet ein den Wanderfaltern ähnliches Muster. Die Verbreitungsstraiegie ist korre-

liert mit der ubiquitären Verbreitung der Raupenfutterpflanzen: Die Art kann es

sich "leisten”, den Ort zu wechseln.

Mythimna I-album

Distanzen: 4 B

Verbreitungsstrategie: r-Stratege, 1. Gruppe

137

265

228

277

102

Leucania comma

97 Individuen 4,2 % 9- Rate

95 Markiert 3 Wiederfänge

Tab. 82: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Leucania comma.

SIEDLUNG WALD HALBTROK- "DA! WALD HALBTROK- "DA |

1987 ! Garten ! KENRASEN Moos‘ 2 1988 Garten Wasserwerk KENRASEN MOOS 2

SiS SIMıSN WaS;ıWaM WaN HO HM HW Mb SN WaS WaN WNw WNo 2 HO HM HW Au We |

3.par.fı 9= 7a - - - 23! 7112 - 36 Zpar. - - - - 1 1: "AB - - 69

& m Me-r EHER ABiHEE nn, DESMetEsta TE 5 € ZHBRE ZA N ENORM. I IF, 22000 Oo |

N N EN NEE: ER, EN PT A a TE ERLES {

9 "ErLan) MaamanT vrniihkt en Serra =0r (, tetler wech Co rs

Mark ee en a 20 Ta TE er TE Mi er re 3

W.f. = = z = = = = = = =

Wiederfang-Quote: relativ hoch, allerdings bei einer kurzen mittleren Verweildauer

von 2 Tagen; es handelt sich um 3 dd. RER $

Distanzen: ?; als Ausnahme kann auch einmal ein Kilometer bewältigt werden, wie

das zugeflogene d im Garten 1983 sowie die dc‘ im Wasserwerk zeigen. Die

letztgenannten Exemplare sind entweder über die Trittsteine im 100-300 m-Ab-

stand zugeflogen oder weisen auf eine sehr kleine Polulation hin, die eine solche

Trittsteinbesiedlung ausgehend von den im Osten liegenden Halbtrockenrasengebie-

ten geschafft hat. } J . . ü

Schon in der 30 m-Ausbuchtung des Habitats an HO zeigt sich ein drastischer

Häufigkeitsabfall, was für niedrige Dispersionsaktivitäten spricht. 4

Populationsbiologie: die Tatsache, daß an HW kein 9 festgestellt wurde, könnte

ahingehend interpretiert werden, daß bei einer höheren Dispersionsaktivität der

dd sogar hier schon, am Rand des Halbtrockenrasengebietes ein höherer Pro-

zentsatz von "eingewanderten Tieren” vorliegt.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

AMPHIPYRINAE

Amphipyra pyramidea

110 Individuen 37,5 % o9- Rate

79 Markiert kein Wiederfang

Wiederfang-Quote: ed auch bei täglichem Fang WaS 1988 von 13 markierten

Exemplaren keines wiedergefangen n.

Distanzen: 4, über biotopfremdes Gebiet nur 2; denkbar wäre auch, daß über bio-

topfremdem Gebiet zwar Dispersion Migration) stattfindet, das Licht jedoch nicht

angeflogen wird. 5 3

Larvalökologie: die Raupe lebt an verschiedenen Laubgehölzen.

Populationsbiologie: proterandrisch

Verbreitungsstrategie: r-Stratege, 3. Gruppe; die Art wird bisweilen als wander-

verdächtig aufgeführt. Im Untersuchungsgebiet entsprach das Häufigkeitsmuster im

Lichtfallennetz weitgehend dem der typischen Wanderfalterarten. Das Auftreten

war, vor allem an HW, oft mit den Wanderfalternächten korreliert.

Amphipyra berbera

Distanzen: 4, vermutlich wie A. pyramidea

Larvalökologie: siehe Bemerkungen zur vorigen Art

Verbreitungsstrategie: r-Stratege, 5. Gruppe; Verhältnisse vermutlich wie bei A.

pyramidea

Amphipyra tragopoginis

190 Individuen 20,0 % 9- Rate

139 Markiert 5 Wiederfänge

Tab. 83: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Amphipyra tragopoginis.

_ MHaurstdad vater bl Sehr Vagina Malen pen 2 Seien nn wald = Wr E ı

HAL - WALD HALBTROK- "DA

1987 ern a KERRASEN DE >= 198 Garten Wasserwerk NRASEN Hass }

SiS SiM | SIN WaS | WaM WaN HO HM HW Mb SN WaS WaN WNw WNo 2 HO HM HW Au We

Z par. ar sBr N! 7 2 9 5 Bigan - 115 I par. 1 8 10 3 3 8 4 22 1 = 6

Z zus. I | 3 - - - - - - 8 % zus. - 7 - & e = = e ® =

dd 7403 6 - 4 4 3 4 - 72 [0102 = 13 7 2 2 4 2.5 10 - - 4

99 zur pe ae a ER ERENTO EIERN TER RZSR SAUERSNESEn EEE ES |

Mark. 47 4 10 - 6 5 6 52 = 94 Mark. = 13 7 2 2 5 211 - — 4

W.f. are - - - - - - - - W.f. — 4 E = - = = 1 = =

138

Wiedertf te: durchschnittlich, drei der WaS-Wiederfänge 1988 erfolgten nach

1 Tag (dc) und sind somit nicht als Hinweis auf Ortstreue zu werten. Auch die

iv: n Verweildauern liegen au we niedrigem Niveau: Ein d nach 3 Tagen

Was) und ein 9 nach 2 Tagen (HW).

Distanzen: 3-4

Larvalökologie: durch die Raupenfutterpflanzen mehr ans Offenland gebunden als die

vorigen Arten. 4

Populationsbiologie: deutlich proterandrisch N

Verbreitungsstrategie: r-Stratege, 3. Gruppe; das Häufigkeitsmuster innerhalb des

Lichtfallennetzes entspricht auch hier den typischen Wanderfaltern, das Auftre-

ten ist oft mit den Wanderfalternächten korreliert, und es waren, vor allem an

HW z.T. starke Nacht-zu-Nacht-Fluktuationen festzustellen, was vielleicht als

Hinweis auf erhöhte Dispersionsaktivitäten gelten mag.

Rusina ferruginea

siehe Fernwiederfänge (8.2.), verringerte Fallendistanzen (8.3.) und Versetzexperiment (8.4.)

559 Individuen 23,3 % 9 - Rate

493 Markiert 21 Wiederfänge

Tab. 84: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Rusina ferruginea.

SIEDLUNG WALD HALBTROK- m WALD HALBTROK- ms

1987 _ ! Garten ! KENRASEN M 2 1988 Garten Wasserwerk KENRASEN M: 0%

SiS SIMıSN WaSıWaM WN HO HM HW Mb SN WaS WaN WNw WNo HO HM HW Au We

Döpar 1a 1) 07 eo a 52 183 per 11, 723), 131, 68, 43) Wesscmenn oyhanaa 233

zus Zeug 207 0-2 7320726: Ku 6 = BI E22 zus, 1-2 7146 - = - ERS EH- - > 46

de 2: 5 20m eine raoltiı: Ehe 25321 Elicsrltce 8 1 BONN 2 EEE 39 LTE alas. 228

99 23 ao 26 1 834099 au 10 Inmitn 4. war 1 .ns6

Markggs13: 211823341167 Krasisi, 1b 10,7 34,.07229,,..Marki 1.917..69,,025 .65,64104438,m, 6 056400) 14 264

wi. Sinne: et a = 1 2 WR. ok ST RE TREE T Se 19

Wiederfang-Quote: relativ niedrig, vor allem im Ort (1988). Hier liegen auch die

Verweildauern auf dem niedrigst möglichen Niveau (1,0 Tage). Ein im Moos 1987

nach 8 Tagen wiedergefangenes JS weist vielleicht auf größere Ortstreue in den

Moorbirkenwäldchen hin; der Fang erfolgte dort zur Flugzeit nur wöchentlich,

sonst wäre dort die Wiederfang-Quote wohl höher ausgefallen.

Die 99 sind im Wiederfangergebnis etwas unterrepräsentiert: 1/21= 4,8 %. Es

handelte sich um einen "Ortswechsler" WaS>SiN (30 m) in 3 Tagen.

Distanzen: 3-4 _ ?

Populationsbiologie: proterandrisch

Verbreitungsstrategie: r-Stratege, 2. Gruppe; es scheinen dennoch Barrieren gegen

die freie Mobilität dieser Art zu existieren: Das polarisierte Häufigkeitsverhältnis

SiN/WanN zeigt beispielsweise, daß R. ferruginea den Schutz von Gebüschen und

Bäumen bevorzust, iotoptreue bedeutet daher nicht zwangsweise Ortstreue!

Hohe Nacht-zu-Nacht-Fluktuationen deuten auch auf hohe Dispersionsaktivitäten

in.

Talpophila matura

21 Individuen 13,3 % 9 - Rate

15 Markiert 1 Wiederfang

Wiederfang- te: hoch, bei einer allerdings kurzen Verweilzeit von 2 Tagen (ein

cd am Standort HW); Stichprobe noch zu klein h

Distanzen: 2; bisher nur an und HW festgestellt, die Distanz zu den nächstge-

legenen anderen Fundorten (1 km) liegt offensichtlich nicht im normalen Bereich

der Dispersionsaktivität. arten

Verbrei strategie: K-Stratege, 6. Gruppe; nach KOCH (1984) treten bei dieser

Art starke Häufigkeitsschwankungen auf, was eigentlich ein Kennzeichen von r-

Strategen ist. Es ist durchaus denkbar, daß diese Art in anderen Gebieten unter

vielleicht anderen Bedingungen auch eine andere Strategie verfolgt.

Euplexia lucipara (41 Individuen)

Distanzen: 2-3

Larvalökologie: das HM-Stück 1986 (cd) stammt al von den Raupenfutterpflan-

zen her beurteilt, vom Ruderal (150-300 m entfernt). Die Distanz Wald> Flugha-

fen-Mitte (1 Em) wird offensichtlich nicht regelmäßig zurückgelegt. Einen weite-

ren Hinweis in dieser Richtung stellt wohl der starke Häufigkeitsgradient zwi-

schen WaN und WNo auf einer Strecke von nur 50 m dar.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

139

Phlogophora meticulosa (15 Individuen)

Distanzen: 4 > alt

Populationsbiologie: bivoltin

Verbrei sstrategie: r-Stratege, 1. Gruppe; die Häufigkeitsschwankungen, die an

einen Schwarm erinnernden 12 Moos-Exemplare in einer Nacht, sowie das den

anderen typischen Wanderfaltern entsprechende Häufigkeitsmuster innerhalb des

Eangnetzes sind weitere Hinweise auf eine Charakteristik dieser Art als hoch-

mobil.

Ipimorpha retusa (18 Individuen)

Distanzen: 2-3

Larvalökologie: oligophag an Weiden, Pappeln und Erle. Im Flughafen-Ruderal be-

findet sich offensichtlich eine relativ stabile Population mit eide als Lebens-

ee die 150-300 m zum Fangplatz HM werden ohne größere Probleme

ewältigt (3d4). .

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Ipimorpha subtusa (1 Individuum)

Distanzen: 1-2

Larvalökologie: die Raupen sind auf Arten der Gattung Populus spezialisiert. Bisher

konnten keine biotopfremden Tiere nachgewiesen werden.

Verbreitungsstrategie: K-Stratege, 6. Gruppe; im Zusammenhang mit der stärkeren

Futterpflanzenspeziaisierung im Vergleich zu /. retusa scheint diese Art auch eine

etwas andere Strategie zu verfolgen.

Enargia paleacea

Distanzen: vermutlich 2-3

Larvalökologie: oligophag an Birke, Pappelarten und Erle

Verbreitungsstrategie: vermutlich intermediärer Typ, 5. Gruppe

Enargia ipsilon (3 Individuen)

Distanzen: vermutlich 2-3

Larvalökologie: die Raupe lebt an Benpelarien und an Bruchweide (Salix fragilis).

Bisher konnten keine biotopfremden Tiere festgestellt werden.

Verbreitungsstrategie: vermutlich intermediärer Typ, 5. Gruppe

Cosmia affinis |

Distanzen: 2: der sehr lokalen Verbreitung in Südbayern entspricht das lokale

Vorkommen im Untersuchungsgebiet: Die Art konnte bisher nur im Korbinianiholz

nachgewiesen werden, die Gebiete außerhalb davon liegen wohl auch nicht im

Bereich der normalen Dispersionsaktivität.

Larvalökologie: auf Ulmenarten spezialisiert

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Cosmia trapezina

245 Individuen 24,3 % o- Rate

177 Markiert 7 Wiederfänge

Tab. 85: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Cosmia trapezina.

WALD

SEDLUNG WALD‘ HALBTROK- "DA

KENRASEN MOO x 1988 ‚Garten Wasserwerk

1987 ' Garten ! EENRASEN Moos J

SIS SIM SIN WaS WaM WaN HO HM HW N WaS WaN WNw WNo 2 HO HM HW Au We

2 par. < - 1 30 0) 5 17 - 2 21 84 X par. 6 40 6 6 1 18 - 6 6 ih

X zus. - - 6 5 - - 1 - - 3 14 X zus. - 40 - - - - - - - -

dd 2 - 4 26 2 3 8 - 1 Ai 63 dd 4 47 6 4 10 - 3 1 4

99 1 - 1 4 2 1 8 - - 8 25 99 = 8 - - 2 4 - 2 - 3

Mark. 2 - 6 30 4 4 15 - 1 14 75 Mark. 4 54 65 4 8 14 - 5 1 7

Wi. je E27 Sie ES = 1 wi - 6 808 N -

Wieder Rogue: sehr niedrig! Sämtliche Wiederfänge erfolgten nach Intervallen

von nur 1 Tag und sind somit keine Hinweise auf Ortstreue. Das 1987 im Garten

rückgefangene Tier war ein "Ortswechsler" WaS>SiN (1 Tag). Auch die Wie-

derfang-Quote von 11 % bei täglichem Fang und die Tatsache, daß kein Zweit-

AESET ang stattfand, zeugen von einem schnellen Verschwinden aus der näheren

‚Umgebung.

Dose, Habitat 3 (-4?), außerhalb 1-2

140

reinen Offenland (HM) bisher noch kein Exemplar festgestellt.

Populationsbiologie: deutlich proterandrisch

Verbreitungsstrategie: r-Straiege, 5. Gruppe; auch hier zeigt sich, daß Biotoptreue

und Ortstreue nicht gekoppelt sein müssen.

Larvalökologie: die a lebt polyphag an Laubbäumen, dementsprechend wurde im

Cosmia pyralina (37 Individuen)

Distanzen: Verhältnisse wahrscheinlich wie bei der vorigen Art

Larvalökologie: polyphag an Laubbäumen, auch hier konnte an HM bisher kein

Exemplar festgestellt werden.

VerbreHungsstzatesie: r-Stratege, 5. Gruppe

Auchmis comma (4 Individuen)

Distanzen: 2-3

Larvalökologie: monophag an Berberitze (Berberis Mugeunı); für die Exemplare, die

im Siedlungsgebiet nachgewiesen wurden, dienten die desöfteren in Gärten an-

genlenzten erberitzen als Lebensgrundlage. Es existiert zusammen mit den im

ereich der Kiefern-Eichen-Wälder eingestreuten Büschen ein Futterpflanzennetz

mit Abständen von ca. 100 m.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Actinotia polyodon (4 Individuen)

Distanzen: vermutlich 3

Larvalökologie: die Raupe, lebt an Johanniskraut (Hypericum) und Tragant (Astra-

zaluıs). Das HM-Stück (C) ist also mindestens vom Ruderal her (150-300 m) zu-

geflogen.

Pepulationsbiolegie bivoltin

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Actinotia hyperici (1 Individuum)

Distanzen: vermutlich 2-3

Larvalökologie: monophag an Hypericum perforatum

Popuiationsbiologie: bivoltin

Verbreitungsstrategie: vermutlich intermediärer Typ, 5. Gruppe; die sehr lokale

erbreitung in Südbayern und das Auffinden dieser Art nur im typischen Biotop

trockene heiße Stellen) lassen an geringe Dispersionsaktivitäten dieser Art unter

en gegebenen Bedingungen im Untersuchungsgebiet denken. Dies dient dazu, nicht

zufällig in ungeeignete Biotope zu gelangen und somit die Fortpflanzungswahr-

scheinlichkeit drastisch herabzusetzen. Die Gefahr eines solchen Ereignisses ist

bei relativ stenöken Bewohnern seltener Biotoptypen am größten.

An anderen Stellen, vor allem südlich der Alpen, wo geeignete Habitate in wei-

taus größerer Anzahl zur Verfügung stehen wird die Strategie dementsprechend

angepaßt sein.

Apamea monoglypha

70 Individuen 17,6 % 9- Rate

68 Markiert 2 Wiederfänge

Wiederfang-Quote: sehr niedrig! Beide Wiederfänge (47) erfolgten 1988 im Garten

WaS) nach I Tag, sind also auf ein Festhalten durch die direkte Lichtwirkung bei

täglichem Fang zurückzuführen. Es sind auch die vergleichsweise niedrige ie-

derfang-Quote von 9,5 % und die fehlenden Zweitwiederfänge an diesem Standort

‚bemerkenswert.

Distanzen: 4 _

Populationsbiologie: proterandrisch , $

Verbreitungsstrategie: r-Stratege, 1. Gruppe; es handelt sich hier allerdings _ nicht

um einen "klassıschen” Vertreter aus der Gruppe der Wanderfalter, eine Korre-

un ga Auftretens mit den sogenannten anderfalternächten war jedoch zu

eobachten.

Apamea lithoxylea (1 Individuum)

Distanzen: vermutlich 3

Verbreitungsstrategie: vermutlich r-Stratege, 3. Gruppe

141

Apamea sublustris

52 Individuen 10,2 % 9 - Rate

47 Markiert 2 Wiederfänge

Wiederfang te: standortabhängig: am Flughafen niedrig, im Wasserwerk relativ

hoch, die Einzel-Stichproben sind noch zu klein.

Ein Ortswiederfang im Garten 1988 (cd) nach I Tag erklärt sich durch ein Fest-

ehalten-Werden am Licht bei täglichem Fang. Interessanter ist dagegen ein cd‘,

as im Wasserwerk nach 5 Tagen wiedergefunden wurde; es war von w nach

WaN (100 m) geflogen. Im Wasserwerk wurden 1988 insgesamt nur 3 Individuen

markiert! Hier könnte eine erhöhte Ortsfestigkeit vorliegen, die ihren Ursprung in

‚der relativ isolierten Lage des Biotops hat.

Distanzen: 2-3; die im Garten vereinzelt nachzuweisenden Stücke sind wohl zuge-

flogen, die 1988 festgestellte größere Anzahl dagegen könnte auf eine erfolgrei-

che "Brut" zurückzuführen sein, zumal am Licht auch sehr frische Exemplare

auftauchten.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Apamea crenata (17 Individuen)

Distanzen: 3 5

Verbreitungsstrategie: r-Stratege, 3. Gruppe

Apamea characterea (13 Individuen)

Distanzen: 2-3 L

Larvalökologie: die Raupen dieser Art sind nach KOCH (1984) mehr auf Waldgrä-

sern und unter anderem auch auf Schilf zu finden. Die beiden HM-Stücke (do)

stammen daher vielleicht aus dem Ruderal (150-300 m).

Verbreitungsstrategie: intermediärer Typ, 3. Gruppe

Apamea lateritia (1 Individuum)

Distanzen: vermutlich 4

Verbreitungsstrategie: vermutlich r-Stratege, 3. Gruppe

Apamea remissa (38 Individuen)

Distanzen: 3; die in der ediunn festgestellten 3 Stücke sind wohl von den Ver-

breitungszentren (Flughafen, Wa

zugeflogen, also ca. | km weit.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Apamea unanimis (7 Individuen)

Distanzen: 2-3

Larvalökologie: oligophag an Glanzgras (Phajaris arundinacea) und selten an Schilf

Phragmites "austra ae: Das HM-Stück (4) stammt daher zumindest aus dem

uderal (150-300 ‚ das S am Standort HO vom Schloßkanal, der Falter an HW

vom Würmkanal (ie

die im Garten beobachtet wurden, vom Ort ihrer Entwicklung aus mindestens 200 m

zurückgelegt haben.

Das 1989 im Wasserwerk (WaN) beobachtete d zeugt von einer Flugdistanz (Ko- '

lonisation?) von mindestens 700 m.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Apamea anceps

siehe verringerte Fallendistanzen (8.3.)

244 Individuen 19,3 % 9- Rate

154 Markiert 5 Wiederfänge

sserwerk und ähnliche Trockengrasbiotope) her |

e ca. 200 m), und schließlich mußten auch die 5 Exemplare,

Tab. 86: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Apamea anceps; die Art wurde 1987 nicht markıert.

W HALBTROK- "DA

n KENRASEN MOOS ?:

1988 Garten Wasserwerk

SN WaS

eN WNw WNo HO M HW Au We

X par. - 2 72 9037 2 37 1 - 165

dd 2 2 53 er: 1 4 33 1 - 130

99 - - 16 2 7 1 1 4 - - 31

Mark. - 2 66 9 36 2 5 34 1 - 154

W.f. - 1 2 - - - - 2 - - 5

142

Wiederfang-Quote: durchschnittlich, lediglich das Wiederfang-Ergebnis am Standort

HW ist als überdurchschnittlich zu bewerten. Es handelt sich dort um zwei dd,

die nach gereil 4 Tagen rückgefangen wurden. Im Garten (WaS), wo A. anceps

vermutlich nur als Gast auftritt, wurde ein d durch die Lichtwirkung einen Te

lang festgehalten.

Distanzen: 3_ _ ;

Populationsbiologie: proterandrisch

Verbrei stralegie. r-Stratege, 3. Gruppe; das Häufigkeitsmuster im Fangnetz des

Untersuchungsgebietes ähnelt dem der typischen Wanderfalter.

Apamea sordens (84 Individuen)

Distanzen: 3

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Apamea scolopacina {35 Individuen)

Distanzen: 2-3 2 :

Larvalökologie: die Raupe lebt an einer Reihe von Gräsern, die bevorzugt in Wäl-

dern wachsen. Davon leitet sich auch die Ökologische Charakteristik der Imagi-

nes ab. Dementsprechend konnte im reinen Offenland HM) die Art nicht nach-

gewiesen werden, die Distanz von 1 km über biotopfremdes Gebiet scheint nicht

ım Bereich der normalen Dispersionsaktivität zu liegen.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe; im Verlauf dieses Jahrhun-

derts hat offensichtlich eine Besiedlung des südbayerischen Faunengebiets stattge-

funden. Hierbei mußten mindestens 2 km pro Jahr zurückgelegt worden sein,

denkbar wäre jedoch auch ein einmaliger größerer Vorstoß.

Apamea ophiogramma (5 Individuen)

Distanzen: 2-3

Larvalökologie: olig phag an einer Reihe von hygrophilen Pflanzen, v.a. an Schilf.

Das WNw-Stück e mußte ca. 100 m von der en Di her

gelogen sein, interessant ist jedoch vor allem die (wahrscheinlich) erfolgreiche

olonisation des Wasserwerkgeländes schon 2-3 Jahre nach dem ersten Auftau-

chen der Raupenfutterpflanzen über 1 km biotopfremdes Gebiet hinweg.

In der Siedlung bildet die Begleitvegetation der Gartenteiche die Lebensgrundla-

e, die beiden HO-cd' zeigen, daß auch 200 m Distanz durch Waldgebiet hin-

durch ee Seren OÖ) bewältigt werden. Das HW-Stück stammt vom

Würmkanal (ca. 100-150 m

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Oligia strigilis (206 Individuen)

Distanzen: 3

Eeaökologie an Gräsern, aber wie alle Oligien im Moos und im Waldinneren

seltener. Der Einflug über Distanzen von 1-3 km scheint hier relativ ineffizient

zu sein. F

Verbreitungsstrategie: r-Stratege, 3. Gruppe

Oligia versicolor (55 Individuen)

Distanzen: 2-3

Larvalökologie: im Vergleich zu den beiden anderen Oligia-Arten ist O. versicolor

noch am ehesten in der Lage, feuchte und stark bewaldete Biotope zu besie-

deln.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Oligia latruncula

497 Individuen 45,5 % o- Rate

31 Markiert kein Wiederfang

Wiederfang-Quote: niedrig, Stichprobe jedoch noch zu klein; die 3l Markierungen

‚erfolgten alle 1988 im Wasserwerk.

Distanzen: 3- . h

Lee sone die Raupe lebt an Gräsern und findet wohl überall eine Lebens-

grundlage. ER i 2

Verbreitungsstrategie: r-Stratege, 3. Gruppe; dies ist ein Beispiel dafür, daß sich

durchaus auch kleine Arten eine r-Strategie zueigen gemacht haben.

143

Miana furuncula (64 Individuen) |

Distanzen: 3; der drastische Häufigkeitsabfall von WaN nach WNw auf einer Strek-

ke von 100 m weist auf Barrieren gegen eine freie Beweglichkeit bei dieser Art

hin. Ein solches Hindernis stellen bei M. furuncula anscheinend feuchte Waldge-

biete bzw. -ränder dar. " 1 4 rs |

Larvalökologie: an verschiedenen Gräsern, die Art scheint lediglich in den etwas

feuchteren Wäldern keine Lebensgrundlage zu finden. |

Verbreitungsstrategie: intermediärer Typ, >. Gruppe |

Mesapamea secalis

63 Individuen 28,3 % 9 - Rate

Distanzen: 4

Larvalökologie: die Raupen dieser und der folgenden Art leben an Gräsern, vor

allem an Getreide; so finden die Mesapamea-Arten so gut wie überall eine Le-

bensgrundlage, lediglich das Waldinnere, insbesondere von feuchteren Wäldern

scheint die Art tendenziell zu meiden.

Populationsbiologie: stark proterandrisch; dies ermöglicht der Art, zumindest in

manchen Jahren eine relativ gute Trennung des Fortpflanzungsgeschehens mit dem

der folgenden Art:

n F

; Er Es ist gut ersichtlich, daß Anfang

M. secalis Im: August 1988 für die secalis-Id

E 3 das Flugmaximum mit dem Zeit-

ai E punkt zusammenfiel, an dem ver-

10 = 2 stärkt 99 emergierten. Für die

£ . Gewährleistung einer möglichst ı

reibungslosen Befruchtung ist es

von Vorteil, wenn schon alle dd

FE vorhanden sind und sozusagen nur

x darauf warten, ein befruchtungsfä-

9 . 5 higes o zu finden.

Abb. 44: Anflugdiagramm für Mesapamea secalis 1988.

n Dieser Zeitpunkt lag, wie aus dem

zweiten Flugdiagramm zu entneh-

M. secalella men ist, = esapamea secalel-

12 te la 1988 1-2 Dekaden früher,

nämlich Mitte bis Ende Juli.

[07 3

4 £ 2, ER

m “€

MER # ze u —

EsivW AT MT ET AG ME E8

Abb. 45: Anflugdiagramm für Mesapamea secalella 1988.

Verbreitungsstrategie: r-Stratege, 3. Gruppe; diese Art unterliegt, wie auch die

folgende, starken Häufigkeitsgeinyen nn was als Hinweis auf starke Disper-

sionsaktivitäten gewertet werden kann. Das Auftreten ist darüber hinaus in einer

besonderen Weise mit den Wanderfalternächten korreliert.

Mesapamea secalella

69 Individuen 25,0 % o- Rate

Distanzen: 4

Larvalökologie: wie M. secalis

Populationsbiologie: siehe vorhergehende Art .

Verbreitungsstrategie: r-Stratege, 3. Gruppe; siehe Hinweise zu M. secalis

144

Photedes minima (25 Individuen)

Distanzen: 1-2; außerhalb der Moorbirkenwäldchen nur in einem Exemplar an HW;

dieses d könnte von den feuchteren Stellen am Würmkanal stammen (an der

kürzesten Stelle 150 m entfernt).

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Photedes extrema (11 Individuen)

Distanzen: 2; die Distanz von 1 km, die bei einem (nicht festgestellten) Zuflug in

den Ort bewältigt werden müßte, Hiest, nicht im Bereich der normalen Disper-

sionsaktivität. Die Kolonisation des asserwerks setzt jedoch Trittsteinsprünge

von mindestens 100-300 m voraus. ;

Larvalökologie: die Raupe lebt monophag an Land-Reitgras (Calamagrostis episer

os), die beiden HM-Stücke stammen daher mit ziemlicher Sicherheit aus dem

uderal, die nächstgelegenen Standorte der Futterpflanze liegen ca. 150 m ent-

fernt.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Photedes fluxa (67 Individuen)

Distanzen: 3, e

Larvalökologie: wie die vorige Art manophag an Land-Reitgras, die Imagines sind

je ungleich weiter verbreitet. Die 6 Exemplare an HM (aus dem Ruderal,

50-300 m) sowie die vereinzelt im, Garten fliegenden Stücke, die aus min-

destens 200 m Entfernung stammen (auch o9), zeigen, daß derartige Distanzen

zur normalen Dispersionsaktivität dieser_Art gehören.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Photedes pygmina

Distanzen: 2

Verbreitungsstrategie: K-Stratege, 6. Gruppe (siehe SPITZER et al., 1984)

Luperina testacea

94 Individuen 17,4 % 9 - Rate

86 markiert 2 Wiederfänge

Wiederfang-Quote: relativ niedrig: 1988 wurde im Garten (WaS) von 13 markierten

Faltern ein d durch die Ta yunz einen Tag lang festgehalten, 1987 erfolgte

‚ein regulärer Wiederfang SiN) eines d nach 2 Tagen.

Distanzen: 3_

Larvalökologie: die Raupe lebt, an Gräsern

Populationsbiologie:, proterandrisch

Verbreitungsstrategie: intermediärer Typ, 3. Gruppe

Amphipoea oculea (? Individuen)

Distanzen: vermutlich 2-3

Verbreitungsstrategie: vermutlich intermediärer Typ, 5. Gruppe

Amphipoea fucosa (180 Individuen)

Distanzen: 3; das stark polarisierte Häufigkeitsverhältniis WaN/WNw auf einer

Strecke von 100 m deutet auf Barrieren gegen eine freie Dispersionsaktivität hin.

Verbreitungsstrategie: intermediärer Typ, 3. Gruppe

Amphipoea lucens (1 Individuum)

Distanzen: 1-2

Larvalökologie: die Raupe lebt nach KOCH (1984) an "Gräsern, mutmaßlich an

Pfeifengras (Molinia caerulea)". Diese Raupenfutterpflanze kommt im Wasser-

werk am Fundort, WNo vor.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Hydraecia micacea (37 Individuen)

Distanzen: 3

Larvalökologie: die Raupe lebt oligophag an einer Reihe mehr oder weniger feuch-

tigkeitsliebender Pflanzen. Die im Garten festgestellten 6 Exemplare sind offen-

sichtlich zugeflogen. Der nächstliegende Bestand einer der von KOCH (1984)

genannten Futterpflanzen liegt ca. 200 m entfernt. In den letzten beiden Jahren

waren an verschiedenen Gartenteichen der näheren Umgebung erstmals Schwertli-

lien (Iris) festzustellen; dies könnte zu einer Kolonisation geführt haben und

würde das Bi Auftreten 1988 im Garten erklären.

Das HM-Stück 6 stammt wohl aus dem Ruderal (150-300 m entfernt).

Verbreitungsstrategie: intermediärer Typ, 3. Gruppe

145

Gortyna flavago (9 Individuen)

Distanzen: 2-3; der starke Häufigkeitsgradient im Wasserwerk könnte auf geringe

Dispersionsaktivitäten dieser Art IchY ten.

Larvalökologie: von den in KOCH 1984) genannten Raupenfutterpflanzen wächst in

der näheren Umgeb des Gartens keine. Das 1986 nachgewiesene J' ist daher

vermutlich vom Rand des Berglwaldes her zugeflogen (mindestens 150-200 m).

Verbreitungsstrategie: intermediarer Typ, 5. Gruppe

Celaena leucostigma (1 Individuum)

Distanzen: vermutlich 2-3; manchmal 4 (EITSCHBERGER & STEINIGER, 1980)

Larvalökologie: die Raupe lebt oligophag an einigen hygrophilen Pflanzen, die alle

nicht am Fundort WaM vorkommen. Das cd ist vermutlich vom 150 m entfernten

Berglbach her zugeflogen. Die Strecke ist bewaldet, der Flug erfolgte nicht

aufgrund der Lichtattraktion, da durch eine hohe Gebüschzeile der direkte Licht-

einfluß erst unmittelbar vor der Falle zum Tragen kam.

Verbreitungsstrategie: vermutlich intermediärer Typ, 5. Gruppe

Nonagria typhae (10 Individuen, dazu eine größere Anzahl von Exuvien)

Distanzen: 1-3; im Wasserwerk weist der starke Häufigkeitsgradient auf eine gerin-

ge Mobilität hin. { ;

Larvalökologie: die Raupe frißt Teichkolben (Typha) und Teichsimse (Schoeno-

lectus Tacustrie), Am Standort WNo, wo sich ein kleiner Bestand von Teichkol-

en befindet, ist eine kleine Population von N. typhae bodenständig, wie durch

Exuvienfunde bestätigt wurde. 1988 konnte trotz 10 an WNo festgestellten Ima-

ame: kein Exemplar beobachtet werden, das die Distanzen von 50 bzw. 120 m an

ie beiden anderen Fallen-Standorte im Wasserwerk bewältigt hätte.

Andererseits wurden am Franzosenhölzl 1989 2 Exemplare d0) gangen, die sich ohne

Sichtkontakt zur Lichtquelle 40 m weit von den Teichkolben-Beständen entfernt hatten.

Denkbar sind als Erklärung aber auch zwei unterschiedliche Flugtypen; en

ist die Anziehungskraft des Lichts auf Tiere, die sich auf "Fernflügen” (Dispersion

befinden, geringer als auf solche im trivial movement.

Der Teichkolbenbestand befindet sich erst seit 2-3 Jahren an dieser Stelle. Es

mußte in diesem Zeitraum eine Kolonisation aus über 1 km Entfernung, dem

nächsten Vorkommen der Raupenfutterpflanze, stattgefunden haben. hnlich

schnelle Kolonisationen über vergleichbare Distanzen ergeben sich auch bei der

Auswert der weiteren Standorte, wo Exuvien gefunden wurden Inepaarie

Teiche im Flughafengebiet und am Franzosenhölzl im Dachauer Moos).

open ouelnlEnie: am Standort WNo, wo ca. 500 Typha-Halme stehen, wurden mit

xuvien besetzte Pflanzen ausgezählt: Auf ca 15 Halme war ein besetzter zu

finden (n = ca. 100). Auf den gesamten Typha-Bestand zurückgerechnet ergäben

sich 33 im Jahr 1988 emergierte Falter. Nur ca. 1/3 davon wurde am Licht nach-

gewiesen. Dies ist wohl vor allem durch nicht optimales Anflugverhalten bedingt;

weitere Gründe können rasche Mortalität vor der nächsten Möglichkeit, die ge-

schlüpften Tiere zu erfassen, sowie ein schnelles Abwandern mancher Exemplare sein.

Verbreitungsstrate e: K-Stratege, 6. Gruppe; die Erschließ neuer Habitate, die

recht schnell erfolgen kann, erfolgt durch Individuen mit höherer Dispersionsakti-

vität, die entweder bei einer starken Vermehr eines Bestandes abwandern oder

bei einer Vernichtung der Lebensgrundlage (z.B. einem Trockenfallen eines

Feuchtgebietes oder einer Vernichtung durch Baumaßnahmen) gezwungen sind, den

Standort zu verlassen.

Nonagria nexa (1 Individuum)

Distanzen: 2-3

Larvalökologie: oligophag an einigen hygrophilen Pflanzen, das HO-Stück (d) stammt

vermutlich vom 200 m entfernten Schloßkanal. Die Flugstrecke führt durch be-

waldetes Gelände. .

Verbreitungsstrategie: K-Stratege, 6. Gruppe; die Verhältnisse ähneln vermutlich

denen bei N. typhae. Das nächstgelegene bisher bekannte Vorkommen dieser Art

liegt in den Isarauen ca. 7 km östlich von Oberschleißheim. Da das Untersu-

chungsgebiet um die Jahrhundertwende entomologisch sehr gut durchforscht war, han-

delt es sich ziemlich sicher um eine Kolonisation über die genannte Mindestdistanz.

Rhizedra lutosa (3 Individuen)

Distanzen: 2, .

Larvalökologie: monophag an Schilf (Phragmites "australis"); die vom Standort HM

aus gesehen nächsten Schilfvorkommen liegen im Ruderal ca. 200 m, entfernt.

Diese Distanz wird mit erstaunlicher Konstanz bewältigt (auch ein 9). Jedoch

schon für die Strecke von 1,2 km nach HO konnte kein Flugnachweis erbracht

werden. Das an HW gefundene N stammt mindestens aus einer Entfernung von

150 m (Würmkanal) ohne Sichtkontakt zur Lichtquelle. j 1

erbreitungsstrategie: K-Stratege, 6. Gruppe; die Strategie dürfte der bei N. typhae

besprochenen ähneln.

146

Abb. 45b: Nonagria typhae & (WNo, 28.9.88), ein Vertreter aus der Gruppe der soge-

nannten Schilf- bzw. Röhrichteulen an der Raupenfutterpflanze Typha latifolia.

147

siehe Fernwiederfänge (8.2.) und Versetzexperiment (8.4.)

300 Individuen 7,37%) 0- Rate

290 markiert 65 Wiederfänge

Tab. 87: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Meristis trigrammica.

SIEDLUNG WALD HALBTROK- DACH, WALD HAI.BTROK- ne)

1987 ' Garten ! KENRASEN M 2 1988 Garten Wasserwerk KENRASEN M

SiS SIMıSN WaS;WaM WaN HO HM HW Mb SN WaS WaN WNw WNo HO HM HW Au We

Ipa. - 2 16 6 1, - Soon - Tape 27 30 4 5.0 33 30 mi

2 zus. FIT 31 a - SEE - 4 Zus - 4 > > Er -

a a Se u 1: Ei - 93 de 23 66 a 60.108 30 29° joe

Same A et - 1.09 1 5 Sul 3 ee x

Merk. - 2 180. sS5 1 Segeaieen - 108 Mark. 24 70 2er 33 30 1 Den

Sr 1 1 15 ee - 2 wi. 4 54 1, wa 4 = wre

Wiederfang-Quote: standortabhängig: Im Garten und an HO meist hoch, sonst_eher

niedrig. An HO wurden 1988 drei Jod‘ nach 2 Tagen, eines erst nach 6 Tagen

miederze angen. Die mittlere Verweildauer im Garten 1988 betrug (bei anderer

Methodik) 3,0 Tage, was verglichen mit anderen Arten einen langen Zeitraum

bedeutet.

Distanzen: 2-3

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe; einer gewissen Ortstreue zu-

mindest von Teilen der jeweiligen Populationen in Eichenwäldern steht eine erhöh-

te Beweglichkeit an den anderen Standorten gegenüber. Im Offenland (HM) sind

überdies starke Nacht-zu-Nacht-Fluktuationen ein weiteres Indiz für starke Di-

spersionsaktivitäten: So wurden am 30.5.88 14 Individuen und am 26.5.86 gar 24

xemplare gefangen; in beiden Fällen erbrachten die Fangnächte davor und danach

kein einziges Stück.

Hoplodrina alsines

siehe Fernwiederfänge (8.2.) und Versetzexperiment (8.4.)

706 Individuen 14,7 % 9- Rate

635 markiert 43 Wiederfänge

Tab. 88: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Hoplodrina alsines.

WALD HALBTROK- _—

Garten Wasserwerk KENRASEN M

SN WaS WaN WNw WNo HO HM HW Au We

1987 ' Garten !

SIEDLUNG WALD HALBTROK- DACH,

| KENRASEN M 2 1988

SiS SIMıSN WaS,WaM WaN HO HM HW Mb

Z par. 9 10 20 64 15 83 26 15 251 I par. 24 69 32 19230 15 en 17, 5

7 2 1

zus WwaAS; 48 6 13210210 - 82 3 zus; (227,161 - Sr Sn Ans

dd 7916 79) S1a MebE rag ee 2 242 dd 22 187 ro le Ve BT

92 sale? 26.4 1. 12 Bert 4 - 57 9 1 25 100.2 ) 0. NE

Mark. Zu 108.177 103,167 Dreresaeene 2 289 Mark. 22 21 30, 10227 1a BT Ben

Wwf. er) 3 ver en - 6 wit. 1 30 1 _ 4 1 Eu I r=

Wiederfang-Quote: durchschnittlich, im Wasserwerk 1988 mit 5,9 % Ortswieder-

fängen vergleichsweise hoch. Es handelte sich dort um 4_d’d' mit einer relativ

niedrigen mittleren Verweildauer von 2,8 Tagen. Auch im Garten 1988 lagen (bei

anderer Methodik) die Verweilzeiten bei einem Mittel von 2,2 Tagen in einem

nicht als hoch zu bezeichnenden Bereich. Das Maximum eines nach 9 Tagen

wiedergefangenen J zeigt, daB durchaus einzelne Exemplare der Population orts-

treu sein können. !

Die 99 sind im Wiederfangergebnis mit 1/43=2,3 % deutlich unterrepräsentiert.

Distanzen: 3-4; die Fernwiederfänge, sowie die Tatsache, daß im Versetzexperi-

ment und im Wasserwerk die Distanzen von 120 m mühelos bewältigt wurden,

deuten auf hohe Dispersionsaktivitäten des Großteils der Population hin.

er tenlage. die Art findet wohl überall, wo niedrige Pflanzen wachsen, eine Le-

bensgrund age. .

a onnio ogie: stark proterandrisch (im Gegensatz zu den Angaben in NOWAK,

974); die 52 scheinen an bestimmten Stellen bevorzugt aufzutreten: Die o-Rate

war an WaS stets 2-3mal so hoch wie am 30 m entfernten Standort SiN, ein

ähnliches Ampnemen zeigte sich im Wasserwerk 1988 auf einer Strecke von 100 m

WaN/WNw). Vielleicht erleichtert ein solches Verhalten das Sich-Auffinden der

eschlechtspartner ("mating sites").

Verbreitungsstrategie: r-Stratege, 2. Gruppe

148

Hoplodrina blanda (26 Individuen)

Distanzen: 2-3; H. blanda scheint im Untersuchungsgebiet weniger dispersionsaktiv

zu sein als die vorige Art. Hinweise darauf sind der bisher fehlende Nachweis an

HM und das stark polarisierte Le zwischen den Standorten WaN

und WNo auf einer Strecke von nur 50 m.

Verbreitungsstrategie: intermediärer Typ, 1. Gruppe

Hoplodrina ambigua (119 Individuen)

Distanzen: 3-4 { E r $

Larvalökologie: H. ambigua findet wohl überall eine Lebensgrundlage. Im Inneren

der Wälder wurde die Art jedoch bisher noch nie festgestellt.

Populationsbiologie:. bivoltin : n

Verbreitungsstrategie: r-Stratege, 5. Gruppe; die Häufigkeitsschwa en und das

Häufigkeitsmuster innerhalb des Fangstellennetzes, das dem der Wanderfalter

entspricht, können als Hinweise auf hohe Dispersionsaktivitäten verstanden wer-

den. H. ambigua hat im südbayerischen Raum im Lauf dieses Jahrhunderts grö-

Bere Arealausweitungen unternommen. Regelmäßige Mindest-Flugleistungen von

1-2 km/Generation sind hierbei zu postulieren.

Atypha pulmonaris (5 Indviduen)

Distanzen: 2-3

Larvalökologie: die Raupe dieses an feuchtwarme Standorte gebundenen Eulenfal-

ters lebt an Lyngenkraut Eamenarıe) und an Beinwell Symphytu officinale);

das HM-Stück stammt daher vermutlich vom Ruderal (150-30 m). das Exem-

plar im Garten 6) zumindest vom Berglwaldrand (150 m).

Verbreitungsstrategie: K-Stratege, 5. Gruppe

Spodoptera exigua (2 Individuen)

Distanzen: 4

Verbreitungsstrategie: r-Stratege, 1. Gruppe

Caradrina morpheus

503 Individuen 8,9 % o- Rate

396 markiert 19 Wiederfänge

Tab. 89: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Caradrina morpheus.

SIEDLUNG WALD HALBTROK- m WALD HALBTROK- Dr

1987 | Garten KENRASEN M > 198 Garten Wasserwerk KENRASEN M 2

SiS SIM|SN WaS|WaM WıN HO HM HW Mb SN WaS WaN WNw WNo HO HM HW Au We

I pa. 7 20 26 2 6 46 27 3 1 3 EB DENE De Na a ce Me R = .9 8

so 5 16 17 320 3e.235 Elise asia 2 170 de 14 9 14 20 36 1.22% Sg

99 sa a a Dr wie 2b = 27.9 2 3 en 21800 SR: 9

Mark. 4 16 21 353 ag 2 1922 Mark. 15 6 Ne 37 MR 7 Pa 208

Es handelte sich um ein cd’, das von SiN nach WaS (30 _m) flog.

Im Wasserwerk wurden 1988 2 dd‘ nach jeweils Tagen, wiedergefangen, was

einer durchschnittlichen Wiederfang-Quote von 2,8 % entspricht. Im Garten 1988

wurden 13. dc nach einem 1-Tages-Intervall und nur 3 do‘ nach 2 Tagen rückge-

fangen; die mittlere Verweilzeit beträgt also nur 1,2 Tage! Zweitwiederfänge

ereigneten sich nicht.

Distanzen: 3, ? .

Larvalökologie: die an HM festgestellten 10 Exemplare (auch ein 2) stammen,

nach den in 1984) gene ten Raupenfutterpflanzen beurteilt, zumindest

vom Ruderal (150-300 m entfernt). Diese Distanzen liegen im normalen Bereich

der Dispersionsaktivität dieser Art.

Populationsbiologie: proterandrisch

Verbrei stra gie: r-Stratege, 2. Gruppe; im Gegensatz zu den Behauptungen

BETTMANNs f 86) handelt es sich im Untersuchungsgebiet um eine recht mobile

nike Nacht-zu-Nacht-Fluktuationen sind vor allem am Flugzeitende zu be-

obachten.

Paradrina clavipalpis

Distanzen: 4

Verbreitungsstrategie: r-Stratege, 1. Gruppe

Wiederfang-Quote: niedrig! 1987 erfolgte der einzige bo-m fang schon nach 1 Tag.

149

Eremodrina gilva (41 Individuen)

Distanzen: 4

Verbreitungsstrategie: r-Stratege, 3. Gruppe; diese ursprünglich alpin verbreitete

Art führt seit ca. 1950 eine nach Norden BEE ce eh Hierbei

müssen zumindest in manchen Jahren Distanzen von über 10 km zurückgelegt

worden sein. Die in den letzten Jahren bekanntgewordenen Funde in Mittel- und

Norddeutschland lassen aber weit höhere Werte vermuten.

Das Häufigkeitsmuster innerhalb des Fallennetzes entspricht interessanterweise

dem der Wanderfalter.

Agrotis venustula (57 Individuen)

Distanzen: 2

Larvalökologie: als Futterpflanzen kommen im Untersuchungsgebiet wohl nur Pfei-

fengras \ olinia) und Frauenmantel (Alchemilla rapie)‘ in Frage. Die an HW

festgestellten Stücke (dd) stammen vermutlich vom Würmkanal (mindestens 150

er dem nächstgelegenen Standort dieser Pflanzen. Es bestand kein Sichtkontakt

zur Lichtquelle. Die Distanz von 1 km (Zuflug nach HM) liegt nicht mehr im

Bereich der Dispersionsaktivität dieser kleinen Art.

Verbreitungsstrategie: K-Stratege, 5. Gruppe

CUCULLIINAE

Cucullia lucifuga (1 Individuum)

Distanzen: vermutlich 2-3

Populationsbiologie: im Untersuchungsgebiet vermutlich bivoltin

Verbreitungsstrategie: vermutlich ıintermediärer Typ, 5. Gruppe; in der unteren

Hochebene Südbayerns ist diese Art nur lokal verbreitet.

Cucullia umbratica (11 Individuen)

Distanzen: 2

Larvalökologie: die Puppe überliegt nach KOCH (1984) "oft 1 bis 2 Jahre”.

Verbreit strategie: K-Stratege, 4. Gruppe; die lange a e 1974)

der Art keine hohe Wachstumsraten der Populationen. GYULAI & VARGA (1974

beschreiben C. umbratica allerdings als Wanderfalter.

Cucullia verbasci (viele Raupenfunde)

Distanzen: 2-3

Larvalökologie: 02 Puppe überliegt nach KOCH (1984) öfters. Die Raupen sind auf

Königskerzen (Verbascum) und Braunwurz (Scrophularia nodosa) spezialisiert.

Verbreitungsstrategie: intermediärer Typ, 3. Gruppe; entsprechend dem relativ un-

steten Auftreten der Raupenfutterpflanzen spiegelten die Raupenfunde hohe Aus-

tauschraten wieder. Im Garten wurden Raupen beispielsweise vor 1981 vereinzelt,

1982, 1984 und 1985 beobachtet.

Cucullia scrophulariae (2 Individuen, dazu einige Raupenfunde)

Distanzen: vermutlich 2-3

Larvalökologie: wie C. verbasci, jedoch vor allem an Mehliger Königskerze (Ver-

bascum Iychnitis) zu finden _

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Calophasia lunula (2 Individuen)

Distanzen: 3-4

Larvalökoiogie: die Raupen sind stark spezialisiert und fressen an Leinkraut (Lina-

ria vulgaris, -repens). Die beiden im Garten festgestellten cd’ sind biotopfremd:

Im Umkreis von mindestens 300 m existieren keine Vorkommen der Futterpilanzen.

Populationsbiologie: bivoltin

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe; C. /unula war um die Jahr-

hundertwende herym im Untersuchungsgebiet noch nicht nachgewiesen worden. Erst

WOLFSBERGER 11950; 1953/1954) erwähnt für Schleißheim ein häufiges Vorkom-

men als zune in den 50er Jahren sowie eine allgemeine »Ausweitung der Wohn-

areale«. Ähnliches berichtet LATTIN (1967). Flugleistungen im oben angegebenen

Rahmen dürften hierzu die Voraussetzung gewesen sein.

Brachionycha sphinx

73 Individuen 5,3 % o-Rate

19 markiert 1 Wiederfang

150

ana Mae: niedrig, Stichprobe noch zu klein. Es konnte lediglich 1988 im

escat aS) ein wegen der Lichtwirkung einen Tag "ortstreu” verbliebenes I

rückgef: en werden.

Larvalökologie: an Tragehiien n Laubgehölzen, das HM-cJ 1987 stammt zumindest

aus dem deral (150-300 m).

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Lithophane socia (11 Individuen)

Distanzen: 2

Larvalökoiogie: an verschiedenen Laubbäumen, bisher konnten keine biotopfremden

Tiere festgestellt menden:

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Lithophane ornitopus (1 Individuum)

Distanzen: vermutlich 2

Larvalökologie: wie L. so

Verbreitungsstrategie: lien K-Stratege, 6. Gruppe

Lithophane furcifera (1 Individuum)

Distanzen: vermutlich 2

Larvalökologie: wie L. socia

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Xylena vetusta (7 Individuen)

Distanzen: 2-3

Earsslökslogie: das 1988 an HM festgestellte cd stammt, nach den Raupenfutter-

pflanzen beurteilt, vermutlich vom Ruderal (150-300 m entfernt).

Verbreitungsstrategie: intiermediärer Typ, 5. Gruppe

Allophyes oxyacanthae

84 Individuen 11,0 % o- Rate

73 markiert 1 Wiederfang

Wiederfang-Quote: niedrig; ein an WaS markiertes cd wurde 1987 nach 2 Tagen 30 m

entfernt an SiN rückgefangen. 1988 konnte bei täglichem Fang am Standort Was

yon) 18; narkierten Faltern keiner mehr ein zweites Mal gefangen werden!

Larvalökoiog ie: die Raupen leben an Schlehe, Apfel, Weißdorn und Pflaume. Für die

drei an HM festgestellten SS kommen nur eine in ungefähr 350-400 m Ent-

fernung verlaufende Weißdornhecke oder ein ca. 100 m entfernier einzelstehen-

der Busch als Herkunft in Frage. Die Distanz von 350-400 m wird also entwe-

der relativ oft zumindest von d’d’ bewältigt, oder sie wurde in der Vergangen-

heit von einem fertilen a zurückgelegt, es kam zu einer Kolonisation des ca. I m?

großen Busches und die Nachkommen wurden dann am Licht nachgewiesen.

Verbreitungsstrategie: intermediärer Typ, 3. Gruppe

Griposia aprilina (\ Individuum)

Distanzen: vermutlich 2

Larvalökol : vor allem an Eiche, selten einige andere Laubgehölze; bisher konn-

ten keine biotopfremden Stücke festgestellt werden.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Blepharita satura

21 Individuen 36,8 % o- Rate

19 markiert kein Wiederfang

Wiederfang- Qunte. niedrig, Stichprobe noch zu klein

Distanzen: 2-3

Verbreitungsstrategie: intermediärer Iyp, 5. Gruppe

Blepharita adusta

Distanzen: vermutlich 2-3

Verbreitungsstrategie: vermutlich intermediärer Typ, 5. Gruppe

Antitype chi (3 Individuen)

Distanzen: vermutlich 2-3

Verbreitungsstrategie: vermutlich intermediärer Typ, 5. Gruppe

151

Eupsilia transversa

66 Individuen 43,8 % 9- Rate

64 markiert 2 Wiederfänge

Wiederfang-Quote: niedrig, a wurden im Garten (SiN) 1987 bei täg-

een ang 2 dd’ nach 1 bzw. Tagen, was keinen Hinweis auf Ortstreue dar-

stellt.

Distanzen: 3

Larvalökologie: die Raupen fressen an Laubhölzern, die 1986 an HM festgestellten

Exemplare stammen vermutlich aus dem Ruderal (150-300 m).

Populationsbiologie: proterandrisch

Verbreitungsstrategie: intermediärer Typ, 3. Gruppe

Conistra vaccinii

78 Individuen 53,2:%..0- Rate

77 markiert 4 Wiederfänge

Wiederfang-Quote: die Wiederfang-Quote ist als durchschnittlich einzustufen. Die

einzelnen Rückmeldungen beziehen sich auf ein 9 1987 im Garten, das bei einer

Verweildauer von 6 Tagen (bei täglichem Fang!) viermal die Falle anflog. Ein

wurde 1987 an WaS nach 2 Tagen (und dazwischenliegender fangfreier Nacht

jaedergelangen.

Distanzen: ?2-

Larvalökologie: auch die Raupenfutterpflanzen von C. vaccinii lassen den Schluß zu,

daß das 0 am Standort HM zumindest vom Ruderal her (150-300 m) zugeflogen

sein muß.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Conistra rubiginosa (4 Individuen)

Wiederfang-Quote: 3 Individuen wurden markiert, Wiederfänge waren keine zu

verzeichnen.

Distanzen: vermutlich 2 .

Larvalökologie: die Raupen sind in den ersten Stadien auf Schlehe, Pflaume, Trau-

benkirsche oder Feld-Ahorn angewiesen. Bisher konnten keine biotopfremden

Exemplare nachgewiesen werden.

Verbrei strategie: vermutlich K-Stratege, 6. Gruppe

Agrochola circellaris

15 Individuen 38,5 % o- Rate

13 markiert kein Wiederfang

Wiederfang-Quote: niedrig, Stichprobe noch zu klein

Distanzen: ?2-

Larvalökologie: im Anfangsstadium leben die Raupen an einer Reihe von Laubbäu-

men, biotopfremde Exemplare konnten bisher noch nicht festgestellt werden.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Agrochola macilenta

16 Individuen 16,7 % 9 - Rate

12 markiert kein Wiederfang

Wiederfang-Quote: niedrig, Stichprobe noch zu klein

Distanzen: ?; zwischen WaS und SiN ist auf einer Strecke von nur 30 m ein deut-

licher Häufigkeitsgradient (7:1) festzustellen.

Larvalökologie: wie A. circellaris

Verbreitungsstrategie: K-Stratege, 6. Gruppe; A. macilenta scheint etwas stärker an

bewaldete Strukturen gebunden zu sein als A. circellaris. Dies steht wohl auch

m Zusammenhang mit dem normalerweise lokalerem Vorkommen von A. macilen-

Agrochola nitida (1 Individuum)

Distanzen: vermutlich 2-3

Verbreitungsstrategie: vermutlich intermediärer Typ, 5. Gruppe

152

Agrochola helvola

8 Individuen 40 % o-Rate (Stichprobe zu klein)

5 markiert kein Wiederfang

Wiederfang-Quote: Stichprobe noch zu klein

Distanzen: 2-3

Larvalökologie: wie A. circellaris, bevorzugt jedoch Weide. Das HM-Stück 1986

stammt mit groBer Wahrscheinlichkeit aus dem Ruderal (150-300 m).

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Agrochola litura

19 Individuen 62,5 % o-Rate

16 markiert kein Wiederfang

Wiederfang-Quote: niedrig, Stichprobe noch zu klein.

Distanzen: 2-3

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Agrochola Iychnidis (4 Individuen)

Wiederfang te: ein am Flügel in charakteristischer Weise leicht deformiertes

Exemplar (cd) flog 1988 an WNo 2 Tage nach dem Erstfang die Falle erneut an.

Distanzen: vermutlich 2-3 i

Larvalökologie: wie A. circellaris, das 1986 HM festgestellte Stück stammte mit

großer Wahrscheinlichkeit aus dem Ruderal (150-300 m).

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Agrochola lota

7 Individuen "0 %" o-Rate (Stichprobe zu klein)

6 markiert kein Wiederfang

Wiederfang-Quote: Stichprobe zu klein

Dissen 2-3 :

84) genannten Raupenfutterpflanzen beurteilt, aus mindestens 200 m Entfer-

zugeflogen. Auch die beiden HW-Stücke stammen wohl vom Würmkanal (150 m

entfernt).

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Parastichtis suspecta

Distanzen: 1-2

Larvalökologie: die Raupen sind in den ersten Stadien an Pappelarten gebunden. Das

im Garten 1983 nachgewiesene Stück e hatte mindestens 300 m vom nächstge-

legenen Standort der Futterpflanze her zurückgelegt. Diese Distanz liegt viel-

leicht schon im oberen Bereich der Dispersionsaktivitäten dieser Art.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Cirrhia aurego (1 Individuum)

Distanzen: vermutlich 2

Larvalökologie: vor allem an Buche, jedoch auch an Eiche, Linde, Pappel und Wei-

de; bisher konnten keine Bonn emden Tiere festgestellt werden.

Verbreitungsstrategie: vermutlic -Stratege, 6. Gruppe

Cirrhia togata

30 Individuen 33,3 % o-Rate

27 markiert 1 Wiederfang

Wiederfang-Quote: hoch; Stichprobe noch zu klein. 1987 wurde im Wasserwerk

(WaN) ein d nach 3 Tagen wiedergefangen.

Distanzen: 1-2

Larvalökologie: in den ersten Larvalstadien an Weidenarten gebunden. Im Wasser-

werk spricht der starke Häufigkeitsgradient schon auf einer Distanz von 50 m für

Barrieren gegen die freie Beweglichkeit dieser Art. Da an .WNo die Falle in

einer Weide poatiert war, an den beiden anderen Standorten jedoch ca. 10-20 m

von der nächstgelegenen entfernt, könnten unter Umständen solch kurze Di-

stanzen bereits nicht mehr zur normalen Dispersionsaktivität dieser Art gehören.

Auch die Tatsache, daß C. togata im Siedlungsbereich noch nie festgestellt wurde,

paßt gut in dieses Bild.

Verbreii strategie: K-Siratege, 6. Gruppe

Leu logie: die 4 im Garten festgestellten Exemplare sind, nach den in KOCH

153

Cirrhia icteritia

16 Individuen 54,5 % o9- Rate

11 markiert kein Wiederfang

Wiederfang-Quote: niedrig, Stichprobe noch zu klein

Distanzen: 2-3

Larvalökologie: die Raupe ai in den len Stadien an Sal-Weide (Salix caprea),

Grau-Weide (Salix cinerea) und Espe as tremula) gebunden. Die vier 1987

an HM gefundenen Stücke (auch ein 9), die mit großer Wahrscheinlichkeit aus

dem Ruderal (150-300 m entfernt) stammen, zeigen, daß die Bewältigung solcher

Distanzen für beide Geschlechter kein besonderes Problem darstellt. Auch die bei-

den im Garten nachgewiesenen Exemplare sind aus mindestens 200 m Entfernung

zugeflogen.

erbreitungsstrategie: intermediärer Typ, 5. Gruppe

Cirrhia gilvago (1 Individuum)

Distanzen: vermutlich 2

Larvalökologie: die Jungraupen sind auf Ulmen angewiesen, seltener akzeptieren sie

auch Pappelkätzchen. In unmittelbarer Nähe beider bisheriger Fundorte wachsen

Ulmen.

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Cirrhia ocellaris

Distanzen: vermutlich 2

Larvalökologie: anfangs sind die Raupen auf verschiedene Pappelarten angewiesen,

welche in unmittelbarer Nähe (20 m) des Fundortes vorkommen.

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Cirrhia citrago

11 Individuen 9,1 % o- Rate

11 markiert kein Wiederfang

Wiederfang-Quote: niedrig, Stichprobe noch zu klein

Distanzen: 2-3

Larralßinlogie: monophag an Linde (Tilia); das HM-Stück (Cd) 1986 stammt also aus

dem Waldgürtel und ist mindestens 1 km weit geflogen. Dies stellt vielleicht

schon die Obergrenze der Reichweite dieser Art dar, die Verteilung der restli-

chen Fundorte laßt jedenfalls eine relativ starke Habitatbindung erkennen.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

MELICLEPTRIINAE

Chloridea viriplaca (? Individuen)

Distanzen: 4

Populationsbiologie: bivoltin

Verbreitungsstrategie: r-Stratege, 1. Gruppe

Chloridea peltigera (2 Individuen)

Distanzen: 4

Populationsbiologie: bivoltin (Einflug und Nachfolgegeneration)

Verbreitungsstrategie: r-Stratege, 1. Gruppe

Pyrrhia umbra (7 Individuen)

Distanzen: vermutlich 2-3

Verbreitungsstrategie: vermutlich intermediärer Typ, 5. Gruppe

Panemeria tenebrata (tags 1 Individuum)

Distanzen: 1-2

Larvalökologie: an Hornkraut (Cerastium) und Sternmiere (Stellaria) gebunden. So-

wohl an SıM als auch 1989 im Mallertshofer Holz wurden die tagaktıven Imagines

in unmittelbarer Nähe der Raupenfutterpflanzen beobachtet. Selbst bei Störung

entfernten sie sich höchstens 10-30 m und kehrten teilweise wieder zurück.

erbreitungsstrategie: K-Stratege, 6. Gruppe

154

Axylia putris

243 Individuen 21,6 % o- Rate

125 markiert 5 Wiederfänge

Tab. 9%: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Axylia putris.

WALD HALBTROK- BE:

1988 Garten ‚Wasserwerk KENRASEN M > L

SN WaS eN WNw WNo 2 HO HM HW A e

Z par. 3 14 43 6 42 4 3 14 6 3 138

2 zus - 8 cr SRAE ee — Ne 8

os 3 10 28 5 3 4 2 8 5 2 98

99 >..8 OB aa. AL28

Mark 3 15 36 539 4 Saa12 5 3 125

W.f. = 2 2 = 1 > > = = = 5

Wiederfang-Quote: vergleichsweise hoch; im Garten 1988 konnte bei täglichem Fang

ein d nach 1 Tag (kein Hinweis auf Ortstreue wiedergefangen werden, ein anderes

ie: nach Tagen. Im Wasserwerk ergab sich eine Wiederfang- Quote von

3,8 % bei „inet mittleren Verweildauer, die sich mit 4,0 Tagen auf hohem Niveau

befindet. Eines der 3 dort wiedergefangenen Stücke war ein 9, das den Standort

wechselte 5 m Distanz).

tanzen: 2 \-3?

Populationsbiologie: leicht proterandrisch

Verbreitungsstrategie: intermediärer Typ, 4. Gruppe

BRYOPHILINAE

Euthales algae (25 Individuen)

Distanzen: 3 (-4?)

Larvalökologie: auf Laubholzflechten spezialisiert, nach KOCH (1984) vorwiegend an

alten Stämmen. Die beiden nach HM zu eflogenen Stücke stammen also vermut-

lich aus den umliegenden Wäldern (Flugdistanz mindestens 1 km). Am Würmkanal

scheint eine größere Population zu existieren, die 150 m Mindestdistanz nach HW

wurde von 10 Individuen bewältigt, obwohl kein Sichtkontakt zur Lichtquelle be-

stand.

Verbreitungsstrategi ie: inermediäner DulND. = I} ‚Gruppe: rück 0 ELDER: ennt von dieser

Art nur einen Einzelfund aus nnsbru Südbayerische

Funde wurden später von WOLFSBERGER "(1945- 19; x 0,100, km gerät entlicht,

Berücksichtigt man die relativ gute Durchforschung Süd bayern em Anfang dieses

Jahrhunderts, insbesondere des Untersuchungsgebietes, läßt dies die Vermu-

tung zu, daß diese Art Arealausweitungen in großem Stil” durchführte.

Bryoleuca raptricula

Distanzen: vermutlich 2-3

Lazralckologie: die Raupe ist an Steinflechten, Schildflechten und Algen zu finden.

Verbre strategie: vermutlich intermediärer Typ, 5. Gruppe; um, die Jahrhun-

dertwende bis ca. 1925 wurde diese Art interessanterweise in Südbayern nicht

nachgewiesen. Eine Häufigkeitszunahme dieser Art, sicherlich verbunden mit Are-

ala Beitunge en, unchsı OLFSBERGER (1945- 1949; 1950; 1953/1954; 1958 und

1960 "Nach L ATTIN (1967) gehört B. raptricula tatsächlich zu den Arealerweiterern.

APATELINAE

Panthea coenobita (1 Individuum)

Distanzen: vermutlich 2

Larvalökologie: an Nadelbäume, vor allem Fichte gebunden. Bisher konnten keine

biotopfremden Tiere festgestellt werden.

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe; nach LATTIN (1967) erwei-

terte P. coenobita in diesem Jahrhundert ihr Areal.

era alpium (1 Individuum)

istanzen: vermutlic h 2 X

arralbenionie die Raupe frißt an verschiedenen Laubbäumen. Biotopfremde Di

konnten ae ıer nicht festgestellt werden. Die Tatsache, daß die Puppe (nach

KOCH, 1984) gelegentlich mehrmals überwintert, verringert die potentielle Wachs-

tumsrate der Populationen dieser Art.

erbreitungsstrategie: vermutlich K-Stratege, 5. Gruppe

155

Colocasia coryli

43 Individuen 0% o9-Rate

25 markiert kein Wiederfang

Wiederfang-Quote: niedrig, Stichprobe noch zu gering; im Wasserwerk wurden 1988

im Versuch der verringerten Fallendistanzen 17 dd‘ markiert.

Distanzen: ?2

te gg an Laubgehölzen. Die Biotopbindung dieser Art scheint relativ stark

zu Bein. iotopfremde Tiere, z.B. an HM, konnten bisher noch nicht beobachtet

werden.

Pogulaianebiologie; im Untersuchungsgebiet mit einer mehr oder weniger starken 2.

eneration u: 2,

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Subacronicta megacephala (16 Individuen)

Distanzen: 2

Larvalökologie: als Futterpflanzen dienen Pappel, Espe und Weide; die vier im Gar-

ten festgestellten Stücken stammen daher aus mindestens 200 m Entfernung.

Durch das von KOCH (1984), erwähnte bisweilen stattfindende UÜberliegen der

Puppen kommt es zu einer Verminderung der potentiellen Populations- Wachs-

tumsrate ("PGR").

Populationsbiologie: bivoltin, die 2. Generation jedoch unvollständig

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Acronicta aceris (18 Individuen)

Distanzen: 2

Larvalökologie: die Raupe lebt an verschiedenen Laubbäumen, biotopfremde Tiere

wurden bisher nicht registriert. Bezüglich der Puppenentwicklung gilt das zur

vorigen Art Gesagte. j }

Populationsbiologie: im Gebiet mit einer unvollständigen 2 Generation

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Acronicta leporina (16 Individuen)

Distanzen: 2

Larvalökologie: die Raupe ist auf Laubbäumen, im eg vor allem auf

Birke zu finden. Biotopfremde Stücke waren bisher nicht zu beobachten.

Verbreitungsstrategie: iniermediärer Typ, 5. Gruppe

Apatele alni (1 Individuum)

Distanzen: 2, ' :

Larvalökoiogie: die Raupen akzeptieren als Futter oligophag einige Laubbaumarten,

biotopfremde Tiere traten bisher nicht auf.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Apatele psi (17 Individuen)

Distanzen: 2

Latraldinlogie- an Laubbäumen, jedoch etwas mehr polyphag als A. al/ni. Das d am

S-Bahnhof mußte mindestens 100 m geflogen sein, sonst konnten bisher keine

buptep re mon Tiere Testzesielit werden. 4 }

Populationsbiologie: abgesehen von den beiden August- Tieren univoltin

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Hyboma strigosa (13 Individuen)

Distanzen: 1-2

Larvalökologie: die Raupe ernährt sich oligophag von einigen Laubhölzern bzw.

-gebüschen, biotopfremde Stücke traten bisher nicht auf. H. strigosa scheint stark

habitatgebunden zu sein, was eine Erklärung für das eher lokale Vorkommen in

Südbayern sein könnte.

Verbrei strategie: K-Stratege, 6. Gruppe

Pharetra auricoma (31 Individuen)

Distanzen: 2-3

Larvalökologie: recht polyphag, im Untersuchungsgebiet be vor allem Wei-

de; die an HM anfliegenden Stücke stammen daher wohl aus dem Ruderal (150-300

m entfernt).

ationsbiologie:, bivoltin

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

156

Pharetra rumicis (33 Individuen)

Distanzen: 3(-4?)

Larvalökologie: relativ polyphag, die HM-Stücke stammen vermutlich aus dem Ru-

deral (150-300 m).

Populationsbiologie: bivoltin

Verbreitungsstrategie: r-Stratege, 3. Gruppe

Craniophora ligustri

112 Individuen 6,9 % o- Rate

102 markiert 1 Wiederfang

Teb. 91: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Craniophora ligustri.

SIEDLUNG WALD HALBTROK- Da WALD HALBTROK- DACH

[OSTSEE Garten | KENRASEN M L 1988 Garten Wasserwerk KENRASEN M L

SiS SIM |SN WaS|WaM Wan HO HM HW Mb SN WaS WaN WNw WNo HO HM HW Au We

2 par. 2.9 8 2 1 8 8 - 2 40 2 par. A 7 14 12 3 7 S 1 5 = 53

zus Zn 1 N Ge N - SEE 2Tzusye- 10 a I a ne 10

[oje] ee 1 = NT Elan, = 1 40 de Ace een ae

99 SE Se Me Bene = 5 09 re“ Ar re nV 2

Mark. - 3 6 Pa 1 ABER Merk EA oa 1003 ht a sr

Wi. ui er a - wi - - Se EN u -

Wiederfang te: niedrig, vor allem 1988 im Garten (WaS) beim täglich durch-

geführten Fang.

tanzen: 3

Larvalökologie: die Raupe ernährt sich von Esche (Fraxinus excelsior), Liguster (Li-

gustrum ulzare) und Flieder. Das 1987 an HM festgestellte @ war also minde-

stens 1 km weit vom Flughafenrand herbeigeflogen.

Verbreitungsstrategie: intermediärer Typ, 3. Gruppe

JASPIDIINAE

Jaspidia deceptoria (22 Individuen + tagaktive Exemplare)

Distanzen: 2,

Larvalökologie: nach KOCH ee) an Wiesengräsern, gern an Picscheee: (Phleum

Dentenee). die Art ist im Untersuchungsgebiet mehr an trockene iesen gebun-

den und wurde außerhalb des Flughafens und des Wasserwerks nur einmal am

Würmkanal gefangen (Au). Die Distanz von 1 km bzw. 1,3 km, die einem Zuflug

in den Garten entspräche, liegt offenbar außerhalb des Bereichs der normalen

Dispersionsaktivität.

Verbreitungsstrategie: intermediärer Ip: 5. Gruppe; nach LATTIN (1967) unternahm

J. deceptoria in diesem Jahrhundert Arealausweitungen.

Jaspidia pygarga (2025 Individuen + tagaktive Exemplare)

Distanzen: 2; das PIEUn ehe Verheltnn SiN/WaS betrug 1988 in den parallel durch-

‚eführten Fängen 22:67. Dieser Gradient läßt Barrieren gegen die freie Bevese

ichkeit dieser Art vermuten; im vorliegenden Fall könnte dies der größere Of-

fenlandcharakter an SiN sein. a

Lergalökolon obwohl sich die Raupe von verschiedenen Gräsern ernährt, ist der

Falter relativ stark an Gebüsch- bzw. Baumstrukturen gebunden. Die 5 an HM

festgestellten Exemplare stammen vermutlich aus dem Ruderal (150-300 m).

ulationsbiologie: je nach Beschaffenheit des Biotops kommt es anscheinend zur

inregulierung einer „upezilischen maximalen Populationsdichte. Jahrweise konstant

bewirkte diese im asserwerk ein Fangergebnis von 50 Individuen, an HO 15

Individuen und im Garten (Was) 5 Individuen, jeweils bezogen auf eine durch-

schnittliche Fangnacht in der Hauptflugzeit. In allen Fällen waren diese Werte

recht konstant, stärkere Nacht-zu-Nacht-Fluktuationen, wie sie für einige hochmo-

bile Arten typisch sind, traten nicht auf.

Verbreitungsstrategie: K-Stratege, 4. Gruppe

Eustrotia uncula

Distanzen: vermutlich 2 SB

Larvalökologie: auf Seggen (Carex) und Zune: ras (et spezialisiert. Am

31.7.83 flogen im Garten bei heißem, windstillem Wetter 2 Exemplare dieses

eigentlich Inpischen Moorbewohners zu. Die minimale Flugdistanz (zum nächst-

gelegenen Futterpflanzen-Vorkommen) beträgt ca. 300 m, ein Moor bzw. ein

naturnahes Feuchtgebiet ähnlicher Prägung existierte zu diesem Zeitpunkt jedoch

in einem Radius von mindestens 3 km nicht.

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

157

Eustrotia olivana (27 Individuen + tagaktive Exemplare)

Distanzen: 2a?)

Larvalökologie: die Raupe lebt an verschiedenen Gräsern, im Untersuchungsgebiet

vor allem an Riedgräsern (Carex) und an Land-Reitgras VlCalamagrosfte epige-

jos). Das SiS-Stück stammt wohl aus dem Flughafengebiet und hatte mindestens

00 m zurückgelegt. Die beiden im Garten festgestellten Stücke sind wohl eben-

falls einer Population außerhalb des en zuzuschreiben und dürften

mindestens 500 m, vielleicht sogar 1 km vom Flughafen her zugeflogen sein.

Populationsbiologie: die tagsüber durchgeführten Beobachtungen dieser Art, die vor

allem im Fiughafen-Ruderal und an der Regattastrecke überaus große Popu-

lationsdichten erreicht, lassen auf ähnliche Verhältnisse schließen, wie sie bei J.

pygarga bereits erläutert wurden. Aufgescheuchte Individuen flogen ca. 3-10 m

weit und setzten sich dann wieder. as Zurücklegen von größeren Distanzen

wurde tagsüber nicht beobachtet.

Verbreitungsstrategie: K-Stratege, 6. Gruppe; nach LATTIN (1967) unternahm E.

olivana ın diesem Jahrhundert Arealausweitungen.

NYCTEOLINAE

Nycteola revayana (6 Individuen)

Distanzen: vermutlich ?2

Larvalökologie: monophag an Eiche; an allen bisherigen Fundorten befinden sich in

unmittelbarer Nähe Eichen.

Populationsbiologie: bivoltin

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe; die sehr ähnliche N. asia-

tica KRUL. ist Ko als hochmobile Art, die groBe Distanzen bewältigen kann,

eat siehe ZBANYAI-RESER,1988 und EITSCHBERGER & STEINIGER,

BENINAE

Earias chlorana (9 Individuen)

Distanzen: 2

Larvalökologie: auf Weide (Salix) spezialisiert, bisher konnten keine biotopfremden

Tiere festgestellt werden. Die Raupen verursachen nach KOCH (1984) "gele-

gentlich Kahlfraß in Weidenkulturen".

P mie Curie a im Untersuchungsgebiet im Gegensatz zu den Angaben in KOCH

.c. ] univoltın

Verbreitungsstrategie: K-Stratege, 6. Gruppe; in anderen Gegenden, wo die Art

bivoltin auftritt, können durch die so erhöhte potentielle achstumsrate gele-

entlich stärkere Vermehrungen stattfinden; unter Umständen ist dies an solchen

tandorten mit einer veränderten Strategie verbunden; es könnte sich dann um

r-Strategen handeln.

Bena prasinana (77 individuen)

Distanzen: ?2

ae die Raupe lebt an einer Reihe von Laubbäumen, vor allem an Rot-

Buche (Fagus sylvatica). Biotopfremde Tiere, z.B. an HM konnten bisher noch

nicht festgestellt werden.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Pseudoips bicolorana (1 Individuum)

Distanzen: 1-2

Larvalökologie: monophag an Eiche, bisher keine biotopfremden Stücke

Verbreitungsstrategie: K-Stratege, 6. Gruppe

PLUSIINAE

Chrysaspidia putnami

Distanzen: vermutlich 2-3

Larvalökologie: oligophag an eine Reihe hygrophiler Pflanzen gebunden, die in ca. 50 m

Entfernung vom Fundort vorkommen.

Populationsbiologie:, bivoltin >

Verbreitungsstrategie: vermutlich intermediärer Typ, 5. Gruppe

158

Autographa gamma

885* Individuen 4,9 % o- Rate

642 markiert 2 Wiederfänge

* dazu tagaktiv zahlreiche weitere Exemplare

Teb. 92: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Autographa gamma.

SEDLUNG WALD HALBTROK- DACH WALD HALBTROK- EDGE

1987 arten KENRASEN M 2 1988 Garten Wasserwerk KENRASEN M

SiS SM |SN WaS|WaM WaN HO HM HW Mb SN WaS WeaN WNw WNo HO HM HW Au We

I par. 7 29 50 53 21 66 66 57 122 2 475 X par. 30 34 28 6923 17 12T AT, 1 11

% zus - 78 28 - 8 S3em13 5 26 171 % zus. - 30 - - - = ei S & z

[oje 2 5 28 90 52 20 56 63 61 80 1 456 [e107 14 37 20 5 21 1677012533 1 -

99 EEE SCENE 1 dar 2 One 720 1 2cthen 26 09 RT ET ER enageenie: SEE

Mark. 5 31 96 65 20 62 63 64 81 1 478 Mark. 15 39 20 6 20 16202122536 1 -

wi. a WEL eRe- Ne San - =. wi. = 1 Sa ame e Re

Wiederfang te: sehr niedrig! 1988 wurde im Garten (WaS) bei täglichem Fang

nur 1 0 (=2,6 %) durch die Lichtwirkung "gefangengehalten”. Der einzige unter

"regulären" Bedi en erfolgte Rückfang ereignete sich bei einem an aN

markierten J, das 2 lage später an WNo (50 m) wiedergefangen wurde. Auch die

sehr kurze mittlere Verweildauer zeigt also, daß A. gamma, die in einer Art

"Nullprobe"” als bekannter Wanderfalter markiert wurde, erwartungsgemäß rea-

giert und die Brauchbarkeit der Methode unterstreicht. BETTMANN (1985a; 1986)

erhielt von A. gamma mehr Wiederfänge zurück, was wohl methodisch zu begrün-

‚den ist al Fang).

en:

rare ivoltin; im Untersuchungsgebiet fällt die nieduns o-Rate auf.

ALICKY (1974 a) hatte in Österreich Werte von nahe 50 %, NOWAK (1974) von

29 % beobachtet. ;

Verbreitungsstrategie: r-Stratege, I. Gruppe; auch die heimischen Populationen sind

hochmobil. Starke Wanderbewegungen (zielgerichteter, rascher Flug) wurden

tagsüber und in der Dämmer vor allem im Moos entlang der Entwässerungs-

graben und entlang des Birket-Randes registriert. Die nächtliche Flugaktivität ist

edoch nicht immer mit den Flugmaxima der anderen typischen migranten Arten

{"Wanderfalternächte") korreliert.

Autographa pulchrina (3 Individuen)

Distanzen: vermutlich 3

Verbreitungsstrategie: vermutlich intermediärer Typ, 3. Gruppe

Autographa bractea (3 Individuen)

Distanzen: 3-4

Verbreitungsstrategie: intermediärer Typ, 1. Gruppe; A. bractea ist eine als Wan-

derfalter bekannte Art, die hauptsächlich alpin verbreitet ist. Von diesem Ver-

breitungszentrum aus unternimmt sie Vorstöße über größere Distanzen hinweg. An

geeigneten Standorten, z.B. im Berglwald bei Oberschleißheim, wird A. bracte

edoch zumindest zeitweilig bodenstandig. EITSCHBERGER & STEINIGER (1980

bezeichnen sie als Arealerweiterer.

Macdunoughia confusa (12 Individuen)

Distanzen: 4

Populationsbiologie: bivoltin : ON

Verbreitungsstrategie: r-Stratege, 1. Gruppe; die starken Häufigkeitsschwankungen

von Jahr zu Jahr sowie von Generation zu Generation sind ein weiterer Hinweis

auf die hohe Mobilität dieser Art.

Plusia chrysitis (58 Individuen)

Distanzen: 3-4

De meh ogie: bivoltin, Flugzeit etwas später als P. tutti (vergleiche REICH-

Verbreitungsstrategie: r-Stratege, 2. Gruppe

Plusia tutti (42 Individuen)

Distanzen: 3-4 3% \

Kup len ivoltin, Flugzeit etwas früher als P. chrysitis (vergleiche

ICHHOLF, 1985).

erbreitungsstrategie: r-Stratege, 2. Gruppe

159

Plusia chryson

Distanzen: vermutlich 2-4

Larvalökologie: die Raupe lebt an Wasserdost (Eupatorium cannabinum) und Kleb-

Salbei (Salvia giutinosa). Die Mindest-Flugdistanz vom nächstgelegenen Standort

der Futterpflanzen zum Fundort beträgt ca. 300 m. KOCH erwähnt, daß die Art

als seltener Irrgast in Gebiete zufliegen kann, die keine eigene Populationen

besitzen, also über viele Kilometer hinweg.

erbreitungsstrategie: vermutlich intermediärer Typ, 5. Gruppe

Polychrysia moneta

Distanzen: 2-4

Larvalökologie: die Raupe ist auf Eisenhut- a und Ritterspornarten (Del/-

hinium) sowie auf Trollblume (Trollius europaeus) zu finden. Eine nicht unerheb-

iche Rolle für die Verbreitungsstrategie dieser Art dürften daher Gärten spielen.

Im Garten des Verfassers standen fie Pflanzen zur Verfügung. Die drei (in drei

verschiedenen Jahren beobachteten) Exemplare wurden also ın ihrem typischen

Habitat gefunden.

Populationsbiologie: bivoltin, die 2. Generation ist oft unvollständig und konnte im

ntersuchungsgebiet noch nicht en werden. _

Verbreitungsstrafegie: intermediärer Iyp, >. Cape die Art unternahm im Lauf

dieses Jahrhunderts Arealausweitungen in den Nordwesten Mitteleuropas hinein,

en) z.T. größere Distanzen bewältigt werden mußten (KOCH, 1984; LATTIN,

Chrysoptera c-aureum (3 Individuen)

Distanzen: 2 (-3?)

Darren. die Raupe ist auf Wiesenrautearten (Thalictrum) und Akelei (Aqui-

legia) spezialisiert. Im Untersuchungsgebiet werden offensichtlich auch die im

Garten angepflanzten Akeleisorten angenommen, wie die regelmäßigen Funde der

letzten 3 Jahre vermuten lassen.

Merbreiiungestzairgi: K-Stratege, 6. Gruppe; C. c-aureum wurde somit im Mün-

chener Norden erstmals außerhalb der Isarauen nachgewiesen; ein solcher Koloni-

sationsvorgang hätte im Lauf dieses Jahrhunderts über eine Mindestdistanz von 7

Kilometer erfolgen müssen.

Abrostola triplasia (12 Individuen)

Distanzen: ? (-3?) “

Larvalökologie: monophag an der Großen Brennessel (ancz dioica); diese war in

der unmittelbaren Nähe aller bisherigen Fundorte zu finden.

Populationsbiologie: bivoltin

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Abrostola asclepiadis (12 Individuen)

Distanzen: ?

rer teserr ie monophag am Schwalbenwurz (Cynanchum vincetoxicum), welcher im

Berglwald und im Schweitzerholz an vielen Stellen recht häufig ist. Die im

Garte ger denen Stücke (nur dc) stammen also mindestens vom Berglwald-

rand (20 m), das WaN-d’‘ mußte auch zumindest 150-200 m von der nächstgele-

genen Raupenfutterpflanze her geflogen sein. h i k

Eupulatiomiolage: im Gegensatz zu den beiden anderen im Gebiet nachgewiesenen

rten der Gattung univoltin

Verbreitungsstrategie: K-Stratege, 5. Gruppe; ee der zerstreuten Ver-

breikunn der Raupenfutterpflanze scheint es sich A. asclepiadis nur in Jahren mit

uter Bestandsentwicklung (z.B. 1987 und 1988) erlauben zu können, Vorstöße

er biotopfremdes Gebiet durchzuführen.

Abrostola trigemina (12 Individuen)

Distanzen: ? (-3?)

Larvalökologie: monophag an der Großen Brennessel; das HM-Exemplar stammt

also zumindest aus dem Ruderal (150-300 m). Der Häufigkeitsgradient im Was-

serwerk könnte auf Barrieren gegen die freie Mobilität hinweisen.

Populationsbiologie:, bivoltin

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

160

CATOCALINAE

Astiodes sponsa (1 Indviduum)

Distanzen: 2-4, aber relativ biotoptreu

Larvalökologie: monophag an Eiche, welche sich auch in unmittelbarer Nähe des

Fundortes befand. PR

Verbreitungsstrategie: intermediärer Typ, 1. Gruppe; A. sponsa ist als Wanderfal-

ter bekannı, dürfte im Untersuchungsgebiet in den älteren Eichenbeständen je-

doch bodenständige Populationen besitzen.

Catocala nupta (4 Individuen)

Distanzen: 2-3

Larvalökologie: an Weiden und Bappein: das an HM gefangene < ist vermutlich aus

dem Ruderal bei einer Distanz von 150-300 m zugeflogen. Aus ähnlichen Minde-

stentfernungen stammen die regelmäßig im Garten festgestellten Stücke.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Callistege mi

Diese tagfliegende Art wurde in der Auswertung unherücksichtigt gelassen, da sie mit

Lichtfallenfängen nicht erfaßbar ist. C. mi ist im Untersuchungsgebiet offensichtlich

univoltin, sie ist wohl als intermediärer Typ der 5. Gruppe einzustufen.

Ectypa glyphica

Wie Callistege mi, es wurde lediglich 1 Exemplar der 2. Generation beobachtet. In-

teressant ist wie bei der vorhergehenden Art der bisher fehlende Nachweis im Garten.

OPHIDERINAE

Scoliopteryx libatrix (15 Individuen)

Distanzen: 2-3

Larvalökologie: die Raupe ernährt sich von Weide und Pappel; das HM-Stück ist

also als Zuflieger aus einer Distanz von mindestens 150-300 m (Ruderal) zu

werten. Auch die vergleichsweise oft im Garten zu beobachtenden Stücke (auch

22) mußten 200-300 m vom nächstgelegenen Standort der Futterpflanzen er

stehende Weiden) geflogen sein.

Populationsbiologie:, bivoltin -

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Lygephila pastinum (38 Individuen)

Distanzen: 2; der siarke Häufigkeitsgradient im Wasserwerk spricht für Barrieren

gegen die freie Beweglichkeit dieser Art.

Larvalökologie: an verschiedenen Schmetterlingsblütlern; im Ort, wo die Vertreter

dieser Pflanzenfamilie nur sehr spärlich auftreten, wurden von L[. pastinum daher

nur vereinzelt (zugeflo ene) Exemplare registriert.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Parascotia fuliginaria (5 Individuen)

Distanzen: 1-2

Larvalökologie: es handelt sich um eine relativ stenöke Art, deren Raupen von

mindestens 200 m vom nächsten möglichen Ort der Larvalentwicklung her ge-

ogen sein.

ulationsbiologie: im Garten (WaS) fällt das regelmäßige Auftreten (dd + 99)

ei sehr geringen Häufigkeitsschwankungen auf. Es handelt sich vermutlich um

eine kleine Population im Gleichgewichtszustand. Ein solches Gleichgewicht auf

niedrigem Niveau dürfte für die in Südbayern nur lokal und selten anzutreffende

P. fuliginaria typisch sein. Schon 30 m entfernt (SiN) konnte bisher kein Stück

nachgewiesen werden.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Phytometra viridaria (3 Individuen + tagaktive Exemplare)

161

Rivula sericealis (219 Individuen)

Distanzen: 2

Larvalökologie: die Raupe ernährt sich yon Gräsern; in den Wäldern ist diese Art

bisweilen recht häufig, im Offenland (HM) war R. sericealis jedoch bisher nicht

zu beobachten, was auf einen Aktionsradius von unter 1 km hinweist.

Populationsbiologie: bivoltin

Verbreitungsstrategie: intermediärer Typ. = FUuNE: die starken Häufigkeitsschwan-

kungen im Garten von Jahr zu Jahr könnten als Indiz dafür gewertet werden, daß

R. sericealis hier verstärkt als zufliegender Gast auftritt.

HYPENINAE

Laspeyria flexula

269 Individuen 15,9 % 9- Rate

43 markiert kein Wiederfang

Wiederfang-Quote: niedrig

Distanzen: 2-3 _

Larvalökologie: die aan ist auf Rindenflechten und -algen vor allem von älteren

Baumbeständen spezialisiert. Dementsprechend sind die beiden an HM registrier-

(D: dd als Gastarten zu charakterisieren, die vermutlich vom Flughafenrand

Distanz: 1 km) stammen. Eines 08 bei ruhigem windstillen Wetter, das andere

e Häufig EIERERRICHE HO/HM 1986 verdeut-

bergrenze der normalen Dis-

bei leichtem Westwind an. Der star

licht jedoch, daß diese Distanz wohl schon die

persionsaktivität von L. flexula darstellt.

Verbreitungsstrategie: intermediärer Typ,, 5. Gruppe

Colobochyla salicalis (1 Individuum + einige tagaktive Exemplare)

Distanzen: 1-2

Larvalökologie: die Raupen sind nach KOCH (1984) auf die zartesten Triebe von

Weide und Pappel spezialisiert. Bisher konnten keine biotopfremden Tiere nach-

gewiesen werden.

Verbreitungsstrategie: K-Stratege, 6. Gruppe; C. salicalis tritt nicht nur im Unter-

suchungsgebiet sondern auch in Südbayern allgemein nur lokal und einzeln auf.

Herminia barbalis (2 Individuen)

Distanzen: 2

Larvalökologie: die Larvalentwicklung spielt sich an Laubhölzern ab; im Offenland

wurde die Art bisher noch nie, an etwas offener strukturierten Standorten (SiM,

Mo/1985) insgesamt nur in 2 Exemplaren beobachtet. Entfernungen

über 500 m über biotopfremdes Gebiet scheinen eine Barriere gegen die Ver-

breitung darzustellen.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Zanclognatha tasipennalis (74 Individuen)

Distanzen: 2-3

Larvalökologie: die Raupen sind nach KOCH (1984) auf "herabgefallene, faule und

modernde Blätter von Gräsern, niedrigen Pflanzen und Laubgehölzen" speziali-

siert. Dies stellt eine interessante Einnischung dar, die Z. tarsipennalis auch von

der folgenden sehr ähnlichen Art, Z. tarsicrinalis unterscheidet. Z. tarsipennalis

ist im Untersuch Eier recht stenök an die Ränder und Gebüschzonen der

etwas trockeneren Wälder gebunden. Das J an HM ist jedoch nicht zwingend als

biotopfremd zu charakterisieren. Im Moos konnte bisher_kein Exemplar nach-

gewiesen werden. Die Distanz von ca. 3 km, die zu einem Zuflug nötig wäre, liegt

außerhalb der normalen Dispersionsaktivität dieser Art.

ala. im Untersuchungsgebiet bivoltin, die 2. Generation ist jedoch un-

vollständig.

Verbrei strategie: intermediärer Typ, 5. Gruppe; OSTHELDER en er-

wähnt für das südbayerische Faunengebiet nur Beuerberg und Herrsching als

Fundort. Ob dies lediglich an Bestimmungschwierigkeiten lag, oder ob tatsächlich

En: on EEE über größere Distanzen stattgefunden hat, muß hier noch

offenbleiben.

162

EEE

Zanclognatha tarsicrinalis (236 Individuen)

Distanzen: 2

Larvalökologie: im Gegensatz zur vorigen Art ernähren sich die Raupen von her-

abgefallenen, trockenen Blättern von Himbeere, Brombeere und Waldrebe (Cle-

matis vitalba). Z. tarsicrinalis ist im Untersuchungsgebiet uy an etwas feuch-

tere Wälder und deren Ränder gebunden. Im Offenland (HM) konnte die Art

bisher noch nicht nachgewiesen werden. ! Be

ationsbiologie: im Gegensatz zu Z. tarsipennalis im Untersuchungsgebiet of-

ensichtlich univoltin

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Zanclognatha grisealis (19 Individuen)

Distanzen: 2

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Trisateles emortualis (32 Individuen)

Distanzen: 2, KR a

Larvalökologie: oligophag an einigen Laubbäumen, vor allem an deren abgefallenen

Blättern. Die Art scheint sehr biotoptreu zu sein: An den Stellen des Vorkommens

befinden sich in unmittelbarer Nähe Eichen oder Buchen.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Hypena obsealis

Distanzen: vermutlich 2

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Hypena proboscidalis

171* Individuen 19,7 % 9- Rate

61 markiert 2 Wiederfänge

* dazu viele tagaktive Exemplare

-Quote: sehr niedrig! Im Garten (WaS) wurden 1988 bei täglichem Fang

21 arleerte Tiere freigelassen, von denen nur 2 dd (9,5 r nach jeweils 1_Tag

wiedergefangen wurden. Im Wiederfangergebnis fehlen also Hinweise auf Orts-

treue.

Distanzen: 3° _

Larvalökologie: die 5 an HM festgestellten Exemplare (auch ein 9) stammen, nach

den in KOCH (1984) genannten Raupenfutterpflanzen beurteilt, zumindest aus dem

Ruderal (150-300 m), derartige Distanzen liegen im Bereich der normalen Disper-

sionsaktivität. _

Populationsbiologie: im Untersuchungsgebiet bivoltin; proterandrisch

Verbreitungsstrategie: 1-Stratege, Zuppe- in der 2. Generation war die Flugak-

tivität mit der der typischen Wanderfalter in den Wanderfalternächten korreliert.

rpenz proboscidalis ist die mobilste der im Untersuchungsgebiet festgestellten

ypeninen, ihre Strategie erinnert an die einiger typischer r-Strategen unter den

Tagfaltern, deren Larvalentwicklung ebenfalls hauptsächlich an wer: abläuft,

als Beispiele mögen hier das Landkärtchen (Araschnia levana) oder der Kleine

Fuchs { glais urticae) genügen.

GEOME TRIDAE

ARCHIEARINAE

Da diese tagaktive Art in Lichtfallenfängen nicht zu erfassen ist, wurde sie bei den

ae ausgeklammert. Es handelt sich um einen monophagen Birkenbewohner

nur selten wird Rot-Buche akzeptiert), der vermutlich als K-Stratege der 6. Grup-

e zu charakterisieren ist. Innerhalb des Birkets wurde A. parthenias jedoch bei

Ben bis zu 300 m Distanz beobachtet (von Baumwipfel zu Baumwipfel).

163

OENOCHROMINAE

Alsophila aescularia

1987

I par.

I zus.

Mark.

W.f.

148 Individuen 0 %* o9-Rate

129 markiert 10 Wiederfänge

* die 99 sind flugunfähig, an WaS wurde 1987 ein 9 an einem Baumstamm sitzend

gefunden.

Tab. 93: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Alsophila aescularia.

Ve niedrig! Mit einer Ausnahme erfolgten die Wiederfänge 1987 im

Garten bei täglichem Betrieb der Lichtfalle und sind auf eine "Gefangenschaft"

im Bann der Lichtquelle zurückzuführen! Die mittlere Verweildauer betrug hierbei

niedrige 1,5 Tage bei einem Maximum von 3 Tagen. Verglichen beispielsweise mit

dera ebenfalls ın einer extremen Jahreszeit und nur im männlichen Geschlecht

fliegenden Frostspanner Operophthera orumata sind diese Werte als sehr niedrig

einzustufen.

Ein d wechselte den Ort (WaS>SiN=30 m) in der Zeitspanne von einem Tag.

Distanzen: Sc 2 -3?), 99 1; im Wasserwerk könnte der Häufigkeitsgradient ein

Hinweis darauf sein, daB Barrieren gegen die Mobilität der dd‘ existieren. Der

Standort WNw zeichnet sich dadurch aus, daß hier unter den 3 Wasserwerk-

Beneplätzen die Schneeschmelze am spätesien erfolgt. Dies behindert unter Um-

ständen den Schlüpfvorgang von A. aescularia.

Larvalökologie: an Laubbäumen und -sträuchern; an HM wurde bisher kein Exem-

plar festgestellt, die Distanz von 1 km scheint außerhalb der normalen Disper-

sionsaktivität zu liegen. Im Moos 1989 wurden in Parallelfängen am Waldrand 7

ESCiDp are, 45 m entfernt davon im Offenland nur noch 2 Exemplare festge-

stellt.

Verbreitungsstrategie: K-Stratege, 5. Gruppe (was die dd betrifft); die dd ge-

währleisten durch die vergleichsweise hohe Mobilität eine gute Gendurchmi-

schung. Für eine Verbreitung über besonders "flugfähige” Eironpen wie bei Orgyia

recens gibt es keine Indizien. Es könnte höchstens eine Verdriltung von sich ab-

seilenden Räupchen eine Rolle spielen. Die erwachsenen een NORD ich im

Juni an der Basis der Futterpflanze (CARTER & HARGREAVES, 1967), Sie

scheiden ais Verbreitungsstadium aus. Vielleicht sind auch die mit langen Beinen

versehenen 99 in der Lage, die für die Verbreitungsstrategie der Art notwendigen

Strecken zurückzulegen oder zumindest die Eier auf eine hohe "Startposition” in

den Bäumen zu bringen.

Der Feinddruck durch Singvögel sowie Störungen durch Parasiten spielen bei die-

ser Art aufgrund der extremen jahreszeitlichen Einnischung eine geringere Rolle

als bei den Sommerarten. Das Gleichgewicht einer Population ist dadurch wohl

SIEDLUNG WALD HALBTROK- "DA WALD HALBTROK- "DA!

| Garten | KENRASEN Moos“ 2 1988 Garten Wasserwerk KENRASEN Moos

SiS SIM ISIN WaS |WaM WN HO HM HW Mb SN WaS WaN WNw WNo HO HM HW Au We

2InSUVB IE 2 = = 24 2 pers (8, 42 I el! „a2, 2 Haze

- mar 30 vw See WM 10 - 65 zzus Sg se Aug ARE

2443 a6 8 50 = 76 Mark. 8 7 Ur 2732 2 = 0 a2

aus far mE er =; = - 10 wi. = 0 = = 0 = 0 UBER

weniger störanfällig. Dies ist eine der Voraussetzungen für eine K-Strategie, denn

der Totalausfall einer lokalen Population könnte nur schwer wieder durch koloni-

sierende Individuen wettgemacht werden.

Odezia atrata

Auch diese tagaktive Art wurde bei der Auswertung ausgeklammert, da sie in Licht- |

fallenfängen nicht erfaßt wurue. Es handelt sich vermutlich um eine Art inter-

mediären Strategietyps die in die 5. Gruppe einzuordnen ist.

GEOMETRINAE

Geometra papilionaria (2? Individuen)

Distanzen: ?

ge oligophag an einer Reihe von Laubgehölzen; das an HM 1986 fest-

gestellte Stück stammt zumindest aus dem Ruderal (150-300 m).

Verbreitungsstrategie: K-Stratege, 6. Gruppe

164

Comibsena pustulata

Distanzen: 1-2

Larvalökologie: monophag an Eiche. Die vier 1986 gefangenen Exemplare flogen die

an HO postierte Falle an, in deren unmittelbaren Nahe größere Eichenbestände

existieren.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Hemithea aestivaria (30 Individuen)

Distanzen: 2; der naufigkeitssradient zwischen WaN und WNo auf nur 50 m Sitrek-

ke weist vielleicht auf Barrieren gegen die Verbreitung hin.

Larvalökologie: die Raupenfutterpflanzen weisen H. aestivaria als Art der Wälder

und (yor Ilem) der Waldränder aus. Demenisprechend konnte bisher im Offen-

land \HM) kein Exemplar nachgewiesen werden. Die normalerweise im Rahmen

A epersionsaktivität und des trivial movement zurückgelegten Distanzen liegen

wohl unter m

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Thalera fimbrialis (16 Individuen)

Distanzen: 2-3 h

Larvalökologie: die Raupen fressen an einer Reihe von niedrigwachsenden Pflan-

zen, die I. fimbrialis als xerothermophil auszeichnen. Im Flughafengebiet stellt

wohl die Gemeine Schafgarbe (Achillea millefolium) die Haupt Nahrunssnucile

zugeflogen sein (Mindestdistanz 700 m). Die Gemeine Schafgarbe wächst jedoch

auch hier vereinzelt, Im Siedlungsgebiet wurde die Art noch nie beobachtet. Die

istanzen von | N (Zuflug in den Garten), vielleicht aber auch von 300 und 700 m

> SiS und SiM), werden über biotopfremdes Gebiet offensichtlich nicht bewäl-

tigt.

Verb räirmusetraiegie K-Stratege, 6. Gruppe

Hemistola chrysoprasaria (2 Individuen)

Distanzen: vermutlich 2; dies gilt vielleicht nur für das trivial movement von H.

ar KOCH 1984) erwähnt ein neuerliches Auftreten in Sachsen, wo-

bei sicherlich größere Distanzen zurückgelegt worden sein müssen.

Larvalökologie: die Raupen dieser Art sind auf Waldrebe (Clematis vitalba, -viti-

Ene) spezialisiert, welche im Wasserwerk, dem Fundort dieser Art, vorkommt.

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe;

Iodis lactearia (16 Individuen + einige dämmerungsaktive Exemplare)

Distanzen: 1-2

Larvalökologie: die Raupenfutterpflanzen charakterisieren diese Art 1 Bewohner

der etwas feuchteren Wälder und deren Ränder. Im Offenland (HM ‚ aber auch

n baum/buschbestandenen Standorten mit etwas stärkerem Offenlandcharakter

SiM, WaN, HW, Mo/1985) konnte kein Exemplar nachgewiesen werden. Die Dis-

ersionsaktivitäten von J. Jlactearia über biotopfremdes Gebiet scheinen sich in

ereichen von wenigen 100 m abzuspielen, wenn nicht sogar unter 100 m.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

STERRHINAE

Sterrha serpentata

Distanzen: 1-2; tagsüber wurden Flüge über ca. 50 m beobachtet.

Verbreitungsstrategie: K-Siratege, 6. Gruppe

Sterrha muricata (15 Individuen)

Distanzen: 1-2; das im Untersuchungsgebiet wie auch überregional nur lokale Vor-

kommen ist ein erster Hinweis auf vergleichsweise geringe Mobilität. Distanzen

von 1,2 Zuflug in den Garten) und 900 m (Zuflug, Wasserwerk? WaM und

HW>SiS) liegen außerhalb der normalen Dispersionsaktivität.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Sterrha biselata (94 Individuen)

Distanzen: 2; im reinen Offenland (HM) wurde diese vor allem an Waldränder ge-

bundene Art noch nie gefunden. Die im "trivial movement” zurückgelegten Di-

stanzen liegen im Bereich von wenigen 100 m und wohl eher darunter. Ein Kilo-

meter wird so gut wie nie bewältigt.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

165

1-2 SIEDLUNG WALD HALBTROK- "DA!

Sterrha inquinata (1 Individuum)

Distanzen: vermutlich 2

Larvalökologie: die Raupen leben an trockenen Pflanzenresten; 5. inquinata wird wie

die folgende Art vor allem in Sekundärlebensräumen, die vom Menschen geschaf-

fen wurden, angetroffen. Siehe Bemerkungen bei S. seriata.

Populationsbiologie: bivoltin

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Sterrha seriata (3 Individuen)

Distanzen: vermutlich 2

Eee die Raupe frißt "dürre, verwelkte oder modernde Reste von nie-

drigen Pflanzen und Laub, Moos und Flechten" (KOCH, 1984). In Mitteleuropa

scheinen sekundäre Lebensräume in der Nähe menschlicher Behausungen bevor-

zugt zu werden. Vielleicht fand $. seriata auf diese Weise einen stabileren Le-

bensraum als das von Natur aus vorprogrammiert war. Beobachtungen des Verfas-

sers in Südeuropa (Süditalien kennzeichnen die dortigen Populationen dieser Art

eher als stark dynamisch, die ubiquitär anzutreffenden Individuen als mobil, und

die Strategie im r-K-Kontinuum mehr in Richtung r-Ende verschoben.

Populationsbiologie: bivoltin

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Sterrha dimidiata (41 Individuen)

Distanzen: 2, wie Sterrha biselata

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Sterrha emarginata (1 Individuum)

Distanzen: vermutlich 2-3 N

Verbreitungsstrategie: vermutlich intermediärer Typ, 5. Gruppe

Sterrha aversata

252 Individuen 10,9 % 9 - Rate

191 markiert 8 Wiederfänge

Tab. 94: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Sterrha aversala.

WALD HALBTROK- "DA

1+2

Garten | KENRASEN MOOS 2 1988 Garten Wasserwerk KENRASEN M ı

SiS SIM |SiN WaS|WaM WN HO HM HW Mb SN WaS WaN WNw WNo HO HM HW Au We

> par 1,031 un, Bl Eor>. SUB 6 68. ,22.pan 22 „37 9,.20 Na. ae

res a2: At na. 43 - oe a ul Sm u ran,

= BouTe RR RE ARE NET. 3 50 de 20 55 8: 16.1057. 200 - Iias RES

a2 DU Hermes == - 609 - 7 SNmaN ia SG SERIE 1; +4

iz 8 19 ARE Va SZ 3 56 Mark. 19 61 Braga Ze Amann

a AN ee - =. WAR. aa\5 er SEE Te

Wiederfang-Quote: in beiden Generationen niedrig! An WaS sind alle 5 Wieder-

& e methodisch zu erklären: Beim täglichen Betrieb der Falle wurden die Stücke

(30) durch die Lichtwirkung einen Tag lang festgehalten. Längere Fang/Rückfang-

intervalle waren hier nicht zu beobachten und es ereignete sich auch kein Zweit-

wiederfang.

Ein & wechselte von einer Nacht zur nächsten den Ort (WaS>SiN) und 2 oc

wurden SiN unter "regulären" Bedingungen (fangfreie Nacht dazwischen) wieder-

‚gefangen. Auch hier betrugen die Verweilzeiten jeweils nur 2 Tage.

Distanzen: ?-3,; 1988 waren an WaS auch starke Nacht-zu-Nacht-Fluktuationen

festzustellen.

Larvalökologie: obwohl S. aversata ihre Raupenfutterpflanzen so gut wie überall

auffinden könnte, ist diese Art im reinen Ofienland (HM) vermutlich nicht boden-

ständig; sie bevorzugt geschütztere Standorte anddeikeg‘ Schutz vor Luft-

feinden), wie dies auch die leichten Häufigkeitsgradienten im Wasserwerk und i

Garten zeigen. Die beiden also offensichtlich nach HM zugeflogenen dd 1987)

wurden in Nächten mit starkem bzw. leichtem Westwind festgestellt.

Populationsbiologie: bivoltin; die 99 sind an offeneren Standorten unterrepräsen-

iert, für sie ist zur Gewährleistung der Fortpflanzung offenbar ein stärkeres

Schutzverhalten entwickelt als bei den dd. h

Die f. remutata L. überwiegt im Untersuchungsgebiet 1986-1988 unabhängig von

SEHEhleehl, Standort, Generation und Jahr mit ca. 80 % der Gesamt-Individuen-

zahl.

Verbreitungsstrategie: intermediärer Typ, 3. Gruppe

166

ee a

——— er

Cyclophora albipunctate (20 Individuen + einige dämmerungsaktive Exemplare)

Distanzen: 1-2

Larvalökologie: die Raupe ist auf Birke spezialisiert und akzeptiert nur selten Eiche

oder Erle; biotopfremde Stücke konnten bisher noch nicht nachgewiesen werden.

Der Häufigkeitsgradient zwischen WaN und WNo (50 m) könnte ein Indiz dafür

sein, daß schon die Distanz von ca. 70 m (=Entfernung von den in der Nähe von

WaN stehenden Birken). eine Barriere gegen die Mobilität dieser Art darstellt.

Populationsbiologie: bivoltin

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Cyclophora punctaria

439 Individuen 42,0 % o- Rate

307 markiert 23 Wiederfänge

Tab. 95: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Cyclophora punctaria.

ı-3 SIEDLUNG WALD HALBTROK- "DACH, 1-3 WALD HALBTROK- esse

1987 | Garten | KENRASEN MOOS” % 1988 Garten Wasserwerk NRASEN M ®

SiS SIM|SIN WaS |WaM WN HO HM HW Mb SN WaS WaN WNw WNo HO HM HW Au We

or TE MEN RR) EN lec) 5 en dor ar ae ea. Er

dc 1 502 BwezInd- Kg 1) Se2 3 24 de 17 89 aa ans Ta a, u 160

99 Sog Kir Sue) 2 1 19.09 16 56 rerineiätg- Hozah 20. 08

Mark. - - 3 La I OR er er 3 AoyE"Markül, 32) Nasa, 1a ae a 267,

Wit. EHEN T- a er a ae S Wi Sr er een 228

Wiederfang-Quote: relativ niedrig; der einzige unter "normalen" Bedingungen, d.h.

bei dazwischenliegenden fangfreien Nächten wiedergefangene Falter war ein d an

HO nach 6 Tagen.

Im Garten ergab sich 1988 bei täglichem Fang folgendes Bild: Sämtliche 19 Erst-

wiederfänge eriolsien nach einem Intervall von nur I Tag! Nur 3 Tiere wurden

ein weiteres Mal gefangen, zwei davon wieder nach einem 1-Tages-Intervall,

eines, ein Weibchen, nach 3 Tagen, was eine Verweildauer von 4 Tagen ergibt.

Die mittlere Verweildauer beträgt demnach sehr niedrige 1,3 Tage.

Die Wiederfangquote 1988 im Garten betrug in der 1. Generation 4/11=36,4 %,

in der 2. Generation 13,6 %. Vielleicht spielen hierbei unterschiedliche Disper-

sionsaktivitäten eine Rolle.

Die 09 sind im Wiederfangergebnis mit 2/19=10,5 %, unterrepräsentiert, sie

‚scheinen j.gseh längere Verweilzeiten zu aufzuweisen (s.o.).

Distanzen:

Larvalökologie: die Raupe lebt monophag an Eiche und nimmt nur selten Birke an.

An HW trat C. punctaria in beiden Jahren erst in der y Generation auf, hier al-

lerdings nicht selten. Einzelstehende Eichen (2-3 m hoch) befinden sich in ca. 100 m

Entfernung. Dies könnte ein Hinweis darauf sein, daB die Falter der I. Gene-

ration recht ortstreu sind, die der 2. Generation in beiden Geschlechtern Distan-

zen von mindestens 100 m jedoch bioblemies bewältigen.

Die Distanz von 1 km über biotopfremdes Gebiet (Zuflug nach HM) liegt in bei-

den Generationen nicht im Bereich der normalen Dispersionsaktivilät.

Populationsbiologie: bivoltin + eine partielle 3. Generation; in beiden Generationen

Wenie proterandrisch!' Während der Flugzeit werden immer wieder Schübe von

frischen Faltern festgestellt, die auf eine kontinuierliche Emergenz hindeuten.

Vielleicht spielt bei dieser Art die Mortalität eine größere Rolle als bei anderen

Arten. l

Verbreitungsstirategie: K-Stratege, 5. Gruppe

Cyclophora linearia (4 Individuen)

Distanzen: vermutlich wie C. punctaria

Larvalökologie: die Raupe ist vor allem an Rot-Buche, jedoch auch an Eiche, Birke

ung. Heidelbeere zu finden. Bisher konnten keine biotopfremden Tiere festgestellt

werden.

Populationsbiologie: im Untersuchungsgebiet bivoltin

Verbreitungsstrategie: vermutlich K-Stratege, 5. Gruppe

Calothysenis griseata (112 Individuen + tagsüber einige weitere Exemplare)

Distanzen: 3

Larvalökologie: an Ampfer- (Rumex) und Knötericharten (Polygonium, Fallopia) ge-

bunden, die 3 HM-Exemplare 1986 stammen zumindest aus dem Ruderal (150-300 m).

Populationsbiologie: bivoltin

Verbreitungsstrategie: r-Siratege, 3. Gruppe

167

Scopula immorata (66 Individuen + tagsüber einige weitere Exemplare)

Distanzen: ?; für die ung der Distanz von 1 km (z.B. ein Zuflug vom Flug-

hafengebiet in den Garten) konnte noch kein Nachweis erbracht werden,

Larvalökologie: im Elushalenneliel. dürfte unter den in KOCH 1984) enannten

Raupenfutierpflanzen vor allem die Gemeine Schafgarbe (Achillea mille olium) in

Frage kommen. Die Art ist somit in einer besonderen Weise an die Schafbe-

weidung angepaßt, da diese Pflanzen von den Schafen stehen gelassen werden.

Populationsbiologie: bivoltin

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Scopula nigropunctata (70 Individuen)

Distanzen: 1-2

LEEVRICHICRE die Raupen uslszpllanzen charakterisieren S. nigropunciala als eine

an die iesen von äldern und Waldrändern gebundene Art. Damit steht der

bisher fehlende Nachweis im reinen Offenland (HM) im Einklang. In diesem Sinn

sind auch die RISRrEgen a ende an SiM und HW, sowie der Häufigkeits-

radient zwischen WaN und o sowie zwischen SiN und WaS auf nur 50 bzw.

0 m Strecke zu beurteilen.

Verbreitungsstrategie: K-Stratege, 6. Giuppe

Scopula ornata (15 Individuen)

Distanzen: 2, selten 3; das im Garten festgestellte Exemplar ist vermutlich vom

Verbreitungszentrum am Flughafen herbeigeflogen (l km). Diese Distanz stellt je-

doch offensichtlich schon die Obergrenze der Dispersionsaktivität dar und wird

nicht regelmäßig geflogen; in der Fangnacht ging auch ein Gewitter nieder, dessen

böige Winde flugunterstützend gewirkt haben könnten.

Larvalökologie: im Untersuchungsgebiet an die Raupen dieser Art wohl vor al-

lem an Schafgarbe und Thymian (Thymus). Schon in einer Einbuchtung von ca. 30

m in den Wald hinein (HO) war ein deutlicher Häufigkeitsabfall festzustellen.

Populationsbiologie: bivoltin

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Scopula rubiginata (?0 Individuen)

Distanzen: ?

Larvalökologie: in einer ähnlichen eise an trockene Wiesen gebunden wie die

vorige Art. Distanzen von I km (Zuflug in das Siedlungsgebiet) liegen offen-

sichtlich außerhalb der normalen Dispersionsaktivität und des trivial movement.

Populationsbiologie: bivoltin

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Scopula immutata

Distanzen: vermutlich 2;

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Scopula lactata (5 Individuen)

Distanzen: vermutlich 2-3

Verbreitungsstrategie: vermutlich intermediärer Typ, 5. Gruppe

LARENTIINAE

Scotopteryx chenopodiata

siehe verringerte Fallendistanzen (8.3.)

754 Individuen 44,6 % 9- Rate

380 markiert 27 Wiederfänge

Tab. 96: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Scotopteryx chenopodiata.

SIEDLUNG WALD HALBTROK- "DA WALD HALBTROK- "DA!

1987 ' Garten ' KERRAÄSEN Da y 1988 Garten Wasserwerk KENRASEN HOSE

SiS SIMISIN WaSıWaM Wa HO HM HW SN WaS WaN WNw WNo HO HM HW Au We

> par er 12,143 et; 37 (8) 214 2 par. 3 = 2oreg

zus: Miss. 8 ul en ze DIL = - smart zus. 95 4 T Frag

dd 2 6 - 2 =

29 = 5 r z Rn

_ 3.702 FBaSg» oe a

1 1 = Ze

168 |

Wiederfang-Quote: ist als hoch einzustufen (siehe Wasserwerk). Auch die Wie-

derfänge am Flughafen deuten_auf lange Verweildauern hin: Der Wiederfang er-

folgte nach durchschnittlich 4,7 Tagen we 322 b

Distanzen: 1-2; die Strecke vom Ruderal (häufiges Vorkommen von chenonodiata)

zum HM-Fangplatz (ca. 150-300 m) kann in den meisten Jahren zurückgeiegt

werden. Im Normalfall werden die 200 m in das Innere des Moorbirkenwäld-

chens nicht bewältigt; an dessen Rand dagegen war S. chenopodiata regelmäßig

anzutreffen. Die Siedlungs-Stücke sind vermutlich zugeflogen.

ulationsbiologie: die 99 sind ortsbeständiger und haben längere Verweilzeiten

Wasserwerk, Flughafen). Im Ort flogen 1988 dementsprechend nur d’d' zu.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Ansitis praeformata (2 Individuen)

Distanzen: vermutlich 2-3 h Bi

Larvalökologie: auf Hariheu (Hypericum) spezialisiert; im Garten wächst diese Fut-

terpflanze erst seit 1988, die Entwicklung, die sich über den Winter erstreckt,

konnte also nicht im Garten abgelaufen sein, das hier nachgewiesene d kam aus

mindestens 150 m Entfernung (Bereiwaldsan))

Verbreitungsstrategie: vermutlich intermediärer Typ, 5. Gruppe; es handelt sich um

die ersten Nachweise für den Münchner Norden. Für die offensichtlich stattge-

fundene Kolonisation durch diese mehr montane Art waren Flugleistungen von

durchschnittlich mindestens 1 km/Generation die Voraussetzung.

Anaitis efformata

197 Individuen 15,4 % 9- Rate

26 markiert 3 Wiederfänge

Wiederfang-Quote: in der 1. Generation hoch, in der 2. Generation niedrig, Stich-

probe noch zu gering; diese Feststellungen besitzen einen noch recht hypotheti-

schen Charakter, da sie auf einem Sinzigen mehrfach gefangenen Individuum

basieren: An WaS wurde 1988 in der 1. Generation bei täglichem Betrieb der

Lichtfalle ein & viermal bei Intervallen von 3, 1 und 4 Tagen gefangen, die

Verweildauer betrug also 8 Tage.

In der 2. Generation wurden 15 Individuen markiert und keines rückgefangen.

Distanzen: 3,

Larvalökologie: monophag an Tüpfel-Hartheu (Hypericum perforatum); an vielen der

Fundorte kommt die Raupenfutterpflanze nicht ın der näheren Umgebung vor (z.B.

Garten und HM). Die Häufigkeit an diesen Stellen ist daher bemerkenswert und

deutet auf, relativ hohe Dispersionsaktivitäten hin.

Erpulat onsbiologie: bivoltin, mit starken Fluktuationen von Generation zu Genera-

ion (die 1. Generation ist meist schwächer), aber auch von Jahr zu Jahr.

Verbreitungsstrategie: r-Stratege, 3. Gruppe; das Häufigkeitsmuster innerhalb des

Fallennetzes entspricht interessanterweise dem der typischen Wanderfalter.

Acasis viretata (2 Individuen)

Distanzen: 1-2

Larvalökologie: bisher konnten, nach den in KOCH (1984) genannten Raupenfutter-

pflanzen beurteilt, keine biotopfremden Stücke nachgewiesen werden.

Populationsbiologie: im Gebiet univoltin

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Nothopteryx polycommata (6 Individuen)

Distanzen: 1-2

Larvalökologie: bisher konnten, nach den in KOCH (1984) genannten Raupenfutter-

pflanzen beurteilt, keine biotopfremden Stücke nachgewiesen werden.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Nothopteryx carpinata (9 Individuen)

Distanzen: 1-2 . i j

Larvalökologie: die Raupe ist an Espe, Birke, Sal-Weide und Hainbuche zu finden.

Bisher keine biotopfremden Tiere

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Lobophora halterata (2 Individuen)

Distanzen: vermutlich 2

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

169

Pterapherapteryx sexalata (50 Individuen)

Distanzen: 2-3

Larvalökologie: die Raupe lebt oligophag an Weide und DADBe.: die 150-300 m-

Distanz vom Ruderal zum Standort HM wird in beiden Geschlechtern konstant

und von zahlreichen Stücken bewältigt; in den Garten (die nächsten einzelste-

henden Weiden befinden sich ca. 200 m entfernt) konnten bisher nur 2 zuge-

flogene Stücke nachgewiesen werden. Der Häufigkeitsgradient im Wasserwerk

spricht für Barrieren gegen eine freie Mobilität dieser Art.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Operophthera brumata

228 Individuen 0 %* o-Rate

169 markiert 30 Wiederfänge

* die 99 sind flügellos und werden daher mit Lichtfallenfängen nicht erfaßt.

Tab. 97: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Operophthera brumata 1988; 1987 war ein schlechtes Flugjahr, es wurden nur 3

Individuen markiert.

WALD HALBTROK- —

198 Garten Wasserwerk M ;”

SN WaS WaN WNw WNo HO HM HW Au We

Ipar. 60 64 Bund ou Br A 322.8‘

I zus. - 18 - - - - - - - - 18

Mark. 53 79 Buip Zus = 16. Rs N

Wi. 7100023 Sr er Sl rae =: 30

Wiederfang-Quote: hoch; in der Flugzeit 1988 erfolgte im Garten an beiden Stand-

orten (SiN und WaS) bei zwei Ausnahmen der Lichtfang parallel und täglich.

Das Ergebnis verdeutlicht die Abhängigkeit vom Standort: An SiN konnten pro-

zentual gesehen weniger J’d' rückgefangen werden, als an WaS.

Die mittlere Verweildauer liegt insgesamt mit 3,0 Tagen auf einem für diese

Methodik hohen Niveau. Interessant sind die 3 längsten festgestellten Intervalle

von 8, 11 und 16 Tagen. Die beiden letztgenannten J’d' überdauerten damit eine

einwöchige Kälteperiode mit Temperaturmaxima von -5 Grad Celsius. Die Nacht-

temperaturen betrugen bis zu 10 Grad Minus und es fielen ca. 10 cm Schnee.

Distanzen: Id 2, 2%]

Larvalökologie: die Raupen fressen an aaab ehölzen. Das an HM nachgewiesene J

stammt vermutlich aus dem Ruderal (150-300 m entfernt).

Verhältnismäßig große Nacht-zu-Nacht-Fluktuationen an SiN, dem mehr offenland-

ähnlichen der beiden Garten-Standorte weisen wohl auf einen erhöhten Anteil

Tiere hin (siehe auch "Wiederfang-Quote”). Im gleichen Zeitraum

waren die Fangergebnisse an WaS (30 m e Lfernt) recht konstant.

Populationsbiologie: siehe REICHHOL (1984)

Verbreitungsstrategie: K-Stratege, 5. Gruppe; es handelt sich um keine K-Strategie

im "klassischen’ Dina. da die 99 vergleichsweise viele Eier produzieren Taiche

REICHHOLF, 1.c.); die Strategie ähnelt in manchen Aspekten der von Alsophila

aescularia (siehe dort) Es zeigten sich allerdings deutlich längere Verweildauern

einem Faden abseilen, spielt bei dieser Art sicherlich eine Rolle t ICHHOLF, |.c.).

Oporinia dilutata

120 Individuen 18,0 % o- Rate

96 markiert 9 Wiederfänge

Wiederfang-Quote: durchschnittlich; 8 Wiederfänge ey im Garten (67 Mar-

kierungen) bei täglichem Feuß und erklären sich zum Großteil durch ein Festge-

halten-Werden im Bann der Lichtwir . Ein d wurde zum zweiten Mal wieder-

efangen (nach 2 Ein-Tages-Intervallen) und ein anderes J ging viermal in die

ichtfalle, wobei es vom 2. auf den 4. Tag den Ort wechselte (Was> SiN = 30 m).

Sonst lagen zwischen Fang und Wiederfang nur Intervalle von einem Tag.

.WNo). Hier waren 1988 10 Individuen markiert worden.

Distanzen: 2, EaRne

Larvalökologie: wie die vorige Art an Laubbäumen, bisher konnte noch kein biotop-

fremdes Stück nachgewiesen werden (z.B. HM). Der Häufigkeitsgradient zwi-

schen WaN und o auf nur 50 m Entfernung könnte für Barrieren gegen die

freie, Beweglichkeit von O. dilutata sprechen.

Verbreitungssirategie: K-Stratege, 6. Gruppe

170

Oporinis autumnata

46 Individuen 10,5 % 9 - Rate

32 markiert 3 Wiederfänge

Wiederfang-Quote: relativ hoch, Stichprobe noch zu klein; es handelt sich, um zwei

dc 1988 am Standort WaS (täglicher Betrieb der Falle). von denen eines nach

einem Intervall von 1 Tag, das andere am 4. und 5. Tag nach der Erstmarkierung

‚wiedergefangen wurden.

tanzen: |- I i \

Larvalökologie: die Raupe wird an Weide, Birke, Ahorn und Lärche gefunden, bio-

topfremde Tiere konnten bisher noch nicht beobachtet werden.

Verbreitungsstrategie: K-Siratege, 6. Gruppe; O. aufumnata scheint im Vergleich

zur vorigen Art lokaler vorzukommen und ortstreuer zu sein, was mit dem weni-

ger breiten Wirtspflanzenspektrum zusammenhängen könnte.

Triphosa dubitate (19 Individuen)

Distanzen: 2. 3 ee

Larvalökologie: der Anflug bzw. die Kolonisation der kleinen gartenähnlichen Fläche

am Sendergebäude muß, nach den in KOCH 1984) Benannten Futterpflanzen

beurteilt, über ca. 350-400 m freies Gelände hinweg erfolgt sein.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Calocalpe cervinalis

110 Individuen 60,4 % 9 - Rate

89 markiert 2 Wiederfänge

Wiederfang-Quote: durchschnittlich; ein d. wechselte 1987 in einem Zeitintervall von

1 Tag den Ort (WaS>SiN = 30 m). Ein 9 konnte im selben Jahr an WaM noch

10 Tage nach der Erstmarkierung nachgewiesen werden! In sich geschlossene Ge-

ländestrukturen scheinen die Verweildauern zu steigern.

tanzen: 2, die oo vielleicht nur 1-2

Larvalökologie: mönophag an Berberitze (Berberis vulgaris); biotopfremde Stücke

konnten bisher noch nicht nachgewiesen werden, lediglich vom Standort HW ist

das nächste Vorkommen der Futterpflanze ca. 150 m entfernt. Dementsprechend

wurde auch nur 1 Individuum nachgewiesen.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Philereme vetulata (154 Individuen)

Distanzen: 3

Larvalökologie: oligophag an Kreuzdorn (Rhamnus cathartica) und Faulhaum (Fran-

gula ae), die regelmäßig im Offenland (HM) auftauchenden Stücke (auch ein 9

sind mindestens 800-1000 m geflogen und deuten auf hohe Dispersionsaktivitäten

hin. Auch die zahlreichen HW-Exemplare stammen aus mindestens 150 m Entfer-

nung.

Populationsbiologie: von Generation zu Generation treten z.T. starke Fluktuationen

auf.

Verbreitungsstrategie: intermediärer Typ, 3. Gruppe

Philereme transversata (19 Individuen)

Distanzen: 1-2; der Häufigkeitsgradient WaS/SiN auf nur 30 m Entfernung deutet

auf geringe Dispersionsaktivitäten i h

| Larvalökologie: Larvalansprüche wie bei P. vetulata, zusätzlich wird noch Schlehe

| akzeptiert. Bisher konnten keine biotopfremden Tiere beobachtet werden.

| Verbreitungsstrategie: K-Stratege, 6. Gruppe

Lygris prunata (36 Individuen)

Distanzen: 1-2; wie P. transversata k

Larvalökologie: die Raupe ernährt sich von Stachelbeere, Johannisbeere, Schlehe,

Weißdorn und Eiche; bisher keine biotopfremden Tiere

Verbreitungsstrategie: K-Stratege, 6. Gruppe

)

Lygris testata (3 Individuen)

Distanzen: vermutlich 2 \ i

| Larvalökologie: die Raupe bevorzugt Heidekraut (Calluna) und Heidelbeere (Vacci-

nium), da diese im ateruchungsgebiet jedoch fast nicht verfügbar sind, weicht

diese Art auf Espe, Weide und Birke aus. Solche Standorte dürften aber wohl als

| suboptimal zu charakterisieren sein. Bisher konnten keine biotopfremden Tiere

festgestellt werden.

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

171

ZT — —m— =

Lygris populata

Distanzen: vermutlich 2

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Lygris mellinata (6 Individuen)

Distanzen: 2-4

Larvalökologie: die Raupe ist auf Johannisbeere und Stachelbeere spezialisiert,

bisher keine biotopfremden Stücke

Verbreitungsstrategie: vermutlich intermediärer Typ, 5. Gruppe; diese Art unter-

nahm in den letzten ca. 50 Jahren eine Arealausweitung quer über ganz Süd-

bayern, wobei zumindest schubweise auch größere Strecken von mehreren Kilome-

tern bewältigt werden mußten.

Lygris pyraliata (116 Individuen)

Distanzen: ?2

Larvalökologie: an Labkraut (Galium) und Bach-Nelkenwurz (Geum rivale); trotz

der weiten Verbreitung im Untersuchungsgebiet konnten an HM keine Stücke

nach eWieacn werden. Die Distanz von 1 km liegt offensichtlich nicht im Bereich

der Dispersionsaktivität und des trivial movement dieser Art.

verbreitungsstrategie: intermediärer Typ, 4. Gruppe

Cidaria fulvata (80 Individuen)

Distanzen: 2(-3)

Larvalökologie: die Raupe ist auf Rosen spezialisiert; im Wasserwerk ist in einer

Entfernung von 100 m von einem Rosenbestand (WaN>WN J ein Häufigkeitsabfall

auf ca. die Hälfte zu beobachten. Das 1986 nach HM (ca. 1 km] zugeflogene

Stück stellt wohl die Obergrenze der normalen Dispersionsaktivität oder vielleicht

schon eine Ausnahme dar.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Plemyria rubiginata (75 Individuen)

Distanzen: 2-3

Larvalökologie: die Raupe lebt an Erle (A/nus); das HM-c' (1988) legte mindestens

800 m vom nächstgelegenen Standort der Raupenfutterpflanze zurück. Die zahl-

reichen im Garten festgestellten Stücke müßten zumindest aus 200-300 m Ent-

fernung gekommen sein, wenn nicht doch zuweilen auch auf andere Futterpflan-

zen ausgewichen wird. ”

Verbreitungsstrategie: intermediärer Typ, 3. Gruppe

Thera variata (64 Individuen)

Distanzen: 2, s

Larvalökologie: T. variata ist an Nadelbäume gebunden. Der Nachweis für ein

Sich-Entfernen von der Futterpflanze um 50 m wurde für eine Reihe von Stand-

orten erbracht, die Distanz von 1 km (Zuflug nach HM) est jedoch außerhalb

der normalen Dispersionsaktivität, Am Franzosenhölzl konnte 1989 ein o ca. 500 m

vom nächsten Nadelbaum (Fichte) entfernt gefangen werden (2. Generation).

Populationsbiologie:, bivoltin, dazu zwei Exemplare einer 3. Generation; proterandrisch

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Thera obeliscata (113 Individuen)

Distanzen: 2 (-3)

Larvalökologie: die Raupen fressen vor allem an Kiefern, vermutlich aber auch an

Fichte und Wacholder; das an HM 11986) in einer windstillen Nacht festgestellte

Exemplar stammt also aus mindestens 800 m Entfernung (Friedhof Hochmutting).

In 5 100 m Entfernung zum Standort HW stehen einige kleine Kiefern (ca. 2 m

hoch), die als Herkunft für die beiden dort nachgewiesenen Falter in Frage kom-

men.

Populetionsbiologie: eine 2. Generation wird zwar nachgewiesen, sie ist jedoch recht

unvollständig.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe; im Zuge einer günstigen Be-

standsentwicklung kam es 1986 offenbar bei den meisten der an Kiefern gebunde-

nen Arten zu Verbreitungsschüben.

Thera juniperata

53 Individuen 11,8 % o- Rate

34 markiert kein Wiederfang

Wiederfang-Quote: niedrig, Stichprobe noch zu klein

Distanzen: 2, bei Besiedelängsmar sungen auch 3

172

Larvalökologie: nach KOCH (1984) monophag an Wacholder (Juniperus communis),

siehe jedoch die Bemerkungen unter "Verbreitungsstrategie”. Biotopfremde Stük-

ke konnten bisher noch nicht beobachtet werden.

Verbreitungsstrategie: intermediärer Tp: 5. Gruppe; eine Arealausweitung, ge-

tützt auf die in Gärten häufig angepflanzten Zierwacholder, erwähnt MEI CKE

(1984). Solche Arealausweitungen fanden im Laufe dieses Jahrhunderts auch in

üdbayern statt. Da Wacholder in der weiteren Umgebung des Untersuchungs-

ebietes nicht in der Natu: vorkommt, bleibt nur eine Trittsteinbesiedlung über

acholder in Gärten und Friedhöfen. Vermutlich kann die Art jedoch unter sub-

optimalen Bedingungen zumindest kurzzeitig auf andere Nadelhölzer zurückgrei-

fen. Dies wird vor allem bei solchen Besiedelungsprozessen eine Rolle spielen.

Thera firmata

249 Individuen 61,2 % o9- Rate

215 markiert 2 Wiederfänge

Tab. 98: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Thera firmata.

SIEDLUNG WALD HALBTROK- "DACH, WALD HALBTROK- "DA

1987 | Garten | M % 1988 Garten Wasserwerk KENRASEN MOO

sis SIM|SIN Was |WaM WaN HO HM HW Mb SiN WaS WaN WNw WNo HO HM HW Au We

apa 66 2 sms us = 139 Zpa. 1 13 Sm 2! N o.9 le

ZA =, 23 - 5 - > - - A zus: - 7 - - - RS HE - -

dd Fun s6 Ale 8 AR RD - 58 dd 4 7 Sa Zn Po] -

99 a 62 23 SEE Sm _ 108 99 SS SE IB = Aa he

Mark. E75 AS 1 32 Du - 162 Mark. 7) 2.20) 3 or 142.02 = 08

Wf. = 1 Ne Suamie _ 1 wi. = 1 Se 2a Sa... o ne

Wiederfang-Quote: niedrig; 1987 wurde ein 9 nach 3 Tagen, 1988 ein J nach 1 Tag

wiedergefangen.

Distanzen: 1-

Larvalökologie: monophag an Kiefer; im Siedlungsbereich spiegelt sich die Ökologi-

sche Trennung im Fangergebnis der verschiedenen Standorte gut wieder. Auch ım

Wasserwerk weist der Häufigkeitsgradient darauf hin, daß schon Entfernungen von

ca. 100 m ein Hindernis für eine Verbreitung darstellen können.

Die immer wieder (in windigen Nächten) im Öffenland (HM) auftauchenden Exem-

plare - 1987 auch ein fertiles 9 - sind mindestens 800-1000 m geflogen und zeu-

en von weiten Vorsioßen über biotopfremdes Gebirt. Im gleichen Sinn ist das

988 am Würmkanal (Au festgestellte @ zu verstehen; in einer Entfernung von

ca. 200 m existiert eine ca. 10 Jahre alte Kiefernschonung.

Populationsbiologie: proterandrisch

Verbreitungsstrategie: K-Stratege, 5. Gruppe

Chloroclysta siterata (14 Individuen)

Distanzen: 2 Er

Larvalökologie: an Laubhölzern, bisher keine biotopfremden Tiere; für die Überwin-

ter scheinen Häuser, Schuppen, Mauern und dergleichen von Vorteil zu sein.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Dystroma truncata (181 Individuen)

Distanzen: 2

Larvalökologie: recht polyphag, D. truncata ist jedoch eine typische Art der Wäl-

er und deren Ränder. Das HM-Stück stammt mindestens aus dem Ruderal

150-300 m). Im Wasserwerk und im Garten besteht ein recht deutlicher Häu-

igkeitsgradient, der auf normale Flugaktivitäten von unter 200 m außerhalb des

Habitats hinweist.

Populationsbiologie: bivoltin

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Dystroma citrata (38 Individuen)

Distanzen: 2 _ h EN:

Populationsbiologie: im Gegensatz zur vorigen Art univoltin

Verbrei strategie: K-Stratege, 6. Gruppe; sowohl im Untersuchungsgebiet als

auch in der unteren Hochebene Südbayerns lokaler verbreitet als D. truncata.

Xanthorhoe fluctuate (88 Individuen)

Distanzen: ?; die Häufigkeitsgradienten im Wasserwerk und im Garten sowie der

bisher fehlende Nachweis an HM könnten als erste Hinweise dafür verstanden

werden, daß im "trivial movement” die Maximaldistanzen einige 100 m betragen.

Populationsbiologie: bivoltin

Verbreitungsstrategie: intermediärer Iyp, 5. Gruppe

Erg sr

173

TE — —

Xanthorhoe montanata (27 Individuen + tagsüber > 500 Exemplare)

Distanzen: 1-2

Larvalökologie: für das lokal so überaus zahlreiche Auftreten im Birket stellt ver-

mutlich die Häufigkeit der Himbeere die Ursache dar.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Xanthorhoe spadicearia (381 Individuen)

Distanzen: 2(-3?); das polarisierte Häufigkeitsverhältns WaN/WNo auf nur 50 m

Distanz deutet vielleicht auf Barrieren gegen die freie Mobilität dieser Art hin. Im

Vergleich zur recht ähnlichen X. ferrugata dominiert X. spadicearia zahlenmäßig

im asserwerk, im Garten sind die Dominanzverhältnisse umgekehrt. Zwischen

diesen beiden Standorten (1,2-1,3 km Distanz) findet vermutlich kein nennens-

werter Austausch statt.

Populationsbiologie:, bivoltin

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Xanthorhoe ferrugata

siehe Versetzexperiment (8.4.)

1023 Individuen 45,2 % o- Rate

406 markiert 25 Wiederfänge

Tab. 99: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Xanthorhoe ferrugata.

SIEDLUNG WALD HALBTROK- nl WALD HALBTROK- Da

1987 ı Garten | KENRASEN MOOS % 1988 Garten Wasserwerk NRASEN M 2

SiS SIM|ISN WaS|WaM WaN HO HM HW Mb SN WaeS WaN WNw WNo HO HM HW Au We

ZI par. 1 9: 15 63 87 33 46 57241 15 236 Xpar. «47 230 42 1929 29 Zem22 16 4 44

Z zus. - 434 En az 9 u 1 TO zus: ) = 1271 - - - ey, Se u = m

dd 1 1 11 22 6 21 17 3 10 4 96 dd 10 9 9 6 6 7 72 1 2 - 134

99 - 3 #1 16 1 3 8 2 7 - 5199 8 123 3 2 2 1 - - - - 13°

Mark. 1 4 22 38 6 24 25 BE 17 4 146 Mark. 15 208 10 7 8 8 1 1 2 - 26(

Wiederfang-Quote: in beiden Generationen niedrig, im Wasserwerk duSeeEn höher;

Fast alle der Wiederfänge erklären sich durch ein Festgehalten-Werden im Bann

des Lichtes: Im Garten erfolgten 21 von 22 Rückfängen nach einem Intervall von

1 Tag, nur 1 S war nach 2 Tagen wiedergefangen worden. Zwei Jd' wechselten

dabei von WaS nach SiN (30 m). Es ereignete sich nur I Zweitwiederfang.

Im Wasserwerk konnten unter "regulären" Bedingungen (dazwischenliegende fang-

freie Nacht) 1987 ein J 3 Tage nach der Markierung BE werden. 1988

wurden zwei den Standort wechselnde JG beobachtet: Es, wurden hierbei die

Strecken, WNo> WaN 50 m) in 2 Tagen und WNw>WaN (100 m) in 10 Tagen

zurückgelegt.

‚Die o-Rate im Wiederfangergebnis entspricht in etwa der der Erstfänge.

Distanzen: in beiden Generationen 2-3; im Offenland wird die Art bevorzugt in

Nächten mit Wind nachgewiesen. j t

Populationsbiologie: bivoltin, in beiden Generationen proterandrisch; der o9-Rate von

9,0 % in der 1. Generation stehen 52,0 % in der 2. Generation gegenüber. Am

Sander WaS war diese Differenz bei einem Verhältnis von 32,2 % / 62,2 % noch

größer.

Verbreitungsstrategie: r-Stratege, 2. Gruppe

Xanthorhoe biriviata (10 Individuen + tagsüber 2 Exemplare)

Distanzen: 2

Larvalökologie: monophag am Echten Springkraut (Impatiens poll fangezei in der

Falle konnten bisher keine biotopfremden Tiere nachgewiesen werden, die beiden

tagsüber im Moos bei prallem Sonnenschein aktiven Exemplare Aeeen jedoch

entlang einer Hecke an einer Stelle, die ca. 100 m von der nächsten Raupenfut-

terpflanze entfernt ist. Langgestreckte Landschaftselemente wie Hecken oder

Bachbegleitfloren können also bei der Verbreitung von relativ spezialisierten

silvicolen Arten eine gewichtige Rolle spielen.

Populationsbiologie: bivoltin

Verbreitungsstrategie: K-Stratege, 5. Gruppe

Xanthorhoe designata (4 Individuen)

Distanzen: vermutlich 2

Populationsbiologie: bivoltin

Verbreitungsstrategie: vermutlich K-Stratege, 5. Gruppe

174

Ochyria quadrifasciata (363 Individuen)

Distanzen: 2; das stark polarisierte Häufigkeitsverhältnis WaS/SiN sowie die Tat-

sache, daß im Offenland (HM ‚nur einmal ein cd’ in einer windigen Nacht gefan-

gen wurde, charakterisieren diese im Vergleich zu den Xanthorhoe-Arten ei-

gentlich flugkräftig erscheinende Art als einen u biotoptreuen Waldbewoh-

ner. Das HM-Stück flog vermutlich aus dem Ruderal (150-300 m) herbei.

Populationsbiologie: im Untersuchungsgebiet abgesehen von 1 Exemplar, das Anfang

eptember gefangen wurde, nur univoltin

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Nycterosea obstipata (1 Individuum)

Distanzen: 4

Verbreitungsstrategie: r-Stratege, 1. Gruppe

Calostigia olivata (14 Individuen)

Distanzen: 2

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Calostigia pectinataria (82 Individuen)

Distanzen: ?2

Larvalökologie: die Raupen fressen an Labkraut, Brennessel und Taubnessel; die

beiden an HM festgestellten Stücke stammen zumindest aus dem Ruderal (150-300 m).

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Lampropteryx ocellata (65 Individuen)

Distanzen: 1-2; die Häufigkeitsgradienten im Wasserwerk (auf 50 m Entfernung) und

im Garten (30 m) könnten Hinweise darauf sein, daß diese Art schon mit der

Bewältigung solch kurzer Distanzen Probleme hat.

Larvalökologie: die Raupen ernähren sich von Labkrautarten. Der fehlende Nach-

weis in der Mitte des Flughafens (HM) deutet auf Dispersionsaktivitäten unter 1 km

hin.

Populationsbiologie: bivoltin

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Lampropteryx suffumata (14 Individuen)

Distanzen: 2.

Larvalökologie: auch diese Art ist auf verschiedene Labkrautarten spezialisiert, die

Habitatbindung scheint stärker zu sein als bei L. ocellata.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Coenotephria berberata (210 Individuen)

Distanzen: 2-3

Larvalökologie: monophag an Berberitze (Berberis vulgaris); dass HM-c won)

mußte vom Ort der Larvalentwicklung aus mindestens 800-1000 m weit geflogen

sein.

Populationsbiologie:, bivoltin

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Euphyia cuculata (70 Individuen)

Distanzen: 2

Larvalökologie: die Raupe ist an Labkrautarten zu finden, die 3 HM-Stücke stam-

men vermutlich aus dem Ruderal 1150-300 m).

Eepnlationsbinlogie: im Untersuchungsgebiet wurde bisher nur 1 Generation festge-

stellt.

Verbreitungsstrategie: K-Stratege, 5. Gruppe

Euphyia molluginata (4 Individuen)

Distanzen: vermutlich 1-2 h 2

Larvalökologie: wie E. cuculata, bisher konnten keine biotopfremden Stücke be-

obachtet werden.

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

175

Euphyia bilinesta (126 Individuen + tagsüber viele weitere Exemplare)

Distanzen: 2-3; das stark polarisierte Häufigkeitsverhältnis im Garten (SiN/WaS)

spricht für Barrieren gegen die freie Mobilität dieser Art. _

Populationsbiologie: E. bilineata fliegt im Untersuchungsgebiet in einer partiellen 2.

eneration

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Diactinia capitata (3 Individuen)

Distanzen: vermutlich 1-2

Larvalökologie: monophag an Echtem Springkraut (/mpatiens noli-tangere); bisher

konnten noch keine biotopfremden Tiere beobachtet werden.

Populationsbiologie: im Untersuchungsgebiet steht der Nachweis einer 2. Generation

noch aus.

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Diactinia silaceata (43 Individuen)

Distanzen: 2, >

Larvalökologie: oligophag an Weidenröschen (Epitobium), Echtem Springkraut und

Großem Hexenkraut [ ircaea han) lebend; bisher wurden alle Exemplare in

der Nähe der Raupenfutterpflanzen gefangen.

Populationsbiologie:, bivoltin

Verbreitungsstrategie: K-Stratege, 6. Gruppe; die Dispersionsaktivitäten dieser Art

scheinen im Vergleich zu D. capilata einhergehend mit dem erweiterten Wirts-

pflanzenspektrum auf einem etwas höheren Niveau zu liegen.

Electrophaes corylata (14 Individuen)

Distanzen: 2,

Larvalökologie: oligophag an einigen Laubhölzern, bisher keine biotopfremden Tiere

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Mesoleuca albicillata (30 Individuen + tagsüber viele weitere Exemplare)

Distanzen: 1-2; auch die in der Dämmerung gemachten Beobachtungen deuten auf

geringe Dispersionsaktivitäten hin. Fmmeal 2

Larvalökologie: die Raupen sind auf Him- und Brombeere spezialisiert; bisher wur-

den alle Exemplare in der unmittelbaren Nähe der Futterpflanzen beobachtet

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Melanthia procellata (5 Individuen)

Distanzen: 2 (-4?)

a monophag an Gemeiner Waldrebe (Clematis vitalba); bisher konn-

ten keine biotopfremden Tiere nachgewiesen werden.

Populationsbiologie: im Untersuchungsgebiet bivoltin

Verbreitungsstrategie: K-Stratege, 6. Gruppe; KOCH (1984) berichtet von der Neu-

kolonisatıon Sachsens innerhalb von einigen Jahren; hierbei müssen größere Strek-

ken zurückgelegt worden sein; solch große Distanzen liegen jedoch vermutlich

nicht im Bereich des "trivial movement’ dieser Art.

Epirrhoe tristata

364* Individuen 20 %** o- Rate

10 markiert kein Wiederfang

* tagsüber viele weitere Exemplare ** Stichprobe zu klein

Wiederfang-Quote: Stichprobe zu klein

Distanzen: 1. Generation 1-2; 2. Generation 2-3; im Wasserwerk fällt der starke

ar llagradien! schon auf einer Distanz von nur 50 m auf (WaN/WNo).

Larvalökologie: die Raupen ernähren sich von Labkrautarten; die beiden HM-Stük-

ke stammen vermutlich aus dem Ruderal (150-300 m). Im Garten konnte E. tris-

tata bisher nur in der 2. Generation nachgewiesen werden. Vielleicht sind dafür

zufliegende Individuen mit erhöhter Dispersionsaktivität (im Vergleich zur 1. Ge-

neration) verantwortlich.

Populationsbiologie:, bivoltin

Verbreitungsstrategie: intermediärer Typ, 3. Gruppe

176

Epirrhoe alternata (766 Individuen + tags viele weitere Exemplare)

Distanzen: 3; ein 9 wurde tagsüber bei einem 500 m weiten Flug beobachtet, diese

Distanzen scheinen innerhalb der Grenzen der normalen Dispersionsaktivität zu

jegen.

Larvalökologie: wie E. tristata; im reinen Offenland (HM) ist diese Art häufiger als

die vorige Art anzutreffen. Im Garten erscheint sie schon in der 1. Generation.

Bevorzugter Lebensraum ist der Brennessel-Giersch-Sau Urtico dioicae-Aego-

podietum),, wenn dieser gut mit Klebkraut (Galium aparine) durchmischt ist.

Populationsbiologie: bivoltin

Verbreitungsstrategie: r-Stratege, 2. Gruppe

Perizoma alchemillata (233 Individuen)

Distanzen: 2 (-3?)

Larvalökologie: nach den in KOCH (1984) genannten Raupenfutterpflanzen beurteilt,

handelt es sich bei den beiden im Offenland HM beobachteten Stücken um Gä-

ste, die zumindest aus dem Ruderal stammen (150-300 m

Verbreitungsstrategie: intermediärer Typ, 3. Gruppe

Perizoma bifaciata (1 Individuum)

Distanzen: vermutlich 2

Larvalökologie; oligophag an Gelbem Zahntrost (Orthanta lutea) und Gemeinem

Augentrost (Eu hrasia officinalis). Letztgenannte Pflanze ist im Flughafengebiet

verbreitet. Ob das Stück von dorther in den Siedlungsbereich eingeflogen ist, läßt

sich, im Moment nicht klären, bodenständig ist die Art jedoch im Garten sicherlich

nicht.

Populationsbiologie: die Bunpe überliegt nach KOCH (1984) meist mehrmals, was die

potentielle Wachstumsrate der Populationen dieser Art drastisch reduziert.

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Perizoma blandiata

Distanzen: vermutlich 2

Larvalökologie: monophag an Gemeinem Augentrost (Euphrasia officinalis); siehe

Bemerkungen zu P. bifaciata

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Perizoma flavofasciata (3 Individuen)

Distanzen: 3

Laryalökologie: auf Lichtnelken (Silene) spezialisiert; auf ein Auftreten der Futter-

pflanzen kann bisweilen sehr rasch mit einem Kolonisationsversuch geantwortet

werden, wie dies 1986 schon im ersten Jahr der Verfügbarkeit der Futterquelle

eschah. Gemäß dem unsteten Auftreten der Wirtspflanzen dürften auch die

eobachteten hohen Austauschraten im Untersuchungsgebiet eine Anpassung an

dieses Phänomen ‚darstellen.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Hydriomena furcata (71 Individuen)

Distanzen: 2

Larvalökologie: H. furcata ist an das Vorhandensein von Weiden oder Heidelbee-

ren (Vaccinium myrtillus) gebunden. Die im Garten festgestellten Tiere müssen

also aus mindestens 200 m Entfernung zugeflogen sein, die HW-Stücke aus 150

m, jeweils ohne Sichtkontakt zur Lichtquelle.

Verbreitungsstrategie: K-Stratege, 5. Gruppe

Hydriomena coerulata

9] Individuen 15,4 % o- Rate

26 markiert 1 Wiederfang

Wiederfang-Quote: hoch, Stichprobe noch zu klein. l Wasserwerk konnte 1988 ein

Ortswechsler (WaN>WNo= 50 m) nach 2 Tagen 5) festgestellt werden. a

Distanzen: 1-2; der Häufigkeitsgradient im Wasserwerk auf 50 bzw. 100 m Di-

stanz, sowie der starke a ehrall im Franzosenhölzi 1989 schon 40 m

außerhalb des Wäldchens (2 Individuen gegenüber 15 am Rand) sprechen für ge-

ringe Dispersionsaktivitäten. \ 4 vn f £

Larvalökologie: die Raune ernährt sich von einigen Laubbäumen und Heidelbeere; im

reinen Offenland (H wurde noch kein Nachweis erbracht.

Populationsbiologie: wenig proterandrisch

Verbreitungsstrategie: K-Stratege, 6. Gruppe

177

Anticlea badiata (6 Individuen)

Distanzen: 1-2

EEE EIPSR die Raupen sind auf Rosen spezialisiert; bisher keine biotopfremden

iere

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Pelurga comitata (6 Individuen)

Distanzen: vermutlich 2 37

Populationsbiologie: die pe überwintert nach KOCH (1984) gelegentlich zweimal,

was die potentielle Wachstumsrate der Populationen dieser Art verringert.

Ve eye vermutlich intermediärer Typ, 5. Gruppe; GYULAI & VARGA

(1978 berichten allerdings von gerichteten Bewegungen innerhalb des Areals die-

ser Art.

Hydrelia testaceata (69 Individuen)

Distanzen: 1-2

Larvalökologie: die Raupe ist auf Erle spezialisiert, vielleicht nimmt sie jedoch

auch Birke und Weide an. Das im Garten gefundene Stück könnte somit von ei-

ner zumindest kurzzeiligen Besiedlung der dortigen Birken zeugen.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Hydrelia flammeolaria (89 Individuen)

Distanzen: 2

Larvalökologie: oligophag an verschiedenen Laubbäumen; im Offenland (HM) bisher

noch nie gefunden, Dispersionsaktivitäten daher wohl deutlich kleiner als 1 km

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Euchoeca nebulata (131 Individuen)

Distanzen: 2

Larvalökologie: Dleuphaz an Erle und Birke; im Offenland IM) bisher noch nie

gefunden, Dispersionsaktivitäten daher wohl deutlich kleiner als | km

Populationsbiologie: bivoltin

Verbreitungsstrategie: K-Stiratege, 6. Gruppe

Asthena albulata (10 Individuen)

Distanzen: vermutlich 2

Larvalökologie: oligophag an einigen Laubbäumen, bisher keine biotopfremden_Tiere

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe; wie auch andere Entomo-

logen berichten, ist diese Art wie auch die folgende in Südbayern in den letz-

ten beiden Jahren Baer häufiger geworden. Die Ela eisiünger der Asthena-

Arten können jedoc achstumsrate der Popu-

lationen nicht so groß sein, daß diese Art in so kurzen Zeitabschnitten zu einer

Be er Südbayerns fähig wäre. Vermutlich ist diese ee einer

un gen Konstellation der Futterpflanzenentwicklung unter den gegebenen Klima-

edingungen zuzuschreiben. Vielleicht spielen weitere Faktoren herein, jedoch

beispielsweise eine Verminderung des Feinddruckes durch Parasiten, der zwar art-

bzw. gattungsspezifisch wäre, müßte dann auch synchron auf einer sehr großen

Fläche stattgefunden haben, was recht unglaubhaft ıst.

Ein derartiges jahrweise häufiges Auftreten synchron in größeren Gebietsteilen ist

auch für viele Eupithecia-Arten typisch.

nicht so stark und die potentielle

Asthena anseraria (3 Individuen)

Distanzen: vermutlich 1-2

Larvalökologie: monophag am Blutroten Hartriegel (Cornus sanguinea), welcher im

Wasserwerk in unmittelbarer Nähe der Fundorte vorkommt.

Verbreitungsstrategie: K-Stratege, 6. Gruppe; siehe Bemerkungen zu A. albulata

Eupithecia tenuiata (19 Individuen)

Distanzen: 2,

Larvalökologie: monophag an Sal-Weide (Salix caprea); die 4 im Garten gefunde-

nen Stücke waren mindestens 200 m, die 6 HW-Stücke mindestens 150 m von der

nächstgelegenen Futterpflanze her geflogen. Das an HM 1986 festgestellte Tier

stammt höchstwahrscheinlich aus dem Ruderal (150-300 m). Diese Daten charak-

terisieren E. tenuiata als eine vergleichsweise mobile Eupithecie.

Populationsbiologie: wenig proterandrisch, Se 50,0 %

Verbreitungsstrategie: intermediärer Typ, >. Gruppe

178

Eupithecia inturbata (16 Individuen)

Distanzen: ?; der starke Häufigkeitsgradient im Wasserwerk spricht für Barrieren

gegen die freie Mobilität dieser Art schon auf 50 m Distanz.

Larvalökologie: monophag am Feld-Ahorn (Acer campestre); die im Garten fest-

gestellten Exemplare müssen mindestens 1 m weit geflogen sein.

erbreitungsstrategie: K-Siratege, 6. Gruppe; E. inturbata gehört zu der Gruppe der

eigentlich als selten geltenden Arten, die in den letzten Jahren ihre Bestände

vergrößern konnten IRiche Bemerkungen zu Asthena albulata).

Eupithecia plumbeolata (5 Individuen)

Distanzen: vermutlich 2

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Eupithecia pini (1 Individuum)

Distanzen: vermutlich 1-2

Larvalökologie: an Nadelbäumen, v.a. Gemeine Fichte (Picea abies) und Kiefer (Pi-

nus); bisher keine biotopfremden Tiere

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Eupithecia bilunulata

Distanzen: vermutlich 1-2 j STR

Larvalökologie: die Kan sind sehr spezialisiert und leben in den Gallen von

Fichtenläusen an der Gemeinen Fichte; alle 1983 und 1989 nachgewiesenen Tiere

wurden in unmittelbarer Nähe der Wirtspflanzen festgestellt.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Eupithecia linariata (4 Individuen)

Distanzen: vermutlich 2

Vase monoplae, am Gemeinen Leinkraut (Linaria vulgaris); das im Garten

festgestellte, etwas geflogene Stück mußte mindestens 300 m weit geflogen sein.

Populationsbiologie: bivoltin

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Eupithecia exiguata (41 Individuen)

Distanzen: 1-2 n #

Larvalökologie: an verschiedenen Laubhölzern, v.a. an Sträuchern, bisher keine bio-

topfremden Tiere, festgestellt

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Eupithecia valerianata (5 Individuen)

Distanzen: vermutlich 2

Larvalökologie: die Raupe ernährt sich monophag vom Echten Baldrian (Valeriana

officinalis);, diese Pflanze kommt im Radius von ca. 200 m um die Fangplätze im

Garten nicht vor. Das hier beobachtete Stück muß also mindestens so weit geflo-

gen sein.

Populationsbiologie: die Puppe überwintert nach KOCH a manchmal zweimal,

was die potentielle Waschstumsrate der Populationen ("PGR") vermindert.

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Eupithecia venosata (6 Individuen)

Distanzen: 1-2 Bisher | abe

Larvalökologie: die Raupe ernährt sich von a (Silene vulgaris), im Gar-

ten jedoch wahrscheinlich von der Roten Lichtnelke (Silene dioica). Bisher konn-

ten keine biotopfremden Tiere festgestellt werden. KEN

Populationsbiologie: die e nn nach KOCH manchmal zwei bis dreimal;

| ies verringert die "PGR" (siehe oben) und kann auch zur Vortäuschung von

Turnover -Ereignissen führen. % h

Verbreitungsstrategie: K-Stratege, 6. Gruppe; das fakultative UÜberliegen der Pup-

Ä pen ist vielleicht eine Möglichkeit der Kompensation von ungünstigen Bestands-

entwicklungen in schlechten Jahren: Wenn einmal eine (apparente) Extinktion

| einer Population eintritt, hat die Art immer noch einige Puppen "auf Vorrat“.

| | 179

Eupithecia egenaria (3 Individuen)

Distanzen: 1

Larvalökologie: monophag an der Sommerlinde (Titia platyphyllos), das HO-J wur-

de 20 m, das im Garten beobachtete Stück 15 m von der Futterpflanze entfernt

gefunden. Das dritte Exemplar wurde direkt an der Futterpflanze gefangen.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Eupithecia extraversaria (14 Individuen)

Distanzen: vermutlich 2

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Eupithecia centaureata (22 Individuen)

Distanzen: 2-3 Aziı

Larvalökologie: vergleichsweise polyphag an niedrigen Pflanzen; im Garten tritt

diese Art jedoch vermutlich nur als Gast auf.

Populationsbiologie:, bivoltin

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Eupithecia selinata (7 Individuen)

Distanzen: vermutlich 2

Populatioasbiologie: im Untersuchungsgebiet univoltin

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Eupithecia trisignaria (6 Individuen)

Distanzen: vermutlich 2

Populationsbiologie: im Untersuchungsgebiet offensichtlich bivoltin, wie in FORSTER

WOHLFAHRT (1981) beschrieben

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Eupithecia intricata

180 Individuen 21.7 2° or Rale

23* markiert 2* Wiederfänge *FE5-M6 1989

Wiederfang- te: die Art wurde nur in einigen Nächten 1989 an den Standorten

WaS und SiN markiert. Es lagen fast stets fangfreie Nächte dazwischen, so daß

die. Wiederfang-Quote als relativ hoch zu werten ist (Stichprobe jedoch noch zu

klein). Je ein J wurde nach ein bzw. zwei Tagen wiedergefangen.

Distanzen: 1-2; das Häufigkeitsverhältnis SIN/WaS war schon auf einer Strecke von

30 m in allen Jahren deutlich polarisiert.

Larvalökologie: nach KOCH (1984) monophag an Wacholder (Juniperus communis),

An einem solchen als Zierpflanze verwendeten Wacholder im Garten wurden

wiederholt in der Dämmerung 2% bei der Eiablage beobachtet. Dieser Strauch

liegt direkt neben dem Standort SiN. Von WaS, an dem die relative Häufigkeit

der Imagines deutlich höher ist, es jedoch das nächste Futterpflanzen-Vor-

kommen weiter entfernt (20-30 m). Entweder nimmt die Art auch Fichte als

Futter an (siehe Bemerkungen bei Thera juniperata) oder die Falter ziehen sich

nach dem Schlüpfen an geschütztere Standorte zurück, wo die Gefahr einer Ver-

frachtung durch Wind und der Feinddruck (z.B. Fledermäuse) kleiner sind.

Populationsbiologie: wenig proterandrisch j

Verbreitungsstrategie: K-Siratege,. 6. Gruppe; E. intricata gehört zur Gruppe der in

Südbayern als selten geltenden Arten, die in den letzten Jahren häufiger ge-

worden sind (siehe Bemerkungen zu Asthena albulata). Die Aussagen zur Strategie

von Thera juniperata gelten vermutlich auch für diese Art.

Eupithecia satyrata (17 Individuen)

Distanzen: ?2

Larvalökologie: vergleichsweise pays an niedrigen Pflanzen

Verbreitungsstrategie: K-Sitratege, 5. Gruppe

Eupithecia tripunctaria (51 Individuen)

Distanzen: 2(32)

Larvalökologie: E. tripunctaria zeichnet sich durch einen Wirtswechsel aus: die

Raupen der 1. Generation fressen an den Blüten des Holunders (Sambucus nie]

rar; die der 2. Generation an den Blüten von Bärenklau (Heracleum) und Engel-

wurz (Angelica); die Flugzeiten liegen jeweils kurz vor den Blütezeiten der be-

treffenden Pflanzen. Das SiM-Stück muß mindestens 100 m weit geflogen sein. Im

reinen Offenland (HM) konnte bisher noch kein Exemplar nachgewiesen werden.

rn nn

180 |

Populationsbiologie: bivoltin

Verbreitungsstrategie: intermediärer Typ, 3. Gruppe; Durch den Wirtswechsel wird

sicherlich in einigen Fällen ein Ortswechsel nötig. So wurde E. Lripunctaria im

Garten in der 1. Generation nur in 2 Exemplaren gefunden, während die 2. Ge-

neration häufig ist. Dieser Befund steht im Einkla mit der Verbreitung der

Futterpflanzen: Holunder kommt 10 m vom Standort WaS entfernt vor, die Um-

belliferen dagegen in einem Radius von mindestens 100 m nicht.

Eupithecia absinthiata (7 Individuen)

Distanzen: vermutlich 2

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Eupithecia assimilata (8 Individuen)

Distanzen: vermutlich 1-2 H :

Larvalökologie: nach KOCH. (1984) nn sich die Raupe nur von wildwachsen-

dem Gemeinem en Humulus lupulus d von Schwarzer Johannisbeere (Ri-

bes nigrum). CAR HARG VES (1987) geben darüber hinaus auch die

Rote Johannisbeere (Ribes rubrum) an, die vermutlich die Lebensgrundlage der

Population im Garten darstellt. Bisher konnten keine biotopfremden Tiere festge-

stellt, werden.

Populationsbiologie: bivoltin

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Eupithecia vulgata (56 Individuen)

Distanzen: 2, }

Larvalökologie: polyphag an niedrigen Pflanzen, im Untersuchungsgebiet ist diese

Art dennoch relativ lokal verbreitet.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Eupithecia castigata (153 Individuen)

Distanzen: 2-3 E

Verbreitungsstrategie: intermediärer Typ, 3. Gruppe

Eupithecia icterata (46 Individuen)

Distanzen: 2; im Wasserwerk fällt der Häufigkeitsgradient zwischen WaN und WNo

schon auf 50 m Distanz auf.

Larvalökologie: an Gemeiner Schafgarbe (Achillea millefolium) und Gemeinem Rain-

farn ( Tanacetum ulgare), dennoch konnte E. icterata bisher nicht an HM nach-

gewiesen werden.

Verbreitungsstrategie: intermediärer I. 5. Gruppe; im Garten fand 1987 offen-

sichtlich eine Neubesiedelung statt. Vorher konnte diese auffällige Art trotz in-

tensiver Fänge nicht nachgewiesen werden.

Eupithecia succenturiata (21 Individuen)

Distanzen: 2

Larvalökologie: wie E. icterata, zusätzlich nehmen die Raupen auch Gemeinen Bei-

Artemisia vulgaris) an.

Populationsbiologie: im Garten sind starke _Häufigkeitsschwankungen festzustellen.

Verbreitungsstrategie: intermediärer Iyp, 5. Gruppe;

Eupithecia subumbrata (30 Individuen)

Distanzen: 1-2

Larvalökologie: an Umbelliferen und Compositen, im Untersuchungsgebiet relativ

stark an die trockeneren Wiesen gebunden; außerhalb davon, z.B. im Siedlungsge-

biet konnte noch kein Stück nachgewiesen werden, was für Dispersionsaktivitäten

Adnan die normalerweise in einem Bereich von unter 500 m liegen. Die (Tritt-

stein-)Besiedelung des Wasserwerks in den letzten 10 Jahren erforderte jedoch

Sprünge von 100-300 m

Verb

reitungsstrategie: K-Stratege, 6. Gruppe

181

Eupithecia millefoliata (4 Individuen)

Distanzen: 2(4); E. millefoliata ist eine in der Ausbreitung begriffene östliche

Steppenart; abgesehen vom Erstnachweis für Südbayern in Paitzkofen bei Straubing

in einer Entfernung von 100 km (WOLFSBERGER, 1974) wurden aus Südbayern

bisher keine Fundorte gemeldet.

Laivalianlöge: monophag an Gemeiner Schafgarbe; bisher konnten keine biotop-

fremden Tiere nachgewiesen werden ;

Verbreit strategie: in einer Arealerweiterung begriffen; wenn die Art an günsti-

en Stellen bodenständig wird, dann handelt es sich um einen K-Strategen der 6.

ruppe.

Eupithecia sinuosaria (4 Individuen)

Distanzen: 3-4; auch diese östliche Art hat in den letzten 30 Jahren seit dem

Erstnachweis in Südbayern ihr Areal stetig nach Südwesten erweitert. Sie kam

hierbei durchschnittlich ca, 387) km pro Jahr voran. Siehe hierzu REZBANYAI-

RESER & WHITEBREAD (1987). j

Verbreitungsstrategie: in einer Arealerweiterung begriffen, sonst vermutlich inter-

mediärer Typ, 5. Gruppe

Eupithecia indigata (12 Individuen)

Distanzen: 1-2

Larvalökologie: auf Wald-Kiefer (Pinus sylvestris) und Gemeine Fichte (Picea_abies)

ge Bisher wurden alle Exemplare an den Standorten der Futterpflanzen

gefangen.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Eupithecia pimpinellata

Distanzen: vermutlich 2

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Eupithecia innotata (4 Individuen)

Distanzen: vermutlich 2-3

Larvalökologie: E. innotata ist - ähnlich E. tripunctaria - durch einen Wirtswechsel

charakterisiert: Die Raupen der 1. Generation leben an einigen Sträuchern und an

Esche (Fraxinus excelsior), die der 2. Generation an Beifuß-Arten Artemiuah

Populationsbiologie: bivoltin

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe; durch den Wirtswechsel wird

sicherlich in einigen Fällen ein Ortswechsel nötig. Die im Garten festgestellten

Stücke stammen aus mindestens 100 m Entfernung.

Eupithecia virgaureata (80 Individuen)

Distanzen: 2 (-3?)

Larvalökologie: Auch diese Art durchläuft einen obligatorischen Wechsel der Wirts-

pflanzen: Von Schlehe (Prunus spinosa) und Weißdorn (Crataegus) ernähren sich

ie Raupen der 1. Generation, die der zweiten dagegen vo chter Goldrute

Solidago virgaurea), Gemeinem Greiskraut (Senecio vul aris) oder Gemeinem

asserdost (Eupatorium na. Da an einer Reihe der Fundorte der Ima-

gines diese Futterpflanzenkombination in einer Entfernung von 100-200 m nicht

vorkommt, ist diese Art als vergleichsweise mobile Eupithecie anzusprechen.

Populationsbiologie: bivoltin; weder Proterandrie noch Protogynie; in der 2. Gene-

ration zeigte sich eine hohe 9-Rate von 76,7 % (1988). >

Verbreitungsstrategie: intermediärer Typ, 3. Gruppe; durch den Wirtswechsel wird

sicherlich in einigen Fällen ein Ortswechsel nötig. E. virgaureata gehört zur

Gruppe der als selten geltenden (und wohl oft nur en Eupithecien, die

in den letzten Jahren häufiger nachgewiesen werden (siehe Bemerkungen zu As-

ihena albulata).

Eupithecia abbreviata

Distanzen: vermutlich 1-2

Larvalökologie: monophag an Eiche, die am Fundort vorkommt

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Eupithecia dodoneata

Distanzen: vermutlich 1-2

Larvalökologie: wie E. abbreviata

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe; die drei 1989 im Garten

Be enen 99 sind die Erstnachweise für das südbayerische Faunengebiet (HAUS-

in Ya E. dodoneata wird wahrscheinlich oft übersehen.

182

Eupithecia sobrinata (1 Individuum)

Distanzen: 1-2 ä

Larvalökologie: monophag am Gemeinen Wacholder; alle Tiere wurden bisher in der

unmittelbaren Nähe der Futterpflanzen gefangen.

Populationsbiologie: die Häufigkeit schwankt im Garten beträchtlich

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Eupithecia lariciata (5 Individuen)

Distanzen: 1-2 \

Larvalökologie: monophag an Europäischer Lärche (Larix decidua), bisher konnten

keine biotopfremden Tiere nachgewiesen werden.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Eupithecia tantillaria (70 Individuen)

Distanzen: 1-2 _

Larvalökologie: die Raupen sind auf Gemeine Fichte spezialisiert, gelegentlich wird

auch Europäische Lärche akzeptiert; die FE ei sera henten im Garten und im

Wasserwerk, die einen starken Häufigkeitsabfall schon in 20-50 m Entfernung zur

nächsten Futterpflanze erkennen lassen, sprechen für Barrieren gegen die freie

eweglichkeit. Im gleichen Sinn ist wohl der fehlende Nachweis im Offenland

HM) zu interpretieren.

ndererseits konnte 1989 im Franzosenhölzl zwei fertile 99 500 m von der näch-

sten Fichtengruppe entfernt gefangen werden.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Eupithecia lanceata (32 Individuen)

Distanzen: 1-2

Larvalökologie: monophag an der Gemeinen Fichte; der deutliche Häufigkeitsgra-

dient im Wasserwerk auf ca. 100 m Distanz spricht für Barrieren, die gegen ei-

ne freie Beweglichkeit gerichtet sind.

Das 1986 an HM nachgewiesene J stammt aus mindestens 800-1000 m Entfer-

nung. Die Bewältigung einer solchen Distanz ist jedoch wohl als absolute Aus-

nahme zu verstehen.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Chloroclystis v-ata (165 Individuen)

Distanzen: 2-3

Larvalökologie: nach den in KOCH (1984) genannten Raupenfutterpflanzen beurteilt,

stammt das HM-Exemplar (1987) mindestens aus dem Ruderal, wo in ca. 200-250

m Entfernung die ersten irtspflanzen wachsen. Die Distanz von 1 km (Zuflug

vom Flughafenrand nach HM) scheint jedoch nicht mehr im normalen Dispersions-

geschehen bewältigt zu werden.

Populationsbiologie: bivoltin

Verbreitungsstrategie: intermediärer Typ, 2. Gruppe; KOCH (1.c.) bezeichnet C. v-

ata als "Arealausbreiter".

Calliclystis chloerata (6 Individuen)

Distanzen: vermutlich 2

Larvalökologie: monophag an. Schlehe (Prunus spinosa); bisher konnten keine bio-

topfremden Tiere, festgestellt werden.

Verbreitungsstrategie: vermutlich K-Stratege, 6. appei der Erstnachweis dieser

Art für Südbayern wurde erst durch WOLFSBERG 1950; 1958) erbracht. Ob

dies auch bei C. chloerata auf Arealerweiterungen zurückzuführen ist, oder ob die

get Auf so lange mit C. rectangulata verwechselt wurde ist derzeit schlecht zu

eurteilen.

Calliclystis rectangulata (111 Individuen)

Distanzen: 2

Larvalökologie: die Raupen sind auf Apfel- und Birnbäume spezialisiert (Malus do-

mestica und Pyrus munie), nach CARTER & HARGREAVES (1987) wird gele-

gentlich auch Schlehe angenommen, was z.B. im Falle der WNo-Stücke als Ent-

wicklungsort der Raupen sehr wahrscheinlich ist. Die beiden ersigenannten Pflan-

zen sind auch von einer Reihe weiterer Fundorte mehrere 100 m entferni. Die

Distanz von 1 km (Zuflug vom Flughafenrand nach HM) liegt nicht im Bereich

des normalen Dispersionsgeschehens von C. rectangulala.

Verbreitungsstrategie: K-Stratege, 4. Gruppe

183

Horisme tersata (16 Individuen)

Distanzen: 1-2

Larvalökologie: die Raupe ernährt sich von _Waldrebe (Clematis) und vpm Pr ouen

Windröschen (Anemone sylvestris), nach CARTER & HARGREAVES (1987 auch

vom Scharfen Hahnenfuß (Ranunculus acris). Biotopfremde Stücke wurden bisher

nicht registriert. Vermutlich akzeptiert die Art im Untersuchungsgebiet die in den

Gärten häufiger BDSEDI RIES Clematis- Ziersträucher.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

BOARMIINAE

Arichanna melanaria (? Individuen)

Distanzen: vermutlich 1-2, in Vorstößen bis 4

Larvalökologie: A. melanaria ist in Südbayern ein in (Hoch-)Mooren sehr (kei ver-

breiteter Spanner, der an das Vorhandensein von Moor-Heidelbeere (Vaccinium

uliginosum), Sumpf-Porst (Ledum palustre) oder Moosbeere (Oxycoccus palu-

stris) gebunden ist. Da keine dieser Pflanzen im Wasserwerk und vermutlich auch

in der weiteren Umgebung vorkommt, handelt es sich bei den beiden in der glei-

chen Nacht gefangenen JT' um ebenso überraschende wie interessante Funde! Die

ER eNIRBTe waren beide stark abgeflogen und stammen vermutlich aus Biotopen in

einer, Entfernung von mindestens 20-30 km.

Verbreitungsstrategie: vermulich K-Stratege, 6. (aivppe: wahrscheinlich handelt es

sich hier um ein Phänomen, dem wir schon bei den Schilfeulen begegnet sind,

nämlich, daß auch orts- und habitatireue Arten bei einer Vernichtung des Le-

bensraumes oft weite Strecken zurücklegen können.

Sehr interessant ist in diesem Zusammenhang eine Publikation HACKERs (1981),

der in Nordbayern bei A. melanaria einmal genau dieselbe Beobachtung machte

und eine zurüc Ser el Distanz von ca. 50 km vermutet.

Auch OSTHELDER (1925-1933) berichtet von einem Stück im (18 km entfernten)

Eichenau, das "offensichtlich verweht" wurde. Auch hier betragt die Entfernung

zum nächstgelegenen von Östhelder genannten Fundort 20 km.

Abraxas grossulariata (1 Individuum)

Distanzen: vermutlich 1-2

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe; in Südbayern tritt diese

Art nur sehr lokal und selten auf.

Calospilos sylvata

siehe Versetzexperiment (8.4.) und Rückschlüsse aus den Ortswiederfängen (8.5.)

1249* Individuen 10,1 % 9- Rate

644** markiert 46** Wiederfänge

* tagsüber viele weitere Stücke

**1989 343 markiert, 53 Wiederfänge

Wiederfang-Quote: sehr hoch; C. sylvata wurde nur 1988 und 1989 markiert, alle

Wiederfänge erfolgten am Fangplatz "We". Hier wurde 1988 im 3- Tage-Rhythmus

ee die Ortswiederfang-Quote von 9,0 % und die mittlere Verweilzeit von

‚8 Tagen kennzeichnen C. sylvata als die ortstreueste aller im Rahmen dieser

Arbeit markierten Arten. Nimmt man die in den Versetzexperimenten rückgefan-

enen Tiere hinzu, ergibt sich mit 4,6 Tagen ebenfalls eine sehr au ttlere

nerweilzeit, Die längsten beobachteten Intervalle waren 3 Exemplare (d’d’9) nach

agen.

Die beiden Geschlechter verhielten sich in den Experimenten ungefähr gleich.

Distanzen: 1-2

Larvalökologie: die Raupe lebt an einer Reihe von Laubbäumen. Im Offenland (HM)

und schon 250 m außerhalb des Birkets (Moos 1985) konnten keine Stücke mehr

nachgewiesen werden.

Verbreitungsstrategie: K-Stratege, 6. Gruppe; abgesehen von den 1988 im Garten

beobachteten Exemplaren wurden alle außerhalb des Mooses nachgewiesenen

Stücke in maximal 200 m Entfernung zum Schloßkanalsystem gefunden. Diese Ka-

näle könnten unter Umständen als "Leitlinien" einer Verbreitung solch wenig mo-

biler Wald(rand)bewohner förderlich sein.

184

Lomaspilis marginata

357 Individuen 5,9 % 9- Rate

236 markiert 5 Wiederfänge

Tab. 100: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Lomaspilis marginata.

SEDLUNG WALD HALBTROK- en WALD HALBTROK- "DACH.

1987 | Garten | KENRASEN M 2 1988 Garten Wasserwerk KENRASEN MOOS”

SiS SM |SN WaS|WaM WaN HO HM HW Mb SN WaS WaN WNw WNo HO HM HW Au We

Spar, 8,1. 6 =, A038 BARS 30 1020 >epar. a 4 531 31 43 Er

% zus. Tem 2 - 10 1 1 4 1 20 8% zus. - 2 = - > = = u = =

[o107 6 - 3 - 228 7 4 5 17 72 [oj 07 3 2 3600255230 3 2 2 12 36

99 Ale eu. warlam 1 8.09 Slas22- 2 asien: 3

Mark. sem a' =... 8%) TANTE 18 80 Mark. 3 a... er ad €

Wi. une sa. e SEN = - - Wi. - 1 1 =. Sa Me a

Wiederfang-Quote: durchschnittlich; ein Wiederfang (on WaS) ist durch den tägli-

chen Betrieb der Lichtfalle bedingt. Die mittlere Verweildauer der anderen vier

wiedergefangenen Jod’ betrug niedrige 2,3 Tage.

Im Wasserwerk, wo die Wiederfang-Quote mit 3,2 % etwas über dem Durch-

schnitt liegt (bedingt durch die bessere Flächenabdeckung durch 3 Falley), wa-

ren der 3 rückgefangenen Tjere Ortswechsler, der eine über 100 m t Nw>

WaN), der andere über 120 m (whw> WNo)

Distanzen: 3,

Larvalökologie: die Raupen sind an einigen Laubbaumarten zu finden; die regelmä-

Bigen und vergleichsweise zahlreichen Nachweise an HM zeigen, daß der Yuflug

aus dem Ruderal (150-300 m) für diese Art (dd und 99) eine völlig normale

Distanz darstellt.

Populationsbiologie: im Untersuchungsgebiet vermutlich teilweise in 2 Generationen,

ie phänologisch jedoch nicht voneinander abzutrennen sind. Proterandrisch

Verbreitungsstrategie: r-Stratege, 3. Gruppe

Ligdia adustata (74 Individuen)

Distanzen: 2,

Larvalökologie: ImunupaR am Europäischen Pfaffenhütchen (Euonprats europaea); an

einigen Standorten ljegt die nächste Raupenfutterpflanze 100-200 m entfernt. Die

Distanz von 1 km (Zuflug nach HM) liegt nicht im Bereich des trivial move-

ment bzw. der Dispersionsaktivität dieser Art.

Populationsbiologie:, bivoltin

Verbreitungsstraiegie: intermediärer Typ, 5. Gruppe

Bapta bimaculata (49 Individuen)

Distanzen: 1-2; die Häufigkeitsgradienten im Wasserwerk und im Garten könnten

auf Barrieren gegen die freie Beweglichkeit hinweisen.

Larvalökologie: an einigen Laubbaumarten; im Offenland (HM) und an einigen

Standorten mit tendenziellem Offenlandcharakter (SiM, SiN, WaN und HW) konn-

te die Art bisher, nicht nachgewiesen werden.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Bapta temerata (196 Individuen)

Distanzen: 2 (-3?)

Larvalökologie: die Raupe ernährt sich von einer Reihe von Laubbaumarten; das an

HM nachgewiesene Jd' stammt zumindest aus dem Ruderal (150-300 m).

Verbreitungsstrategie: intermediärer Typ, 3. Gruppe

Cabera pusaria

232 Individuen 21,6 % o- Rate

72* markiert 1 Wiederfang * nur 1988

Wiederfang-Quote: niedrig! Unter "regulären" Bedingungen (fangfreie Nächte zur

Ermöglichung der freien Durchmischung) konnte kein Tier wiedergefangen wer-

den. Lediglich an WaS flog ein JS nach 1 Tag zum zweitenmal die Falle an; die

Wiederfans "Quote liegt hier mit 4,0 % auf einem für die Methodik sehr niedrigen

iveau!

Distanzen: im Habitat 2-3, außerhalb 1-2; der Häufigkeitsgradient zwischen WaN

und WNo (50 m) deutet wohl auf Barrieren gegen eine freie Dispersion hin. _

Larvali rolöpe: die Raupen sind an Weide, Birke, Ulme Erle und Eiche zu finden;

das HM-Stück (c) stammt vermutlich aus dem Ruderal (150-300 m). 5

ulationsbiologie: im Untersuchungsgebiet mit einer partiellen 2. Generation

Verbreitungsstrategie: intermediärer Iyp, 5. Gruppe

185

Cabera exanthemata (273 Individuen)

Distanzen: 3

Larvalökologie: die Raupe ist an einigen Laubbaumarten zu finden; die HM-Stücke

stammen wahrscheinlich aus dem Ruderal (150-300 m). Diese Distanzen liegen im

absolut normalen Bereich der Dispersionsaktivitäten.

Populationsbiologie: bivoltin

Verbreitungsstrategie: r-Stratege, 2. Gruppe

Plagodis dolabraria (26 Individuen)

Distanzen: ?

Larvalökologie: ebenfalls an einigen Laubbaumarten; bisher konnten keine biotop-

fremden Tiere nachgewiesen werden.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Ellopia fasciaria (1 Individuum)

Distanzen: vermutlich 1-2 ?

Larvalökologie: monophag an Kiefer (die Angaben beziehen sich bei dieser und de

folgenden Art auf das Artverständnis wie es in FORSTER & WOHLFAHRT 1981)

niedergelegt ist. Das an HW festgestellte Stück muß aus 150 m Entfernung (dem

nächstgelegenen Standort ca. 2 m hoher Kiefern) ohne Sichtkontakt herbeigeflogen

sein.

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Ellopia prasinaria (6 Individuen)

Distanzen: 1-2

Larvalökologie: die Raupen ernähren sich von Fichte, Lärche und Weißtanne; bis-

her wurden alle Stücke in der Nähe der Raupenfutterpflanzen gefangen.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Campaea margaritata (27 Individuen)

Distanzen: 1-2

Larvalökologie: die Raupe ist an Een, Laubbaumarten zu finden, biotopfremde

Tiere konnten bisher nicht festgestellt werden. Der Häufigkeitsgradient 1988 im

Garten auf nur 30 m Entfernung hängt vermutlich mit dem an SiN etwas weni-

ger ausgeprägten Waldcharakter zusammen.

Populationsbiologie:, bivoltin

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Ennomos autumnaria (4 Individuen)

Distanzen: vermutlich 2

Larvalökologie: relativ polyphag an Laubhölzern, bisher keine biotopfremden Tiere

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Deuteronomos alniaria (? Individuen)

Distanzen: vermutlich 2

Latvalöknlogi: an einigen Laubbaumarten, das an HM gefangene 7 stammt vermul-

lich aus dem Ruderal (150-300 mi:

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Deuteronomos fuscantaria (? Individuen)

Distanzen: vermutlich 2

Larvalökologie: die Raupen sind auf Gemeine Esche (Fraxinus excelsior) und Ge-

meinen Liguster (Ligustrum vulgare) spezialisiert. Die in manchen Jahren in den

Garten zufliegenden Exemplare stammen aus mindestens 80 m Entfernung.

Verbreitungsstrategie: vermutlich K-Stratege, 6. Gruppe

Deuteronomos erosaria (13 Individuen)

Distanzen: 2-3

Larvalökologie: Raupenfutterpflanzen sjnd vor allem Eiche, aber auch Birke, Linde

und Rot-Buche. Das HM-Exemplar (1986) stammt also zumindest vom ae

rand (800-1000 m). Dies scheint in Jahren mit günstiger Bestandsentwicklung (z.B.

1986) ungefähr die Obergrenze der normalen Dispersionsaktivität darzustellen. _

Populationsbiologie: im Garten fallen starke Häufigkeitsschwankungen auf, die viel-

eicht ein weiterer Hinweis dafür sein könnten, daß D. erosaria im Vergleich mit

den anderen Arten der Gattung die dynamischste Strategie aufweist.

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

186

Selenia bilunaria (47 Individuen)

Distanzen: ?2

Le: ralökslogıe: nach den in KOCH (1984) genannten Raup

stammen die beiden HM-Stücke vom Ruderal (150-300 m

Populationsbiologie: bivoltin

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

ee beurteilt,

Selenia tetralunaria (45 Individuen)

Distanzen: 2

Larvalökologie: nach den in KOCH a gena ten Raupenfutterpflanzen beurteilt,

stammt das HM-Stück vom Ruderal (150-300 a)

Populationsbiologie: bivoltin

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Apeira syringaria (11 Individuen)

Distanzen: 1-2 #

Larvalökologie: A bisher festgestellten Stücke wurden in unmittelbarer Nähe der

in KOCH 11984 genannten Futterpflanzen gefangen.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Gonodontis bidentata (31 Individuen)

Distanzen: 2

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Colotois pennaria

89 Individuen 7,1% 9- Rate

28 markiert 6 Wiederfänge

Wiederfang-Quote: durchschnittlic 9 Stichprobe noch zu klein. Alle Wiederfänge

stammen aus dem Garten (WaS) 1988 bei täglichem Betrieb der Lichtfalle. 5 der

6 Rückfänge (dc, darunter zwei, Zweitwieder änge) erfolgten nach einem Intervall

von 1 Tag und sind kein Hinweis auf eine mögliche Ortstreue der Imagines. Ein

cd’ flog die Falle nach einem Intervall von 2 Tagen wieder an.

ie mittlere Verweilzeit liegt mit 1,8 Tagen auf einem relativ niedrigen Niveau

(wie beispielsweise auch beim Wanderfalter Noctua Bromieaı

Distanzen: im Habitat 2-3, außerhalb 1-2; die etwas pump wirkenden 99 viel-

leicht auch nur 1-2; ihre schlechtere Flugtauglichkeit könnte auch in Zusammen-

hang mit einem nicht optimalen Anflugverhalten an das Licht stehen.

Larvalökologie: relativ golyphae an Laubbäumen; das polarisierte Häufigkeitsver-

hältnis im Garten 198 12/2 Individuen in den Parallelfängen) ist wohl als Hin-

weis darauf zu verstehen, daß zunehmender Offenlandcharakter (SiN) schon nach

30 m als Barriere gegen die Verbreitung zu wirken beginnt.

Verbreitungsstrategie: K-Stratege, 5. Gruppe; für die Verbreitung ist das Verhalten

der 99, nicht der dc’ relevant. Eine höhere Mobilität der dd‘ kann jedoch zu ei-

ner besseren Gendurchmischung nützlich sein.

Crocallis elinguaria (104 Individuen)

Distanzen: 2

Larvalökologie: von den in KOCH (1984) m ten Ran I on enzen kommt in

der näheren Umsebung des Fangplatzes M (2 Exemplare 1986) nur der Weiß-

dorn in Frage. Es gilt hierbei analog das bei Allophyes oxyacanthae (Noctuidae,

Cuculliinae, S. 136) Gesagte. Der Häufigkeitsgradient im Garten 1988 ist vermut-

lich wie bei Colotois pennaria zu interpretieren.

Verbreitungsstrategie: K-Stratege, 5. Gruppe

Angerona prunaria (19 Individuen)

Distanzen: ?; 1989 wurde im Mallertshofer Holz (Hochwald) ein 4 in der Abend-

dämmerung bei einem ziemlich geradlinigen Flug über 200 m beobachtet.

Larvalökologie: nach den in KOC (1984 genannten Wirtspflanzen beurteilt, wur-

den im Uni eszuchunses@biet bisher keine biotopfremden Tiere registriert.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

187

Ourapteryx sambucaria (10 Individuen)

Distanzen: 1-2 }

Larvalökologie: nach den in KOCH 11984) genannten Wirtspflanzen beurteilt, wur-

den im Untersuchungsgebiet bisher keine biotopfremden Tiere registriert.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Opisthograptis luteolata (39 Individuen)

Distanzen: 2

Larvalökologie: die Raupe lebt an verschiedenen Laubgehölzen, v.a. an Sträuchern;

die beiden im, Offenland gefundenen Exemplare (9) stammen wahrscheinlich aus

dem Ruderal (150-300 m).

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Epione repandaria (31 Individuen)

Distanzen: 2-3

Larvalökologie: diese Art kann sich an Weide, Espe, Erle, Schlehe und Pappel

entwickeln. Die Distanz von 150-300 m (Ruderai>HM) liegt im Bereich der

normalen Dispersionsaktivität von E. repandaria.

Populationsbiologie:, bivoltin

Verbreitungsstrategie: intermediärer Typ, 5. Gruppe

Cepphis advenaria (23 Individuen)

Distanzen: 1-2

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Lozogramma chlorosata (1 Individuum)

Distanzen: vermutlich 1-2

Larvalökologie: die Raupen ernähren sich von Adlerfarn (Pteridium aquilinum), viel-

leicht auch von Gemeinen Wurmfarn (Dryopteris filix-mas). Das nachgewiesene

Eaenpler wurde in der Nähe der Futterpflanzen gefangen.

Verbreitungsstrategie: K-Stratege, 6. Gruppe

Macaria notata (39 Individuen)

Distanzen: 2,

Larvalökologie: an Birke, Erle, Sal-Weide und Eiche zu finden; das 1986 im Offen-

land gefundene Stück stammt vermutlich aus dem Ruderal (150-300 m)

Populationsbiologie: die 2. Generation dieser Art ist nur partiell ausgeprägt.

Verbreitungsstrategie: intermediärer Typ, 6. Gruppe

Macaria alternaria

279 Individuen 10,4 % 9- Rate

77 markiert 1 Wiederfang

Wiederfang-Quote: durchschnittlich, an WaN erfolgte 1988 ein Ortswiederfang (C)

nach 2 Tagen.

Distanzen: 2,

Larvalökologie: die Raupe frißt an Sal-Weide, Eiche, Erle, Schlehe und Traubenkir-

sche. Die beiden wahrscheinlich vom Ruderal (150-300 m) her zugeflogenen HM-

Stücke (dd) zeigen, daß eine solche Distanz wohl innerhalb der Grenzen der

normalen Dispersionsaktivität liegt.

Eu oa die 2. Generation dieser Art ist nur partiell ausgeprägt; prote-

randrisc

Verbreitungsstrategie: intermediärer Typ, 3. Gruppe

Macaria signaria (31 Individuen)

Distanzen: 1-2 i 1

Larvalökologie: die Art ist an das Vorkommen von Gemeiner Fichte oder Heide-

kraut gebunden. Im Untersuchungsgebiet vor allem an Fichte, wie Raupenfunde

bestätigten. Der bisher am weitesten von der Futterpflanze entfernte Nachweis ist

ein d an WNOo (ca. 50 m)

Verbreitungsstrategie: K-Stratege, 6. Gruppe

188

Mecaria liturata

243 Individuen 29,9 % o- Rate

163 markiert 5 Wiederfänge

Tab. 101: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Macaria liturata.

EN ee TE a ES

SIEDLUNG ALD HALBTROK- eE WALD HALBTROK- "DACH.

1987 | Garten KENRASEN M 2 1988 Garten Wasserwerk KENRASEN MOOS’

SiS SIM|SIN WaS |WaM Wan HO HM HW Mb SIN WaS WaN WNw WNo HO HM HW Au We

Ipr.. - 29 17 15 25 Vote - 80 par 5 26 25 28 10 if 1 -

De zusue I =D 11 - 3 10 - 2 - 28 2% zus - 29 = = - = = = =

de = Mm on Aa 9 Meet - 48 dd 25 Bo 6 5 m

9 Ss.) A Snwlr is Zoe 2 - 24 99 3 8 EEE <) I a

Mark. - 2 55 Es ars - 71 Mark. 2033 190722069 6 Er =. 8

Wit. EINEN ie SW esse = =. We. 1 Se Eee Eee EEE

Wiederfang-Quote: niedrig! Alle Rückfänge erfolgten nach einem Intervall von nur

Tag und erklären sich also durch ein Festgehalten-Werden am Licht. Ein o

wechselte den Ort von WaS nach SiN, wo es tags darauf noch einmal gefangen

wurde. Die mittlere Verweildauer liegt mit 1,3 Tagen sehr niedrig. Die o-Rate

im Wiederfangergebnis entspricht ungefähr der der Erstfänge.

Distanzen: 2-3

zu, ee : auf Nadelhölzer spezialisiert; an HM wurden 1986 2 biotopfremde

Stücke (do) nachgewiesen, die mindestens 800-1000 m geflogen waren. Die Häu-

figkeitsgradienten im Wasserwerk und im Garten 1988 sprechen jedoch für Bar-

rieren (O fenlandeharakt,) Sehen die Verbreitung dieser Ärt.

Populationsbiologie: bivoltin, 2. Generation nicht vollständig; proterandrisch

Verbreitungsstrategie: intermediärer Ip 5. Gruppe; im Zusammenhang mit der

günstigen Bestandsentwicklung fast aller an Kieiern lebenden Arten 1986 erfolg-

te auch bei liturata ein "Verbreitungsschub”, der sich in Vorstöße auf das

Offenland hinaus bemerkbar machte. In normalen Jahren liegt dagegen die Ober-

grenze der Dispersionsaktivität über biotopfremdes Gebiet niedriger.

Chiasmia clathrata

siehe verringerte Fallendistanzen (8.3.)

157 Individuen 3,3 % 9- Rate

112 markiert 11 Wiederfänge

Tab. 102: Fangergebnisse, Geschlechterverteilung und Markierungsergebnisse von

Chiasmia clathrata.

1987 SIEDLUNG WALD HALBTROK- DACH 19897 SIEDLUNG WALD HALBTROK- "DACH.

!G KENRASEN MOO L KENRASEN MOOS”

| Garten |

1.6 arten 2. Gen

. Gen. sis sim|sin Was|WaM WaN HO HM HW M i SiS SIM|SIN WaS|WaM WaN HO HM HW Mb

Spar - Ay BB A 2 1 A Zpa. I - - ir 18127459 ) 1

% zus. her = - - 3 - - - - su e27zus: Zu - - 1 1 - 2 -

ds - a - 23 dd I er 8 6 ae =

99 =. ayıo Er N = 109 Es Ei en So 1

Mark. = 2 0. Shi A, N aa‘ - 2A Mark I 8 9% BURE-IIENA: 1

W.f. au a ee Sun - 3 wi. ee En (ER EN 2

1988 WALD HALBTROK- "DA! 1988 WALD HALBTROK- "DACH.

Garten Wasserwerk KENRASEN MOO: DB Garten Wasserwerk KENRASEN MOOS”

1. Gen. sy Was WaN WNw WNo HO_ HM HW Au We SIN WaS WaN WNw WNo HO HM HW Au We

I par. - 5 2 19 gr N 3 1 - 36 2% par. - 2 6 SEm15 Ders" >)

& zus. = = > - = = = = - - = 2% zus. = 1 = > S - = = - -

[oje] - - 3 1 16 3 1 2 - - 26 dd - 2 5 3 10 6 - 3 - -

29 z 7 EUREN Se Frelern = =109 = 7 at Zur Den =

Mark. > = 3 1 15 3 1 2 _ - 25 Mark. = 2 5 2 10 6 - 3 -