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24 Contributions

of the

Volume 24, 1988

eye

No. 1. Harbach, Ralph, E. The mosquitoes of the

subgenus Culex in Southwestern Asia and Egypt

(Diptera: Culicidae). July 8, 1988. 240 pages.

No. 2. Weaver, John S. Ill. A synopsis of the

North American Leptosomatidae (Trichoptera).

November 1, 1988. 141 pages.

No. 3. Menke, Arnold S. Pison in the New World:

a revision (Hymenoptera: Sphecidae: Trypoxylini).

November 21, 1988. 171 pages.

JUN 12 1989

LIBRARIES

Contributions

of the

American Entomological Institute

Volume 24, Number 1, 1988

oy"

| THE MOSQUITOES OF THE SUBGENUS CULEX IN

SOUTHWESTERN ASIA AND EGYPT (DIPTERA: CULICIDAE)

Ralph E. Harbach

Dedicated to

Kenneth L. Knight,

my much admired mentor, colleague and

friend, whose collections made in

Egypt and the Yemen Arab Republic

considerably enhanced this work.

PREFACE

The mosquito fauna of southwestern Asia is limited in taxonomic diversity. Only

98 species are recorded from the region, but at least 25 of these are reportedly involved

in the transmission of pathogens that cause malaria, filariasis, and arboviral diseases of

importance to man. Epidemiological studies and the control of the associated vectors

are extremely limited due to difficulties in making precise species determinations. These

difficulties are largely attributed to the absence of up-to-date taxonomic treatments for

the various species. This work is a contribution toward defining part of the mosquito

fauna of the region. It is a preliminary treatment of the species belonging to the

subgenus Culex. Its primary objective is to provide modern keys, descriptions, and

illustrations for recognizing these species in the adult, pupal, and larval stages.

| consider this work to be preliminary because too few specimens were available

to determine the actual distributions and the total range of morphological variation for

most of the species. Most of the nomenclatural problems have been solved, but many

biosystematic questions require further investigation. The taxonomic status of some

forms remains uncertain. Perhaps the greatest contribution of taxonomic study is not in

filling gaps in our Knowledge about species, but in showing how many gaps still exist. If

this manual allows the user to make accurate species determinations, then its primary

purpose will be fulfilled. It is my hope that this work will also serve as a basis for further

research on the included species.

This report is based on data and observations accumulated at the Walter Reed

Biosystematics Unit (WRBU) (housed in the National Museum of Natural History,

Smithsonian Institution) between July 1980 and December 1984. The work began as a

study of Culex pipiens Linnaeus. It was broadened in January 1982 to include all

members of the subgenus Culex in southwestern Asia and Egypt.

| am indebted to many persons for the successful completion of this manual, and |

am most happy to acknowledge their help. | owe much to the stimulating discussions

about systematic problems and taxonomic procedures | had with Bruce A. Harrison, E.L.

Peyton, Curtis W. Sabrosky, F. Christian Thompson, Michael E. Faran, and Thomas J.

Zavortink while we were working together at the Smithsonian Institution. Many

individuals have been most generous in loaning specimens or otherwise assisting me

with the study of collections under their care: Peter S. Cranston, British Museum

(Natural History), London; John Lane, London School of Hygiene and Tropical

Medicine; Francois Rodhain, Institut Pasteur, Paris; Jean Clastrier, Museum National

d'Histoire Naturelle, Paris; A. Rickenbach, Office de la Recherche Scientifique et

Technique Outre-Mer, Bondy; Paul H. Arnaud, California Academy of Sciences, San

Francisco; J. Callot, Institut de Parasitologie, Strasbourg; Amnon Freidberg, Tel Aviv

University; Joel Margalit, Ben-Gurion University of the Negev, Beer Sheva; M.W.

Service, Liverpool School of Tropical Medicine; Thomas Gaffigan and James Pecor,

WRBU, National Museum of Natural History, Smithsonian Institution; and staff members

of the Dokki Institute and the Entomological Society of Egypt, Cairo. Sherif el Said, Ain

Shams University, Cairo, and Joel Margalit (affiliation given above) are especially

acknowledged for their special effort in assisting Bruce A. Harrison and myself in

collecting a considerable amount of material in Egypt and Israel, respectively, that was

examined during the study. All of the illustrations were meticulously and skillfully

prepared by Taina Litwak, WRBU. They are a most valuable asset to this work. The

entire manuscript was read critically by Ronald A. Ward. His comments and suggestions

have resulted in many important improvements. Finally, | warmly acknowledge the con-

stant love and support of my wife Roberta, son Paul, and daughter Andrea who patiently

tolerated my use of hundreds of hours of family time to complete this manual.

Bangkok, Thailand Ralph E. Harbach

October, 1987

CONTENTS

RVCIACE a kok ee Ree te Ce ac eee ee, il

BCU oe acesccn iB costs, tse. sal EP ee aa be pl RG ae Fe ra ctl, tae ihe omen aa 1

PVF RAVINE GY koe ave, Ea Mo We We ol. De ab so cng mes RU ac eats tltay eve el ay Tae. ae ce 2

HAA STONY iG sepa ich ew gd oR ah AD Ole te an cal a ta oe Ge ae te ep a 2

PAGHOTAL c.g ears ci eo eg hac INR lee tea ee at 2

5200.8 0) 0,0) Anas ar ar amen aaeee Won eeenape iy ety mene re Ree orc) Aero en y Lie heater ap hin 3

fe.” circa aheneeet Te prc mets aR (ir RRA T RENE iting trp ie nis Rr eae ie Aer UN 3

IS ass sal a pyle i taeda tite cae en I a eet he nee it ecu ate Is 3

SPECIES TEA TIONUS in sateanets Ud aire Autres rate Pilg ital Ua aunt cig eee feac a Nadel ELON, 4

TYNE GRO SIONS ie an. 5. sect te cas, Gh ah ace a ee i gc ka pac a 4

Hustrations cigs Aa eat 58 Fey Gee Lee ae SOE a i es 4

Keys to generac Culitidae’. setiris ak Oren, See, Mae Sie ere. Be <

AOS... CPI APO EMG, Je Pies aa). Ss eee ee 5

PUG nasa, eee et A PR a Sa eg ree ee eee SENT SUM Oe: ee hoe 5

Larvae caiieus. ARMY Sy AE Oe Se or ee Be ee 6

Keyetosubdenera-etCuley Sock pega eine ies . eek, Se etree Leen, e 7

PS 2 i ARR FBS). Ee ed a es ee a 7

Puget ei. Fe eR othe. GRD US a i Le PN ae 8

AIRS 52 shs ig eT FL Ry ee a ees oe eee eagle 8

Gane Cape. .chwituc. wacker atl ehiey Sie Rs Soe erate are comer 10

Subgenus Culers oiihs+ Srereuces, AF ae en ae cept ree, batt aE oe alg Mere Tam, 10

linear arrangement-olinciunedtaxan: sou er... Oeeele ea. Rei ee Ses 11

Pistribulonalist etthespecles 6) gists, ae Rees Sree ws. chek 11

Loocenoranhiogiinities . fil. teak EN SI ORS EE ee 12

Keysto siecies:ef Culex (ule i t.%:.. gupcion:, Weeien. ties Ae iene do Fee 13

PONG el 6 6 RRR ORR) BU PU. Ga ae! BGS 13

PUNO ry ha Se etek. ae ae Bhan ey ea Ram aie d, Mere eeaR a. keen Sohne 17

LAIN AO cise aad held CAS Sian MOEN Dla Geli f Maia Alas aa cig get PRA nig 19

Wie SOCIO SH STINGS cee cee ki le oe el a a a eh cee ana ah aa hg | Ue cr 23

CHIEN TC UIA PIMA TARGUS 6 Sic Gh bay vn tgs pect ie Ie Le ple cas eee doin ae tk 23

Culex (CUE) CUIPBUBIASCIAN IG EY ops hogs «ae dew ape ae aah Oa eae a a1

Culex (Caper) Vag ans VEGAN: ce ck 2 eles, fie wig kN ws wl ack, ABUL AL meas Ae aes 36

Culex (Calas) Tomer a RNe a. kh, bs ae qae u H. Raa oa ea bees ae a 40

Cinlox (Culler) CeCe se Veg as a) og oy be wa ver pn moe ROBICRE ot “ALIBI a SE lovelah Ge al al Phin 44

Culex (CUIOK) STOR IBE ie RBG ae heed a aiclgl ae oh oe geben ar gs eps 48

Culex (COUN) URIS TIVE ge g wh Gels pi cela AUR. Seaways Ho iano aug Uh Blak 52

Culex (Culex peremis TSG ANG nek ee ED. we ke le la a 57

Cislex (Cue POR VN, eae fee ies Ty. ov sgeeie eels gc epih ga ott bs ee orm Re 61

CHEK (CHIEN) TACIT TANTS fo da. glum OA, we ek Aight “eb Chas gue Walaa 67

CALAN (OUIO) TIT Te coi thsgil a tee ak ae! ag lecto hind erat ooag ey azo pel ca sl 72

Culex (OU) SING TOGA 6 isi bw. oe Gwe kw a laa Sp ae eae ae 75

Calox (Culexperiali cus Kingsathick® 6 S68 MAA PORN FL ae eS, 78

Culex (Culex Guinot THeebalg Soe PP Re ee ee ea Le eee 82

Ce TOUNa) SHIGIS WACO Golscde y tas es ee sila ce he tg ea ae a eee 87

Cuter (C NON BOOS TGA ie yu sue k's dw uc yy cau Wig) Oi anuwea oly lw morula 92

Ce Cree eC re ge ie ao kei aS iatlan! we ice gals ee cc 97

Culex (Culex) DIGGMICM AICS. HES, soap ay ahold 8 acensPAsons orl ene ating BULK) aoeattes « 101

Culex (OGIEX) THAENIOMYNCHIS GES OR ee a a 106

Cex (CUE PSCUGOVISINIET CONGSS ak a ke le ee “IB.

Pee tr ye Sh Oe ie Ryo mca anne noe ma Mey anes kt ig aa. 8 alas as 116

ie eh i Re eae a tana wee SORE a We tala le kA wa a 142

Ue ANG ae eS Pam AS Once vce GN. Mi le ane a 186

Oa CN uit 0d eR rea tan AB eh ABE) CSU feet a peu ORS el Sg Pn a 227

A. Species of Culicidae other than Culex (Culex) known

fo OCGurr ify SoutWweSteIn Asia ANG EQVON. a be ee. 227

B. Complete synonymy of Culex (Culex) pipiens Linnaeus ........... 229

C. Complete synonymy of Culex (Culex) quinquefasciatus Say ......... 232

D. Complete synonymy of Culex (Culex) sitiens Wiedemann .......... 235

OES CES OP ROTI CPO ee eee a! oe 236

ee ee eet eR Ue tele s G a am ae Sb ia gains 237

THE MOSQUITOES OF THE SUBGENUS CULEX IN SOUTHWESTERN

ASIA AND EGYPT (DIPTERA: CULICIDAE)

Ralph E. Harbach'.2

ABSTRACT

Twenty species of Culex of the subgenus Culex are recognized in southwestern

Asia and Egypt: pipiens Linnaeus, guinquefasciatus Say, vagans Wiedemann, for-

rentium Martini, decens Theobald, antennatus (Becker), univittatus Theobald, per-

exiguus Theobald, theileri Theobald, /aticinctus Edwards, mattinglyi Knight, simpsoni

Theobald, sinaiticus Kirkpatrick, duttoni Theobald, sitiens Wiedemann, poicilipes

(Theobald), mimeticus Noe, bitaeniorhynchus Giles, tritaeniorhynchus Giles, and

pseudovishnui Colless. Keys, descriptions, and illustrations are provided for the

identification of the adult, pupal, and larval stages of each species. Keys are also

included for the genera of mosquitoes and the subgenera of Cu/ex occurring in the

region. The individual treatment of each species also includes a complete synonymy, a

summary of previous literature, a systematic discussion, information on bionomics and

distribution, and a synopsis of material examined. Lectotype specimens are designated

for decens, simpsoni, duttoni, and 16 junior synonyms. A neotype specimen is desig-

nated for bitaeniorhynchus. Six new junior synonyms are recognized: alpha Séguy

(= theileri); calloti Rioux and Pech, erectus lglisch, and forridus Iglisch (= pipiens);

ethiopicus Edwards (= bitaeniorhynchus ); and mauritanicus Callot (= simpsoni). The

recognized species are arranged into three groups and 10 subgroups. The aduttoni

group and the decens and simpsoni subgroups are recognized for the first time.

'The opinions or assertions contained herein are the private views of the author and are not to be

construed as official or as reflecting the views of the Department of the Army or the Department

of Defense.

2Walter Reed Biosystematics Unit, Department of Entomology, Walter Reed Army Institute of

Research, Washington, DC 20307-5100. Present address: U.S. Army Medical Component,

Armed Forces Research Institute of Medical Sciences, APO San Francisco 96346-5000.

2 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

INTRODUCTION

The masquito fauna of southwestern Asia is very poorly known. No modern,

comprehensive treatment exists for any group in any part of the region and most of the

species descriptions and keys are widely scattered through the literature. There are a

number of major works on the mosquitoes of certain countries and specific parts of the

region (Edwards, 1921, 1926; Séguy, 1924; Kirkpatrick, 1925; Monchadskii, 1951;

Mattingly and Knight, 1956; Senevet and Andarelli, 1959; Gutsevitch et al. 1974), but all

of these are limited in scope and treatment, recognize junior synonyms, or include

misidentifications or confused species concepts. The majority of the species known to

occur in the region were originally described from elsewhere.

This work began in July 1980 as a study of Culex pipiens Linnaeus, 1758 in the

Middle East. The study was prompted by the alleged involvement of pipiens in the 1977

and 1978 epizootics of Rift Valley Fever in Egypt. At that time the concept of pipiens in

Egypt was confused by the uncertain status of mo/estus Forskal, 1775. It was unknown

whether one or more forms identifiable as pipiens may have played a role in

transmission. In January 1982 a decision was made to expand the study to include all

members of the subgenus Culex in southwestern Asia. This decision was made in

response to the need for up-to-date reliable means of identifying medically important

species in this rapidly developing part of the world. The subgenus Culex in this region

includes a number of actual and potential vector species which could not be identified

accurately with existing keys and taxonomic analyses. The decision to expand the study

was made at a time when it was becoming apparent that pipiens in Egypt was no more

than a single behaviorally and physiologically variable species. _

Southwestern Asia is defined here as the tract of land south of the USSR

between the Mediterranean Sea and the Indus River of Pakistan (Fig. 1). The area

includes all of Asian Turkey and the Arabian Peninsula. The eastern part of Pakistan

lying along the arid plain of the Indus River is excluded because its mosquito fauna is

almost entirely Oriental. Egypt is included in its entirety although only the Sinai

Peninsula is actually a part of Asia. All of the species of the subgenus Culex found in

Egypt west of the Gulf of Suez also occur in parts of southwestern Asia.

The mosquito fauna of the region consists of only 98 species representing seven

genera and 18 subgenera (Appendix A). The genus Culex is represented by 34

species assigned to seven subgenera. The subgenus Culex includes 20 species.

Most of southwestern Asia lies within the Mediterranean Subregion of the Palaearctic.

The southern part of the Arabian Peninsula belongs to the Afrotropical Region. The

majority of the species of the subgenus Culex have Palaearctic and Afrotropical

affinities. Seven species are clearly Afrotropical, seven Palaearctic, and three Oriental.

Three species are broadly distributed in both Asia and Africa.

TAXONOMY

The composition and organization of this section is unusual. It includes all

information of taxonomic relevance. Explanatory information is given followed by keys to

the genera of mosquitoes and the subgenera of Culex in the region. These are

followed by a brief taxonomic treatment of the subgenus Culex, keys to the species of

the subgenus, and the individual species treatments.

Material. The material examined during this study largely came from the collec-

Harbach: Subgenus Culex in Southwestern Asia 3

tions of the British Museum (Natural History), the London School of Hygiene and

Tropical Medicine, and the National Museum of Natural History, Smithsonian Institution.

A large proportion of the material in the National Museum of Natural History came from

collections made in Egypt and the Yemen Arab Republic by Kenneth L. Knight

(1949-51) and in Egypt by Bruce A. Harrison and myself (1983). Important additional

material was examined from the collections of the Institut Pasteur (Paris); Office de la

Recherche Scientifique et Technique Outre-Mer (Bondy, France); California Academy

of Sciences; Museum National d'Histoire Naturelle (Paris); Institut de Parasitologie,

Faculte de Medicine, Universite de Strasbourg (Strasbourg, France); Dokki Institute

(Cairo); Entomological Society of Egypt (Cairo); and the personal collections of Amon

Freidberg, Tel Aviv University, and Joel Margalit, Ben-Gurion University of the Negev,

Beer Sheva. Some 18,600 specimens (5,129 females, 3,703 males, 1,245 male

genitalia, 4,220 pupal exuviae, 2,576 larval exuviae, and 1,727 fourth-instar larvae) were

examined, including the type specimens of 50 nominal forms. A total of 14,305

specimens (3,571 females, 2,741 males, 874 male genitalia, 3,641 pupal exuviae,

2,123 larval exuviae, and 1,355 fourth-instar larvae) were examined from southwestern

Asia. The remainder (1,558 females, 962 males, 371 male genitalia, 579 pupal exuviae,

453 larval exuviae, and 372 fourth-instar larvae) were examined from neighboring areas,

mainly in Africa.

Morphology. The morphological terminology recommended by Harbach and

Knight (1980) is used throughout this work. No new terminology is introduced. Terms

for some details of the male genitalia are taken from Harbach (1986 [1987]) and Harbach

et al. (1983 [1984]). The system of lettering the setae borne on the subapical lobe of

the male gonocoxite is adopted from Edwards (1941) and Belkin (1962). Siphon indices

were calculated using the basal width of the larval siphon. Trumpet indices were

determined using the greatest width of the pupal trumpet.

Taxa. The species is the taxonomic unit of primary importance in this study. The

species are arranged into groups and subgroups largely for convenience. No

subspecies are recognized in this work. Subspecies are largely meaningless biological

concepts which have little recognition in modern culicid taxonomy. Most recent workers

have either synonymized subspecific names with specific epithets or recognized them

for distinct biological species. Users will find that | have followed Sirivanakarn (1976) in

considering Culex quinquefasciatus Say, 1823 as a species distinct from pipiens, even

though these forms apparently interbreed in some areas of southwestern Asia.

Subspecies are not included among the taxa listed in Appendix A because | believe that

descrete populations currently regarded as subspecies in southwestern Asia will

eventually prove to be differentiated species.

Keys. Keys are provided for recognizing the adult, pupal, and larval stages of

the seven genera of mosquitoes, seven subgenera of Culex, and the 20 species of the

subgenus Culex currently known to occur in southwestern Asia. The keys were

prepared especially for the limited fauna of the region (Appendix A) and some species of

the included genera of mosquitoes and subgenera of Culex from other areas will not key

out properly. The keys were prepared for specimens in reasonably good condition.

Badly rubbed or damaged specimens will be difficult to identify. The pupal keys for the

subgenera of Culex and the species of Culex (Culex ) were difficult to prepare and are

not as reliable as they appear. Identifications made in the pupal stage should be

confirmed in the adult and larval stages when possible. The keys for adults were

prepared for both males and females. Keys are not included for the male genitalia. Such

keys are normally difficult to use, especially for Culex, and more often than not genitalia

are identified by comparing dissections with published figures. Detailed descriptions

4 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

and illustrations are provided in lieu of cumbersome keys for the male genitalia.

Species treatments. The format of the species treatments is traditional. A

complete synonymy is given for each species followed by a summary of previous work,

detailed descriptions of the adult, pupal, and larval stages, a systematic discussion,

information on bionomics and distribution, and a synopsis of material examined. A cross

(+) preceding type data in the synonymies indicates that the type specimen or

specimens of the associated nominal taxon were examined. Exceptionally long

synonymies are included as appendices. References to previous taxonomic works are

exhaustive for the region. Some ecological papers and most major taxonomic analyses

for the included species in neighboring geographic areas are also included. The

references are listed in chronological order under the biomen, triomen, or other

combination used by the authors. Most of the abbreviations used in the synonymies

and literature summaries are self-evident. The letters A, M, F, P, L, and E indicate adult,

male, female, pupa, larva, and egg, respectively. An asterisk (*) after one of these letters

indicates that at least part of the life stage was illustrated in the publication cited.

Abbreviations used for type depositories follow the end of this paragraph. The species

descriptions include a brief diagnosis for each life stage. All measurements and counts

are based on at least 10 specimens from southwestern Asia, when available. Diagnostic

characters were checked in most of the positively determined specimens on hand. The

chaetotaxy of the immatures was studied using a differential interference contrast

microscope. Tables are included which summarize the complete chaetotaxy of the pupa

and fourth-instar larva of each species. The names of localities given in the material

examined sections were taken from the labels accompanying the specimens. For the

most part, no attempt was made to determine the modern spellings of the names.

Type depositories. BC - Bigot Collection (in the collection of J.E. Collin, New

Market, Great Britain); BM - British Museum (Natural History), London, Great Britain; Dl -

Dokki Insitute, Cairo, Egypt; ESE - Entomolgical Society of Egypt, Cairo, Egypt; HNM -

Magyar Nemzeti Muzeum, Budapest, Hungary; IPS - Institut de Parasitology, Faculte de

Medicine, Universite de Strasbourg, Strasbourg, France; MM - Macleay Museum of

Natural History, University of Sydney, Sydney, Australia; MNHP - Museum National

d'Histoire Naturelle, Paris, France; NMM - National Museum of Victoria, Melbourne,

Australia; NMNH - National Museum of Natural History, Smithsonian Institution,

Washington, DC, USA; NMW - Naturhistorisches Museum Vienna, Austria; TMRI - Taiwan

Provincial Malaria Research Institute, Nankang, Taipei County, Taiwan; TMS - Tomsk

Malaria Station, Tomsk, Siberia, USSR; US - School of Public Health and Tropical

Medicine, University of Sydney, Sydney, Australia; ZM - Zoologisches Museum der

Humboldt Universitat, Berlin, East Germany; ZMC - Universitetets Zoologiske Museum,

Copenhagen, Denmark.

Illustrations. The illustrations are original. Drawings are included for the male

genitalia, pupa, and larva of each species. Special features of the adult ornamentation of

some species are illustrated on a single plate. Structures of the male genitalia are

illustrated in a manner consistent with the standard arrangement and positioning

adopted in my earlier works (Harbach, 1985b, 1986 (1987); Harbach et al., 1984, 1985).

This arrangement includes the mesal aspect of the gonocoxopodite, the dorsal aspect

of the intact phallosome, lateral and mesal aspects of the fused lateral plate and

aedeagal sclerite, and dorsal aspects of the proctiger and tergum IX. The pupal and

larval stages are illustrated in the traditional manner.

Harbach: Subgenus Culex in Southwestern Asia 5

KEYS TO GENERA OF CULICIDAE

(Not treated further except for Culex)

ADULTS

Maxillary palpi of female about as long as proboscis; scutellum evenly

rounded; abdominal scales largely or completely absent . . . . Anopheles

Maxillary palpi of female much shorter than proboscis; scutellum

distinctly trilobed; abdomen completely covered withscales .......... 2

Cell R2 of wing shorter than vein R2+3; wing membrane without

cIStiNCLMIcrOtiChia (a dee cA AMG ee nies Win) Bla? eee. Uranotaenia

Spiracular setae present; wing with setae present ventrally at base

CE RUDODSEA Si Wa EO he ree PIN are rattan acta laa avd Culiseta

Tarsomere 1 of fore- and midlegs distinctly longer than tarsomeres

2-5 combined; tarsomere 4 shorter than tarsomere 5. . . Orthopodomyia

Tarsomere 1 no longer than combined length of tarsmeres 2-5,

usually shorter; tarsomere 4 longer thantarsomere5.............. 5

Postspiracular setae present; foreungues toothed. ............. Aedes

Postspiracular setae absent; ungues simple... 0... 000.0000. 20k e. 6

Hindungues large and conspicuous; pulvilliabsent ........ Coquillettidia

Hindungues small and inconspicuous; pulvillipresent........ Culex (p. 7)

PUPAE

Seta 9-IV-VII spiniform, arising at or near extreme posterolateral

corner of tergum; trumpet short, flared, meatus with broad

slit:extending nearly tobase 45... wreaks eres enliends hetiiginn ah) Anopheles

Seta 9-IV-VIl weakly developed, arising distinctly before postero-

lateral corner of tergum; trumpet otherwise, meatus at most with

narrow slit extending a short distance frompinna ................ 2

Trumpet without pinna, apex modified for piercing plant

PUG OLN D> veo sec RP RRR mel Ek) Ota alla Lili yt Coquillettidia

Inner part of paddle deeply excavated toward base, distinctly

WIGENEC MOAR ITUCID 6 POC WE Ee a var tees bee stag Ath ich Uranotaenia

Inner part of paddle little if at all excavated toward base, slightly

VORTEC VONAGE TAO Net tenia eee at alta yA ta te 4

Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Tracheoid area of trumpet well developed, extending a consider-

able distance from base; seta 2-P usually present. ........ Culex (p. 8)

Tracheoid area absent or at most weakly developed at extreme

DASESScIa 2-F acon a Make reese a PR 5

Seta 9-VIlIl strongly developed, about 0.5 length of paddle; paddle

somewhat rectangular with buttress distinctly ending beyond

middle of outer margin; seta 1-P branched .......... Orthopodomyia

Seta 9-VIII not as well developed; paddle more or less ovoid, but-

tress ending before middle of outer margin; seta 1-P simple .......... 6

Setae 8- and 9-CT in line perpendicular to mediankeel......... Culiseta

Seta 8-CT distinctly more anterior than9-CT ................. Aedes

LARVAE

Vignes cat Swany hh SH Raa Ag Soy Anopheles

SOLIDER ete ete Rt the See HEY OI Sra aie HS EE 2

Spiracular apparatus at apex of siphon modified for piercing plant

tissues, with anterior toothed plate. ............... Coquillettidia

Spiracular apparatus unmodified, comprised of 5 lobes, without

AIMEMOH OCU icgler minis room eme Thre ely Pah SS SAP 3

SIPHON WiIOUE DECISN (ei A Se ar aN, sat Pe a Dg Orthopodomyia

SPC] 0) ai 971 5 ccs 1c 11 AC aM Ne Mn Re OS AL A See 4

Head longer than wide; comb scales arising from a sclerotized

CG OUD ag Ns LIE) 2's a aie a a eae NG CN aL ae cc oo Uranotaenia

Head at least as wide as long; comb plate absent, scales arising

HOMTUNSGIE CU se NOOEINIENE: SPL, CSR OO OS WE ae 5

Siphon with at least 3 pairs of seta 1-S, usually more ......... Culex (p. 8)

Sino willl dove pale! Sola t-o SS CIP IG PP TEP SO oe 6

seta 1-9 alising Near. base of siphons... ee OPP ee Culiseta

Seta 1-S arising near last pecten spine in middle partofsiphon...... Aedes

Harbach: Subgenus Culex in Southwestern Asia Z

KEYS TO SUBGENERA OF CULEX

(Not treated further except Culex)

ADULTS

No lower mesepimeral setae; proboscis with pale ring in

Middle: i: Reyna ahs Bae ad ee rele AV IE ee ie Culex (in part, p. 13)

One or more lower mesepimeral setae; proboscis either

entirey dane 0t pale beneallt.. ia fer ee Co, ae ee 2

Three or more lower mesepimeral setae and prealar scales present;

occurring in southern part of Arabian Peninsula. ..........2.. Lutzia

One to 4 lower mesepimeral setae present; prealar scales present

or absent; normally no prealar scales if 3 or 4 lower mesepimeral

Solas Qrerareserin 6 BO, SR ee CED e Guan Slices. Sikes ea a ae 3

Proboscis shorter than forefemur; hindtarsomere 1 short, not more

ihalre-o6 lenguyor Rindge hs HG hier OG Ah Eee OE a, hes Barraudius

Proboscis longer than forefemur; hindtarsomere 1 usually as long

or only slightly shorter than hindtibia, not less than 0.85 length

Of TIMER aL ele. a es ey ay Tee ee a Rta apen ge 4

Abdominal terga with basal pale bands or basolateral pale

spots (lateral pale stripes on terga VI and VII in

antennas, Pe see OT, ST) RRO Ear ee Saha Culex (in part, p. 13)

Abdominal terga with apical pale bands, apicolateral pale

Spots oPembraly Gane firraveetr Se a Mr es Beh TY 5

Decumbent scales of vertex mainly narrow and dark, those on ocular

line broad and white; scutal scales entirely dark and very fine;

occurring in southern part of Arabian Peninsula ......... Culiciomyia

All decumbent scales of vertex narrow and pale; scutal scales

predominently pale and rather coarse (except in

NRGCAOR DIN hk Oe ag SE Pe eg eth ce Nees 6

Prealar, and usually postspiracular, scales present. ........... Maillotia

Prealar and posteniracuiar scales aise. a PA eee be ee we 7

Small yellowish-brown species; apical markings of abdominal terga

yellowish and indistinct against pale brown background;

maxillary palpi of male shorter than proboscis .......... Lasiosiphon

Larger, darker species; apical markings of abdominal terga white

and well contrasted against black or nearly black background;

maxillary palpi of male longer than proboscis ............. Neoculex

Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Ota Be WISI CUSHIONS sie We ARMING Ra 2

Seta 6-Ill-VI usually all branched (except sometimes Quttoni)............ 5

All or most of setae 1-9-CT single; seta 9-VIII well removed cephalad

from caudolateral angle of tergum, about 0.7 frombase......... Lutzia

All or most of setae 1-9-CT branched; seta 9-VIII immediately

adjacent to caudolateral angle of tergquM 6... ee ee 3

Seta 5-CT very long, longer than trumpet; paddle distinctly

NORCO Rhus taurine uot oenanihe Gi Bee esta MEE ARREARS iene te wh Culiciomyia

Seta 5-CT short; paddle evenly rounded or only slightly

POLST UD aistink ughee ius knee da iohia Loa ca CURE Dee ew kk 4

Trumpet short and broad, length less than 0.6 mm, index about

SO) eelag tt andie-F a0Sermt! Gl a aoe de a Lasiosiphon

Trumpet longer, length normally more than 0.6 mm, index more

than 3.0; setae 1-IX and 2-P usually present. ...... Culex (in part, p. 17)

Seta 5-V,VI at least nearly as long as 2 terga following, usually

BOI COMICS OBO isa eo ee eS real wre Meneame Pe he

Seta 5-V,VI 1.5 or less length of tergum following (except in

COU SHAN aie ene ut: witiiatantociniay dimseaue Alaa RU i Merle igyintate «+ fs vf

Trumpet long and slender, index more than 7.0; seta 1-P not

much stronger than 2-P, about twice aslong............. Neoculex

Trumpet not as long and slender, index less than 7.0; seta 1-P

clearly stronger than 2-P, about fourtimes aslong.......... Maillotia

Trumpet short and broad, length less than 0.6 mm, index

OU RE UC Ceili a tad gal tah i Nes yin eonluholl dues ayy eMiyhigy ws Barraudius

Trumpet variable, length and index usually more than indicated,

if length less than 0.6 mm then index more than 4.0. . Culex (in part, p. 17)

LARVAE

Lateral palatal brushes developed for predation; saddle longer than

siphon; pecten extending to apex of Siphon ................ Lutzia

Lateral palatal brushes developed for filter-feeding; saddle shorter

than siphon; pecten not extending to apex of siphon. ............. 2

Ventral brush (seta 4-X) with 4 pairs of setae; antenna shorter than

0.5 length of head; siphon with lateral pair of transversely-

alioned setae adiacenttO Pecten ke a ae oad ee Culiciomyia

Ventral brush with 5 or more (usually more) pairs of setae; antenna

longer than 0.5 length of head; siphon without lateral pair of

rococo HME Es ELL gio) ce IND ARE Bn RG a en vn AO 3

Harbach: Subgenus Culex in Southwestern Asia y

Setae 5,6-C very short and inconspicuous, about same thickness

and length as seta Ars a ay ea Pe a Lasiosiphon

Setae 5,6-C long and conspicuous, much thicker and longer than

SORES ieee er Gry eRe GUN a IRE te el ar ey a AL 4

Ventral brush with 1 or more setae borne anteriortogrid.............. 5

Ventral brush with-dllsetae-bome ongrid siti Pot RAI ee. 6

Seta 3-P nearly length of setae 1,2-P; siphon with 2 or more

(usually more) anterolateral setae on each side; seta 1-S

unpaired, occurring in single median posteriorrow. ......... Maillotia

Seta 3-P much shorter than 1,2-P, about 0.5 as long; siphon

without anterolateral setae; seta 1-S more or less paired,

occurring in 2 posterolateralrowe ik WARES aaa WOU ea Neoculex

Seta 1-S in single row, with all elements arranged in more or less

SIIOT IE UD UAL PETE RN CT Re ROT RD Barraudius

Seta 1-S in 1 or 2 rows, with 1-3 elements distinctly out of line with

the others (inserted laterally)... .......0.....2....0.2. Culex (p. 19)

10 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Genus Culex Linnaeus

Subgenus Culex Linnaeus

Culex Linnaeus, 1758: 602. Type species: pipiens Linnaeus, 1758: 602

(subsequent designation, Latreille, 1810: 442).

Heteronycha Lynch Arribalzaga, 1891a: 373; 1891b: 155. Type

species: dolosa Lynch Arribalzaga (monotypy).

Lasioconops Theobald, 1903a: 236. Type species: poicilipes

Theobald (monotypy).

Heptaphlebomyia Theobald, 1903a: 336. Type Seles simplex

Theobald (monotypy).

Pseudoheptaphlebomyia Ventrillon, 1905b:427. Type species:

madagascariensis Ventrillon (monotypy).

Trichopronomyia Theobald, 1905b: 98. Type species: annulata

Theobald (monotypy).

Aporoculex Theobald, 1907: 316. Type species: punctipes Theobald

(monotypy).

Leucomyia Theobald, 1907: 372. Type species: gelidus Theobald

(original designation).

Oculeomyia Theobald, 1907: 515. Type species: sarawaki Theobald

(monotypy).

Theobaldiomyia Brunetti, 1912: 462. Type species: Culex gelidus

Theobald, 1901a: 20 (fixed type of a replacement name).

Phalangomyia Dyar and Knab, 1914: 58. Type species: debilis Dyar

and Knab (monotypy).

Laiomyia Izquierdo, 1916: 65. Type species: Culex stigmatosoma Dyar,

1907: 123 (subsequent designation, Edwards, 1932a: 200).

Culex subgenus Transculicia Dyar, 1917(1918): 184. Type species:

eleuthera Dyar (monotypy).

Culex subgenus Cacoculex Dyar, 1918: 100. Type species: habilitator

Dyar and Knab, 1906: 212 (original designation).

The subgenus Culex is a large and diverse group with more than 200 species

which are distributed throughout the world. The concept of the subgenus defined by

Edwards (1932a, 1941), Belkin (1962), Bram (1967a), Sirivanakarn (1976), and Tanaka

et al. (1979) is followed here although this concept will probably change once the

phylogenetic relationships of the species are understood. These authors have

adequately diagnosed the subgenus and it does not need to be characterized here.

The subgenus Culex in southwestern Asia is represented by 20 nominal

species which fall into three groups and 10 subgroups. The groups, subgroups, and

species are arranged and treated in the order indicated in the chart below. The species

and subgroups, with the exception of the trifilatus subgroup, are arranged

chronologically within their respective higher categories. The species are placed in

subgroups based on presumptive phylogenetic relationships. | have made no special

effort to determine natural affinities and little is said or conjectured about evolutionary

relationships between species or groups of species. My primary aim has been to

provide a strong descriptive base for the identification and future study of each species.

The supraspecific groups are not described. They are adequately defined by the

shared characters noted for the included species.

Harbach: Subgenus Culex in Southwestern Asia

Linear Arrangement of Included Taxa

pipiens group (Edwards, 1932a: 201; excluding duttoni

Theobald)

pipiens subgroup (Sirivanakarn, 1976: 29)

1) pipiens Linnaeus, 1758 (p. 23)

2) quinquefasciatus Say, 1823 (p. 31)

trifilatus subgroup (Mattingly and Rageau, 1958: 241)

3) vagans Wiedemann, 1828 (p. 36)

4) torrentium Martini, 1925 (p. 40)

decens subgroup (New Subgroup)

5) decens Theobald, 1901c (p. 44)

6) antennatus (Becker, 1903) (p. 48)

univittatus subgroup (Sirivanakarn, 1976: 47)

7) univittatus Theobald, 1901b (p. 52)

8) perexiguus Theobald, 1903a (p. 57)

theileri subgroup (Sirivanakarn, 1976: 43)

9) theileri Theobald, 1903a (p. 61)

10) /aticinctus Edwards, 1913a (p. 67)

11) mattinglyi Knight, 1953b (p. 72)

simpsoni subgroup (New Subgroup)

12) simpsoni Theobald, 1905c (p. 75)

13) sinaiticus Kirkpatrick, 1924(1925) (p. 78)

duttoni group (New Group)

14) duttoni Theobald, 1901c (p. 82)

sitiens group (Edwards, 1932a: 201)

sitiens subgroup (sitiens series of Edwards, 1932a: 201;

including poicilipes (Theobald))

15) sitiens Wiedemann, 1828 (p. 87)

16) poicilipes (Theobald, 1903b) (p. 92)

mimeticus subgroup (mimeticus series of Edwards,

1932a: 201)

17) mimeticus Noe, 1899 (p. 97)

bitaeniorhynchus subgroup (bitaeniorhynchus series of

Edwards, 1932a: 201, excluding poicilipes

(Theobald))

18) bitaeniorhynchus Giles, 1901a (p. 101)

vishnui subgroup (Bram, 1967a)

19) tritaeniorhynchus Giles, 1901a (p. 106)

20) pseudovishnui Colless, 1957 (p. 111)

Distributional List of the Species

Too few collections have been made in most areas of southwestern Asia to plot

the distributions of the species accurately and completely. Members of the subgenus

Culex known to occur in the region are listed by country below. The list is compiled from

material examined and literature records. Species mistakenly or doubtfully recorded in

the literature are not included.

12 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

AFGHANISTAN: Culex perexiguus, pipiens, quinquefasciatus, theileri, and

vagans.

BAHRAIN: Culex quinquefasciatus.

EGYPT: Culex antennatus, laticinctus, mimeticus, perexiguus, pipiens,

poicilipes, sinaiticus, theileri, and tritaeniorhynchus.

IRAN: Culex antennatus, bitaeniorhynchus, laticinctus, mimeticus, perexiguus,

pipiens, pseudovishnui, quinquefasciatus, sinaiticus, sitiens, theileri, torrentium,

tritaeniorhynchus, and vagans.

IRAQ: Culex antennatus, laticinctus, mimeticus, perexiguus, pipiens,

pseudovishnui, quinquefasciatus, theileri, torrentium, and tritaeniorhynchus.

ISRAEL: Culex antennatus, laticinctus, mimeticus, perexiguus, pipiens,

poicilipes, sinaiticus, theileri, and tritaeniorhynchus.

JORDAN: Culex antennatus, laticinctus, mimeticus, perexiguus, pipiens,

sinaiticus, theileri, and tritaeniorhynchus.

KUWAIT: Culex quinquefasciatus.

LEBANON: Culex laticinctus, mimeticus, perexiguus, pipiens, theileri, and

tritaeniorhynchus.

OMAN: Culex laticinctus, perexiguus, quinquefasciatus, sinaiticus, sitiens, and

tritaeniorhynchus.

PAKISTAN (west of Indus River): Culex bitaeniorhynchus, mimeticus,

perexiguus, pseudovishnuli, quinquefasciatus, sitiens, theileri, tritaeniorhynchus, and

vagans.

P.D.R. YEMEN: Culex bitaeniorhynchus, decens, duttoni, laticinctus, mattinglyi,

pipiens, quinquefasciatus, sinaiticus, sitiens, theileri, tritaeniorhynchus, and univittatus.

QATAR: Culex quinquefasciatus.

SAUDI ARABIA: Culex laticinctus, mattinglyi, perexiguus, pipiens,

quinquefasciatus, sinaiticus, sitiens, theileri, and tritaeniorhynchus.

SYRIA: Culex laticinctus, mimeticus, perexiguus, pipiens, theileri, and

tritaeniorhynchus.

TURKEY: Culex antennatus, laticinctus, mimeticus, perexiguus, pipiens, theileri,

torrentium, tritaeniorhynchus, and vagans.

UNITED ARAB EMIRATES: Culex pipiens, quinquefasciatus, and sitiens.

YEMEN ARAB REPUBLIC: Culex bitaeniorhynchus, decens, l/aticinctus,

mattinglyi, pipiens, quinquefasciatus, simpsoni, sinaiticus, sitiens, theileri,

tritaeniorhynchus, and univittatus.

Zoogeographic Affinities

The Yemen Arab Republic, the P.D.R. Yemen, and the southwestern part of

Saudi Arabia belong to the Afrotropical Region. The rest of southwestern Asia and

Egypt lie within the southern Palaearctic. The known distributions of the included

species can be examined to determine their zoogeographic affinities. Culex decens,

univittatus, mattinglyi, simpsoni, and duttoni are strictly Afrotropical species. All of these

species except mattinglyi are widely distributed south of the Sahara and occur in the

Afrotropical part of the Arabian Peninsula. Culex mattinglyi is endemic to southwestern

Asia. Culex antennatus and poicilipes are also widely distributed in sub-Saharan Africa,

but they do not occur in the Afrotropical part of Arabia. Culex poicilipes extends

northward along the Nile River into Egypt and Israel. Culex antennatus is common in

Harbach: Subgenus Culex in Southwestern Asia 18

lands around the eastern end of the Mediterranean. Seven Afrotropical species are

present in the region.

Culex theileri appears to be a southern Palaearctic species which has spread

southward through the temperate highlands of eastern Africa and eastward through

India. Both /aticinctus and sinaiticus are mainly Palaearctic with limited distributions in the

Afrotropical Region. Culex perexiguus is very widely distributed across northern Africa

and southwestern Asia and extends eastward into the Oriental Region. This species is

apparently replaced by univittatus in southwestern Arabia. Culex mimeticus occurs in a

relatively narrow zone which extends from the shores of the Mediterranean, through the

Middle East, and beyond. Populations of this species in eastern Asia occur in both the

Palaearctic and Oriental regions. Culex vagans is widespread in the northeastern

Palaearctic. Its known range includes southern China and India in the Oriental Region

and northern countries of southwestern Asia. Culex torrentium is the only species of

the subgenus in southwestern Asia which is exclusively Palaearctic. Its range lies mainly

north of southwestern Asia. A total of seven Palaearctic species occur in the region.

Oriental species in southwestern Asia include sifiens, tritaeniorhynchus, and

pseudovishnui. Culex bitaeniorhynchus perhaps should be included among these

species, but its wide distribution in the Oriental Region is offset by its nearly equally

broad distribution in the Afrotropical Region. Culex tritaeniorhynchus is very widely

distributed in southwestern Asia, but the main part of its range lies within the Oriental

Region. Isolated populations occur in Egypt and it is known from certain coastal areas of

eastern and western Africa where it was probably introduced by man. Both sitiens and

pseudovishnui have limited distributions in the region. The former is a coastal species

which extends from the eastern coast of Africa to the South Pacific. Culex

pseudovishnui is an indigenous member of the Oriental fauna which has spread into the

eastern part of the region.

Culex pipiens and quinquefasciatus are common widespread sister species

whose distributions undoubtedly have been influenced by man. Their natural ranges

cannot be known for certain. Culex pipiens is very widely distributed in the Palaearctic

and eastern areas of the Afrotropical Region. This species may have evolved in Africa

and dispersed to more northern latitudes. It is replaced at lower elevations in warmer

areas by quinquefasciatus. Culex quinquefasciatus is cosmotropical.

KEYS TO SPECIES OF CULEX (CULEX)

ADULTS

+. One or more lower mesepimeral setae present (Fig. 2A) (note

presence of alveoli if setae are missing); proboscis and tarsi

WIDOLIL ale NNSA Cre OA I Se Cy! Res a, 2

No lower mesepimeral setae (note absence of alveoli); proboscis

with median pale ring; tarsi with narrow pale rings (Sitiens group)... . . 18

2(1). Tarsi with narrow pale rings (Guttoni group); midtibia with anterior

pale stripe (Fig. 2H); postspiracular scales present. .... duttoni (p. 82)

Tarsi without pale rings (pipiens group); midtibia with or without

anterior pale stripe; postspiracular scales present orabsent......... 3

Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

pipiens group

Postspiacularand prealar scales present (Fig. 2A). 2. Glib ee. 4

NO POSTS IACUIAISCdies sr O as ees oe oe S 8

All tibiae with anterior pale stripes (Fig. 2G); prealar and upper

and lower mesokatepisternal scale-patches confluent

(Fig. 2B); basal pale bands of abdominal terga produced

DOSIeHOry In mae re Deo eee Oe ee theileri (p. 61)

Fore- and midtibiae normally without anterior pale stripes

(univittatus and perexiguus usually have an anterodorsal

pale stripe on the midtibia); prealar and upper and lower

mesokatepisternal scale-patches separate; basal pale

bands of abdominal terga not produced posteriorly inmiddile........ 5

Hindtibia with anterior pale stripe (Fig. 2K); wing with short line

Ol Pale scales ar ase CCCs a i ae ee en 6

Hindtibia without anterior pale stripe; wing entirely dark-scaled .......... 7

Anterior surface of midfemur with complete narrow pale stripe

(Fig. 21); wing usually with row of scales on vein 2A ( an

unique character among mosquitoes) (Fig. 2D); occurring

in southern part of Arabian Peninsula .......... univittatus (p. 52)

Anterior surface of midfemur entirely dark-scaled or with

incomplete narrow pale stripe; wing seldom with few

scales on vein 2A; occurring throughout southwestern

PAS Ree GIy ROU A A te ie Shad perexiguus (p. 57)

Hindfemur with complete or nearly complele anterodorsal

dark stripe (Fig. 2L); females: abdominal sterna with

apical dark bands; forecoxa with some dark scales;

males: abdominal sterna mainly dark-scaled, with

basolateral pale spots; maxillary palpus normal ...... simpsonii (p. 75)

Hindfemur with anterodorsal dark stripe on distal 0.5 or less

(Fig. 2M); females: abdominal sterna usually entirely

pale-scaled; scales of forecoxa usually all pale; males:

abdominal sterna mainly pale-scaled, posterior sterna

usually with dark scales posteriorly; maxillary palpus

SPaRSehSCIOS ORE ee ais, Pe GN gai sinaiticus (p. 78)

Prealar scales Present TGs 2s ee Re ee Mea ee

PReAIARECATES AICS tat ee era sh crt eu eas i IM A oe 10

Thorax pale; wing with short line of pale scales at base of costa;

scales of forecoxa all pale; frequently with 2-4 lower

Mesepimeralisetae aaa seer Gees PR oa laticinctus (p. 67)

Thorax darker; wing entirely dark-scaled; forecoxa with some

dark scales; normally with only one lower mesepimeral

7 RR Me SOUS A SEREIRRM ENC TFA SD Ls 4 tae Sry SuPer VR ca torrentium (p. 40)

10(8).

ASO):

T2044).

13(12).

14(13).

15(14).

16(14).

Harbach: Subgenus Culex in Southwestern Asia 15

Fore- and midfemora and all tibiae with anterior pale stripes

(FIG Sa Ha Ne ee et ay aes mn eg range Ne yy vagans (p. 36)

These with anterior surfaces entirely dark-scaled ................. 11

Abdominal sterna with apical dark bands; scutal scales with

distinct reddish tint; occurring in southern part of Arabian

PEPSI ee ai Re 8 PN Cette ce FU decens (p. 44)

Abdominal sterna unbanded, dark scales confined to midline

and/or posterolateral corners when present; color of scutal

SCAICS VaNablIe eee aed Fe a Fa hg ANE rR ue ik, 12

Abdominal terga unbanded, terga VI and VII with lateral pale

stripe; proboscis of male with ventral cluster of setae at

LESSON es Is ee se i OR TR ae ee Aah Pah BA ae antennatus (p. 48)

Abdominal terga with basal pale bands; proboscis of male

without ventral cluster of setae at false joint. ee a. 13

Two to 4 lower mesepimeral setae present (note presence of

alveotit Selae are MISSING) ok re eS ee nes laticinctus (p. 67)

Only one lower mesepimeral seta present ............2..... .14

Scales of forecoxa all pale; wing with short line of pale scales

at base of costa; proboscis all dark or faintly pale beneath. ......... 15

Forecoxa with some dark scales; wing entirely dark-scaled;

proboscis usually distinctly pale beneath in middle ............. 16

Thoracic integument pale; scutal scales light yellowish brown;

scutum with short diagonal line of whitish scales along

prescutal suture; dorsal dark stripe of hindfemur beginning

beyond base and expanded over distal 0.2-0.3 of anterior

LUT (= Mele ac a as i a al i si zi i er ly dle gras: Ce laticinctus(p. 67)

Thoracic integument and scutal scales darker; scutum with

curved line of pale scales along prescutal suture and

posterior dorsocentral setae; dorsal dark stripe of

hindfemur beginning at base and expanded over

distal'0.3-0.4 OV atienior SUN ACO oe ete ee sw at mattinglyi (p. 72)

Scutal scales more or less buff-colored; basal bands of

abdominal terga nearly white, usually slightly paler than

sternal scaling; females: tergal bands slightly if at all

darker than basolateral white spots, subcosta normally

intersects costa before furcation of R2+3; males: dorsal

arms of phallosome nearly parallel in dorsal aspect

(Fig. 5), DV/D usually more than 0.6". .... . quinquefasciatus (p. 31)

*These characters are unreliable for separating pipiens and quinquefasciatus in central and

northeastern areas of the Arabian Peninsula where introgressed populations (hybrids) occur.

16 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Scutal scales golden brown with reddish tint; basal bands of

abdominal terga yellowish, usually same color as sternal

scaling; females: tergal bands distinctly darker than baso-

lateral white spots; subcosta normally intersects costa

at or beyond furcation of R2+3; males: dorsal arms of

phallosome divergent in dorsal aspect (Figs. 3, 9);

VAD UStia Ose An: Zen ay Are Ba See. ive

17(16). Females: cell R2 more than 4.0 length of vein R2+3; integument

and scales between supraala and posterior dorsocentral

setae usually noticeably darker than surrounding integument

and scales, evident as pair of ovoid spots; males: cell R2

2.4-3.6 length of vein R2+3; dorsal arm of phallosome

truncate in dorsal aspect, lateral plate without apicolateral

SPICES TODS FIG SB) yuna Puig CO NUR Ras Ie ay emer pipiens (p. 23)

Females: cell R2 less than 4.0 length of vein R2+3; integument _

and scales between supraalar and posterior dorsocentral

setae not appreciably darker than surrounding integument

and scales; males: cell R2 2.0-2.5 length of vein R2+3;

dorsal arm of phallosome pointed in dorsal view, lateral

plate with apicolateral spiculate lobe (Fig.9) ....... torrentium (p. 40)

sitiens group

18(1). | Wing with pattern of pale spots (Fig. 2E); midtibia with anterior

DAI SOS CIO CER SI EVAR OR OR OS mimeticus (p. 97)

Wing either speckled, entirely dark-scaled or with variable

amount of pale scaling along anterior veins; no anterior

Re AR a iL ee UN NE ia HM eakigblarg 6d vas wwe. a 19

19(18). Anterior surfaces of femora and tibiae with rows of small pale

spots (Fig. 2J); lower proepisternal scales present . . . poicilipes (p. 92)

Femora and tibiae without rows of pale spots; lower

ICHIGO SG yi DG WW GE oh oe ccd at gee dibvitok w e! 20

20(19). Abdominal terga with apical pale bands; wing with numerous

scattered pale scales (Fig. 2F); proboscis with pair of

dorsolateral pale spots before labella. .... bitaeniorhynchus (p. 101)

WVRIORE GGG CAE ACO oc a atin RR bh erg kw ke ee 21

21(20). Femora with numerous scattered pale scales on anterior surfaces;

scutum with indefinite mottled pattern; cell M1 long, furcation

of vein M1+2 proximal to furcation of vein R2+3 ........ sitiens (p. 87)

RV OLT 10TAGO CONIA oi ee hte NO cea SV ee eH a a ely Ua 22

*These characters are unreliable for separating pipiens and quinquefasciatus in central and

northeastern areas of the Arabian Peninsula where introgressed populations (hybrids) occur.

29101),

Harbach: Subgenus Culex in Southwestern Asia 17

Females: pale ring of proboscis extended proximally on ventral

surface”; forked scales of vertex dirty yellow to brown in

middle; cell R2 more than 3.0 length of vein R2+3; males:

proboscis with ventral cluster of 10 or more setae at false

joint; forked scales as infemale. ........ tritaeniorhynchus (p. 106)

Females: pale ring of proboscis without proximal extension on

ventral surface; forked scales of vertex white to yellow in middle;

cell R2 less than 3.0 length of vein R2+3; males: proboscis

without ventral cluster of setae (1 or 2 setae sometimes

present); forked scales asinfemale........ pseudovishnui (p. 111)

PUPAE

Median area of scutum, metanotum and terga | and II with con-

spicuous vesicles; seta 5-VI as long as terga VII and VIII

combined; Usually singigdrign2o ai iuisae a eh duttoni (p. 82)

Median area of scutum, metanotum and terga | and II without

vesicles; seta 5-VI distinctly shorter than combined length

of terga Viliand Vill, normally double or triple la. Po Vee we br 2

Seta 8-CT usually with 3 or more branches, seldom double; seta

6-III-VI also usually with 3 or more branches, sum of their bran-

ches on one side of abdomen equals 9 or more, usually more ....... 3

Set 8-CT usually double, rarely single or triple; seta 6-III-VI nor-

mally single or double, sum of their branches on one side

of abdomen Not excesning Scar Ue NN CA ed 14

pipiens group

Seta 6-1,ll short, not much longer than 7-I,Il; trumpet strongly

flared, index normally less than 4.0; tracheoid area short,

0:3 OF 1eSS-OF TGel WERTH Ey PLS ee IN Sa ers FANNY hy ag ok iy 4

Seta 6-I,Il long, at least twice as long as 7-I,II; trumpet not so

strongly flared, index normally more than 4.0; tracheoid

area long, usually more than 0.3 of trumpetlength..........002. 5

Seta 6-I single or double; seta 5-Il with 4-7 branches (usually 5);

seta 4-VIll double or triple (Fig. 2TH ee ee laticinctus (p. 67)

Seta 6-I with at least 3 branches; seta 5-II double; seta 4-VIII

single (Fig. 23) (based ona single specimen) ...... mattinglyi (p. 72)

Trumpet sharply bent at distal end of tracheoid area; seta 5-IV

at least. 1.5 lengin of tergqum V (Pig, 7) i Pe ae vagans (p. 36)

Trumpet gently curved; seta 5-IV less than 1.5 length of

RTM ITS EN tele aad ak WD hit ci eared Wei a he ali Me IMAM Tg 6

*In many specimens, the ventral extension is weak or absent in the middle and the proboscis bears

an isolated spot proximal to the ring.

10(6).

11(10).

PoC T yh).

Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Setae 6-IIl and 3-VII normally double or triple, sum of their branches

on both sides of-abdomen rarelwexceeding la xsisiceha 2... ew. . Ys

Setae 6-IIl and 3-VII normally with 3-5 branches, sum of their

branches on both sides of abdomenrarely 12orless ............ 10

Seta 1-Il with at least 25 branches, usually more; trumpet long and

slender, index usually more than 6.0 (Fig. 11); occurring in

southern part of Arabian Peninsula. ............. decens (p. 44)

Seta 1-ll with fewer than 25 branches, usually fewer than 20;

trumpet moderate, index usually lessthan6.0................. 8

seta 5-VI at least 1.5 length of tergum VII; tracheoid area usually

more than 0.36 of trumpet length; pinna short, usually less

than 0.25 of trumpet length (Fig. 9); occurring in Turkey,

CPV Arar) Sea Sioa sae Warnes: omni) chy Wc es torrentium (p. 40)

Seta 5-VI less than 1.5 length of tergum VII; tracheoid area

usually less than 0.36 of trumpet length; pinna longer,

normally more than 0.25 of trumpet length; more

Widespread GbeGieS oc ta ee ee Ue ted ee 2 ke 9

Trumpet distinctly flared, index less than 5.0; pinna long, about

0.4 or more of trumpet length; seta 8-CT usually triple*

(Fei iD) edt Sie reat. eee ae awrite Ge ti eee ies quinquefasciatus (p. 31)

Trumpet widened very little distally, index more than 5.0;

pinna shorter, usually less than 0.35 of trumpet length;

seta 8-CT with 4-7 branches* (Fig.3)............. pipiens (p. 23)

Caudolateral angle of tergum VIII bluntly rounded; trumpet

lightly to moderately tanned, length usually more than

0.58 mm; seta 4-CT usually with 3or4 branches ............... 11

Caudolateral angle of tergum VIII produced and acute, if not

produced, then trumpet heavily tanned and short, usually

less than 0.58 mm, and seta 4-CT normally double ............. 16

Seta 1-I| dendritic, usually with fewer than 12 branches;

trumpet flared, length more than 0.7 mm (Fig. 19). ..... theileri (p. 61)

Seta 1-ll fanlike, with 12 or more branches (usually more);

trumpet almost cylindrical or very little widened distally,

lerot eee tees iat.0. 7 MN eee ee dle eles ke 12

Seta 5-IV shorter than 1-IV; seta 5-V,VI about as long as

following tergum; paddle ovoid, inner part not noticeably |

narrowed toward base (Figs. 15,17) ........... univittatus (p. 52)

perexiguus (p. 57)

*These characters are unreliable in central and northeastern areas of the Arabian Peninsula where

introgressed populations (hybrids) occur.

13(12).

14(2).

15(14).

16(10).

17(16).

Harbach: Subgenus Culex in Southwestern Asia 19

Seta 5-IV at least as long as 1-IV, usually longer; seta 5-V,VI about

1.5 or more length of following tergum; paddle more or less

pear-shaped, inner part distinctly narrowed toward base ........... 13

Abdominal length usually more than 2.6 mm; paddle length

usually more than 0.8 mm; pinna of trumpet usually longer

TAPP. VO VARIA I aay esa eS ates encanta ay ilar antennatus (p. 48)

Smaller species, these measurements usually less than

Indicated (Figs; ZS27yi ts ea simpsoni (p. 75); sinaiticus (p. 78)

sitiens group

Trumpet very strongly flared, funnel-shaped; pinna long, more

than 0.36 mm; setae11-CT, 3-I-IIl and 1-VII normally single

(PIGCST PE: Gre U8 295 Oe ee Ghee Sees bitaeniorhynchus (p. 101)

Trumpet not so strongly flared; pinna shorter, less than

0.36 mm; setae 11-CT, 3-I-Ill and 1-Vil usually double... ......2.2.. 15

Seta 1-Il with 1-4 branches; setae 6-III-VI usually single;

seta 9-VIlIl with 4-9 branches (Fig. 35)... 0... ee. mimeticus (p. 97)

Seta 1-Il with 4-10 branches; setae 6-III-VI usually double;

seta S-Vil wiht S14 oranches (Fiqvaty eer ee as, sitiens (p. 87)

Paddle with conspicuous dark spot on inner part; trumpet

length more than 0.75 mm; paddle length more than

0.87 mm; seta 1-VI with 3 or 4 branches (Fig. 33)... . . poicilipes (p. 92)

Paddle without dark spot on inner part; trumpet length less

than 0.75 mm; paddle length less than 0.87 mm;

Sséta: 1-ViwitvS-2 Oramciics aie uh eeerrerae re vite ug

Seta 1-Il with more than 18 branches; seta 8-CT with 6 or 7

branches; caudolateral angle of tergum VII bluntly

MURUES TF IG Oo) ee ee eee tritaeniorhynchus (p. 106)

Seta 1-Il with fewer than 18 branches; seta 8-CT with

3-5 branches; caudolateral angle of tergum VIII

OU oc) ee era ties ns (ook at ee Ute See ep eee g pseudovishnui (p. 111)

LARVAE

Siphon strongly swollen, sides markedly convex, with subapical

dark band and short spine anteriorly near apex (Fig. 30) . . duttoni (p. 82)

CHT OUNONWIGE oi ice SA aie eR ae ee ee er RR, ne 2

All scales of comb evenly fringed at sides andapex ................. 3

Some or all scales of comb spinelike, with pointed apex

ANG IGS AL GIGSS WOKITIANY os caesar e eke wes 13

Distal pecten spines with 2-5 ventral denticles of different sizes

arising proximally; seta 1-C thin, scarcely if at all thicker than

10(7).

Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

PRSRCE SOT Seta SO a ed i ee 4

Distal pecten spines with 7 or more ventral denticles of similar

size arising along entire length; seta 1-C stouter, distinctly

WICKGF Cin Olanichee OF Selae oor em 16

pipiens group

Siphon very long and slender, index more than 8.5; seta 1-IV,V

long and single, much longer than length of 2 abdominal

SOG Sis Pe ee eM gl eg decens (p. 44)

Siphon not as long and slender, index less than 8.5; seta 1-IV,V

shorter, less than length of 2 abdominal segments, with

ONG Or FOS DTANCNES: CSOT TRO Pe a er 5

Siphon with 6-8 pairs of seta 1-S, with one pair arising laterally

and 5-7 irregular pairs arising relatively close to posterior

midline; with 2 pairs arising within pecten (Figs. 22,24)............ 6

Siphon with 3-6 pairs of seta 1-S, with 1-3 pairs arising laterally

and 2-4 pairs arising posterolaterally; sometimes with one

ANISTON) De ClOn ee a a SMe em ri

Seta 2-X with 4 or 5 branches; seta 6-VI single; ventral brush

with 13 or 14 setae; usually14 (Pig? 22). So ae laticinctus (p. 67)

Seta 2-X only double or triple; seta 6-VI double; ventral brush

with 11-138 setae usually T2:(Pig. 24s oe eae mattinglyi (p. 72)

Seta 1-S no longer than diameter of siphon at point of attachment,

usually in 5 pairs; seta 6-VI normally single; seta 5-C usually

double or triple (occasionally with 4branches) ................. 8

Seta 1-S longer than diameter of siphon at point of attachment,

usually in 4 pairs; seta 6-VI normally double; seta 5-C with

4-6 branches oye EE a Te ar Ga ee tty MI 10

Seta 1-C long and slender, length about 0.1 mm; siphon with 3

lateral pairs of seta 1-S; seta 1-M usually double or triple,

SOV SS LEI Pe ee eae antennatus (p. 48)

Seta 1-C not as long, slightly stouter; siphon with 2 lateral

pairs of seta 1-S; seta 1-M usually single, sometimes

Seta 1-S distinctly shorter than diameter of siphon at point of

attachment (Fig. 16); occurring in southern part of Arabian

NE 0s AR UR aS nV ST MR OU RR NL Ona univittatus (p. 52)

Seta 1-S about as long as diameter of siphon at point of

attachment (Fig. 18); occurring throughout southwestern

PSI AN EAGER EPPING SAAN ROR StS eR Shey. eh anliceaks grt perexiguus (p. 57)

Seta 1-Ill-V with 3-6 branches (usually 4 or 5), sum of their

branches on one side of abdomen 10 or more (usually

Harbach: Subgenus Culex in Southwestern Asia 21

more); seta 1-M normally double or triple; seta 1-X usually

COUDIO FIG. FO) ac) an) Gee aa Oa teal ia eek torrentium (p. 40)

Seta 1-IIIl-V with 1-4 branches (usually 1 or 2), sum of their

branches on one side of abdomen not exceeding 10

(usually 6 or less); seta 1-M normally single; seta 1-X

LISLIAMY SITS Fe LY RA Ae et Pe ols Veh ert A A Ts 11

11(10). Seta 1-C pigmented, usually spiculate in middle; integument of

thorax and abdomen with rows of minute vesicles; seta 13-T

as long as 12-T (Fig. 8); occurring in Pakistan, and perhaps

PAIQHAIUSTAN GIN HAR iy! Bel OE al May leatiat agian vw tome ae vagans (p. 36)

Seta 1-C not pigmented, sides smooth; integument of thorax

and abdomen without vesicles; seta 13-T distinctly

shorter than 12-T; more widespread species... ........0.0....0.. 12

12(11). Siphonal saddle index less than 3.45; seta 1-III,1V usually

SHI” CES er Ge he aia RM Ae Ade quinquefasciatus (p. 31)

Siphonal saddle index greater than 3.45; seta 1-lII,1V

USLIANY-GOUbie Rip: Apiary ane Pari ai ee pipiens (p. 23)

1312). Seta 7-I distinctly shorter than 6-1, usually double; seta 14-C

with 2 or more branches, rarely single; seta 1-C slender,

usually not thicker than branches of setae 5,6-C ............00.. 14

Seta 7-| about as long as 6-1, usually single; seta14-C single;

seta 1-C stout, usually much thicker than branches of

SOUS ler 0 a an a aa Meee hy goes RARE GPU Vir ea ER SEN Ww

14(13). Seta 5-C with 3 or 4 branches; siphon with most elements of

seta 1-S arising relatively close to posterior midline,

these with 4-11 branches; seta 14-P single (Fig. 20) . . . . theileri (p. 61)

Seta 5-C with 1 or 2 branches (usually single);

siphon with all elements of seta 1-S arising laterally

and posterolaterally, these with 1-4 branches; seta

14-F double on alleaet ONG Sidley eich ee aed ey poe es 15

15(14). Pecten borne on proximal third or less of siphon; all scales of

comb spinelike (Fig. 26); siphon index usually less than

6.0; occurring in southern part of Arabian Peninsula . . simpsoni (p. 75)

Pecten longer, extending about 0.4 of siphon length;

some anterior scales of comb evenly fringed at sides

and apex (Fig. 28); siphon index usually greater

than 6.0; more widespread species ............ sinaiticus (p. 78)

“These characters are unreliable in central and northeastern areas of the Arabian Peninsula where

introgressed populations (hybrids) occur.

17118).

18(17).

19(18).

Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

sitiens group

Seta 5-C with 3 or 4 branches; seta 1-S only slightly longer

than diameter of siphon at point of attachment; seta 1-X

with 2-4 branches (Fig. 40); siphon index greater

PATA Pe a RR ie che tritaeniorhynchus (p. 106)

Seta 5-C with 6-8 branches; seta 1-S nearly twice as

long as diameter of siphon at point of attachment;

seta 1-X single (Fig. 32); siphon index less

HS ek ee ee ran ck ee th Sitiens (p. 87)

Comb with more than 25 scales; seta 2-S long and curved;

most elements of ventral brush with 4 branches

Comb with no more than 10 scales; seta 2-S long or short,

always straight; most elements of ventral brush with

more tinale 4 branches) Sass ao ie a aN eee). sk 18

Median labral plate indistinguishably fused with dorsal

apotome; dorsomentum a straight-sided triangle with

minutely serrated margins; pecten very short, less

than0.1.Jength.ot siphon (Fig: 38)vier 6 2 bitaeniorhynchus (p. 101)

Median labral plate distinct, separated from dorsal

apotome by well-developed suture; dorsomentum

an imperfect triangle with large teeth; pecten

MUCH ONGet eeu O: SION OLSIDNON cites be cai a eens 19

Anterior margin of siphon distinctly curved in lateral view;

length of seta 1-S 2.5-3.0 diameter of siphon; pecten

spines with 4-9 basal denticles; setae 2,3-A inserted

alapex ol amenna(Fidi B4)24 dite ite take os poicilipes (p. 92)

Anterior margin of siphon more or less straight; length of

seta 1-S not more than 1.5 diameter of siphon; pecten

spines with complete ventral row of denticles; setae

2,3-A inserted subapically (Fig.42)........ pseudovishnui (p. 111)

Harbach: Subgenus Culex in Southwestern Asia 23

THE SPECIES TREATMENTS

Culex (Culex) pipiens Linnaeus

pipiens Linnaeus, 1758: 602. tNeotype male: Silvakra farm, Veberéd,

Scania, Sweden; designated by Harbach et al., 1985: 9 (NMNH).

molestus Forskal, 1775: 85. tNeotype male: Rosetta, Buhayrah Gov.,

Eygpt; designated by Harbach et al. 1984: 523 (NMNH). Synonymy

with pipiens by Harbach et al., 1984: 540.

bicolor Meigen, 1818: 7. *Type female: ? Africa (MNHP). Synonymy

with pipiens by Edwards, 1921: 345.

pallipes Macquart, 1838a: 37. Type specimen(s) (non-extant): Egypt.

Synonymy with pipiens by Edwards, 1932a: 210.

melanorhinus Giles, 1900: 342. Nomen novum for pallipes Macquart,

1838b, non Waltl, 1835: 110.

longefurcatus Becker, 1903: 68. Type female: Cairo, Egypt (ZM).

Synonymy with pipiens by Edwards, 1932a: 209.

For complete synonymy, see Appendix B.

Culex fatigans Wiedemann of Theobald, 1904: 76 (in part; Egypt); Theobald, 1905b: 92

(Egypt); Gough, 1914: 135 (Egypt).

Culex molestus Forskal of Marshall and Staley, 1937: 17-26 (Egypt, M, F, L, beh.,

physiol.); Buttiker,1981: 473 (in part; Saudi Arabia, bionomics).

Culex (Culex) fatigans Wiedemann of Abdel-Malek, 1960: 113-115, 118 (Syria, L

bionomics).

Culex (Culex) laticinctus Edwards of Séguy, 1924: 37 (in part; L* in Figs. 20-23, PI. Il,

only).

Culex (Culex) molestus Forskal of Parr, 1943: 246-251 (Syria, Lebanon, A, P, L keys,

bionomics); Knight, 1951: 354 (Mediterranean Subregion, M, F, L, E, beh.,

physiol, distr.); Senevet and Andarelli, 1954: 49 (North Africa, M, F, A key); Sen-

evet and Andarelli, 1959: 185 (North Africa, A, Mgen., P, L, keys, biol.); Abdel-

Malek, 1960: 113-121 (Syria, bionomics, distr.); Lotfi, 1970: 401 [author listed as

"(Wiedemann)"] (Iran); Lotfi, 1973: 206 (Iran); Harbach et al., 1984: 523 (Egypt,

M*,F*, P*, L*, biol., beh./physiol.).

Culex pipiens Linnaeus of Theobald, 1901b: 132 (World, M*, F, syn., bionomics, distr.);

Theobald, 1904: 76 (Egypt); Patton, 1905: 635 (P.D.R. Yemen, L habitat);

Theobald, 1905a: 28 (Egypt, syn.); Theobald, 1905b: 93 (Egypt, Israel, Jordan);

Edwards, 1913a: 49 (Israel, A); Edwards, 1913b: 53 (Africa, Syn., A, M gen.*);

Gough, 1914: 135 (Egypt); Storey, 1918(1919): 87-99 (Egypt, A, L keys, A,

bionomics); Barraud, 1920: 324 (Iraq); Barraud, 1921: 393 (Egypt, Israel, Jor-

dan, A, L, distr.); Edwards, 1922: 324 (Oriental Region, A key); Buxton, 1923:

317 (Israel, Jordan, bionomics); Theodor, 1923: 341 (Israel, Jordan, P*, key);

Kirkpatrick, 1924 (1925): 368, 371 (Egypt, A, L keys); Salem, 1938: 27 (Egypt);

Lewis, 1945: 17 (Sudan, M, F, L, distr.); Knight and Abdel Malek, 1951: 178

(Egypt, M, F, E, L, beh., physiol., bionomics); Dow, 1953: 688 (Iran, L assoc.);

Gad, 1956: 135 (Egypt, bionomic note); Gad and Darwish, 1957: 538 (Sinai);

Senevet et al., 1957a: 86 (N. Africa, L*); Hussain, 1963: 64 (Iraq); Gad and Sait,

1972: 581 (Egypt, bionomics); Margalit et al., 1973: 32 (Israel); Doyle et al., 1980:

24 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

676 (Egypt, crossings); El-Said and Kenawy, 1983a (Egypt); Zimmerman et al.,

1985: 84 (Egypt, bionomics).

Culex pipiens molestus Forskal of Lewis, 1945: 17 (Sudan, M, F, L, bionomics); Theo-

dor, 1952: 113 (Middle East, zoogeogr.); Margalit and Tahori, 1970a: 142 (Israel,

bionomics); Margalit et al., 1971: 323 (Israel, bionomics); Margalit and Tahori,

1973: 90 (Sinai, L assoc.); Zaini et al., 1983: 117 (Iraq).

Culex pipiens pipiens Linnaeus of Zaini et al., 1983: 117 (Iraq).

Culex (Culex) pipiens Linnaeus of Edwards, 1921: 345 (Palaearctic Region, A, L keys,

tax., syn., distr.); Séguy, 1924: 35 (Palaearctic Region, A, L keys, M", L", tax.,

distr., med. imp., syn., distr.); Kirkpatrick, 1925: 133 (Egypt, A, P, L keys, syn.,

M*, F, P*, L*, bionomics); Edwards, 1926: 142 (Palaearctic Region, A, L keys,

syn., A, L, distr.); Stackelberg, 1927: 166 (Palaearctic Region, M, F keys, A, M

gen.”, distr.); Martini, 1931: 374 (Palaearctic Region, A, L keys, syn., M*, F, L*,

distr.); Edwards, 1941: 314, 420, 482 (Afrotropical Region, A key, M’*, F*, P,

distr.); Senevet, 1947a: 119 (N. Africa, L*, distr., L assoc., L key); Knight, 1951:

359 (Mediterranean Subregion, M, F, L, E, beh., physiol., distr.); Knight, 1953a:

230 (Yemen Arab Republic, M, L, coll. rec., bionomics); Senevet and Andarelli,

1954: 48 (N. Africa, M, F, A key); Khattat, 1955: 166, 183 (Iraq, L*, distr., syn.,

bionomics); Lewis, 1956: 710 (Egypt, Sudan, Eritrea, L, coll. rec., bionomics);

Abdel-Malek, 1956: 102 (Sinai, bionomics, L key); Mattingly and Knight, 1956:

102 (Saudi Arabia, P.D.R. Yemen, United Arab Emirates, Yemen Arab Republic,

A, L* keys, distr., bionomics); Senevet et al., 1957b: 92 (N. Africa, L); Parrish,

1959: 266 (Turkey); Senevet and Andarelli, 1959: 167 (North Africa, A, L, P keys,

M*, F, P*, L*); Abdel-Malek, 1960: 121 (Syria, bionomics, distr.); Khalaf, 1962: 46

(Iraq, A, M*, P*, L, bionomics); Derwesh, 1965: 44 (Iraq); DuBose and Curtin,

1965: 352 (Mediterranean area, A, L keys); Abul-hab, 1968: 246 (lraq, L key);

Lotfi, 1970: 401 (Iran); Lotfi, 1973: 206 (Iran); Margalit and Tahori, 1974: 87

(Israel); El-Said and Kenawy, 1983b (Egypt, distr.); Danilov, 1985a: 71 (Afghani-

stan, A key); Danilov, 1985b: 55 (Afghanistan, L key); Harbach et al., 1985: 9

(Sweden, M*, F*, P*, L*, tax.); Harbach, 1985a: 86, 95, 105 (SW Asia [excluding

Pakistan], Egypt, distr., A, L keys).

Culex (Culex) pipiens var. molestus Forskal of Mattingly and Knight, 1956: 103 (Saudi

Arabia, A, L keys, zoogeogr.).

Culex (Culex) pipiens molestus Forskal of Monchadskii, 1951: 274 (Middle East, L key,

L*, biol.); Hopkins, 1952: 302 (Afrotropical Region, L key, L*, bionomics); Khalaf,

1962: 49 (Iraq, M, F, L, biol.); Margalit and Tahori, 1970b: 152 (Israel); Lotfi, 1976:

84 (Iran, L key, L*); Ibrahim et al., 1983: 92 (Iraq, L* key).

Culex (Culex) pipiens pipiens Linnaeus of Monchadskii, 1951: 272 (Middle East, L",

key, biol.); Hopkins, 1952: 300 (Afrotropical Region, L key, L*, bionomics);

Senevet and Andarelli, 1959: 184 (North Africa, A, M gen.*, P, L, biol.); Khalaf,

1962: 48 (Iraq, M, F, L, biol.); Lotfi, 1976: 83 (Iran, L key, L*); Ibrahim et al., 1983:

92 (Iraq, L* key). |

Culex quinquefasciatus Say of Dow, 1953: 688 (in part; northern localities in Iran, L

bionomics).

Culex (Culex) quinquefasciatus Say of Parrish, 1959: 266 (Turkey).

Adult. A medium-sized mosquito without striking features and special

ornamentation. This species closely resembles quinquefasciatus with which it has

introgressed in certain areas of the Arabian Peninsula. It can be distinguished from

quinquefasciatus by the following combination of characters: (1) scutal scales golden

Harbach: Subgenus Culex in Southwestern Asia 25

brown with reddish tint, (2) integument and scales between supraalar and posterior

dorsocentral setae noticeably darker, forming a pair of more or less conspicuous spots,

(3) cell R2 at least 4.0 length of vein R2+3, subcosta intersecting costa just beyond

furcation of R2+3; (4) basal bands of abdominal terga yellowish, usually same color as

sterna, contrasted with white basolateral spots in females.

FEMALE. Dark scaling with metallic blue reflections, reflections lost in aged

specimens. Head: Length of antennal flagellum about 2.0 mm, entirely dark,

flagellomere 1 yellowish to brownish on proximal 0.5, with some tiny, inconspicuous pale

scales mesally; pedicel yellowish to brownish, darker mesal surface with or without some

tiny pale scales. Proboscis length 1.9-2.3 mm, mean 2.1 mm; dorsal surface dark, darker

distally; ventral surface with cream-colored scales on proximal 0.7, distal 0.3 dark; labella

yellowish. Maxillary palpus entirely dark-scaled; length 0.4 mm, about 0.2 proboscis

length. Forked scales of vertex rather short, dark, some pale medially; falcate scales

narrow, pale yellow, paler laterally; lateral spatulate scales creamy white. Ocular setae

dark, reddish brown to black, antrorsely curved. Interocular space narrow, with pale

falcate scales continuous with those of vertex, and 2 large yellowish-brown setae

projecting ventrally over clypeus. Thorax: Pleural integument yellowish brown to

reddish, faded posteriorly; scutal integument dark brown. Scutal scales fine, golden

brown with reddish tint; finer on fossae and supraalar areas; integument and scales

between supraalar and posterior dorsocentral setae usually noticeably darker, evident as

a pair of ovoid spots, particularly when viewed in dorsal aspect, acrostichal and fossal

scales often as dark as ovoid spots; whitish to pale yellow scales on anterior promotory

and outer margins of supraalar and prescutellar areas. Scutal setae dark reddish

brown to nearly black. Scutellum with whitish to pale yellow falcate scales; 3-6 large

setae on each lateral lobe, 7 or 8 on median lobe. Antepronotum with upper and lower

patches of whitish to pale yellow falcate scales, scales of lower patch coarser and paler;

setae mainly dark, pale ventrally. Postpronotum with golden-brown falcate scales,

usually paler and slightly coarser posteriorly; usually with 6 setae (5-9) on posterodorsal

margin, longer posteriorly. Pleural setae golden to golden brown: 7-15 upper

proepisternal in more or less single row, 5-12 prealar, 3-7 upper mesokatepisternal, 5-13

lower mesokatepisternal, 4-12 upper mesepimeral, and 1 (rarely 2) lower mesepimeral.

Pleural spatulate scales nearly white: few below upper proepisternal setae, patches on

upper corner and lower posterior border of mesokatepisternum, anterior patch on

mesepimeron at level of upper mesokatepisternal patch, and small patch before upper

mesepimeral setae. Wing: Length 3.6-4.5 mm, mean 4.0 mm; length of cell R2/length

of vein R2+3 4.6-6.0, mean 5.3; subcosta intersects costa at or just beyond furcation of

vein R2+3; length of cell M1/length of cell R2 0.7-0.8; scales entirely dark. Dorsal scaling:

broad squame scales on costa, subcosta, R, R1, and CuA; relatively narrow squame

scales on R4+5, M1, M2, M3+4, mcu, and proximally on 1A; linear plume scales on Rs,

R2+3, R2, R3, M, M1+2, and distally on 1A; remigium with 2 distinct rows of scales and

usually 2, sometimes 3, setae distally. Ventral scaling: squame scales on costa,

subcosta, base of R1, Rs, R2+3, bases of R2 and R3, M1+2, and bases of M1 and M2;

plume scales on other veins and parts of veins except CuA before mcu and proximal 0.5

of 1A which are without scales. Halter: Yellowish, capitellum usually darker with pale

scales. Legs: Anterior surface of forecoxa mainly black-scaled, with small basal patch of

yellowish scales, anterior surface also with many long golden-brown to nearly black

ventrally curved setae, apex with 3 or 4 shorter setae on posterior margin, most proximal

seta projecting nearly perpendicular to surface, others project ventrally; midcoxa with

midiateral longitudinal row of 4 or 5 long golden to golden-brown setae, setae margined

anteriorly by longitudinal patch of white spatulate scales, anterior surface with small patch

26 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

of black scales and several ventrally projecting setae at apex; posterolateral surface of

hindcoxa with longitudinal row of 5-11 golden to golden-brown setae becoming

gradually shorter from base to apex, anterolateral surface with narrow longitudinal row of

whitish spatulate scales and about 4 short ventrally projecting setae at apex, mesal

surface with 2 dark setae at apex. Ventral surfaces of trochanters with white spatulate

scales; anteroventral surface of fore- and midtrochanters usually with some black

spatulate scales. Apices of all femora with narrow dorsal knee spot of yellowish scales;

anterior surface of forefemur black-scaled, posterior surface white-scaled; midfemur like

forefemur but black scales extend over dorsal surface toward apex; hindfemur mainly

white-scaled, with complete anterodorsal stripe of black scales gradually widening to

extend over entire anterior and posterior surfaces just before apex. Foretibia mainly

black-scaled, posteroventral surface with whitish scales; midtibia mainly black-scaled,

posteroventral surface with whitish scales, anterior and dorsal surfaces with whitish spot

at apex. Tarsi black-scaled, slightly paler ventrally, particularly tarsomere 1. Pulvilli pale.

Ungues small, simple, black. Abdomen: Terga mainly black-scaled; tergum | golden

setose, with posteromesal patch of black scales; posterior margins of terga II-VI with row

of golden setae, median setae about length of basal band of next tergum, lateral setae

longer, about 0.75 of tergum length; tergum II with basomedian spot of yellowish scales

(often nearly or entirely lost) and lateral patches of white scales; terga III-VII with basal

bands of yellowish scales and basolateral spots of white scales, spots becoming

gradually larger on succeeding posterior terga and actually cover entire lateral surfaces

of tergum VII, bands 0.15-0.35 tergum length, bands usually convex on terga Ill-V and

separated from or just touching basolateral spots, bands normally straight or slightly

concave on terga VI and VII and contiguous with basolateral spots, bands of terga III and

VIl sometimes nearly or entirely lost; tergum VIII golden setose, usually with whitish

scales, sometimes with yellowish scales, scales normally paler than those of basal bands

of preceeding terga, sometimes with some dark scales posteriorly in middle. Sterna II-VII

mainly or entirely yellowish-scaled (scales same color as basal bands of terga), often with

some median black scales, particularly in less arid areas; sternum VIII with whitish scales

on lateral margins, broad median area without scales; all sterna golden setose.

MALE. Like female except as follows. Metallic reflections of dark scaling

subdued or scales without reflections, sometimes more opalescent than metallic. Head:

Length of antennal flagellum 1.4-1.6 mm; flagellomeres 1-12 pale between whorls;

flagellomeres 13 and 14 0.6-0.7 mm, approximately 0.4 flagellum length. Proboscis

without ventral cluster of setae near false joint; false joint about 0.6 from base; ventral

pale scales at level of false joint, usually extending 0.5-0.7 from base, sometimes

covering most of ventral surface. Length of maxillary palpus 2.2-3.0 mm, mean 2.6 mm,

approximately 1.3 proboscis length, extending beyond tip of proboscis by about length

of palpomere 5; palpus mainly dark-scaled, integument between palpomeres 2 and 3

pale; lateral surface of palpomere 3 with stripe of cream-colored to white scales in middle,

stripe bordered ventrally by row of long black setae on distal 0.5 of palpomere, ventral

surface devoid of scales and pale, with ventromesal row of small antrorsely curved pale

setae; palpomere 4 with nearly complete ventral stripe of white scales; base of

palpomere 5 with small ventral patch of white scales; lateral surfaces of palpomeres 4 and

5 with long black setae that are longest at base of 4 and become gradually shorter toward

apex of 5. Thorax: Ovoid spots of scutum usually distinct. Proepisternum with 9-25

setae in more or less double row. Wing: Length 2.8-3.7 mm, mean 3.3 mm; length of

cell R2/length of vein R2+3 2.4-3.6, mean 2.8; subcosta intersects costa at or before

furcation of vein R2+3; length of cell Mi/length of cell R2 0.70-0.85. Legs: Ungues

black; anterior foreunguis larger than posterior foreunguis, both stout, anterior foreun-

Harbach: Subgenus Culex in Southwestern Asia 27

guis with small ventral tooth near midlength, posterior foreunguis with small tooth nearer —

base; anterior midunguis like that of foreleg, posterior midunguis shorter and more

slender than posterior unguis of foreleg but with tiny ventral tooth near midlength:

hindungues very small, simple. Abdomen: Terga without basolateral spots; basal bands

0.25-0.40 tergum length, usually straight, or nearly so, and produced posteriorly along

lateral scale-free areas, particularly on terga V-VII, giving bands a concave appearance:

lateral scale-free areas with long laterally-projecting golden setae. Sterna more

frequently with median dark scales; sternum VIII (dorsal in position) entirely pale or with

dusky scales posteriorly in middle. Genitalia (Fig. 3): Ninth tergal lobes small, each with

7-14 unevenly-spaced setae. Gonocoxite normal, ventrolateral setae strongly

developed, these longer and stouter than lateral setae, mesal surface with 5 rows of

small setae extending from base to level of subapical lobe; subapical lobe undivided,

setae a-f in more or less straight row with gap between c and d, seta g immediately

lateral to d-f, seta h lateral to g; a-c slightly curved, a stout and rodlike with blunt apex, b

and c each with stout base, tapering distally and with apex hooked and pointed; a-f

shorter than a-c, hooked apically, d and e slender, f laterally flattened and appearing

broad in lateral view; g foliform, longer than broad, slightly asymmetrical, apex not sharply

produced; h slender, bent distally. Gonostylus stout, curved, dorsal surface concave

with 2 small slender setae on distal 0.3; gonostylar claw short, broadest apically,

troughlike. Phallosome longer than broad with lateral plates and aedeagus of nearly

equal length; lateral plate with definite dorsal, lateral and ventral arms, dorsal arm broad,

apex nearly truncate, diverging laterally from its mate of the opposite side, appearing

Slightly sinuous in lateral view; lateral arm broad in lateral view, its posterior margin more

or less trilobed, the ventral lobe more prominent than the others and bent

ventrolaterally, base of lateral arm with thumblike basal articulatory process (articulates

with basal piece), base of this process continuous mesally with dorsal aedeagal bridge:

ventral arm narrow and sharply curved with apex directed laterally; DV/D minus0.14 to

zero, mean minus 0.09. Aedeagus subcylindrical, narrowed distally; ventral aedeagal

bridge relatively wide, joining aedeagal sclerites just beyond midlength. Proctiger

without distinctive features; paraproct with small, conical basal lateral arm, crown dark with

numerous short, spinelike spicules. Cercal sclerite elongate, irregularly shaped; 3-6

cercal setae. Tergum X rectangular, adjoining paraproct below and behind basal lateral

arm.

Pupa (Fig. 3). Placement and attributes of setae as figured, range and modal

number of branches in Table 1; diagnostic characters as in the key. Cephalothorax:

Lightly tanned, legs, scutum, metanotum and metathoracic wings darker. Setae 1,2-CT

with 3-5 branches, most often with 4; 3,4-CT usually triple (2-5); 5-CT usually with 5

branches (3-7); 6-CT usually triple (1-5); 7,9,11-CT double; 8-CT most often with 4

branches (4-7); 10-CT frequently with 8 branches (5-13); 12-CT usually triple (2-5).

Trumpet: Moderately tanned, subcylindrical, gradually widened distally; index 4.9-6.8,

mean 5.4; tracheoid area darker, extending about 0.33 from base; pinna oblique,

about length of tracheoid area. Abdomen: Lightly tanned, terga I-VI darker in middle:

length 2.3-3.7 mm, mean 3.1 mm. Seta 6-l,Il normally single; 7-1,1I usually double; 1-II

multiple (10-24); 1-IIl-V frequently with at least 6 branches, 1-IIl,IV most often with 7

branches (5-10), 1-V most often with 6 branches (4-8); 2-II,VIl lateral to seta 1, 1-III-VI

mesal to 1; 5-IV often with 4 branches, rarely double (2-6); 5-V-VII almost always double:

6-III,IV most often triple, 6-V, VI most often with 4 branches. Genital lobe: Lightly tanned;

length about 0.20 mm in female, 0.34-0.39 mmin male. Paddle: Lightly tanned, midrib

and buttress darker; outer margin without distinct spicules; midrib distinct except at

apex; length 0.80-1.30 mm, mean 0.93 mm, width 0.62-0.78 mm, mean 0.69 mm, index

28 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

1.2-1.6, mean 1.4.

Larva (Fig. 4). Character and positions of setae as figured, range and modal

number of branches in Table 21; diagnostic characters as inthe key. Head: Wider than

long; length 0.70-1.04 mm, mean 0.87 mm; width 1.00-1.389 mm, mean 1.25 mm; mainly

lightly tanned, labiogula and posterior portions of lateralia and dorsal apotome

moderately tanned, dorsal apotome often with variably developed, moderately tanned

spots as follows: crescentic spot just anterior to each seta 8-C, small median spot

immediately posterior to bases of seta 5-C, median transverse oval spot midway

between bases of seta 5-C and posterior margin of head, latter with small spot on either

side laterally. Median labral plate narrow but distinct, anterior margin slightly emarginate

between insertions of seta 1-C. Labiogula narrower anteriorly than posteriorly, length

about same as posterior width; hypostomal suture complete, extended posterolaterally

from posterior tentorial pit to near collar. Collar moderately developed along lateralia,

heavily tanned. Dorsomentum most often with 10 teeth (8-14) on either side of median

tooth. Seta 1-C slender, tapered distally, slightly bent mesad, not tanned; 2-C normally

absent; 3-C distinct; 4-C single, rather long; 5-C frequently with 5 branches (4-7); 6-C

most often with 4 branches (3-6); 7-C resembles 5,6-C, most often with 10 branches

(8-13); 8-C usually double (2-4); 9-C most often with 5 branches (3-7); 10-C usually

double (2,3); 11,12,13-C double or triple, 13-C infrequently with 4 branches; 14-C

normally single, rarely double. Antenna: Length 0.38-0.55 mm, mean 0.45 mm, about

0.5 length of head; moderately tanned, mesal surface with dark spot at base; seta 1-A

0.7 from base; part proximal to seta 1-A with strongly developed aciculae mainly on

dorsal and lateral surfaces, distal part slender and smooth except for few short aciculae

laterally near seta 1-A. Seta 1-A with about 22 branches (11-27). Thorax: Integument

hyaline, tubercles of all large setae moderately tanned; setae 1-3-P and 9-12-P,M,T on

common tubercles. Setae 1-3-P single, nearly of equal length; 4,7,8-P normally double,

occasionally triple; 11-P most often with 5 branches (3-8). Seta 1-M single, about 0.5

length of 3-M; 3-M single; 4-M usually double, occasionally single. Seta 1-T short, 0.5 or

less length of 2-T, frequently triple (2-5); 2-T usually single or double, infrequently triple.

Abdomen: Integument hyaline, tubercles of setae 7-I, 6-I-VI and 1,3-VIll moderately

tanned, tubercle of setae 2,3-X heavily tanned. Seta 3-1,VIl usually single, occasionally

double; seta 6-I-VI long, 6-I,11 most often triple, sometimes with 4 branches, 6-II rarely

double, 6-III-VI double, 6-III rarely triple; 7-1 resembles 6-l, almost always double (1-3);

1-IIl-VI usually double, one branch longer than the other. Segment Vill: Comb with

33-58 scales, mean 45; scales short, evenly fringed on sides and apex, arranged in 4

irregular rows. Seta 1-VIIIl most often with 5 branches (4-9); 3-VIll frequently with 8

branches (6-9); 5-VIIl commonly with 4 branches (3-5). Siphon: Index 3.00-5.80, mean

4.58; subcylindrical, broadest at base, slightly sigmoid in lateral view; moderately tanned,

darker at base; acus attached, longer on posterior side of attachment. Pecten on basal

0.3 of siphon, with 10-18 spines, mode 14; spines increasing in size from base of

siphon, larger spines with 3 or 4 long basal denticles. Seta 1-S usually in 4 pairs (a fifth

seta sometimes occurs on one or both sides of siphon),1a-S very near most distal

pecten spine, 1c-S distinctly out of line with the others. Segment X: Saddle complete;

moderately tanned, darker dorsally; posterodorsal area with minute spicules; length

0.31-0.45 mm, mean 0.36 mm, siphon/saddle index 3.48-4.63, mean 4.08. Seta 1-X

usually single (1-3); 2-X usually double, infrequently triple; 4-X almost always in 6 pairs, all

setae borne on grid, setae increasing in length posteriad, most posterior seta about 7.5

of most anterior. Anal papillae elongate, subacutely tapered, length variable.

Systematics. Culex pipiens is a highly plastic species which is widely distri-

buted in the Holarctic Region and cooler parts of Africa. It occurs throughout Egypt and

Harbach: Subgenus Culex in Southwestern Asia 29

generally north of a line transecting Saudi Arabia between Jeddah in the west and Al

Hasa Oasis in the east. It occurs north of about 30° N. in countries east of the Persian

Gulf. Some isolated populations occur in the southwestern mountains of the Arabian

Peninsula.

This species exhibits considerable variation in all stages and it cannot always be

distinguished from quinquefasciatus and torrentium with certainty. In the adult stage,

pipiens is reliably differentiated from these species by the character of the male genitalia.

The immatures are usually separable by the characters given in the keys. Additional

characters for separating the larvae of pipiens and torrentium are discussed by Harbach

et al. (1985). The separation of all stages of pipiens and quinquesfasciatus is

complicated by the fact that these species introgress in certain areas of Saudi Arabia and

lraq (see discussion under quinquefasciatus ).

A relatively small, dark variant of pipiens occurs in southwestern Asia and Egypt

which might be confused with antennatus. This variant consists chiefly of females in

which the basal pale bands of the abdominal terga are reduced or absent. The

occurrence of similar specimens were reported previously by Edwards (1921) and

Harbach et al. (1985). Unbanded specimens also occur in Africa (Edwards, 1941). A

sporadic larval variant in which the head and siphon are darker than usual is sometimes

found in the region. Some dark larvae develop into dark adults (Harbach et al., 1984). A

single female from northern Sinai deserves special mention. This specimen (collected

by E.E. Austen at El Arish and deposited in the BM) is unusually pale, the wing has a well

developed patch of pale scales at the base of the costa, the pale bands of the abdominal

terga are white and convex as in quinquefasciatus, and the scutal scales are somewhat

coarser and paler than usual.

There is tremendous variation in the length of the siphon in the larval stage.

Specimens from polluted water tend to have a shorter siphon than specimens from

cleaner water, but there is a considerable range in siphon length in almost every

collection.

Despite the extensive amount of morphological variation which occurs in all life

stages of this species, there is no indication of subspecific or racial differences in the

various geographically separated populations examined. The only exception to this

might be those populations which occur at higher elevations in the southwestern

mountains of the Arabian Peninsula and the mountainous countries of eastern Africa.

Insufficient material was available for a critical examination of these populations, but they

do not appear to introgress with populations of quinquefasciatus, indicating that they

may not be as closely related to this species as populations occurring at more northern

latitudes. Introgression occurs where populations of pipiens and quinquefasciatus

overlap in the lower Tigris-Euphrates Valley, the coastal plains east of the Persian Gulf,

and a narrow zone extending across the Arabian Peninsula from the Gulf of Bahrain to

the Red Sea coast in the vicinity of Jeddah. The parental epiphenotypes are distinct

outside the zone of introgression, indicating that species-specific traits are being

selected outside the zone. This is evidence for independent species cohesion. It

supports the taxonomic conclusion that pipiens and quinquefasciatus are distinct

species. The genetic data provided by Urbanelli and Bullini (1985) accords this

conclusion. These authors found that allopatric populations exhibit an average Nei's

genetic distance of 0.17. Values of similar magnitude have been found among some

members of the Aedes (Ochlerotatus) caspius (Pallas, 1771), Anopheles (Cellia)

gambiae Giles, 1902, and Anopheles (Anopheles) maculipennis Meigen, 1818 species

complexes (Bullini and Coluzzi, 1982).

Bionomics. Culex pipiens is the most common species of the subgenus Culex

30 Contrib. Amer. Ent..Inst., vol. 24, no. 1, 1988

in southwestern Asia. It utilizes numerous and variable breeding places ranging from

highly polluted cesspits to clear water pools and containers. The species usually breeds

in stagnant water in either shaded or unshaded situations. The water may be fresh or

slightly brackish. Populations reach their highest densities in areas occupied by man. It

is not uncommon to find larvae in underground habitats such as drains, wells, and septic

tanks.

Culex pipiens feeds on a variety of warm-blooded vertebrates from birds to man.

Specific host feeding is influenced by host density and availability. In southwestern

Asia, the species feeds predominantly on mammals. Females feed at night and readily

attack man both indoors and outdoors.

Culex pipiens is an important vector of human pathogens nearly everywhere it

occurs. It has been found naturally infected with Sindbis and West Nile viruses in Israel

(Samina et al., 1986) and West Nile and Rift Valley Fever viruses in Egypt (Taylor et al.,

1953; Taylor et al., 1955; Tahori et al., 1955; Meegan et al., 1980; Meegan, 1981; Turell

et al., 1982). The importance of pipiens as a primary vector of periodic Bancroftian

filariasis in Egypt is well known (see Southgate, 1979 for a thorough review). Culex

pipiens is a very serious pest in southwestern Asia because of its close association with

man.

Distribution. Culex pipiens inhabits the temperate regions of Europe, Asia,

Africa, Australia, and North and South America.

Material examined. 7,994 specimens. A total of 6,384 specimens (1,386

females, 1,059 males, 205 male genitalia, 1,889 pupal exuviae, 1,129 larval exuviae,

and 716 fourth-instar larvae) were examined from southwestern Asia — AFGANISTAN:

(Bolla Quichi, Garwargin, He'ral, Kataghan); EGYPT (including the neotype series of

molestus ): Aswan (Abu Simbil, Aswan, Dar el Salam, El Aqaba el Saghira, Ezbet el

Silsila, Khour Abu Subeira, Kom Ombo, Nag el Ashbab, Nag el Idwa el Bahari, Nag el

Ritag, Nag el Shalabab, Nag Bami' el Gezira, Nag’ Tingar, Sahara City), Buhayrah

(Alexandria, El Tarh, Kafr el Dauwar, Rosetta), Eastern Deserts (Farouk Field), E/ Fayyum

(Abhit el Haggar, El Nazla, Tubhar), E/ Giza (Abu Rauwash, Barnasht, Dokki, El

Badrshein, El Rahawi, Ezbet Bosna Sharawy, Imbaba, Kafr Tohormos, Mena Road,

Moneeb, Nazlet el Ashtar, Nazlet el Saman, Saqgara, Talbia, Tanash, Tirsa), E/ Isma’ iliya

(Abu Khalifa, Isam' iliya, Nifisha), E/ Qahira (Abbassia, Cairo, Ezbet Said), E/ Qalyubiya

(Bahtim, El Ga'afara, El Qash-Shish, El Salmaniya, Hashim, Kafr Abo-Seer, Kaha,

Khanka, Qalyubiya, Sindiwa), E/ Suweis (Kantara, Suez), Luxor (Luxor), Marsa Matruh

(Siwa Oasis — Abul el Leef, Aghurmi, Ain Bundi, Masouse), Port Said (Bur Fu'ad, El

Raswa, Port Said), Red Sea (Bernece, Ghubbetel Bos, Marsa Alaam, Quseir, Wadi

Qiseib, St. Anonious Monastery), Sinai Peninsula (Dahab, El Arish, Wadi Feiran),

unknown localities; IRAN: (Afzulabad, Alchangi, Amirabad, Ardebil, Behbehan,

Bisotun, Bushehr, Cham Asbi, Enzeli, Ghasem Gheshlari, Kamroud, Kermanshah,

Khorramshuhr, Khorroniabad, Kivi, Natchi Bahran, Rasht, Sari, Shemiran, Teheran,

Zarjub); IRAQ: (Al Basrah); ISRAEL: (Acre, Beer Sheva, Beit Gubrin, Ben Sheman,

Coastal plains, Deir el Belah, Dhahiriya, Ein Gedi, Ein Hajla, Ein Meshoshim, El Athroun,

En Agev, Gennesaret, Golan Nafech, Hadera, Haifa, Jaffa, Jericho, Jerisheh, Jerusalem,

Kafr Shammai, Kalia, Kindana, Kishon Marsh, Lake Huleh, Lake Tiberias, Ludd, Nahr

Namin, Nazareth, Nulata, Ramat Razi'el, Revivim, Sarona, Shafat, Tell el Nagilla, Tul

Keram, Wadi Ghuzze, Wadi Latron, Wadi Shikma, Wassat, Yazur, unknown localities);

JORDAN: (Amman, Az Zarqa); LEBANON: (Amioum, unknown localities); P.D.R.

YEMEN: (Seiyum); SAUDI ARABIA: (Al Hasa Oasis, Al Hufuf, Al Kharj, Al Qatif, Jeddah);

TURKEY: (Istanbul, Izmir, Konya, Sinop, Tatvan); YEMEN ARAB REPUBLIC: (Hada,

Hamman ‘Ali, San'a, Ta'izz). An additional 1,610 specimens (599 females, 354 males, 45

Harbach: Subgenus Culex in Southwestern Asia SI

male genitalia, 344 pupal exuviae, 237 larval exuviae, and 31 fourth-instar larvae) were

examined from ALGERIA, ETHIOPIA, KENYA, LIBYA, MAURITANIA, MOROCCO,

SOUTH AFRICA, SUDAN, SWEDEN (including the neotype series), TANZANIA,

TUNISIA, and ZIMBABWE.

Culex (Culex) quinquefasciatus Say

quinquefasciatus Say, 1823: 10. tNeotype male: New Orleans, Loui-

siana, USA; designated by Sirivanakarn and White, 1978: 362

(NMNH).

fatigans Wiedemann, 1828: 10. Lectotype female: East Indies

[Indonesia]; designated by Belkin, 1968a: 68 (NMW). Synonymy

with quinquefasciatus by Stone, 1956(1957): 342.

pallidocephala Theobald, 1904: 73. Holotype female: Sennar, Blue

Nile, Sudan (BM). Synonymy with fatigans by Edwards, 1941: 316.

cartroni Ventrillon, 1905b: 429. +Lectotype male, hereby designated,

bearing following data: "6 // MUSEUM PARIS / MADAGASCAR /

VENTRILLON 1904 // Culex / cartroni ventr. / type"; genitalia on

acetate strip (MNHP). Synonymy with fatigans by Edwards, 1932a:

208.

Pygmaeus Neveu-Lemaire, 1906: 256. Syntypes (non-extant): Imi,

Ehtiopia. Synonymy with fatigans by Edwards, 1932a: 208.

zeltneri Neveu-Lemaire, 1906: 251. Syntypes (non-extant): Harar,

Ethiopia. Synonymy with quinquefasciatus by White,1975: 322.

For complete synonymy, see Appendix C.

Culex fatigans Wiedemann of Theobald, 1901b: 151 (World, A key, syn., M*,F*, distr.);

Theobald, 1904: 76 (in part; Sudan); Patton, 1905: 635 (P.D.R. Yemen, L

habitat); Giles, 1906: 131 (Bahrain); Edwards, 1913b: 55 (Africa, syn., M’*, F,

distr.); Barraud, 1920: 324 (Iraq); Edwards, 1922: 279 (Oriental Region, A key);

Barraud, 1924b: 1264 (India, syn., M*, F, distr.); Barraud, 1924c: 430 (India, ys

Lewis, 1945: 19 (Sudan, M, F, L, distr.); Leeson and Theodor, 1948: 228

(Socotra, L habitat; A, L keys); Hussain, 1963: 64 (Iraq).

Culex (Culex) fatigans Wiedemann of Edwards, 1921: 345 (Iran, Iraq, A, L keys, tax.,

distr.); Séguy, 1924: 35 (Iran, Iraq, A, L keys, A, M gen.*, tax., distr., med. imp.,

syn.); Edwards, 1926: 141 (Iran, Iraq, A, L keys, syn., A, distr.); Stackelberg,

1927: 165 (Iran, Iraq, M, F keys, A, M gen.*, distr.); Martini, 1931: 364 (Palaearctic

Region, A, L keys, M’*, F, L*, distr.); Barraud, 1934: 420 (India, A, L keys, M’*, F,

L", syn., L habitat, distr.); Edwards, 1941: 316, 421, 482 (Afrotropical Region, A

key, M", F*, P, syn., distr., bionomics); Lewis, 1943a: 282 (Eritrea, Lay

Monchadskii, 1951: 261 (Middle East, L*, biol., distr.); Hopkins, 1952: 304

(Afrotropical Region, L*, bionomics); Senevet and Andarelli, 1959: 187 (North

Africa, A, P, L keys, A, M gen.*, P*, L*, biol.); Derwesh, 1965: 44 (Iraq); Abul-hab,

1968: 249 (Iraq, L key); Lotfi, 1970: 401 (Iran); Lotfi, 1973: 206 (Iran).

Culex molestus Forskal of Buttiker, 1981: 473 (in part; Saudi Arabia, bionomics).

Culex pipiens fatigans Wiedemann of Zaini et al., 1983: 117 (Iraq, bionomics).

Culex (Culex) pipiens fatigans Wiedemann of Lewis, 1956: 715 (Sudan, Eritrea (?), coll.

rec.); Mattingly and Knight, 1956: 103 (Arabian Peninsula, A, L keys, distr.,

zoogeogr., bionomics); Khalaf, 1962: 50 (Iraq, M*, F, L, A biol., L habitat): Ward,

32 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

1972: 96 (Afghanistan); Lotfi, 1976: 84 (Iran, L key, L*); Ibrahim et al., 1983: 92

(Iraq, L* key).

Culex quinquefasciatus Say of Dow, 1953: 688 (in part; southern localities in Iran, L

bionomics).

Culex (Culex) quinquefasciatus Say of Knight, 1953a: 230 (Yemen Arab Republic, M

gen., L, coll. rec., L habitat); Sirivanakarn, 1976: 30 (Oriental Region, A, P, L

keys, M*, F*, P*, L*, tax., distr., bionomics, med. imp.); Danilov, 1985a: 71

(Afghanistan, A key); Danilov, 1985b: 55 (Afghanistan, L key); Harbach, 1985a:

86, 95, 105 (SW Asia, distr., A, L keys).

Adult. A medium-sized species closely resembling Culex pipiens with which it

has introgressed in northeastern and central areas of the Arabian Peninsula. This

species is distinguishable from pipiens by the following combination of characters: (1)

scutal scales more or less buff-colored, relatively long and sparse, giving scutum a rather

shaggy appearance, (2) integument and scales between supraalar and posterior

dorsocentral setae usually not markedly shorter or noticeably darker than surrounding

scales, (3) length of cell R2 less than 4.0 length of vein R2+3, subcosta intersects costa

before furcation of vein R2+3; (4) basal bands of abdominal terga whitish to

cream-colored, slightly if at all darker than basolateral white spots, usually slightly paler

than sterna.

FEMALE. Dark scaling with metallic blue reflections, reflections lost in aged

museum specimens. Head: Antennal length 1.6-2.0 mm, mean 1.8 mm; flagellum dark,

base of flagellomere 1 and most of pedicel whitish to yellowish orange; mesal surface of

pedicel dark with some inconspicuous scales and/or setae. Proboscis length 1.7-2.1

mm, mean 1.9 mm; mainly black-scaled, ventral surface with cream-colored scales on

proximal 0.7, pale scaling most pronounced broadly in middle. Maxillary palpus short,

less than 0.2 proboscis length, entirely black-scaled. Falcate scales of vertex narrow,

whitish to pale yellow, slightly paler along margin of eye; forked scales pale medially, dark

laterally; lateral spatulate scales narrow, white; ocular setae golden brown. Interocular

space narrow, with some ventrally projecting falcate scales coarser and slightly paler than

those of vertex, with 2 long golden to golden-brown setae projecting downward over

clypeus. Thorax: Integument yellowish brown, scutum slightly darker than pleura.

Scutal scales slightly coarser and less dense than in pipiens, more or less buff-colored

(at low magnification), slightly paler on margins and prescutellar area; integument and

scales between supraalar and posterior dorsocentral setae usually not distinctly shorter

or darker than surrounding integument and scales. Each lobe of scutellum with falcate

scales concolorous with prescutellar scales; lateral lobes each with 5 or 6 large setae and

several smaller setae, large setae in 2 rows with 2 setae in dorsal row and 3 or 4 in ventral

row; middle lobe with 6 or 7 large setae in single row and several scattered small setae.

Antepronotum with whitish to pale yellow falcate scales confined mainly to dorsal and

ventral patches, dorsal patch coarser; setae concolorous with scutal setae, paler

ventrally. Postpronotum with pale yellow to golden-yellow falcate scales, scales paler

and coarser posteriorly; with curved posterodorsal row of 7 or 8 setae, setae more or less

same color as scutal setae, posterior 2 or 3 setae paler. Pleural setae pale yellow to

golden yellow, numbers follow: usually 6 upper proepisternal, 8-10 prealar, 5 or 6 upper

mesokatepisternal, 7-9 lower mesokatepisternal, usually about 7 upper mesepimeral

and 1 lower mesepimeral. Proepisternum with small number of elongate white spatulate

scales below upper proepisternal setae, usually few dark scales mesal to setae.

Mesopleuron with the usual patches of white spatulate scales in upper and lower

positions on katepisternum and anterior and upper positions on anepimeron. Wing:

Harbach: Subgenus Culex in Southwestern Asia 33

Length 3.1-3.8 mm, mean 3.4 mm; length of cell R2 2.8-3.3 length of vein R2+3;

subcosta usually intersects costa before furcation of vein R2+3; length of cell M1 0.8

length of cell R2; entirely dark-scaled. Dorsal scaling: broad squame scales on costa and

subcosta; narrow spatulate to near fusiform (decumbent) scales on R, R1, R3+4, distally

on M1, M2, mcu, M4+5, CuA, and all but distally on 1A; linear plume scales on Rs, R2+3,

R2, R3, M, M1+2, proximally on M1, and distally on 1A; remigium with 2 rows of scales and

2 or 3 setae distally. Ventral scaling: broad squame scales on costa, Rs, R2+3, M1,

M1+2, and proximally on M1; linear plume scales on R, R1, R2, R3, R4+5, distally on M1,

M2, M2+3, mcu, CuA distad of mcu, and about distal 0.3 of 1A; CuA proximal to mcu and

about proximal 0.7 of 1A without scales. Halter: Pedicel and scabellum pale; capitellum

dark with pale scales. Legs: Forecoxa with patch of whitish scales at base continuous

with predominantly black scales of anterior surface, variable number of whitish scales

usually near middle of anterior surface, ventrally curved anterior setae golden brown,

usually with 2 more or less prominent laterally projecting and several less conspicuous

ventrally projecting setae apically on posterolateral margin; midcoxa with narrow

longitudinal patch of white spatulate scales on anterior side of longitudinal row of usually

4 large golden-brown and several smaller paler setae on lateral midline, pale scales

continuous with small apical patch of black scales, number of small ventrally projecting

pale setae anterolaterally at apex; hindcoxa with somewhat inconspicuous narrow

longitudinal patch of whitish scales on anterolateral margin, with 6-8 golden

posterolateral setae and about 4 short golden setae at apical end of scale patch.

Trochanters with white scales on ventral surface extended onto anterior and posterior

surfaces; fore- and midtrochanters often with some black scales anteromesally. Femora

with inconspicuous narrow yellowish knee spots; fore- and midfemora black-scaled

anteriorly, white-scaled posteriorly, black scaling of midfemur gradually expanded over

dorsal and onto posterior surface toward apex; hindfemur white-scaled with anterodorsal

stripe of black scales gradually expanded distally over anterior and posterior surfaces,

sometimes nearly forming complete subapical ring. Tibiae mainly black-scaled, foretibia

with whitish scales ventrally; midtibia with whitish scales posteroventrally; hindtibia like

foretibia with somewhat indistinct apical pale spot mainly on dorsal and anterior surfaces.

Tarsi dark, slightly paler ventrally; tarsomeres 1-4 with hint of pale scaling dorsally at apex.

Pulvilli pale. Ungues small, simple. Abdomen: Tergum | golden setose, with median

posterior patch of black, white or mixture of black and white scales; terga II-VII with long

golden setae on lateral and posterior margins, setae short medially on posterior margin;

tergum II with median basal triangular patch of whitish or very pale yellow scales and

lateral patches of white scales; terga III-VIl with nearly white convex basal bands and

white basolateral spots, bands about 0.4 tergum length, spots becoming larger on

succeeding posterior terga, bands not touching or narrowly continuous with spots on

terga IIl-V, bands continuous with spots on terga VI and VII, tergum VII with some pale

scales on posterior border; tergum VIII entirely pale-scaled. Sterna II-VII nearly white to

yellowish, usually more yellow (slightly darker) than basal bands of terga, often with some

median black scales which sometimes form rather large patch not reaching posterior

border, occasionally with few black scales on posterolateral corners.

MALE. Like female except as follows. Head: Proboscis without ventral cluster of

setae near false joint; false joint about 0.6 from base; pale scaling mainly in middle

ventrally, forming indistinct more or less complete narrow ring at false joint. Length of

maxillary palpus about 2.0-2.6 mm, mean 2.4 mm, extending beyond tip of proboscis by

length of palpomere 5 (palpus of some males from Djibouti only slightly longer than

proboscis or exceed its length by no more than 0.5 length of palpomere 5); integument

pale across joint between palpomeres 2 and 3; middle of palpomere 3 with elongate

34 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

patch of cream-colored scales on dorsal and lateral surfaces, palpomere with about 14

long ventrolateral setae on distal 0.33 and longitudinal row of short antrorsely curved

setae on ventromesal margin; palpomere 4 with complete ventral line of white scales,

distal scales usually darker with bluish hue; palpomere 5 with small ventral patch of white

scales at base. Thorax: Scutal scales slightly longer and narrower, of more uniform size

than in female. Propleuron with 2 irregular ventromesally oriented rows of upper

proepisternal setae. Wing: Length 2.6-3.2 mm, mean 3.0 mm; length of cell R2 1.8-2.2

length of vein R2+3; length of cell M1 0.75-0.85 length of cell R2. Legs: Anterior unguis

longer than posterior unguis on fore- and midlegs, each with ventral tooth near middle.

Abdomen: Terga without basolateral spots, basal bands produced posteriorly on lateral

margins, particularly on terga VI and VII; tergum VIII (ventral in position) largely or entirely

pale-scaled. Sterna more frequently with median black scales than in female; sternum

VIII (dorsal in position) variable, often like tergum VII, sometimes largely pale-scaled.

Genitalia (Fig. 5): Form as figured; essentially as in pipiens, differing chiefly in details of

the phallosome. Ninth tergal lobe with 4-14 setae (mode 8). Seta f of subapical lobe of

gonocoxite sometimes duplicated, distinctly hooked. Lateral plate of phallosome

conspicuously broader at base; dorsal arm not as broad, apex pointed, nearly parallel

with its mate of the opposite side; dorsal and ventral lobes of lateral arm more prominent;

ventral arm broad and long; DV/D 0.56-1.89, mean 1.03. Paraproct with short, slender

basal lateral arm.

Pupa (Fig. 5). Character and placement of setae as figured, range and modal

number of branches in Table 2; very similar to pipiens, but usually separable by the

character of the trumpet and seta 8-CT. Cephalothorax: Seta 5-CT usually with 4

branches (2-4); 6-CT often double (2-4); 8-CT usually triple (2,3). Trumpet: Relatively

short and flared apically; index 2.9-4.8, mean 4.2; pinna oblique, long, length 0.4 or

more of trumpet length. Abdomen: Length 3.2-4.0 mm, mean 3.6 mm; seta 1-Ill-V

frequently with 4 or 5 branches, 1-IIl most often with 5 branches (4-7), 1-IV,V most often

with 4 branches (3-6); 5-IV usually double or triple, sometimes with 4 branches; 6-III,IV

double or triple, more often double. Paddle: Length 0.98-1.16 mm, mean 1.04 mm;

width 0.74-0.87 mm, mean 0.82 mm; index 1.1-1.4, mean 1.3.

Larva (Fig. 6). Character and positions of setae as figured, range and modal

number of branches in Table 22; closely resembling pipiens from which it cannot always

be separated, the most reliable diagnostic features from pipiens include the

siphon/saddle index and character of seta 1-Ill,IV. Head: Length 0.74-0.88 mm, mean

0.83 mm; width 1.13-1.30 mm, mean 1.21 mm. Dorsomentum most often with 12 teeth

(10-13) on either side of median tooth. Seta 2-C normally present, very small; 6-C most

often with 5 branches (3-5); 7-C most often with 8 branches (7-10); 8-C often triple (2-4);

10-C most often triple (1-3). Antenna: Length 0.43-0.50 mm, mean 0.45 mm. Thorax:

Seta 1-T frequently double (2-5). Abdomen: Seta 1-lll,IV normally single, seldom

double (more seldom double on both segments). Siphon: Generally shorter and

stouter than in pipiens, index 2.88-4.64, mean 3.70. Pecten with 5-17 spines, mode

10. Seta 1-S usually in 4 pairs, occasionally in 3 pairs, seldom with a fifth seta on one

side; number of branches variable, tending to have a greater number than in pipiens,

particularly 1a- and 1b-S (cf. in Tables 21 and 22). Segment X: Saddle length 0.35-0.48

mm, mean 0.41 mm; siphon/saddle index 2.77-3.41, mean 3.11.

Systematics. Culex quinquefasciatus is a variable but fairly well differentiated

species which has introgressed with pipiens in areas where the two species come

together. During this study, 180 mongrels (57 females, 67 males, and 56 male genitalia)

were examined from the lower Tigris-Euphrates Valley in Iraq, the eastern coastal region

of Saudi Arabia along the Persian Gulf, Al Kharj Oasis near Riyadh, and Jeddah on the

Harbach: Subgenus Culex in Southwestern Asia 35

Red Sea coast. Hybrids are morphologically more variable than the parental species.

Some hybrids are recognizable in having the scutal scaling of one species and the

abdominal banding of the other species, but in general the only reliable means of

distinguishing hybrids from the recurrent parental species is by the male genitalia. The

form of the dorsal and ventral arms of the phallosome is intermediate between that found

in pipiens and quinquefasciatus. Hybrid males examined from southwestern Asia have

DV/V ratios ranging from 0.08 to 0.43 with a mean of 0.18. Mongrels of partial hybrid

ancestry may be indistinguishable from the parental species. For this reason, some

hybrid specimens collected in and around the zone of introgression were probably

misidentified as parental epiphenotypes during the course of this study. There is no

doubt, however, that the parental epiphenotypes show independent species cohesion

outside the zone. The two epiphenotypes are closely related, distinct species. The

capacity for pipiens and quinquefasciatus to interbreed should not be considered as

prima facie evidence for incomplete speciation.

Culex quinquefasciatus is widely distributed in the tropical and subtropical zones

of the world where it has no doubt been spread by commerce. However, the taxon has

not been sufficiently studied thoughout its range and it is possible that it consists of two

or more biological species. There is currently no evidence to suggest that more than

one species occurs in southwestern Asia.

Bionomics. Culex quinquefasciatus is a common domestic and peridomestic

species which has received considerable attention in the literature. It breeds mainly in

polluted water but is also found in clean fresh water and brackish water habitats. The

immatures occur in virtually all types of man-made and natural breeding places. Females

are undoubtedly anthropophilic but readily feed on a variety of other mammals and birds

depending on the availability of hosts. Biting occurs both indoors and outdoors at night,

and engorged females can be found resting either inside or outside dwellings and

shelters.

Although it is well known that quinquefasciatus is an important vector of filarial and

viral agents of human disease in many parts of the world, there are no published reports

of natural infection in southwestern Asia. Culex quinquefasciatus is the predominant

vector of urban filariasis in areas of Africa and the Orient. It has been found infected with

Japanese encephalitis virus in Vietnam (Nguyen et al., 1974) and Chikungunya virus in

Tanzania (White, 1971). It is capable of transmitting West Nile virus in India (Rao, 1975)

and Murray Valley encephalitis virus in Australia (Karabatsos, 1985).

Distribution. Culex quinquefasciatus is widely distributed in the tropical and

subtropical regions of the world.

Material examined. 1,113 specimens. A total of 907 specimens (306

females, 308 males, 256 male genitalia, 10 pupal exuviae, 5 larval exuviae, and 22

fourth-instar larvae) were examined from southwestern Asia — BAHRAIN: (specific

localities unknown); IRAN: (Alchangi, Bushehr, Enzeli, Halileh, Hizan, Kish Island,

Manyuhi, Nahr Mohsen, Shush); IRAQ: (Amara, Baghdad, Basrah); OMAN: (Muscat,

Salalah); PAKISTAN (including localities east of the Indus River): (Jhang, Jhelum,

Lahore, Peshawar, Rawalpindi); P.D.R. YEMEN: (Anag, Lahej, Makhzan, N'Air, Wadi

Duan); SAUDI ARABIA: (Al Hasa Oasis, Al Khobar, Al Qatif, Al Ta'if, Dhahran, Jeddah,

Mecca, Saihat); YEMEN ARAB REPUBLIC: (Murawah). An additional 206 specimens

(102 females, 46 males, 31 male genitalia, 12 pupal exuviae, 12 larval exuviae, and 3

fourth-instar larvae) were examined from DJIBOUTI, ETHIOPIA, MAURITANIA,

MOZAMBIQUE, NEPAL, SENEGAL, SOCOTRA, SOUTH AFRICA, SUDAN, UGANDA,

and the UNITED STATES (the neotype series).

36 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Culex (Culex) vagans Wiedemann

vagans Wiedemann, 1828: 545. Syntypes (non-extant); Foochow

(Barraud, 1934: 416), China.

tipuliformis Theobald, 1901b: 325. *Holotype female: Bakloh, N. W.

Provinces (? Pakistan) (BM). Synonymy with vagans by Edwards,

1926: 136.

virgatipes Edwards, 1914b: 126. *tHolotype male: Hong Kong (BM).

Synonymy with vagans by Edwards, 1926: 136.

exilis Dyar, 1924: 127. +Lectotype male: Vladivostok, [Maritime

Territory], USSR; designated by Stone and Knight, 1957: 49

(NMNH). Synonymy with vagans by Edwards, 1926: 136.

Culex tipuliformis Theobald of Barraud, 1924b: 1269 (India, A key, syn., M’*, F, distr.);

Barraud, 1924c: 430 (India, L"*).

Culex vagans Wiedemann of Theobald, 1901a: 411 (Oriental Region, A key, F*).

Culex (Culex) vagans Wiedemann of Edwards, 1926: 136 (Palaearctic Region, A, L

keys, syn., A, distr.); Martini, 1931: 391 (Palaearctic Region, A, L keys, M", F,

syn.); Barraud, 1934: 416 (India, A, L keys, syn., A, M gen.*, L*, L habitat, distr.);

Monchadskii, 1951: 283 (Middle East, L key, L*, bionomics, distr.); (?) Parrish,

1959: 266 (Turkey); DuBose and Curtin, 1965: 352, 354 (Mediterranean area, A,

L keys); (?) Lotfi, 1970: 400, 402 (Iran); Aslamkhan, 1971: 155 (Pakistan); (7?)

Lotfi, 1976: 73, 76 (Iran, L key, L*); Sirivanakarn, 1976: 37 (Oriental Region, A, P,

L keys, M*, F, P*, L*, tax., bionomics, distr.); Harbach, 1985a: 86, 94, 105

(Afghanistan, Iran [not Egypt], Pakistan, A, L keys).

Adult. This medium-sized species resembles thei/eri in having anterior pale

stripes on the fore- and midfemora and all tibiae, but is easily distinguished by the

absence of postspiracular and prealar scales. The species otherwise resembles pipiens,

which of course lacks the striped legs. The male genitalia are strikingly similar to those of

torrentium.

FEMALE. Head: Length of antenna about 2.2 mm; pedicel usually mainly yellow,

mesal surface black, both pedicel and flagellomere 1 with some small white scales on

mesal surface; flagellum normal, whorls normally with 6 setae. Proboscis length 2.0-2.4

mm, mean 2.2 mm; sides and venter with cream-colored scales extending from 0.15 to

0.70 from base, proximal and distal portions distinctly black-scaled. Maxillary palpus

dark-scaled with white scales mainly confined to dorsal stripe on palpomeres 3 and 4;

length about 0.4 mm, about 0.2 proboscis length. Vertex with whitish to pale yellow

falcate scales, usually paler medially and along ocular line; forked scales mainly brown,

some pale ones usually in middle; lateral spatulate scales white or nearly so. Interocular

space with few pale scales, concolorous with ocular scales. Ocular setae dark brown with

golden or reddish sheen; 2 yellowish interocular setae project ventrally over clypeus.

Thorax: Pleural integument yellowish brown to brown; scutal integument brown. Color

of scutal scales ranging from brownish yellow to golden brown, often mainly yellowish

brown, surface sometimes appearing somewhat mottled with fossal, sublateral supraalar

and occasionally posterior acrostichal scales darker; scales pale yellow, whitish or even

white on anterior and lateral margins and on prescutellar area. Scutal setae dark brown

with golden or reddish sheen. Scales of scutellum same color as prescutellar scales;

lateral lobes each with 4or5 large setae, median lobe with 6-8. Antepronotum mainly

Harbach: Subgenus Culex in Southwestern Asia a7

with dark brown setae, some yellowish ones ventrally; with rather coarse falcate scales in

upper and lower patches, upper patch brownish yellow, lower whitish. Postpronotum

with brownish-yellow falcate scales, some paler (frequently whitish) posteriorly; with

usually 6 or 7 dark setae on posterodorsal margin. Pleural setae golden: 8-12 upper

proepisternal, 10-12 prealar, 4-6 upper mesokatepisternal, 9-13 lower mesokat-

episternal, 10-14 upper mesepimeral, and 1 lower mesepimeral. Pleura with white,

elongate spatulate scales in patches as follows: small patch below upper proepisternal

setae, patches on upper corner and lower posterior border of mesokatepisternum,

anterior patch on mesepimeron at level of upper mesokatepisternal patch, and patch

before and among upper mesepimeral setae; lower mesokatepisternal and upper

mesepimeral scales semierect, others decumbent. Wing: Length 3.5-5.4 mm, mean

4.5 mm; length cell R2 2.9-3.8 length of vein R2+3, mean 3.3; length cell M1 0.8 length

of cell R2; scales predominantly dark, pale scales on subcosta and posterior side of costa

on dorsal and ventral surfaces, dorsal surface also with pale scales on anterior and

posterior sides of R1 distal to furcation of vein R2+3. Halter: Integument usually

yellowish, dorsal areas of pedicel and capitellum often dark; scales entirely pale. Legs

(midleg, Fig. 2G): Anterior surface of forecoxa dark-scaled in middle, with patches of

white scales at base and apex; midcoxa with longitudinal patch of white scales on

anterior side of midlateral row of strong setae, anterior surface with spot of scales at

apex; hindcoxa with longitudinal patch of white scales on anterior side of lateral midline,

posterolateral margin with irregular row of strong setae. Lateral surfaces of trochanters

without scales, other surfaces white-scaled, anterior surface of foretrochanter often with

some dark scales. Femora mainly white-scaled, apices with pale knee spots, spots often

indistinct on fore- and midfemora; anterior and dorsal surfaces of fore- and midfemora

dark-scaled, anterior surface with narrow longitudinal white stripe, forefemur sometimes

with ventral 0.5 of anterior surface completely white-scaled, or nearly so, because

anteroventral dark scaling partially or completely absent; hindfemur with complete

anterodorsal stripe of dark scales, stripe gradually widening distally, expanded over

approximately distal 0.5 of dorsal surface but may or may not be expanded over whole of

anterior surface very near apex. Anterior and posterior surfaces with dark scales: dorsal

surface with spot of white scales at apex; ventral dark scales of foretibia sometimes partly

or entirely replaced by white scales. Tarsi dark-scaled with faint anterior and posterior

pale stripes on tarsomere 1. Abdomen: Tergum | with median posterior patch of

yellowish scales; terga II-VII mainly clothed in dark brown to blackish scales; terga II-VII

with rather large basolateral patches of white scales, sometimes covering most or all of

lateral surface, particularly on terga Il, VI and Vil; tergum Il with basomedian patch of

yellowish scales, patch sometimes lengthened posteriorly; terga III-VII with basal bands

of yellowish scales, bands 0.3-0.5 of tergum length, bands of terga III-V (and often VI

and sometimes VII) convex, those of VI and VII often straight or nearly so, bands of terga

Il and IV (and frequently V and VI) not reaching basolateral white patches; tergum VIII

usually entirely pale-scaled, sometimes with some dark scales posteriorly. Sterna II-VII

mainly clothed in yellowish scales, with a median area of dark scales; sternum VIII with

lateral patches of pale scales, median area without scales.

MALE. Like female except for the following principal differences. Head:

Proboscis with indefinite pale band (narrowly incomplete dorsally) about 0.5-0.7 from

base, pale scaling of ventral surface extending from band to about 0.2 from base: false

joint 0.6 from base; without ventral cluster of setae at false joint. Maxillary palpus mainly

dark-scaled above; lateral surface of palpomere 3 with rather broad stripe of white scales

0.2-0.8 from base, mesal surface with complete or nearly complete narrow line of white

scales which is broader and more noticeable near apex, ventrolateral margin with row of

38 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

about 28 long dark setae on distal 0.5, ventromesal margin with complete row of short,

pale, antrorsely curved setae; palpomere 4 with complete ventral stripe of white scales

and dorsal spot of rather subtle pale scales at base; dorsal surface of palpomere 5 like

that of 4, ventral surface with distinct basal spot of white scales; lateral and mesal

surfaces of palpomeres 4 and 5 densely setose; palpus exceeds length of proboscis by

length of palpomere 5. Thorax: Scutal scales finer. Upper proepisternal setae more

numerous. Wing: Length of cell R2 1.5-2.0 length of vein R2+3; subcosta intersects

costa before furcation of vein R2+3; length of cell M1 0.8-0.9 length of cell R2.

Abdomen: Terga II-VIl without lateral white patches; basal bands of terga III-VII either

convex or nearly straight, bands of terga V-VII produced posteriorly along lateral

scale-free areas but only slightly so on V; tergum VIII (ventral in position) with pale scales

before median posterior setose area. Sterna II-VIl usually with median dark scales

forming distinct stripe; sternum VIII (dorsal in position) entirely pale-scaled. Genitalia (Fig.

7): Form as figured; in general as in pipiens and related species. Ninth tergal lobe short

and broad, with 6-14 unevenly spaced setae. Gonocoxopodite essentially as in pipiens;

subapical lobe indistinctly divided, seta g on small prominence with h at its base laterally;

usually 2 setae in group d-e, sometimes only 1, sometimes 3; seta g not as broad,

pointed; / flattened and shorter. Phallosome longer than broad with lateral plates

slightly longer than aedeagal sclerites; dorsal arm slightly twisted and pointed apically,

diverging laterally from its mate of the opposite side but not appearing sinuous in lateral

view; lateral arm longer than broad, with prominent triangular laterally directed apical lobe

and a short dorsal process, apical lobe minutely spiculate on posterior surface, dorsal

process relatively short and borne on dorsal margin between apical lobe and rather

weakly developed basal articulatory process; ventral arm short, bent as in pipiens.

Aedeagus short, somewhat conical; aedeagal sclerites about as long as broad; ventral

aedeagal bridge rather narrow, joining aedeagal sclerites near midlength. Proctiger

normal; paraproct with long, rather stout, ventrally curved basal lateral arm, crown with

numerous spinelike spicules and some short lateral blades. Cercal sclerite elongate,

with 2-5 cercal setae. Tergum X developed as usual.

Pupa (Fig. 7). Placement and attributes of setae as figured, range and modal

number of branches in Table 3; differing little from pipiens and allied species, but usually

recognized without much difficulty by the bent trumpet and relatively long seta 5-IV.

Cephalothorax: Seta 1,2,4,6-CT usually with 3 or 4 branches; 3-CT usually triple (2,3);

7-CT double; 8-CT tending to have more branches than usual (3-9); 9-CT with 2-4

branches; 11-CT usually double, sometimes single. Trumpet: Distinctly bent at distal

end of tracheoid area; index 5.0-6.9, mean 5.6; pinna short, 0.15-0.20 length of

trumpet. Abdomen: Length 2.1-2.6 mm, mean 2.4 mm. Seta 1-IlI-VIl tends to have

more branches than in pipiens and related species, 1-lll frequently with 9 or 10 branches

(6-11), 1-IV often with 8 branches (5-10); 5-IV as strong as 5-V,VI, normally double or

triple (2-4). Paddle: Length 0.73-0.81 mm, mean 0.78 mm; width 0.51-0.60 mm, mean

0.56 mm; index 1.2-1.6, mean 1.4.

Larva (Fig. 8). Placement and character of setae as figured, range and modal

number of branches in Table 23; very similar to pipiens, but usually easily recognized by

the character of seta 1-C and the presence of rows of minute vesicles on the thoracic

and abdominal integument. Head: Length 0.76-0.92 mm, mean 0.82 mm; width

1.17-1.30 mm, mean 1.24 mm; lightly to moderately tanned, darker behind eyes, gula

also usually darker; collar and dorsomentum heavily tanned. Dorsomentum usually with

9 teeth (8-11) on either side of median tooth. Seta 1-C long, tapered, tanned, and

usually spiculate in middle; seta 2-C absent. Antenna: Generally slightly longer than in

pipiens, length 0.52-0.58 mm, mean 0.57 mm. Seta 1-A with about 25 branches

Harbach: Subgenus Culex in Southwestern Asia 39

(20-33). Thorax: Integument with rows of minute vesicles. Seta 1-M single, normally

slightly longer than 0.5 length of 3-M. Seta 1-T about 0.5 length of 2-T; 2-T frequently

single, sometimes double; 13-T rather long, about same length as 12-T. Abdomen:

Integument with rows of tiny vesicles, changing to rows of tiny spicules on segment VIII.

Seta 6-| usually triple (2-4), 6-II usually with 3 or 4 branches (3-5), 6-III-VI almost always

double, occasionally triple. Segment Vill: Comb with 36-47 scales, mean 41. Siphon:

Generally longer than in pipiens, index 3.92-6.72, mean 5.39; lightly to moderately

tanned. Pecten with 10-15 spines, mode 13; larger spines with 2-5 denticles, usually 3

or 4. Seta 1-S usually in 4 pairs, occasionally with a fifth seta on one side; 1b-S often

inserted laterally in line with 1c-S. Segment X: Saddle length 0.38-0.47 mm, mean 0.41

mm; siphon/saddle index 3.79-4.59, mean 4.19. Seta 1-X single or double, more often

single. Anal papillae equal in length, slightly longer than saddle.

Systematics. Culex vagans and the next species (torrentium ) bear a

pronounced resemblance to members of the pipiens subgroup, but based on the male

genitalia they unmistakingly belong to the trifilatus subgroup of Mattingly and Rageau

(1958). Culex vagans is the most distinctly marked member of the subgroup, the adults

being readily distinguished from all other species of the group, as well as from pipiens

and quinquefasciatus, by the striped femora and tibiae. This species bears a superficial

resemblance to theileri with which it has been confused in the past.

This species occurs in a relatively broad belt which stretches from the Middle East

to Hong Kong, Japan, and Korea. Its exact distribution in southwestern Asia is

unknown. | have seen specimens only from Afghanistan and Pakistan, but the species

has been reported to occur in Iran (Lotfi, 1970; 1976) and Turkey (Parrish, 1959). | am

somewhat skeptical about the records from Iran because they are based on limited

collections of larvae. Characters which have been used to separate the larva of vagans

from the larvae of pipiens, quinquefasciatus, and torrentium are certainly not as reliable

as they appear to be in previously published keys (see the list of earlier works following

the synonymy). In fact, the characters used in my key are probably not completely

reliable because of the very limited number of specimens examined. Concerning the

record from Turkey, it is not clear whether Parrish (1959) based his identification on larval

or adult specimens. Parrish apparently collected much of his material in light traps. It is

possible that his record of vagans from Turkey is based on damaged and/or partially or

completely denuded females of theileri and/or males of torrentium collected in light

traps. | would have the same concern about the identification of larvae of vagans from

Turkey that | have for the identification of larvae from Iran. Mattingly (1955a: 31)

identified three male genitalia and a larva of torrentium from Turkey which were labelled

previously as vagans by Martini (1931).

The adult and immature specimens examined during this study seem to conform

quite well with the recent descriptions of this species based on material from eastern

Asia (Sirivanakarn, 1976; Tanaka et al., 1979). The only obvious difference is the

development of seta 1-C in the larva, and this may be within the normal range of

geographic variation. At present there seems to be no reason to suspect that more than

one species exists under the name of vagans, but this will not be known for certain until

considerably more material from southwestern and central Asia is studied in detail.

Bionomics. Little specific information is available on the bionomics of vagans.

In eastern Asia, the immature stages are found in permanent or semipermanent bodies

of fresh ground water, including streams, stream pools, ponds, lakes, and marshes.

Larvae have been collected in Iran in ponds with or without vegetation (Lotfi, 1976). In

Japan and Korea, females are known to bite man at night (Hsiao and Bohart, 1946;

Barnett and Toshioka, 1951). Culex vagans has been reported tobe naturally and

40 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

experimentally infected with larvae of Wuchereria bancrofti (Yamada, 1927, as

tipuliformis Theobald; Manson-Bahr, 1959; Hsiao, 1948; Hsiao and Bohart, 1946) and

should be regarded as a potential vector of this pathogen in southwestern Asia.

Distribution. This species is widespread in eastern and southcentral areas of

the Palaearctic Region. It is Known to occur in eastern Siberia, Korea, Japan, Hong

Kong, China, Mongolia, Bangladesh, Nepal, northern India, Pakistan, (7?) Iran,

Afghanistan, and (7?) Turkey.

Material examined. 83 specimens. A total of 68 specimens (11 females, 10

males, 12 male genitalia, 8 pupal exuviae, 19 larval exuviae, and 8 fourth-instar larvae)

were examined from southwestern Asia — AFGHANISTAN: Bamiyan (Sayedabat);

PAKISTAN (including areas in eastern Pakistan): (Gharian, Gujranwalia, Jhelum,

Khanoharni, Lahore, Sattoki). An additional 15 specimens (3 females, 6 males, and 6

male genitalia) were examined from HONG KONG (the holotype male of virgatipes ),

INDIA (including the holotype female of tipuliformis ), and the USSR (the lectotype male

of exilis).

Culex (Culex) torrentium Martini

torrentium Martini, 1925: 336. *+Hololectotype male: Schwartza River,

[Thueringia], Germany; designated by Mattingly, 1955a: 31 (BM).

pavlovsky Shingarev, 1928: 52. Type specimen(s) (male): Naryn

District, Romsk, Siberia, USSR (TMS). Synonymy with torrentium by

Edwards, 1932a: 110.

Culex (Culex) exilis Dyar of Martini, 1931: 386 (Palaearctic Region, A, L keys, syn., M’, F,

L); Monchadskii, 1951: 260 (Middle East, L, key, syn., distr.).

Culex torrentium Martini of Zaini et al., 1983: 117 (Iraq).

Culex (Culex) torrentium Martini of Edwards, 1926: 136 (Palaearctic Region, A key, A,

distr.); Senevet et al., 1957b: 92 (E. Mediterranean (?), L); Parrish, 1959: 266

(Turkey, distr.); Senevet and Andarelli, 1959: 203 (Mediterranean Basin, A, L, P

keys, syn., M*, F, P*, L*, distr., biol., L assoc.); DuBose and Curtin, 1965: 352,

354 (Mediterranean area, A, L keys); Abul-hab, 1969: 249 (Iraq, L key); Lotfi,

1970: 402 (Iran); Lotfi, 1976: 73, 77, 83 (Iran, L key, ecol. note, L*); Ibrahim et al.,

1983: 91 (Iraq, L key); Harbach, 1985a: 86, 94, 105 (Iran, Iraq, Turkey, distr., A, L

keys).

This description is based almost entirely on specimens from Sweden.

Adult. A medium-sized species closely resembling pipiens, but slightly larger,

with prealar scales and different male genitalia.

FEMALE. Description based on specimens from Sweden. Head: Antennal

length about 2.2 mm; pedicel yellow or orange, mesal surface black with some tiny setae

and whitish scales; flagellum normal, whorls with 6 setae, proximal part of flagellomere 1

with some tiny whitish scales on mesal surface. Proboscis mainly black-scaled, ventral

surface with pale scales extending from 0.1 to 0.7 from base. Maxillary palpus usually

entirely black-scaled, palpomere 4 sometimes with some indistinct pale scales on

dorsomesal surface; length 0.4 mm, about 0.2 proboscis length. Falcate scales of

vertex mainly pale yellow, narrower and darker laterally, whitish along margin of

compound eye and on interocular space; forked scales yellow medially, black postero-

Harbach: Subgenus Culex in Southwestern Asia 41

laterally; lateral spatulate scales yellowish white. Ocular setae dark reddish brown:

interocular setae golden brown. Thorax (Fig. 2C): Pleural integument yellowish brown,

faded posteriorly; scutal integument dark brown. Scutal scales uniformly golden brown

with slight reddish tint, paler on anterior promontory, among supraalar setae, and on

margins of prescutellar area; scales finer on fossa and supraalar area between supraalar

and posterior dorsocentral setae, but not appreciably darker. Scutal setae dark reddish

brown. Scutellum with fine pale yellow falcate scales; 4-7 large setae on each lateral

lobe, 7 or 8 on median lobe. Antepronotum with yellowish falcate scales in upper and

lower patches, upper patch often darker, sometimes golden brown; setae dark reddish

brown dorsally, becoming golden brown ventrally. Postpronotum with fine

golden-brown falcate scales on dorsal 0.5, paler and slightly coarser posteriorly; with

5-10 dark setae on posterodorsal margin, longer posteriorly. Pleural setae golden

brown: 7-9 upper proepisternal, 8-11 prealar, 5-7 upper mesokatepisternal, 7-11 lower

mesokatepisternal, 9-12 upper mesepimeral, and 1 lower mesepimeral. Pleura with

white or nearly white elongate spatulate scales in patches as follows: small patch below

upper proepisternal setae, few to small patch below prealar setae, patches on upper

corner and lower posterior border of mesokatepisternum, anterior patch on

mesepimeron about same level and size as upper mesokatepisternal patch, and patch

mainly before upper mesepimeral setae; rarely with some postspiracular scales. Wing:

Length about 4.7 mm; length of cell R2 3.0-3.8 length of vein R2+3; subcosta intersects

costa at or slightly beyond furcation of vein R2+3; length of cell M1 0.7-0.8 length of cell

R2; scales entirely dark. Dorsal scaling: broad decumbent scales on costa, R and R1:

narrow decumbent scales on R4+5, M3+4, mcu, CuA and proximal 0.5 of 1A;

subcosta with broad semidecumbent scales; linear plume scales on Rs, R243, 2,

R3, M, and M1+2; near-linear semiplume scales on M1, M2 and distal 0.5 of 1A; remigium

with 2 distinct rows of broad decumbent scales and 2 or 3 setae distally. Ventral scaling:

narrow decumbent scales on costa, subcosta, Rs, R2+3, M, M1+2, bases of M1 and M2,

and distal 0.5 of 1A; linear plume scales on R1, R2, R3, R4+5, beyond bases of M1 and

M2, mcu, M3+4, and CuA distal to mcu; R, CuA proximal to mcu, and proximal 0.5 of 1A

without scales. Halter: Entirely pale, capitellum slightly darker dorsally. Legs: Anterior

surface of forecoxa mainly black-scaled, with small patch of cream-colored scales at base

and usually some scattered pale scales distally, also with many long, dark, ventrally

curved setae, posterior margin with 3 or 4 shorter setae at apex, most proximal of these

more or less perpendicular to surface, others project ventrally; midcoxa with midlateral

longitudinal row of 4 or 5 long golden-brown setae and several shorter ones margined

anteriorly by longitudinal patch of whitish spatulate scales, anterior surface with small

patch of black scales and several short ventrally projecting setae at apex; posterolateral

surface of hindcoxa with longitudinal row of 6-8 golden-brown setae of different lengths,

anterolateral surface with narrow longitudinal row of white spatulate scales and several

ventrally projecting setae at apex. Lateral surfaces of trochanters without scales,

foretrochanter with black scales anteriorly and white scales posteriorly, anterior, mesal

and posterior surfaces of mid- and hindtrochanters with white scales. Apices of femora

with knee spots of yellowish scales; forefemur black-scaled anteriorly, white-scaled

posteriorly; midfemur like forefemur but distal portion with black scales spread over

dorsal surface; hindfemur with complete anterodorsal stripe of black scales, Stripe

gradually widening distally to cover about distal 0.5 of dorsal surface but not covering

whole of anterior surface except near apex. Foretibia mainly black-scaled, with

posteroventral stripe of white scales, apex with inconspicuous dorsal spot of whitish

scales; anterior surface of midfemur black-scaled with some faint pale scales at apex,

posterior surface white-scaled; anterior and dorsal surfaces of hindtibia black-scaled with

42 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

conspicuous spot of whitish scales at apex, posterior and ventral surfaces with whitish

scales. Tarsi black-scaled, with some indistinct pale scaling posteriorly, particularly on

tarsomere 1 of fore- and midlegs. Pulvilli pale. Ungues black, small, simple. Abdomen:

Terga mainly black-scaled; tergum | golden-brown setose, with posteromesal patch of

black scales; posterior and lateral margins of terga II-VIII with golden to golden-brown

setae; tergum II with small basomedian yellowish spot and lateral patches of white scales;

terga III-VII with yellowish basal bands and white basolateral patches, spots slightly

increasing in size on succeeding posterior terga, bands of terga IIl-V convex and usually

not reaching spots, bands of terga VI and VII reaching spots, band of VI usually straight,

that of VII slightly concave; lateral scales of tergum VIII white, dorsal scales entirely

yellowish or black posteriorly. Sterna II-VIl mainly yellowish-scaled, with blackish scales

forming a median spot or streak; sternum VIII with lateral patches of whitish scales,

devoid of scales medially; all sterna golden to golden-brown setose.

MALE. Differs from female as follows. Head: Proboscis length about 2.4 mm;

false joint 0.6 from base; ventral pale scales usually extending from 0.4 to 0.7 from base;

without ventral cluster of setae at false joint. Maxillary palpus length about 3.4 mm,

usually extending beyond tip of proboscis by more than length of palpomere 5; mainly

dark-scaled; ventral surface of palpomeres 2 and 3 without scales, integument pale;

palpomere 3 with or without some indistinct pale scaling on lateral surface, with

ventrolateral row of 15-25 long setae on distal 0.33 and a complete row of short

antrorsely curved setae on ventromesal margin; palpomere 4 with complete or nearly

complete ventral line of white scales, distal scales usually losing their color; palpomere 5

with some white scales at base ventrally; lateral and mesal surfaces of palpomeres 4 and

5 densely setose. Falcate scales of vertex of uniform size and color; forked scales less

numerous and shorter, particularly posterolaterally. Thorax: Scutal scales slightly

coarser, scales and integument between supraalar and posterior dorsocentral setae

noticeably darker in dorsal view. Propleuron with 11-16 upper proepisternal setae more

or less in 2 rows. Wing: Length about 3.9 mm; length of cell R2 2.0-2.5 length of vein

R2+3; length of cell M1 0.75-0.85 length of cell R2; subcosta intersects costa before

furcation of vein R2+3. Legs: Forecoxa without scattered pale scales distally. Anterior

unguis longer than posterior unguis on fore- and midlegs, each with midventral tooth;

posterior unguis of midleg smaller than posterior unguis of foreleg, both with small

ventral tooth before middle; hindungues small, simple. Abdomen: Terga without

basolateral spots; basal bands usually 0.4 tergum length (0.25-0.40); bands of terga III-V

usually straight, those of terga VI and VII produced posteriorly along lateral scale-free

margins, particularly on tergum VII; tergum VIII (ventral in position) pale-scaled before

posteromesal setose area. Sterna II-VII frequently with dark scales confined to median

area and posterolateral corners; sternum VIII (dorsal in position) pale-scaled, often with

some subtle dark scales which tend to form a pair of submedial spots. Genitalia (Fig. 9):

Form as figured; very much as in vagans, differing in minor details as follows. Ninth tergal

lobe with 5-11 setae. Subapical lobe of gonocoxite always with 2 setae in group d-e;

seta g slightly longer, rounded apically. Dorsal arm of lateral plate of phallosome broader,

0.25 distinctly twisted and curved mesad; lateral arm also broader, apical lobe with more

numerous and larger spicules on posterior surface, dorsal process more strongly

developed.

Pupa (Fig. 9). Character and arrangement of setae as figured; range and modal

number of branches in Table 4; closely resembling pipiens from which it cannot be

distinguished with certainty, seta 5-VI apparently stronger than in pipiens, trumpet

exhibiting slight differences in length of tracheoid area and pinna. Cephalothorax:

Lightly tanned, scutum, legs, and metathorax darker. Setae 1,4,6,8,12-CT usually triple,

Harbach: Subgenus Culex in Southwestern Asia 43

1-CT with 2-4 branches, 4,6-CT with 3-5 branches, 8-CT with 1-4 branches and 12-CT _

with 2-4 branches; 2,5-CT usually with 4 branches, 2-CT with 2-5 branches and 5-CT

with 3-6 branches; 3,7,11-CT usually double, sometimes triple; 9-CT double. Trumpet:

Moderately tanned, subcylindrical, gradually widened distally; index 5.2-6.1, mean 5.5:

tracheoid area darker, extending about 0.4 from base; pinna oblique, shorter than in

pipiens, 0.5-0.7 length of tracheoid area. Abdomen: Lightly tanned, terga I-IV dark in

middle, particularly on anterior margin; length 2.6-3.3 mm, mean 2.9 mm. Seta 1-Il most

often with 15 branches (12-16); 1-IIl-V frequently with at least 5 branches (5-9 on IV; 4-9

on V); 5-IV usually triple, sometimes with 4 branches; 5-V-VII normally double, 5-VII rarely

single; 6-III-VI most often triple, 6-III,1V double or triple, 6-V with 3 or 4 branches, 6-VI

usually with 3 or 4 branches, infrequently with 2 or 5 and generally longer than 6-lll-V.

Genital lobe: Tanning and length same as in pipiens. Paddle: Essentially as in pipiens;

length 0.80-1.05 mm, mean 0.91 mm; width 0.53-0.74 mm, mean 0.63 mm: index

1.3-1.7, mean 1.5.

Larva (Fig. 10). Placement and character of setae as figured, range and modal

number of branches in Table 24; extremely similar to pipiens and vagans, differing

chiefly in having setae 1-III-V and 1-M more branched, also usually distinguishable from

pipiens by the character of setae 1-T, 3-I,VIl, and 1-X. Head: Length 0.76-0.86 mm,

mean 0.83 mm; width 1.06-1.23 mm, mean 1.14 mm; moderately tanned, lateralia

slightly darker posterior to eyes; black spot area, dorsomentum and collar heavily

tanned. Dorsomentum usually with 8 teeth (8-10) on either side of median tooth. Seta

7-C most often with 8 or 9 branches (8-10); 9-C usually triple (2-5). Antenna: Generally

longer than in pipiens, length 0.50-0.64 mm, mean 0.57 mm; lightly tanned before seta

1-A, darker at level and distal to 1-A, base with dark spot on mesal surface. Seta 1-A with

about 26 branches (21-29). Thorax: Seta 1-M usually double or triple, infrequently with

4 branches, length normally about 0.6 length of 3-M. Seta 1-T normally slightly longer

than 0.5 length of 2-T, often with 4 branches (3-7); 2-T double. Abdomen: Seta 3-I,VII

usually double, sometimes single on one side, seldom single on both sides: 6-| usually

triple (3,4), 6-Il most often with 3 or 4 branches, sometimes with 5 branches: 1-III-V

usually with 4 branches, 1-VI normally triple, sum of branches for 1-III-VI always greater

than for pipiens and vagans. Segment Vill: Comb with 31-46 scales, mean 39. Seta

1-VIIl frequently with 6 branches (5-9); 3-VIII most often with 7 branches (6-9). Siphon:

Index 4.40-5.67, mean 5.23; shape and tanning as in pipiens. Pecten with 10-16

spines, mode 13. Seta 1-S usually in 4 pairs; siphon often with a fifth seta 1-S on one or

both sides, rarely with 3 on one side. Segment X: Saddle length 0.32-0.38 mm, mean

0.37 mm; siphon/saddle index 3.92-4.65, mean 4.17. Seta 1-X usually double,

sometimes single on one or both sides, rarely triple; 2-X usually double, occasionally

triple; 4-X in 6 pairs; anal papillae subequal, longer than saddle.

Systematics. The occurrence of torrentium in southwestern Asia is based on

the examination of a few very poor specimens from Iran and reports from the literature.

The description here is based almost entirely on material from Sweden. Although the

occurrence of forrentium in Turkey is probably correct (see Mattingly, 1955a: 31), some

doubt remains about the occurrence of this species in Iran and Iraq. Previous records

from these countries are based on the identification of larvae (Lotfi, 1970, 1976: Ibrahim

et al., 1983). Since it has been almost impossible to differentiate the larvae of torrentium

and pipiens using previously published keys, it is not unlikely that these records actually

refer to pipiens. | have seen only one denuded male with associated genitalia, an

unassociated male genitalia, and a single fourth-instar larva from Iran, and | am not

absolutely certain that these are specimens of torrentium. The genitalia may be those of

vagans and the larva that of pipiens. These specimens are only provisionally assigned to

44 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

torrentium.

The female and larva of forrentium have never been easy to distinguish from

pipiens. These species, however, are not as closely related as the outward similarity

would suggest. The two species are well separated in the structure of the male

phallosome. If the morphological variation observed in the different life stages of

torrentium during this study is representative of the holomorphology of the species,

then the characters mentioned in the keys should reliably distinguish most specimens of

torrentium from pipiens in southwestern Asia, assuming of course that torrentium

indeed occurs there.

Bionomics. Very little is Known about the biology of forrentium. The immatures

frequently occur with pipiens in Europe. Larvae are commonly found in ground pools,

but are also known to occur in artificial containers such as tin cans, metal tanks, rain

barrels, and concrete ornaments (Service, 1968). Harbach et al. (1985) reported their

occurrence with pipiens in an old boat. This species apparently tolerates some organic

matter and salt in its breeding sites. Collections have been made in rock pools in which

the water was dark brown with decaying leaves (Ribeiro et al., 1977) and in ground pools

with salinities of 0.3-0.4 that of sea water (Lever, 1954). Females are not attracted to

man and apparently feed on birds (Callot, 1957; Service, 1968).

Distribution. This species is widely distributed in Europe and western Asia,

including Turkey, (?) Iran, and (’?) Iraq.

Material examined. 449 specimens. Only 4 specimens (1 male, 2 male

genitalia, and 1 fourth-instar larva) were examined from southwestern Asia — IRAN:

(Kivi, Sarab-bahran Kazeroon). A total of 445 specimens (73 females, 71 males, 36

male genitalia, 140 pupal exuviae, 102 larval exuviae, and 23 fourth-instar larvae) were

examined from EAST GERMANY (the hololectotype male, allolectotype female, and 2

paralectotypes), PORTUGAL, SWEDEN, and WEST GERMANY.

Culex (Culex) decens Theobald

decens Theobald, 1901c: vii. t*tLectotype male, hereby designated,

bearing following data: "Bonny [Nigera] / 29 V 00 // Type // Culex /

decens / (Type). Theobald.", genitalia on acetate strip (BM).

masculus Theobald, 1901b: 125. tLectotype male, hereby designated,

bearing following data: "Wilberforce, / Free Town. / 3.9. 99./ Fr. larva

in / wood-side puddle. // Type // Free Town. / Sierra Leone. / |X. 99. /

E.E. Austen. 99. 267. / Culex / masculus / (Type) Theobald";

genitalia on acetate strip (BM). Synonymy with decens by Edwards,

TOT 2ES

minutus Theobald, 1905c: 30. +Lectotype male, hereby designated,

bearing following data: "Type // Transvaal / Dr. Simpson // Culex /

minutus. / (Type). F.V.T. // Recd. from / F.V. Theobald / 1907-29.";

genitalia on acetate strip (BM). Synonymy with decens by Edwards,

1OT1) 263:

nigrocostalis Theobald, 1909: 16. +Lectotype male, hereby

designated, bearing following data: "Accra / 7-6-08 / c in latrine /

WMG // Culex / nigrocostalis / Type. [male symbol]. F.V.T / 30. 3. 09.

// Accra, / Gold Coast, / W. Africa. / 7-VI. 1908. / Dr. W.M. Graham. /

1909-271."; genitalia on acetate strip (BM). Synonymy with decens

by Edwards, 1911: 263.

Harbach: Subgenus Culex in Southwestern Asia 45

lividocostalis Graham, 1910: 269. +Lectotype male, hereby

designated, bearing following data: "Yaba / 3-7-09 / fr. larvae / No 16

/ WMG // Type // Yaba, / Lagos, / W. Africa. / 3. VII. 1909. / D.W.M.

Graham. / 1910-80."; genitalia on acetate strip (BM). Synonymy with

decens by Edwards, 1911: 263.

Culex decens Theobald of Edwards, 1912c: 381 (Africa, L key); Edwards, 1913b: 58

(Africa, M gen.).

Culex (Culex) decens Theobald of Edwards, 1941: 336, 425, 484 (Afrotropical Region,

A key, M*; F*; P*, syn., distr.); Lewis, 1943a: 282 (Eritrea, L bionomic note);

Lewis, 1943b: 72 (Sudan, A beh.); Lewis, 1945: 21 (Sudan, distr.); Abbott,

1948: 45 (Sudan, L habitat); Lewis, 1948: 144 (Sudan, L habitat); Hopkins, 1952:

320 (Afrotropical Region, L*, bionomics); Knight, 1953a: 229 (Yemen Arab

Republic, A, L, distr.); Lewis, 1956: 715 (Sudan, distr.); Mattingly and Knight,

1956: 110, 113, 120, 127-128, 132 (P.D.R. Yemen, Yemen Arab Republic, A, L

keys, distr., zoogeogr., bionomics); Mekuria, 1968: 78 (Ethiopia); Harbach,

1985a: 86, 94, 103 (P.D.R. Yemen, Yemen Arab Republic, distr., A, L key).

Adult. Culex decens is distinguished from all other species of the subgenus

Culex occurring in southwestern Asia by the following combination of characters: (1)

proboscis entirely dark-scaled, (2) scutal scales uniform golden brown with reddish tint,

(3) abdominal sterna with broad apical black bands, and (4) maxillary palpus of male

completely dark-scaled. Specimens examined from the Sudan and Uganda differ from

Arabian material in having finer scutal scales which are more evenly dispersed and darker

in color.

FEMALE. Head: Antennal length 1.6-2.0 mm, mean 1.8 mm; entirely dark.

Proboscis length 1.8-2.0 mm, mean 1.9 mm; entirely clothed in brownish-black scales,

ventral surface slightly paler; labella yellowish brown. Maxillary palpus entirely

dark-scaled; length about 0.4 mm, approximately 0.2 of proboscis length. Forked scales

of vertex dark; falcate scales pale yellow; lateral spatulate scales off-white; with 2 very

long interocular setae projecting downward over clypeus. Thorax: Integument brown,

pleura paler. Scutum mainly with golden-brown scales with reddish tint, with pale yellow

or whitish scales on anterior and lateral margins and prescutellar area (scutum of

Sudanese and Ugandan specimens with finer, more evenly dispersed, uniform

reddish-brown scales; without distinctly paler scales at margins); setae dark reddish

brown, prominent. Scutellum with pale yellow falcate scales on all 3 lobes; 3 or 4 large

dark setae on each lateral lobe, 5-8 on median lobe. Antepronotum with rather broad

pale yellow falcate scales and dark setae, setae longer anteriorly; postpronotum with

pale yellow to golden-brown falcate scales somewhat intermediate in form between

scutal and antepronotal scales, usually with 4 (sometimes more) well-developed setae in

row along posterior margin at level of mesothoracic spiracle. Pleura with creamy-white

spatulate scales as follows: patch below and mesal to upper proepisternal setae,

patches on upper corner and lower posterior border of mesokatepisternum, and anterior

and upper patches on mesepimeron, upper mesepimeral patch larger than in most

species, these scales being among and around the upper mesepimeral setae; numbers

of pleural setae: 7-11 proepisternal, 7-12 prealar, 5-9 upper mesokatepisternal, 10-14

lower mesokatepisternal, 7-13 upper mesepimeral, and 1 (rarely 2) lower mesepimeral.

Wing: Length 3.3-3.6 mm, mean 3.5 mm; length ofcell R2/length of vein R2+3 2.4-3.0,

mean 2.6; length of cell M1 0.8-1.0 length of cell R2; entirely dark-scaled. Halter: Pale

brown. Legs: anterior surface of forecoxa dark-scaled, dark scales continuous with

46 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

patch of creamy-white spatulate scales at base, coxa with few pale yellow spatulate

scales posterolaterally at apex; mid- and hindcoxae with longitudinal patch of

creamy-white spatulate scales on anterior side of lateral midline. Posterior surface of

trochanters with small, pale yellow spatulate scales. Femora with narrow pale knee

spots; forefemur dark-scaled anteriorly, pale-scaled posteriorly; midfemur dark-scaled

anteriorly and dorsally, pale-scaled posteroventrally; hindfemur largely pale-scaled, with

more or less complete dorsal stripe of dark scales that widens distally onto anterior and

posterior surfaces. Foretibia mainly dark-scaled, paler ventrally, with pale spot

anterodorsally at apex; midtibia also mainly dark, paler posteriorly, with spot of pale scales

dorsally at apex; hindtibia like foretibia but with prominent spot of pale scales at apex.

Tarsi entirely dark-scaled. Abdomen: Tergum | with posteromedial patch of brownish

scales; terga II-VII brown with narrow basal pale bands of off-white scales connected to

large lateral spots of same color, bands 0.01-0.25 tergum length; tergum VIII with narrow

basal pale band. Sterna II-VII with broad apical black bands; sternum VII with pale scales

laterally, without scales medially.

MALE. Differing from female as follows. Head: Maxillary palpus entirely dark;

palpomere 3 with 7-10 ventral setae at apex; palpomeres 4 and 5 densely setose. Fal-

cate scales of vertex less numerous and paler in color. Thorax: Scutal scales paler and

longer. Wing: Length 3.0-3.2 mm, mean 3.1 mm; length of cell R21.8-1.9 length of vein

R2+3; length of cell M1 0.7-0.9 length of cell R2; scales usually paler, particularly along

anterior veins. Abdomen: Terga essentially as in female, terga without basolateral pale

spots, basal bands produced posteriorly along lateral scale-free margins; tergum VIII

(ventral in position) mainly dark-scaled, with narrow basal pale band, posterior margin

slightly emarginate, densely setose, posterolateral setae very long. Apical dark bands

of sterna II-VII broader, more or less convex, proximal sterna with only basolateral corners

pale-scaled; sternum VIII (dorsal in position) usually nearly completely pale-scaled,

caudal margin narrowly dark-scaled, particularly in middle. Genitalia (Fig. 11): Ninth tergal

lobe small, with 9-17 setae (mode 10) in 1 or 2 irregular rows. Gonocoxite normal,

ventrolateral setae strongly developed, longer and stouter than lateral setae, mesal

surface with 4 irregular rows of small setae extending from base to level of subapical

lobe; subapical lobe indistinctly divided, setae a-f borne on slight prominence on mesal

side of main lobe; setae a-c rodlike, a shorter than b and c and more or less straight, b

and c each with stout base, tapered and slightly curved, D slightly stouter than the

others; 3 setae in group d-e, 2 simple, 1 flattened and resembling f, f longer than a-e,

flattened, slightly broader distally; g foliform, broad and strongly asymmetrical; h long,

slender, bent in middle. Gonostylus slender, curved, slightly widened before tip on

lateral side, tip rugose dorsally. Phallosome not much longer than broad, lateral plates

slightly longer than aedeagus; dorsal arm long, tip pointed and bent mesad, diverging

from its mate of the opposite side; lateral arm with 3-5 denticles (mode 4) in more or less

Straight line on mesal side of caudal margin, also with a prominent flat lateral lobe, a short

conical dorsal process, and a weak basal articulatory process; ventral arm a sinuous

mesal process with tip pointed and bent caudad. Aedeagus semispherical; ventral

aedeagal bridge narrow. Proctiger normal; paraproct with long curved basal lateral arm,

crown dark with numerous short spinelike spicules and some shorter simple blades.

Cercal sclerite elongate, irregular in shape; 3-5 cercal setae, usually 4. Tergum X

Straplike, joining base of paraproct below basal lateral arm.

Pupa (Fig. 11). Character and positions of setae as figured, range and modal

number of branches in Table 5; similar to pipiens and closely related species, the most

obvious distinctions being the form of the trumpet, which is long and slender, and the

character of seta 1-Il, which has at least 25 branches. Cephalothorax: Setae 3,4-CT

Harbach: Subgenus Culex in Southwestem Asia 47

usually triple, sometimes with 4 branches; 6-CT with 3 or 4 branches, more often with 4:

12-CT usually with 3 or 4 branches (3-5). Trumpet: Long and slender, index 5.5-9.1,

mean 7.1; tracheoid area about 0.4 trumpet length; pinna 0.15-0.25 trumpet length.

Abdomen: Length 2.6-2.9 mm, mean 2.8 mm. Seta 1-Il with 25 or more branches,

usually more (25-32); seta 5-IV of same order of thickness and length as 5-V,VI, with 3 or

4 branches, more often with 3; 6-IIl double or triple, more often double, 6-IV-VI usually

triple, sometimes with 4 branches, rarely double. Genital lobe: Rather short in male,

length about 0.33 mm. Paddle: Length 0.74-0.92 mm, mean 0.82 mm: width 0.48-0.60

mm, mean 0.57 mm; index 1.3-1.5, mean 1.4.

Larva (Fig. 12). Placement and character of setae as figured, range and modal

number of branches in Table 25; immediately distinguished from all other Culex in

southwestern Asia by the long, slender siphon and very long, single seta 1-IV,V. Head:

Length 0.70-0.81 mm, mean 0.76 mm; width 1.00-1.21 mm, mean 1.13 mm; lightly

tanned, darker behind eyes, dorsal apotome sometimes with small dark median spot

posterior to bases of seta 5-C, less frequently with a pair of dark sublateral spots before

median spot. Dorsomentum moderately tanned; usually with 8 teeth (6-8) on either side

of median tooth, lateral teeth longer. Seta 0-C well developed, distinct; 1-C lightly to

moderately tanned, tapered to sharp point, length less than 0.1 mm; 2-C absent; 5,6-C

double, 5-C seldom triple; 7-C usually with 6 or 7 branches (5-7); 14-C normally double

with strongly divergent branches (1-3). Antenna: Length 0.52-0.61 mm, mean 0.55

mm, lightly tanned, distal part darker, base with dark spot on mesal surface. Seta 1-A

with about 27 branches (21-32). Thorax: Seta 7-P triple. Seta 1-M usually single,

seldom double, short, about 1.5 length of 2-M; 9-M with 3 or 4 branches, more often

with 4. Seta 1-T usually single, sometimes double; 2-T normally double (1,2).

Abdomen: Seta 3-1,Vll double, 3-VII rarely triple; 6-1,I triple, 6-III,V,VI single; 6-IV normally

double (1-3); 1-III,VI developed as usual, 1-Ill either single, double or triple, 1-VI usually

triple, sometimes double; 1-IV,V strongly developed, single and very long, much longer

than length of 2 abdominal segments. Segment VIII: Comb with 43-65 scales, mean 52.

Siphon: Very long and slender, index 8.75-11.27, mean 9.96; lightly to moderately

tanned. Pecten on basal 0.2, with 11-16 spines, mode 14; larger spines with 4 or 5

denticles. Seta 1-S usually in 4 pairs (a fifth seta occasionally occurs on one or both

sides of siphon), usually double or triple (1-3), length about same as diameter of siphon

at point of attachment. Segment X: Saddle lightly to moderately tanned; length

0.36-0.48 mm, mean 0.39 mm; siphon/saddle index 5.43-7.20, mean 6.36. Seta 1-X

usually double, occasionally single. Anal papillae subequal, slightly shorter than saddle.

Systematics. | am recognizing decens as the nominotypical member of a

subgroup which includes invidiosus Theobald, 1901c, antennatus, ornatothoracis

Theobald, 1909, trifoliatus Edwards, 1914a, perfuscus Edwards, 1914a, perfidiosus

Edwards, 1914a, telesilla De Meillon and Lavoipierre, 1945 (in De Meillon et al., 1945)

and litwakae Harbach, 1985b. Except for antennatus, the members of this group are

restricted to the Afrotropical Region. The group as a whole is poorly known and it may

be necessary to divide it into two or more subgroups when the species are studied in

detail. The subgroup is recognized primarily on the basis of overt similarities in adult

ornamentation and the structure of the male genitalia. There appear to be some striking

differences in the larvae.

The occurrence of decens in southwestern Asia (the Yemen Arab Republic) was

first recorded by Knight (1953a) who noted that Yemenite material resembles the African

material examined by Edwards (1941) except that the ventral surface of the proboscis

and the scutal scaling are lighter in some specimens. | have noticed that the scutal

scales of some specimens are slightly coarser and without the distinct reddish tint found

48 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

in specimens from Sudan and Uganda.

The larva of decens is quite different from other Culex occurring in southwestern

Asia. It is easily recognized by the long, slender siphon and the long, single seta 1-IV,V.

Hopkins (1952) did not mention seta 1-IV,V in his description of decens so it is possible

that this character may serve to distinguish decens from invidiosus, a species which is

supposedly indistinguishable from decens in Africa. Seta 1-IV,V is similarly developed in

larvae of decens examined from several African countries. No specimens identified as

invidiosus were examined during this study.

Bionomics. Larvae of decens have been taken from ground waters, such as

pools, swamps, rock holes, ditches, and streams, and a variety of artificial containers,

including cement basins, tubs, drains, wells, troughs, and household pots. The early

reports of decens breeding in tree holes, cut bamboo, and crabholes probably refer to

another species. This species has been collected in association with /aticinctus and

Culex (Lutzia) tigripes De Grandpre and De Charmoy in the Yemen Arab Republic

(Knight, 1953a). Collection data seen during this study indicate that resting adults have

been collected in a house and an unused well. Lewis (1943b) observed that decens is

common in the Anglo-Egyptian Sudan, but is never seen biting man. Lumsden and van

Someren (1953) reported that a single female was collected during landing catches

made 11 m above the ground in riverain forest in Uganda. This species probably is not

involved in disease transmission.

Distribution. Culex decens is very widely distributed in the Afrotropical Region.

Material examined. 129 specimens. A total of 50 specimens (8 females, 9

males, 11 male genitalia, 9 pupal exuviae, 10 larval exuviae, and 3 fourth-instar larvae)

were examined from southwestern Asia — P.D.R. YEMEN: (Mukalla); YEMEN ARAB

REPUBLIC: (Ta’izz). An additional 79 specimens (17 females, 18 males, 15 male

genitalia, and 29 fourth-instar larvae) were examined from GHANA (type specimens of

nigrocostalis ), \VORY COAST, KENYA, LIBERIA, MAURITANIA, NIGERIA (type

specimens of decens and lividocostalis ), SENEGAL, SIERRA LEONE (including the

type series of masculus ), SOUTH AFRICA (the type male of minutus ), SUDAN,

UGANDA, and ZAIRE.

Culex (Culex) antennatus (Becker)

antennatus Becker, 1903: 68 (Anopheles ). Type specimen(s) (male):

Cairo, Egypt (ZM).

laurenti Newstead, 1907 (in Newstead et al., 1907: 24). tLectotype

male, hereby designated, bearing following data: "SYN- / TYPE //

COTYPE / [male symbol] // SCHOOL OF TROP. MED., /

LIVERPOOL UNIVERSITY / Culex laurenti. / Leopoldville, / Belgian

Congo, / December 1903. / Drs. Dutton & / Todd // SYNTYPE of /

Culex laurenti / Newstead / det. J. Chainey 1975"; genitalia on ace-

tate strip (BM). Synonymy with antennatus by Hopkins, 1936: 224.

Culex antennatus (Becker) of Lewis, 1945: 20 (Sudan, distr.); Theodor, 1952: 113

(Middle East, zoogeogr.); El-Said and Kenawy, 1983a (Egypt, distr.); Zimmerman

et al., 1985: 84 (Egypt, bionomics); Kitron and Pener, 1986 (Israel, L bionomics).

Culex (Culex) antennatus (Becker) of Edwards, 1941: 333, 425, 484 (Afrotropical

Region, A key, M’*; F*; P, distr.); Lewis, 1943b: 72 (Sudan, A beh.); Lewis, 1948:

145-148 (Sudan, A beh., L habitat); Hopkins, 1952: 320 (Afrotropical Region, L

Harbach: Subgenus Culex in Southwestern Asia 49

key, L, bionomics); Lewis, 1956: 715 (Sudan, distr., zoogeogr.); Ovazza et al.,

1956: 176 (Ethiopia); Senevet et al., 1957b: 92 (North Africa, L); Senevet and

Andarelli, 1959: 215 (North Africa, A, P, L keys, syn., M*, F, P*, L*, distr., biol., L

assoc.); DuBose and Curtin, 1965: 352, 354 (Mediterranean area, A, L keys):

Mekuria, 1968: 77 (Ethiopia, distr.); Lotfi, 1970: 401 (Iran); Margalit and Tahori,

1970b: 151 (Israel, coll. sites); Margalit and Tahori, 1974: 56 (Israel, coll. rec.);

Lotfi, 1976: 73, 76, 82 (Iran, L key, ecol. note, L*); El-Said and Kenawy, 1983b

(Egypt, distr.); Harbach, 1985a: 86, 94, 104 (Egypt, Iran, Israel, Egypt, distr., A, L

keys).

Culex (?) invidiosus Theobald of Gough, 1914: 135 (Egypt, bionomic note, coll. sites).

Culex invidiosus Theobald of Storey, 1918(1919): 100 (Egypt, A, L keys, A, bionomics).

Culex laurenti Newstead of Buxton, 1923: 317 (Israel, Jordan, coll. rec.); Kirkpatrick,

1924(1925): 369, 372 (Egypt, A, L keys); Salem, 1938: 27 (Sinai); Abdel-Malek,

1956: 105 (Sinai, L key).

Culex (Culex) laurenti Newstead of Edwards, 1921: 344 (Egypt, Israel, Jordan, A key,

tax., distr.); Seguy, 1924: 39, 190 (Egypt, Israel, Jordan, A key, M’*, F, tax., distr.,

syn.); Kirkpatrick, 1925: 129 (Egypt, A, P, L keys, M*, F, P*, L*, bionomics);

Edwards, 1926: 141 (Egypt, Israel, Jordan, A, L keys, A, distr.); Stackelberg,

1927: 164 (Middle East, M, F keys, A, M gen.*, distr.); Martini, 1931: 370 (Egypt,

Jordan, A, L keys, M’*, F, L", distr.); Monchadskii, 1951: 265 (Middle East, L key,

L*, bionomics, distr., syn.).

Adult. A small pale brown species easily recognized by the prominent lateral

pale stripes on abdominal terga VI and VII and lateral pale spots on the other terga. The

proboscis of the male bears a ventral cluster of setae near the middle. This species

superficially resembles pusillus Macquart of the subgenus Barraudius, but the

abdominal terga of the latter are darker, black or nearly so, and the proboscis of the male

lacks a midventral cluster of setae.

FEMALE. Head: Antenna blackish; pedicel with some tiny scales and/or setae

on mesal side. Proboscis dark-scaled dorsally, with paler scaling ventrally, particularly on

distal 0.5; length 1.6-1.8 mm, mean 1.7 mm. Maxillary palpus dark-scaled, darker at tip;

length approximately 0.2 proboscis length. Forked scales of vertex numerous, usually

dark; falcate scales pale yellowish white; lateral spatulate scales narrow, white. Thorax:

Scutum brown; uniformly clothed in fine, golden-brown falcate scales, with some pale

yellow or whitish scales on anterior promontory and prescutellar area. Scutellum with

very fine pale yellow falcate scales on lateral and median lobes. Pronotum brown:

antepronotum with fine pale yellow falcate scales; similar scales on postpronotum but

Slightly darker anteriorly. Pleura brownish yellow; numbers of pleural setae: 6-12 upper

proepisternal, 6-9 prealar, 4 or 5 upper mesokatepisternal, 5-9 lower mesokatepisternal,

5-8 upper mesepimeral, and 1 lower mesepimeral; patches of narrow cream-colored

Spatulate scales present as follows: small patch below upper proepisternal setae, upper

and lower patches on mesokatepisternum, and anterior and upper patches on

mesepimeron. Wing: Length 2.7-3.0 mm, mean 2.9 mm; infrequently with some pale

scales on posterior side of costa before humeral crossvein, otherwise entirely

dark-scaled; length of cell R2 approximately 4.5 length of vein R2+3; length of cell M1

about 0.8 length of cell R2. Halter: Pale, more or less same color as pleura. Legs:

Anterior surface of forecoxa with dark scales in middle, off-white scales at base and apex:

mid- and hindcoxa each with longitudinal patch of small cream-colored scales on anterior

side of lateral midline. Forefemur dark brown-scaled anteriorly, pale-scaled posteriorly,

with inconspicuous knee spot of yellow scales; midfemur same but dark brown scaling of

50 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

anterior surface gradually expanded over dorsal surface on distal 0.5; hindfemur mainly

pale-scaled, with dorsal line of dark scales (often faint proximally; widened distally,

particularly on distal 0.5) and narrow yellow knee spot (usually more distinct than on fore-

and midfemora). Foretibia dark brown to blackish-scaled dorsally, with off-white scales

ventrally; anterior surface of midtibia dark brown-scaled, posterior surface with yellowish

white scales; hindtibia same as foretibia but sometimes with faint apical spot of pale

scales. All tarsi dark-scaled, somewhat paler scaling ventrally. Abdomen: Tergum | with

inconspicuous median posterior patch of cream-colored scales; terga II-VII almost

entirely brown-scaled; terga II-V with small basolateral patches of cream-colored scales

which are usually not visible in dorsal aspect, those of tergum V sometimes partly visible;

lateral borders of terga VI and VII with stripes of cream-colored scales; tergum VIII largely

pale-scaled. Sterna with cream-colored scales; sternum VIII with lateral patches only,

median area without scales.

MALE. Like female except as follows. Head: Ventral surface of proboscis usually

entirely pale-scaled, conspicuously so on distal 0.5; with ventral cluster of setae on

proximal side of false joint; length 1.7-1.9 mm, mean 1.8 mm. Maxillary palpus dark,

palpomeres 4 and 5 darker; length about 1.3 proboscis length, extending beyond tip of

proboscis by length of palpomere 5; palpomere 3 with lateral patch of pale scales before

apex; ventral surface of palpomere 4 with spot of white scales at base and short line of

similar scales before apex, these sometimes joined; palpomere 5 noticeably longer than

4 (about 1.4 length of 4), with spot of white scales ventrally at base. Lateral spatulate

scales of head slightly smaller and more numerous. Thorax: Scutal scales slightly finer.

Wing: Length 2.5-3.0 mm, mean 2.8 mm; length of cell R2 about 1.6 length of vein

R2+3. Abdomen: Terga II and Ill normally entirely dark-scaled; tergum VIII (ventral in

position) and sternum VIII (dorsal in position) largely or completely with cream-colored

scales. Genitalia (Fig. 13): Form as figured; basic structure as in decens, differing chiefly

as follows. Ninth tergal lobe with 6-14 setae (mode 8). Subapical lobe of gonocoxite

with 3 setae in group d-e, 2 longer setae with apical series of 2 or 3 (usually 3) recurved

projections, shorter seta simple and slightly flattened; seta g much as in decens,

acuminate. Gonostylus broader distally. Phallosome longer than broad with lateral

plates slightly longer than aedeagus; dorsal arm slightly sinuous in lateral view; caudal

margin of lateral arm with tight row of 3-5 denticles (usually 4) projecting over prominent

lateral ridge, end of ridge differentiated as a conspicuous dorsal process with narrowed

and slightly bent apex, ventral arm without prominent flat lateral lobe, basal articulatory

process more strongly developed; ventral arm a prominent rounded ventrocaudal lobe

with a short dorsally directed hornlike projection, ventrocaudal surface with many minute

spinules, dorsal projection slightly bent caudad and resembling denticles of lateral arm in

dorsal view. Aedeagus conical; ventral aedeagal bridge slightly broader. Basal lateral

arm of paraproct shorter, flattened, and broad. Cercal sclerite usually with 2 setae (1-3)

on posterolateral corner.

Pupa (Fig. 13). Character and positions of setae as figured, range and modal

number of branches in Table 6; resembling pipiens in most respects, but generally

smaller and setae 3-VIl and 6-IIl with more branches; extremely similar to members of the

univittatus and simpsoni subgroups, from which it cannot be distinguished with

confidence. Cephalothorax: Seta 5-CT frequently with 5, 6, or 7 branches (4-8); 8-CT

frequently with 6 or 7 branches (4-9); 9-CT often with 3 branches (2-4); 12-CT often with

5 branches (2-5). Trumpet: Much as in pipiens, index 4.3-6.5, mean 5.2; tracheoid area

about 0.4 trumpet length; pinna oblique, 0.2-0.3 trumpet length. Abdomen: Length

2.6-3.3 mm, mean 2.9 mm. Seta 1-Il with considerable variation in number of branches

(7-41); 5-IV not developed quite as strongly as 5-V,VI, often with 5 branches (4-6); 6-Ill,

Harbach: Subgenus Culex in Southwestern Asia 51

IV most often with 3 or 4 branches (38-6), 6-V,VI most often with 5 or 6 branches (3-6).

Paddle: Length 0.80-1.01 mm, mean 0.90 mm; width 0.56-0.70 mm, mean 0.63 mm;

index 1.3-1.5, mean 1.4.

Larva (Fig. 14). Placement and form of setae as figured, range and modal

number of branches in Table 26; differing from pipiens and allied species and

resembling univittatus and perexiguus in having seta 5-C usually double or triple, seta

6-VI normally single, and seta 1-S no longer than diameter of siphon. Head: Length

0.64-0.75 mm, mean 0.67 mm; width 1.02-1.16 mm, mean 1.10 mm; tanning, including

spots on dorsal apotome, almost exactly as in pipiens. Dorsomentum usually with 8

teeth (7-9) on either side of median tooth. Seta 1-C long, slender, moderately tanned,

length about 0.1 mm; 2-C absent or represented by a small bump; 5,6,11-C double or

triple; 7-C with 6, 7, or 8 branches occurring in nearly equal frequencies; 10-C usually

triple, sometimes with 4 branches; 14-C single or double, branches strongly divergent

when double. Antenna: Length 0.44-0.58 mm, mean 0.50 mm; moderately tanned at

base and on distal part from just before seta 1-A; spicules rather broad, not true aciculae;

scape developed; puncture not evident. Seta 1-A with about 20 branches (17-28).

Thorax: Seta 7-P triple. Seta 1-M most often double, frequently triple, sometimes

single, very short, about same length as 2-M. Seta 1-T short, about same length as 1-M,

usually double or triple (1-3); 2-T double or triple. Abdomen: Segments III and V with

black pigment granules giving abdomen a 2-banded appearance to the unaided eye.

Seta 3-I usually triple (2-5), 3-VIl usually with 4 or 5 branches (3-5); 6-I,II normally triple, 6-II

occasionally with 4 branches, 6-IIl usually double (1-3), 6-IV triple, 6-V double or triple,

more often double, 6-VI single; 1-IIIl with 3 or 4 branches, 1-IV-VI usually triple, 1-IV,V

seldom with 4 branches, 1-V,VI sometimes double. Segment VI/l: Comb with 32-46

scales, mean 38. Siphon: Long, slender, slightly tapered; index 5.47-8.14, mean 6.78.

Pecten on approximately basal 0.25, with 10-14 spines, mode 13; larger spines with 2 or

3 basal denticles. Seta 1-S normally in 5 or 6 pairs, 2 or 3 posterolateral and 3 lateral,

length about 0.5 diameter of siphon at point of attachment. Segment X: Saddle length

0.36-0.40 mm, mean 0.38 mm; siphon/saddle index 3.77-4.99, mean 4.35. Seta 1-X

with 3 or 4 branches, more often with 4; seta 2-X double or triple, more often double.

Anal papillae subacutely tapered, about length of saddle.

Systematics. Culex antennatus is an interesting species which shares some

features with the adults of both the pipiens and the decens subgroups. It is included as

a member of the decens subgroup because the male genitalia suggest a closer affinity

with decens than pipiens. The larva of this species bears a close resemblance to those

of univittatus and perexiguus which suggests a relationship with these species. It is

possible, therefore, that antennatus has retained some generalized features of an

ancestral stock which may have given rise to the pipiens, decens and univittatus

subgroups. There is some suggestion of a distant relationship with members of the

sitiens group in the structure of the male phallosome. Once the Culex fauna of Africa

has been studied thoroughly and species affinities are more clearly understood, it may

be necessary to include antennatus and the closely related litwakae Harbach, 1985b, in

a separate subgroup.

The adults of antennatus are superficially very similar to those of Culex

(Barraudius ) pusillus Macquart, 1850, but are easily distinguished by the smaller

forefemur/proboscis ratio and the much longer hindtarsomere 1. These species are

sympatric thoughout the range of antennatus in southwestern Asia and Egypt. Females

of antennatus are almost indistinguishable from unbanded females of pipiens, but the

males, larvae, and pupae of these species are easily differentiated. The larva of

antennatus is extremely similar to that of perexiguus which usually, but not always, dif-

Pd Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

fers in the character of setae 1-C, 1-M, and 1-S. Live larvae of antennatus are readily

distinguished from those of perexiguus by the two-banded appearance of the

abdomen.

Bionomics. Culex antennatus mainly breeds in stagnant bodies of fresh water,

but is sometimes found in brackish water pools. It rarely occurs in polluted water.

Common larval habitats include rice fields, stream pools, springs, ponds, swamps,

ditches, seepages, and animal footprints. The breeding places usually contain

emergent vegetation. Larvae are often found in association with theileri, poicilipes,

perexiguus, and pipiens. Females feed on mammals. They appear to feed primarily on

bovids, but also attack sheep, goats, horses, donkeys, and man (Zimmerman et al.,

1985). The species bites both indoors and outdoors in the evening, and seeks resting

sites away from buildings (Lewis, 1948; Zimmerman et al., 1985). Culex antennatus has

been found naturally infected with West Nile virus in Egypt (Taylor et al., 1953; Taylor et

al., 1956) and Rift Valley Fever virus in Nigeria (Lee, 1979). Successful experimental

transmission has been demonstrated for both of these viruses in the laboratory (Taylor

et al., 1953; Gad et al., 1987, respectively). Arumowat virus was isolated from a pool of

unfed females in the Sudan (Karabatsos, 1985), and there is serological evidence for

human infection of this virus in Egypt, Sudan, and Somalia (Tesh et al., 1976). Immature

filarial infections have been found in this species in Tanzania (Smith, 1955) and Egypt

(A. Gad, personal communication).

Distribution. This species occurs in the Middle East and the Afrotropical

Region south to Botswana.

Material examined. 885 specimens. A total of 849 specimens (211 females,

175 males, 13 male genitalia, 310 pupal exuviae, 112 larval exuviae, and 28 fourth-instar

larvae) were examined from southwestern Asia — EGYPT: Aswan (Aswan, El Aqaba el

Saghira, Ezbet el Silsila, Knour Abu Subeira, Nag el Kagug, Nag el Ritag el Qibli), Asyut

(Beni Shigeir, El Qusiya), E/ Fayyum (Abhit el Haggar, Birket Qarum, Biyahmu, El Nazla,

Sinero el Bahria, Tubhar, unknown localities), E/ Foadia (Baltim, Nimra el Basal), E/

Gharbiya (Saft Turab), E/ Giza (Ezbet Bosna Sharawy, Harania, Koniessa, Nazlet el

Ashtar, Talbia), E/ Isa’ iliya (Abu Khalifa, Nifisha), E/ Qahira (Cairo), E/ Qalyubiya (Bahtim,

El Salmaniya, Ga'afara, Kaha, Kafr Hamza, Sindiwa), E/ Shargiya (Inshas el Ram,

Khirbetrama), Kafr el Sheikh (Kafr el Sheikh, Makallet el Qasab); IRAN: (Bisotun,

Kamroud, Kernianshah); IRAQ: (unknown localities); ISRAEL: (Lake Tiberias, Ludd,

Rosh Hal Ayn, many unknown localities); JORDAN: (Az Zarqa, unknown localities);

TURKEY: (unknown localities). An additional 36 specimens (18 females, 7 males, 9

male genitalia,1 pupal exuviae, and 1 larval exuviae) were examined from ALGERIA,

MAURITANIA, SENEGAL, SUDAN, TUNISIA, ZAIRE (type specimens of /aurenti), and an

unknown country.

Culex (Culex) univittatus Theobald

univittatus Theobald, 1901b: 29. +Lectotype female: Salisbury, Zim-

babwe; designated by White, 1975: 321 (BM).

simplex Theobald, 1903a: 337 (Heptaphlebomyia ). +tHolotype female:

Salisbury, Rhodesia (BM). Synonymy with univittatus by Edwards,

1911: 262.

montforti Ventrillon, 1905a: 448 (Heptaphlebomyia ). tLectotype male:

Andajobé, Madagascar; designated by White, 1975: 321 (BM).

Synonymy with univittatus by Edwards, 1911: 262.

Harbach: Subgenus Culex in Southwestern Asia 53

ataeniatus Theobald, 1911: 261. *Holotype female: Onderstepoort,

Transvaal (BM). Synonymy with univittatus by White, 1980: 137.

Culex univittatus Theobald of Edwards, 1912b: 28, 32 (in part; Afrotropical Region, A

key, distr.); Edwards, 1912c: 381 (in part; Africa, L key); Edwards, 1913b: 58

(inpart; Africa, A, M gen.); Edwards, 1914a: 67 (Africa, M gen.*); Mattingly, 1954:

56 (typical form, in part; Africa, zoogeogr., M gen.”*).

Culex (Culex) univittatus Theobald of Séguy, 1924: 39, 190 (in part; Africa, tax., M

gen.*, distr.); Edwards, 1941: 306, 419, 482 (typical form, in part; A key, M*, F’,

P, syn., distr.); Lewis, 1943a: 282 (in part; Eritrea, coll. rec.); Hopkins, 1952: 291

(? in part; Afrotropical Region, L); Mattingly and Knight, 1956: 104, 122 (in part:

Yemen Arab Republic, A, L keys); Jupp, 1970: 9-16 (in part, Highveld form;

south Africa, M*, F, distr., crossmatings, beh.); Jupp, 1971: 339-356 (South

Africa, M gen.*, F*, distr., crossmatings); Jupp, 1972 (excluding short-spined

form; Africa, M gen.*, F); White, 1975: 320-322 (Afrotropical Region, syn., distr.,

A key); Harbach, 1985a: 86, 93, 104 (P.D.R. Yemen, Yemen Arab Republic,

distr., A, L keys).

Culex (Culex) univittatus var. neavei Theobald of Knight, 1953a: 232 (Yemen Arab

Republic, coll. rec.).

The description of this species is based largely on the examination of topotypic

material and the type series. The occurrence of univittatus in southwestern Asia is

based on the examination of a single male from the P.D. R. Yemen and a single female

from the Yemen Arab Republic.

Adult. This species might be confused with sinaiticus or simpsoni, but differs in

the character of the scutal scaling and by the presence of an anterior pale stripe on the

hindtibia. It is closely allied to perexiguus from which it is usually distinguished by the

presence of a complete anterior pale stripe on the midfemur, and a line of scales on vein

2A in the female, and slightly different genitalia in the male.

FEMALE. Head: Length of antenna 1.5-1.9 mm, mean 1.7 mm; pedicel pale

laterally, dark mesally with patch of white scales; flagellum dark, flagellomere 1 with some

white scales mesally just above base, most flagellar whorls with 4 long dark setae.

Proboscis black-scaled, with whitish scales ventrally in middle; length 1.8-2.3 mm, mean

2.0 mm. Maxillary palpus black-scaled, with some white scales mesally at apex,

palpomeres 3 and 4 often with some white scales laterally; length about 0.2 proboscis

length. Falcate scales of vertex pale golden brown to white, usually paler medially;

interocular scales concolorous with white or whitish ocular scales; forked scales

cream-colored medially, golden brown to brown posterolaterally; lateral spatulate scales

white. Thorax: Integument brown. Scutal setae golden brown to brown; scutal scales

narrow, somewhat dense, golden brown; some pale yellow to white scales on margins

and prescutellar area, pale scales often forming pair of faint to distinct submedian spots

near middle of scutum. Scutellar scales same color as prescutellar scales. Falcate scales

of antepronotum golden brown medially, paler and slightly coarser dorsally and

especially ventrally, color of setae same as scales; postpronotum with narrow,

golden-brown scales becoming paler posteriorly, with 4 or 5 long dark setae along

posterodorsal margin. Pleural setae golden: 6 or 7 upper proepisternal, 10 or 11

prealar, 4-6 upper mesokatepisternal, 7-10 lower mesokatepisternal, 7 or 8 upper

mesepimeral, and 1 lower mesepimeral. Pleura with broad white spatulate scales: small

patch below upper proepisternal setae extending well mesad of setae on anterior

surface, patch of narrower scales on upper portion of postspiracular area sometimes se-

54 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

parated into anterior and posterior clusters, small patch below prealar setae continuous

with upper mesokatepisternal scales, large patches on upper corner and lower posterior

border of mesokatepisternum, large anterior patch on mesepimeron at level of upper

mesokatepisternal scales and patch before and among upper mesepimeral setae. Wing

(Fig. 2D): Length 3.6-4.1 mm, mean 3.8 mm; length of cell R2/length of vein R2+3

2.7-3.8, mean 3.3; length of cell M1/length of cell R2 0.75-0.79, mean 0.78; scales

entirely dark except for short line of white scales on posterior margin of costa at level of

humeral crossvein; vein 2A usually with scales on dorsal surface, scales usually

extending entire length of vein. Ha/ter: Pedicel and scabellum whitish; capitellum

blackish. Legs (mid- and hindlegs, Fig. 2!1,K): Mainly black-scaled. Anterior surface of

forecoxa with white scales at base and apex and black scales in middle, with many long

golden-brown to brown setae among median black and distal white scales, with 3 or 4

shorter setae and few pale scales posteriorly at apex; midcoxa with midlateral row of 5 or

6 well developed golden-brown to brown setae margined anteriorly by longitudinal patch

of white spatulate scales, with small patch of black scales and several short ventrally

projecting setae anteriorly at apex; hindcoxa with posterolateral row of normally 8 long

golden to golden-brown setae, anterolateral surface with longitudinal patch of white

spatulate scales and several short ventrally projecting setae at apex. Trochanters with

white scales on posterior and dorsal surfaces. Each femur with narrow pale knee spot;

anterior surface of forefemur black-scaled, usually with complete or incomplete narrow

ill-defined or broken anteroventral white stripe, posterior surface entirely white-scaled;

midfemur like forefemur but with complete distinct or ill-defined anteroventral white

stripe and black scales extending over dorsal surface toward apex; hindfemur largely

white-scaled, with anterodorsal stripe of black scales beginning beyond base and

widening distally, stripe abruptly expanded over anterior and posterior surfaces on distal

0.2 to form subapical band. Anterior and dorsal surfaces of foretibia black-scaled,

posterior and ventral surfaces whitish-scaled; frequently with small dorsal whitish spot at

apex; midtibia mainly black-scaled, whitish-scaled posteriorly, usually with narrow

anterodorsal stripe of whitish scales that usually reaches small dorsal whitish spot at

apex; hindtibia black-scaled with narrow anterior and posterior whitish stripes on proximal

0.8, with distinct white spot at apex. Tarsi black-scaled, tarsomeres 1 and 2 usually

noticeably paler ventrally. Abdomen: Terga mainly black-scaled; tergum | with median

posterior patch of black scales usually bordered anteriorly by some whitish scales; terga

II-VIl with convex basal bands and basolateral spots of white scales, bands normally

0.25-0.35 tergum length; bands usually not reaching spots on tergum Il, barely touching

spots on terga II-IV and joining them on terga V-VII, spots progressively larger from

anterior to posteror terga; tergum VIII frequently with posterior band of black scales, but

often entirely whitish-scaled. Sterna II-VII mainly with cream-colored to white scales,

usually with black scales on midline and posterolateral corners; sternum VIII with dark

integument and lateral patches of white scales.

MALE. Differing from the female as follows. Head: Proboscis usually entirely

black-scaled, occasionally with some whitish scales at false joint, without ventral cluster

of setae at false joint. Maxillary palpus mainly dark; midregion of palpomere 3 with narrow

ventrolateral line of dingy-white scales, with few whitish scales ventromesally at apex;

palpomere 4 with ventral white stripe narrowing to apex, also with indistinct or faint

dorsolateral line of whitish scales; palpomere 5 with ventral spot of white scales at base.

Falcate scales of vertex coarser, forked scales shorter. Thorax: Submedial spots of

scutum extremely faint or absent. Wing: Length 3.1-3.6 mm, mean 3.3 mm; length of

cell R2/length of vein R2+3 1.6-2.8, mean 2.1; basal line of white scales on costa ex-

Harbach: Subgenus Culex in Southwestern Asia 55

tremely reduced or absent; vein 2A usually without scales. Legs: Anterior white stripe

of midfemur sometimes less distinct. Abdomen: Basal white bands of terga generally

broader, less convex; basolateral spots absent; terga V-VII with posterior extension of

bands along lateral scale-free margins. Black scaling of sterna more extensive, pale

scaling often reduced to large basolateral spots; sternum VIII (dorsal in position) with

basal pale band or entirely pale-scaled. Genitalia (Fig. 15): Form as figured; lateral plate

of phallosome generally constructed as in vagans and torrentium. Ninth tergal lobe

small, with 6-16 (mode 11) setae. Gonocoxite normal; ventrolateral and mesal surfaces

with usual complement of large and small setae respectively, lateral surface with 1 or 2

rows of moderately long slender setae and several rows of small setae between these

and the large ventrolateral setae; subapical lobe slightly divided, proximal part

prominent, bearing setae a-c of usual form, b slightly stouter than a and c; setae dand e

fine, hairlike and inconspicuous, shorter than f, fa blunt rod with slightly enlarged tip; g

foliform as usual, asymmetrical, longer than broad, apex not sharply produced; h

slender, bent in middle. Gonostylus slightly expanded laterally near tip. Phallosome

longer than broad, aedeagal sclerite shorter than lateral plate; dorsal arm of lateral plate

rather long, slender, and slightly sinuous in dorsal view; lateral arm strongly developed,

with conspicuous angled scooplike caudal extremity and prominent dorsal process;

ventral arm spinelike, curved dorsad, and projecting slightly beyond caudal margin of

lateral arm. Proctiger normal; paraproct with long, curved, distally flattened basal lateral

arm, crown mostly of small spinelike spicules but with some inconspicuous lateral blades.

Cercal sclerite and tergum X unmodified; 1-3 cercal setae, usually 2.

Pupa (Fig. 15). Not associated with adults from southwestern Asia, specimens

examined primarily from South Africa; character and placement of setae as figured, range

and modal number of branches in Table 7; not immediately distinguishable from pipiens

and related species, differing chiefly in the number of branches of setae 6-IIl and 3-VIl,

resembling theileri, perexiguus, antennatus, simpsoni, and sinaiticus in this respect.

Cephalothorax: Seta 5-CT rather short, about same length and thickness as 4-CT; 6-CT

with 3-5 branches occurring in nearly equal frequencies; 9-CT double or triple; 11-CT

single or double; 12-CT triple. Trumpet: Generally longer than in pipiens, index 5.6-7.3,

mean 6.4; pinna shorter than tracheoid area, usually about 0.25-0.30 trumpet length;

tracheoid area normally about 0.35-0.40 trumpet length. Abdomen: Length 2.7-3.2

mm, mean 3.0 mm; seta 6-III normally with 3 or 4 branches, rarely double, 6-IV-VI most

often with 4 branches (3-5); 5-IV very nearly same length and thickness as 5-V,VI, with 5

or 6 branches, 5-V with 3 or 4 branches, 5-VI double or triple; 3-VIl most often with 4

branches (3-5). Paddle: Length 0.82-0.94 mm, mean 0.88 mm; width 0.56-0.71 mm,

mean 0.63 mm; index 1.3-1.5, mean 1.4; outer margin with tiny serrations on proximal

0.8; |

Larva (Fig. 16). Not associated with adults from southwestern Asia, specimens

examined include topotypic material; form and positions of setae as figured, see Table

2/7 for range and modal number of branches; very closely resembling antennatus, but

usually distinguished by the combination of shorter seta 1-C, siphon with 2 lateral pairs

of seta 1-S, seta 1-M usually single, and abdomen without obvious 2-banded

appearance in life. Head: Length 0.69-0.84 mm, mean 0.76 mm; width 1.02-1.24 mm,

mean 1.15 mm; lightly tanned, darker behind eyes. Dorsomentum usually with 7 teeth

(6-8) on either side of median tooth. Seta 1-C normally shorter than in antennatus,

tapered, not heavily tanned; 2-C occasionally present; 5,6,11-C as in antennatus but

5-C occasionally with 4 branches. Antenna: Length 0.50-0.64 mm, mean 0.57 mm;

lightly tanned, distal part slightly darker, dark on mesal surface at base. Seta 1-A with

about 27 branches (17-30). Thorax: Integument hyaline; with rows of vesicles on ventral

56 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

surface. Seta 7-P almost always triple, seldom double; 8-P normally double, rarely triple.

Seta 1-M single or double, more often single, not much longer than 2-M. Seta 2-T most

often triple (2-4). Abdomen: Ventral surface with rows of vesicles; segments Ill and V

without conspicuous pigmentation. Chaetotaxy resembles that of antennatus in nearly

all respects; seta 6-VI usually single, occasionally double. Segment VIII: Comb with

32-55 scales, mean 43. Siphon: More lightly tanned and generally slightly shorter than

in antennatus; index 4.93-7.15, mean 5.72. Pecten with 8-15 spines, mode 11; larger

spines with 3 or 4 basal denticles. Seta 1-S in 5 or 6 pairs, la-S (when present) usually

borne within pecten, 2 pairs (Id, le) borne laterally; length distinctly less than diameter of

siphon, usually 0.5 or less diameter at point of attachment; branching variable, distal

elements usually with more branches than proximal elements (see table). Segment x:

Saddle length 0.37-0.42 mm, mean 0.40 mm; siphon/saddle index 3.56-4.30, mean

3.82. Seta 1-X most often with 3 or 4 branches, occasionally double; 2-X usually double,

sometimes triple.

Systematics. Culex univittatus is a member of a very interesting species

aggregate. A thorough comparative study of this group has not been made yet but it

appears from a superficial examination of available material from Africa that three nominal

taxa and a possible fourth species are involved. The group includes univittatus, which

appears to be restricted to the temperate highlands of southern and eastern Africa and

the southwestern corner of the Arabian Peninsula; neavei Theobald, 1906, which

seems to occur throughout the subtropical and tropical lowlands south of the Sahara;

perexiguus, which inhabits the arid areas of northern and eastern Africa and

southwestern Asia; and another species which occurs in the evergreen rain forest of the

Congo Basin. Jupp (1972) identified specimens of univittatus from Benin, Niger, and

Upper Volta, but | do not believe that this species occurs in western Africa. These

records probably refer to neave/ or an undescribed species.

The members of the univittatus aggregate are remarkably similar in all life stages.

The three nominal forms are characterized by slight and variable differences in adult

ornamentation and features of the male genitalia. These characters are contrasted for

univittatus, perexiguus, and neavei in Table 41. Some of these were noted previously

(Jupp, 1970, 1971, 1972) while others are noted here for the first time. No definite

differences have been found in the immatures, but the larva of univittatus, based on the

examination of topotypic material, appears to be separable from perexiguus and neavei

by the development of seta 1-S. It is hoped that this information will provide a starting

point for a critical objective study of the group throughout its range. Few specimens are

currently available from most areas, and there are very few individual rearings from

anywhere other than Egypt. A comprehensive study of the group must include

extensive sampling throughout Africa and southwestern Asia.

The occurrence of univittatus in southwestern Asia is based on the examination

of a male and a female from the Yemen Arab Republic and the P.D.R. Yemen,

respectively. These are undoubtedly the specimens which Mattingly and Knight (1956)

recorded from the Yemen Arab Republic, the male erroneously. These specimens

agree well with the concept of univittatus presented here.

A very striking feature of univittatus is the presence of a line of scales marking the

position of vein 2A. To my knowledge, this character is unique among mosquitoes and

appears to be a primitive feature in the family Culicidae. Culex univittatus appears to be

somewhat of a relict species.

Bionomics. Very little is known about the biology of univittatus, partly because

it has been confused with neave/ in the past. The species seems to be fairly abundant in

more temperate areas of the Afrotropical Region. Larvae are found in fresh water

Harbach: Subgenus Culex in Southwestern Asia 5/

ground pools, marshes, ditches, swamp margins, stream margins, and cattle hoofprints

under some degree of shade. Females feed primarily on birds (Anderson, 1967), but

will enter houses and bite humans (Jupp and Mcintosh, 1967). Sindbis, West Nile, and

Wesselsbron viruses have been isolated from wild-caught specimens in South Africa

(Weinbren et al., 1956; McIntosh et al., 1967; Worth and De Meillon, 1960:

respectively). Mcintosh et al. (1967) regard univittatus as the primary feral vector of both

Sindbis and West Nile infections in that country.

Distribution. Culex univittatus is generally distributed in the temperate

highlands of the Afrotropical Region.

Material examined. 306 specimens. Only 3 specimens (1 F, 1M, and 1M

genitalia) were examined from southwestern Asia — P.D.R. YEMEN: (Mukaira); YEMEN

ARAB REPUBLIC: (Usaifira). A total of 303 specimens (110 females, 76 males, 41 male

genitalia, 6 pupal exuviae, 3 larval exuviae, and 67 fourth-instar larvae) were examined

from ANGOLA, ETHIOPIA, KENYA, MADAGASCAR (the type series of montforti ),

SOUTH AFRICA (the holotype female of ataeniatus ), TANZANIA, ZAIRE, and

ZIMBABWE (including the type series of univittatus and the holotype female of simplex).

A total of 253 specimens (111 females, 52 males, 44 male genitalia, 14 pupal

exuviae, 18 larval exuviae, and 14 fourth-instar larvae) of Culex neavei from the

following countries were examined and compared with univittatus and perexiguus

during this study: BENIN, ETHIOPIA, GAMBIA, GHANA, KENYA, MALAWI,

MOZAMBIQUE, NIGER, NIGERIA, SOUTH AFRICA, SUDAN (including the type series of

neavei), TANZANIA, UGANDA, UPPER VOLTA, ZAIRE, and uncertain locations.

Culex (Culex) perexiguus Theobald

perexiguus Theobald, 1903a: 199. +Lectotype male: Sidon, Palestine

[Lebanon]; designated by White, 1975: 318 (BM).

Culex decens Theobald of Storey, 1918(1919): 100 (Egypt, A, L keys, A); ? Abul-hab,

1968: 249 (Iraq, L key).

Culex (Culex) decens Theobald of ? Abdel-Malek, 1960: 113-118 (Syria, L bionomics,

distr.); Ibrahim et al., 1983: 91 (Iraq, L key, L”*).

Culex pallidocephalus Theobald of Gough, 1914: 135 (Egypt, coll. sites).

Culex perexiguus Theobald of Theobald, 1905a: 28 (Israel, Jordan); Edwards, 1922:

278 (Oriental Region, A key); Buxton, 1923: 316 (Israel, Jordan, coll. rec., A, L

bionomics); Theodor, 1923: 344 (Israel, Jordan, P*, key); Barraud, 1924b: 1263

(India, A key, M’, F, distr.); Barraud, 1924c: 431 (India, L*); Kirkpatrick,

1924(1925): 368, 372 (Egypt, A, L keys).

Culex (Culex) perexiguus Theobald of Edwards, 1921: 342 (Egypt, Israel, Jordan,

Oman, A, L keys, L*, tax., distr.); Séguy, 1924: 38, 190 (Lebanon, Egypt, A, L

keys, L", tax., distr., syn.); Kirkpatrick, 1925: 125, 191 (Egypt, A, P, L keys, M’, F,

P", L*, bionomics, distr.); Stackelberg, 1927: 163 (Middle East, A key, A, M gen.*,

distr.); Senevet and Andarelli, 1954: 63 (Egypt, A key, M, F); White, 1975: 318,

322 (Africa, SW Asia, A key); Harbach, 1985a: 86, 93, 104 (SW Asia, Egypt, distr.,

A, L keys).

Culex univittatus Theobald of Barraud, 1921: 394 (Israel, Jordan, Egypt, L habitat,

distr.); Parr, 1943: 247-250 (Syria, Lebanon, A, P, L keys, bionomics); Theodor,

1952: 113 (Middle East, zoogeogr.); Gad, 1956: 135 (Egypt, distr., bionomic

note); Senevet et al., 1957a: 86 (North Africa, L*); Nielsen and Nielsen, 1958:

58 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

311 (Iraq, swarming beh.); Margalit and Tahori, 1970a: 142 (Israel, bionomics);

Margalit et al., 1971: 323 (Israel, bionomics); Margalit et al., 1973: 32 (Israel, coll.

rec.); El-Said and Kenawy, 1983a (Egypt, distr.); Kitron and Pener, 1986 (Israel, L

bionomics).

Culex univittatus form perexiguus Theobald of Mattingly, 1954: 56 (eastern

Mediterranean form, zoogeogr.).

Culex (Culex) univittatus Theobald of Edwards, 1926: 138 (Egypt, Israel, Jordan, Oman,

A, L keys, syn., A, L", distr.); Martini, 1931: 389 (Turkey, Oman, A, L keys, M’, F,

L", distr.); Barraud, 1934: 418 (India, A, L keys, M*, F, L*, L habitat, distr.); Lewis,

1943b: 72 (in part; bionomic note); Senevet, 1947a: 126 (Turkey, Egypt, L*,

distr., L habitat, bionomics, L key); Abbott, 1948: 45 (Sudan, L); Senevet, 1949:

64 (North Africa, M gen.*, key); Monchadskii, 1951: 279 (Middle East, L*, key,

bionomics, distr., syn.); Hopkins, 1952: 291 (in part; Afrotropical Region, L*, key,

bionomics); Abdel-Malek, 1956: 101 (Sinai, L bionomics, L key); Lewis, 1956,

708 (at least in part; Sudan, coll. sites); Mattingly and Knight, 1956: 104, 122 (in

part; Oman, A, L keys); Senevet et al., 1957b: 92 (North Africa, L); Parrish, 1959:

266 (Turkey, distr.); Abdel-Malek, 1960: 113-125 (Syria, coll. sites, L bionomics);

DuBose and Curtin, 1965: 252, 254 (Mediterranean area, A, L keys); Abul-hab,

1968: 246 (Iraq, distr., L key); Lotfi, 1970: 402 (Iran, coll. rec.); Margalit and

Tahori, 1970b: 153 (Israel); Aslamkhan, 1971: 155 (Pakistan); Jupp, 1972:

105-113 (short-spined form, M gen.*, F, distr.); Ward, 1972: 96 (Afghanistan);

Lotfi, 1973: 207 (Iran, coll. sites); Gutsevich et al., 1974: 385 (Middle East, A, L

keys, A, M gen.*, L*, L bionomics, med. imp.); Margalit and Tahori, 1974: 88

(Israel, coll. rec.); Lotfi, 1976: 73, 76, 82 (L key, ecol. note, L*); Sirivanakarn,

1976: 48 (Oriental Region, A, P, L keys, M*, F*, P, L*, tax., distr., bionomics);

El-Said and Kenawy, 1983b (Egypt, coll. rec.); Danilov, 1985a: 71 (Afghanistan, F

key); Danilov, 1985b: 55 (Afghanistan, L key).

Culex (Culex) univittatus form perexiguus Theobald of Lewis, 1956: 708 (Sudan, distr.);

senevet and Andarelli, 1959: 212 (North Africa, Mediterranean basin, A, L keys,

syn., A distr., bionomics).

Culex (Culex) univittatus var. perexiguus Theobald of Mattingly and Knight, 1956: 104,

135 (Oman, distr., zoogeogr.); Ovazza et al., 1956: 171 (Ethiopia, M gen.*, coll.

sites); Mekuria et al., 1978: 80 (Ethiopia, distr.).

Adult. This common and widespread species closely resembles univittatus in

almost all respects, but the midfemur lacks a complete anterior pale stripe, vein 2A is

normally without scales, and the ventral arm of the male phallosome is short.

FEMALE. Head: Antenna dark, length 1.4-1.6 mm; pedicel with patch of pale

scales on dorsomesal surface; flagellomere 1 with few pale scales on mesal surface near

base. Proboscis dark-scaled, pale-scaled ventrally except at base, weakly pale on distal

0.25; length 1.7-1.8 mm. Maxillary palpus dark-scaled; palpomere 4 with pale scales

dorsomesally at apex, palpomeres 3 and 4 often with some lateral pale scales; length

0.3-0.4 mm, about 0.2 proboscis length. Falcate scales of vertex narrow, pale ochreous

to whitish, those of ocular line off-white and concolorous with interocular scales; forked

scales pale yellow medially, becoming brownish laterally; lateral spatulate scales white.

Thorax: \Integument brown. Scutal scales narrow, somewhat dense, golden brown;

scales very pale yellow or whitish along extreme margins and around margin of

prescutellar area; pair of submedial, very faint to distinct pale spots on middle of scutum;

occasionally with trace of median anterior pale stripe. Falcate scales of antepronotum

pale yellow, paler ventrally; postpronotum with golden-brown falcate scales. Pleural

Harbach: Subgenus Culex in Southwestern Asia 59

setae golden: 6-8 proepisternal, 12-16 prealar, 4-6 upper mesokatepisternal, 7-11

lower mesokatepisternal, 9 or 10 upper mesepimeral, and 1 (rarely 2) lower mesepimeral.

Pleura with white spatulate scales as follows: small patch below upper proepisternal

setae extending mesad onto anterior surface above level of base of coxa; postspiracular

area with patch on dorsal 0.5; posterior anepisternum with small patch below prealar

knob, usually more or less contiguous with upper katepisternal scales; katepisternum

with patches on upper corner and posterior border above coxa, extensive but well

separated; anterior and upper patches on anepimeron. Wing: Length 3.0-3.5 mm,

mean 3.2 mm, length of cell R2 2.2-2.8 length of vein R2+3, mean 2.4; length of cell

M1 0.7-0.8 length of cell R2; dark-scaled with short line of dorsal pale scales on

posterior margin of costa at humeral crossvein; rarely with few pale scales at extreme

base of R; vein 2A occasionally with few scales distally, rarely with scales on entire

length. Halter: Pedicel and scabellum pale yellow; capitellum brown. Legs (hindleg,

Fig. 2K): Anterior surface of forecoxa with white spatulate scales at base and mixed pale

and dark scales distally, dark scales mostly in middle; mid- and hindcoxae with vertical

patch of white spatulate scales on anterior side of lateral midline. Each femur with narrow

pale knee spot; forefemur dark-scaled anteriorly, pale-scaled posteriorly, sometimes with

trace of anteroventral pale stripe; midfemur dark-scaled anteriorly, dark scales

progressively expanding over dorsal surface toward apex, with or without narrow

incomplete anteroventral longitudinal pale stripe, stripe faint, broken or distinct when

present; hindfemur pale-scaled with dorsal dark stripe usually beginning about 0.3 from

base and broadening apically. Anterior and dorsal surfaces of foretibia dark-scaled,

posterior and ventral surfaces pale-scaled; midtibia dark-scaled with anterodorsal and

posteroventral pale stripes; hindtibia dark-scaled dorsally, with broad anterior and ventral

pale stripes on proximal 0.8, stripes joined ventrally for at least part of their length distally,

separated on proximal 0.5 or less by narrow, often faint, distally diminished, dark stripe,

with distinct apical pale spot. Tarsi dark-scaled dorsally, pale-scaled ventrally; with very

faint indication of paler scales at apices of tarsomeres 1-4. Abdomen: Tergum | with

median posterior scale patch usually comprised of pale and dark scales; terga II-VIII with

slightly convex basal white bands, bands 0.20-0.35 length of terga and continuous with

elongate basolateral patches of same color. Sterna usually completely pale-scaled,

occasionally with dark scales on posterolateral corners, less frequently also with median

patch of dark scales.

MALE. Differing from female as follows. Head: Proboscis essentially lacking

midventral pale scales; generally slightly longer, length 1.8-2.0 mm, mean 1.9 mm.

Maxillary palpus dark brown-scaled, length 2.4-2.7 mm, mean 2.6 mm; palpomere 2 with

some lateral pale scales; lateral surface of palpomere 3 with pale scales beginning above

base and few pale scales ventromesally at apex; palpomere 4 with narrow line of pale

scales tapering distally on ventral surface and dorsolateral line of pale scales ending just

short of apex; palpomere 5 with spot of white scales ventrally at base and somewhat faint

dorsolateral line of pale scales tapering distally to near apex. Falcate scales of vertex

very nearly white; forked scales paler than in female but still brownish laterally; lateral

spatulate scales broader dorsally than ventrally and posteriorly. Thorax: Pair of pale

spots extremely faint or absent on middle of scutum. Upper proepisternal scales more

extensive, usually well extended mesally on anterior surface. Wing: Length of cell R2

shorter, length of cell R2/length of vein R2+3 variable, 1.5-2.1, mean 1.8. Legs:

Midfemur usually without anterior pale stripe. Abdomen: Terga as in female except VI

and VII only with middle area dark-scaled, borders with pale scales. Sterna usually with

median steak and posterolateral spots of dark scales. Genitalia (Fig. 17): Almost exactly

as in univittatus, the only obvious distinctions being the following: seta g of subapical

60 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

lobe of gonocoxite broader; ventral arm of lateral plate of phallosome shorter, not

projecting beyond caudal margin of lateral arm.

Pupa (Fig. 17). Character and placement of setae as figured, range and modal

number of branches in Table 8; no specific differences from univittatus are apparent,

setae 6,12-CT and 6-lll-VI tend to have fewer branches. Cephalothorax: Seta 6-CT

usually triple (2-4); 12-CT most often double (2-4). Trumpet: As in univittatus, index

5.4-7.5, mean 6.5. Abdomen: Length 2.7-3.0 mm, mean 2.8 mm; setae 1-lII-VI and

5-IV-VI tend to be rather shorter than in univittatus; 6-III usually triple, seldom double or

with 4 branches, 6-IV-VI most often triple (3-5). Paddle: Length 0.84-0.94 mm, mean

0.89 mm; generally not as broad as in univittatus, width 0.53-0.61 mm, mean 0.57 mm;

index 1.5-1.7, mean 1.6.

Larva (Fig. 18). Form and positions of setae as figured, range and modal number

of branches in Table 28; characters almost the same as in univittatus, the only apparent

distinction being the development of seta 1-S; the siphon/saddle index is generally

larger. Head: Length 0.71-0.86 mm, mean 0.78 mm; width 1.12-1.38 mm, mean 1.28

mm. Dorsomentum most often with 7 teeth (6-9) on either side of median tooth.

Antenna: Length 0.54-0.64 mm, mean 0.60 mm. Seta 1-A with about 22 branches

(19-27). Thorax: Seta 7-P normally triple, seldom with 4 branches. Seta 2-T usually

double, sometimes triple. Abdomen: Setae 3-l-VIl and 7-VI tend to have fewer

branches than in univittatus. Segment Vill: Comb with 37-62 scales, mean 51. Siphon:

Generally slightly longer than in univittatus (see); index 5.45-7.87, mean 6.77. Pecten

with 9-15 spines, mode 13. Seta 1-S normally in 5 pairs, sometimes only 4.5 pairs

present, all normally borne beyond pecten; length about as long as diameter of siphon

at point of attachment; always with fewer branches than in univittatus, each element

usually double or triple (cf. tables). Segment X: Saddle length 0.35-0.46 mm, mean

0.41 mm; siphon/saddle index 4.20-4.87, mean 4.44. Seta 1-X double or triple in nearly

equal frequencies.

Systematics. The recognition of perexiguus as a species distinct from

univittatus is somewhat provisional. The taxonomic status of these forms cannot be

settled unequivocally without a comprehensive study of individually reared material from

sympatric populations. An important analogy exists between these forms and pipiens.

The distribution of these forms coincides exactly with the distribution of pipiens in

southwestern Asia and Africa, yet pipiens is considered to be a single plastic species

while univittatus and perexiguus are regarded as separate species. The reason for this

is that univittatus and perexiguus exhibit slight anatomical differences (see Table 41),

whereas none are apparent in pipiens. For the present | am treating perexiguus as a

distinct species primarily because these differences, although variable, were observed

in a limited number of specimens from Ethiopia where the forms appear to be sympatric.

However, these forms seem to occur at different altitudes, and the observed differences

may be due in large part to environmental influences.

Bionomics. Culex perexiguus is a common mosquito which is particularly

abundant during the summer and autumn months. The immature stages are found ina

variety of standing water habitats including swamps, springs, stream pools, ponds, and

wells. Breeding occurs in clean to moderately polluted water which usually contains

quantities of emergent vegetation. This species tolerates some degree of salinity and

occasionally utilizes artificial containers. It generally breeds away from dwellings, but has

the ability to adapt to breeding places modified by man. It is frequently found in

association with pipiens, antennatus, and Aedes (Ochlerotatus) caspius (Pallas).

Although perexiguus probably feeds mainly on birds, it apparently occasionally

enters houses and bites man (Buxton, 1923; Kirkpatrick, 1925). West Nile virus has

Harbach: Subgenus Culex in Southwestern Asia 61

been isolated from this species in Israel (Nir et al., 1968; Samina et al., 1986) and Egypt

(Taylor et al., 1953; Taylor et al., 1956). Isolations of Sindbis virus have been made in

Israel (Samina et al., 1986), Egypt (Taylor et al., 1955), and Saudi Arabia (Wills et al.,

1985). Experimental transmission of Sindbis virus has been demonstrated for Egyptian

populations (Taylor et al., 1955).

Distribution. Culex perexiguus is widely distributed in northern Africa and

southwestern Asia, and extends eastward into India.

Material examined. 2,696 specimens. A total of 2,546 specimens (689

females, 445 males, 65 male genitalia, 889 pupal exuviae, 429 larval exuviae, and 29

fourth-instar larvae) were examined from southwestern Asia — AFGHANISTAN:

(Garwargin); EGYPT: Aswan (Aswan, El 'Aqaba el Saghira, Ezbet el Silsila, Knour Abu

Subeira, Kom Ombo, Nag el Higab, Nag el Idwa el Bahari, Nag el Kagug, Nag el Ritag el

Qibli, Nag el Shalabab, Nag' Tingar, Sahara City), Asyut (Beni Shigeir), Beni Suef (Biba),

Eastern Deserts (Bir el Maskhara, Wadi Digla, Farouk Field), E/ Buhayra (Wadi Natrun), E/

Fayyum (Abhit el Hagar, Birket Qarum, Sanhur Bahariya, Sinero el Bahria), E/ Girgha (EI

Birba), E/ Giza (Abu Ragwan el Bahari, Abu Rauwash, El Badrshein, Ezbet Bosna

Sharawy, Harania, Kafr el Ghataty, Koneissa, Nazlet el Ashtar, Saqqara, Talbia), El Isma’

iliya (Abu Khalifa, Isma’iliya), E/ Qahira (Abbassia, Kirdasa, Ezbet Said), E/ Qalyubiya

(Ga'afara, Kaha, Khanka), E/ Shargiya (Inshas el Raml), Port Said (Port Said), Qena (Nag'

Hammadi); IRAN: (Andimeshk, Behbehan, Bisotun, Ghosse, Kermanshah,

Khorramabad, Natchi Bahran, Shirin, Teheran); IRAQ: (Daurah); ISRAEL: (Akka,

Baniass, Beer Sheva, Beisan, Deir el Belah, Ein el Howl, Eliot, Galilee, Gaza, Genin,

Hadera, Inlerl, Jerisheh, Khirbet Hadrah, Kishon River, Lake Huleh, Latron, Lhallal,

Ludd, Nahala, Nahr es Zerga, Nahr Rubin, Shivta, Sidrat el Mallaha, Tel Aviv, Wassat,

Zichron); JORDAN: (Amman, Tafile, Taflah); LEBANON: (the lectotype male and

paralectotype female); OMAN: (Itelba, Se’ Harr); PAKISTAN: (Peshawar); SAUDI

ARABIA: (Al Hasa Oasis, Al Khobar, Al Qatif, Riyadh). An additional 150 specimens (46

females, 29 males, 27 male genitalia, 6 pupal exuviae, 17 larval exuviae, and 25

fourth-instar larvae) were examined from ALGERIA, CHAD, DJIBOUTI, ETHIOPIA,

MOROCCO, SENEGAL, SUDAN, TUNISIA, UPPER VOLTA, and unknown localities.

Culex (Culex) theileri Theobald

theileri Theobald, 1903a: 187. +Lectotype male: Pretoria, Transvaal,

[South Africa] (BM), designated by Sirivanakarn, 1976: 45.

creticus Theobald, 1903a: 189. +Holotype female: Crete, Greece (BM).

Synonymy with theileri by Edwards, 1932a: 210.

pettigrewii Theobald, 1910a: 15. +Holotype female: Ukhrul, Manipur,

6400 ft., [Assam], India (BM). Synonymy with theileri by Edwards,

1911: 262.

onderstepoortensis Theobald, 1911: 265. +Holotype female:

Onderstepoort, Transvaal, [South Africa] (BM). Synonymy with

theileriby Edwards, 1932a: 210.

annulata Theobald, 1913: 321(theileri var.). Type specimens (female):

Onderstepoort, Transvaal, [South Africa] (? BM, see Harbach, 1983:

103, 104). Synonymy with theileri by Edwards, 1932a: 210.

alpha Séguy, 1924: 41. Lectotype larva, hereby designated, larva

illustrated by Séguy (1924: PI. VII, Fig. 34), specimen non-extant (7)

(see Harbach, 1983: 99). NEW SYNONYMY.

Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Culex theileri Theobald of Gough, 1914: 135 (Egypt, coll. site); Storey, 1918(1919): 98

(Egypt, A, L keys, bionomics); Barraud, 1924b: 1266 (India, A key, syn., M’*, F,

distr.); Barraud, 1924c: 429 (India, L*); Scott, 1927: 88 (Ethiopia, coll. rec.);

salem, 1938: 27 (Sinai); Salem, 1940: 15 (Egypt, A, M gen.*); Parr, 1943:

246-251 (? Syria, Lebanon, A, P, L keys, bionomics); Senevet, 1947: 212 (North

Africa, L key); Theodor, 1952: 113 (Middle East, zoogeogr.); Dow, 1953: 688

(Iran, coll. rec., L bionomics); Mattingly, 1954: 58 (Africa, Asia, zoogeogr.); Gad,

1956: 135 (Egypt, distr., bionomic note); Senevet et al., 1957a: 85-86 (North

Africa, L"); Nielsen and Nielsen, 1958: 312 (Iraq, swarming beh.); Derwesh, 1965:

44 (Iraq); Margalit and Tahori, 1970a: 142 (Israel, bionomic note); Margalit et al.,

1971: 323 (Israel, bionomics); Gad and Salit, 1972: 581 (Egypt, coll. site); Margalit

and Tahori, 1973: 92, 93 (Sinai, distr., L assoc.); Margalit et al., 1973: 32 (Israel,

coll. rec., L assoc.); Buttiker, 1981: 476 (Saudi Arabia, coll. site); El-Said and

Kenawy, 1983a (Egypt, distr.); Zaini et al., 1983: 117 (Iraq, bionomic note); Kitron

and Pener, 1986 (Israel, L bionomics).

Culex (Culex) theileri Theobald of Edwards, 1926: 133 (Mediterranean Region, A, L

Culex

keys, A, L", syn., distr.); Martini, 1931: 384 (Egypt, Israel, Jordan, Syria, Turkey,

A, L keys, M*, F, L*, distr.); Barraud, 1934: 414 (India, A, L keys, M*, F, L*, syn., L

habitat, distr.); Edwards, 1941: 305, 419, 482 (Afrotropical Region, A key, M’*, F*,

P, syn., distr.); Lewis,1943a: 281 (Eritrea, coll. rec., L bionomic note); Lewis,

1945: 15 (Sudan, A, distr.); Senevet, 1947a: 124 (Egypt, L’*, distr., L bionomics,

L key); Senevet, 1949: 57, 64 (North Africa, M gen.*, key); Monchadskii, 1951:

2/75 (Middle East, L*, key, L assoc., distr.); Hopkins, 1952: 289 (Afrotropical

Region, L*, key, bionomics); Knight, 1953a: 232 (Yemen Arab Republic, coll.

rec.); Senevet and Andarelli, 1954: 60 (North Africa, A key, M, F); Khattat, 1955:

165 (Iraq, syn., L bionomics, distr., L*); Abdel-Malek, 1956: 101 (Sinai, L

bionomics, distr.); Lewis, 1956: 708-718 (Sudan, Egypt, Eritrea, L, coll. sites,

distr.); Mattingly and Knight, 1956: 106-129 (P.D.R. Yemen, Yemen Arab

Republic, L*, A, L keys, distr., zoogeogr.); Ovazza et al., 1956: 171 (Ethiopia, L*,

coll. sites); Parrish, 1959: 266 (Turkey, distr.); Senevet and Andarelli, 1959: 195

(North Africa, Mediterranean basin, A, P, L keys, syn., M*, F*, P*, L*, distr.,

bionomics); Abdel-Malek, 1960: 113-123 (Syria, coll. sites, L bionomics, distr.):

Khalaf, 1962: 54 (Iraq, A, L, L habitat); Derwesh, 1965: 44 (Iraq); DuBose and

Curtin, 1965: 352, 354 (Mediterranean area, A, L keys); Abul-hab, 1966: 281

(Iraq, distr., L bionomics); Abul-hab, 1968: 245 (Iraq, L habitat, distr., L key); Lotfi,

1970: 402 (Iran, coll. rec.); Margalit and Tahori, 1970b: 153 (Israel, coll. sites);

Aslamkhan, 1971: 155 (Pakistan); Ward, 1972: 96 (Afghanistan); Lotfi, 1973: 206

(Iran, coll. sites); Gutsevich et al., 1974: 388 (Asia, A, L keys, M’*, F, L*, distr.,

bionomics); Margalit and Tahori, 1974: 87 (Israel, coll. rec.); Lotfi, 1976: 72, 74,

80 (Iran, L key, ecol. note, L*); Sirivanakarn, 1976: 43 (Oriental Region, A, P, L

keys, M*, F*, P, L*, tax., distr., bionomics); El-Said and Kenawy, 1983b (Egypt,

coll. rec.); Ibrahim et al., 1983: 91 (Iraq, L*, key); Danilov, 1985a: 71 (Afghanistan,

F key); Danilov, 1985b: 55 (Afghanistan, L key); Harbach, 1985a: 86, 92, 106

(SW Asia, Egypt, distr., A, L keys).

tipuliformis Theobald of Edwards, 1912b: 31 (Africa, A key); Barraud, 1920: 324

(Iraq, bionomic note); Barraud, 1921: 394 (Israel, Jordan, Egypt, distr., L assoc.);

Edwards, 1922: 278 (Oriental Region, A key); Buxton, 1923: 317 (Israel, Jordan,

coll. rec., bionomic note);Theodor, 1923: 344 (Israel, Jordan, P*); Kirkpatrick,

1924(1925): 367, 371 (Egypt, A, L keys).

Culex (Culex) tipuliformis Theobald of Edwards, 1921: 339 (Mediterranean Region, A, L

Harbach: Subgenus Culex in Southwestern Asia 63

keys, L", tax., distr.); Séguy, 1924: 40, 192 (Egypt, Iran, Turkey, A, L keys, L"*,

tax., distr., syn.); Kirkpatrick, 1925: 115, 188 (Egypt, A, P, L keys, syn., M*, F, P*,

L*, distr., bionomics); Stackelberg, 1927: 159 (Middle East, M, F keys, A, M

gen.", distr.).

Adult. A distinct species with postspiracular and prealar scales and anterior white

stripes on all tibiae and the fore- and midfemora. Easily distinguished from all other

Culex species of southwestern Asia and Egypt, except perhaps univittatus, perexiguus,

and vagans. Culex univittatus and perexiguus are normally without an anterior white

stripe on the foretibia and vagans is without postspiracular and prealar scales.

FEMALE. Scaling of proboscis, maxillary palpus, legs, and abdominal terga

predominantly brownish black, legs blacker. Head: Antenna with some nearly white

scales on mesal surface of pedicel and flagellomere 1; length 1.6-2.0 mm. Proboscis

length 2.2-2.7 mm, mean 2.5 mm; dorsal surface largely black-scaled but often with

scattered pale scales, particularly in middle; ventral surface entirely with cream-colored

scales except at base. Maxillary palpus about 0.2 of proboscis length, with variable

amount of pale scaling; palpomere 4, and usually 3, with distinct dorsomesal patch of

white scales at apex, sometimes entire dorsomesal surface of both white-scaled:

scattered pale scales laterally, sometimes lateral surface entirely white-scaled. Falcate

scales of vertex whitish to light brown, usually largely pale yellow posteriorly, narrower

and darker anteriorly and laterally, narrowly white along margin of eye; forked scales pale

yellow medially, brown to blackish laterally; lateral spatulate scales white, narrow. Thorax

(Fig. 2B): Scutal integument dark brown; scales dense, rather fine to moderately coarse,

yellowish brown to golden brown, whitish or pale yellow on extreme anterior and lateral

margins and prescutellar area. Lobes of scutellum with scales same as prescutellar

scales. Falcate scales of antepronotum yellow, becoming pale yellow to white

posteriorly; falcate scales on anterior 0.67 of postpronotum fine, golden yellow, larger

and pale yellow to white on posterior 0.33. Pleural integument light to moderate brown,

faded posteriorly, frosted with dense covering of aculeae, with patches of white narrow

spatulate scales on upper area of proepisternum, postspiracular area near mesothoracic

spiracle, lower part of prealar area, upper and lower areas of mesokatepisternum, and

anterior and upper areas of mesepimeron, prealar patch continuous with upper

mesokatepisternal patch, latter usually continuous with lower mesokatepisternal patch,

anterior and upper mesepimeral patches often continuous; pleural setae golden: 10-16

upper proepisternal, 7-14 prealar, 4-7 upper mesokatepisternal, 7-13 lower

mesokatepisternal, 5-16 upper mesepimeral, and 1 lower mesepimeral. Wing: Length

3.9-4.8 mm, mean 4.4 mm; length of cell R2 2.8-3.5 length of vein R2+3, mean 3.3:

length of cell M2 about 0.8 length of cell R2; scales almost entirely black or nearly black,

pale scales on posterior side of costa, distal 0.67 of subcosta, anterior side of distal 0.5

of R, and anterior side and approximately distal 0.5 of posterior side of R1. Halter: Same

color as thoracic integument, with pale yellow to white scales on capitellum. Legs

(midleg, Fig. 2G): Anterior surface of forecoxa largely white-scaled, with black scales in

middle; midcoxa with small patch of white scales at base, these more or less distinctly

separated anteriorly from longitudinal patch of concolorous scales on anterior side of

lateral midline; hindcoxa with patch of white spatulate scales on anterior side of lateral

midline. All femora with narrow knee spots of white scales; posterior surface of fore- and

midfemora white-scaled, anterior surface dark-scaled with narrow longitudinal stripe of

white scales, white stripe of forefemur placed anteroventrally and sometimes more or

less broken up into spots, white stripe of midfemur borne medially with dark scales not

reaching base and anterodorsal line of dark scales progressively expanding onto dorsal

64 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

surface toward apex; hindfemur white-scaled with dorsal and anteroventral stripes of

dark scales, dorsal stripe beginning beyond base and gradually widening distally,

anteroventral stripe on approximately distal 0.5. Anterodorsal surface of foretibia

dark-scaled with complete longitudinal stripe of white scales, posteroventral surface

white-scaled; anterior and posterior surfaces of mid- and hindfemora with complete

stripe of white scales, narrowly dark-scaled on dorsal and ventral surfaces. Tarsomere 1

of all tarsi more or less distinctly pale-scaled with anterodorsal longitudinal stripe of dark

scales; other tarsomeres more or less completely dark-scaled, scales appearing opales-

cent under light of certain angles. Abdomen: Tergum | with median posterior patch of

pale scales; tergum Il with median longitudinal patch of creamy-white scales that is

broadest basally and may extend entire length of tergum; terga III-VIl with more or less

triangular basal patches of creamy-white scales, lateral patches of white scales and

usually with narrow band of pale scales on posterior border, triangular patches broader,

produced less in middle and often continuous with lateral patches on posterior terga,

sometimes posterior angle of triangular patches extends to posterior margin or this area

with pair of submedial spots of pale scales; tergum VIII largely or entirely pale-scaled.

Sterna usually with scattered dark scales but sometimes with all whitish or yellowish

scales or a mixture of both; dark scales sometimes fairly numerous and arranged in more

or less distinct submedial longitudinal stripes and posterolateral spots that are often

joined posteriorly.

MALE. Differing from female as follows. Head: Maxillary palpus approximately

1.25 length of proboscis; integument dark brown; palpomeres 1 and 2 with scattered

pale scales ventrally; palpomere 3 with extensive patch of white scales on dorsolateral

margin of distal 0.67 and small mesal patch near apex, setose ventrally; dorsal and

ventral surfaces of palpomere 4 each with line of white scales; palpomere 5 white-scaled

dorsally, with small patches of white scales ventrally at base and apex; palpomeres 4 and

5 densely setose, each with lateral rows of dark setae. Dorsal falcate scales of vertex

more numerous; color more uniformly pale yellow. Thorax: Scutal scales finer; with

22-27 upper proepisternal setae. Wing: Length of cell R2 1.9-2.3 length of vein R2+3,

mean 2.1. Abdomen: Terga Ill-VIl without lateral patches; basal triangular patches

generally broader; posterior terga usually with pale scales along lateral scale-free areas

and posterior border, these sometimes rather broad and more or less continuous with

submedial pale spots, thus occasionally significantly reducing dark areas. Genitalia (Fig.

19): Form as figured; structure of phallosome somewhat as in decens. Ninth tergal

lobes developed as usual, each with 7-14 (mode 9) irregularly spaced setae.

Gonocoxite normal, ventrolateral surface with usual complement of strong setae, mesal

setae longer and denser at base than usual, mesal surface also with 4 or 5 setae at apex,

lateral surface with cluster of moderately long setae at and below level of subapical lobe

(similar to /aticinctus and mattinglyi); subapical lobe slightly divided, setae a-c long and

tapered, a shorter and blunt at tip, b and c hooked apically; seta d absent, e simple, f

flattened and bladelike with one edge thickened; seta g foliform, rather short and

narrow, rounded apically; h simple, about same length as g, slightly curved distally.

Gonostylus developed as usual. Phallosome longer than broad with lateral plates and

aedeagal sclerites of nearly equal length; dorsal arm straight or bent laterad, tapered

distally, apex pointed; lateral arm with 2-5 (mode 3) laterally-directed denticles at tip, a

stubby dorsal process, and a rather weakly developed basal articulatory process; ventral

arm poorly developed, slender, sinuous, only slightly longer than one of the larger

denticles of lateral arm. Aedeagus rather conical; crest of aedeagal sclerite rather long,

about same length as proximal part. Proctiger not unusual; paraproct with moderately

long curved basal lateral arm and prominent ventral acetabulum, crown dark with numer-

Harbach: Subgenus Culex in Southwestern Asia 65

ous spinelike spicules mesally and short blades laterally. Cercal sclerite elongate,

irregular in outline; cercal setae 2-6 (mode 3). Tergum X straplike, articulating with

paraproct below basal lateral arm.

Pupa (Fig. 19). Form and positions of setae as figured, range and modal number

of branches in Table 9; resembling univittatus, perexiguus, antennatus, simpsoni, and

sinaiticus in the character of setae 6-IIl and 3-VIl, but usually distinguishable in having

seta 1-ll dentritic and the trumpet longer and flared. Cephalothorax: Lightly tanned; scu-

tum with pair of irregular submedial moderately tanned spots, these sometimes

subdivided; metanotum moderately tanned. Setae 1,2-CT usually with 4 branches (3-5);

6-CT with 3 or 4 branches; 8-CT usually with 5 branches (4,5); 9-CT double or triple;

10-CT generally with fewer branches than other species of the pipiens group (2-4);

11-CT double; 12-CT usually with 4 or 5 branches (3-5). Trumpet: Moderately tanned,

pinna darker than usual; rather long and flared distally, index 4.1-6.2, mean 4.4; pinna

oblique, rather long, 0.2-0.3 total length of trumpet. Abdomen: Lightly tanned, often

with dark V-shaped stripe beginning at base of tergum | and gradually tapering to

posterior margin of tergum IV, width of stripe on tergum | equals width of metanotum;

length 3.0-3.7 mm, mean 3.5 mm. Seta 1-Il dendritic, usually with fewer than 12

branches (7-16); 1-III-VII multiple, number of branches progressively fewer on each

succeeding posterior segment, 1-IIl often with 10 or 11 branches (6-13), 1-VII frequently

with 4 branches (3-6); 5-IV usually with 4 branches (2-4), 5-V,VI double, both distinctly

longer than 5-IIl, about 1.5 length of following tergum; 6-III-VI variable, often with 3-5

branches (2-6). Paddle: Developed and tanned as usual; length 0.79-1.08 mm, mean

0.94 mm; width 0.72-0.82 mm, mean 0.74 mm; index 1.2-1.4, mean 1.3.

Larva (Fig. 20). Character and placement of setae as figured, range and modal

number of branches in Table 29; resembles simpsoni and sinaiticus but differs from

other members of the pipiens group in having comb scales with a distinct sharp apical

spine, differs conspicuously from simpsoni and sinaiticus in the character of setae 5-C

and 1-S. Head: Length 0.78-1.05 mm, mean 0.88 mm; width 1.28-1.54 mm, mean 1.37

mm; mainly lightly tanned, lateralia with rather distinctly delineated dark area behind each

eye, dorsal apotome with 0-9 dark spots of variable size and distinctness behind seta

5-C (in 3 rows of 3 when all are present, largest at bases of seta 5-C and medially

between bases of seta 5-C and posterior margin of head, 3-5 spots usually present).

Dorsomentum most often with 6 teeth (5-8) on either side of median tooth. Seta 1-C

stout, moderately tanned, sometimes barbed; 2-C absent; 4-C often double or triple,

sometimes single; 5-C usually triple, occasionally with 4 branches; 6-C almost always

double, seldom triple; 7-C often with 7 branches (6-10); 10-C with 3 or 4 branches

occurring in nearly equal frequencies; 11-C normally double, seldom single; 12-C most

often with 4 branches (3-6); 14-C usually double, infrequently single. Antenna: Length

0.60-0.80 mm, mean 0.70 mm; proximal part lightly tanned, distal part moderately

tanned, with dark spot mesally at base; proximal part aciculate, aciculae coarser just

before seta 1-A. Seta 1-A with about 22 branches (18-32). Thorax: Integument hyaline,

surface smooth, tubercles of pleural setal groups moderately tanned; with distinctive

somewhat hourglass-shaped internal body of black pigment extending from anterior

margin to middle of metathorax, easily seen in larva resting at water's surface. Seta 7-P

usually triple, infrequently with 4 branches; 8-P most often double, frequently triple.

Seta 12-T usually double, sometimes single. Abdomen: Integument hyaline, lightly

tanned, sometimes slightly darker dorsally; tubercles of larger setae lightly to moderately

tanned. Seta 3-l usually double or triple (2-5), 3-VIl most often with 5 branches (3-8);

6-I-VI almost always triple, 6-I-IIl distinctly longer than 6-IV-VI; 1-III,IV usually with 4

branches, 1-lIl occasionally with 5 branches, 1-lV infrequently with 3 or 5 branches,

66 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

1-V,VI usually triple, 1-V often with 4 branches, 1-VI sometimes double and infrequently

with 4 branches. Segment VIII: Comb with 28-45 fringed scales (mean 35) with sharp

apical spine. Seta 1-VIII most often with 7 branches (5-9); 3-VIII usually with 8 branches

(7-10). Siphon: Index 4.15-5.36, mean 4.79; straight or very slightly sigmoid in lateral

view; moderately tanned, sometimes slightly darker in middle, proximal 0.5 often very

dark. Pecten of 8-13 long slender spines (mode 12) on basal 0.3 of siphon; spines with

3-5 short basal denticles. Seta 1-S usually comprised of 10 setae (7-13), two forming lat-

eral pair, others imperfectly paired close to posterior midline. Segment X: Saddle

complete; moderately tanned, sometimes dark along posterior margin; length 0.44-0.56

mm, mean 0.48 mm, siphon/saddle index 3.15-3.46, mean 3.33. Seta 1-X usually triple,

occasionally double, sometimes with 4 branches; 2-X most often with 4 branches, often

triple, sometimes with 5 branches; 4-X usually in 7 pairs, sometimes less 1 or 2 setae.

Systematics. Sirivanakarn (1976) placed theileri in a subgroup of its own and

suggested that Afrotropical species such as andersoni Edwards, 1914a, and simpsoni

might also be members of the group. It is still not known whether andersoni belongs to

this group, but simpsoni and its close allies are sufficiently distinct from theileri to be

placed in a separate subgroup. There are a number of intermediate Afrotropical forms

which are not definitely assignable to either subgroup, so these subgroups may have to

be redefined once the species are better known. The affinities of laticinctus and

mattinglyi are not clear, but they resemble theileri in a number of characters and | am

placing them provisionally in the thei/eri subgroup.

Culex theileri is a clearly marked species which is widely distributed in the

Afrotropical, southern Palaearctic, and western Oriental regions. The larvae and adults,

excluding the male genitalia, are extremely variable and exhibit the same degree of

variation within local populations that is observed throughout the entire range.

Therefore, it appears that only a single widespread and variable species is involved, yet it

is possible that the taxon actually consists of an aggregate of biologically distinct,

isomorphic species. This form needs to be thoroughly studied throughout its range.

| am certain that Culex alpha Séguy, 1924 is conspecific with theileri. There are

two possible type larval specimens labelled as a/poha in the Museum National d'Histoire

Naturelle, Paris (Harbach, 1983) which are easily identified as theileri. Since there is no

clear evidence to indicate that these larvae are type specimens, | have designated the

larva illustrated by Séguy as the lectotype of a/pha. This figure agrees in all essential

details with the current concept of theileri.

Bionomics. Culex theileriis an adaptable species which utilizes a wide range of

breeding sites, but it is also a focal species, reaching high densities in some areas while

appearing to be rare or absent in others. The immature stages can breed in permanent

or temporary bodies of fresh, brackish, or foul water. They are commonly found in

Stagnant or slow streams and irrigation ditches, swamps, marshy pools, ground pools,

ponds, springs, disused wells, open cisterns, and seepage water. The breeding sites

are usually sunlit and often contain little or no floating or emergent vegetation. Larvae

are often found alone or in association with pipiens, perexiguus, or antennatus.

In southwestern Asia, this species reaches peak densities in late spring and early

summer. Females feed mainly on mammals and are known to enter houses and bite man

(Kirkpatrick, 1925; Gad, 1956; Anderson, 1967). This species does not seem to be

involved in the transmission of any human diseases in southwestern Asia, but its

importance as a vector has not been thoroughly studied. Mcintosh et al. (1967) found it

naturally infected with West Nile and Sindbis viruses in South Africa, but easy

experimental transmission has been demonstrated only for Sindbis virus (Jupp et al.,

1966).

Harbach: Subgenus Culex in Southwestem Asia 67

Distribution. Culex theileri seems to have a discontinuous distribution in

southern and eastern Africa, lands around the Mediterranean, southwestern Asia,

southern USSR, southern China, India, and Burma.

Material examined. 1,452 specimens. A total of 1,337 specimens (311

females, 221 males, 62 male genitalia, 282 pupal exuviae, 161 larval exuviae, and 300

fourth-instar larvae) were examined from southwestern Asia — AFGHANISTAN: (Angur

Abajhr, Bolla Quchi, Garwargin, Karukh, Sayedabat); EGYPT: Aswan (Aswan, El ‘Aqaba

el Saghira, Nag’ Tingar, Sahara City), E/ Burhayra (Wadi Natrun — Bir Hooker, Ezbet Kafr

Dawud), El Fayyum (Abhit el Haggar, Birket Qarun), E/ Giza (Saqqara), El Isma' iliya

(Ismaiiliya), Marsa Matruh (Siwa Oasis — Abul el Leef, Aghurmi, Ain Bundi, Masouse,

Mshendit), Sinai Peninsula (unknown localities); IRAN: (Bisotun, Cham Asbi, Dorud,

Ghasem Gheshlari, Kivi, Maragheh, Mazandaran, Mehrdasht, Mish Kazeroon, Natchi

Bahran, Semnan, Sharaf Khaneh); IRAQ: (Abu Ghraib, Amara, Baghdad, Barari Bela

Musal, Basrah, Daurah, Erbil Town, Hadji Geul, Rawanduz Nahia, Shubaichas, unknown

localities); ISRAEL: (Acre, Deir el Belah, Dhahiriya, Ein Ghridyan, Khedeira, Kishon

River, Latron, Lake Huleh, Ludd, Marakeb, Ramat Razi'el, Shivta, Wadi Sharia);

JORDAN: (Az Zarqa); PAKISTAN (localities east of the Indus River): (Gujranwalia,

Jhelum, Lahore); P.D.R. YEMEN: (Jebel Jihaf); SAUDI ARABIA: (Gatib Oasis, Sabo);

SYRIA: (Damascus, Deir el Aachair, Doumar, Kissoue); TURKEY:\ (Adana, Istanbul,

Konya, Koyna-Eregli, Sinop-Boyobat, Tatvan); YEMEN ARAB REPWBLIC: (EI Amra,

Hada, San’a, Ta'izz). An additional 115 specimens (61 females, 38 males, 5 male

genitalia, 1 pupal exuviae, 6 larval exuviae, and 4 fourth-instar larvae) were examined

from ALGERIA, GREECE (CRETE) (including the holotype female of creticus ), KENYA,

INDIA (the type series of pettigrewii ), LIBYA, NEPAL, SOUTH AFRICA (including the

types of theileri and onderstepoortensis), TUNESIA, ZIMBABWE, and unknown

localities.

Culex (Culex) laticinctus Edwards

laticinctus Edwards, 1913a: 49. +tHolotype male: Tiberias, Palestine,

[Israel] (BM).

Culex sp. no. 2518 of Storey, 1918(1919): 101 (Egypt, A, L keys, A, M gen.*, L*).

Culex laticinctus Edwards of Barraud, 1921: 394 (Israel, Jordan, Syria, Egypt, L habitat,

distr.); Buxton, 1923: 317 (Israel, Jordan, coll. rec., L bionomics); Theodor, 1923:

341 (Israel, Jordan, P*, key); Kirkpatrick, 1924(1925): 368, 371 (Egypt, A, L

keys); Salem, 1938: 27 (Sinai); Parr, 1943: 246-250 (Lebanon, coll. sites,

bionomics, A, P, L keys); Lewis, 1945: 17 (Sudan, distr.); Senevet, 1946:

317-321 (? North Africa, L*); Leeson and Theodor, 1948: 228 (Socotra, L habitat,

coll. sites, A, L keys); Theodor, 1952: 113 (Middle East, zoogeogr.); Senevet et

al., 1957a: 86 (North Africa, L*); Abul-hab, 1968: 249 (Iraq, L key); Margalit and

Tahori, 1973: 90-93 (Sinai, distr., coll. sites, L bionomics); Margalit et al., 1973: 32

(Israel, coll. rec., L assoc.); Dimentmand and Margalit, 1981: 124-127 (Israel, L

bionomics); Kitron and Pener, 1986 (Israel, L bionomics).

Culex (Culex) laticinctus Edwards of Edwards, 1921: 342 (Egypt, Israel, Jordan, Oman,

A, L keys, tax., distr.); Séguy, 1924: 37, 190 (Egypt, Israel, Jordan, Oman, A, L

keys, M gen.*, tax., distr., syn., not Lin Figs. 20-23, PI. Il); Kirkpatrick, 1925: 119,

190 (Egypt, A, P, L keys, M*, F, P*, L*, bionomics, distr.); Edwards, 1926: 137

(Egypt, Israel, Jordan, Oman, A, L keys, A, L, distr.); Stackelberg, 1927: 160

68 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

(Middle East, M, F keys, A, M gen.*, distr.); Martini, 1931: 368 (Arabia ?, Egypt,

Israel, Jordan, Syria, A, L keys, M*, F, L", distr.); Edwards, 1941: 313, 420, 482

(Afrotropical Region, excluding records from Dhahran and San‘a, A key, M*, F*, P,

distr.); Lewis, 1943a: 282 (Eritrea, coll. rec., bionomic note); Senevet, 1947a:

123 (Egypt, Saudi Arabia, Syria, Turkey, L*, distr.); Abbott, 1948: 45 (Sudan, L, L

habitat, distr.); Senevet, 1949: 56 (North Africa, M gen.*, key); Monchadskii,

1951: 263 (Middle East, L*, key, L assoc., distr.); Hopkins, 1952: 298 (Afrotropi-

cal Region, L*, key, bionomics); Knight, 1953a: 229 (Yemen Arab Republic, coll.

rec.); Senevet and Andarelli, 1954: 58 (Israel, Jordan, A key, M, F); Abdel-Malek,

1956: 100 (Sinai, L bionomics, L key); Lewis, 1956: 710 (Sudan, Egypt, Eritrea,

coll. sites, L habitat, distr.); Mattingly and Knight, 1956: 102-132 (Oman, P.D.R.

Yemen, Socotra, Yemen Arab Republic, L*, A, L keys, distr., bionomics); Senevet

et al., 1957b: 92 (North Africa, L); Parrish,1959: 266 (Turkey, distr.); Senevet and

Andarelli, 1959: 158 (North Africa, Mediterranean basin, A, P, L keys, M*, F*, P*,

L*, distr., bionomics); Abdel-Malek, 1960: 113-120 (Syria, coll. sites, L bionomics,

distr.); DuBose and Curtin, 1965, 352, 354 (Mediterranean area, A, L keys);

Mekuria, 1968: 78 (Ethiopia, distr.); Lotfi, 1970: 401 (Iran, coll. rec.); Margalit and

Tahori, 1970b: 152 (Israel, coll. sites); Lotfi, 1973: 206 (Iran, coll. site); Gutsevich

et al., 1974: 345 (Middle East, L key); Lotfi, 1976: 73, 76, 83 (Iran, L key, ecol.

note, L*); Margalit and Tahori, 1974: 87 (Israel, coll. rec., L assoc.); Harbach,

1985a: 86, 95, 103 (SW Asia, Egypt, distr., A, L keys).

Culex (Culex) mattinglyi Knight of ? Abdel-Malek, 1960: 120 (Syria, coll. sites, L

bionomics, distr.); ? Abul-hab, 1968: 248 (Syria [based on report by Abdel-Malek,

1960: 120], L key).

Adult. A medium-sized mosquito easily recognized by the light-colored thorax,

dark proboscis, absence of postspiracular and prealar scales, and the relatively broad

basal pale bands of the abdominal terga. Individuals usually possess 2 or 3 lower

mesepimeral setae.

FEMALE. Head: Length of antenna about 2.0 mm; flagellum normal; pedicel and

flagellomere 1 with some small pale scales on mesal surface. Proboscis blackish, slightly

paler on proximal 0.6 of ventral surface; length 2.2-3.1 mm, mean 2.7 mm. Maxillary

palpus usually entirely black-scaled, palpomere 3 sometimes with few apical pale scales;

length 0.4-0.6 mm, about 0.2 proboscis length. Forked scales of vertex long and

slender, light yellowish brown; falcate scales white to pale yellowish, distinctly white

along margin of eye; lateral spatulate scales white. Thorax: Pleural integument yellowish

with tinge of brown, scutum not much darker. Scutal scales moderately coarse, light

yellowish brown, white scales on anterior promontory, lateral margins, along prescutal

suture and on prescutellar area; scutal setae prominent, brown, well contrasted with

light-colored scales. Scutellum with narrow whitish falcate scales on all 3 lobes. Ante-

and postpronota with long whitish to yellowish falcate scales; with 4-8 setae on

posterodorsal margin. Pleura with yellowish to golden-brown setae and patches of

narrow white spatulate scales: upper part of proepisternum with small patch of scales

below 5-10 setae, postspiracular and prealar areas without scales, prealar area with 7-14

setae, upper part of mesokatepisternum with scale-patch and 5-7 setae, lower posterior

border of mesokatepisternum with patch of semierect scales and row of 5-13 setae,

mesepimeron with anterior and upper scale-patches and 5-14 setae on upper area;

number of lower mesepimeral setae variable, commonly 2 or 3, often 1 (at least on one

side) and occasionally 4. Wing: Dark-scaled with short line of whitish scales on posterior

margin of costa near humeral crossvein; length 3.7-5.1 mm, mean 4.4 mm; length of cell

Harbach: Subgenus Culex in Southwestern Asia 69

R2 2.6-4.3 length of vein R2+3, subcosta intersects costa beyond furcation of vein R2+3;

length of cell M1 0.65-0.78 length of cell R2. Ha/ter: Pedicel and scabellum pale;

capitellum dark. Legs: Anterior surface of forecoxa with whitish scales, mid- and

hindcoxae with longitudinal patch of narrow whitish scales on anterior side of lateral

midline. All femora with inconspicuous whitish knee spots; anterior surface of fore- and

midfemora with blackish scales, posterior surface white-scaled; hindfemur largely white-

scaled, with dorsal stripe of black scales beginning 0.1-0.3 from base and gradually

widening to encircle approximately distal 0.2. All tibiae black-scaled anteriorly,

whitish-scaled posteriorly, apices of fore- and midtibiae inconspicuously pale-scaled,

apex of hindtibia with conspicuous pale spot. Tarsi black-scaled with hint of paler scales

on posterior surface of tarsomere 1. Abdomen: Tergum | with median posterior patch of

white to light brown scales; terga II-VIl black-scaled with basal 0.4-0.6 white-scaled;

tergum VIII with lateral patches of cream-colored scales, broad median area without

scales. Scaling of sterna I-VII entirely cream-colored; sternum VIII usually entirely white-

scaled.

MALE. Differing from female as follows. Head: Antennal length 1.5-2.1 mm,

mean 1.8 mm. Length of proboscis 2.3-2.9 mm, mean 2.6 mm; without ventral cluster of

setae. Maxillary palpus barely if at all longer than proboscis, length 2.5-2.9 mm, mean

2.7 mm; scaling mainly dark, palpomere 3 with subtle subapical stripe of pale scales on

ventrolateral margin, palpomere 4 with ventral stripe of white scales, and palpomere 5

with ventral spot of white scales at base. Wing: Length 3.3-4.2 mm, mean 3.7 mm;

length of cell R2 1.3-2.3 length of vein R2+3, subcosta intersects costa before furcation

of vein R2+3; length of cell M1/length of cell R2 0.6-0.8. Abdomen: Basal bands

0.45-0.75 tergum length; posterior edge of bands irregular, usually with submedial

indentations of posterior dark scaling; bands produced posteriorly along lateral setae,

particularly on terga VI and VII. Genitalia (Fig. 21): Form as figured; phallosome much as

in theileri. Ninth tergal lobe small, with 4-9 (mode 5) small setae. Gonocoxite stout,

ventral and mesal surfaces with usual complement of large and small setae respectively,

mesal surface also with conspicuous clump of moderately long setae near base of

gonostylus, lateral surface with rather dense covering of moderately long setae and

some long slender setae at base; subapical lobe distinctly divided but not prominent,

setae a-c stout, slightly flattened and bent distally, a about half as long as c, b

intermediate; seta d absent, e and fresemble a-c but are more slender and distinctly

flatter and broader distally, e about as long as a, fabout as long as c; setaga

crescent-shaped blade with pointed tip; MN slender, curved, and inconspicuous.

Gonostylus relatively short, rather more sharply bent in middle than usual, with crest of

small sharp ridges before tip. Phallosome constructed as in theileri but lateral plate

longer than aedeagal sclerite, crest of aedeagal sclerite shorter, lateral arm of lateral plate

with 3-7 (mode 4) apical denticles, and ventral arm more strongly developed, much

longer than denticles of lateral arm. Proctiger almost as in theileri, ventral acetabulum of

paraproct inconspicuous; cercal setae more numerous, 5-8 (mode 7).

Pupa (Fig. 21). Form and placement of setae as figured, range and modal

number of branches in Table 10; distinguished from all the other species except

mattinglyi by the strongly flared trumpet with its short tracheoid area and unusually short

seta 6-1,l|. Cephalothorax: Lightly to moderately tanned, scutum, legs, and metathorax

darker. Setae 1,3,7,9-CT double or triple, more often double; 8-CT usually with 4 or 5

branches, sometimes with 6; 10- and 11-CT close set, 10-CT with variable number of

branches (4-10), 11-CT usually double, occasionally single; 12-CT with 3 or 4 branches.

Trumpet: Moderately tanned, tracheoid area darker; rather short and distinctly flared,

index 2.8-3.7, mean3.2; pinna 0.3-0.4 trumpet length; tracheoid area less than 0.3

70 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

trumpet length. Abdomen: Lightly tanned, anterior terga and sterna darker in middle,

anterior and posterior margins of these darker still; length 3.5-4.0 mm, mean 3.8 mm.

Seta 6-I,Il short, only slightly longer than 7-I,Il, single or double; 7-I usually triple,

sometimes double; 7-I| usually double, sometimes single; 1-Il with short stem and only

7-14 branches; 10-Il occasionally present, often only on one side, usually single, some-

times double or triple; alveolus of 11-II present; 6-III-VI usually triple (2-4); 5-IV shorter

than 5-V,VI, about length of following tergum, most often with 4 branches (2-5); 5-V,VI

normally double, distinctly longer than following tergum. Genital lobe: Moderately

tanned; larger than usual, length about 0.25 mm in female, about 0.50 mm in male;

female with short ventromesal pointed process projecting cephalad from base; male with

pair of bumps at base ventrally. Paddle: Very lightly tanned, buttress and midrib darker;

inner part widest distally; length 1.00-1.15 mm, mean 1.09 mm, width 0.83-1.00 mm,

mean 0.91 mm, index 1.1-1.3, mean 1.2.

Larva (Fig. 22). Character and positions of setae as figured, range and modal

number of branches in Table 30; easily distinguished from all other Culex in

southwestern Asia by the combination of seta 6-IIl-VI normally single and 2-X with 4 or 5

branches, resembling decens and auttoni in the character of seta 6-III-VI, and many

specimens of theileri, sitiens, poicilipes, and bitaeniorhynchus in the branching of seta

2-X. Head: Length 0.77-1.04 mm, mean 0.91 mm; width 1.10-1.54 mm, mean 1.33 mm;

mainly lightly tanned, darker ventrally (mainly gula) and posteriorly (particularly lateralia);

ventral ecdysial line distinct from level of seta 15-C to posterior margin of gula.

Dorsomentum usually with 7 or 8 teeth (7-9) on either side of noticeably larger median

tooth. Seta 1-C long and rather stout; 2-C present or absent; 5,6-C shorter than usual,

reaching only slightly beyond anterior margin of median labral plate; 5-C usually with 4 or

5 branches, seldom with 6; 6-C usually with 4 branches, sometimes triple; 8-C frequently

with 4 or 5 branches (3-6); 10-C usually triple (2-5); 14-C double or triple, more often

triple. Antenna: Length 0.53-0.68 mm, mean 0.62 mm; lightly tanned before seta 1-A,

distal part darker, with usual dark mesal spot at base; aciculate as usual, aciculae slightly

shorter and broader near seta 1-A. Seta 1-A with about 23 branches (19-26); 2,3-A

distinctly subapical. Thorax: Integument hyaline; surface smooth. Seta 7-P with 3 or 4

branches, more often with 3; 8-P double. Setae 1,2-M about same length, both

frequently triple (1-4 and 1-5 respectively). Seta 12-T double or triple, more often

double; 13-T smaller than usual, with 7-11 branches. Abdomen: Integument hyaline;

surface smooth. Seta 3-I usually with 3 or 4 branches (2-7), 3-VIl frequently double or

triple, sometimes with 4 branches; 6-I,ll usually triple, infrequently with 4 branches,

6-II1,Vl normally single, 6-III,I1V occasionally double; 1-Ill most often single, sometimes

double, 1-IV,V single, long, about 1.5 length of segment, 1-VI almost always single and

long, seldom double, 1-VII with fewer branches than usual, double or triple, more often

double. Segment Vill: Comb with 24-38 (mean 32) evenly fringed scales, fringe longer

at apex. Siphon: Index 3.07-4.18, mean 3.48; subcylindrical, broadest at base and

tapering distally; moderately tanned, slightly darker at base and apex. Pecten on

approximately basal 0.4, composed of 10-16 (mode 12) long curved spines with 3 or 4

basal denticles. Seta 1-S usually represented by 14 setae but 12-15 may be present, 2

paired laterally near tip, others more or less paired along posterior midline, 2 proximal

pairs within pecten, lateral and most distal pair smaller and with fewer branches than the

others. Segment X: Saddle complete; moderately tanned, darker on anterior and

posterior margins; length 0.36-0.48 mm, mean 0.42 mm, siphon/saddle index

2.78-3.24, mean 3.02. Seta 1-X single or double, more often double; 2-X with 4 or 5

branches (Kirkpatrick, 1925 reported 5 or 6); 4-X almost always in 7 pairs, infrequently

with 6 paired and 1 unpaired seta.

Harbach: Subgenus Culex in Southwestern Asia 71

Systematics. The adults of /aticinctus bear little resemblance to those of theileri

except for a remarkable similarity in the structure of the male phallosome. The larva of

this species is generally similar to that of theileri, but the comb scales and setae 1,6-III-VI

are very different. In spite of these differences | am including both /aticinctus and

mattinglyi in the theileri subgroup because mattinglyi, which seems to be very closely

related to /aticinctus, exhibits some characters in the adult and larval stages which are

intermediate in development between those in /aticinctus and theileri.

Culex laticinctus and theileri are sympatric throughout the arid lands of northern

Africa and southwestern Asia where they were once equally common species. Today,

laticinctus seems to be less common than theileri. The reason for this is unknown, but it

may be due to human influence. In 1983 | had the opportunity to collect mosquitoes all

over the Nile Delta area of Egypt with Bruce A. Harrison and we never found /aticinctus ,

not even in the vicinity of Alexandria where it was once quite common (Kirkpatrick,

1925). We examined a number of specimens in the Dokki Institute and the Entomo-

logical Society of Egypt in Cairo which came from near Alexandria during Kirkpatrick's

time, but no other specimens of /aticinctus were present in either repository.

The presence of two or three lower mesepimeral setae has been used

extensively to distinguish this species from other species of Culex known to occur in the

Afrotropical and Mediterranean regions. This character, however, is not as reliable as it

was previously thought to be. Many specimens examined during this study, especially

from Israel, have only one lower mesepimeral seta on both sides of the thorax. This

condition is reflected in the key presented above.

Bionomics. Culex laticinctus seems to have been more common in the past

than it is today. It was once collected frequently in artificial containers such as cisterns,

tanks, barrels, wells, concrete basins, and similar structures. Now it seems to be found

more often in stream pools, rock pools, swamps, springs, irrigation ditches, and

temporary ground water. It also uses seepages (Kitron and Pener, 1986) and has been

found in animal hoofprints supplied with water by a spring (Abdel-Malek, 1960). This

species usually occurs in fresh water, but is found occasionally in slightly brackish water.

It is frequently collected in association with Anopheles (Anopheles) claviger (Meigen)

and Culiseta (Allotheobaldia) longiareolata (Macquart). It has also been found in

association with Anopheles (Cellia) hispaniola (Theobald), ? An. (Cel.) multicolor

Cambouliu, An. (Cel.) sergentii (Theobald), Aedes (Ochlerotatus) caspius (Pallas),

Culiseta (Culiseta) annulata (Schrank), Uranotaenia (Pseudoficalbia) unguiculata

Edwards, Culex (Barraudius) pusillus Macquart, Cx. (Maillotia) hortensis Ficalbi, and

several members of the subgenus Culex, including mimeticus, perexiguus, pipiens,

sinaiticus, theileri, and tritaeniorhynchus. Culex laticinctus is mainly a summer species

but specimens can be found during most of the year. Resting adults have been

captured in tents, a tub, and an ice factory, but it is not known whether females enter

houses or bite man.

Distribution. This species is apparently widespread from the Canary Islands

eastward through lands around the Mediterranean, Equatorial Africa, Sudan, Ethiopia,

Somalia, the Arabian Peninsula, and the Middle East.

Material examined. 481 specimens. A total of 400 specimens (103 females,

109 males, 21 male genitalia, 67 pupal exuviae, 53 larval exuviae, and 47 fourth-instar

larvae) were examined from southwestern Asia — EGYPT: Sinai Peninsula (Mt. Abas),

unknown (Ibrahimien, Moharem Bey); IRAN: (Nanrizac Borazjun); ISRAEL (including the

holotype male): (Ein Turieba, Haifa, Jericho, Jerusalem, Mont Fort, Mt. Carnish, Rahm

Alla, Lake Tiberias, Wadi Faria, Yeroham); JORDAN: (Aqaba, Wadi Ram); LEBANON: (N.

Damdur); OMAN: (Muscat, unknown localities); P.D.R. YEMEN: (Amd Town, Anag,

Tz Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Awabil, Geidun, Khirbat Bakarman, Khalla, Mikhuras, Mukalla, Nafhun, N'Air, Saha,

Shu'abat Amudi, Wadi Amd, Wadi Duan); SAUDI ARABIA: (Hima, Jeddah, Qunfida);

SYRIA: (Baalbelk); YEMEN ARAB REPUBLIC: (Ta’'izz). An additional 81 specimens (35

females, 24 males, 4 male genitalia, 1 pupal exuviae, 6 larval exuviae, and 11

fourth-instar larvae) were examined from ALGERIA, GREECE (CRETE), ETHIOPIA,

LIBYA, MOROCCO, SOCOTRA, SUDAN, TUNISIA, and an unknown locality.

Culex (Culex) mattinglyi Knight

mattinglyi Knight, 1953b: 320. *tHolotype male: Birket Shiekh Kunnaf,

san'a, Yemen Arab Republic (NMNH).

Culex (Culex) laticinctus of Edwards, 1941: 313 (records from Zahran, Saudi Arabia, and

San’a, Yemen Arab Republic).

Culex (Culex) mattinglyi Knight of Knight, 1953b: 214-219 (Yemen Arab Republic, coll.

rec.); Mattingly and Knight, 1956: 94-134 (Saudi Arabia, Yemen Arab Republic,

A, L keys, distr., bionomics); Harbach, 1985a: 86, 95, 103 (Saudi Arabia, Yemen

Arab Republic, distr., A, L keys).

Described chiefly from the type series.

Adult. A medium-sized yellowish-brown species markedly similar to /aticinctus,

differing in having normally only 1 lower mesepimeral seta and a line of pale posterior

scutal fossal scales that is continous with a posterior line of pale outer dorsocentral

scales. The thoracic integument is generally darker and the male genitalia are distinctive.

FEMALE. Head: Antennal length 2.0-2.5 mm, mean 2.3 mm; pedicel and first

flagellomere largely yellow or yellowish brown, rest of flagellum dark; mesal surface of

pedicel dark with patch of tiny setae and pale scales; ventromesal surface of first

flagellomere with patch of whitish scales. Proboscis dark-scaled, labella lighter; with 3-5

large basal labial setae and number of smaller decumbent setae just anterior to them;

long and slender, length 2.9-3.2 mm, mean 3.1 mm. Maxillary palpus dark-scaled; length

about 0.6 mm, approximately 0.2 proboscis length; palpomere 2 usually with 2 or 3

strong lateral setae. Vertex with narrow pale yellow falcate scales, whitish on ocular line

and interocular space; forked scales brownish, darker posteriorly; lateral spatulate scales

narrow, yellowish white. Thorax: Integument yellowish brown, darker dorsally. Scutum

with narrow golden-brown falcate scales; scales whitish to pale yellow on margins and

prescutellar area; pale line of posterior scutal fossal scales continuous with concolorous

posterior line of outer dorsocentral scales; broad strip of bare integument between

distinct lines of acrostichal and dorsocentral scales; approximately 13 well developed

lateral prescutellar setae on each side. Scutellum with narrow pale yellow falcate scales;

lateral lobes each with 4-10 and median lobe with 7-10 large setae. Ante- and

postpronota with mainly broad yellowish-white spatulate scales, some narrow ones

anteriorly; postpronotum with 5-8 setae on posterodorsal margin. Pleura with rather

small patches of yellowish-white spatulate scales below upper proepisternal setae, on

upper corner and lower posterior border of mesokatepisternum, at middle of anterior

area of mesanepimeron and among upper mesepimeral setae; pleural setae golden

brown: 5-8 upper proepisternal, 8-13 prealar, 5-7 upper mesokatepisternal, 5-10 lower

mesokatepisternal, 10-16 upper mesepimeral, and normally 1 lower mesepimeral (2 on

one side of thorax in 1 female and 1 male examined). Wing: Length 5.4-5.7 mm, mean

5.5mm; length of cell R2 3.3-4.4 length of vein R2+3, mean 3.6; length of cell M1

Harbach: Subgenus Culex in Southwestern Asia rh)

0.8-0.9 length of cell R2; dark-scaled with small patch of pale scales on posterior margin

at base of costa, all scales narrow, those on veins Rs, R2+3, R1 and R2 noticeably longer.

Halter: \Integument dark with whitish scales covering scabellum and capitellum. Legs:

Anterior surface of forecoxa pale-scaled, with about 6 small setae and several

indistinct pale scales in short row posteriorly at apex; midcoxa with anterior patch of pale

scales; hindcoxa with narrow longitudinal stripe of pale scales on anterior side of lateral

midline. Ventral and posterior surfaces of trochanters covered with small white spatulate

scales. Fore- and midfemora dark-scaled anteriorly, pale-scaled posteriorly, with narrow

knee spots of pale scales; hindfemur mainly pale-scaled with dorsal stripe of dark scales

beginning at base and expanding over anterior and posterior surfaces on distal 0.3-0.4,

dark scaling sometimes forming complete ring distally, apex with knee spot of yellowish

scales. Tibiae dark-scaled, slightly paler posteriorly, apex of each with ring of yellowish

scales. Tarsi entirely dark-scaled. Abdomen: Tergum | with median posterior patch of

pale scales; tergum II with more or less convex basal pale band not reaching lateral

margins (one female in BM with median posterior extension of pale band); terga III-VII

with broad basal white bands, bands 0.3-0.5 tergum length, straight on terga Ill-V,

slightly concave on terga VI and VII; tergum VIII with concave basal pale band, band

sometimes obsolete in middle. Sterna entirely whitish-scaled; sternum VIII with lateral

scale-patches only, broad median area void of scales.

MALE. Differing from female as follows. Head: Length of antenna about 2.0-2.5

mm; pedicel and flagellomere 1 without scales. Maxillary palpus dark-scaled; with some

indistinct pale scales in dorsomesal line on distal 0.33 of palpomere 3, on ventral surface

of palpomere 4, and ventrally at base of palpomere 5; palpomeres 4 and 5 sparsely

setose, long setae mainly on palpomere 4; length about same as proboscis. Wing: Pale

scales absent at base of costa (few present at humeral crossvein in one specimen

examined). Abdomen: Terga and sterna same as in female except those of segment VIII

entirely pale-scaled. Genitalia (Fig. 23): Form as figured; phallosome resembling that of

theileri and /aticinctus. Ninth tergal lobes more widely separated than usual, each with

3-8 unevenly spaced setae. Gonocoxite enlarged, ventral and mesal surfaces with

setae developed as usual, dorsolateral surface with very dense vestiture of moderately

long hairlike setae; subapical lobe prominent, very near tip of gonocoxite, undivided;

seta a well removed proximally from b and c, rather slender and curved, blong, sinuous

and stout at base, c shorter, more slender and slightly hooked at tip; d-f represented by

3 fine simple setae on mesal side of lobe; g narrow and pointed; h rather short, slender

and nearly straight. Gonostylus notably expanded, with a conspicuous recurved

thornlike process on mesal side at tip. Lateral plate of phallosome differing from that of

laticinctus in having a smaller dorsal arm, stouter ventral arm, and lateral arm with variable

number of tiny elevations in addition to 2-6 (mode 5) distinct small denticles. Proctiger

not much different than in /aticinctus; basal lateral arm of paraproct not as large; cercal

setae generally fewer in number (2-6, mode 4).

Pupa (Fig. 23). Known from a single specimen in the type series; form and

placement of setae as figured, number of branches in Table 11; apparently similar to

laticinctus, seemingly differing in most of the following. Cephalothorax: Setae 8,12-CT

double; 11-CT with 4 branches; 13-CT present, small. Trumpet: Index about 4.0; pinna

slightly longer. Abdomen: Length 3.8 mm. Seta 6-I,II multiple; 10-ll with distinct stem

and many branches; alveolus of 11-Il absent; 5-V,VI single, 5-VIl much shorter than 4-VII.

Paddle: Length about 1.20 mm; width about 0.95 mm; index about 1.3.

Larva (Fig. 24). Character and positions of setae as figured, range and modal

number of branches in Table 31; resembles J/aticinctus but differs constantly in having

the thorax covered with minute spicules, seta 6-VI double, and seta 2-X double or triple.

74 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Head: Length 0.94-1.05 mm, mean 0.99 mm; width 1.26-1.48 mm, mean 1.37 mm;

moderately tanned, slightly to distinctly darker behind eyes. Dorsomentum with 9-12

(mode 10) teeth on either side of median tooth. Seta 1-C long and slender; 2-C

present; 5,6-C long, 5-C usually with 4 branches (3-5), 6-C normally triple, infrequently

with 4 branches; 8-C most often triple (2-4); 10-C forked, often triple (2-5); 14-C single.

Antenna: Long and slender, length 0.70-0.78 mm, mean 0.73 mm; lightly to moderately

tanned, distal part darker, basal rim and mesal spot at base more heavily tanned. Seta

1-A with about 21 branches (18-26); 2,3-A subapical as in /aticinctus. Thorax:

Integument hyaline; covered with tiny spicules. Seta 7-P appreciably longer than usual,

usually double (2-4). Seta 1-M longer than 2-M, single; 2-M most often with 4 branches

(2-4). Seta 12-T single; 13-T short as in /Jaticinctus but with fewer branches (4-7).

Abdomen: \Integument hyaline, smooth. Seta 3-I usually triple, seldom double, 3-VII

usually double, sometimes single; 6-I-IV with 2-5 branches, 6-1,IV usually double, 6-II, III

usually triple; 6-V,VI usually double, sometimes triple; 1-IV-VI single or double, 1-VI more

often single. Segment VIII: Integument with minute spicules, particularly around comb.

Comb with 29-48 (mean 38) scales. Siphon: Index 3.40-4.55, mean 4.03; more or less

distinctly swollen before middle, apex narrowed. Pecten with 13-22 (mode 16) spines;

most spines with 2 basal denticles (1-3). Seta 1-S usually composed of 14 setae (7

pairs) but 11-16 may be present, paired and developed as in /aticinctus. Segment X:

Saddle complete, broader than long; moderately tanned, darker dorsally; posterodorsal

margin conspicuously spiculate; length 0.37-0.42 mm, mean 0.39 mm, siphon/saddle

index 3.90-4.64, mean 4.19. Seta 1-X double or triple, more often triple; 2-X double or

triple; 4-X usually comprised of 12 setae (6 pairs), sometimes 11 or 13 are present.

Systematics. Culex mattinglyi is clearly indigenous to the southwestern

mountains of the Arabian Peninsula where it is restricted to higher terrain. This species

is undoubtedly derived from the same stock as /aticinctus. The two species are so similar

that Edwards (1941) failed to recognize specimens of mattinglyifrom San'a, Yemen Arab

Republic, and Zahran, southwestern Saudi Arabia, as a species different from /aticinctus.

In fact, the author of mattinglyi did not realize that the type series was distinct from

laticinctus until he removed and examined the male genitalia (Kenneth L. Knight,

personnal communication). The adults of mattinglyi differ in normally having only one

lower mesepimeral seta and a diffuse line of pale scales on the scutum. The males differ

markedly in the development of the gonocoxite and gonostylus. Conspicuous

differences in the larval stage include the minutely spiculate thorax and the differently

developed setae 6-VI and 2-X. The pupa of mattinglyi is known from only one specimen

in the type series so it is probably much more difficult to distinguish from that of

laticinctus than it appears to be in the key.

Culex mattinglyi and laticinctus differ conspicuously from ftheileri in the

ornamentation of the adult, the structure of the comb scales of the larva, and the

development of setae 1,6-ll of the pupa. Culex mattinglyi seems to be intermediate

between theileri and /aticinctus with respect to the general pigmentation of the adult, the

structure of the gonocoxopodite of the male, and the development of setae 1,6-III-VI of

the larva. | consider mattinglyi and laticinctus to be members of the theileri subgroup

because of the intermediate nature of mattinglyi. The male phallosome of these species

is undoubtedly of the type found in theileri, and it is difficult to believe that this striking

similarity is due to convergence.

Bionomics. Knight (1953b) found the immature stages of this species in a

cement watering trough, a broad open well, a large cement basin, and a large cement

tank. Adults misidentified as /aticinctus by Edwards (1941) were collected in a house.

Other than this nothing is known about the bionomics or disease relations of this

Harbach: Subgenus Culex in Southwestern Asia 75

species.

Distribution. Culex mattinglyi is known only from the highlands of the Yemen

Arab Republic and a single locality near its northern border in Saudi Arabia.

Material examined. 42 specimens (11 females, 5 males, 5 male

genitalia, 1 pupal exuviae, 11 larval exuviae, and 9 fourth-instar larvae) were examined

from southwestern Asia — SAUDI ARABIA: (Zahran); YEMEN ARAB REPUBLIC

(mostly type specimens): (Al-Asr, Rauda, San'a, Wadi Dhahr).

Culex (Culex) simpsoni Theobald

simpsoni Theobald, 1905c: 28. t+tLectotype male, hereby designated,

bearing following data: "46 // Type // Transvaal / Simpson // Culex /

simpsoni / (Type) F.V.T. // Recd. from / F.V. Theobald, / 1907-29.";

genitalia on acetate strip (BM).

richteri Ingram and De Meillon, 1927: 72. +Lectotype male, hereby

designated, bearing following data: Museum type label with "Culex"

written above and "richteri / [illegible mark]" written below the word

"Type" // "[Begamuzi River], Zululand. / 1927. / Dr. A. Ingram. / B.M.

1928-27." (BM). Synonymy with simpsoni by Edwards, 1932a: 210.

mauritanicus Callot, 1940: 362. +Cotypes (larvae): Taghjicht, Morocco

(IPS; slide with 3 larvae retained in NUNH). NEW SYNONYMY.

Culex simpsoni Theobald of Edwards, 1913b: 58 (Africa, A, M gen.); Lewis, 1945: 16

(Sudan, distr.).

Culex (Culex) simpsoni Theobald of Edwards, 1941: 309, 420, 482 (Afrotropical

Region, A key, M*, F, P, syn., distr.); Lewis, 1943a: 282 (Eritrea, coll. rec., L

bionomic note); Hopkins, 1952: 293 (Afrotropical Region, L key, L*, bionomics);

Knight, 1953a: 215, 218, 230 (Yemen Arab Republic, coll. rec., M, distr.); Lewis,

1956: 709 (Sudan, coll. sites); Mattingly and Knight, 1956: 94-132 (Yemen

Arab Republic, A, L keys, distr., bionomics); Ovazza et al., 1956: 172 (Ethiopia,

coll. sites, L assoc.); Harbach, 1985a: 86, 93, 106 (Yemen Arab Republic only,

distr., A, L keys).

Described chiefly from specimens collected in eastern Africa.

Adult. This and the following species resemble univittatus and perexiguus in

most respects, but are easily distinguished by the absence of an anterior pale stripe on

the hindtibia. They also lack a spot of pale scales at the base of the costa.

FEMALE. Proboscis, maxillary palpus, legs and abdominal terga largely clothed in

dark brown scales. Head: Antenna dark, pedicel with pale scales on dorsomesal surface;

length 1.6-1.9 mm, mean 1.7 mm. Proboscis with pale scaling in middle of ventral

surface, sometimes forming more or less complete ring; length 1.6-2.0 mm, mean 1.8

mm. Maxillary palpus with some pale scales dorsomesally at apex; length about 0.4 mm,

approximately 0.2 proboscis length. Falcate scales of vertex coarse, yellowish white:

forked scales long, yellowish medially, dark laterally; ocular scales elongate, white,

continuous with elongate white spatulate scales of postgenal area. Thorax: Integument

brown. Scutum with yellowish and golden-brown falcate scales, often forming pattern

consisting of median stripe on anterior 0.5, patches on fossae and pair of diffuse

patches behind middle; with paler scaling on anterior and lateral margins, posterior

dorsocentral area and lateral prescutellar area. Scutellum with white to very pale yellow

76 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

falcate scales mainly on median lobe, scales larger on median lobe and projecting laterad

on either side of midline. Ante- and postpronota with some golden-yellow falcate scales,

longer on antepronotum. Pleura with patches of broad white spatulate scales as follows:

patch ventral and mesal to upper proepisternal setae, patch on postspiracular area near

spiracle, small patch below prealar knob, larger patches on upper corner and lower

posterior border of mesokatepisternum, anterior patch on mesepimeron about same

level and size as upper mesokatepisternal patch, and small patch before upper

mesepimeral setae. Pleural setae golden brown, numbers follow: 4-6 upper

proespisternal, 6-10 prealar, 4-6 upper mesokatepisternal, 4-7 lower mesokatepisternal,

3-5 upper mesepimeral, and 1 lower mesepimeral. Wing: Length 3.0-3.5 mm, mean 3.2

mm; length of cell R2 2.0-2.5 length of vein R2+3; length of cell M1 about 0.8 length of

cell R2; entirely dark-scaled. Halter: Pedicel and scabellum pale; capitellum dark brown.

Legs (hindleg, Fig. 2L): Forecoxa pale-scaled with dark scales in middle; mid- and

hindcoxae with longitudinal line of pale scales on anterior side of lateral midline, scales

slightly more numerous on hindcoxa. Femora with narrow knee spots of white scales;

anterodorsal surface of forefemur dark-scaled, posteroventral surface white-scaled;

midfemur dark anteriorly, dark scaling expanded over dorsal surface toward apex,

posterior surface white-scaled; hindfemur largely white-scaled, with narrow dorsal stripe

of dark scales beginning near base and abruptly widening to encircle femur on distal 0.2.

Tibiae with spots of white scales dorsally at apex, large and conspicuous on hindtibia;

fore- and midtibiae dark-scaled anteriorly, pale-scaled posteriorly; hindtibia very dark,

nearly black on dorsal surface. Tarsi entirely dark-scaled. Abdomen: Tergum | with large

median patch of brown scales; terga Il-VII with distinct, slightly convex, white basal bands

that are narrowly joined to basolateral spots of same color, bands 0.20-0.35 tergum

length; tergum VIII normally entirely white-scaled. Sterna with broad dark apical bands

(0.45-0.50 sternum length), usually produced anteriorly in middle; sternum VIII with

lateral patches of pale scales only.

MALE. Like female except as follows. Head: Proboscis sometimes with few pale

scales ventrally at base; length shorter, about 1.4 or 1.5 mm. Maxillary palpus dark

brown; lateral surface of palpomere 3 with median patch of white scales; palpomere 4

with ventral line of white scales on proximal 0.5; palpomere 5 with ventral spot of white

scales at base. Scales of vertex generally paler. Wing: Slightly shorter, 2.6-3.1 mm,

mean 2.9 mm; length of cell R2 1.6-2.1 length of vein R2+3. Legs: Midfemur sometimes

with very narrow line of pale scales on anterior surface. Abdomen: Terga without

basolateral pale spots. Sterna sometimes almost entirely dark-scaled, with basolateral

pale spots; tergum VIII (ventral in position) entirely dark-scaled or with narrow posterior

band of pale scales. Genitalia (Fig. 25): Form as figured; general structure of

phallosome as in decens. Ninth tergal lobe not produced, with 5-10 (mode 7) short

setae. Gonocoxite normal, ventral and mesal setae developed as usual, lateral setae in 2

rows of 4 or 5 setae each; subapical lobe undivided, setae a-c of the usual form, group

d-e absent, fstout (on order of magnitude of a-c), tapered and flattened apically, g

asymmetrical (triangular) and rounded apically, h gently curved. Gonostylus narrow,

tapered; gonostylar claw rather broad, flattened and curled. Phallosome longer than

broad, lateral plate larger than aedeagal sclerite; dorsal arm more or less straight,

diverging slightly from its mate of the opposite side; lateral arm with graded series of

laterally directed denticles on caudal margin, 2-5 (mean 4) larger denticles project over

13-23 (mean 18) smaller denticles which grade into small bumps laterally, small denticles

extend onto prominent dorsal process; ventral arm much as in decens. Proctiger

unmodified; paraproct with long, curved, rodlike basal lateral arm and distinct ventral

acetabulum. Cercal setae most often 3 (2-4).

Harbach: Subgenus Culex in Southwestern Asia 77

Pupa (Fig. 25). Not associated with adults from southwestern Asia; form and

placement of setae as figured, range and modal number of branches in Table 12; much

the same as antennatus, but generally smaller, as reflected in the length of the abdomen

and paddle, and pinna of trumpet usually shorter. Cephalothorax: Not distinguishable

from antennatus. Trumpet: |Index 4.6-6.3, mean 5.2; pinna short, usually less than 0.2

trumpet length. Abdomen: Length 2.2-2.6 mm, mean 2.4 mm. Seta 1-II not so variable,

generally with fewer branches (12-19); 6-III,IV like 6-V,VI, often with 5 branches (4-6).

Paddle: Length 0.63-0.77 mm, mean 0.69 mm; width 0.48-0.59 mm, mean 0.50 mm;

index 1.3-1.6, mean 1.4.

Larva (Fig. 26). Not associated with adults from southwestern Asia; character

and positions of setae as figured, range and modal number of branches in Table 32; this

and the next species are readily separable from all other species of Culex in

southwestern Asia by the combined character of seta 5-C, seta 14-P, and the comb

scales. Head: Length 0.58-0.72 mm, mean 0.64 mm; width 1.01-1.14 mm, mean 1.06

mm; lightly tanned, darker behind eyes. Dorsomentum with 4-6 teeth on either side of

median tooth, third from center distinctly larger. Seta 1-C long, slender, pale; 2-C

absent; 4-C rather smaller than usual; 5-C single or double, most often single; 6-C

usually double, occasionally single; 10-C usually triple (2,3); 11-C double; 13-C most

often triple, sometimes with 4 branches; 14-C normally double with markedly divergent

branches (2-4). Antenna: Length 0.50-0.66 mm, mean 0.60 mm; proximal part lightly

tanned with dark basal ring contiguous with dark basomesal spot, distal part moderately

tanned. Seta 1-A with about 27 branches (17-33). Thorax: Integument hyaline,

smooth. Setae 4,7-P with 3 or 4 branches, 4-P more often with 4, 7-P most often triple;

14-P normally double, sometimes single on one side. Seta 1-M usually single, seldom

double, longer than 2-M; 5-M shorter than usual. Seta 1-T normally single, seldom

double, 0.5 or less length of 2-T; 2-T usually double, rarely single, occasionally triple;

13-T usually with 5 or 6 branches (5-7). Abdomen: |Integument hyaline, smooth. Seta

3-I-IV double (3-1 occasionally triple), 3-V,VI single, 3-VIl variable (2-5); 6-I-VI normally

triple (see table); 7-| usually double, sometimes single; 1-ll-V with 4 or 5 branches, 6-VI

with 3 or 4 branches, 6-VII often with 5 or 6 branches (4-7). Segment Vill: Comb with

17-36 scales, mean 28; most scales with sharp apical spine, some anterior scales fringed

entirely, apical spine usually larger on posterior scales. Siphon: Index 5.14-5.67, mean

5.40; slender, tapered, broadest at base; lightly tanned, basal rim and acus moderately

tanned. Pecten on proximal 0.33 or less, composed of 9-14 (mode 12) rather long

slender spines with tiny basal denticles. Seta 1-S in 4 pairs, 1a-S occasionally within

pecten, 1b- and 1c-S out of line with the others; usually double or triple (see table).

Segment X: Saddle lightly tanned, with very inconspicuous posterodorsal spicules;

length 0.35-0.39 mm, mean 0.37 mm, siphon/saddle index 2.98-3.97, mean 3.85. Seta

1-X usually triple, sometimes double; 2-X double, triple, or with 4 branches occurring in

nearly equal frequencies; 4-X almost always in 6 pairs.

Systematics. The simpsoni subgroup is erected for simpsoni and sinaiticus,

and also undoubtedly includes Striatipes Edwards, 1941, terzii Edwards, 1941, and

seldeslachtsi Wolfs, 1947 from the Afrotropical Region. The group is homogeneous in

all essential adult and larval features. The adults resemble univittatus in ornamentation

and coloration, but differ markedly in the male genitalia. The male genitalia resemble the

type found in the decens subgroup, but the species are otherwise distinct in all life

stages. The larvae share some characteristics with theileri, particularly the form of the

comb scales, but differ conspicuously in the position and development of seta 1-S.

Culex simpsoni shows relatively little variation in adult characters, particularly the

male genitalia. The larva is somewhat variable in details of the comb and siphon, but

78 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

there is no indication that more than one species is involved in the various separated

populations examined. During this study, several individually reared specimens from

Morocco previously identified as Culex mauritanicus Callot, 1940, and three cotype

larvae of this nominal species, were examined and found to be remarkably similar to the

type specimens of simpsoni and richteri in all important features. The similarities are so

evident in all stages that for the present | cannot consider mauritanicus as a distinct

species.

The occurrence of simpsoni in southwestern Asia is based upon two males

collected in the Yemen Arab Republic (Knight, 1953a). The association of the immature

stages with the adults is based on a small number of individual rearings from Tanzania

and Morocco, and the type series of richteri. The unknown stages of simpsoni in

southwestern Asia should agree with the descriptions above based on the overall

conformity of the males with African specimens.

Bionomics. Culex simpsoni is found in a wide variety of habitats, but its original

breeding places were probably sunlit pools of slow or stagnant water along streams or

streambeds. Besides these habitats, simpsoni is commonly found in lakes, ponds,

marshes, puddles, furrows, seepage pools, and artificial pools such as tubs, wells,

troughs, and tanks. Ribeiro and Mexia (1966) collected larvae in relatively saline

marshes and ground pools with somewhat turbid water and muddy bottom in Angola.

Larvae have been found in association with Anopheles (Cellia) cinereus Theobald, An.

(Cel.) pharoensis Theobald, An. (Cel.) sergentii (Theobald), Culex theileri, and Cx.

(Maillotia) arbieeni Salem (as jenkinsi Knight) in southwestern Asia (Knight, 1953a), with

An. (Cel.) arabiensis Patton (as gambiae Giles) in Uganda (Harris, 1942), and with Culex

antennatus, Cx. species near infula Theobald (as bitaeniorhynchus Giles), and Cx.

(Culex) thalassius Theobald in Angola (Ribeiro and Mexia, 1966). Nothing is known

about the vector potential of this species, but it is not likely to be of importance in

disease transmission because it only rarely bites man (Mattingly and Brown, 1955).

Distribution. This species is widely distributed in the Afrotropical Region and is

known to occur in the northwest African countries of Mauritania and Morocco.

Material examined. 229 specimens. Only 4 specimens (1 female, 2 males,

and 1 male genitalia) were examined from southwestern Asia —- YEMEN ARAB

REPUBLIC: (El-Hauban , Wadi el-Malak near Ta’izz). A total of 225 specimens (49

females, 52 males, 24 male genitalia, 15 pupal exuviae, 13 larval exuviae, and 72

fourth-instar larvae) were examined from African countries which include ETHIOPIA,

KENYA, MALAGASY REPUBLIC, MAURITANIA, MOROCCO (including 3 cotype larvae

of mauritanicus ), SOUTH AFRICA (the type specimens of simpsoni and richteri),

TANZANIA, ZAIRE, ZIMBABWE, and unknown countries.

Culex (Culex) sinaiticus Kirkpatrick

sinaiticus Kirkpatrick, 1924(1925): 383. *+Syntypes (male; female):

Kossaima, El Moweilleh, Ain Kadeis, Ain Gedeirat, and Ain Musa,

Sinai , Egypt (DI; ESE). |

Culex sinaiticus Kirkpatrick of Salem, 1938: 27 (Sinai, coll.); Lewis, 1945: 16 (Sudan, L

distr.); Leeson and Theodor, 1948: 228 (Socotra, A, L keys, coll. sites, L habitat);

Senevet et al.,1949: 46, 47 (Egypt [probably Sinai], A, M gen., L); Theodor,

1952: 113 (Middle East, zoogeogr.); Senevet et al., 1957a: 88 (North Africa, L);

Margalit and Tahori, 1973: 90-93 (Sinai, distr., L bionomics): Margalit et al., 1973:

Harbach: Subgenus Culex in Southwestern Asia 79

32 (Israel, coll. rec., L assoc.).

Culex (Culex) sinaiticus Kirkpatrick of Kirkpatrick, 1925: 122, 191 (Sinai, A, P, L keys, M*,

F, P*, L*, bionomics, distr.); Edwards, 1926: 138 (Sinai, A, L keys, A, L, distr.);

Stackelberg, 1927: 162 (Sinai, M, F keys, A, M gen.*, distr.); Martini, 1931: 381

(Sinai, A, L keys, M*, F, L*); Edwards, 1941: 311, 420, 482 (P.D.R. Yemen, Sinai,

Somalia, Sudan, A key, M", F, P, distr.); Lewis, 1943a: 282 (Eritrea, coll. rec., L

morphol. note, L bionomic note); Lewis, 1943b: 69 (Sudan); Abbott, 1948: 45

(Sudan, L, L habitat, distr.); Hopkins, 1952: 295 (Afrotropical Region, L key, L,

bionomics); Knight, 1953a: 216, 230 (Yemen Arab Republic, coll. rec., L, L

habitat, distr.); Lewis, 1956: 709 (Sudan, Eritrea, A, coll. sites, distr.);

Abdel-Malek, 1956: 100 (Sinai, L bionomics, distr., L key); Mattingly and Knight,

1956: 94-132 (Oman, P.D.R. Yemen, Saudi Arabia, Socotra, Yemen Arab

Republic, A, L keys, distr., bionomics); Senevet et al., 1957b: 92 (North Africa, L);

Senevet and Andarelli, 1959: 192 (SW Asia, A, P, L keys, M*, F, P, L, distr.,

bionomics); DuBose and Curtin, 1965: 352, 354 (Mediterranean area east of

Suez Canal, A, L keys); Lotfi, 1970: 401 (Iran, coll. rec.); Lotfi, 1973: 206 (Iran,

coll. sites); Margalit and Tahori, 1974: 87 (Israel, coll. rec.); Lotfi, 1976: 72, 74, 79

(Iran, L key, ecol. note, L"); Harbach, 1985a: 86, 93, 106 (SW Asia, Egypt, A, L

keys).

Adult. A rather small species closely resembling simpsoni, but with an

incomplete dark dorsal stripe on the distal half or less of the hindfemur, the abdominal

sterna largely pale-scaled, and different maxillary palpi and genitalia in the male.

FEMALE. Head: Antenna dark, pedicel paler; pedicel and flagellomere 1 with

some pale scales on mesal side; length about 1.5 or 1.6 mm. Proboscis dark with

indefinite broad area of pale scaling in midregion of ventral surface, sometimes forming

more or less complete ring; length about 1.6 or 1.7 mm. Maxillary palpus about 0.2

proboscis length; dark-scaled with pale scales dorsally on palpomere 3 and base and

apex of palpomere 4. Falcate scales of vertex coarser than in simpsoni, yellowish white;

ocular and interocular scales white; forked scales mainly yellowish, paler anteriorly, dark

posterolaterally; lateral spatulate scales white. Thorax (Fig. 2A): Integument brown.

Scutum with pale golden-brown falcate scales, whitish on margins; scutal setae dark

brown with reddish and golden tint. Scutellar scales whitish, few on lateral lobes,

confined mainly to median lobe. Antepronotum with large, almost white falcate scales

ventrally; postpronotum with similar but slightly yellowish scales, with 4 golden setae on

posterodorsal border. Pleural setae golden to golden brown, numbers follow: 4 or 5

upper proepisternal, 6-9 prealar, 3-5 upper mesokatepisternal, 2-6 lower meso-

katepisternal, 3-5 upper mesepimeral, and 1 lower mesepimeral. Pleura with broad white

spatulate scales as follows: patch ventral and mesal to upper proepisternal setae, patch

on anterodorsal border of postspiracular area just behind spiracle, small patch below

prealar knob, larger patches at upper corner and on lower posterior border of

mesokatepisternum, mesepimeron with anterior patch about same level and size as

upper mesokatepisternal patch, and small upper patch before upper mesepimeral setae.

Wing: Length about 3.0 mm; length of cell R2 2.0-2.3 length of vein R2+3, mean 2.1;

length of cell M1/length of cell R2 0.7-0.8; entirely dark-scaled; dorsal scaling: elongate

spatulate scales on costa, subcosta, R4+5, mcu, M3+4, CuA and proximal 0.3 of 1A,

linear or near linear scales on other veins and distal part of 1A; ventral scaling: elongate

spatulate scales on costa, subcosta, R1, R2+3, bases of R2 and R3, M, M1+2, and

proximally on M1 and M2, scales absent from R, CuA before mcu and proximal 0.5 of 1A,

other veins and distal parts of CuA and 1A with linear or near linear scales. Halter: Pedi-

80 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

cel and scabellum pale; capitellum dark. Legs (hindleg, Fig. 2M): Anterior surface of

forecoxa completely white-scaled or with dark scales in middle; mid- and hindcoxae with

line of white or slightly yellow scales on anterior side of lateral midline. Femora with

distinct white knee spots; fore- and midfemora with anterior dark scaling expanded

distally over dorsal surface; forefemur often with narrow anteroventral line of white scales

on proximal 0.5; midfemur with nearly complete white anteroventral stripe that terminates

before apex, stripe often reduced to narrow line on distal 0.5; hindfemur largely white,

with dorsal dark stripe on distal 0.5 or less (usually about 0.4) that abruptly expands over

anterior and posterior surfaces on distal 0.1-0.2 (normally about 0.2). Tibiae with

conspicuous white anterodorsal spots at apex; anterior and dorsal surfaces of foretibia

dark-scaled, posterior and ventral surfaces pale-scaled; midtibia pale-scaled dorsally,

dark-scaled ventrally; hindtibia mainly dark-scaled, posteriorly with narrow longitudinal

stripe of pale scales not reaching base or apex (about 0.2 from both). Tarsi mainly

dark-scaled; posteroventral surface of tarsomere 1, and often 2, with pale scaling.

Abdomen: Tergum | with median posterior patch of black, black and white or white

scales; terga II-VIl with white basal bands 0.3-0.4 tergum length, slightly produced

laterally, especially on terga VI and VII; tergum VIII almost entirely pale-scaled. Sterna

pale-scaled, infrequently with some subtle dark scales on posterolateral corners of

sterna VI and VII; sternum VIII with scales on lateral borders only.

MALE. Like female except as follows. Head: Antenna shorter, length 1.2-1.5

mm, mean 1.4 mm. Length of maxillary palpus 1.7-1.9 mm, mean 1.8 mm, extending

beyond tip of proboscis by length of palpomere 5; palpomere 3 with broad line of white

scales on dorsolateral margin not reaching base or apex, also with 3 or 4 long setae and

several shorter ones distally beyond pale scaling on ventrolateral margin; palpomere 4

with ventral line of white scales not always reaching apex; palpomere 5 with ventral spot

of white scales at base. Legs: Anteroventral pale stripe of midfemur poorly developed

or absent. Abdomen: Sterna usually with some dark scaling on posterior margin or

posterolateral corners. Genitalia (Fig. 27): Form as figured; phallosome developed as in

simpsoni. Ninth tergal lobe with 4-7 (mode 5) setae. Gonocoxite unmodified, lateral

setae more numerous than in simpsoni (about 21 in 4 rows, increasing in length toward

base of gonocoxite), apex with conspicuous cluster of about 9 moderately long setae on

mesal side of gonostylus; subapical lobe undivided, setae a-f similarly developed, rather

short, all about same length (fthe shortest), all flattened and hooked at tip; g a rather

long, crescentic blade; hlong and sharply bent distally. Gonosylus rather short and

narrow, distinctly flattened distally, rugose dorsally before tip. Phallosome very much as

in simpsoni, lateral plate with fewer denticles, 3-5 (mode 4) larger denticles project over

7-16 (mode 11) smaller denticles, smaller denticles not as well developed, often

reduced to mere bumps. Proctiger and cercal setae as in simpsoni.

Pupa (Fig. 27). Form and positions of setae as figured, range and modal number

of branches in Table 13; not separable from simpsoni unless by the possible distinctions

noted below. Cephalothorax: As in antennatus and simpsoni. Trumpet: \|ndex 4.0-7.3,

mean 5.6; pinna usually longer, 0.2-0.3 length of trumpet. Abdomen: Length 2.1-2.6

mm, mean 2.4mm. Setae 1-lII-VI, 5-Ill-V, and 6-III-VI generally shorter. Paddle: Length

0.72-0.82 mm, mean 0.77 mm; width 0.49-0.58 mm, mean 0.54 mm; index 1.3-1.6,

mean 1.4.

Larva (Fig. 28). Character and placement of setae as figured, range and modal

number of branches in Table 33; resembles simpsoni in most respects, differing mainly

in characters of the siphon. Head: Length 0.61-0.75 mm, mean 0.69 mm; width

1.01-1.18 mm, mean 1.08 mm; lightly tanned, not noticeably darker behind eyes.

Dorsomentum usually with5 teeth (4,5) on either side of median tooth, shape as in

Harbach: Subgenus Culex in Southwestern Asia 81

simpsoni. Seta 1-C long, very slender, pale; 5-C single, smaller than in simpsoni, 10-C

more often double (2-4); 11-C usually double, seldom single; 13-C with 3 or 4 branches, |

more often with 4. Antenna: Length 0.60-0.69 mm, mean 0.63 mm. Seta 1-A with

about 25 branches (21-28). Thorax: Seta 4-P double, triple or with 4 branches, seldom

single. Abdomen: Segments III and V with pigmentation giving abdomen a 2-banded

appearance similar to antennatus. Seta 3-| double or triple, 3-II-IV double (3-II, III seldom

single), 3-V,VI single (8-V seldom double), 3-VII with 3 or 4 branches; 1-IIl usually with 3

or 4 branches (3-5), 1-IV,V with 4 or 5 branches, 1-VI often with 4 branches (2-5).

Segment Vill: Comb with 32-44 scales, mean 37; anterior scales small and evenly

fringed, posterior scales larger with long apical spine. Siphon: Index 5.97-8.07, mean

6.97; long, slender, very slightly expanded apically; lightly tanned, slightly darker

distally, basal rim dark. Pecten on proximal 0.4, composed of 11-19 (mode 17) spines

similar to those in simpsoni. Seta 1-S in 4 pairs, first pair occasionally within pecten,

individual elements in straight row on posterolateral margin; normally double or triple

(see table). Segment X: Saddle longer than broad, length 0.33-0.40 mm, mean 0.38

mm, siphon/saddle index 3.35-3.80, mean 3.60. Seta 1,2-X double or triple; 4-X usually

in 6 pairs, occasionally in 7, sometimes with 1 seta missing on 1 side.

Systematics. Culex sinaiticus is very similar in all stages to simpsoni, but is

normally distinguishable in the adult and larval stages by the characters given in the

keys. The larva also apparently differs consistently in the positions of seta 1-S. The

pupae cannot be differentiated at present. The males are easily recognized by the

uniquely developed setae of the subapical lobe of the gonocoxite. The latter is

probably the most derived feature of the species.

Three larvae from southeastern Iran were available during this study which differ in

vague details from typical sinaiticus. The comb scales are larger than usual and more

closely resemble those of simpsoni. These specimens were not examined critically, but |

suspect that another species may be involved. The single male from Muscat, Oman,

noted by Mattingly and Knight (1956) seems to differ little from specimens collected

elsewhere in Arabia.

Culex beta Séguy, 1924 could be a synonym of sinaiticus. This nominal form was

described from an undisclosed number of larvae collected in Algiers. The partial

illustration of the larva published with the original description bears a definite

resemblance to sinaiticus and | would regard beta as a synonym of this species except |

have never seen a specimen identified as beta. There are no labelled specimens in the

alleged type depository (Harbach, 1983) and sinaiticus has never been recorded from

northern Africa. A thorough survey of the type locality is necessary before it can be

decided whether beta is a valid species or a synonym of Sinaiticus.

Bionomics. The immature stages of Sinaiticus are commonly found in quiet

permanent or semipermanent bodies of clean water such as streams, stream pools,

springs, wells, and seepages. The breeding sites often contain green algae or higher

plants. The species tolerates water with relatively high salt content, but occurs most

frequently in fresh water. Associated species of the subgenus Culex in southwestern

Asia include bitaeniorhynchus, laticinctus, mimeticus, perexiguus, pipiens, theileri, and

tritaeniorhynchus. The species has also been collected in association with Anopheles

(Cellia) arabiensis Patton, An. (Cel.) dthali Patton, An. (Cel.) hispaniola (Theobald), An.

(Cel.) multicolor Cambouliu, An. (Cel.) pharoensis Theobald, An. (Cel.) pretoriensis

(Theobald), An. (Cel.) rhodesiensis Theobald, An. (Cel.) sergentii (Theobald), An. (Cel.)

Superpictus Grassi, An. (Cel.) turkhudi Liston, Culex (Maillotia) arbieeni Salem, Cx. (Mal.)

deserticola Kirkpatrick, Culiseta (Allotheobaldia) longiareolata (Macquart), and

Uranotaenia (Pseudoficalbia) unguiculata Edwards. Kirkpatrick (1925) found adults of

82 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Sinaiticus in a rest-house and reported that females bite at night. Nothing else is known

about the bionomics of the adults, but the species does not appear to be involved in

disease transmission.

Distribution. Culex sinaiticus is known from the Arabian Peninsula, the Sinai

Peninsula of Egypt, Israel, Jordan, Iran, Sudan, Ethiopia, Somalia, and Socotra.

Material examined. 196 specimens. A total of 171 specimens (38 females,

40 males, 19 male genitalia,12 pupal exuviae, 10 larval exuviae, and 52 fourth-instar

larvae) were examined from southwestern Asia — EGYPT (including type specimens):

Sinai Peninsula (Ain Gedeirat, Ain Musa, Moweilleh, Wadi Feiran, unknown localities);

IRAN: (Hajiabad, Sarbaz Zahedan, Shah-Kahoor Zahedan); ISRAEL: (En Avedat,

Kadesh Baruea, Neot Hakikar, St. Caltrie, Tel Nagila, unknown localities); JORDAN:

(Amman, Aqaba, Wadi Hasa); OMAN: (Muscat); P.D.R. YEMEN: (Harshiyat, Hami, Wadi

Ma’adin); SAUDI ARABIA: (Al Hasa); YEMEN ARAB REPUBLIC: (unknown locality). An

additional 25 specimens (5 females, 9 males, 7 male genitalia, and 4 fourth-instar larvae)

were examined from ETHIOPIA, SOCOTRA, and SUDAN.

Culex (Culex) duttoni Theobald

duttoni Theobald, 1901c: v. tLectotype female, hereby designated,

bearing following data: "Type // Culex / Duttoni [sic] / (Type)

Theobald // C. No Il /fr. larvae / 15. 4. 00 / Canoes Creek / Duke Town

[Nigeria]" (BM).

dissimilis Theobald, 1901a: 376. tLectotype male, hereby designated,

bearing following data: "Mt. Aureol, / Free Town. / 9. 9. 99. / Bred fr.

water / in drain. // Type // Free Town. / Sierra Leone. / VIll. 99. / E.E.

Austen. / 99. 267. // Culex / dissimilis / (Type). Theobald" (BM).

Synonymy with duttoni by Edwards, 1911: 259.

hirsutipalpis Theobald, 1901a: 378. +Lectotype male, hereby

designated, bearing following data: "Type / SALISBURY /

Mashonaland [Rhodesia] / Feb. 1900 / G.A.K. Marshall / 79 / Culex /

hirsutipalpis / (Type) Theobald." (BM). Synonymy with duttoni by

Edwards, 1911: 259.

anarmostus Theobald, 1903a: 170. +Holotype female: Freetown,

Sierra Leone (BM). Synonymy with duttoni by Edwards, 1911: 259.

bifoliata Theobald, 1905c: 31. +Lectotype male, hereby designated,

bearing following data: "Type // Culex / bifoliata. / Type. F.V.T. //

Transvaal / Dr. Simpson // 54~ // Recd. from / F.V. Theobald, /

1907-29." (BM). Synonymy with duttoni by Edwards, 1941: 300.

condylodesmus Grunberg, 1905: 385. Syntypes (male; female):

Victoria, Kamerum, [Nigeria] (ZM). Synonymy with duttoni by

Edwards, 1911: 259.

albovirgatus Graham, 1910: 264. *+Lectotype male, hereby designated,

bearing following data: "Yaba / 26-5-09 / from larva in/ a fetish pot /

WMG // Type // Yaba, / Lagos, / W. Africa. [Nigeria] / 26. V. 1909. / Dr.

W.M. Graham. / 1910-80." (BM). Synonymy with duttoni by Edwards,

1911: 259.

Culex dissimilis Theobald of Wesche, 1910: 40 (W. Africa, P, L keys, .P* i"),

Culex duttoni Theobald of Wesche, 1910: 34 (W. Africa, P, L keys, P*, L*); Lewis, 1945:

Harbach: Subgenus Culex in Southwestem Asia 83

15 (Sudan, distr.).

Culex (Culex) duttoni Theobald of Edwards, 1941: 300, 418, 482 (Afrotropical Region,

A key, M*, F*, P*, syn., distr.); Lewis, 1943a: 281 (Eritrea, coll. rec., L bionomic

note); Lewis, 1943b: 69, 72 (Sudan, L habitat, bionomic note); Abbott, 1948: 45

(Sudan, distr.); Hopkins, 1952: 287 (Afrotropical Region, L key, L*, bionomics);

Lewis, 1956: 707 (Sudan, Eritrea, coll. sites); Mattingly and Knight, 1956: 94-135

(P.D.R. Yemen, A, L keys, distr. bionomics); Ovazza et al., 1956: 168 (Ethiopia,

coll. sites); Mekuria, 1968: 78 (Ethiopia, distr., rec.); Harbach, 1985a: 86, 92, 102

(SW Asia, distr., A, L keys).

Adult. A rather large species easily distinguished from all other species of Culex

known to occur in southwestern Asia by the presence of an indefinitely ringed

proboscis, ringed tarsi, postspiracular scales, an anterior white stripe on the midtibia, and

usually 2 to 4 lower mesepimeral setae.

FEMALE. Head: Antenna dark, length about 2.4 mm; pedicel pale laterally,

mesal surface usually with some pale scales; flagellum normal, most whorls with 5 or 6

setae. Proboscis black-scaled at base and apex, middle with ring of whitish scales

beginning 0.2-0.4 from base and extending to point about 0.7 from base, proximal

boundary of ring ill-defined, distal boundary sharply defined, ring sometimes incomplete

or nearly so on dorsal surface; length about 2.2 mm. Maxillary palpus mainly

black-scaled, palpomere 4 with dorsomesal stripe of white scales; length about 0.5 mm,

about 0.2 length of proboscis. Forked scales of vertex mainly pale, darker laterally;

falcate scales mainly pale, usually yellowish white in middle, somewhat brownish laterally

and white along margins of compound eyes; lateral spatulate scales white, often dark

posteriorly and in middle of lateral surface. Thorax: Pleural integument brown, paler on

middle of mesokatepisternum and posterior area of mesepisternum, pale areas and

scale-patches separate dark areas to give pleura a spotted appearance; scutal

integument dark brown. Scutum with fine, rather densely matted falcate scales, scales

mainly golden brown to brown but with an indefinite and irregular mottling of pale yellow

to golden scales, usually with whitish scales on anterior and lateral margins, prescutellar

area and forming indefinite middorsal pair of submedial spots; scutal setae prominent

only on supraalar area. Scutellar scales same as prescutellar scales; lateral lobes each

usually with 4 or 5 large setae, median lobe normally with 8 large setae. Antepronotum

with upper and lower patches of falcate scales, upper patch golden to golden brown,

lower patch paler, usually whitish. Postpronotum mainly with golden-brown scales,

scales usually whitish posteriorly; with 6-9 dark setae in posterodorsal row, often paler

posteriorly. Pleural setae golden to golden brown: 5-9 upper proepisternal, 12-14

prealar, 5 upper mesokatepisternal, 5-7 lower mesokatepisternal, 6-12 upper mes-

epimeral and 1-4 (usually 2; often 3) lower mesepimeral. Pleural scales white, anterior

part of postspiracular area with falcate scales, other areas with narrow spatulate scales as

follows: few below upper proepisternal scales, upper and lower patches on meso-

katepisternum and anterior and upper patches on mesepimeron. Wing: Length 4.2-4.5

mm, mean 4.4 mm; length of cell R2/length of vein R2+3 3.0-3.9, mean 3.4; subcosta

intersects costa just before furcation of R2+3; length of cell M1/length of cell R2 slightly

more than 0.8; dark-scaled with line of pale scales on posterior margin of costa at base.

Dorsal scaling: narrow spatulate scales on costa, subcosta, R and R1; very narrow

spatulate scales on R4+5, M1, M2, mcu, M3+4, CuA and approximately proximal 0.5 of

1A; linear scales on Rs, R2+3, R2, R38, M, M1+2 and distally on 1A. Ventral scaling:

narrow spatulate scales on costa, subcosta, Rs, R1+2, M, M1+2, CuA proximal of mcu and

proximal 0.5 of 1A; linear scales on R, R1, R2, R38, R4+5, mou, M344 and CuA

84 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

beyond mcu. Halter: Entirely pale. Legs (midleg, Fig. 2H): Mainly with dark brown to

black scales. Anterior surface of forecoxa mainly dark-scaled, with basal and distal

patches of white scales; anterolateral margin of midcoxa with longitudinal stripe of white

scales margined anteriorly by longitudinal stripe of dark scales, white stripe broadest at

base, dark stripe broadest at apex; integument of hindcoxa nearly colorless, lateral

surface with longitudinal stripe of white scales and small spot of dark scales at apex.

Trochanters mainly with white scales, anteroventral surface of fore- and midtrochanters

usually with some dark scales. Femora each narrowly pale at base and with indistinct

narrow pale knee spot; forefemur with narrow anteroventral line of pale scales on

proximal 0.5, posterior surface with posterodorsal pale stripe and more or less dintinct

narrow median line of pale scales; posterior surface of mid- and hindfemora pale-scaled

except on distal 0.3 or less. Tibiae each with narrow white ring at base; posterior surface

of foretibia with complete white stripe; midtibia with narrow white anterodorsal and

broader posterior white stripes; hindtibia with incomplete posterior pale stripe. Tarsi with

narrow white rings across joints, rings may be indistinct or absent between tarsomeres 4

and 5; tarsomere 1 of each leg with narrow basal white ring, tarsomere 1 of fore- and

midlegs with pale scales on posterior surface, hindtarsomere 1 with pale scales on

ventral surface. Pulvilli distinct, white. Ungues dark, small, simple. Abdomen: Tergum |

with median posterior patch of black scales; terga II-VIIl mainly dark brown- to

black-scaled, tergum II with small basomedian triangular patch of cream-colored scales,

terga IlI-VIII with narrow, convex, cream-colored bands 0.20-0.35 tergum length and

elongate basolateral white patches which become longer on more posterior terga,

bands and spots not connected on anterior terga, narrowly if at all connected on terga

V-VIII, band usually present and continuous with spots on tergum VIII. Sterna II-VII mainly

with cream-colored scales, with black scales on posterior corners; sternum VIII with lateral

patches of cream-colored scales, broad median area without scales.

MALE. Like female except as follows. Head: Antenna strongly plumose, pale

between flagellar whorls; length about 1.8 mm. Proboscis with narrow pale ring

0.55-0.70 from base; false joint about 0.65 from base; length 2.3-2.9 mm, mean 2.5 mm.

Length of maxillary palpus 3.1-3.4 mm, mean 3.3 mm, extending beyond tip of

proboscis by more than length of palpomere 5; palpus mainly black-scaled, integument

between palpomeres 2 and 3 pale; palpomere 3 white-scaled in middle, ventromesal

margin at apex with few white scales, ventral surface without scales, with ventrolateral

row of about 20 long black setae on distal 0.5 and complete or nearly complete

ventromesal row of short antrorsely curved setae; palpomeres 4 and 5 with small dorsal

and ventral patches of white scales at base, ventral surface of palpomere 4 also with

some white scales just beyond middle, palpomere 5 also with dorsomesal line of white

scales on distal 0.4-0.5; lateral and mesal surfaces of palpomeres 4 and 5 densely

setose. Thorax: Upper proepisternal setae more numerous, 16-26. Wing: Length

3.8-4.0 mm, mean 3.9 mm; length of cell R2/length of vein R2+3 1.8-2.4, mean 2.1;

subcosta intersects costa well before furcation of R2+3. Abdomen: Terga without

basolateral patches of white scales, basal pale bands normally 0.35-0.45 tergum length;

tergum II with basomedian triangular pale patch or narrow basal pale band produced

posteriorly in middle; basal bands of terga III-VIl usually straight, sometimes slightly

produced posteriorly in middle on terga V-VII, bands of terga V-VII and occasionally IV

produced posteriorly along lateral scale-free areas, particularly on terga VI and VII;

tergum VIII (ventral in position ) with broad concave basal whitish band. Sternum VIII

(dorsal in position) variable, similar to tergum VII but basal band sometimes obsolescent

in middle or produced medially to posterior border. Genitalia (Fig. 29): Ninth tergal lobe

poorly developed, with 3-8 (mode 6) rather widely spaced setae. Gonocoxite rather long

Harbach: Subgenus Culex in Southwestern Asia 85

and slender, mesal and ventral setae developed as usual, lateral setae in 3 or 4

longitudinal rows, these shorter in each row toward large ventral setae; subapical lobe

indistinctly divided, setae a-c equally developed, long and tapered with hooked tips; d-e

slender with hooked tips, f flattened with hooked tip; g more or less symmetrical, rather

narrow, rounded apically, borne on distinct prominence; h straight or bent apically,

tapered with narrow pointed distal part. Gonostylus normal; gonostylar claw flat, slightly

broader distally. Phallosome as broad as long with lateral plate and aedeagal sclerite

nearly completely fused into a single simple sclerite, crest of aedeagal sclerite not

distinguished, dorsal arm of lateral plate absent, ventral arm a long slender dorsally

directed and laterally bent sickle-shaped process. Proctiger slightly elongated;

paraproct long and slender, without basal lateral arm, crown of spinelike spicules grading

into short blades and series of bumps laterally. Cercal sclerite with ribbonlike

caudolateral extension joining paraproct near base of crown; 3-6 (mode 4) cercal setae.

Tergum X developed as usual.

Pupa (Fig. 29). Not definitely associated with adults; form and placement of

setae as figured, range and modal number of branches in Table 14; immediately

distinguished from all other species of Culex in the region by the vesiculation of the

scutum, metanotum, and first two abdominal terga. Cephalothorax: Lightly tanned,

scutum and metathorax moderately tanned; middle of scutum behind trumpets and

metanotum covered with distinct vesicles. Setae 1,2,8,9-CT frequently double,

sometimes triple; 3,4,6,11,12-CT often triple (2-5); 7-CT double. Trumpet: Lightly to

moderately tanned, tracheoid area darker; relatively short and flared, index 3.3-4.5,

mean 3.8; pinna with distinct shallow rounded emargination at tip. Abdomen: Lightly

tanned, terga I-IV darker medially; middle of terga | and II and intersegmental membrane

between terga Il-Ill and Ill-IV with conspicuous vesicles. Seta 6-I,II short as in /aticinctus

and mattinglyi, 7-|,I| usually with 3 or 4 branches, 7-| occasionally double or with 5

branches; 9-ll frequently anterior to 7-Il; 1-III,IV most often with 4 or 5 branches (see

table), 1-V-VII variable (see table) but usually double or triple, more often double; 5-IV

usually multiple (1-4) and about length of following tergum, very different in appearance

from 5-V,VI which are usually single and much longer than following tergum; 6-IlI-V

normally single, seldom double (6-V triple on 1 side in 1 specimen examined), 6-VI most

often triple (1-3). Genital lobe: Lightly tanned; length 0.2-0.3 mm in female, about 0.4

mm in male. Paddle: Very lightly tanned, buttress and midrib darker; midrib with obvious

curvature, inner part not as broad as outer part; length 0.98-1.16 mm, mean 1.07 mm,

width 0.72-1.02 mm, mean 0.88 mm, index 1.1-1.4, mean 1.2. Seta 1-P usually double,

sometimes single; 2-P almost always absent (double on 1 paddle in 1 specimen

examined).

Larva (Fig. 30). Not definitely associated with adults; positions and attributes of

setae as figured, range and modal number of branches in Table 34; easily recognized by

the unusual siphon. Head: Length 0.87-1.07 mm, mean 0.95 mm; width 1.14-1.54 mm,

mean 1.35 mm; moderately tanned, darker posteriorly. Labiogula very nearly

parallel-sided; hypostomal suture rather poorly indicated but complete, extended to

collar from posterior tentorial pit. Dorsomentum more strongly produced anteriorly in

middle than usual; with 12-15 teeth on each side of median tooth, teeth progressively

larger toward sides. Seta1-C long, slender, tapered, and tanned; 2-C absent; 5,6-C

usually with 7 or 8 branches (see table); 7-C usually with 8 or 9 branches (7-11); 8-C

normally double, seldom single; 9-C variable (1-6); 10-C forked, usually bifurcate,

infrequently trifurcate. Antenna: Length 0.53-0.57 mm, mean 0.55 mm; lightly to

moderately tanned, distal part and basal rim darker. Seta 1-A with about 23 aciculate

branches (18-26); 2,3,4-A of equal length, distinctly aciculate. Thorax: Integument

86 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

hyaline; rather sparsely covered with fine spicules. Seta 4-P double or triple; 7-P

double; 8-P most often with 4 branches (3-6). Seta 1-M single; 2-M variable (1-4); 6,7-M

on common tubercle. Seta 1-T very small, usually double (1-3); 2-T long, single.

Abdomen: |Integument hyaline; smooth. Seta 3-I-VIl single, 3-VIl sometimes double; 6-|

usually double, infrequently triple, 6-IIl not as strong as 6-I, with 2-4 branches occurring in

nearly equal frequencies; 7-| most often triple (2-5), nearly as long as 6-l, 7-Il more

strongly developed than in other species, resembling 7-| but shorter, with 3 or 4

branches; 5-IlI-VIl and 6-III-VI long and single, 5-III-V1 distinctly longer than 2 abdominal

segments; 13-lll-V normally double (see table). Segment Vill: Comb with 25-44 (mode

42) rather short fringed scales. Setae 1,5-VIll distinctly shorter than 2,4-VIll, most often

with 5 or 6 branches. Siphon: Index 3.37-4.24, mean 3.70;strongly inflated, narrowed

distally, with short spine anteriorly near apex; moderately tanned, with dark subapical

band, basal rim heavily tanned. Pecten reduced, beginning well above base of siphon,

composed of 3-5 (Hopkins, 1952 reported 3-6) slender spines with ventral row of short

denticles. Seta 1-S in 4 or 5 pairs, usually 4, 1c-S often absent; 1a,b,c,d-S long and

single (1d-S double on 1 side in 1 specimen examined), 1e-S usually triple (1-4); 1a-S

within pecten, 1d-S either within or below dark band, 1e-S within band. Segment xX:

Relatively small; saddle complete, length 0.44-0.51 mm, mean 0.48 mm, siphon/saddle

index 3.76-4.63, mean 4.09. Seta 1-X long and single; 2-X single; 4-X in 6 pairs. Anal

papillae stout, about twice length of saddle, ventral pair shorter than dorsal pair.

Systematics. Culex duttoni is a bizarre species which combines features of the

pipiens and sitiens groups and is recognized here as the name-bearing member of a

separate group. The group also includes the closely related Culex watti Edwards, 1920.

These are rather large species which are characterized by the indefinitely ringed

proboscis, ringed tarsi, and the usual presence of two to four lower mesepimeral setae in

the adult. The male genitalia are distinctive: the lateral plate is a simple sclerite without a

dorsal arm, the ventral arm is enormously developed, yet reminiscent of the type found

in the pipiens and trifilatus subgroups; and the paraproct has no trace of a basal lateral

arm. The larva resembles members of the pipiens group in most respects, but setae 1-C

and 3-P and the pecten spines approach the condition in the sitiens group and setae

8-P, 7-Il, and 1-S are uniquely developed. The distinct form of the larval siphon is

suggestive of that in Culex hutchinsoni Barraud, 1924b of the Oriental Region. The

larva of watti has never been described, but is said to be indistinguishable from duttoni

(Hopkins, 1952). The pupal stage seems to differ from most members of the pipiens

and sitiens groups in having seta 7-I,Il usually with 3 or 4 branches, setae 6-IIl-V and

5-V,VI usually single, 1-P usually double, and 2-P normally absent. The pupa of watti is

unknown.

The association of the immature stages with the adults of this species is

presumptive only. No individual rearings are available, but the association seems

correct. It is based on the sympatric occurrence of larvae and adults in the P.D.R. Yemen

and associated larval and pupal exuviae from Ghana. There is a possibility, however, that

the unknown larva and pupa of wattiwere mixed in the material examined.

Bionomics. Culex duttoni is a semidomestic species which uses a wide variety

of breeding places. It has been found in pools in streams, ground pools (including

puddles), and a variety of artificial containers, including drums, pots, sunken barrels,

boats, and wells. It is possible that the dispersal of this species to the Arabian Peninsula

was accomplished by man. The occurrence of this species in southwestern Asia is

based on two larvae and two adults from the P.D.R. Yemen. The adults were captured in

acave. According to Lewis (1943b), females are never seen biting man. Culex duttoni

is probably not a medically important species.

Harbach: Subgenus Culex in Southwestem Asia 87

Distribution. Culex duttoni is widely distributed in the Afrotropical Region. It

has been recorded and described from a single larva collected in the south of Morocco”

(Bailly-Choumara, 1968).

Material examined. 85 specimens. Only 5 specimens (1 female, 1 male, 1

male genitalia, and 2 fourth-instar larvae) were examined from southwestern Asia —

P.D.R. YEMEN: (Mlkhuras, Wadi Gheil el Khumeira). A total of 80 specimens (36

females, 22 males, 5 male genitalia, 9 pupal exuviae, 4 larval exuviae, and 4 fourth-instar

larvae) were examined from ANGOLA, BENIN, CAMEROON, ETHIOPIA, GHANA,

KENYA, MALAWI, NIGERIA (including the type specimens of duttoni and albovirgatus ),

SENEGAL, SIERRA LEONE (the type specimens of dissimilis and anarmostus ), SOUTH

AFRICA (the lectotype male of bifoliata ), SUDAN, TANZANIA, UGANDA, ZIMBABWE

(the type specimens of hirsutipalpis ), and unknown localities.

Culex (Culex) sitiens Wiedemann

sitiens Wiedemann, 1828: 542. Holotype female: Sumatra, Indonesia

(ZMC).

microannulatus Theobald, 1901a: 353. +Lectotype male: Quilon,

Travancore, [Madras State], India (BM), designated by Bram, 1967a:

243. Synonymy with sitiens by Edwards, 1913c: 232.

somaliensis Neveu-Lemaire, 1906: 254. Syntypes (male; female):

Djibouti, Djibouti (non-extant?). Synonymy with sitiens by Edwards,

1913¢):232.

salus Theobald, 1908b: 256. +Lectotype male, hereby designated,

mounted on two slides (see Harbach, 1983: 104) bearing following

data: Slide 1 (with maxillary palpi, proboscis, and part of head cap-

sule) — "The / Salt Water. / Mosquito. / TYPE // Culex / salus. /

Theobald / Port Sudan // Recd. from F.V. Theobald. / 1910-396":

Slide 2 (with genitalia) — "Salt water / mosquito. / TYPE // Culex /

salus / Theobald / Port Sudan. // Recd. fr[om] / F.V. Theo[bald] /

1910-395." (BM). Synonymy with somaliensis by Edwards, 1911:

261.

For complete synonymy, see Appendix D.

Culex annulus Theobald of Becker, 1907(1910): 140 (Socotra, A).

Culex impellens Walker of Giles, 1901a: 606 (India, A*, L*, distr.); Theobald, 1901a: 362

(India, F"); Christophers, 1906: 8 (? India, L*).

Culex sitiens Wiedemann of Edwards, 1913c: 232 (Oriental Region, syn., tax., A);

Edwards, 1922: 277, Pl. VI (Oriental Region, A key, M gen.*); Barraud, 1924a:

993 (India, A key, syn., M*, F, distr.); Barraud, 1924c: 427 (India, L*); Lewis, 1945:

15 (Sudan, L, distr.); Leeson and Theodor, 1948: 227 (Socotra, A, L keys, L, L

bionomics, coll. sites).

Culex (Culex) sitiens Wiedemann of Barraud, 1934: 398 (India, A, L keys, M’*, F, L*, L

habitat, distr., syn.); Edwards, 1941: 296, 417, 482 (Afrotropical Region, A key,

M*, F, P, syn., distr.); Hopkins, 1952: 284 (Afrotropical Region, L key, L,

bionomics); Knight, 1953a: 231 (Yemen Arab Republic, A, L, distr., L habitat);

Lewis, 1956: 707 (Sudan, L, coll. rec., distr.); Mattingly and Knight, 1956:

103-137 (Arabian Peninsula, L*, A, L keys, distr., bionomics); Ovazza et al., 1956:

167 (Ethiopia, M gen.*); Lotfi, 1970: 401 (Iran, coll. rec.); Aslamkhan, 1971: 155

88 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

(Pakistan); Lotfi, 1973: 206 (Iran, coll. rec.); Lotfi, 1976: 72, 75, 81 (Iran, L key,

ecol. note, L*); Sirivanakarn, 1976: 95 (Oriental Region, A, P, L keys, M*, F*, P*,

L*, tax., distr., bionomics); Harbach, 1985a: 86, 97, 102 (SW Asia, distr., A, L

keys).

Culex somaliensis Neveu-Lemaire of Edwards, 1911: 261 (Africa, A, key); Edwards,

1912c: 28 (Africa, A key).

Culex thalassius Theobald of Leeson and Theodor, 1948: 228 (in part; Socotra, L key).

Adult. A medium-sized species distinguished from all other Culex occurring in

southwestern Asia by having speckled femora and base of cell R2 distal to base of cell

M1.

FEMALE. Head: Antenna blackish; pedicel yellow, mesal surface dark brown to

black with tiny setae and scales; length 1.7-2.2 mm, mean 1.9 mm. Proboscis dark, with

distinct median whitish ring about 0.2 proboscis length, extending from 0.4 to 0.6 from

base; frequently with scattered pale scales proximal to ring, particularly ventrally;

sometimes also with scattered pale scales faintly evident distal to ring; with indefinite

narrow pale fringe of scales at apex of prementum; length 1.8-2.1 mm, mean 1.9 mm.

Maxillary palpus dark with pale scales laterally at apex, sometimes also with pale scales

laterally at base or on entire lateral surface; palpomeres 2 and 3 with 3-5 long lateral

setae; length about 0.6 mm, approximately 0.3 proboscis length. Forked scales of

vertex pale yellow to yellowish brown, darker posterolaterally; falcate scales pale yellow

to brownish, darker laterally; interocular scales yellowish white and projecting ventrally

between eyes; lateral spatulate scales yellowish white. Thorax: Integument dark brown

to black. Scutum mottled with variable spots and blotches of coarse dark and pale

scales; dark brown to golden-brown scales usually predominant with pale yellow to

golden scales on anterior and lateral margins, across middie and on prescutellar area;

acrostichal and dorsocentral setae short, slender and indistinct; supraalar setae robust

and dense. Scutellum with rather numerous pale yellow falcate scales; lateral lobes each

with 4 or 5 strongly developed setae, 6-9 weaker ones; median lobe with 5-8 large

setae, 9-12 smaller ones. Antepronotum with coarse yellowish falcate scales, often with

some brownish ones in middle; postpronotum with dark brown or golden-brown scales

in middle, pale yellow scales anteriorly and posteriorly, posterior scales coarser.

Proepisternum with elongate subfusiform scales laterally below upper proepisternal

setae; shorter apically rounded pale yellow spatulate scales among and extending mesal

to setae. Mesopleuron with white spatulate scales: distinct upper and lower patches on

katepisternum, distinct anterior patch on anepimeron, and few scales usually present

among upper mesepimeral setae. Numbers of pleural setae: 6-8 upper proepisternal,

9-17 prealar, 4-8 upper mesokatepisternal, 5-10 lower mesokatepisternal, 7-12 upper

mesepimeral, no lower mesepimeral. Wing: Length 3.2-4.0 mm, mean 3.7 mm; length of

cell R2 2.1-2.5 length of vein R2+3, mean 2.3; length of cell M1/length of cell R2 about

0.9; furcation of vein R2+3 normally distal to furcation of vein M1+2; mainly dark-scaled,

with pale spatulate scales along posterior margin of costa (except at apex), posterior

margin of approximately distal 0.5 of subcosta, and anterior margin of R1; usually with

small spot of pale scales at base of remigium. Ha/ter: Pedicel and scabellum pale;

capitellum dark, with white scales. Legs: Anterior surface of forecoxa white-scaled, with

few pale scales posterolaterally at apex; midcoxa with broad longitudinal patch of white

scales on anterior side of lateral midline, extending onto anterior surface distally;

hindcoxa with slightly curved lateral longitudinal line of pale scales. Trochanters with

white spatulate scales on ventral and posterior surfaces; mid- and hindtrochanters also

with white scales on dorsal surface. Femora with whitish knee spots; anterior surface of

Harbach: Subgenus Culex in Southwestern Asia 89

forefemur predominantly dark-scaled with more or less definite whitish stripe on proximal

0.5, distal 0.5 speckled with pale scales, posterior surface whitish-scaled; anterior —

surface of midfemur dark-scaled exept at base, speckled with pale scales, posterior

surface whitish-scaled; hindfemur primarily whitish-scaled, with narrow band of dark

scales at apex, sometimes with indistinct dorsal dark stripe extending proximad from

band, frequently with scattered dark scales on anterior surface. Dorsal surface of

foretibia dark-scaled with indistinct spot of pale scales at apex, pale-scaled ventrally;

anteroventral surface of midtibia with dark scales confluent with patch of dark scales on

0.25 to 0.50 of dorsal surface except at extreme base, with spot of pale scales dorsally at

apex; hindtibia mainly dark-scaled dorsally, with dark scales dissipated in middle and

extended over sides at ends, with anterodorsal spot of pale scales at apex. Tarsi mainly

dark-scaled; tarsomere 1 pale-scaled ventrally; joints with narrow whitish rings mainly on

bases of tarsomeres, rings distinct between tarsomeres 1-2 and 2-3, often indistinct

between 3-4 and 4-5, tarsomere 1 with narrow pale ring at base; hindtarsomere 2 faintly

pale beneath. Abdomen: Tergum | with median posterior patch of scales, scales usually

dark anteriorly and pale posteriorly; terga II-VIII with basal white bands continuous with

elongate basolateral spots of same color, band of tergum II produced in middle, others of

even width (0.2-0.4 length of tergum); terga II-VIIl also with very narrow white bands on

posterior margin, that of tergum VII often strongly produced in middle and occasionally

united with basal band. Sternum | bare; sternum II usually somewhat sparsely clothed in

white scales, particularly on median and lateral areas; sterna III-VIIl white-scaled with

posterolateral dark spots, median area of posterior margin with yellowish scales; sternum

VIII with lateral patches of white scales, median area void of scales.

MALE. Like female except as follows. Head: Proboscis with ventral cluster of

setae at base of pale ring; false joint about 0.55 from base of proboscis. Length of

maxillary palpus about 2.1-2.8 mm, mean 2.6 mm, extending beyond tip of proboscis by

length of palpomere 5; integument of palpomere 2 pale apically, with pale scales on

dorsal and lateral surfaces; palpomere 3 with broad median pale ring comprised of

whitish scales on dorsal and lateral surfaces and pale bare integument on ventral and

mesal surfaces, ventromesal margin with row of about 25 short apically curved and

flattened setae on distal 0.8, ventrolateral margin beyond median pale ring with about 15

long setae; palpomeres 4 and 5 with basal pale rings, densely setose; palpomere 5 with

apical 0.5 or less (usually 0.383) pale-scaled. Thorax: Upper proepisternal setae more

numerous, 9-12. Wing: Pale scales usually absent at base of remigium. Abdomen:

Terga without basolateral pale spots; basal bands of terga V-VII produced posteriorly

along lateral scale-free areas; tergum VII (and occasionally VI) with large triangular

posteromedian pale spot, spot often connected medially to basal band; tergum VIII

(ventral in position) with broad basal pale band. Sterna | and II same as in female; sterna

III-VIl with narrow posterior band of white scales, rather broad subapical dark band and

broad basal pale band, dark band of more posterior sterna often divided in middle by

narrow longitudinal line of pale scales connecting basal and apical bands; sternum VIII

(dorsal in position) like tergum VII. Genitalia (Fig. 31). Ninth tergal lobe poorly

developed, narrowly connected with its mate of the opposite side, with line of 2-10

(mode 6) short setae. Gonocoxite normal, lateral area with 3 or 4 rows of moderately

developed setae extending from above base to point just above subapical lobe;

subapical lobe undivided, setae a-c rodlike as usual, a shorter than b andc and blunt at

tip, band c hooked at tip; setae d-e with small barblike denticle before recurved tip, f

hooked at tip; seta g a simple, symmetrical, apically rounded leaf; h about length of g,

flattened, bent distally. Gonostylus rather stout basally, distinctly tapered apically,

slightly rugose dorsally before tip; gonostylar claw short, broadened apically, troughlike.

90 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Phallosome not much longer than broad, aedeagal sclerite shorter than lateral plate;

lateral plate with definite inner and outer divisions, inner division (= ventral arm) a broad

minutely spiculate lobe borne on ventromesal margin of outer division (= lateral arm),

ventrocaudal angle rounded, dorsocaudal angle rather pointed, dorsal margin with 3

prominent caudally directed teeth before apex; outer division rather simple,

posteromesal margin with a conspicuous laterally curved tooth which appears broad in

lateral view, ventrocaudal angle giving rise to a heavy flattened laterally directed and

dorsally curved pointed process, dorsal process a prominent dorsolaterally projecting

earlike lobe; dorsal arm absent (unless represented by the posteromesal tooth of the

outer division). Proctiger unmodified; paraproct with moderately long, gently curved

basal lateral arm and distinct ventral acetabulum. Cercal sclerite more or less rectangular;

2-5 (mode 3) cercal setae present. Tergum X developed as usual.

Pupa (Fig. 31). Character and positions of setae as figured, range and modal

number of branches in Table 15; without obvious distinctive features, most closely

resembling mimeticus from which it can usually be separated by the larger number of

branches of setae 1-ll, 6-III-VI, and 9-VIIl. Cephalothorax: Lightly tanned, legs and

metathorax darker. Setae 1-9,11,12-CT usually double or triple (see table); 1-CT longer

and slightly stronger than 2,3-CT; 10-CT frequently with 7 or 8 branches (5-9), distinctly

metathorax darker. Setae 1-9,11,12-CT usually double or triple (see table); 1-CT longer

and slightly stronger than 2,3-CT; 10-CT frequently with 7 or 8 branches (5-9), distinctly

longer than 11,12-CT. Trumpet: Moderately tanned, tracheoid area only slightly darker;

index 4.1-5.7, mean 4.7; pinna long, 0.3-0.4 trumpet length. Abdomen: Lightly

tanned, anterior terga darker; length 3.1-3.7 mm, mean 3.5 mm. Seta 7-I triple, 7-II

double or triple, more often double; 1-II with 4-10 aciculate branches; 1-III-VI moderately

developed, with fewer branches on each successive posterior tergum, 1-IIl usually with 5

or 6 branches (5-8), 1-IV normally with 4 or 5 branches (3-5), 1-V with 3 or 4 branches,

1-VI double or triple; 6-III-VI single or double, more often double, 6-V,VI larger than

6-I11,1V; 5-IV-VI equally developed, usually double, longer than following tergum; 9-VIII

commonly with 10-12 branches (9-14). Genital lobe: Lightly tanned; length about 0.20

mm in female, about 0.35 mm in male. Paddle: Lightly tanned, buttress and midrib

darker; length 0.90-1.00 mm, mean 0.94 mm, width 0.68-0.80 mm, mean 0.74 mm,

index about 1.3.

Larva (Fig. 32). Form and placement of setae as figured, range and modal

number of branches in Table 35; bearing a strong resemblance to theileri, laticinctus,

and mattinglyi but immediately distinguished by the form of seta 1-C and the pecten

spines. Head: Length 0.72-0.88 mm, mean 0.80 mm; width 1.13-1.35 mm, mean 1.24

mm; lightly tanned, labiogula and posterior areas of lateralia and dorsal apotome darker.

Hypostomal suture strongly indicated, diverging posteriorly from its mate of the opposite

side, ending about halfway between posterior tentorial pit and collar. Dorsomentum with

7 or 8 teeth on either side of center. Seta 1-C short, thick, usually spiculate at sides,

dark; 2-C present or absent; 5-C usually with 6 or 7 branches (6-8); 6-C often with 4, 5 or

6 branches (4-7); 7-C frequently with 9 or 10 branches (7-11); 10,11,12,15-C double or

triple, 10,11-C occasionally with 4 branches; 13-C normally double (1-3). Antenna:

Length 0.41-0.56 mm, mean 0.49 mm; lightly tanned, distal part darker, with dark ring at

base; proximal part with aciculae on dorsal and lateral surfaces, distal part with few

spicules laterally. Seta 1-A with about 27 branches (22-32); 2,3-A subapical. Thorax:

Integument hyaline, smooth. Seta 4-P double; 7-P double or triple; 8-P single or

double, distinctly shorter than 4,7-P. Seta 1-M single, about twice length of 2-M; 4-M

single or double. Seta 1-T single, nearly as long as 2-T which is usually single,

sometimes double; 13-T distinctly longer than12-T, with 2-5 branches. Abdomen:

Harbach: Subgenus Culex in Southwestern Asia 91

Integument like that of thorax. Seta 3-I usually single, seldom double, 3-II-VI single, 3-VII

usually double (1-4); 6-1,Il usually triple (2-4), 6-III-VI normally double; 7-I usually long and.

single, about as long as 6-I, seldom double; 1-III-VIl rather short, about 0.5 length of

segment, double or triple, more often double; 13-III-V,VII developed like 1-III-VII, usually

double (see table). Segment Vill: Comb with 30-54 (mean 42) rather short, fringed

scales. Siphon: Index 3.61-4.55, mean 3.80; lightly tanned, basal ring and acus darker.

Pecten composed of 9-17 (mode 13) spines with complete ventral row of denticles,

larger spines with 7-13 (mode 10) denticles, tendency for most basal spines to fuse.

Seta 1-S in 6, 6.5 or 7 pairs, 1 distal pair (1e-S) borne laterally, others imperfectly paired

near posterior midline, proximal 3 or 4 pairs often with 6 or 7 branches and about twice as

long as diameter of siphon at point of attachment. Segment X: Saddle complete; lightly

tanned; posterodorsal margin with distinct short spicules; length 0.29-0.35 mm, mean

0.33 mm, siphon/saddle index 3.03-4.29, mean 3.71. Seta 1-X single; 2-X usually with 4

or 5 branches, occasionally triple; 4-X usually in 6 pairs. Anal papillae short, globose.

Systematics. Culex sitiens is the nominotypical member of a large and variable

group of species which probably should be divided into several different groups. The

group is widespread in the warmer parts of the Old World with species occurring in the

Afrotropical, Oriental, and Australian regions, and the Mediterranean and Manchurian

subregions of the Palaearctic. The group is currently subdivided into six subgroups

(Sirivanakarn, 1976). Six members belonging to the sitiens, mimeticus,

bitaeniorhynchus, and vishnui subgroups occur in southwestern Asia. The sitiens

subgroup is represented by sitiens and poicilipes.

Culex sitiens is a highly variable species which breeds in brackish coastal waters

from East Africa to the South Pacific. It is very likely that this nominal form is actually

composed of several distinct species. There is considerable variation in all stages and

there seems to be many isolated populations. Adults from southwestern Asia have

much lighter scutal scales and more extensive pale scaling on the wings than specimens

from the Oriental Region. There is an extreme range of variation in larval characters,

especially in the size and shape of seta 1-C and the siphon, but there are no apparent

differences in the form of the male genitalia.

There should be no difficulty in identifying specimens of sitiens in southwestern

Asia unless the very similar Culex thalassius Theobald, 1903a also occurs in the

Afrotropical part of the Arabian Peninsula. Adults of this species differ in having darker

scutal scales and no speckling of pale scales on the femora. The male genitalia differ

slightly in details of the phallosome. The larva of thalassius at present cannot be

distinguished with certainty from that of sitiens. The record of thalassius from Syria

(Abdel-Malek, 1960) cannot be correct, and probably is based on misidentified larvae of

tritaeniorhynchus.

Bionomics. Culex sitiens breeds primarily in coastal brackish ground pools,

mangrove swamps, and salt marshes, but is also found in artificial containers and

collections of fresh water not far from the seacoast. Specimens from the Yemen Arab

Republic have been collected in open cement basins some distance inland and in

brackish water seepage pools near the seashore (Knight, 1953a). Females feed mainly

on birds and pigs (Colless, 1959), but are also known to bite cattle, dogs, and man.

Culex sitiens is a potential vector of human pathogens. Experimental transmission of

Japanese encephalitis virus has been demonstrated in the laboratory (Hodes, 1946, as

jepsoni Theobald) and it has been found naturally infected with larvae of Brugia malayiin

Thailand (lyengar, 1953).

Distribution. This species occurs in the coastal lands of the Oriental Region,

southwestern Asia, eastern Africa, and Madagascar. Populations are also known from

92 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

the Ryukyu Archipelago, Korea, northern Australia, and many islands in the South

Pacific.

Material examined. 544 specimens. A total of 455 specimens (90 females,

85 males, 31 male genitalia, 105 pupal exuviae, 105 larval exuviae, and 39 fourth-instar

larvae) were examined from coastal areas of southwestern Asia —- OMAN: (Dhofar,

ltelba, Salalah, Trucia); P.D.R. YEMEN: (Al Ais, Buaish, Dis Town, Geregeri, Gheil

Bawazir, Hami, Hamial Sharg, Harshiyab, Khon, Makhzan, Meifa, Mukalla, Qaiti, Shihr,

Sukhal al Dis, Thrilla, Urfat Subai, Wadi Maseila, Wasira, unknown localities); SAUDI

ARABIA: (Jeddah); UNITED ARAB EMIRATES: (Sharja); YEMEN ARAB REPUBLIC:

(Hodeidah). An additional 89 specimens (40 females, 15 males, 4 male genitalia, 10

pupal exuviae, 4 larval exuviae, and 16 fourth-instar larvae) were examined from

DJIBOUTI, INDIA (the type specimens of microannulatus ), SOCOTRA, SUDAN

(including the type specimens of sa/us ), and unknown localities. An unknown number

of specimens from Southeast Asia and the type specimens of impellens, gnophodes,

and jepsoni were also examined.

Culex (Culex) poicilipes (Theobald)

poicilipes Theobald, 1903b: ix (Lasioconops ). *Holotype female:

Bonny, [Eastern Provinces], Nigeria (BM).

quasigelidus Theobald, 1903a: 181. +Lectotype female, hereby

designated, bearing following data: "Type // Entebbe / Dr. Row //

Culex / quasigelidus / (Type) Theobald // Entebbe. / bred from /

larvae. / Sep. 20. 02"; left wing mounted on slide (BM). Synonymy

with poicilipes by Edwards, 1932a: 203; incorrectly considered as

senior synonym of poicilipes by Edwards, 1911:258.

taeniorhynchoides Giles, 1904: 369. +Holotype female: Benguela,

Angola (BM). Synonymy with quasigelidus by Edwards, 1911: 258.

maculipes Theobald, 1904: 79 (Taeniorhynchus tenax var.). *Holotype

female: Kenissa, White Nile and Middle Sobat, Sudan (BM).

Synonymy with quasigelidus by Edwards, 1911: 258.

madagascariensis Ventrillon, 1905b: 427 (Pseudoheptaphlebomyia).

+L_ectotype female, hereby designated, bearing following data: "9 //

MUSEUM PARIS / [Tananarive,] MADAGASCAR / VENTRILLON

1905 // Pseudohepta- / phlegomyia [sic] / madagasca- / riensis Ventr.

/ type." (MNHP). Synonymy with quasigelidus by Edwards, 1911:

256:

par Newstead, 1907 (in Newstead et al., 1907: 25). +tHolotype female:

Tshumbiri, [Leopoldville], Belgian Congo (BM). Synonymy with

guasigelidus by Edwards, 1912b: 29.

punctipes Theobald, 1907: 316 (Aporoculex ). tHolotype female:

Chinde, [Zambezia, Mozambique] (BM). Synonymy with quasigeli-

dus by Edwards, 1911: 258. :

auritaenia Enderlein, 1920: 49. tLectotype female, hereby designated,

bearing following data: "Madagaskar / Tanararivo [sic] / Friederichs

S.V. // Type // Culex / auritaenia / Type Enderl. [female symbol] / Dr.

Enderlein det. 1920 // 1920. 375." (BM). Synonymy with poicilipes

by Edwards, 1932a: 203.

Harbach: Subgenus Culex in Southwestem Asia 93

Culex poicilipes (Theobald) of Lewis, 1945: 14 (Sudan, distr.); Theodor, 1952: 113

(Egypt, zoogeogr.); Senevet et al., 1957a: 86 (North Africa, L*); El-Said and

Kenawy, 1983a (Egypt, distr.).

Culex (Culex) poicilipes (Theobald) of Edwards, 1941: 289, 415, 482 (Afrotropical

Region, A key, M*, F*, P*, syn., distr.); Lewis, 1943a: 281 (Eritrea, coll. rec.);

Lewis, 1943b: 68 (Sudan); Abbott, 1948: 44 (Sudan, distr.); Lewis, 1948:

144-148 (Sudan, A, L bionomics); Hopkins, 1952: 280 (Afrotropical Region, L

key, L", bionomics); Mattingly, 1954: 58 (Africa, zoogeogr.); Lewis, 1956: 705

(Sudan, distr., bionomics); Ovazza et al., 1956: 167 (Ethiopia, coll. sites):

Senevet et al., 1957b: 92 (North Africa, L); Senevet and Andarelli, 1959: 135

(Egypt, A, L, P keys, syn., M*, F*, P*, L*, distr., biol., L assoc.); DuBose and

Curtin, 1965: 351-354 (Mediterranean area, A, L keys); Mekuria, 1968: 78

(Ethiopia, distr.); El-Said and Kenawy, 1983b (Egypt, coll. rec., distr.); Harbach,

1985a: 86, 96, 107 (Egypt, Israel, A, L keys.).

Culex quasigelidus Theobald of Wesche, 1910: 38 (West Africa, P*, L*, keys); Edwards,

1912c: 381 (Africa, L, key); Gough, 1914: 135 (Egypt, coll. rec.); Storey,

1918(1919): 86-98 (Egypt, A, L keys, A); Ingram and Macfie, 1919: 66 (W. Africa,

P*, L); Kirkpatrick, 1924(1925): 367, 371 (Egypt, A, L keys); Scott, 1927: 88

(Ethiopia, coll. rec.); Abdel-Malek, 1956: 105 (Sinai, L key).

Culex (Culex) quasigelidus Theobald of Edwards, 1921: 337 (Egypt, A, L keys, tax.,

bionomic note, distr.); Séguy, 1924: 29, 185 (Egypt, A key, A, L, tax., distr.,

syn.); Kirkpatrick, 1925: 107, 186 (Egypt, syn., M*, F, P*, L*, distr., bionomics);

Edwards, 1926: 129 (Egypt, A, L keys, A, L, distr.); Stackelberg, 1927: 152

(Egypt, M, F keys, A, M gen.*, distr.); Martini, 1931: 380 (Egypt, A, L keys, M’, F,

L, distr.).

Adult. A rather large species easily distinguished from other species of the

subgenus Culex by the presence of rows of pale spots on the femora and tibiae. It bears

a superficial resemblance to tigripes De Grandpre and De Charmoy of the subgenus

Lutzia which also has spotted legs, but these species do not occur together in

southwestern Asia and are distinguished without difficulty by the characters given in the

key to subgenera.

FEMALE. Head: Antenna dark; pedicel with small patch of pale scales on mesal

surface; length 1.5-1.9 mm, mean 1.7 mm. Proboscis dark-scaled with sharply defined

median pale ring extending from 0.4 to 0.6 from base; length 1.8-2.1 mm, mean 1.9 mm.

Maxillary palpus dark-scaled, with pale scales at apex and also usually on dorsal surface

of palpomere 2; length 0.4-0.5 mm, about 0.25 proboscis length. Forked scales of

vertex dark; narrow falcate scales yellowish and yellowish brown; postocular and

postgenal areas with white spatulate scales, these often divided into dorsal and ventral

patches by anterior extension of posterior black scales, white patches usually connect-

ed along margin of eye. Thorax: Integument usually black, sometimes dark brown.

Scutum mainly with dark brown scales, with variable amount of whitish or yellowish to

yellowish-brown scaling, principal areas of pale scaling include median area of anterior

0.33, paired sublateral areas at middle, and transverse band behind middle; small but

conspicuous patch of long white near-linear scales on antealar area; some whitish or pale

yellow scales around bare part of prescutellar area. Scutellum largely covered with

yellowish-brown falcate scales. Ante- and postpronota with golden-brown falcate scales,

antepronotum with paler scales dorsally and/or ventrally, postpronotum with few slightly

lighter scales at extreme ventral angle. Proepisternum with lower patch of pale scales in

addition to usual upper patch, upper patch rather large, extending ventromesally on

94 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

anterior surface; with 9-13 setae in upper area. Mesopleuron with usual patches of white

spatulate scales on katepisternum and epimeron; with numbers of setae as follows:

8-14 prealar, 3-8 upper mesokatepisternal, 6-12 lower mesokatepisternal, 6-10 upper

mesepimeral, and no lower mesepimeral. Wing: Length 3.4-4.6 mm, mean 4.0 mm;

length of cell R2 3.6-4.7 length of vein R2+3, mean 4.0; length of cell M1/length of cell

R2 0.75-0.85, mean 0.81; mainly brown-scaled, with variable pale scaling; pale scaling

sometimes absent or indistinct, sometimes limited to patches at base and apex of costa,

but usually more extensive (particularly in dark individuals) as follows: patch at base of

costa before humeral crossvein; scattered along posterior margins of costa and

subcosta; line on posterior margin of costa distal to intersection of subcosta; scattered

along anterior margin of R1 before level of furcation of R2+3; lines on anterior and

posterior margins of R1 beyond furcation of R2+3. Halter: Pedicel and scabellum yellow;

capitellum dark. Legs (midleg, Fig. 2J): Anterior surface of forecoxa with mixture of dark

and pale scales, sometimes almost completely pale-scaled, often nearly all dark-scaled;

anterolateral surface of midcoxa with mixture of dark and pale scales; hindcoxa with

lateral line of pale scales. Femora with pale scales encircling base and covering posterior

surface; anterior surfaces dark brown-scaled with row of 7-9 yellowish-white spots, spots

fewer and larger on hindfemur; with narrow pale knee spots. Tibiae like femora; pale

spots more distinct on fore- and midtibiae than on hindtibia; spots slightly dorsal on

foretibia and in more or less double row; with pale scales at apex. Tarsi with pale rings on

base and apex of tarsomeres 1-3 and base of tarsomere 4, basal rings broader.

Abdomen: Terga II-VI mainly dark brown-scaled with variable basal pale bands and lateral

pale spots; basal bands of terga II and III produced medially; most of middorsal surface

pale on terga VI and VII; basal bands continuous with lateral spots except perhaps on

more proximal terga; lateral spots progressively larger from anterior to posterior terga.

Sterna largely pale-scaled; often with indistinct V-shaped patch of dark scales, base of V

at middle of anterior margin.

MALE. Same as female except as follows. Head: Proboscis paler; with many pale

scales mixed with dark ones; mostly pale beneath; often pale at sides before labella.

Maxillary palpus with 6 pale areas as follows: (1) lateral pale area at base of palpus without

scales, (2) narrow pale ring across junction of palpomeres 2 and 3 without scales, (3)

broad dorsolateral pale area at middle of palpomere 3 with pale scales, (4) pale area at

apex of palpomere 3 and base of palpomere 4, latter with pale scales dorsally and

ventrally, (5) pale area at apex of palpomere 4 and base of palpomere 5, latter with small

patch of pale scales dorsally and ventrally; (6) pale area on approximately distal 0.5 of

palpomere 5 with line of pale scales on dorsal and ventral surfaces; palpomeres 4 and 5

often with scattered pale scales on ventral surface; length 2.6-3.3 mm, mean 3.0 mm.

Forked scales of vertex pale medially, becoming dark laterally and posteriorly; falcate

scales paler, whitish to yellow; spatulate scales becoming yellowish ventrally. Thorax:

Upper proepisternal scales numerous, about 35. Scutal scales smaller, more numerous

and paler, those of fossae largely golden yellow; band behind middle pale yellow.

Scutellar scales narrower and paler. Pleural scale-patches generally paler. Wing:

Length 3.1-4.2 mm, mean 3.7 mm; length of cell R2/length of vein R2+3

variable, 2.1-4.1, mean 2.7; pale scaling less distinct, wing paler overall; pale scaling

reduced along posterior margin of R1. Abdomen: Terga II and Ill with medial

extension of basal band often reaching posterior margin; terga VI and VII sometimes

entirely pale-scaled. Genitalia (Fig. 33): Form as figured; lateral plate of phallosome

somewhat as in sitiens. Ninth tergal lobe scarcely produced, with row of 5-9 (mode 8)

irregularly spaced setae. Gonocoxite rather elongate, lateral setae extending from near

base to apex, in 2 or 3 rows increasing to about 5 rows at level of subapical lobe; subapi-

Harbach: Subgenus Culex in Southwestern Asia 95

cal lobe undivided, not very prominent; setae a-c slender, about same length, a blunt

at tip, b and c hooked at tip; only 2 simple setae in group a-f, one acute, the other (f ?)

slightly flattened and hooked at tip; g and / essentially as in sitiens; a noteworthy

auxiliary seta (aux) present basally on proximal side of lobe. Gonostylus not unusual.

Phallosome longer than broad; aedeagal sclerite slightly longer and narrower than lateral

plate, ventral margin deeply excavated in middle; inner division of lateral plate closely

fused to ventromesal margin of outer division, shape much as in sitiens but with 3-5

more massive teeth arising between dorsocaudal angle and mesal surface of outer

division, teeth resemble a hand with stubby fingers directed dorsolaterally, dorsomesal

margin with tiny bumps and spicules; outer division with large concavo-convex flap

arising from dorsocaudal margin, dorsal process and basal articulatory process poorly

developed. Proctiger large; paraproct long and slender, without basal lateral arm, crown

a large mass of short spinelike spicules. Cercal sclerite elongate, somewhat triangular;

2-6 (mode 4) cercal setae. Tergum X small.

Pupa (Fig. 33). Form and positions of setae as figured, range and modal number

of branches in Table 36; usually easily recognized by the presence of a darkened area

on the paddle (not illustrated). Cephalothorax: Setae 1,3,8-CT usually with 3 or 4

branches (see table); 2-CT usually with 4 or 5 branches (3-5); 4,6-CT usually triple (2-4);

5-CT usually with 4 or 5 branches (4-6), not much longer than 4-CT; 7,11-CT double or

triple, more often double; 10-CT multiple (7-16); 12-CT with 3-6 branches occurring in

nearly equal frequencies, longer than 10,11-CT. Trumpet: Lightly to moderately

tanned, tracheoid area and pinna darker; index 5.7-7.5, mean 6.8; pinna short, 0.1-0.2

length of trumpet. Abdomen: Length 3.1-3.6 mm, mean 3.4 mm; caudolateral angle of

terga VII and VIIl acute. Seta 7-I double or triple, 7-Il usually double, sometimes single or

triple; 1-Ill-V strongly developed, multiple, about length of following tergum; 6-IlI-VI

moderately developed, usually with 4 or 5 branches (3-6), 6-VI distinctly longer than the

others; 5-IV usually triple (3-5), about length of 1-IV, shorter than 5-V,VI which are

double and considerably longer than following terga. Paddle: Lightly tanned, with

darkened area on distal 0.5 or less and mainly on inner part; length 0.91-1.16 mm, mean

1.00 mm, width 0.68-0.82 mm, mean 0.74 mm, index 1.2-1.5, mean 1.4.

Larva (Fig. 34). Character and placement of setae as figured, range and modal

number of branches in Table 36; recognized at once by the shape of the siphon, and

black pigmentation of posterior part of thorax and abdominal segments III and V which

give the larva a 3-banded appearance in life. Head: Length 0.82-0.95 mm, mean 0.89

mm; mainly lightly tanned, darker posteriorly, ventral part of lateralia adjacent to

hypostomal suture sometimes with dark triangular spot. Median labral plate noticeably

produced forward at point of insertion of seta 1-C. Dorsomentum usually with 6 teeth

(5-7) on either side of middle tooth. Seta 1-C short, stout, pointed, dark; 2-C

occasionally present; 4,5,6-C inserted more posterior than usual, 5,6-C only extending

to near base of 1-C; 4-C short, usually double (1-3); 5-C most often with 4 branches

(3-5); 6-C distinctly shorter than 5-C, often with 4 branches (4-6); 7-C strongly

developed, usually with 10, 11 or 12 branches (10-15); 8,9,10-C generally with more

branches than usual, 10-C usually with 4 or 5 branches (3-5); 11-C double, seldom triple;

14-C rather spiniform, normally single (1,2). Antenna: Length 0.76-0.86 mm, mean

0.80 mm; tanning and spiculation as usual; scape developed. Seta 1-A large, with

about 34 branches (29-37); 2,3-A inserted near apex. Jhorax: Integument hyaline,

smooth; with concentration of black pigment granules under dorsal surface of meso- and

metathorax. Setae 4,8-P double, about same length; 7-P with 3 or 4 branches, perhaps

slightly longer than 4,8-P. Setae 1,4-M similar, both with more branches than usual, 1-M

often with 5 branches (5-8), 4-M usually with 4or5 branches (4-6). Seta 1-T multiple

96 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

about 0.5 length of 1-T; 2-T also multiple, most often with 4 branches (3-5); 12-T double

or triple; 13-T with more branches than usual (12-23), not much longer than 12-T.

Abdomen: \Integument hyaline, smooth; dark pigment granules present, particularly

noticeable in segments III and V (as in antennatus and sinaiticus). Seta 3-| multiple (2-7),

infrequently double or triple, 3-Il-IV usually with 3 or 4 branches (see table), 3-V,VI

double or triple, 3-VII highly branched (7-10); 6-I-VI normally triple, some occasionally

with 4 branches, 6-V usually distinctly shorter than the others; 7-1 usually double (1,2)

but as long as 6-l, 7-II-VIl generally with more branches than usual (see table); 1-III-VI|

usually with 7 or 8 branches (see table). Segment Vill: Comb consisting of 6-10 (mode

7) large spinelike scales, fringed on either side at base. Seta 2-VIII double or triple; 5-VIII

generally with more branches than usual (5-7). Siphon: Index 3.59-4.66, mean 4.22;

moderately tanned; slightly but distinctly bent anteriorly, posterior side longer than

anterior side; spiracular lobes particularly large. Pecten composed of 8-14 (mode 11)

short spines with 4-9 (mode 5) basal denticles, most basal denticles tend to be fused.

Seta 1-S in 5 pairs, middle pair borne laterally, others on posterior midline, length of latter

more than twice diameter of siphon at point of attachment. Segment X: Saddle

complete, rather long, length 0.43-0.52 mm, mean 0.48 mm, siphon/saddle index

2.54-3.07, mean 2.78. Seta 1-X usually with 4 branches (2-5); 2-X often with 5 branches

(3-7); 4-X in 6 or 7 pairs, branches numerous. Length of anal papillae variable, ventral

pair usually, but not always, longer than dorsal pair.

Systematics. Culex poicilipes is widely distributed south of the Sahara and

along the Nile River into Egypt. The species occurs in the Sinai Peninsula and the

Negev of Israel where it probably spread from the Nile Delta in relatively recent times.

The species is strikingly distinct in all stages and cannot be confused with any other

species of Culex occurring in Egypt or Israel.

This is one of the most variable species of Culex in Africa. Remarkable variation

occurs in the ornamentation of the adults and the chaetotaxy of the larva. Much of this

variation is undoubtedly individual, but there are indications that some populations differ

constantly from others. A noticeably darker adult form with less extensive pale markings

is common in some areas. Whether this is due to genetic of environmental factors

cannot be determined without sufficient individually reared material collected

throughout the recorded range of the taxon. The adult and immature stages of

poicilipes in Egypt and Israel are not so variable and it is unlikely that more than one

species is involved in these countries.

The placement of poicilipes in the sitiens subgroup is provisional. It is included

chiefly because of overt similarites in the structure of the male phallosome. There are

important differences in all life stages. The adults differ in having proepisternal scales

and ornate markings. The larval and pupal stages differ in the development of many

setae. Despite the differences in chaetotaxy, the larval head capsule, comb scales, and

siphon are constructed almost like those in the aberrant Culex whitmorei (Giles, 1904)

which is currently recognized as a member of this subgroup (Sirivanakarn, 1976). Itis

very likely that both poicilipes and whitmore/ will have to be placed in separate

subgroups after members of the sitiens group are studied more thoroughly.

Bionomics. The immatures of this species generally inhabit clean bodies of

standing water with thick vegetation. The breeding places are normally exposed to full

sunlight. Rice fields, shallow pools, and ditches are common breeding sites in Egypt.

This mosquito is often associated with antennatus, perexiguus, and Anopheles (Cellia)

pharoensis Theobald, but rarely with other species. Adults enter houses in Egypt

(Kirkpatrick, 1925) and readily attack man near swamps in the Sudan (Lewis, 1956), but

are not known to vector any pathogens of human disease.

Harbach: Subgenus Culex in Southwestem Asia 97

Distribution. Culex poicilipes is widely distributed south of the Sahara and

enters Egypt along the Nile. Populations occur in the Sinai Peninsula and the Negev

Desert of southern Israel.

Material examined. 294 specimens. A total of 122 specimens (74 females,

14 males, 5 male genitalia, 10 pupal exuviae, 9 larval exuviae, and 10 fourth-instar

larvae) were examined from southwestern Asia — EGYPT: Beni Suef (Biba), E/

Bahayrah (Alexandria), E/ Buhayra (Mansura, Wadi Natrun), E/ sma’ iliya (Abu Khalifa,

Isma'iliya), E/ Fayyum (Birket Qarum), E/ Giza (Harania, Kafr el Ghataty, Kerdasa), E/

Qulyubiya (Ezbet Ayoub), E/ Shargiya (Inshas el Raml); ISRAEL: (Ha'arana, Iddan, Neot

Hakikar, Yeroham, Zikim). An additional 172 specimens (95 females, 39 males, 7 male

genitalia, 3 pupal exuviae, 5 larval exuviae, and 23 fourth-instar larvae) were examined

from ANGOLA (the holotype female of taeniorhynchoides ), BENIN, ETHIOPIA,

GAMBIA, GHANA, KENYA, MADAGASCAR (including the types series of

madagascariensis and auritaenia), MALAWI, MALI, MAURITANIA, MOZAMBIQUE (the

holotype female of punctipes), NIGERIA (including the holotype female), SENEGAL,

SOUTH AFRICA, SUDAN (including the holotype female of maculipes), TANZANIA,

UGANDA (including the holotype female of par), ZIMBABWE, and unknown localities.

Culex (Culex) mimeticus Noé

mimeticus Noé, 1899: 240. Type(s) (female): Grassano in Basilicata,

[Italy] (non-extant).

pseudomimeticus Sergent, 1909: 445. Type(s) (adult): Oued Chiffa,

Algeria (non-extant). Synonymy with mimeticus by Edwards, 1932a:

205.

Culex mimeticus Noé of Giles, 1901a: 605 (India, A, distr.); Theobald, 1901a: 329 (India,

A key, F"); Christophers, 1906: 9 (? India, L); Joyeux, 1918: 533 (Macedonia,

coll.); Waterston, 1918: 9 (Macedonia, A, L habitat); Barraud, 1921: 395 (Israel,

Jordan, Syria); Edwards, 1922: 284 (Oriental Region, A, key); Barraud, 1923: 941

(India, L*); Buxton, 1923: 316 (Israel, Jordan, coll. rec., biol. note); Theodor,

1923: 341 (Jordan, Israel, P*, key); Barraud, 1924a: 991 (India, A key, M’*, F);

Parr, 1943: 246-250 (Syria, Lebanon, A, P, L keys, bionomics); Senevet, 1947b:

212 (North Africa, L key); Theodor, 1952: 113 (Middle East, zoogeogr.); Dow,

1953: 688 (Iran, coll. rec.); Senevet et al., 1957a: 86 (North Africa, L*); Margalit

and Tahori, 1973: 90-93 (Sinai, distr., L assoc.); Margalit et al., 1973: 32 (Israel,

coll. rec., L assoc.); Kitron and Pener, 1986 (Israel, L bionomics).

Culex (Culex) mimeticus Noé of Edwards, 1921: 337 (E. Mediterranean Region, Israel,

Jordan, A, L keys, tax., distr.); Séguy, 1924: 34, 186 (E. Mediterranean Region,

Israel, Jordan, A, L keys, L*, tax., distr., syn.); Edwards, 1926: 130 (E. Mediter-

ranean Region, Israel, Jordan, A, L keys, A, L, L assoc., distr.); Stackelberg,

1927: 155 (Middle East, M, F keys, A, M gen., distr.); Martini, 1931: 371 (Israel,

Jordan, Turkey, A, L keys, M’, F, L*, distr.); Barraud, 1934: 409 (India, A, L keys,

M*, F, L*, distr.); Edwards, 1934: 451 (India, F, distr.); Senevet, 1947a: 121

(Egypt, L*, distr., L assoc.); Senevet, 1949: 56 (North Africa, M gen.*, key);

Monchadskii, 1951: 266 (Middle East, L*, L assoc., distr.); Senevet and Andarelli,

1954: 57 (North Africa, M, F, A key); Senevet et al., 1957b: 92 (North Africa, L);

Parrish, 1959: 266 (Turkey, distr.); Senevet and Andarelli, 1959: 151 (North

Africa, A, P, L keys, syn., M*, F*, P*, L*, distr., biol., L assoc.); Abdel-Malek, 1960:

98 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

112-121 (Syria, coll. sites, L bionomics); Derwesh, 1965: 44 (Iraq); DuBose and

Curtin, 1965: 351, 354 (Mediterranean area, A, L keys); Abul-hab, 1966: 283

(Iraq, distr., L bionomics); Abul-hab, 1968: 246 (Iraq, L key, distr.); Lotfi, 1970:

401 (Iran, coll. rec.); Margalit and Tahori, 1970b: 152 (Israel, coll. sites, distr.);

Aslamkhan, 1971: 154 (Pakistan); Lotfi, 1973: 206 (Iran, coll. sites); Gutsevich et

al., 1974: 375 (Middle East, F, M gen., L keys, M’*, F*, L*, distr., bionomics);

Margalit and Tahori, 1974: 87 (Israel, coll. rec.); Lotfi, 1976 (Iran, L key, L*, ecol.

note); Sirivanakarn, 1976: 145 (Oriental Region, A, P, L keys, M*, F*, P*, L*, syn.,

tax., distr., bionomics); Ibrahim et al., 1983: 91 (Iraq, L*, key); Harbach, 1985a: 86,

96, 107 (SW Asia [excluding records for Afghanistan and Saudi Arabia], Egypt,

distr., A, L, key).

Adult. The presence of pale spots on the wings of this medium-sized species

readily distinguishes it from all other species of Culex known to occur in southwestern

Asia and Egypt. This character evokes a superficial resemblance to Anopheles species

of the subgenus Cellia.

FEMALE. Head: Antenna dark; length 2.0-2.3 mm, mean 2.1 mm; pedicel with

small patch of tiny pale scales on dorsal surface. Proboscis black-scaled with well

defined median whitish ring extending from 0.4 to 0.6 from base; length 1.9-2.1 mm,

mean 2.0 mm. Maxillary palpus dark with pale scales at apex. Forked scales of vertex

pale yellow medially, dark brown to black laterally; falcate scales white, yellowish

dorsolaterally; lateral spatulate scales white. Thorax: Integument brown, pleura with

darker spots on proepimeron, postspiracular area, prealar area, anterior and posterior

margins of mesokatepisternum, and anterior and upper areas of mesepimeron; meron

also dark. Scutal scales coarse, predominantly pale yellow to golden yellow, with white

to pale yellow scales on anterior promontory, lateral margins and prescutellar area, scales

on fossa and middorsal area sometimes golden brown to brown. Scutellum with white

falcate scales, large patch on median lobe, few on lateral lobes. Ante- and postpronota

with golden-yellow falcate scales; postpronotum with about 6 setae on posterior border.

Pleura with patches of white spatulate scales as follows: proepisternum with patch

immediately above forecoxa, mesopleuron with upper and lower patches on

katepisternum and anterior and upper patches on anepimeron, scales on katepisternum

more elongate; numbers of pleural setae: 4-8 upper proepisternal, 10-12 prealar, 5-7

upper mesokatepisternal, 9-13 upper mesepimeral; without lower mesepimeral seta.

Wing (Fig. 2E): Length 3.8-4.0 mm, mean 3.9 mm; black-scaled with white or yellowish-

white spots, pattern follows: costa dark with sector, subcostal and preapical pale spots;

R1 dark with subcostal and preapical pale spots; pale scales at apex of R2+3 continuous

with pale scales at base of R2 and R3; R2 with preapical pale spot; R4+5 with dark base

followed by large median pale spot and small apical dark spot; pale spot at furcation of

M1+2; M3+4 with large median pale spot; CuA with very small pale spot at apex

confluent with pale fringe spot; 1A with large pale spot near middle. Ha/ter: Pedicel and

scabellum yellow; capitellum brown with pale scales. Legs (Fig. 2H): Forecoxa

black-scaled with patch of white scales at base; mid- and hindcoxae with longitudinal

stripe of white scales on anterior side of lateral midline. Anterior surface of forefemur

entirely black-scaled with ill-defined narrow line of white scales on proximal 0.5, with

distinct posterodorsal stripe of white scales not reaching apex; anterior surface of

midfemur black-scaled with white scales at base, posterior surface white-scaled with

small subapical area of black scales; hindfemur white-scaled except for dorsal stripe of

black scales beginning near base and broadening distally to cover approximately distal

0.25 of anterior and posterior surfaces. Tibiae and tarsi mainly black-scaled; all tibiae with

Harbach: Subgenus Culex in Southwestem Asia 99

posterior white stripe, mid- and hindtibiae with narrow anterior white stripe, anterior stripe

of midtibia incomplete distally, anterior stripe of hindtibia incomplete at ends.

Tarsomeres 1-4 of all legs with basal and apical white rings, basal rings broader.

Abdomen: Terga mainly black-scaled; tergum | with submedian patches of black scales

on posterior border that may be separated by a few median pale scales: terga II-VII with

basal bands of yellowish-white scales and large basolateral spots of white scales, spots

may cover entire lateral surface, bands straight or slightly convex, 0.15-0.35 tergum

length and barely if at all reaching spots. Sternum | with some dark scales near

posterolateral corners; sterna II-VII with posterolateral dark spots that may join to form

broad apical band.

MALE. Resembling female, differences follow. Head: Maxillary palpus with broad

subapical ring of pale scales on palpomere 3, narrow ring of pale scales at bases of

palpomeres 4 and 5, and pale scales on dorsal surface of palpomere 5. Vertex with

fewer forked scales; falcate scales fewer, longer and all yellowish white; larger patch of

pale lateral spatulate scales. Thorax: Scutal scales longer, predominantly whitish to

golden yellow with spots of golden-brown scales on fossae and middorsal area, lightest

scales on lateral margins and posterior 0.33 of scutum. Scutellar scales fewer and

longer. Upper proepisternal setae more numerous, usually 8-10. Wing: Length 2.9-3.3

mm, mean 3.1 mm; pattern of spots differs as follows: sector pale spot on anterior

margin of subcosta; pale spots at apex of costa, R1 and R2; pale spots absent from

furcation of R1+2, furcation of M1+2 and apex of CuA; M3+4 with small basal dark spot

followed by small pale spot and large dark spot; 1A pale spot extending further proximad

than in female. Abdomen: Without basolateral spots; basal pale bands concave,

0.35-0.45 tergum length on dorsal midline. Genitalia (Fig. 35): Form as figured;

Structure much as in sitiens, differing chiefly as follows. Ninth tergal lobe poorly

differentiated, with 4-7 short setae. Subapical lobe of gonocoxite with 3 or 4 setae in

group d-e, 3 usually present, these flattened with recurved apex and 1 or 2 recurved

denticles before tip; seta f flattened and bladelike, rounded apically; h longer. Inner

division of phallosome more closely fused to outer division, not as broad, dorsocaudal

angle elongate and slender, with 1-3 teeth (usually 2) arising very near tooth on

posteromesal margin of outer division; outer division with much smaller process

projecting laterally from ventrocaudal margin, dorsal process not as prominent. Proctiger

essentially as in sitiens; basal lateral arm of paraproct generally shorter and more slender.

Cercal sclerite with 2 or 3 setae on posterolateral corner. Tergum X small.

Pupa (Fig. 35). Placement and form of setae as figured, range and modal

number of branches in Table 17; without striking distinctive features, closely resembles

sitiens but setae 1-ll, 6-III-VI, and 9-VIII usually with fewer branches. Cephalothorax:

Lightly tanned. All setae except 10-CT usually double or triple, more often double:

10-CT usually with 4 or 5 branches (4-6). Trumpet: Moderately tanned: slender,

pinna slightly widened; index 4.7-7.0, mean 5.9; pinna 0.25-0.40 trumpet length.

Abdomen: Lightly tanned; length 2.6-3.9 mm, mean 3.2 mm. Seta 6-I,IIl short, not more

than twice length of 7-1,II; 7-1,ll usually double (1-3); 1-Il weak, with only 1-4 branches,

1-III-V generally with progressively fewer branches on each successive posterior tergum,

1-IIl frequently with 4 or 5 branches (3-6), 1-IV most often triple (2-5), 1-V usually double

(1-3), 1-VI commonly single (1,2), 1-VIl like 1-V; 6-III-VI usually single (see table), 6-III,1V

distinctly shorter than 6-V,VI; 5-IV-VI strongly developed, about 1.5 length of tergum

following, 5-IV most often double (2-4), 5-V,VI double; 2-VII usually mesal to 1-VII; 9-VIII

usually with 6 or 7 branches (4-9). Paddle: Lightly tanned; length 0.76-1.03 mm, mean

0.94 mm; width 0.59-0.78 mm, mean 0.67 mm; index 1.3-1.6, mean 1.5.

Larva (Fig. 36). Character and positions of setae as figured, range and modal

100 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

number of branches in Table 37; resembling theileri, simpsoni, and sinaiticus in

the form of the comb scales but differing conspicuously from these species in the

placement of seta 1-S and in having setae 2,3-A inserted approximately midway

between seta 1-A and apex of antenna, easily distinguished from other members

of the sitiens group by the character of the comb scales and the reduced

branching of seta 4-X. Head: Length 0.60-0.72 mm, mean 0.66 mm; width

0.95-1.10 mm, mean 1.05 mm; lightly tanned, darker posteriorly. Hypostomal

suture ending before collar; collar well developed, heavily tanned. Dorsomentum

almost shape of an equilateral triangle; usually with 7 or 8 teeth (6-8) on each side

of middle tooth. Seta 1-C moderately long, stout, pointed, tanned; 2-C absent;

4-C double, triple, or with 4 branches occurring in nearly equal frequencies; 5-C

with 3 or 4 branches, more often with 4; 6-C double or triple, more often double;

14-C rather large and spinelike. Antenna: Length 0.46-0.62 mm, mean 0.55 mm;

lightly tanned, distal part dark, with dark ring at base. Seta 1-A with about 25

branches (21-27); 2,3-A inserted almost halfway between 1-A and apex. Thorax:

Integument hyaline, smooth. Setae 1-6-P equally long and single; 7-P usually

triple, seldom double; 8-P double or triple in nearly equal frequencies, distinctly

shorter than 7-P. Seta 1,2-T usually single, occasionally double, 2-T 2 or 3 times

as long as 1-T; 13-T weak, shorter than 12-T, usually double, occasionally single

or triple. Abdomen: \Integument hyaline, smooth. Seta 3-I,IV single or double,

3-II,11,V,Vl almost always single, 3-VII often double but frequently triple or with 4

branches; 6-I-V usually triple, 6-I,Il often with 4 branches, 6-VI usually double,

occasionally triple; 7-l as in sitiens; 1-II| short and single, 1-IV-VI usually double,

6-V,VI stronger than 6-IV, 1-VIlI triple or with 4 branches, more often with 4;

13-III-V,VIl weakly developed, much shorter than seta 1 of same segment, almost

always single or double. Segment VI//: Comb with 27-50 (mean 33) sharply

pointed scales with short lateral fringes on proximal part. Siphon: Index

4.55-7.41, mean 6.07; lightly tanned, slightly darker at apex, basal rim moderately

to heavily tanned. Pecten composed of 13-20 (mode 15) long slender spines

with rather short line of fused denticles at base. Seta 1-S in 5-7 pairs (10-14

setae), 1 subapical pair borne laterally, others along posterior midline, proximal 4

or 5 pairs stronger, most often with 4 or 5 branches (see table), lateral pair and

most distal pair shorter and usually double or triple; 2-S rather long and curved.

Segment X: Saddle complete; lightly tanned, slightly darker posteriorly; with rows

of minute spicules; length 0.37-0.42 mm, mean 0.41 mm, siphon/saddle index

3.90-4.65, mean 4.22. Seta 1-X normally double (1-3); 2-X normally double,

occasionally triple; 4-X usually in 6 pairs, 1 seta sometimes unpaired, most setae

commonly with 4 branches (3-6). Anal papillae long and slender, about twice

length of saddle.

Systematics. This species is the nominate member of an interesting

subgroup which is largely confined to the Oriental Region. Culex mimeticus is the

only species of the group which is known to occur in the Mediterranean

Subregion of the Palaearctic. It is apparently mainly a chaparral and temperate

grassland species which is excluded from the Arabian Peninsula and the Sahara.

The adults of mimeticus are unique among the Culex of southwestern Asia

in having spotted wings much as in species of the subgenus Cellia of Anopheles.

The larva is variable but distinct from all other species in the region. It bears a very

superficial resemblance to the larva of Culex deserticola Kirkpatrick, 1924(1925),

but is distinguished at once from this species by the placement of setae 2,3-A

and the character of the comb scales.

Harbach: Subgenus Culex in Southwestem Asia 101

It is possible that two or more species are currently confused under the concept

of mimeticus in the Mediterranean Subregion. Topotypic material from Italy, for example,

differs noticeably in the size and shape of the pupal trumpet. There also seems to be a

number of geographically isolated populations in the region. Individual rearings are

lacking from almost all localities and the available material is much too scanty to permit a

detailed comparative analysis. The above diagnosis is based mainly on reared material

from the Negev of Israel.

Bionomics. The natural breeding sites of mimeticus appear to be springs and

residual pools in drying streams. The water in these habitats is fresh and clear and

usually contains growths of Spirogyra. Other collections have been made at the margins

of streams, pools in swampy ground, irrigation ditches, and animal hoofprints. Larvae are

frequently found with Anopheles (Cellia) hispaniola (Theobald), An. (Cel.) sergentii

(Theobald), and An. (Cel.) superpictus Grassi. Adults apparently occasionally enter

human habitations (Hsiao and Bohart, 1946; Ribeiro et al., 1977), but females have

never been seen landing on or biting man or animals. Nothing is known about the

biology of the adults.

Distribution. This species occurs in the southern Palaearctic and Oriental

regions. Its range extends from lands around the Mediterranean to eastern and

southeastern Asia. Culex mimeticus seems to be less common than it once was,

probably as a result of pesticide usage, organic pollution, and the destruction of

preferred breeding places. The species has a very spotted distribution.

Material examined. 215 specimens. A total of 128 specimens (26 females,

27 males, 24 male genitalia, 15 pupal exuviae, 17 larval exuviae, and 19 fourth-instar

larvae) were examined from southwestern Asia —- IRAN: (Amirabad, Chalus,

Izeh-Ahwaz, Kanroud, Kazeroon, Khorramabad, Natchi Bahran, Tehran, Zanjan); IRAQ:

(Mirgasur); ISRAEL: (Dalia, Ein Amud, En Avedat, Jericho, Jirou, Latron, N. Bet

Shearim, N. Moran, Wadi Kabalah, Wadi Kurn); JORDAN: (Jerash, Tafile, Tafilan, Wadi

Hasa, Wadi Musa, unknown localities); TURKEY: (Yenice). An additional 87 specimens

(24 females, 28 males, 12 male genitalia, 10 pupal exuviae, 11 larval exuviae, and 2

fourth-instar larvae) were examined from ALGERIA, GREECE (CRETE), ITALY (topotypic

material), MOROCCO, and TUNISIA.

Culex (Culex) bitaeniorhynchus Giles

bitaeniorhynchus Giles, 1901a: 607 . Type(s) (adult): Travancore,

India (non-extant).

ager Giles, 1901b: 196 (Taeniorhynchus ). Type(s) (female): Madras,

Presidency, India (non-extant, lectotype designation of Bram,1967a:

263 invalidated by Harbach, 1983: 105). Synonymy with bitaenio-

rhynchus by Edwards, 1913c: 231.

ethiopicus Edwards, 1912b: 30 (Taeniorhynchus ager var.). +tHolotype

female: Bole, Gold Coast, West Africa (BM). NEW SYNONYMY.

abdominalis Taylor, 1913: 53 (Culicelsa ). Type(s) (female): Ayr and

Townsville, Queensland, Australia (non-extant). Synonymy with

bitaeniorhynchus by Edwards, 1913c: 231.

maculipesarabiensis Patton, 1905: 635 (Taeniorhynchus tenax var.).

Type(s) (adult; larva): D'thala, Hardeba and Nobat, P.D.R. Yemen

(non-extant). Synonymy with ethiopicus by Edwards, 1941: 291.

karatsuensis Mochizuki, 1913: 28. Type(s) (male; female; egg): Karat-

102 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

su, Kyushu, Japan (location unknown). Synonymy with bitaenio-

rhynchus by Edwards, 1932a: 202.

Culex ager var. ethiopicus Edwards of Edwards, 1912c: 381 (Africa, L*, key); Ingram and

Macfie, 1919: 65 (W. Africa, P*, L*, L habitat).

Culex bitaeniorhynchus Giles of Barraud, 1923: 936 (India, L*); Barraud, 1924a: 984 (in

part; India, A key, syn., M’*, F, distr.).

Culex bitaeniorhynchus var. ethiopicus Edwards of Edwards, 1920: 135 (Africa, M gen.).

Culex (Culex) bitaeniorhynchus Giles of Edwards, 1921: 337 (in part; Palaearctic

Region, A, L keys, tax., bionomics, distr.); Edwards, 1926: 129 (in part;

Palaearctic Region, A, L keys, syn., A, distr.); Stackelberg, 1927: 153 (in part; S.

Asia, M, F keys, A, M gen.", distr.); Martini, 1931: 362 (in part; S. Asia, A, L keys,

M*, F, L*, distr.); Barraud, 1934: 391 (in part; India, A, L keys, M*, F, L*, syn., L

habitat, distr.); Monchadskii, 1951: 258 (in part; S. Asia, L*, L assoc., distr.); Lotfi,

1970: 401 (Iran, coll. rec.); Aslamkhan, 1971: 154 (in part; Pakistan); Lotfi, 1973:

206 (Iran, coll. rec.); Gutsevich et al., 1974: 367 (in part; S. Asia, M*, F, L’, distr.,

bionomics, med. imp.); Lotfi, 1976: 72, 74, 79 (Iran, L key, bionomic note, L*):

Sirivanakarn, 1976: 65 (Oriental Region, A, P, L keys, M*, F*, P*, L*, tax., distr.,

bionomics, med. imp.); Harbach, 1985a: 86, 98, 107 (Iran, Pakistan, P.D.R.

Yemen, Yemen Arab Republic, A, L keys).

Culex ethiopicus Edwards of Lewis, 1945: 14 (Sudan, L, distr.).

Culex (Culex) ethiopicus Edwards of Edwards, 1941: 291, 415, 482 (Afrotropical

Region, A key, M"*, F, P*, syn., distr.); Lewis, 1943a: 281 (Eritrea, coll. rec., L

bionomic note); Lewis, 1943b: 69 (Sudan, bionomic notes); Abbott, 1948: 44

(Sudan, distr.); Lewis, 1948: 144 (Sudan, L habitat); Hopkins, 1952: 282

(Afrotropical Region, L key, L*, bionomics); Knight, 1953a: 229 (Yemen Arab

Republic, coll. rec., L habitat); Lewis, 1956: 706 (Sudan, coll. rec.); Mattingly and

Knight,1956: 106-132 (P.D.R. Yemen, Yemen Arab Republic, L*, A, L keys,

distr., bionomics); Mekuria, 1968: 78 (Ethiopia, distr.).

Adult. A rather large species easily recognized by the speckled wings and

rather broad apical yellow bands of the abdominal terga.

FEMALE. Head: Antenna dark, pedicel and flagellomere 1 paler; pedicel

sometimes with some pale scales on mesal surface; length 1.8-2.4 mm, mean 2.0 mm.

Proboscis with distinct median pale ring about 0.25 proboscis length, pair of dorsolateral

pale spots at apex before labella and scattered pale scales proximal to ring; length

1.6-2.1 mm, mean 1.9 mm. Maxillary palpus dark-scaled; palpomere 4 with pale scales

apically, sometimes with scattered pale scales proximally; length 0.4-0.5 mm, slightly

more than 0.2 length of proboscis. Dorsal falcate scales of vertex coarse, very pale

yellow; forked scales largely pale yellow to yellowish brown, some darker

posterolaterally; lateral and ventral spatulate scales yellowish white. Thorax: Integument

brownish yellow, scarcely darker dorsally. Scales on anterior 0.7 of scutum mainly pale

yellow to golden yellow, somewhat mottled by presence of some golden-brown to

brownish scales, particularly on anteromedian area and sublaterally at middle; posterior

0.3 clothed in golden-brown and dark brown scales with some coarser pale scales on

antealar area and posteriorly on prescutellar area; scutal setae golden brown except for

distinctly paler (yellowish) antealar setae, supraalar setae dark, dense (numerous) and

long. Scutellum with pale scales on lateral lobes, median lobe mainly pale-scaled with

some brownish-yellow to dark brown scales anteriorly; 5 or 6 long setae on lateral lobes,

7-11 on median lobe. Ante- and postpronota with coarse yellowish to golden falcate

Harbach: Subgenus Culex in Southwestem Asia 103

scales, scales golden brown to brown centrally. Pleural setae mainly pale, usually

entirely pale: 8-11 upper proespisternal, 6-13 prealar, 5-7 upper mesokatepisternal,

6-10 lower mesokatepisternal, 4-14 upper mesepimeral, and no lower mesepimeral.

Pleural scales yellowish white: proepisternum with upper patch comprised of falcate

scales laterally and spatulate scales anteriorly, mesokatepisternum with upper and lower

patches of elongate spatulate scales, mesepimeron with anterior patch of elongate

spatulate scales at level of upper katepisternal patch and patch of similar scales before

upper mesepimeral setae. Wing (Fig. 2F): Length 3.6-4.6 mm, mean 4.1 mm; length of

cell R2 2.4-3.5 length of vein R2+3, mean 3.0; length of cell M1 about 0.9 length of cell

R2; dorsal surface with broad apically rounded yellowish-white and brown spatulate

scales evenly intermixed on all veins; ventral surface with mixture of pale and dark

spatulate scales on costa, subcosta, Rs and M, other veins with pallid brownish linear to

very narrow spatulate scales, scales on R1, R4+5, M2 and M3+4 linear and near linear,

scales on other veins near linear and very narrow spatulate. Halter: Integument

yellowish to brownish yellow; capitellum pale-scaled, distal part of scabellum and base of

capitellum usually with some dark scales. Legs: Forecoxa with dark and pale spatulate

scales anteriorly, latter dominant at base and apex; mid- and hindcoxae with anterior

patches of pale scales, some dark scales distally. Anterior, posterior, and ventral

surfaces of trochanters with mixture of pale and dark scales. Femora with more or less

even mixture of dark and pale scales on anterior surface, posterior surface

predominantly pale with interspersed dark scales mainly on distal 0.5 or less. Dorsal

Surface of foretibia and anterior surface of mid- and hintibiae with interspersed dark and

pale scales, ventral surface of foretibia and posterior surface of mid- and hindtibiae

mainly pale-scaled with intermingled pallid brown scales, especially proximally. Tarsi

dark-scaled; tarsomeres 1-4 with narrow apical and broader basal rings of pale scales;

tarsomere 1 of all legs and tarsomere 2 of hindleg with some scattered pale scales.

Abdomen: Tergum | with rather large median posterior patch of yellow scales,

specimens from Asita, Wallo Province, Ethiopia with some rather diffuse dark and pale

scales on lateral borders; terga II-VIII with broad apical yellow bands (about 0.33 tergum

length) not reaching lateral margins, scattered yellowish scales among proximal dark

scales, and large basolateral white spotsdiagonally opposed to apical bands, tergum VIII

also with basal pale band contiguous with basolateral spots that may join apical band

medially. Sterna II-VII with narrow yellow apical bands joined medially to broad whitish

basal bands, apicolateral corners with dark scales, sometimes with some scattered pale

scales, particularly on proximal sterna where dark patches are larger and less definite:

sternum VIII with lateral pale patches, middle devoid of scales.

MALE. Like female, differing mainly in sexual characters, major differences follow.

Head: Median pale ring of proboscis narrow, 0.1-0.15 proboscis length; with ventral

setal cluster at base of ring before false joint; false joint about 0.55 from base. Maxillary

palpus mainly dark-scaled; with narrow ring of pale integument across junction of

palpomeres 2 and 3; palpomere 3 with rather long median pale patch with dispersed dark

scales on dorsal and lateral surfaces, sometimes with scattered pale scales distal to pale

patch, dense patch of long setae on ventrolateral surface immediately distal to pale

patch, and ventromesal surface with line of short antrorsely curved setae along entire

length; palpomeres 4 and 5 densely setose with basal pale rings, palpomere 4 with

scattered pale scales distal to ring, distal 0.5 of palpomere 5 with pale setae and scales;

length 2.6-2.9 mm, mean 2.8 mm, extending beyond tip of proboscis by length of

palpomere 5 and part of 4. Wing: Length of cell R2 1.6-1.8 length of vein R2+3; length

of cell M1 1.0-1.1 length of cell R2; scales fewer and smaller, particularly posteriorly.

Abdomen: Terga same as in female but basolateral white spots smaller and sometimes

104 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

less distinct (comprised of fewer scales in loose cluster) on anterior terga (II-IV or V).

Sterna as in female, tergum VIII (ventral in position) like sterna. Genitalia (Fig. 37): Form

as figured; closely resembling those of infula Theobald of the Oriental Region. Ninth

tergal lobe not produced, indicated by line of 3-7 (mode 6) setae. Gonocoxite rather

short, lateral setae as in poicilipes; subapical lobe moderately prominent, part bearing

setae a and b more strongly produced; setae a and b rather short and stout, c slender

and removed from a and b, all usually hooked at tip; 4 or 5 simple setae in group a-f, 1 (f

?) longer and heavier (flattened) than the others; seta g short and lanceolate (sometimes

duplicated in specimens from the Oriental Region); h not distinctly differentiated, short

and acuminate. Gonostylus rather short, simple, tapered distally; gonostylar claw of the

usual form. Phallosome much longer than broad; aedeagal sclerite larger than lateral

plate, angled ventrad, crest about as large as proximal part, proximal part with patch of

minute bumps along ventral margin of mesal surface; lateral plate with strongly

differentiated inner and outer divisions, inner division (= ventral arm) a laterally

compressed lobe with a fuzzy covering of minute spicules, dorsocaudal angle rounded,

ventrocaudal angel produced ventrad and pointed, dorsal margin with a flat laterally

directed single or bilobed process (= dorsal arm 7?); outer division (= lateral arm)

developed as a large ventrolateral winglike flap, dorsal process and dorsal articulatory

process indicated by a thickened ridge. Proctiger rather long; paraproct long and

narrow, basal lateral arm short and slender, crown a relatively small cluster of rather short

spinelike spicules and a few small lateral blades. Cercal sclerite with 2 or 3 setae on

posterolateral margin. Tergum X small.

Pupa (Fig. 37). Character and positions of setae as figured, range and modal

number of branches in Table 18; well distinguished from other species of the subgenus

in the region by the darkened cuticle around the bases of most setae, differs from other

species of the sitiens group in having setae 11-CT, 3-I-Ill, and 1-VII normally all single.

Cephalothorax: Lightly to moderately tanned a bright yellowish color. Setae short, all

except 5,10,11-CT usually double (see table); 5-CT double or triple, more often triple;

10-CT frequently with 7 branches (5-10); 11-CT usually single, sometimes double.

Trumpet: Moderately to heavily tanned, brown, tracheoid area darker; strongly flared,

funnel-shaped, slightly bent at tracheoid area; index 4.0-5.7, mean 4.6; pinna large,

about 0.4 trumpet length; tracheoid area about 0.3 trumpet length. Abdomen: Lightly

to moderately tanned a bright yellowish color, cuticle around bases of almost all setae

strongly tanned a dark brown (not shown in figure); length 3.3-3.9 mm, mean 3.6 mm;

caudolateral corners of terga VII and VIII sharply produced as in poicilipes. Seta 7-| single

to triple, 7-Il single or double; 1-Il single or double, aciculate; 1-IIl-V! moderately

developed, 1-IIl often with 6 or 7 thin branches (6-10), branches of 1-IV-VI thicker, 1-IV

very often with 4 branches (4-6), 1-V double or triple, 1-VI single or double; 1-VII single,

thinner than 1-IV-VI; 5-IV-VI resemble 1-IV-VI respectively, slightly longer, about length

of following tergum; 6-III-VI single, 6-III,IV relatively short and thin, 6-V,VI longer and

thickened, markedly different from 6-III,IV. Paddle: Inner part darkened; length

0.98-1.05 mm, mean 1.02 mm, width 0.76-0.81 mm, mean 0.79 mm, index 1.3-1.4.

Larva (Fig. 38). Form and placement of setae as figured, range and modal

number of branches in Table 38; bearing little resemblance to other members of the

subgenus in southwestern Asia, easily recognized by the triangular dorsomentum with

its minutely serrated edges and the absence of a discrete median labral plate. Head:

Length 0.61-0.64 mm, mean 0.63 mm; width 0.90-1.02 mm, mean 0.97 mm; lightly

tanned, dorsomentum and collar moderately to heavily tanned. Median labral plate

indistinguishably fused to cranium. Hypostomal suture complete, extended posteriorly

from posterior tentorial pit but ending well before collar. Collar large and well developed,

Harbach: Subgenus Culex in Southwestern Asia 105

heavily tanned only on posterior margin. Dorsomentum an equilateral or isosceles

triangle with numerous minute teeth on lateral sides. Seta 1-C long, stout, slightly

swollen just beyond midlength, sharply pointed; 2-C absent; 4-C usually double (1-4);

5-C double or triple; 6-C double; 7-C distinctly shorter than 5,6-C, branches thinner

(4-7); 10,15-C double or triple; 11,13-C both usually single (see table). Antenna:

Length 0.36-0.48 mm, mean 0.41 mm; lightly tanned, distal part darker near tip; proximal

part with few spicules on dorsal and ventral surfaces; seta 1-A inserted near middle, with

about 19 branches (16-27); 2,3-A subapical. Thorax: Integument hyaline, smooth.

Seta 4-P short, about half length of 1-P, usually double, sometimes single; 7-P triple;

8-P usually double (2,3), only slightly larger than 4-P. Tubercles of meso- and

metathoracic pleural setal groups large and heavily tanned; seta 13-T small, with 3-6

branches. Abdomen: |Integument like that of thorax. Dorsal and ventral setae all shorter

than usual, especially 1 and 13; seta 3-I usually double (1-3), 3-II-VII usually double or

triple (see table); 6-I-V normally triple (see table), 6-VI normally double; 1-III-VII normally

double or triple; 13-III-V,VII usually double, sometimes triple, 13-VIl seldom with 4

branches. Segment Vill: Comb with 5-7 (mode 6) large spinelike scales with tiny

spicules on sides of expanded base. Setae 2,4-VIlIl normally double, 2-VIlIl seldom triple.

Siphon: \Index 5.01-8.35, mean 6.83; long and slender, broadest at base, sigmoid in

lateral view; lightly tanned, basal rim darkened before acus. Pecten very short, confined

to basal 0.1; comprised of 5-8 (mode 7) inconspicuous spines with 0-3 tiny spicules on

ventral side at base. Seta 1-S in 4 pairs, widely spaced, in line, usually double or triple

(see table), no longer than diameter of siphon at point of attachment; 2-S rather longer

than usual, straight, stiff, darkly tanned. Segment X: Saddle complete; lightly tanned;

smooth; length 0.38-0.45 mm, mean 0.43 mm, siphon/saddle index 3.89-5.69, mean

4.84. Seta 1-X often triple (2-4); 2-X often with 3 or 4 branches (2-5); 4-X usually in 6

pairs. Anal papillae subequal, longer than saddle.

Systematics. The concept of bitaeniorhynchus presented here is essentially

that of Sirivanakarn (1976). This concept includes populations in southwestern Asia and

Africa which Edwards (1941) and all later workers recognized under the name of Culex

ethiopicus Edwards, 1912b. | could find no distinct differences in any stage to separate

these populations from typical bitaeniorhynchus in the Oriental Region. The form which

Edwards (1941) and later authors confused for bitaeniorhynchus in Africa appears to be

an unnamed species related to Culex infula Theobald, 1901a which occurs in countries

east of Pakistan. This species remains unnamed because none of the currently

available material is individually reared and is inadequate for description.

Culex bitaeniorhynchus and Taeniorhynchus ager Giles, 1901b are objective

synonyms. They are based on the same name-bearing type female which is apparently

lost (Harbach, 1983). Nevertheless, there are two males in the British Museum which

were labelled as types of ager by Theobald (1901b). These are the specimens which

Edwards (1922), Bram (1967b), and Sirivanakarn (1973; 1976) used to establish the

concept of "typical" bitaeniorhynchus in the Oriental Region. Bram (1967b) designated

one of these specimens as the lectotype of ager. Unfortunately, these specimens have

absolutely no taxonomic standing and the lectotype designation is invalid (see Harbach,

1983). But since these specimens already serve as a standard for the current

application of the name of this species, the specimen bearing the invalid lectotype label

of ager is hereby designated as the neotype of Culex bitaeniorhynchus Giles. This

action ensures taxonomic stability and restricts the type locality to Madras, India. The

neotype is identified by the following labels: "Paddy field / [two illegible marks] /

24.12.99 // Taeniorhynchus / ager / (Type). Theobald // Capt. Cornwall. / Madras. // Type

// LECTOTYPE / Taeniorhynchus / ager Giles / By R. A. Bram '66"; with genitalia on an

106 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

acetate strip attached to the pin that supports the specimen.

Culex bitaeniorhynchus is widely distributed in the Afrotropical and Oriental

regions. Its range also includes part of the Australian Region and the eastern part of the

Mediterranean Subregion of the Palaeactic. It is possible that bitaeniorhynchus consists

of more than one biological species, but there is no indication of geographical

differentiation. The same degree of variation occurs within individual populations that

occurs throughout the known range of the taxon.

Bionomics. The immature stages of bitaeniorhynchus occur in permanent or

semipermanent bodies of standing water containing quantities of filamentous green

algae (Spirogyra). This species apparently bites man in the Oriental Region where it is

suspected of being involved in disease transmission. It has been found naturally

infected with larvae of Wuchereria bancrofti in India (lyengar, 1938) and Brugia malayi in

Sri Lanka (Carter, 1948). It also is associated with Murray Valley encephalitis in Australia

and with Batai virus in India (Karabatsos, 1985). Laboratory studies have shown that

Japanese encephalitis virus can be transmitted transovarially (Soman and Mourya, 1985)

and Sindbis virus has been isolated from populations in the Philippines and Australia

(Rudnick et al., 1962; Doherty et al., 1963). Females have not been reported to bite

man in southwestern Asia or Africa (as ethiopicus Edwards).

Distribution. Culex bitaeniorhynchus has a very wide distribution in the tropical

and subtropical areas of Palaeogaea. Populations occur in the Afrotropical Region,

eastern and southern areas of the Palaearctic Region, and the Oriental and Australian

regions. They occur in the southern coastal lands of southwestern Asia.

Material examined. 276 specimens. A total of 76 specimens (20 females, 18

males, 8 male genitalia, 9 pupal exuviae, 6 larval exuviae, and 15 fourth-instar larvae)

were examined from southwestern Asia — IRAN: (Hajiabad, Shah-Kahoor); PAKISTAN

(locations east of the Indus River): (Kahnakacha, Kasur, Lahore, Sargodna, Sattoki);

P.D.R. YEMEN: (Wadi Ma’adin); YEMEN ARAB REPUBLIC: (Wadi Siham near 'Obal).

Another 200 specimens (81 females, 47 males, 27 male genitalia, 3 pupal exuviae, 8

larval exuviae, and 34 fourth-instar larvae) were examined from BENIN, ETHIOPIA,

GAMBIA, GHANA (including the type series of ethiopicus ), INDIA (including the

neotype), MAURITANIA, NEPAL, NIGERIA, SUDAN, TANZANIA, UGANDA, ZAIRE,

ZIMBABWE, and unknown localities in Africa.

During this study 56 specimens (18 females, 13 males, 12 male genitalia, 1 pupal

exuviae, 6 larval exuviae, and 6 fourth-instar larvae) representing a species previously

regarded as bitaeniorhynchus in Africa were examined from BENIN, ETHIOPIA,

GAMBIA, GHANA, SENEGAL, SUDAN, TANZANIA, UGANDA, UPPER VOLTA, ZAIRE,

and ZIMBABWE.

Culex (Culex) tritaeniorhynchus Giles

tritaeniorhynchus Giles, 1901a: 606. +Holotype female: Travancore,

[Madras State], India (BM).

biroi Theobald, 1905b: 82. Syntypes (male; female): Bombay, India

(HNM). Synonymy with tritaeniorhynchus by Edwards, 1913c: 233.

summorosus Dyar, 1920: 180. Holotype male: Los Banos, [Laguna,

Luzon], Philippines (NMNH). Synonymy with tritaeniorhynchus by

Bram, 1967a: 225.

siamensis Barraud and Christophers, 1931: 283. Lectotype male:

Chiang Mai, Thailand; designated by Mattingly, 1956: 37 (BM). Syn-

Harbach: Subgenus Culex in Southwestem Asia 107

onymy with summorosus by Colless, 1957: 98.

Culex tritaeniorhynchus Giles of Theobald, 1901a: 364 (India, A key, M*, F*); Edwards,

1913¢: 233 (Oriental Region, syn., A, tax.); Barraud, 1920: 324 (Iraq, coll. rec.);

Austen, 1921: 116 (Israel, Jordan, coll. rec.); Barraud, 1923: 940 (India, L*);

Buxton, 1923: 316 (Israel, Jordan, coll. rec., A, L bionomics); Barraud, 1924a:

995 (India, syn., M", F, distr.); Kirkpatrick, 1924(1925): 367, 372 (Egypt, A, L

keys); Parr, 1943: 247-251 (Syria?, Lebanon, L bionomics, A, P, L keys); Lewis,

1945: 23 (Sudan, L key); Theodor, 1952: 113 (Middle East, zoogeogr.); Dow,

1953: 689 (Iran, Coll. rec., L bionomics); Gad, 1956: 136 (Egypt, distr., bionomic

note); Reuben, 1969: 650 (India, A key, M*, F*, L*); Margalit et al., 1971: 323

(Israel, bionomics); El-Said and Kenawy, 1983a (Egypt); Zaini et al., 1983: 117

(Iraq, bionomic note); Kitron and Pener, 1986 (Israel, L bionomics).

Culex (Culex) thalassius Theobald of ? Abdel-Malek, 1960: 113-123 (Syria, L coll. sites,

bionomics).

Culex (Culex) tritaeniorhynchus Giles of Edwards, 1921: 339 (Iraq, Israel, Jordan, A, L

keys, tax., distr.); Edwards, 1926: 132 (Iraq, Israel, Jordan, A, L keys, A, L, distr.);

Barraud, 1934: 404 (India, A, L keys, M*, F, L*, syn., L habitat, distr.); Séguy,

1924: 32, 186 (Iraq, Israel, Jordan, A key, M, F, L, tax., distr., syn.); Kirkpatrick,

1925:° 4140 (eoypt, A, Pb Keys, syn, Mi, hy Peo Le odistr.,, bionomics):

Stackelberg, 1927: 157 (Middle East, M, F keys, A, M gen.*, distr.); Martini, 1931:

387 (Egypt, Iraq, Israel, Jordan, Turkey, A, L keys, M*, F, L*, distr.); Edwards,

1941: 299, 282 (Afrotropical Region, A key, M, F’, P, distr.); Monchadskii, 1951:

2/7 (Middle East, L key, L*, L assoc., distr.); Hopkins, 1952: 286 (L key, L,

bionomics); Knight, 1953a: 232 (Yemen Arab Republic, coll. rec., L habitat,

distr.); Khattat, 1955: 165 (Iraq, L*, L habitat, distr.); Mattingly and Knight, 1956:

104-137 (Oman, P.D.R. Yemen, Saudi Arabia, Socotra, Yemen Arab Republic, A,

L keys, L*, distr., bionomics); Parrish, 1959: 266 (Turkey); Senevet and Andarelli,

1959: 142 (North Africa, A, P, L keys, syn., M*, F, P*, L*, distr., bionomics, med.

imp.); Abdel-Malek, 1960: 113-124 (Syria, L, bionomics); Khalaf, 1962: 52 (Iraq,

M, F, L, bionomics); Derwesh, 1965: 44 (Iraq); DuBose and Curtin, 1965: 352,

354 (Mediterranean area, A, L keys); Abul-hab, 1966: 281 (Iraq, distr., L

bionomics); Abul-hab, 1968: 245 (Iraq, L key, L habitat, distr.); Lotfi, 1970: 402

(Iran, coll. rec.); Margalit and Tahori, 1970b: 153 (Israel, coll. sites); Aslamkhan,

1971: 155 (Pakistan); Lotfi, 1973: 206 (Iran, coll. sites); Gutsevich et al., 1974:

382 (Middle East, M, F, L keys, M*, F*, L*, distr., bionomics, med. imp.); Margalit

and Tahori, 1974: 88 (Israel, coll. rec.); Lotfi, 1976: 73, 76, 82 (Iran, L key, ecol.

note, L*); Sirivanakarn, 1976: 129 (Oriental Region, A, P, L keys, syn., M*, F*, P*,

L*, tax., distr., bionomics, med. imp.); Ibrahim et al, 1983: 91 (Iraq, L*, key);

Harbach, 1985a: 86, 97, 102 (SW Asia, Egypt, distr., A, L keys).

Adult. A rather drab species resembling sitiens in most respects, but readily

distinguished by its smaller size, shorter cell R1 of the wing, lack of speckling on the fore-

and midfemora, and the presence of ventral pale scaling proximal to the median pale ring

of the proboscis.

FEMALE. Head: Antenna dark, pedicel yellowish to light brown; length 1.2-1.9

mm, mean 1.5 mm. Proboscis dark-scaled with narrow median pale ring, ring 0.15-0.20

proboscis length, proximal margin of ring about 0.45 from base; proximal part of

proboscis usually with scattered pale scales above and proximal extension of median

pale ring below (extension of ring often weak or absent in middle leaving an isolated spot

108 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

proximal to ring), sometimes giving appearance of a second pale ring; labella yellowish;

length 1.4-1.7 mm, mean 1.6 mm. Maxillary palpus dark-scaled, pale scales at apex of

palpomere 4; length about 0.3 mm, about 0.2 length of proboscis. Forked scales of

vertex dirty yellow to brown, darker posteriorly and laterally, nearly linear anteriorly;

falcate scales pale yellow, long and relatively narrow; lateral spatulate scales yellowish

white, rather broad above, narrower below. Thorax: Integument reddish brown to

brown. Scutum with shiny golden-brown falcate scales, with paler scales on anterior

promontory, prescutellar area and often on lateral margin of fossa and supraalar area.

Scales of scutellum same as prescutellar scales. Antepronotum with some pale to dirty

yellow falcate scales confined largely to anterior area, with 2 more or less distinct rows of

setae; postpronotum with fine falcate scales more or less same color as scutal scales,

with 3-5 setae in vertical row on posterior margin. Pleura with pale scales as follows:

proepisternum with small patch of narrow spatulate scales below and mesal to upper

proepisternal setae; mesokatepisternum with rather small patches of spatulate scales on

upper corner and lower posterior border at level of katepimeron; mesepimeron with small

anterior patch of spatulate scales adjacent to upper katepisternal scales, and sometimes

with few near linear or fusiform scales among upper mesepimeral setae. Pleural setae:

2-7 upper proepisternal, 4-9 prealar, 3 or 4 upper mesokatepisternal, 7-10 lower

mesokatepisternal, 5-9 upper mesepimeral, and no lower mesepimeral. Wing: Length

2.6-3.4 mm, mean 2.9 mm; length of cell R2 2.6-3.4 length of vein R2+3, mean 2.9;

length of cell M1 0.8-0.9 length of cell R2; dark-scaled with short line of pale scales at

base of costa, pale line often weakly developed or absent; dorsal surface with spatulate

scales on costa, subcosta, R and R1, R3, M3+4 and CuA, dorsal surface of other veins

with long linear scales; ventral surface with linear scales on all veins except costa and

subcosta. Halter: Pale with distal part of scabellum and base of capitellum usually

dark-scaled, capitellum sometimes largely dark-scaled. Legs: Forecoxa always with pale

scales at base, rest of scales either entirely dark, dark proximally and pale distally, or with

dark and pale scales mixed, frequently with few narrow pale scales posterolaterally at

apex; midcoxa with vertical patch of pale scales on anterior side of lateral midline and

small patch of dark scales distally on anterior surface; hindcoxa with vertical line of

indistinct pale scales on anterior side of lateral midline. Trochanters each with small

patch of dark scales on posteroventral surface. Forefemur largely dark-scaled, poorly

contrasted but darker anteriorly than posteriorly, with narrow rather indistinct knee spot;

midfemur dark-scaled anteriorly, pale-scaled posteriorly and ventrally, dark and pale

areas better contrasted than on forefemur, dark area forming narrow band at apex, with

very indistinct knee spot; hindfemur pale-scaled with variable, often indefinite,

anterodorsal dark stripe that abruptly widens distally to form more or less distinct narrow

apical band, with very narrow knee spot. Foretibia dark-scaled dorsally (darker than

femur), pale-scaled ventrally, with narrow dorsal pale spot at apex; midtibia dark-scaled

anteriorly, pale-scaled posteriorly, with faint indication of pale scaling at apex, particularly

on anterior surface; hindtibia dark-scaled dorsally, pale-scaled ventrally, with indistinct

dorsal pale spot at apex and indication of dorsal pale spot at base. Tarsi with narrow,

indistinct basal and apical pale rings on tarsomeres 1-4; if pale ring present on apex of

midtarsomere 4, then pale ring usually also evident at base of midtarsomere 5; rings

between midtarsomeres 3-4 and 4-5 often absent. Abdomen: Tergum | with median

posterior patch of dark scales; terga II-VIl with narrow, slightly convex basal pale bands

(0.15-0.35 tergum length) and basolateral pale spots that become larger in progression

from anterior to posterior terga, tergum VII often with median posterior patch of pale

scales; tergum VIII largely pale-scaled with narrow apical dark band. Sterna pale-scaled,

frequently with dark scales on posterolateral corners; sternum VIII with lateral patches of

Harbach: Subgenus Culex in Southwestem Asia 109

pale scales, without scales medially.

MALE. Like female except as follows. Head: Proboscis with ventral cluster of

setae on proximal side of false joint. Maxillary palpus extending beyond tip of proboscis

by at least length of palpomere 5, length 1.8-2.8 mm, mean 2.4 mm; palpomere 3 with

subapical stripe of pale scales on dorsal and lateral surfaces, often only subapical line on

lateral surface, ventromesal margin with short setae in row extending length of pale

scaling, ventral surface with long setae distal to pale scaling; palpomeres 4 and 5 each

with narrow basal ring of pale scales and numerous long ventrolateral setae, basal rings

actually very narrowly broken on lateral and mesal margins by row of setae, palpomere 5

also usually with some pale scales dorsally at apex. Forked scales of vertex broader and

somewhat shorter. Wing: Length 2.2-3.2 mm, mean 2.8 mm; length of cell R2 1.6-2.7

length of vein R2+3, mean 2.2; scaling same as in female but scales smaller and less

dense, spatulate scales not as short and broad. Abdomen: Basal pale bands of terga

generally slightly broader, 0.22-0.40 tergum length. Genitalia (Fig. 39): Form as figured;

very much as in sitiens and mimeticus. Ninth tergal lobe with row of 3-10 (mode 6) setae.

Gonocoxite with 3 or 4 rows of lateral setae extending from just above subapical lobe to

point about midway between subapical lobe and base of gonocoxite; subapical lobe with

3 setae in group d-e, all flattened and hooked, 2 with subapical denticles, f flattened and

broad, broadest distally. Inner division of phallosome with 3-5 (mode 4) caudally curved

teeth. Proctiger not unusual; paraproct with slightly larger basal lateral arm, smaller

crown, and well developed ventral acetabulum. Cercal sclerite usually with 2 setae (1-4)

on posterolateral corner.

Pupa (Fig. 39). Form and placement of setae as figured, range and modal

number of branches in Table 19; different from other members of the sitiens group in

southwestern Asia in having setae 8-CT and 1-Il more branched. Cephalothorax: Lightly

tanned. Setae 1-3-CT usually triple (see table); 4,7,11-CT double, 4-CT rarely single;

8-CT shorter and more branched than usual, same length as 9-CT, with 6 or 7 branches;

9-CT usually double (2,3); 12-CT longer than 10,11-CT, usually with 4 or 5 branches

(3-5). Trumpet: Dark and rather short; index 5.88-6.25, mean 6.04; pinna 0.2-0.3

trumpet length. Abdomen: Lightly tanned; length 2.19-2.42 mm, mean 2.32 mm. Seta

7-| usually with 3 or 4 branches (2-5), 7-Il usually double or triple (2-4); 1-II-VII and 6-IlI-VI

tend to be more branched than usual, especially 1-Il (see table); 1-III-VIIl about length of

tergum following; 5-IV multiple, different in appearance from 5-V,VI which are double

with heavier and longer branches; 6-V,VI developed like 6-III,1V but distinctly longer;

4-VIill with 3 or 4 branches. Paddle: Pale; length 0.66-0.77 mm, mean 0.70 mm, width

0.46-0.54 mm, mean 0.51 mm, index 1.3-1.5, mean 1.4.

Larva (Fig. 40). Character and positions of setae as figured, range and modal

number of branches in Table 39; resembling members of the pipiens group, but

differing conspicuously in the character of seta 1-C and the pecten spines; more closely

resembling si/tiens than other members of the sitiens group occurring in southwestern

Asia, but easily distinguished by the characters given in the key. Head: Length

0.68-0.78 mm, mean 0.72 mm; width 1.03-1.11 mm, mean 1.07 mm; lightly to

moderately tanned, darker behind eyes, collar moderately to heavily tanned, dorsal

apotome with variable number of moderately tanned spots distributed somewhat as in

pipiens. Hypostomal suture complete, extended to collar from posterior tentorial pit.

Dorsomentum with 6-9 teeth (mode 7) on either side of center. Seta 1-C tapered,

sharply pointed, dark; 2-C absent; 5-C normally triple (3,4); 6-C normally double (2,3);

7-C with 5-10 branches, about as long as 5,6-C; 10,12,13-C usually triple (2-4); 11-C

double. Antenna: Length 0.52-0.58 mm, mean 0.55 mm; tanning and spiculation as in

sitiens. Seta 1-A with about 24 branches (21-35); 2,3-A subapical. Thorax: Integument

110 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

normal. Seta 4-P double; 7-P normally triple (2,3), slightly longer than 4-P; 8-P double,

weaker than 4-P. Seta 1-M single, longer than 2-M; 2-M multiple (4-8); 4-M usually triple

(2,3); 5-M shorter than usual. Seta 1-T single, not more than 0.5 length of 2-T; 2-T

double or triple; 13-T with 7-11 branches, not much longer than 12-T. Abdomen:

Integument normal. Seta 3-1,II,1V usually triple (2,3), 3-I,1l rarely with 4 branches; 3-Ill

commonly double (2,3), 3-V single or double, 3-VII with 4-6 branches, often with 5; 6-I,II

normally triple (2,3), 6-Il shorter than 6-1; 6-III-VI usually double (2,3), 6-III, VI same length

as 6-II, 6-IV,V shorter; 7-1 most often double (1-3); 1-III-VII about 0.5 length of segment,

1-III most often triple (2-4), 1-IV,V with 4 branches, 1-VI most often with 4 branches (3-6),

1-VIl usually with more than 4 branches (4-7); 13-III-V resemble 1-III-V but usually with

more branches (see table). Segment VIII: Comb with 26-60 (mode 38) evenly fringed

scales. Siphon: Index 5.77-7.00, mean 6.30; moderately tanned, slightly darker at

apex, basal ring and acus darker. Pecten of 12-18 (mode 14) spines with ventral row of

denticles, larger spines with 7-12 denticles. Seta 1-S normally in 6 pairs, occasionally in

5 Or 5.5 pairs, 1 subapical pair (1e-S) borne laterally, others borne posterolaterally, 3

proximal pairs (1a,b,c-S) commonly with 4 branches (2-6), 3 distal pairs (1d,e,f-S) most

often triple (2-4), only slightly shorter than proximal pairs, all a little longer than diameter

of siphon at point of attachment. Segment X: Saddle complete; lightly to moderately

tanned, darker posterodorsally; with conspicuous spicules posteriorly; length 0.29-0.36

mm, mean 0.33 mm, siphon/saddle index 4.23-6.14, mean 4.73. Seta 1-X often triple

(2-4); 2-X usually with 3 or 4 branches, seldom double; 4-X usually in 6 pairs, an

additional unpaired seta occasionally present. Anal papillae elongate, subacutely

tapered, about as long as saddle.

systematics. This and the next species are members of the complicated

vishnui subgroup of Bram (1967b). The group is almost wholly Oriental, but some

members also occur in adjacent areas. Culex tritaeniorhynchus and pseudovishnui are

the only members of the group which occur in southwestern Asia. Culex

tritaeniorhynchus is common throughout the area and many coastal countries of Africa.

Culex tritaeniorhynchus should not be confused with any other species in

southwestern Asia except pseudovishnui. It is readily differentiated from this species by

the development of the comb scales and pecten spines of the larva. The pupa differs in

the character of setae 8-CT and 1-Il and the posterolateral corners of tergum VIII. There

is considerable overlap in the adults, but specimens of tritaeniorhynchus usually can be

recognized relatively easily by the more extensive ventral pale scaling of the proboscis.

The male genitalia are very nearly congruent, differing essentially only in the size of the

teeth of the inner division of the phallosome.

Culex tritaeniorhynshus is a peculiar species which, as pointed out by

Sirivanakarn (1976), is similar to members of the barraudi subgroup in the larval stage.

This situation parallels that of poicilioes and whitmorei which are included as members of

the sitiens subgroup in spite of conspicuous larval disparity. It is apparent that the

present system of grouping species of the sitiens group does not accurately reflect

species affinities based on all life stages. This group more than any other in the

subgenus Culex brings to mind Belkin's (1962) contention that hybridization seems to

have been an important mechanism of speciation in mosquitoes.

Bionomics. Culex tritaeniorhynchus is a common species in southwestern

Asia. The immature stages are typically found in rice fields, flood waters, and marshy

areas with floating or emergent vegetation. They also are found in ponds, swamps,

streams, springs, irrigation ditches, grassy pools, seepages, and animal hoofprints. The

water may be fresh or slightly brackish but is always clear and clean.

Females of tritaeniorhynchus are known to enter dwellings and bite man during

Harbach: Subgenus Culex in Southwestem Asia 174

any time at night (Kirkpatrick, 1925; Khalaf, 1962). In Iran and Iraq, tritaeniorhynchus is

considered to be a vector of Japanese encephalitis virus (Travis and Labadan, 1967).

The susceptibility and transmission efficiency for West Nile virus in the laboratory

indicate that tritaeniorhynchus is an excellent potential vector of this pathogen in

Pakistan (Hayes et al., 1980). A single isolation of Sindbis virus was made from a pool of

specimens collected at Al Khobar, Saudi Arabia (Wills et al., 1985). This species has

been associated with various viruses and filarial pathogens in many areas of eastern and

southeastern Asia.

Distribution. Culex tritaeniorhynchus occurs in eastern Asia, the Oriental

Region, southwestern Asia, and Africa.

Material examined. 570 specimens. A total of 536 specimens (221 females,

137 males, 69 male genitalia, 23 pupal exuviae, 40 larval exuviae, and 46 fourth-instar

larvae) were examined from southwestern Asia — IRAN: (Aliabad, Basra, Cham Asbi,

Chomaisaray, Hassan Kiadeh, Kanroud, Karken River, Sabzab, Shour River, Zahedan,

Zarjub); IRAQ: Basrah, Galala, Magil, Rawasnduz, Tal A'far, Thallal); ISRAEL: (Beisan,

Galilee, Khirbet Hadrah, Neot Hokikar); JORDAN: (Yabes); OMAN: (Salalah); PAKISTAN

(including localities east of the Indus River): (Balloki, Kasur, Khulna, Lahore, Peshawar):

P.D.R. YEMEN: (Bateis, Dis Town, Hami, Mukalla, Urfat Subai); SAUDI ARABIA: (Alajam,

Al Khobar, Al Qatif, Dhahran, Saihat); TURKEY: (Adana, Silifke). An additional 34

specimens (19 females, 6 males, 5 male genitalia, and 4 fourth-instar larvae) were

examined from BANGLADESH, ETHIOPIA, INDIA (the holotype female), KENYA,

SOCOTRA, and unknown localities.

Culex (Culex) pseudovishnui Colless

pseudovishnui Colless, 1957: 88. Holotype female: Singapore (BM).

neovishnui Lien, 1968: 230. Holotype male: Peiyuan, Tungho,

Taitung, Taiwan (TMRI). Synonymy with pseudovishnui by

Sirivanakarn, 1976: 116.

Culex pseudovishnui Colless of Reuben, 1969: 648 (India, M*, F*, L*, A key); Zaim and

Cranston, 1984: 179 (Iran, L, coll. sites).

Culex (Culex) pseudovishnui Colless of Aslamkhan, 1971: 155 (Pakistan); Sirivanakarn,

1976: 116 (Oriental Region, M*, F, P*, L*, syn., distr., bionomics, med. imp.);

Harbach, 1985a: 86, 97, 107 (Iran and Pakistan only, distr., A, L keys).

Culex vishnui Theobald of Barraud, 1923: 938 (India, L*); Barraud, 1924a: 996 (in part:

India, A key, syn., M*, F, distr.).

Culex (Culex) vishnui Theobald of Edwards, 1921: 339 (Iraq, A key, bionomics, distr.):

Edwards, 1926: 132 (Iraq, A, L, keys, A, distr.); Séguy, 1924: 30, 186 (Iraq, A

key, M, F, L, tax., distr., syn.); Stackelberg, 1927: 158 (in part; S. Asia, M, F keys,

A, M gen.*, distr.); Martini, 1931: 392 (in part; S. Asia, A, L keys, M*, F, L*);

Barraud, 1934: 400 (in part; India, A, L keys, M*, F, L*, syn., L habitat, distr.); ?

Monchadskii, 1951: 284 (in part; S. Asia, L key, L*, bionomic note, distr.); Khalaf,

1962: 55 (Iraq, A, M gen., L); Abul-hab, 1968: 246 (Iraq); Lotfi, 1973: 206 (Iran,

bionomics, distr., coll. sites); Gutsevich et al., 1974: 371 (in part; S. Asia, L key):

Lotfi, 1976: 72, 78, 79 (Iran, L key, ecol. note, L*).

Adult. A small brown species closely resembling tritaeniorhynchus, but the

median pale ring of the proboscis is not extended proximally on the ventral surface and

142 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

there is often an indefinite anterior pale stripe on the hindtibia. The proboscis of the

male may have one or two short setae at the base of the median pale ring but is without a

distinct ventral cluster of setae.

FEMALE. Head: Pedicel of antenna yellowish laterally, mesal surface dark brown

with some tiny inconspicuous scales; flagellum normal, dark, length about 1.8 mm.

Proboscis length 1.5-1.8 mm, mean 1.7 mm; dark brown to blackish-scaled with median

pale ring of cream-colored scales 0.40-0.65 from base. Maxillary palpus dark-scaled with

few pale scales at apex; length about 0.3 mm, approximately 0.2 proboscis length.

Vertex with whitish to pale golden falcate scales, paler along margin of eye, sometimes

darker laterally; forked scales white to yellow in middle, dark brown to black

posterolaterally; lateral spatulate scales white, or nearly so. Interocular space with few

scales, same color as ocular scales. Ocular setae golden brown to brown, often paler

near middorsal line of cranium; 2 golden interocular setae project ventrally over clypeus.

Thorax: Scutal integument brown; pleural integument yellowish brown or paler, with

dark brown areas on proepimeron, anterior margin of mesokatepisternum, postspiracular

area, prealar area, between upper and lower mesokatepisternal scale-patches, below

anterior mesepimeral scales and on upper portion of mesepimeron. Scutal scales rather

long and narrow, usually predominantly pale, color ranging from silvery white to golden,

frequently pale golden with whitish scales along margins and on prescutellar area; darker

scales, when present, either indistinct or distinct, occurring on acrostichal, median scutal

fossal and/or posterior dorsocentral areas, usually golden brown. Scutal setae dark

brown with reddish and/or golden sheen. Lobes of scutellum with scales resembling

prescutellar scales; each lateral lobe with 3-5 large setae, median lobe with 5-7.

Antepronotum with brown setae, some paler, often yellowish, ventrally; with whitish to

pale golden falcate scales, coarser and paler ventrally. Postpronotum with whitish to

pale golden falcate scales on about dorsal 0.5, slightly coarser and paler posteriorly; with

5 or 6 dark setae on posterodorsal margin. Pleural setae yellow to brownish yellow: 4-6

proepisternal, 8 or 9 prealar, 4 or 5 upper mesokatepisternal, 7 or 8 lower meso-

katepisternal, 4 or 5 upper mesepimeral; lower mesepimeral seta absent. Pleura with the

usual patches of elongate white spatulate scales: small patch below proepisternal setae,

patches on upper corner and lower posterior margin of mesokatepisternum, anterior

patch on mesepimeron at level of upper patch on mesokatepisternum, and small patch

before upper mesepimeral setae; upper proepisternal and upper mesepimeral patches

usually with only 2 or 3 scales, or sometimes absent. Wing: Length 2.8-3.2 mm, mean

3.0 mm; length of cell R2 1.5-2.8 length of vein R2+3, mean 2.3; length of cell M1

0.8-0.9 length of cell R2; scales predominantly dark, with variable amount of pale scaling

on costa, subcosta, R and R1, pale scales always present on posterior side of costa near

humeral crossvein. Halter: Integument and scales entirely pale. Legs: Anterior surface

of forecoxa entirely white-scaled or with variable amount of black scaling in middle, most

often with black scales; midcoxa with longitudinal patch of white scales on anterior side

of midlateral row of strong setae; hindcoxa with longitudinal patch of white scales on

anterolateral surface, posterolateral surface with irregular row of strong setae. Lateral

surfaces of trochanters without scales, other surfaces white-scaled, anterior surface of

foretrochanter sometimes with dark scales. Apices of all femora with narrow white knee

spots; anterior surface of forefemur brown-scaled, posterior surface white-scaled;

midfemur like forefemur but brown scaling gradually expanded over dorsal surface

toward apex and white scaling sometimes expanded over ventral surface onto lower part

of anterior surface; hindfemur mainly white-scaled, with anterodorsal stripe of brown

scales, stripe gradually spreading over dorsal surface distally and then abruptly

expanded onto anterior and posterior surfaces near apex. Foretibia whitish-scaled with

Harbach: Subgenus Culex in Southwestern Asia ia,

complete dorsal stripe of brown scales; midtibia mainly whitish-scaled, ventral surface

with complete stripe of brown scales, dorsal surface with narrow, usually obscure brown

stripe that is largely obsolescent in middle; hindtibia mainly brown-scaled, narrowly pale

at base and apex, often with ill-defined anterior and posterior stripes of whitish scales

usually not reaching ends of tibia and normally joined ventrally for most or part of their

length. Tarsi with narrow pale basal bands on tarsomeres 1-3, frequently on 4, and

occasionally on 5; tarsomeres 1 and 2 also frequently with very narrow pale apical bands;

posterior surface of tarsomere 1 usually pale-scaled. Abdomen: Tergum | with

posteromesal patch of blackish scales; terga II-VIl mainly clothed in dark brown to

blackish scales, with basolateral patches of white scales, tergum II with basomedian spot

of pale scales; terga III-VII with straight or convex basal bands of white or nearly white

scales, bands usually about 0.25 tergum length but often narrower and sometimes not

reaching basolateral pale patches, particularly on more anterior terga; tergum VIII usually

with broad basal pale band and narrow apical dark band. Sterna II-VII mainly pale-scaled

with large apicolateral dark patches, dark patches sometimes more or less joined mesally,

particularly on more posterior sterna; sternum VIII with lateral pale patches, broad median

area devoid of scales.

MALE. Resembling female except for following sexual differences. Head:

Proboscis with narrow pale band about 0.50 to 0.65 or 0.70 from base, sometimes with 1

or 2 long setae ventrally at base of band; false joint inconspicuous. Maxillary palpus with

bands of pale scales and/or pale integument on about distal 0.5 of palpomere 2, middle

of palpomere 3, bases of palpomeres 4 and 5 and apex of palpomere 5; ventrolateral

margin of palpomere 3 with 10-20 long dark setae at apex; ventromesal margin with

complete row of short, translucent, antrorsely curved, scalelike setae that increase

progressivly in length from base to apex of palpomere; lateral and mesal surfaces of

palpomeres 4 and 5 densely setose; longer than proboscis by length of palpomere 5

and part of 4. Thorax: Scutal scales finer, more uniformly colored. Wing: Length of cell

R2 1.5-2.1 length of vein R2+3, mean 1.8; subcosta intersects costa before furcation of

R2+3. Abdomen: Terga Il-VIl without basolateral pale patches; basal bands of terga III-VII

broader, usually 0.30-0.45 tergum length, bands usually straight or nearly so, those of

terga VI and VII produced posteriorly along lateral scale-free areas; tergum VII sometimes

with pale scales apically in middle; tergum VIII (ventral in position) with approximately

basal 0.5 pale-scaled. Sterna Il-Vil with dark apical bands that are broadest in middle,

sometimes with few pale scales centrally on posterior border; sternum VIII (dorsal in

position) like tergum VIII of female but sometimes with pale scales along posterior

margin. Genitalia (Fig. 41): Form as figured; constructed as in sitiens, mimeticus, and

tritaeniorhynchus. Ninth tergal lobe with row of 3-7 (mode 4) short setae. Gonocoxite

and phallosome closely resembling those of tritaeniorhynchus, inner division of

phallosome differs in having distinctly larger teeth (4 or 5, mode 5) and the dorsocaudal

angle usually notched. Proctiger as in tritaeniorhynchus; cercal sclerite with 2 or 3,

usually 2, setae on posterolateral corner.

Pupa (Fig. 41). Character and positions of setae as figured, range and modal

number of branches in Table 20; similar to tritaeniorhynchus, distinguished by setae

8-CT and 1-ll with fewer branches and posterolateral angle of tergum VIII acute.

Cephalothorax: Lightly tanned. Seta 2-CT usually with 3 or 4 branches, more often with

4 (3-5); 8-CT with 3-5 branches, distinctly longer than 9-CT; 12-CT about as long as

10,11-CT, usually triple (2-5). Trumpet: Moderately tanned, tracheoid area darker; index

5.45-8.10, mean 6.61; pinna 0.15-0.35 trumpet length. Abdomen: Lightly tanned;

length 2.34-2.96 mm, mean 2.69 mm. Seta 7-I usually double (2-4); 1-II-VII distinctly

shorter than tergum following; 6-III-VI commonly with 4 or 5 branches (see table), all

114 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

about same length; 5-IV shorter and 5-V,VI about the same as in tritaeniorhynchus; 4-VIII

usually double or triple (2-4). Paddle: Very lightly tanned; length 0.72-0.83 mm, mean

0.75 mm, width 0.47-0.56 mm, mean 0.53 mm, index 1.4-1.6, mean 1.4.

Larva (Fig. 42). Form and placement of setae as figured, range and modal

number of branches in Table 40; distinct from sitiens, tritaeniorhynchus, and members

of the pipiens group in southwestern Asia in the character of the comb scales, easily

distinguished from other members of the sitiens group in the region by the characters

given in the key. Head: Length 0.71-0.88 mm, mean 0.76 mm; width 1.08-1.22 mm,

mean 1.16 mm; lightly tanned, darker posteriorly. Dorsomentum heavily tanned; teeth

large and distinct, 5-7 (mode 6) on either side of center. Seta 1-C long, relatively

slender, tapered, dark; 2-C absent; 5-C usually triple (2-4); 6-C usually double (1-3); 7-C

often with 8 or 9 branches (8-12), about as long as 5,6-C; 10-C with 3 or 4 branches on

short stem; 11,13-C usually double (1-3); 12-C frequently double but often more

branched (2-5). Antenna: Length 0.60-0.71 mm, mean 0.66 mm; tanning and

spiculation as usual. Seta 1-A with about 30 branches (27-30); 2,3-A distinctly

subapical. Thorax: Integument hyaline, with rather distinct thumbprint pattern, lateral

surfaces and dorsal surface anteriorly with minute spicules. Seta 4-P normally double

(1-3), nearly as long as 7-P; 7-P triple; 8-P normally double (1,2). Seta 1-M single or

double in nearly equal frequencies; 4-M usually triple (2,3). Seta 1-T single or double,

much shorter than 2-T; 2-T double or triple; 13-T often with 9 or 10 branches (7-12), not

much longer than 12-T. Abdomen: Hyaline, thumbprint pattern evident. Seta 3-|

variable, with 3-7 branches, most often with 3; 3-ll-IlV usually double or triple, 3-Il, III

occasionally with 4 branches; 3-V usually single (1,2); 3-VIl with 3-7 branches; 6-I,II triple,

6-III-VI usually triple (see table), 6-1 distinctly longer than the others, 6-III,IV not much

shorter than 6-II,V,VI; 7-1 single; 1-lII-VII slightly longer than 0.5 length of tergum, 1-Ill,IV

frequently with 5 or 6 branches, 1-V,VI often with 6 or 7 and 1-VII often with 7 or 8 (see

table). Segment VIII: Comb with 5-10 (mode 6) scales developed as in poicilipes and

bitaeniorhynchus. Siphon: \|Index 4.41-6.87, mean 5.40; lightly tanned, basal rim dark;

surface with rows of minute vesicles; very slightly bent forward before apex. Pecten

composed of 8-12 (mode 11) large distally curved spines with complete ventral row of

denticles. Seta 1-S in 6.5, 7 or 7.5 pairs, middle pair (1d-S) borne laterally, others borne

posterolaterally; most proximal pair (1a-S) 1.5-2.0 times as long as diameter of siphon at

point of attachment, others gradually shorter toward tip of siphon, most distal pair (1g-S)

no longer than diameter of siphon at point of attachment. Segment X: Saddle lightly

tanned, with prominent spicules on posterodorsal margin; length 0.36-0.42 mm, mean

0.38 mm, siphon/saddle index 3.13-4.29, mean 3.61. Seta 1-X with 3 or 4 branches;

2-X usually triple (2,3); 4-X in 6 pairs, often with 7-9 branches. Anal papillae rather

slender, longer than saddle.

Systematics. This species has been confused in the past with Culex vishnui

Theobald, 1901a, which is mainly restricted to the Oriental Region and occurs no farther

west than eastern India. Although pseudovishnui closely resembles vishnui, it is

definitely a distinct species which is well differentiated in the larval stage. Zaim and

Cranston (1984) corrected the previous record of vishnui in Iran on the basis of larval

characters. All life stages from populations in southwestern Asia seem to agree in all

essential respects with pseudovishnui in the Oriental Region, but too few specimens

are currently available for a detailed comparative study.

To my knowledge, pseudovishnui has never been recorded from Afghanistan.

My previous report (Harbach, 1985a) was in error, but the species probably occurs there

because it has been collected very near its borders in both Iran and Pakistan.

Culex pseudovishnui is very similar to tritaeniorhynchus in the adult stage, and

Harbach: Subgenus Culex in Southwestern Asia 115

the two forms are often difficult to differentiate. The most reliable characters that will

separate the females of these forms are the less extensive pale scaling of the proboscis

and the shorter vein R2+3 in pseudovishnui. The male genitalia cannot be differentiated

with absolute certainty even though the teeth of the inner division of the phallosome are

definitely larger in this species. The larvae are strikingly different in the form of the comb

scales and pecten spines, and the pupae differ in the development of setae 8-CT and

1-Il and in having more acute posterolateral corners on tergum VIII.

Bionomics. This species breeds in permanent standing water and transient

ground pools. Larvae are commonly found in rice fields, ditches, stream margins,

marshes, and ponds in association with tritaeniorhynchus and bitaeniorhynchus.

Females are attracted to birds, pigs, and other domestic animals and occasionally bite

man (Colless, 1959; Reuben 1971a,b). Culex pseudovishnui is a potential vector of

Japanese encephalitis virus and has been associated with Kunjin and Tembusu viruses

in Southeast Asia (Karabatsos, 1985).

Distribution. Culex pseudovishnui is widely distributed in the Oriental Region

and occurs in Japan, Korea, China, Pakistan, and Iran.

Material examined. 73 specimens. Only 38 specimens (6 females, 7 males, 7

male genitalia, 2 pupal exuviae, 7 larval exuviae, and 9 fourth-instar larvae) were

examined from southwestern Asia — IRAN: (Shemian, Zahedan); PAKISTAN (including

localities east of the Indus River): (Kohat, Lahore, Peshawar, Sargodha, Wahga).

Another 35 specimens (17 females, 10 males, 5 male genitalia, and 3 pupal exuviae)

were examined from INDIA.

116 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

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142 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

FIGURE 1

Map of southwestern Asia and Egypt

tdAOFT UNV

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FIGURE 2

Some Adult Vestiture

Thoraces, lateral (left) aspects. MeSL = lower mesanepimeral seta; MScL = lower

mesokatepisternal scales; MScU = upper mesokatepisternal scales; PoSc = post-

spiracular scales; PrSc = prealar scales.

Wings, dorsal aspects of right wings. 2A = vein 2A.

Midlegs, anterior aspects.

Hindlegs, anterior aspects.

univittatus

mimeticus

| sinaiticus

bitaeniorhynchus

Midlegs Hindlegs

I-— 2.0 —4 recon ae erie

theileri

univittatus

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theileri

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is Culex (Culex) pipiens FI Q. 3

Israel

dorsal dorsal

Fig.4

Culex (Culex) pipiens

Israel

Va a

\\ Culex (Culex) guinquetasciatus Fig.d

pupa - India

genitalia - Oman

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lateral

dorsal

Culex (Culex) vagans

Pakistan

Culex (Culex) vagans

Pakistan

\\\W : \\

\ \ \

. Culex (Culex) torrentium

Sweden

dorsal

Fig.10

Culex (Culex) torrentium

Sweden

ST Litwak

Culex (Culex) decens

Yemen Arab Republic

dorsal

Culex (Culex) decens Fig. 12

Yemen Arab Republic

Culex (Culex) antennatus Fig.14

Egypt

Culex (Culex) univittatus |

pupa - South Africa

genitalia - Salisbury, Zimbabwe (type locality)

lateral

dorsal

Rh ae %

Culex (Culex) ea uails F | Q: 16

Salisbury, Zimbabwe (type locality)

S L dita

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=, SW :

aid 4

3 a 6

| esd Ww? &

po a JN

i ] al

| ih ’ ‘

i mi ,

/ \ \

—

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lt Bhahe..

7 Culex ( Culex) perexiguus

Egypt) isd

Culex (Culex) theileri

CT pupa - Egypt

G = genitalia - Egypt and: Israel

lateral

dorsal

Culex (Culex) theileri

\ Culex (Culex) laticinctus

pupa - Yemen Arab Republic

genitalia - Israel

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nd = H]

Ss!

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cat y

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Fig. 22

Culex (Culex) laticinctus

Israel

® +; 34

LAAT

iif

Culex (Culex) mattinglyi

Yemen Arab Republic

dorsal

Culex (Culex) mattinglyi

Yemen Arab Republic ©

Culex (Culex) simpsoni

Ethiopia and Tanzania

lateral”

FER

Reames

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dorsal

ee ae

Culex (Culex) sinaiticus

Israel

Culex (Culex) duttoni

Gt genitalia - P.D.R. Yemen and Ethiopia

pupa - Uganda

dorsal

Scere

dorsal

eye ean

Culex (Culex) duttoni

Pe pR: prrectiarn and Ethiopia

Culex (Culex) sitiens

pupa - Yemen Arab Republic

ei genitalia - P.D.R. Yemen and Saudi Arabia

dorsal Mi V8 ae

Culex (Culex) sitiens

Yemen Arab Republic

(

Culex (Culex) poicilipes

Egypt

dorsal

u ' wiih OE

eta i nails Sr

TENG eB Py,

ay aad

dorsal

\\\

a fig aakonen

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Culex (Culex) poicilipes

Egypt /

Culex (Culex) mimeticus

Israel

lateral

dorsal

Fig. 36

Israel and Iran

Culex (Culex) mimeticus

Culex (Culex) bitaeniorhynchus

pupa - Pakistan

genitalia - Yemen Arab Republic and Sudan

= s

ee SENOS

V/A Som | GG

Si aly

dorsal

Fig. 38

Culex (Culex) bitaeniorhynchus

Pakistan

ZY,

Culex (Culex) tritaeniorhynchus

Pakistan

7S |

¢ 1 ‘ | jf ' <

ne CME Seay l

Hein, Why i

vines

en Te oe

Past Bane 4

444

NOs TE ERY 75 toe

ee AO ae? g

! MLE ait COE

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\ ae =

war! Ved Pe eece

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Fig.40

Culex (Culex) tritaeniorhynchus

Iran and Saudi Arabia

8 “ i ‘ i

a Pf

13 P

sav \ ‘\ \ . . N

\\\

AAR

Lak.

\ Culex (Culex ) pseudovishnui

W) ;

CT Pakistan

ES ~\ re e D

: 8 =

dorsal

rn ae

Culex (Culex) pseudovishnui

Pakistan

186 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Table 1. Number of branches for pupal setae of Culex pipiens Linnaeus.#

4 Based primarily on specimens from Egypt and Israel.

b Range (mode).

d infrequently present.

Seta Cephalothorax Abdominal Segments

No. GT | i I

0 . : 1 1,2(1)

1 3-5(4)> 36-75 10-24(18) 5-10(7)

Q 3-5(4) 1-4(1) 1 1

3 2-5(3) 2,3(2) 2,3(2) 2,3(2)

4 2-5(3) 4-8(5) 2-7(4) 3-8(4)

5 3-7(5) 1-8(5) 4-9(5) 4-10(6

6 1-5(3) 1,2(1) 1 2-4(3)

7 2,3(2) 2-4(2) 2,3(2) 4-8(5)

8 4-7(4) : . 3-5(3)

9 1,2(2) 1,2(1) 1 1

10 5-13(8) “ 2,3(2)¢ 2,3(2)

11 2,3(2) 1-3(2) 3-5 1

12 2-5(3) . - -

13 : . - -

14 : ; j

Seta Abdominal Segments

No. V VI Vie ae Vill x

0 1 1,2(1) 1 1 :

1 4-8(6) 3-7(5) 3-7(5) : 1

2 1 1 1,204) ‘ :

3 2 312) 2-4(2) 1-3(2) : .

4 3-6(4) 2-6(4) 1212) 1-3(2) :

5 2,3(2) 1-3(2) 1-3(2) : :

6 3-6(4) 3-7(4) 3-10(5) - .

7 3-6(4) 1,2(1) 1,2(1) : :

8 2-4(3) 2-5(3) 2-5(4) . :

9 1 1 4-6(5) 5-11(7) ,

10 1 1,2(1) 1 : :

14 1,2(1) 1-3(2) 1-4(2) : :

12 - . - - -

13 - - - - -

14 1 1 1 1,2(1) .

Harbach: Subgenus Culex in Southwestem Asia

Table 2. Number of branches for pupal setae of Culex quinquefasciatus Say.2

Seta Cephalothorax

No. CT

0 es

1 2,3(2)°

2 3,4(3)

3 2,3(3)

4 2-5(3)

5 2-4(4)

6 2-4(2)

Fj 1-3(2)

8 2,3(3)

9 1,2(2)

10 4-11(7)

1] 1,2(2)

12 2-4(3)

13 -

14 :

Seta

No. V

0 1

1 3-5(4)

2 1

8 1,2(2)

4 3-5(3)

5 1,2(2)

6 2-4(3)

7 3-5(4)

8 2-5(3)

9 1

10 1

11 1,2(1)

12 -

13 -

14 1

22...

aAnpawada

ae)

Pas tk

omnch

Peaicey Sees

Ae)

~=——

Abdominal Segments

12-22(16)

Vil

1,2(2)

3,4(3)

3,4(4)

4 Based on specimens from southwestern India.

b Range (mode).

d Alveolus only.

1 Qa NNeANNaN =

W O1M OOP

a DD as Ss OO as On to

MMMM = ©

aot,

7“

~=———

187

188

Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Table 3. Number of branches for pupal setae of Culex vagans Wiedemann.@

Seta Cephalothorax Abdominal Segments

No. CT i il

0 - - 1 1

1 3,4(4) -¢ 9-16(10) 6-11(10

2 3,4(4) 1,2(1) 1 1

2 2,3(3) 2-4(2) 2,3(2) 1,2(2)

4 3,4(3) 4-7(6) 2-5(4) 4-7(5)

5 4-7(4) 2-6(3) 4-6(5) 5-8(6)

6 2-4(3) 1 1 2-5(3)

Ps 2 2,3(2) 1,2(2) 4-9(6)

8 3-9(7) - - 3-5(3)

9 2-4(2) 1,2(1) 1 1

10 5-14(7 -d . 1,2(2)

11 1,2(2) 1,2(1) . 1

12 4-6(4) . . .

13 - - - :

14 - - -

Seta Abdominal Segments

No. V Vi Vil Vill xl

0 1 1 1 1

1 5-9(7) 5-7(6) 4-7(5) -

2 1 1 1 -

3 1,2(2) 2 1,2(2) .

4 3-6(4) 3,4(4) 1,2(2) 1202)

5 1,2(2) 2 2-5(2) ;

6 3-6(3) 4,5(4) 4-7(5) -

7 4-6(4) 1,2(1) 1 -

8 1-3(2) 2,3(2) 2,3(3) "

9 1 1 3-6(4) 7-11(8)

10 1 1 1 -

11 1 1,2(1) 2,3(2) .

12 : : . -

13 - - ; :

14 1 1 1

4 Based on specimens from Pakistan.

b Range (mode).

d Alveolus only.

Harbach: Subgenus Culex in Southwestern Aisa 189

Table 4. Number of branches for pupal setae of Culex torrentium Martini.4

seta Cephalothorax Abdominal Segments

No. GT | l il IV

0 - - 1 1 1

1 2-4(3) -¢ 12-16(15) 6-10(7) 5-9(5)

2 2-5(4) 1 1 1 1

3 2,3(2) 2,3(2) 2,3(2) 2 2-6(4)

4 3-5(3) 3-5(3) 3-5(3) 4,5(4) 1-3(1)

5 3-6(4) 3-5(4) 3,4(3) 2-6(5) 3,4(3)

6 3-5(3) 1 1,2(1) 2,3(2) 2,3(3)

7 2,3(2) 2,3(2) 2 5,6(5) 2-5(3)

8 1-4(3) - - 2,3(3) 2-4(2)

S 2 1,2(1) 1 1 1

10 5-12(5) .d : 2 1,2(2)

11 2,3(2) 1,2(1) - 1 1

12 2-4(3) . - - -

13 - - . - -

14 - m i j

Seta Abdominal Segments Paddle

No V VI Vil Vill x P

0 1 1 1 1 : :

1 4-9(5) 2-6(4) 2-5(4) - 1 1

2 1 1,2(1) 1 . . 1,2(1)

3 1,2(2) 1,2(2) 1,2(2) . : :

4 2-6(4) 2,3(2) 1,2(2) 1,2(2) . .

5 2 2 1,2(2) . . .

6 3,4(3) 2-5(3) 3-6(6) - - -

7 3-7(5) 1 1 - - .

8 2-5(3) 2-4(3) 2-4(3) - ~ -

9 1 1 3-5(4) 5-10(7) - .

10 1 1 1 . . .

11 1 1,2(1) 1-3(2) . . .

12 - - - . . .

13 - - - - . -

14 1 1 1 . -

4 Based on specimens from Sweden.

b Range (mode).

d Alveolus only.

190 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Table 5. Number of branches for pupal setae of Culex decens Theobald.4

Seta Cephalothorax

No. CT i

0 - - 1

1 3-5(3) -¢ 25-32(30)

2 3-5(4) 1-3(1) 1

3 3,4(3) 2 2

4 3,4(3) 4-7(5) 3-6(4)

5 3-6(4) 4-6(5) 3-6(5)

6 3,4(4) 1,207) 1-3(1)

7 2 2,3(2) 2

8 3-6(4) . -

) 2,3(2) 1-3(2) 1

10 5-10(7) -d :

17 2,3(2) 1-3(1) -

12 3-5(4) - -

13 - - -

14 : : :

Seta Abdominal Segments

No. V VI Vil VIII

0 1 1 1 1

1 3-5(5) 4-6(4) 3,4(4) -

2 1 1 { -

3 2 2 2,3(2) -

4 3-5(4) 2-4(4) 2 2

5 2 2 2 -

6 3,4(3) 3,4(3) 4-9(6) -

7 3,4(4) 1 1 -

8 2:3i3) 3,4(4) 3-5(4) -

9 1 1 2-5(4) 5-8(8)

10 1 1 1 .

11 1,2(2) 1-3(2) 1-4(2) -

12 - - ~ -

13 - - : -

14 1 1 1 1

& Based on specimens from the Yemen Arab Republic.

b Range (mode).

d Alveolus only.

Abdominal Segments

—

PR = © —

QO ONY

1 RNAQRWADH

SONGS

= a

— NO

—— ~~

~~ ~

Harbach: Subgenus Culex in Southwestern Asia

Table 6. Number of branches for pupal setae of Culex antennatus (Becker).4

Seta Cephalothorax Abdominal Segments

No. CT | ll lil

0 7 - 1 1

1 3-5(3) -¢ 7-42(31) 6-13(7)

2 3-5(4) 1 1 1

3 2-5(3) 2-5(2) 2,3(2) 2-4(2)

4 1-4(3) 5-9(7) 3-6(4) 2-8(4)

5 4-8(6) 4-10(8) 5-7(6) 6,7(7)

6 2-5(4) 1 1 3-6(4)

7 2,3(2) 2,3(2) 1,2(2) 2-8(6)

8 4-9(7) : : 2-5(4)

9 2-4(3) 1-3(2) 1 1

10 3-10(9) -d . 2

11 1,2(2) 1 . 1

12 2-5(5) - - -

13 - - - -

14 - 3

Seta Abdominal Segments

No. V VI Vil VIII x

0 1 1 1 1 -

1 5-9(6) 4-8(7) 4-7(5) : 1,2(1)

Ps 1 1 1 - -

3 2-4(3) 2,3(3) 3-5(5) : :

4 4-8(4) 3-6(4) 2 2,3(2) ;

5 2,3(2) 2 1,2(2) g *

6 3-6(6) 3-6(5) 4-12(10) ; :

7 4-8(6) 1 1 ‘ ‘

8 2-5(4) 2-5(4) 2-4(3) : 2

9 1 1 2-4(3) 6-8(8) .

10 1 1 1 - -

11 1,2(1) 1-3(2) 1-5(2) : ;

12 - - - - -

13 - - - - -

14 1 1 1 1 -

&@ Based on specimens from Egypt.

b Range (mode).

d Alveolus only.

191

192 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Table 7. Number of branches for pupal setae of Culex univittatus Theobald.4

Seta Cephalothorax

No. CT

0 i mt

1 2-3(3)> -c

2 3-6(4) 1

3 3,4(3) 1,2(2)

4 3-5(3) 5-8(5)

> 3-6(4) 6-9(7)

6 3-5(5) 1

7 2 2,3(2)

8 3-5(5) -

2) 2,3(2) 2,3(2)

10 4-10(4) -

11 1,2(2) 2

12 3 -

13 - -

14 . .

Seta

No. V VI Vil

0 1 1 1

1 5-8(7) 5-7(5)

2 1 1 1

3 2 ale) Z1G(3)

4 4-6(5) 3-5(4) 2

5 3,4(3) 2;3(3) 2

6 4,5(4) 3-5(4)

7 4-7(6) 1 1

8 1-4(3) 2-4(3) 2-4

9 1 1 3,4

10 1 1 1

11 1 1 1

12 - - -

13 - - -

14 1 1 1

&@ Based on specimens from South Africa.

b Range (mode).

d Alveolus only.

Abdominal Segments

, |

1 ,2(2)

3-5(4)

14-20(17)

QM os

2 nma do kOonn

SSESB

— © -—

ieaNDouaKRW AOD WO

Harbach: Subgenus Culex in Southwestem Asia

Table 8. Number of branches for pupal setae of Culex perexiguus Theobald.#

Abdominal Segments

Seta Cephalothorax

No. CT | il

0 - - 1

1 3,4(3)> -c 13-22(16)

2 3-5(3) 1 1

3 2-4(3) 2 2

4 2,3(3) 5-7(5) 3-6(5)

5 3-5(4) 4-7(6) 5-7(6)

6 2-4(3) { 1

7 2 1-3(2) 1-3(1)

8 4-6(4) : .

9 2,3(2) 1,2(2) 1

10 4-9(6) -d -

11 1,2(2) 1 .

12 2-4(2) . -

13 - . .

14 : j ;

Seta Abdominal Segments

No. V VI Vil Vill

0 1 1 1 1

1 4-8(5) 4-6(5) 4,5(4) -

2 1 1 1,2(1) ,

5 2 2 3,4(4) -

4 3-5(4) 2-4(3) 1,2(2) 2,3(2)

5 2-4(3) 2,3(2) 2 .

6 3,4(3) 3-5(3) 4-6(5) :

Fy 2-5(4) 1 1 i

8 2,3(3) 2-A(3) 2-4(3) :

9 1 1 3-6(3) 5-8(7)

10 1,211) 1 1 -

11 1 1,2(1) 1-3(2) .

12 - . - -

13 - . - -

14 1 1 1 1,2(1)

2 Based on specimens from Egypt.

b Range (mode).

d Alveolus only.

Rh) =

OfhNM® &

(tha QnkONN

NS GSSbSG

193

194 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Table 9. Number of branches for pupal setae of Culex theileri Theobald.2

Seta Cephalothorax

No. CT

0 =

1 3,4(4)°

2 3-5(4)

3 3,4(3)

4 2-5(4)

5 2-6(4)

6 3,4(3)

7 1,2(2)

8 4,5(5)

9 2,3(2)

10 2-4(2)

11 2

12 3-5(5)

13 -

14 -

Seta

No. V

0) 1

1 4-7(6)

2 1

2 2,3(2)

4 5,6(6)

5 2

6 2-6(3)

7 4-7(6)

8 2-6(5)

9 1

10 1

11 1,2(2)

12 .

13 -

14 1

-c 7-16(10) 6-13(10)

1203) 1 1

2 2 2,3(2)

5-8(6) 4-8(4) 5-8(7)

3-9(4) 5-10(6) 7-10(9

1 1 2-5(3)

2,3(2) 2 5-7(7)

- - 3-7(4)

1-3(2) 1 1

-d : 1,2(2)

1,2(2) - 1

Abdominal Segments

Vil Vill x

1 1,2(1)

3-6(4) .

| ‘

2-5(4) -

2 2,3(2)

2 is

6-9(8) :

| i

3-5(4) -

4-8(4) 5-10(6)

| ¢

2,3(2) .

Abdominal Segments

&@ Based on specimens from Egypt and the Yemen Arab Republic.

b Range (mode).

d Alveolus only.

Harbach: Subgenus Culex in Southwestern Asia

Table 10. Number of branches for pupal setae of Culex laticinctus Edwards.2

Seta Cephalothorax Abdominal Segments

No. CT | Hl It IV

0 : : 1 1 1

1 2,3(2) -¢ 7-14(9) 4-7(5) 4

2 2-4(3) 1-3(1) 1 eae 1

3 2,3(2) 2 2,3(2) 2,3(2) 4-7(7)

4 4-6(4) 6-9(6) 3-6(4) 4,5(4) 1-3(2)

5 3-8(4) 5-8(5) 4-7(5) 4-8(5) 2-5(4)

6 4-6(4) 1,2(1) 1,2(?) 3,4(3) 2,3(3)

7 2,3(2) 2,3(3) 1,2(2) 4-9(8) 3-5(4)

8 4-6(4) : : 4,5(4) 2-4(3)

9 2,3(2) 2 1 1 1

10 4-10(7) _d 1-3° 1-3(2) 2,3(2)

11 1,2(2) 1,2(1) -¢ 1 1

12 3,4(3) : : . -

13 : : . : .

14 - : : 1

Seta Abdominal Segments Paddle

No. V VI vil vill xl P

0 1 1 1 1 ,

{ 3-5(4) 3,4(4) 3,4(3) ‘ \

2 1 1,2(1) 1,2(1) : 1

3 202) 1,2(2) 2-4(3) : .

4 3-5(4) 2-4(3) 1,2(2) 2,3(2) : -

5 2 2,3(2) 1,2(2) : : :

6 2-4(3) 2-4(3) 4-8(6) - : :

7 4-6(4) 1 1 ; : ;

8 1-3(2) 3,4(3) 3-5(4) : 3 :

9 1 1 3-5(4) 6-13(6) : ;

10 1,2(1) 1 1 : : :

11 1 1 1-3(1) ; ; :

12 : : : ; ‘ :

13 . - . : : :

14 1 1 1 1 : -

4 Based on specimens from the Yemen Arab Republic.

b Range (mode).

d Alveolus only.

® Frequently absent, often present on one side only.

196 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Table 11. Number of branches for pupal setae of Culex mattinglyi Knight.

Seta Cephalothorax

No. CT

0 us

1 3/-5

2 a3

3 3/-

4 3/-

5 7/7

6 3/2

7 2/-

8 2/2

9 2/2

10 8/10

11 4/4

12 2/2

13 3/2

14 -

Seta

No. V

0 1/1

1 4/4

2 1/1

3 2/1

4 2/3

5 1/1

6 3/4

7 4/2

8 3/2

9 1/1

10 1/2

11 1/1

te -

13 -

14 1/1

1/1

Abdominal Segments

- 1/1

9/6 ¢ 17/14

1/1 1/1

2/2 2/2

5/4 3/2

6/6 2/2

7/3 5/5

3/4 1/1

3/2 1/1

- oy 10/9

Abdominal Segments

Vil Vill

1/1 1/1

3/2 “

1/1 “

1/2 .

1/1 1/1

2/4 ‘

-/9 q

4 :

fe x"

-/2 /12

1/1 ‘

1/1 .

/1 1/1

1/1

14/10

4 Based on a single specimen in the type series from the Yemen Arab Republic.

5 Branches of setae on right/left sides.

d Alveolus only.

Harbach: Subgenus Culex in Southwestem Asia

Table 12. Number of branches for pupal setae of Culex simpsoni Theobald.4

Seta

No.

OON OAR WBN =|“ ©

COON OAARWN $C

Cephalothorax

os & to or =

i aWmPNWWODWuAO um

(ee)

SOGNOW

oe,

‘

~~“

erate

Abdominal Segments

12-19(16)

Vil

Onnhy

=AOMNADwWWARUPU

;

3-7(6)

5-7(6)

4 Based on specimens from Ethiopia and Tanzania.

b Range (mode).

Vill

12-15(12)

197

8-12(12)

198 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Table 13. Number of branches for pupal setae of Culex sinaiticus Kirkpatrick.2

seta Cephalothorax Abdominal Segments

No. CT | it IV

0 - : 1 1 1

1 3,4(4)> -¢ 15-22(18) 8-16(11) 7-12(7)

2 4,5(5) 1,2(1) 1,2(1) 1 1

3 2,4(2) 1212) 1,2(2) 2 5-7(5)

4 2-4(4) 4-6(6) 4-7(6) 4-6(5) 1-3(2)

5 3-6(5) 2-5(3) 5,6(5) 5-8(6) 5,6(6)

6 3-5(4) 1 1 3-5(4) 3-6(5)

7 2 2 1,2(2) 4-7(6) 2-5(3)

8 4-6(5) : - 2-6(4) 2-5(3)

9 2,3(3) 1-3(2) 1 1 1

10 4-7(5) -d : 2 1,2(2)

41 2 1 : 1,2(1) 1,2(1)

12 3,4(4) - - : -

13 - . . . -

14 - : - 1

Seta Abdominal Segments Paddle .

No V VI vil Vill x P

0 1 1 1 1 : :

1 5-10(7) 5-7(5) 3-5(4) : 1 1

2 1 1 1,2(1) : : 1

3 1-3(2) 2 3,4(4) : : :

4 4-6(5) 3,4(3) 2,3(2) 2,3(2) : .

5 2,3(2) 2 2 : . :

6 3-5(5) 4-6(5) 5-10(6) . . -

7 4-6(6) 1 1 : : :

8 2-4(4) 3-5(3) 3-5(4 : : :

g 1 1 3-5(4) 6-10(7) : :

10 1 1 1 - - -

11 1,2(1) 1-3(1) 1-3(2) : : .

12 . - . - -

13 - - - ~ -

14 1 1 1 . -

&@ Based on specimens from Egypt (Sinai) and Israel.

b Range (mode).

d Alveolus only.

Harbach: Subgenus Culex in Southwestern Asia 199

Table 14. Number of branches for pupal setae of Culex duttoni Theobald.

Seta Cephalothorax Abdominal Segments

No. CT | ! Il IV

0 - - 1 1 1

1 2,3(2) -¢ 17-26(18) 4-6(4) 2-6(4)

2 2,3(2) 1,2(1) 1,2(1) 1 1

3 2-4(3) 2 1,2(2) 2 4-8(6)

4 2-5(3) 6-11(9) 4-7(6) 3-6(4) 1-3(2)

5 2-6(4) 3-6(4) 3-5(3) 4,5(4) 1-4(4)

6 2-4(3) 1,2(1) 1,2(1) 1,2(1) 1,2(1)

7 2 2-5(3) 3,4(4) 3-8(4) 3-5(4)

8 2,3(2) : . 2-5(4) 1-4(2)

9 2,3(2) 1,2(2) 1 1 1

10 5-13(8) : ; 2,3(2) 2

11 2-5(3) 2-5(2) ‘ 1-3(1) 1,2(1)

12 2-5(3) , : : :

13 . - - - :

14 . A {

Seta Abdominal Segments Paddle

No. V VI Vil Vill XI P

0 1 1 1 1 ‘ :

1 1-4(2) 2,3(2) 2-4(2) . 1 1,2(1)

2 1 1 1 ; : 0,2(0)

3 1,2(2) 1-3(2) 2 : : :

4 4-6(5) 3-5(4) 2 2-4(2) : 5

5 1-3(1) 1 1-3(2) . : :

6 1-3(1) 1-3(3) 6-16(11) - - -

7 4-9(6) 1 1 - - -

8 2-4(4) 2-5(4) 4-9(4) ; ‘ ‘

9 1 1 3-6(3) 7-11(10) - -

10 1 1 1,2(1) ? . :

11 1-3(1) 1-3(1) 2,3(2) ; : .

12 . . . : . .

1g - : - - -

£&

—_ 1

oth

—_.

4 Based on specimens from Uganda and Cameroon.

b Range (mode).

d Alveolus only.

200 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Table 15. Number of branches for pupal setae of Culex sitiens Wiedemann.@

Seta Cephalothorax

No. CT | ll

0 - - 1

{ 2,3(2)> -¢ 4-10(6)

2 3,4(4) 1 1

3 1-3(2) 2,3(2) 1,2(2)

4 2-4(3) 5-8(7) 4-6(4)

5 3,4(3) 4-7(6) 4,5(4)

6 2,3(2) 1 1

Pe 2 3 2,3(2)

8 2 - -

9 2,3(2) 1-3(2) 1

10 5-9(7) -d :

11 2-4(3) 1,2(1) -

12 2-5(3) - -

13 - - -

14 - - :

Seta Abdominal Segments

No V VI Vil Vill

0 1,2(1) 1,2(1) 1 1

1 3,4(4) 2,3(2) 2 ~

2 1 1 1 .

3 2 1,2(2) 2-4(2) :

4 Oo) 2-4(3) 1,2(1) 2,3(2)

5 1,2(2) 2 1,2(2) .

6 4212) 1,2(2) 6-11(8) -

7 5-7(6) 1 1 -

8 2-4(3) 2-4(3) 2-4(4) ’

9 1 1 3-5(4) 9-14(12)

10 1,2(1) 1,2(1) 1 :

11 Tj20T) Tart)”. 2,3(2) -

12 : : ; *

13 : - - -

14 1 1 1

4 Based on specimens from the Yemen Arab Republic.

b Range (mode).

d Alveolus only.

Abdominal Segments

Co =k

4 «

O1 PO

oh,

OE OE

BS a ee

r-aOaAOND WW

Harbach: Subgenus Culex in Southwestern Asia 201

Table 16. Number of branches for pupal setae of Culex poicilipes (Theobald).4

Seta Cephalothorax Abdominal Segments

No. og | l It IV

0 : ‘ 1 1 1

1 3-5(4)9 -¢ 8-15(11) 7-14(9) 5-8(8)

2 3-5(4) 1-3(1) 1,2(1) 1 1

3 2-5(3) 2,3(2) 2,3(2) 2 5-11(6)

4 2-4(3) 8-17(14) 5-10(7) 6,7(6) 3-6(4)

5 4-6(6) 4-13(4) 7-14(8) 8-20(10) 3-5(3)

6 2-4(3) 1 1 3-5(3) 3-6(4)

7 2,3(2) 2,3(2) 1-3(2) 6-10(6) 4-8(7)

5 2-4(4) , ; 5-8(5) 4-7(5)

9 2-4(2) 1,2(1) 1 1 1

10 7-16(9) -d ‘ 2-4(3) 2,3(3)

11 2,3(2) 1,2(2) ’ 1,2(2) 1,2(1)

12 3-6(4) . : ‘

13 i " : : ,

14 j : ; 1 1

seta Abdominal Segments Paddle

No “coe VI vil vill xl pP

0 1 1 1 1-mé ‘ ‘

1 5,6(5) 3,4(3) 2,3(3) : 1 1

2 1,2(1) 1,2(1) 1,2(1) , ‘ 1

3 2-5(3) 2,3(3) 3-5(4) , ; :

4 5-10(6) 3-7(5) 2-4(3) 2-A(3) : ‘

5 2 2 2-4(4) ‘ : 4

6 3-6(4) 3-6(5) 6-17(8) : ‘ :

7 6-13(7) 1,2(1) 1 ‘ : ‘

8 3-7(5) 3-5(4) 4-8(7) : ; ‘

9 1 1 2-5(3) 5-10(8) . .

10 1 1 1 5 , f

11 1,2(2) 1-3(3) 2-4(3) : . .

12 : ‘ ‘ : . .

13 _ . . . . .

14 1 1 1 1-4(1) . 4

4 Based on specimens from Egypt.

b Range (mode).

d Alveolus only.

202 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Table 17. Number of branches for pupal setae of Culex mimeticus Noé.4

Seta Cephalothorax Abdominal Segments

No. CT | ll Ill

0 - - 1 1

1 2,3(2)° -c 1-4(3) 3-6(4)

2 2-4(3) 1 1 1

3 2-4(2) 2 1,2(2) 2

4 1-3(2) 5-9(5) 3-6(4) 5-10(6)

5 2,3(2) 2-7(3) 3-5(3) 2-5(3)

6 2-4(2) 1 1 1,2(1)

7 2 1-3(2) 1-3(2) 5-9(6)

8 2,3(2) 1 . 2,3(3)

g 2-4(2) : 1 1

10 4-6(5) -d a/1e 1,2(2)

11 2,3(2) 1 : 1

12 2-4(2) - - :

13 : ; ~ -

14 : : : 1

Seta Abdominal Segments

No. V VI Vil Vill XI

0 1 1 1 -

1 1-3(2) 1,2(1) 1-3(2) : 1

2 1 1 1 - -

3 1/201) 1 ,2(2) 1-3(2) - -

4 2-5(4) 2-4(2) 1-3(1) 1,201) -

5 2 2 1 : -

6 1-3(1) 1,204) 3-7(3) . -

7 2-8(3) 1 1 . :

8 1-3(2) 2,3(2) 2-4(2) : :

9 1 1 2-4(2) 4-9(6) -

10 1 1 1 - .

si 1 1 1,2(2) - -

12 - . . - .

13 - - - -

14 1 1 1 .

&@ Based on specimens from Israel.

b Range (mode).

d Alveolus only.

Harbach: Subgenus Culex in Southwestern Asia 203

Table 18. Number of branches for pupal setae of Culex bitaeniorhynchus Giles.

Seta Cephalothorax Abdominal Segments

No. CT | l il IV

0 - - 1 1

1 4-3(2)° -¢ 1-4(2) 6-10(7) 2-6(4)

2 2-4(2) 1 1 1

3 1-4(2) 1 1 1 3-7(4)

4 1-3(2) 4-8(6) 2-6(4) 4-6(6) 2,3(3)

5 2,3(2) 3-9(3) 2-5(4) 3-6(5) 2,3(3)

6 1-4(2) 1,2(1) 1,2(1) 1 1

7 1,2(2) 1-3(2) 1,2(2) 3-6(5) 1-5(3)

8 1,2(2) . : 2-5(4) 2-4(3)

9 1,2(2) 1-3(2) 1 1 1,2(1)

10 5-10(7) -d : 2,3(2) 2

11 1,2(1) 1 : 1 1,2(1)

12 1-3(2) : - : :

13 - - - . :

bias P ; : 1 1

Seta Abdominal Segments Paddle

No. V VI Vil Vill x Pp

0 1 1 1 1 : :

1 2,3(2) 120) 1,2(2) : 1 1

2 1 1 1 : : 1

3 1-3(1) 1,2(1) 1,2(1) - : :

4 2-5(4) 2,3(3) 1-3(2) 2 : ;

5 2,3(2) 2,3(2) 1,2(1) : ‘

6 1 1 3-8(5) . - -

7 4-7(5) 1,2(1) 1 : : :

8 1-3(2) 2-4(3) 2-4(3) - . .

9 1 1 3-5(3) 6-10(6) - -

10 1 1 1,2(1) : . .

11 1 1,2(1) 1-3(2) ; . :

12 - . : . : .

13 - . - - - -

14 1 1 1 1 . :

4 Based on specimens from Pakistan and the Yemen Arab Republic.

b Range (mode).

d Alveolus only.

204

Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Table 19. Number of branches for pupal setae of Culex tritaeniorhynchus Giles.

Seta Cephalothorax Abdominal Segments

No. cr | l T IV

0 : ; 1 1 1

1 3,4(3)° -¢ 20-36(28) 10-15(11) 7-12(10)

2 3-5(3) 1 1 1 1

3 2,3(3) 2 2 2 4-7(6)

4 1,2(2) 7-10(9) 4-7(5) 4-7(6) 1-4(3)

5 4-6(5) 4-7(6) 5-9(6) 6-8(8) 4-7(5)

6 3-6(4) 1 1 4-7(6) 3-6(5)

7 1,2(2) 2-5(4) 2-4(2) 4-8(5) 2-5(3)

8 6,7(7) : : 2-6(5) 3,4(3)

9 2,3(2) 1-3(1) 1 1 1

10 6-12(9) -d : 1,2(2) 2

& 2 1 ’ 1,2(1) 1

12 3-5(4) : : ' :

13 : : _ : :

14 : : : 1 1

Seta Abdominal Segments Paddle

No. V vl vil Vill x Pp

0 1 1 1 1 :

1 7-10(8) 7-9(7) 5,6(5) : 1 1

2 1 1 1 ‘ 1

3 2,3(2) 2,3(2) 4-6(5) : :

4 5-7(6) 4-6(5) 2,3(3) 3,4(3) .

5 2 2 1,2(2) ’ :

6 6,7(6) 6-8(7) 9-13(12) . ,

7 5,6(6) 1 1 ‘ : ‘

8 2-5(4) 2-5(3) 4-6(4) : : *

9 1 1 3-5(5) 4-8(5) : .

10 1 1 1 : : '

a, 1,2(1) 1,2(1) 2-5(2) ‘ j :

12 : . ! : " .

13 : : : : . .

14 1 1 1 1 : .

&@ Based on specimens from Pakistan.

b Range (mode).

d Alveolus only.

Harbach: Subgenus Culex in Southwestern Asia

Table 20. Number of branches for pupal setae of Culex pseudovishnui Colless.@

Seta Cephalothorax Abdominal Segments

No. CT | l ll

0 - - 1 1

1 2-4(3) -c 9-15(12) 10-16(10)

2 3-5(4) 1 1,2(1) 1

3 3 2,3(2) 2,3(2) 2

4 1-3(2) 6-12(6) 4-8(7) 3-8(6)

5 3-5(4) 3-7(4) 6-8(7) 5-9(6)

6 2-5(3) 1 1 3-5(4)

7 2 2-4(2) 2 3-9(7)

8 3-5(5) - - 3-6(4)

9 2,3(2) 2 1 1

10 3-10(9) -d : 2

11 2 1,2(1) . 1

12 2-5(3) . i ;

Ve - - ~ -

14 : : ‘ 1

Seta Abdominal Segments

No. V VI VII Vill Xx

0 1 1 1 1 -

1 4-8(7) 5-8(6) 3-5(4) - 1

2 1 1 1 - -

3 2,3(2) 2,3(2) 3-5(4) : ,

4 5-7(5) 4,5(4) 2,3(2) 2-4(2) 4

5 2-4(2) 2,3(2) 1,2(2) : ‘

6 5,6(5) 5-7(5) 5-11(8) , 4

7 5-8(7) 1,2(1) 1 - :

8 3-5(4) 4,5(4) 2-5(3) - -

9 1 1 3-6(4) 6-9(8) -

10 1 1 1 ; -

11 1,2(1) 1-3(1) 2-4(3) - >

12 , : ‘ ‘ ‘

13 . . : " .

14 1 1 1 1 -

&@ Based on specimens from Pakistan and India.

b Range (mode).

d Alveolus only.

7-10(9)

205

206 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Table 21. Number of branches for fourth-instar larval setae of Culex pipiens

Linnaeus.@

Seta Head Thorax Abdominal Segments

No C P M a ll Il

0 1 9-26(17) - : : 1 1

1 1 1 1 2-5(3) 1-5(5) 1-4(2) 1-4(2)

2 0-1(0) 1 2-5(3) 1-3(2) 1 1 1

3 1 1 1 2-5(4) 1,2(1) 1201) 1,2(1)

4 1 2,3(2) 1,2(2) 2-5(4) 7-12(8) 3-7(4) 1-4(2)

5 4-7(5) 1 1 1 2-7(5) 1-3(2) 1-4(2)

6 3-6(4) 1 1 3,4(3) 2-4(3) 2,3(2)

7 8-13(10) 2,3(2) 1 6-10(10) 1-32) 3-6(4) 4-9(6)

8 2-4(2) 2,3(2) 5-9(5) 8-18(14) - 1 1

9 3-7(5) 1 4-8(5) 4-9(6) 1-5(2) 1 1

10 2,3(2) 1 1 1 1-3(1) 1 1

11 2,3(2) 3-8(5) 1-4(2) 1-3(2) 3-10(4) 2-5(3) 1-3(2)

12 2,3(2) 1 1 1 1-3(2) 1-3(2) 1-3(1)

13 2-4(3) - 14-27(23) 3-6(5) 1-3(2) 7-23(15) 2-5(3)

14 1.214) 1,2(1) | 15-26(20) - . - 1

15 3-5(3) - - - - - -

Seta Abdominal Segments

No. IV V VI Vil Vill X

0 1 1 1 1

1 2-4(2) 1-3(2) 2,3(2) 2-5(3)

2 1 1 1,2(1) 1

3 1,2(1) 1 1 1,2(1)

4 1,201) 2-5(3) 1-3(2) 1

5 1-3(2) 1-4(2) 1-4(3 1-4(2)

6 2 2 2 10-20(10)

us 5-10(7) 4-8(6) 1 ,2(1) 1

8 1 1,2(1) 2-5(3) 2-6(4)

9 1 1 1 2-8(3)

10 1 1 1 1

11 1-4(3) 1-4(3) 2-4(3) 2-4(2) - -

12 1,2(1) 1 1 1 - .

13 2-4(3) 2-4(3) 22-43(28) 2-4(3) - -

14 1 1 1 1 1 .

15 - - - - - -

42 Based primarily on specimens from Egypt and Israel.

b Range (mode) obtained from counts made on at least 40 setae.

Harbach: Subgenus Culex in Southwestern Asia 207

Table 22. Number of branches for fourth-instar larval setae of Culex quinquefasciatus

Say.?

Seta Head Thorax Abdominal Segments

No. C P M T | l ll

0 1 11-20(15)° . - 3 1 1

1 1 1 1,2(1) 2-5(2) 2-6(2) 1-4(2) 1

2 1 1 1,2(1) 1,2(1) 1 1 1

3 1 1 1,2(2) 2-4(3) 1,2(1) 1 1

4 1 2 1-3(2) 3-6(5) 6-10(8) —-2-5(4) 1-4(1)

5 4-7(5) 1 1 1 5-10(6) 1-3(2) 1,2(2)

6 3-5(5) 1 1 1 3,4(3) 3,4(3) 2

7 7-10(8) 2 1 6-8(7) 2 2-5(4) 4-7(6)

8 2-4(3) 2 4-6(5) 9-16(11) ; 1 1

) 2-7(4) 1 4-6(5) 4-7(5) 120g) 1 1

10 1-3(3) 1 1 1 1 1 1

11 2-4(3) 4-7(5) 2-4(2) 1-3(3) 3-7(6) 2-4(3) 1-3(2)

12 1-4(2) 1 1 1 1-3(2) 1,2(2) 1

13 2-5(4) : 16-36(22) 3-7(6) 1,201) 10-2607) 2-413)

14 1,2(1) 1 15-33(19) - - . 1

15 2-4(3) - - - - - -

Seta Abdominal Segments

No IV V VI VII Vill X

0 1 1 1 1

1 1-20 1,2(2) 2 3-5(3)

2 1 1 1 1

3 Leu 1 1 1,2(1)

4 1.2¢%) 2-4(3) 1,2(2) 1

5 1,2(2) 1-3(2) 2-6(3) 2-5(4)

6 2 2 2 8-15(12)

7 5-10(6) 5-7(6) 1 1

8 1,219) 1,2(1) 2-5(3) 2-5(4)

9 1 1 1 3-6(4)

10 1 1 1 1

11 2,3(2) 2-4(2) 2-4(2) 2,3(2) ; :

12 1,2(1) 1 1 1 ‘ .

13 1-4(3) 2-4(3) 22-46(31) 2-4(3) - -

14 1 1 1 1 1 -

15 - ; - - - .

2 Based on specimens from the Yemen Arab Republic and India.

b Range (mode) obtained from counts made on at least 10 setae.

208

Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Table 23. Number of branches for fourth-instar larval setae of Culex vagans

Wiedemann.?

QOS

O10) —

weer ee ee

SS3seS Se

owed,

OD fe ak ak. ee A ee

= 1

WD —- —

' a Ce Re cg

NO PO

cond,

i he = ee Se oe CR oe Ss a

“~

—s

: no 19

qo as

\ -F- WD ANNNAAAHD

Thorax Abdominal Segments

M T | ll IN

- - - 1 1

1 2-5(3) 3-10(6) 2-4(3) 1-3(2)

1-3(2) 1,2(1) 1,2(1) 1 1

1 2-6(4) Feat) 1 1

1-3(2) 3-6(4) 8-12(9) 3,4(4) 1-3(2)

1 1 4-7(6) 1-3(2) 1-3(2)

1 1 2-4(3) 3-5(4) 2,3(2)

1 6-9(6) 2,3(2) 3-6(4) 4-8(5)

4-8(6) 11-17(11) : 1 :

4-7(5) 4-8(6) 1-3(2) 1 1

1 1 1 1 1

2,3(2) 23(2) 4-9(6) 2,3(2) 1-3(2)

1 1 1-3(1) 1 ,2(2) 1,29)

10-31(24) 3-7(4) 1-3(2) 10-25(18) 2-6(4)

15-30(24) - - 2 1

Abdominal Segments

VI Vil Vill X

1 1

1271) 3-5(3)

1 1

Tate) 1

2-4(3) 2-4(3)

2 ,a(2) 10-16(10)

Vet) 1,2(1)

1-4(3) 3-6(6)

1,2(%) 3-6(5)

1 1

2-4(2) 1,3{2) - -

1 1 - -

27-36(29) 3-5(4) - -

1 1 1 -

&@ Based on specimens from Pakistan.

b Range (mode) obtained from counts made on at least 10 setae.

Harbach: Subgenus Culex in Southwestem Asia 209

Table 24. Number of branches for fourth-instar larval setae of Culex torrentium

Martini.

Seta Head Thorax Abdominal Segments

No. C P M HS ll il

0 1 9-13(11)° : ‘ ’ 1 1

1 1 1 2-4(3) 3-7(4) 2-6(5) 1-3(2) 4,5(4)

2 ; 1 1-4(2) 2 1 1 1

3 1 1 1 3,4(4) 1-3(2) 1,2(1) 1,2(1)

4 1,2(1) 2 2 ,3(2) 2-4(3) 7-12(9) 3-5(4) 1

5 4-7(5) 1 1 1 4-7(5) 1-4(1) 1-3(1)

6 3-5(4) 1 1,461) 1 3,4(3) 3-5(4) 2

7 8-10(8) 2,3(2) 1 4-8(6) 2,3(2) 3-5(4) 3-7(4)

8 2-4(2) 2 3-7(5) 8-15(12) - 1 1

g 2-5(3) | 4-6(6) 5-7(6) 1-3(2) 1 1,2(1)

10 2,3(2) 1 1 1 1 1 1

{4 2-4(3) 4-8(6) 1-3(2) 1,2(2) 4-6(4) 2-4(3) 2,3(2)

12 2:3(3) 1 1 1 1,2(2) aiate) 1,2(1)

13 2,3(3) . 13-20(18) 3-8(5) 1,2(1) 12-19(17) 4-6(4)

14 1-3(1) 1 12-25(19) . - . 1

13 3,4(4) - - - - - -

Seta Abdominal Segments

No. IV V VI Vil Vill X

0 1 1 1 1

1 3-6(4) 3-5(4) 2-5(3) 4-6(4)

2 1 1 1 1

3 1,2(1) 1 1 2

4 1 3-5(3) 1,2(2) 1,2(1)

5 1-4(2) 1-4(2) 2-5(3) 1-4(3)

6 2 2 2 8-13(10)

7 4-7(6) 4-7(5) 1 1

8 1 1,2(1) 2-4(2) 3-8(6)

9 1,29} 1,2(1) 1,2(1) 2-6(3)

10 1 1 1 1

11 1,2(2) 2 1-3(2) 1,2(1) . :

12 1 1 1 1 - ~

13 4-6(4) 3-5(4) 17-34(28) 3,4(3) . -

1 1 1 1 ~

4 Based on specimens from Sweden.

b Range (mode) obtained from counts made on at least 10 setae.

210 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Table 25. Number of branches for fourth-instar larval setae of Culex decens

Theobald.@

Seta Head Thorax Abdominal Segments

No. C P M if | ll Il!

0 1 13-19(15)> : : : 1 1

1 1 1 1,2(1) 1,2(1) 1-3(1) 1 1-3(2)

2 : 1 2-4(3) 1,2(2) 1,2(1) 1,2(1) 1

3 1 1 1 3-5(3) 2 1,2(2) 1,2(1)

4 1 2,3(2) 1 ,2(2) 4-7(6) 6-11 (9) 4-6(5) 1,200)

5 2,a(2) 1 1 1,2(1) 7-9(8) 2,3(2) 1-4(2)

6 2 1 1 1 3 3 1

7 5-7(6) 3 1 5-7(6) 2 4-7(5) 4-7(5)

8 4-6(5) 2 4,5(4) 10-18(11) ~ 1,2(2) 1,2(2)

9 4-8(7) 1 3,4(4) 5,6(6) 3,4(4) 1 1

10 1-3(2) 1 1 1 1 1 1

14 tie) 5-7(6) 2-4(3) 2-4(3) 4-8(5) 2-5(3) 2 joe)

12 2-4(3) 1 1 Tat) 1,2(2) 2-4(3) 1,2(1)

12 3-5(4) - 18-28(24) 5-7(6) 1-3(2) 14-23(17) 2-5(4)

14 1-3(2) 1 16-24(21) : . : 1

1 3-6(4) - - - “ - -

Seta Abdominal Segments

No. IV V VI Vil Vill X

0 1 1 1 1

1 1 1 2 B(3) 2-5(4)

2 1 1 1 1

3 2 1 1 2,3(2)

4 1 4-6(5) 2,3(3) 1

5 2-4(2) 2 oi2) 2-4(3) 2-4(2)

6 1-3(2) 1 1 11-21(15)

a 5-7(6) 4-8(6) 1 1

8 2 2 2-4(4) 2-4(2)

9 1 1 1 1 ,2(2)

10 1 1 1 1

11 2,3(2) 2,3(2) 2 3(3) 2-4(3) -

12 1 1 1 1 : -

13 3-6(4) 3-5(4) 32-56(42) 2-4(3) - -

14 1 1 1 1 1 .

15 - - - - - .

4 Based on specimens from the Yemen Arab Republic (primarily) and Senegal.

b Range (mode) obtained from counts made on at least 10 setae.

Harbach: Subgenus Culex in Southwestem Asia 211

Table 26. Number of branches for fourth-instar larval setae of Culex antennatus

(Becker) .4

Seta Head Thorax Abdominal Segments

No C P M T ll IN

0 1 17-29(23)> : : : 1 1

1 1 1 1-3(2) 2-4(2) 1-4(4) 1-3(2) 3,4(4)

2 : 1 4-8(5) 2,3(2) 1 1 1

3 1 1 1 4-7(5) 2-5(3) 2-4(3) 2,3(2)

4 1,2(1) 2 1-3(2) 3-8(4) 8-14(11) 5-9(6) 2-4(2)

5 2,312) 1 1 1.211) 4-9(5) 2,3(3) 2-5(3)

6 2,312) 1 1 1 3 3,4(3) 1-3(2)

7 6-8(8) 3 1 5-8(7) 2 5-9(6) 7-10(9)

8 3-5(5) 2 4,5(5) 9-16(12) - 1,2(1) 1,2(1)

9 4-7(5) 1 4,5(4) 4-6(5) 3,4(3) 1 1

10 3,4(3) 1,2(1) 1 1 1 1 1

11 2,3(2) 6-9(6) 2-4(2) 2-5(3) 4-6(5) 2-4(3) 2,3(2)

12 3-5(4) 1 1 1-3(2) 1-4(2) 2,3(2) 1,202)

13 2-4(3) - 26-43(36) 7-14(8) 2,3(2) 15-30(20) 4-7(5)

14 1,2(1) 1 14-27(21) - . - 1

15 5-8(7) - - - - - -

Seta Abdominal Segments

No IV V VI Vil Vill X

0 1 1 1 1

1 3,4(3) 2-4(3) 2,3(3) 3-6(5)

2 1 1 1 1

3 2-4(3) 1,2(1) 1 3-5(4)

4 1-3(2) 5-8(6) 3,4(3) 1

5 2,3(2) 1-3(2) 2,3(2) 2,3(2)

6 3 2,3(2) 1 12-20(15)

7 6-12(9) 8-13(9) 2,3(3) 1,2(1)

8 1,2(2) 2 1-4(2) 4-7(5)

9 1 1 1 2-4(2)

10 1 1 1 1

11 2,3(2) 1,2(2) 2,3(3) 2-4(3)

12 2 1 1 1

13 3-6(5) 4-6(5) 31-47(39) 3-6(4) : -

14 1 1 1 1 1 -

15 - - . - - -

4 Based on specimens from Egypt.

b Range (mode) obtained from counts made on at least 10 setae.

212 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Table 27. Number of branches for fourth-instar larval setae of Culex univittatus

Theobald.4

Seta Head Thorax Abdominal Segments

No. C P M i | ll IN

0 1 8-26(19)° . . 1 1

1 1 1 1,2(1) 1-3(2) 2-5(3) 1-3(2) 2-4(3)

2 0,1(0) 1 2-7(2) 2-4(3) 1,2(1) 1 1

3 1 1 1 2-6(5) 2-4(2) 1-3(2) 1-3(2)

4 1 e 1-4(3) 2-7(4) 5-15(12) 3-7(5) 1-4(3)

5 2-4(3) 1 1 1-3(1) 4-12(5) 1-3(3) 1-4(2)

6 2,3(3) 1 1 1 3,4(3) 3 2,3(2)

7 6-8(6) Pope Fo 1 5-7(6) 2,3(2) 2-7(5) 4-9(8)

8 2-6(5) 2,3(2) 5 ,6(5) 7-16(10) - 1,2(1) 1,2(2)

9 3-9(7) 1 3-6(4) 5,6(5) 2-4(2) 1,2(1) 1

10 2-4(3) 1,2(1) 1 1 1 1 1,2(2)

11 2,3(2) 6-10(7) 1-4(2) 2-5(3) 2-7(4) 3,4(3) 1-3(2)

12 2-5(4) 1 1 1-5(2) 1-3(2) 1-3(2) 1-3(2)

13 3-5(4) - 20-41(29) 5-10(7) 1-3(2) 9-22(20) 3,4(4)

14 1 ,2(2) 1 10-28(21) - - - 1

15 3-8(6) - - - . - -

Seta Abdominal Segments

No. IV V VI Vil Vill X

0 1 1 1 1

1 2-4(3) 2-4(3) 2-4(3) 3-7(5)

2 1 1 1 1

3 2-4(3) 1,242) 1-3(2) 3-6(4)

4 1,2(1) 3-9(6) 1-5(4) 1

5 1-3(2) 1-3(2) 1-4(3) 1-5(4)

6 2,3(2) 2 1,2(1) 7-23(16)

7 3-11(8) 5-12(8) 1-3(3) 1

8 1-3(2) 53(2) 2-4(4) 3-7(3)

9 1 1 1 2-5(4)

10 1,2(2) 1 1 1

11 1-3(2) 1-3(2) 2,3(2) 2-4(3)

12 1,2(1) 1 1 1

13 3-5(4) 3-5(4) 16-42(30) 3-6(4) - -

14 1 1 1 1 1 -

16 - - - - - -

4 Based on specimens from Zimbabwe (type locality) and South Africa.

b Range (mode) obtained from counts made on at least 10 setae.

Harbach: Subgenus Culex in Southwestern Asia 213

Table 28. Number of branches for fourth-instar larval setae of Culex perexiguus

Theobald.@

Seta Head Thorax Abdominal Segments

No. C P M i A | l IN

0 1 10-23(19)® : : , 1 1

1 1 1 1,2(1) 2,3(2) 2-4(2) 1,2(2) 3,4(3)

2 - 1 3-5(4) 2,3(2) 1 1 1

3 1 1 1,2(1) 3-5(4) 2-4(2) 1,2(1) 1,2(2)

4 1 2 1-3(2) 4-6(4) 6-14(8) 4-8(5) 1-3(2)

5 2:3(3) 1 1 1 4-9(5) 3,4(3) 2,3(3)

6 2,3(2) 1 1 1 2,363) 3 2

7 4-8(6) 3,4(3) 1 5-8(8) 1,2(2) 3-6(5) 5-8(6)

8 3-5(4) 2 3-6(6) 6-18(9) - 1,2(1) 1,2(2)

9 5-9(5) 1 3-5(3) 4-6(5) 2-4(3) 1 1

10 2-4(3) 1 1,2(1) 1 1 1 1,2(2)

11 1-3(2) 4-8(5) 1-4(2) 2-4(2) 3-6(4) 2,3(2) 1,2(2)

12 2-5(3) 1 1 1-3(1) 2 2,3(3) 1-3(2)

13 2-4(2) - 21-42(29) 5-8(6) 2,3(2) 11-26(15) °3-5(3)

14 2 1 15-28(21) . . , 1

15 3-7(6) - - - - -

Seta Abdominal Segments

No IV V VI Vil Vill X

0 1 1 1 1 1 .

1 2-4(3) 2-4(3) 3,4(3) 3-5(4) 5-8(6) 2,3(2)

2 1 1 1 1 1 2

3 1-3(2) 1,204} 1,2(2) 2-4(3) 5-8(8) 1

4 1 4-6(6) 2-4(3) 1 1 4-9(7)

5 2 ,3(2) 2,3(2) 2,3(2) 2-4(3) 3-5(4) -

6 2 ,3(2) 2 1,2(1) 9-18(11) - -

7 4-8(7) 5-10(5) 1,2(2) 1

8 1-3(2) 2 2-4(3) 4-6(4)

9 1 1 1 1-3(2)

10 aby 1 1 1

11 1,207) 2 1-3(2) 2,3(2)

12 1,2(1) 1 1 1 : -

13 3-5(4) 3-5(4) 26-44(29) 2-5(3) - -

14 1 1 1 1 1 .

15 - - ~ - - -

& Based on specimens from Egypt.

b Range (mode) obtained from counts made on at least 10 setae.

214

Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Table 29. Number of branches for fourth-instar larval setae of Culex theileri Theobald.4

Seta

No.

OONDOAWNM HO

No —

aA s-O=>A WIM

ONROHANARON

2 eelkele2 {aces

jae

an s

Beh

awh,

<a O

— &

= PD =

G) —& [\) =

[

1m OTM BND =

13-25(16)°

1 —™ »; me COM @AW DB SB | PS SA |

“_—_

ol —

—

Ol —

<_<

~~”

Thorax

M T |

1-4(1) 2-4(2) 4-6(4)

3-9(3) 1-3(2) 1

1 3-8(5) 2-5(3)

232) 3-6(4) 10-18(1

1 1 3-11(5

1 1 3

1 5-7(6) 2

4-6(5) 12-19(15) -

4-6(5) 4-6(5) 1-5(3)

1 1 1

3-5(3) 2-6(3) 3-6(5)

1 1,2(2) 1-3(3)

30-48(32) 6-10(7) 2-5(2)

21-39(34) - .

Abdominal Segments

Vl VII

{ 1

2-4(3) 3-5(4)

1 1

1 3-8(5)

2-5(4) 1

3-5(4) 4-6(4)

3 14-22(18)

1 {

3-6(3) 5-9(5)

{ 2-7(4)

1 {

3,4(3) 2-4(3)

38-52(46)

Abdominal Segments

1 1

1-3(2) 1,2(2)

4-7(6) 1-4(3)

3-8(4) 3-6(5)

3,4(3) 3

4-9(7) 9-20(9)

1,2(2) 2,3(2)

1 1

1 1,2(1)

2-7(3) 2,3(2)

2,3(2) 1-3(2)

21-35(29) 3-5(5)

‘ 1

Vil Xx

@ Based on specimens from Egypt, Israel, and the Yemen Arab Republic.

b Range (mode) obtained from counts made on at least 10 setae.

C Siphon usually with 10 seta 1-S, not distinctly paired; sometimes with as few as 7 or as

many as 13.

Harbach: Subgenus Culex in Southwestern Asia 215

Table 30. Number of branches for fourth-instar larval setae of Culex laticinctus

Edwards.@

Seta Head Thorax Abdominal Segments

No. C P M ile | ll ll

0 1 11-19(14) . - - 1 1

1 1 1 1-4(3) 1-3(2) 1-3(2) 1,2(1) 1,2(1)

2 0,1 1 1-5(3) 2,3(2) 1 1 1

3 1 1 1 3-5(4) 2-7(3) 1-3(2) 1-3(2)

4 1 2 1,2(2) 3-6(5) 12-15(12) 3-8(5) 1,2(1)

5 4-6(5) 1 1 ,2(1) 1 4-6(4) 3-8(4) 3-6(5)

6 3,4(4) 1 1 1 3,4(3) 3,4(3) 1,2(1)

7 7-448) 3,4(3) 1 6-8(6) 2 2-5(4) 5-10(7)

8 3-6(5) 2 4-7(5) — 8-18(13) ! 1/2¢4) 1,2(1)

9 4,5(4) 1 4-6(5) 4-8(5) 2-4(2) 1 1

10 2-5(3) 1,2(1) 1 1 1 1 1

11 1,212) 2-6(4) 3-5(3) 2-4(4) 5-7(5) 2 a(2) 1-3(2)

t2 2-4(3) 1 1 2,3(2) 1-3(1) 2 ate) 1

13 2,3(2) - 20-30(24) 7-11(10) 1 13-29(22) 3-6(4)

14 2,3(2) 1 16-21(20). - . . 1

15 3-5(4) - - - - - -

Seta Abdominal Segments

No IV V VI Vil VII X

0 1 1 1 1

1 1 1 1,2(1) 2,02)

os 1 1 1 1

3 4,212) 1 1 2-4(2)

4 1 4-6(4) 1,2(2) 1

5 ti2(1) 1,2(1) 1;2(1) 1,2(2)

6 1,2(1) 1 1 10-15(10)

7 6-10(9) 5-8(7) 1 1

8 1,2(1) 1,2(2) 1-3(2) 4-6(4)

9 1 1 1 1-4(2)

10 1 1 1 1

11 1-3(2) 1-3(2) 1-3(2) 1-3(1)

12 1 1 1 1

13 3-6(4) 2-6(4) 25-39(31) 3-6(4)

14 1 1 1 1

15 - ; - - - -

4 Based on specimens from Israel and the Yemen Arab Republic.

b Range (mode) obtained from counts made on at least 10 setae.

216 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Table 31. Number of branches for fourth-instar larval setae of Culex mattinglyi Knight.2

Seta Head Thorax Abdominal Segments

No. C P M 7 | I IN

0 1 7-12(9)° ‘ : ; 1 1

1 1 1 1,2(1) 2-6(5) 1-3(2) 1,2(2)

2 1 1 2-4(4) 2-4(3) 1 1 1

3 1 1,2(2) 1 3-6(4) 2,3(3) 1 ,2(2) 1,2(1)

4 1 1 ,2(2) 3-5(3) 6-10(7) 3-6(5) 1

5 3-5(4) 1 1 1 3-5(3) 1,207) 1,2(1)

6 3,4(3) ‘ 1 1 2-5(2) 2-4(3) 2-5(3)

7 7-10(8) 2-4(2) 1 5-9(8) 2,3(2) 3-5(4) 5-9(7)

8 2-4(3) 2-4(2) 6,7(7) 9-14(12) - 1 1

9 4-7(4) 1 5-8(6) 6-10(8) 1,2(1) 1,2(1) 1

10 2-5(3) 1 1 1 1 1 1

a 2-4(3) 2-4(3) 2-4(3) 2,3(3) 2-7(3) 1-4(2) 1,2(1)

12 2-4(3) 1 1 1 1,2(1) 2,3(2) 1

13 2,3(2) - 12-22(17) 4-7(5) 1,2(1) 9-14(13) 2-6(4)

14 1 1 16-27(21) : . : 1

16 2-5(4) - - - - - -

Seta Abdominal Segments

No IV V VI Vil Vill X

0 1 1 1 1 1 -

1 1,2(1) 1,2(1) 1,2(1) 2 5-8(6) 2,3(3)

2 1 1 1 1 1 2,3(2)

3 1,2(1) 1 1 1,2(2) 12-14(12) 1

4 1 3-5(4) 2,3(2) 1 1 7-12(9)

5 1-4(2) 1-4(2) 1-3(2) 4-3(2) 3,4(4) “

6 2-5(2) 2,at2) 2;0(2) 6-9(8) - -

7 5-8(7) 6-8(7) 1,2(1) 1,2(1)

8 1 1 1-4(2) 2-4(3)

9 1 1 1 1-4(3)

10 1 1 1 1

11 1-3(2) 1-3(2) 1-4(2) 1-3(1)

12 1,2(1) 1 1 ei

13 4,5(5) 3-5(4) 16-29(21) 2-5(2)

14 1 1 1 1

15 - - . . *

@ Based on specimens from the Yemen Arab Republic.

b Range (mode) obtained from counts made on at least 10 setae.

Harbach: Subgenus Culex in Southwestem Asia 217

Table 32. Number of branches for fourth-instar larval setae of Culex simpsoni

Theobald.@

Seta Head Thorax Abdominal Segments

No. C P M T ll li

0 1 13-19(16)° : . : 1 1

1 1 1 1,2(1) 1,2(1) 1-6(3) 1 4,5(4)

2 , 1 2-4(3) 1-3(2) 1,2(1) 1 1

3 1 1 1 3,4(3) 2,3(2) 2 2

4 1 3,4(4) 2 2-5(3) 9-13(10) 3-84) 1-3(2)

5 1,2(1) 1 1 1 2-6(4) 1-3(2) 1-4(1)

6 1,2(2) 1 1 1 3 3 3,4(3)

7 7-10(9) 3,4(3) 1 6-8(6) 1,2(2) 4-6(5) 8-13(8)

8 3-5(4) 2,3(2) 4-6(6) 8-19(11) ” 1,2(1) 2,3(2)

9 4-8(6) 1 3-5 (5) 4-8(5) 2,3(3) 1 1

10 2,3(3) 1 1 1 1 1 1,2(1)

11 2 3-5(4) 1-3(2) 1-3(2) 3-6(4) 2-4(3) 2

12 4-7(5) 1 1 1,2(1) 1,2(1) 2 2

13 3,4(3) : 23-38(30) 5-7(5) 2 15-24(19) 3-5(4)

14 2-4(2) 1212). 14-220) . . - 1

15 4-6(5) - . - - - -

Seta Abdominal Segments

No. IV V VI Vil Vill X

0 1 1 1 1 1 -

1 4,5(4) 4,5(5) 3,4(4) 4-7(5) 5-7(5) 2.3(3)

2 1 1 1,2(1) 1 1 2-4(2)

3 2 1 1 2-5(3) 6-10(7) 1

4 1,2(1) 3-7(5) 1-3(2) 1 1 4-11(10)

5 1,2(1) 1-3(1) 1-3(2) 1-3(2) 3,4(4) -

6 3-5(3) 2-4(3) 3,4(3) 14-25(19) - -

7 6-10(9) 7-10(8) 1 1

8 2 2 3 3-8(5)

1 1 1 1-3(1)

10 1,2(1) 1 1 1

11 2,3(2) 2,3(2) 2,3(2) 1-3(2) : ‘

C2 1,2(1) 1 1 1 - .

13 3-6(4) 3-5(4) 29-45(33) 3-6(4) ~ -

14 1 1 1 1 1 .

15 - . : - - -

42 Based on specimens from Ethiopia, Kenya, and Tanzania.

b Range (mode) obtained from counts made on at least 10 setae.

218

Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Table 33. Number of branches for fourth-instar larval setae of Culex Sinaiticus

Kirkpatrick.

Seta Head Thorax Abdominal Segments

No. C P M i, | l ill

0 1 13-22(18)9 = - : : 1 1

1 1 1 1,2(1) 1 2-4(3) 1 3-5(3)

2 - 1 2-5(3) 2 1,2(1) 1 1

3 1 1 1 3-5(3) 2,3(2) 1 ,2(2) 1,2(2)

4 1 1-4(3) 1-3(2) 3-5(4) 8-11(10) —3-6(5) 2,3(2)

5 1 4 1 1 2-6(3) 1-3(2) 1-3(2)

6 1,2(2) 1 1 1 2,3(3) 3 3

7 6-8(7) 3 1 5-7(5) 1,2(2) 4-6(4) 6-10(9)

8 3-7(4) 1 ,2(2) 4-6(5) 7-12(8) - 1 2

9 3-6(3) 1 3,4(4) 4-6(5) 2-4(2) 1 1

10 2-4(2) 1 1 1 1 1 jee

11 1,2(2) 2-6(4) 1-3(3) 1-3(2) 2-6(3) 2,32) 2

12 3-6(4) 1 1 1,2(1) 1,2(2) 2,3(2) 2

13 3,4(4) - 19-28(23) 5-8(5) 2,312) 14-22(17) 3,4¢3)

14 2-4(2) 1,2(2) 10-23(15) ~ ~ - 1

15 4-6(5) - - - : -

Seta Abdominal Segments

No. IV V VI Vil Vill Xx

0 1 1 1 1

1 4,5(4) 4,5(5) 2-5(4) 4-7(5)

2 1 1 1 1,2(1)

3 2 1,2(1) 1 3,4(3)

4 1,2(1) 4-6(5) 2,3(2) 1

5 1,2(2) 1-3(2) 1-4(2) 2,3(2)

6 3 3 3 12-20(17)

F< 7-11(9) 6-9(7) 1 1

8 2,02) 2,3(2) 3-5(3) 4-7(4)

9 1 1 1 1,2(1)

10 1,2(1) 1 1 1

44 2,3(2) 2 2,3(2) 2,3(2) : .

12 1,2(2) 1,2(1) 1 1 . .

13 3,4(3) 3,4(4) 22-43(35) 4,5(4) - -

14 1 1 1 1 1 -

15 - - - “ - >

&@ Based on specimens from Israel, Egypt (Sinai), and Iran.

b Range (mode) obtained from counts made on at least 10 setae.

Harbach: Subgenus Culex in Southwestem Asia 219

Table 34. Number of branches for fourth-instar larval setae of Culex duttoni

Theobald.@

Seta Head Thorax Abdominal Segments

No. C P M T | ll IN

0 1 18-25(20)9 —- - - 1 1

1 1 1 1 1-3(2) 1-5(2) 1,2(1) 1

2 . 1 1-4(3) 1 1 1 1

3 1 1,20 1 4-6(6) 1 1 1

4 1 2,3(2) 2 2-7(4) 10-13(11) 5-10(7) 2,3(2)

5 6-8(7) 1 ,2(1) 1 1 5-14(7) 1-4(4) 2-5(4)

6 7-9(8) 1 1 1 2,3(2) 2-4(2) 1

7 7-11(9) 2 1 5-11(8) 2-5(3) 3,4(3) 6,7(6)

8 1,2(2) 3-6(4) 5-10(5) 16-24(20) - 1 1

9 1-6(4) 1 4-7(5) 5-9(6) 1-5(3) 1 1

10 2,3(2) 1 1 1 1 1 1

11 2,3(2) 2-7(6) 2-4(2) 1-3(2) 2-15(9) 2,3(2) 1-3(2)

12 3-5(3) 1 1 1 1,2(1) 1 1

13 2-5(3) - 18-37(28) 3-6(5) 1-4(2) ~ 15-34(25) 2,3(2)

14 1,2(1) 1 21-37(27) - - - 1

15 3-5(3) - - - - - -

Seta Abdominal Segments

No. IV V VI Vil Vill X

0 1 1 1 1 1 -

1 1 1 1 1 4-9(6) 1

2 1 1 1 1 1 1

3 1 1 1 1,2(1) 7-11(9) 1

4 2,3(2) 1-7(5) 2-5(2) 1 1 4-7(5)

5 2-7(3) 2-6(2) 3-9(5) 2-6(4) 3-6(5) -

6 1 1 1 12-27(20) - :

7 8-16(9) 7-12(9) 1 1 1

8 1 1,2(1) 3-7(4) 7-15(10) 1

9 1 1200) 1 2-5(3) 1

10 1 1 1 1 2

11 2-4(2) 2-4(3) 2-5(3) 1-4(3)

12 1:20) 1 1 1

13 2 1,2(2) 21-52(37) 2-5(4) - -

14 1 1 1 1 1 -

15 - - - - - -

4 Based on unassociated specimens from P.D.R. Yemen, Ethiopia, Cameroon, Ghana,

and Uganda.

b Range (mode) obtained from counts made on 12 setae (in most cases).

were present in specimens from other areas.

220

Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Table 35. Number of branches for fourth-instar larval setae of Culex sitiens

Wiedemann.

Seta Head

No. C

0 1

1 1

2 i

3 1

4 1

5 6-8(7)

6 4-7(4)

7 7-11(9)

8 3-5(4)

9 3,4(3)

10 2-4(2)

11 2-4(2)

12 2:3(3)

13 1-3(2)

14 1

15 2,3(3)

Seta

No. IV

0 1

1 2,3(2)

2 1

3 1

4 1,2(1)

5 2,3(2)

6 2

7 5-8(6)

8 2

9 1

10 |

11 2,3(2)

12 1-3(2)

13 1-3(2)

14 1

15 -

9-16(12)

> —

tak ek OD eh ee FD 6 ok a kk

ee)

PR BW

QNYWHA+~+~wWal <

S SS 8B

eS =] 2, = =

(2)

Abdominal Segments

Thorax

M t |

1 1 1-3(1)

2-4(3) 1,2(1) 1

1 2-4(3) 1,2(1)

1,212) 9-6fA) «55-1449

1,2(1) 1 2-7(4)

1 1 3,4(3)

1 6-9(7) 1,2(1)

4-6(6) 8-15(14) z

4-6(5) 4-7(5) 4,2(2)

1 1 1

2-4(3) 2-518) 1-3(2)

1 1,2(1) 1,2(1)

13-21(16) 2-5(3) 1-3(2)

i@-2ne) ;

Abdominal Segments

mhyA Gs 4 © =

NO SS Eee Gee ya Oper

wWeauwAT r

SS ee eee

oud,

(ee)

{[ = GE —

Vil

bY 1

Ot eo a Sot

Lies

nm —

r ~

cub,

—

-3(1)

&@ Based on specimens from the Yemen Arab Republic, United Arab Emirates, and

Oman.

b Range (mode) obtained from counts made on at least 10 setae.

Harbach: Subgenus Culex in Southwestem Asia 221

Table 36. Number of branches for fourth-instar larval setae of Culex poicilipes

(Theobald) .@

Seta Head Thorax Abdominal Segments

No. C P M T | ll IN

0 1 15-30(22)> —- : , 1 1

1 1 1 5-8(5) 3-6(5) 7-10(9) 5-9(6) 6-8(7)

2 - 1 4-10(5) 3-5(4) 3-5(3) 2-4(3) 1-3(2)

3 1 1 1,2(2) 6-11(S) 2-7(5) 2-4(3) 3-5(3)

4 1-3(2) 2 4-6(4) 5-10(6) 11-19(15) 7-10(8) 3-5(4)

5 3-5(4) 1 1 2-4(3) 5-10(7) 5-16(12) 7-13(11)

6 4-6(4) 1 1 1:22) 3 3,4(3) 3,4(3)

7 10-15(10) = 3,4(3) 1 5-8(6) 1,2(2) 8-18(11) 9-27(17)

8 3-7(5) 2 4-8(6) 8-17(12) ; 1,2(1) 2,3(2)

9 6-12(8) 1-3(2) 4-6(4) 4-7(5) 3-6(5) 1,2(1) 1

10 3-5(4) 1,2(1) 1 1 1-3(2) 1 2,3(2)

11 2,3(2) 4-8(8) 2-4(4) 2-4(2) 6-10(7) 5-9(6) 4-6(4)

12 3-7(5) 1 1 2,3(2) 2-4(3) 2-4(4) 2-5(3)

13 2 - 24-45(32) 12-23(21) 2-5(4) 18-40(29) 5-9(6)

14 1,2(1) 1 31-43(37) . - - 1

15 5,6(5) - - : : . -

Seta Abdominal Segments

No. IV V VI Vil Vill X

0) 1 1 1 |

1 6-10(9) 7-9(8) 6-9(7) 5-10(7)

2 1-3(2) 1-3(1) 1-3(2) 1,2(1)

3 3,4(4) 2,3(2) 2,3(2) 7-10(8)

4 2-5(4) 6-12(9) 2-6(4) 1-3(3)

5 6-10(8) 4-8(5) 3-7(5) 2-8(5)

6 3,4(3) 2 3,4(3) 13-27(19)

7 10-16(13) 10-16(10) 2-6(3) 3,4(3)

8 3,4(3) 3-5(3) 3-7(4) 5-12(7)

2) 1,2(1) 1 1 2-5(3)

10 1,2(2) 1,2(1) 2,3(2) 4-6(5)

11 3-6(5) 3-7(4) 3-6(4) 2-6(4)

12 2-4(3) 1-3(1) 1 1 : ;

13 5-7(6) 5-7(6) 43-57(50) 6-10(6) - -

14 1 1 1 1 1 -

16 - - : - - -

&@ Based on specimens from Egypt.

b Range (mode) obtained from counts made on at least 10 setae.

222

Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Table 37. Number of branches for fourth-instar larval setae of Culex mimeticus Noé.2

Seta

No.

OON ODA AWN OO

Head Thorax Abdominal Segments

C P M cl | I lll

1 8-16(11)° . . : 1 1

1 1 1 1,2(1) 1,2(1) 1 1

: 1 2-7(4) 1,2(1) 1 1 1

1 1 1 3-5(3) 1,2(1) 1,2(1) 1

2-4(3) 1 2,3(2) S64 64201) 374) 2-413)

3,4(4) 1 1 1-3(1) 2-6(3) 1-4(2) 1-3(1)

2,3(2) 1 1 1 3,4(3) 3,4(3) 3

5-8(6) 23(3) 1 5-8(8) 1,2(1) 2-5(4) 5, 6(6)

2-4(3) 2,3(2) 5,6(5) 9-16(11) : 1 1

3-6(4) 1 3-5(5) 5,6(5) 2,3(2) 1,2(1) 1

2-4(3) 1 1 1 1 1 1

2-4(3) 4-10(5) 3,4(3) 2-4(3) 2-A(2) 2,3(2) 2

2,3(3) 1 1 1,2(1) 4-3(2) 122 © 1,2/2)

2,3(2) ; 20-34(25) —1-3(2) 249) 13-88(15) 1,212)

1 1 15-2619) «<3 : : 1

23(3) - - - - - -

Abdominal Segments

IV V VI Vil Vill X

1 1 1 1

1,2(2) 2,3(2) 2,3(2) 3,4(4)

1 1 1 1

1,2(1) 1 201) 2-4(2)

2,3(2) 4-6(4) 2,3(2) 1

1-3(1) 1-3(1) 1,2(1) 2-5(2)

3,4(3) 2,3(3) 2,3(2) 10-17(14)

5-9(6) 5-7(6) 1 1

4,2(1) 2 1,2(2) 2-5(3)

1 1 1 2-A(3)

1 1 1 1

2,312) 1-3(2) 2,3(2) 2,3(2)

1,2(2) 1 1 1

1,2(1) 1,2(1) 25-41 (34) 1-3(1) : :

1 1 1 1 1 .

&@ Based on specimens from Israel and Iran.

b Range (mode) obtained from counts made on at least 10 setae.

C Siphon most often with 12 seta 1-S (6 pairs), but 10-14 may be present.

Harbach: Subgenus Culex in Southwestem Asia 223

Table 38. Number of branches for fourth-instar larval setae of Culex bitaenio-

rhynchus Giles.

Head Thorax Abdominal Segments

C P M x | ll IN

1 10-20(14)> —- , - 1 1

1 1 1,214) 1 1-4( 1,2(1) 2,3(2)

oie 1 3-7(4) 1-3(2) 1,2( 1 1,2(1)

1 1 1 3,4(3) 1-3( 2,3(3) 2,012)

1-4(2) 1 ,2(2) 2302) 2-6(5) 8-11(8) 3-8(8) 1-3(2)

2,;3(3) 1 1 1,2(1) 4-8(8) 2-4(3) 2-4(3)

2 1 1 1 3,4(3) 2,3(3) 3,4(3)

4-7(6) 3 1 6-9(7) 1 3-7(5) 5-10(6)

2-4(2) 2,3(2) 4-6(5) 8-23(16) - 1 1,2(1)

4,5(4) 1-3(2) 5-8(6) 5-7(5) 2,33) 1 1

2,3(2) 1-3(1) 1 1 1 ,2(2) 1,2(1) 2

1,2(1) 3-7(5) 2-4(2) 2-6(4) 2-4(3) 2-4(2) 2,3(2)

2-5(3) 1 1 1,2(1) 1,2¢2) 2 2

1-3(1) ; 23-45(35) 3-6(4) 2,32) 13-18(15) 2,3(2)

1 1 16-34(20) - - - 1

2,3(3) - - : - -

Abdominal Segments

IV V VI Vil Vill X

1 1 1 1

2,3(2) 2,3(3) 1-3(2) 2,3(2)

1 1 1-3(1) 1

2-4(3) 1-3(2) 2-5(3) 2-4(2)

1,2(2) 4-7(6) 2-5(3) 2,3(2)

2,3(2) 2,3(2) 2,3(2) 2-4(3)

2-5(3) 2-5(3) 2,3(2) 10-19(11)

4-10(6) 4-9(7) 23(2) 1-3(2)

2 2 1-3(3) 2-5(5)

1 1 1 1-3(1)

1,2(2) 1,2(2) 1 ,2(2) 2-6(3)

1,2(2) 2 2,3(2) 1,2(1) - -

2 1 1,2(2) 2,3(2) - -

22) 232) 23-35(25) 2-4(2) : -

1 1 1 1 1 :

4 Based on specimens from Iran, Pakistan, and the Yemen Arab Republic.

b Range (mode) obtained from counts made on at least 10 setae.

224

Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Table 39. Number of branches for fourth-instar larval setae of Culex tritaenio-

rhynchus Giles.

Seta Head

No. C

0 1

1 1

2 =

3 1

4 1

5 3,4(3)

6 2,3(2)

m 5-10(8)

8 3-8(6)

9 3-7(3)

10 2-4(3)

11 2

12 2-4(3)

13 2-4(3)

14 1

15 3-5(4)

Seta

No. IV

0 1

1 4

2 1

3 2)0(3)

4 1-3(2)

5 2,3(2)

6 2,3(2)

Fm 6-11(9)

8 2,3(2)

9 1

10 1

11 2,3(2)

12 1,2(2)

13 4-6(4)

14 1

15 -

Thorax Abdominal Segments

e M - l Il

17-23(19)> —- . ‘ 1 1

1 122(4) 1,2(1) 2-4(3) 1,200) 2-4(3)

1 4-8(5) 2,3(2) 121%) 1 1

1 1 5-8(5) 2-4(3) 2-4(3) 2,3(2)

1 ,2(2) 2,3(3) 3-6(4) 11-16(13) 6-10(8) 1-3(2)

1 1 1 4-8(5) 2-4(2) 2,3(2)

1 1 1 2,3(3) 2,3(3) 2,3(2)

2,3(3) 1 5-7(6) 1-3(2) 5-8(7) 6-12(9)

2 3-5(5) — 9-18(11) . 1 1,2(1)

1 3-5(4) 4,5(5) 2-4(2) 1 1

1 1 1 1 1 1

5-9(6) 2-5(3) 1-4(3) 3,4(3) 2-4(3) 1,2(2)

1 1 1-3(2) 1-3(2) 2,3(2) 2

: 20-25(28). Fey: «2,3(2) 11-2317) 2-55)

VO Ot. O0s) - . 1

Abdominal Segments

V VI Vil Vill X

1 1 1

4 3-6(4) 4-7(5)

1 1 1

1,2(1) 1 4-6(5)

5-9(6) 2-4(3) 1,2(2)

2,3(3) 2-4(3) 3-6(3)

2,3(2) 2 10-17(17)

6-10(10) 2-4(3) 2,3(2)

2,3(2) 3-5(4) 4-13(7)

1 1 2-4(3)

1 1 1,2(2)

2-4(2) 2,3(2) 2-4(3)

1 1 1

2-5(5) 21-48(38) 2-6(5)

1 1 1

4 Based on specimens from Iran, Pakistan, and Saudi Arabia.

b Range (mode) obtained from counts made on at least 10 setae.

Harbach: Subgenus Culex in Southwestern Asia 225

Table 40. Number of branches for fourth-instar larval setae of Culex pseudovishnui

Colless.@

Seta Head Thorax Abdominal Segments

No. C P M T | ll I

0 1 11-24(15)5—- : ’ 1 1

1 1 1 1,2(1) 1,2(1) 1-3(2) 1,2(2) 4-8(5)

2 i 1 3-5(4) 2,3(2) 1,2(1) 1 1

3 1 1 1 4-7(5) 3-7(3) 2-4(2) 2-4(2)

4 1-3(1) 1-3(2) 2,3(3) 2-613) aA 4-107) - 2,812)

5 2-4(3) 1 1 1,2(1) 3-5(4) 2,3(2) 2,a(2)

6 1-3(2) 1 1 1 3 3 3,4(3)

7 8-12(9) 3 1 — -4-6(5) 1 4-9(6) 7-12(8)

8 2-6(4) 1,2(2) 3-5(4) 11-20(14) ; 1 1,2(1)

9 5-10(5) 1 3-5(4) 3-6(4) 2-4(3) 1 1

10 3,4(3) 1 1 1 1 1 1

11 1-3(2) 4-6(4) 3,4(4) 2,3(3) 3-5(4) 2,3(3) 2,3(2)

12 2-5(2) 1 1 1-3(2) 1,2(2) 1-3(2) 2,3(2)

13 1-3(2) . 35-44(40) 7-12(9) 2,3(2) 13-32(21) 3-6(4)

14 1 1 14-32(15) : . ; 1

15 2-5(3) - - - - - -

Seta : Abdominal Segments

No. IV V VI VII VIII X

0 1 1 1 1

1 4-7(6) 4-9(7) 4-8(6) 6-9(8)

2 1 1 1 1

3 2,3(2) 1,2(1) 1 3-7(4)

4 1-3(2) 4-7(6) 2-4(3) 1

5 2-4(2) 2,3(2) 2-A(2) 2-5(3)

6 2-4(3) 3,4(3) 2,3(3) 12-23(16)

7 6-12(8) 7-13(8) 3,4(3) 1-3(2)

8 2-4(2) 2,3(2) 3-5(3) 4-11(5)

9 1 1 1 1-5(2)

10 1 1 1 1

11 2,3(2) 2-4(2) 2-5(2) 2-4(3)

12 2,3(2) 1 1 1

12 4,5(5) 4-7(5) 29-55(42) 5-7(5) - -

14 1 1 1 1 1 -

15 - - - - - -

@ Based on specimens from Pakistan.

b Range (mode) obtained from counts made on at least 10 setae.

226

Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

Table 41. Comparison of potentially diagnostic characters observed in members of the Culex

univittatus species aggregate.

Character

1) Ventral surface

of proboscis

2) Postspiracular

area

3) Forefemur

4) Midfemur

5) Hindtibia

6) Wing — Costa

— Vein 2A

7) Abdomen —

Pale bands

of terga

8) Male genitalia

—Setag

— Ventral

arm

univittatus

pale in middle

tendency for scales

to cover less than

dorsal 0.5 —

sometimes with

indistinct anterior

pale stripe

with complete distinct

or indistinct anterior

pale stripe

with distinct anterior

and posterior pale

stripes on proximal 0.8,

separated ventrally by

complete dark stripe;

with distinct apical

pale spot

with short line of pale

scales at base

female usually with

line of scales

normal

narrow

long

perexiguus

pale except at base,

weakly pale on distal

0.25

tendency for scales

to cover more than

dorsal 0.5

usually with indistinct

anterior pale stripe

with or without incom-

plete faint or distinct

anterior pale stripe

with distinct anterior

and posterior pale

stripes on proximal 0.8,

partly separated on

proximal 0.5 or less by

weak ventral dark

stripe; with distinct

apical pale spot

with short line of pale

scales at base

female occasionally

with few scales

normal

broad

short

neavei

inconspicuously

pale in middle

tendency for scales

to occur in small patch

near spiracle

no anterior pale stripe

normally without

anterior pale stripe,

weakly indicated

when present

with rather indistinct

anterior and posterior

pale stripes ending

before base; with

rather indistinct

apical pale apot

without pale scales

at base

female occasionally

with few scales

reduced or absent

broad

long

Harbach: Subgenus Culex in Southwestern Asia

APPENDIX A

Species of Culicidae other than Culex (Culex) known to occur in

southwestern Asia and Egypt

(subspecies and varieties not recognized)

Anopheles (Anopheles) algeriensis Theobald, 1903a

Anopheles (Anopheles) claviger (Meigen, 1804)

Anopheles (Anopheles) coustani Laveran, 1900

Anopheles (Anopheles) habibi Mulligan and Puri, 1936

Anopheles (Anopheles) hyrcanus (Pallas, 1771)

Anopheles (Anopheles) maculipennis Meigen, 1818

Anopheles (Anopheles) marteri Senevet and Prunnelle, 1927

Anopheles (Anopheles) melanoon Hackett, 1934

Anopheles (Anopheles) plumbeus Stephens, 1828

10. Anopheles (Anopheles) sacharovi Favre, 1903

11. Anopheles (Anopheles) tenebrosus Dénitz, 1902

12. Anopheles (Cellia) apoci Marsh, 1933

13. Anopheles (Cellia) arabiensis Patton, 1905

14. Anopheles (Cellia) azaniae Bailly-Choumara, 1960

15. Anopheles (Cellia) cinereus Theobald, 1901a

16. Anopheles (Cellia) culicifacies Giles, 1901c

17. Anopheles (Cellia) demeilloni Evans, 1933

18. Anopheles (Cellia) dthali Patton, 1905

19. Anopheles (Cellia) fluviatilis James, 1902

20. Anopheles (Cellia) hispaniola (Theobald, 1903a)

21. Anopheles (Cellia) moghulensis Christophers, 1924

22. Anopheles (Cellia) multicolor Cambouliu, 1902

23. Anopheles (Cellia) pharoensis Theobald, 1901a

24. Anopheles (Cellia) pretoriensis (Theobald, 1903a)

25. Anopheles (Cellia) pulcherrimus Theobald, 1902

26. Anopheles (Cellia) rhodesiensis Theobald, 1901a

27. Anopheles (Cellia) sergentii (Theobald, 1907)

28. Anopheles (Cellia) splendidus Koidzumi, 1920

29. Anopheles (Cellia) squamosus Theobald, 1901a

30. Anopheles (Cellia) stephensi Liston, 1901

31. Anopheles (Cellia) subpictus Grassi, 1899

32. Anopheles (Cellia) superpictus Grassi, 1899

33. Anopheles (Cellia) turkhudi Liston, 1901

34. Aedes (Aedimorphus) hirsutus (Theobald, 1901a)

35. Aedes (Aedimorphus) natronius Edwards, 1932b

36. Aedes (Aedimorphus) vexans (Meigen, 1830)

37. Aedes (Aedimorphus) vittatus (Bigot, 1861)

38. Aedes (Finlaya) echinus (Edwards, 1920)

39. Aedes (Finlaya) geniculatus (Olivier, 1791)

40. Aedes (Ochlerotatus) berlandi Séguy, 1921

41. Aedes (Ochlerotatus) caballus (Theobald, 1912)

42. Aedes (Ochlerotatus) caspius (Pallas, 1771)

oN oo > Oy =

ge/

228 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

43. Aedes (Ochlerotatus) chelli (Edwards, 1915)

44. Aedes (Ochlerotatus) detritus (Haliday, 1833)

45. Aedes (Ochlerotatus) duplex Martini, 1926

46. Aedes (Ochlerotatus) flavescens (Muller, 1764)

47. Aedes (Ochlerotatus) lepidonotus Edwards, 1920

48. Aedes (Ochlerotatus) leucomelas (Meigen, 1804)

49. Aedes (Ochlerotatus) nigrocanus Martini, 1927

50. Aedes (Ochlerotatus) phoeniciae Coluzzi and Sabatini, 1968

51. Aedes (Ochlerotatus) pulchritarsis (Rondani, 1872)

52. Aedes (Ochlerotatus) rusticus (Rossi, 1790)

53. Aedes (Rusticoidus) refiki Medschid, 1928

54. Aedes (Stegomyia) aegypti (Linnaeus, 1762)

55. Aedes (Stegomyia) cretinus Edwards, 1921

56. Culex (Barraudius) modestus Ficalbi, 1889(1890)

57. Culex (Barraudius) pusillus Macquart, 1850

58. Culex (Culiciomyia) nebulosus Theobald, 1901c

59. Culex (Lasiosiphon) adairi Kirkpatrick, 1926

60. Culex (Lutzia) tigripes De Grandpre and De Charmoy, 1900(1901)

61. Culex (Maillotia) arbieeni Salem, 1938

62. Culex (Maillotia) deserticola Kirkpatrick, 1924(1925)

63. Culex (Maillotia) hortensis Ficalbi, 1889a

64. Culex (Maillotia) quettensis Mattingly, 1955b

65. Culex (Maillotia) salisburiensis Theobald, 1901a

66. Culex (Neoculex) impudicus Ficalbi, 1890

67. Culex (Neoculex) judaicus Edwards, 1926

68. Culex (Neoculex) martinii Medschid, 1930

69. Culex (Neoculex) territans Walker, 1856

70. Culiseta (Allotheobaldia) longiareolata (Macquart, 1838a)

71. Culiseta (Culicella) morsitans (Theobald, 1901a)

72. Culiseta (Culicella) setivalva (Monchadskii, 1936)

73. Culiseta (Culiseta) annulata (Schrank, 1776)

74. —Culiseta (Culiseta) subochrea (Edwards, 1921 [in Wesenberg-Lund,

1921])

75. Coquillettidia (Coquillettidia) buxtoni (Edwards, 1923)

76. Coquillettidia (Coquillettidia) richiardii (Ficalbi, 1889b)

77. Orthopodomyia pulcripalpis (Rondani, 1872)

78. | Uranotaenia (Pseudoficalbia) unguiculata Edwards, 1913a

Harbach: Subgenus Culex in Southwestern Asia 229

APPENDIX B

Complete synonymy of Culex (Culex) pipiens Linnaeus

pipiens Linnaeus, 1758: 602. Type locality: Sweden, Scania, Veberdéd, Silvakra farm.

*Neotype male (NMNH), designated by Harbach et al., 1985: 9.

bifurcatus Linnaeus, 1758: 603. Type locality: France. Lectotype male (non-extant,

illustrated by Réaumur, 1738: Pl. 40, Fig. 2), designated by Harbach et al., 1985:

7. Synonymy with pipiens by Martini, 1922: 107.

fasciatus Miller, 1764: 87. Type locality: none designated. Location of type(s) (adult)

unknown. Synonymy with pipiens by Edwards, 1921: 345.

molestus Forskal, 1775: 85. Type locality: Egypt, Buhayrah Gov., Rosetta. *+Neotype

male (NMNH), designated by Harbach et al., 1984: 523. Synonymy with pipiens

restored by Harbach et al., 1984: 540; prior synonymy with pipiens by Edwards,

1921« 345,

trifurcatus Fabricius, 1794: 401. Type locality: "Europae paludosis." Type(s) (adult)

non-extant. Synonymy with pipiens by Edwards, 1932a: 210.

luteus Meigen, 1804: 6. Type locality: Europe. Type(s) (female) non-extant.

Synonymy with pipiens by Edwards, 1932a: 310.

domesticus Germar, 1817: 290. Type locality: Yugoslavia, Dalmatia. Location of

type(s) (adult) unknown. Synonymy with molestus by Marshall and Staley, 1937:

23; prior synonymy with pipiens by Edwards, 1921: 345.

rufus Meigen, 1818: 7. Type locality: none designated. +Syntypes (male: female)

(MNHP), see Harbach, 1983: 103. Synonymy with pipiens by Edwards, 1921:

345.

bicolor Meigen, 1818: 9. Type locality: ? Africa. +Type female (MNHP), see Harbach

1983: 99. Synonymy with pipiens by Edwards, 1921: 345.

marginalis Stephens, 1825: 455. Type locality: England, London. Type(s) (female)

(BM). Synonymy with pipiens by Edwards, 1921: 345.

meridionalis Leach, 1825: 292. Type locality: France, Nice. Location of type(s)

(adult) unknown. Synonymy with pipiens by Edwards, 1921: 345.

consobrinus Robineau-Desvoidy, 1827: 408. Type locality: U.S.A., Pennsylvania.

Type(s) (adult) non-extant. Synonymy with pipiens by Edwards, 1932a: 209.

calcitrans Robineau-Desvoidy, 1827: 409. Type locality: France, Paris; Germany.

Type(s) (female) non-extant. Synonymy with pipiens by Edwards, 1921: 345.

230 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

thoracicus Robineau-Desvoidy, 1827: 409. Type locality: France, Paris. Type(s)

(female) non-extant. Synonymy with pipiens by Edwards, 1921: 345.

pallipes Waltl, 1835: 110. Type locality: Spain. Type(s) (female) (MNHP). Synonymy

with pipiens by Edwards, 1921: 345.

unistriatus Curtis, 1837: 231. Type locality: England. Type(s) (adult) (NMM).

synonymy with pipiens by Stone et al., 1959: 254.

pallipes Macquart, 1838a: 37. Type locality: Egypt. Type(s) (adult) non-extant.

Synonymy with pipiens by Edwards, 1921: 345.

rufinus Bigot, 1888: 7. Type locality: Tunisia, El-Djerid. Type(s) (female) (non-extant).

Synonymy with pipiens by Edwards, 1921: 345.

agilis Bigot 1889: 122. Type locality: Algeria. Type(s) (female) (BC). Synonymy with

pipiens by Edwards, 1921: 345.

phytophagus Ficalbi, 1889 (1890): 126. Type locality: Italy, Sardinia, Romagna,

Toscana. Location of types (male; female) unknown. Synonymy with pipiens by

Edwards, 1921: 345.

haematophagus Ficalbi, 1893: 143. Type locality: Europa (e d'Italia). Location of

type(s) (adult) unknown. Synonymy with mo/estus by Marshall and Staley, 1937:

23; prior synonymy with pipiens by ? Edwards, 1921: 345.

melanorhinus Giles, 1900: 342. Nomen novum for pallipes Macquart, 1838b: 37, non

Waltl, 1835: 110.

varioannulatus Theobald, 1903a: 198. Type locality: Azores, St. Michaels. Type(s)

(female) (BM). Synonymy with pipiens by Edwards, 1913b: 53.

azoriensis Theobald, 1903a: 210. Type locality: Azores, St. Michaels. Types (male;

female) (BM). Synonymy with pipiens by Edwards, 1913b: 53.

longefurcatus Becker, 1903: 68. Type locality: Egypt, Cairo. Type(s) (female) (ZM).

Synonymy with pipiens by Edwards, 1932a: 209.

quasimodestus Theobald, 1905b: 88 (bicolor var. 7). Type locality: Tunisia, Sfax.

Type(s) (female) (HNM). Synonymy with molestus by Stone,1957: 172; prior

synonymy with pipiens by Edwards, 1921: 246.

doliorum Edwards, 1912a: 263 (pipiens var.). Type locality: England, Devon,

Kingswear. Type(s) (male) (BM).

autogenicus Roubaud, 1935: 444 (pipiens ssp.). Type locality: France. Location of

type(s) (egg) unknown (probably lost), see Harbach,1983: 106. Synonymy with

molestus by Marshall and Staley, 1937: 18.

berbericus Roubaud, 1935: 445 (pipiens ssp.). Type locality: North Africa [Algeria].

Harbach: Subgenus Culex in Southwestem Asia 231

Location of type(s) (egg) unknown (probably lost), see Harbach 1983: 106.

Synonymy with molestus by Knight and Abdel-Malek, 1951: 184.

Sternopallidus Roubaud, 1945: 54 (autogenicus ssp.). Type locality: France,

Aigues-Mortes and Bages (Pyrenees-Orientales). Location of type(s) (adult)

unknown (probably lost), see Harbach, 1983: 106.

Sternopunctatus Roubaud, 1945: 54 (autogenicus ssp.). Type locality: France,

Arles and Paris. Location of type(s) (adult) unknown (probably lost), see Harbach,

1983: 106.

disjunctus Roubaud, 1957: 3116 (pipiens ssp.). Type locality: France, Normandy, Ifs.

Type(s) non-extant. Synonymy with pipiens by Stone et al., 1959: 254.

calloti Rioux and Pech, 1959: 117 (pipiens ssp.). NEW SYNONYMY. Type locality:

France and Tunisia. Location of type(s) (adult) unknown.

erectus lglisch, 1977: 269 (pipiens ssp.). NEW SYNONYMY. Type locality: West

Germany, Berlin-Zehlendorf, Krummer Fenn. No type(s) designated.

torridus Iglisch, 1977: 269 (pipiens ssp.). NEW SYNONYMY. Type locality: West

Germany, Berlin-Zehlendorf, Krummer Fenn. No type(s) designated.

Qn2 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

APPENDIX C

Complete synonymy of Culex (Culex) quinquefasciatus Say

quinquefasciatus Say, 1823: 10. Type locality: New Orleans, Louisiana, USA.

*+Neotype male (NMNH), designated by Sirivanakarn and White, 1978: 362.

pungens Wiedemann, 1828: 9. Type locality: New Orleans, Louisiana, USA.

Lectotype female (NMW), designated by Belkin, 1968b: 19. Synonymy with

guinquefasciatus by Edwards, 1932a: 208.

fatigans Wiedemann, 1828: 10. Type locality: East Indies [Indonesia]. Lectotype

female (NMW), designated by Belkin, 1968a: 68. Synonymy with quinque-

fasciatus by Dyar and Knab, 1909a: 34.

aestuans Wiedemann, 1828: 11. Type locality: Salvador, Bahia, Brazil. Lectotype

male (NMW), designated by Stone, 1958: 186. Synonymy with guinquefasciatus

by Stone, 1958: 186.

acer Walker, 1848: 8. Type locality: New Zealand. Holotype female (BM). Synonymy

with fatigans by Edwards, 1924: 395.

cingulatus Doleschall, 1856: 405. Type locality: Ambarawa, Java. Location of type(s)

(adult) unknown. Synonymy with fatigans by Edwards, 1932a: 208.

cubensis Bigot, 1857: 329. Type locality: Cuba. Lectotype female (MNHP),

designated by Belkin, 1968b: 15. Synonymy with fatigans by Edwards, 1932a:

208.

anxifer Bigot, 1859: 117. Type locality: Madagascar. Location of types (male; female)

unknown. Synonymy with fatigans by Edwards, 1932a: 208.

serotinus Philippi, 1865: 595. Type locality: Santiago and Valdivia, Chile. Types

(male; female) non-extant. Synonymy with fatigans by Edwards, 1932a: 209.

autumnalis Weyenbergh, 1882: 23. Type locality: Primero River, Cordoba, Argentina.

Location of type(s) (male; female; pupa; larva) uncertain (? Universidad de

Cordoba). Synonymy with fatigans by Edwards, 1932a: 208.

penafieli Sanchez, 1885: 213. Type locality: Mexico City, Mexico. Location of types

(male; female) unknown. Synonymy with fatigans by Edwards, 1932a: 208.

macleayi Skuse, 1889: 1746. King George's Sound, Western Australia, Australia.

Type(s) (female) (MM). Synonymy with fatigans by Edwards, 1924: 395.

skusii Giles, 1900: 292. Type locality: Australia. Location of type(s) (male; female)

unknown. Synonymy with fatigans by Edwards, 1924: 395.

Harbach: Subgenus Culex in Southwestern Asia 233

doleschallii Giles, 1900: 338. Nomen novum for cingulatus Doleschall, 1856: 405.

synonymy with fatigans by Edwards, 1932a: 208.

albolineatus Giles, 1901a: 609. Type locality: Shahjahanpur, India. Lectotype female

(BM), designated by Sirivanakarn, 1976: 33. Synonymy with fatigans by Barraud,

1924b: 1264.

quasipipiens Theobald, 1901b: 136. Type locality: Sambalpur, Central Provinces,

India. Lectotype female (BM), designated by Bram, 1967b: 327. Synonymy with

fatigans by Edwards, 1913b: 55.

fouchowensis Theobald, 1901b: 137. Type locality: Foochow, China. Lectotype

male (BM), designated by Bram, 1967a: 196. Synonymy with fatigans by

Edwards, 1913b: 55.

reesi Theobald, 1901b: 145. Type locality: Hong Kong. Type(s) non-extant (Bram,

1967a: 196). Synoymy with fatigans by Edwards, 1913b: 55.

sericeus Theobald, 1901b: 147. Type locality: Hong Kong. Location of type(s)

(female) unknown. Synonymy with fatigans by Edwards, 1913b: 55.

luteoannulatus Theobald, 1901b: 159 (fatigans ssp.). Type locality: none

designated. Type(s) (adult) (BM). Synonymy with fatigans by Edwards, 1932a:

208.

trilineatus Theobald, 1901b: 159 (fatigans ssp.). Type locality: none designated.

Location of type(s) (adult) unknown. Synonymy with fatigans by Edwards, 1932a:

208.

pallidocephala Theobald, 1904: 73. Type locality: Sennar, Blue Nile, Sudan.

Holotype female (BM). ? Synonymy with fatigans by Edwards, 1941: 316.

cartroni Ventrillon, 1905b: 429. Type locality: Majunga, Moroudara, Madagascar.

*Lectotype male (MNHP), designated herein (see treatment of quinquefasciatus

Say). Synonymy with fatigans by Edwards, 1932a: 208.

barbarus Dyar and Knab, 1906: 210. Type locality: Trinidad. Holotype female

(NMNH). Synonymy with fatigans by Edwards, 1932a: 208.

zeltneri Neveu-Lemaire, 1906, 251. Type locality: Harar and Comboltcha, Ethiopia.

Type(s) (male; female) non-extant. Synonymy with quinquefasciatus by White,

1975: 322.

didieri Neveu-Lemaire, 1906: 257. Type locality: Leopoldville, Belgian Congo.

Type(s) (female) non-extant. ? Synonymy with fatigans by Edwards, 1932a: 208.

pygmaeus Neveu-Lemaire, 1906: 256. Type locality: Imi, Ethiopia. Type(s) (male;

female) non-extant. ? Synonymy with fatigans by Edwards, 1932a: 208.

234 Contrib. Amer. Ent. Inst., vol. 24, no.1, 1988

quasilinealis Theobald, 1907: 415. Type locality: Adelaide, Australia. Type (female)

(BM). Synonymy with fatigans by Edwards, 1932a: 208.

stoehri Theobald, 1907: 419. Type locality: British Central Africa. Type (female) (BM).

synonymy with fatigans by Edwards, 1932a: 209.

christophersii Theobald, 1907: 453. Type locality: India. Lectotype male (BM),

designated by Bram, 1967b: 327. Synonymy with fatigans by Edwards, 1913b:

55:

raymondii Tamayo, 1907 [in Tamayo and Garcia, 1907: 36]. Type locality: Huacachina,

Peru. Location of type(s) (male; female; pupa; larva) unknown. Synonymy with

fatigans by Edwards, 1932a: 209.

aikenii Dyar and Knab, 1908: 61. Type locality: New Amsterdam, Surinam. Lectotype

male (NMNH), designated by Stone and Knight, 1957: 42. Synonymy with

fatigans by Edwards, 1932a: 208.

minor Theobald, 1908a: 298. Type locality: Sylhet and Lungleh, Lushai Hills, Assam

and Calcutta, Bengal, India. Types (male; female) (BM). Synonymy with fatigans

by Barraud, 1924b: 1264.

revocator Dyar and Knab, 1909b: 256. Type locality: Hope Gardens, Newcastle,

Jamaica. Lectotype male (NMNH), designated by Stone and Knight, 1957: 55.

synonymy with quinquefasicatus by Dyar, 1918: 99.

lachrimans Dyar and Knab, 1909b: 259. Nomen novum for aikenii Dyar and Knab,

1908: 61, non Aiken and Rowland, 1906: 34.

goughii Theobald, 1911: 268. Type locality: Onderstepoort, Transvaal. Lectotype

male (BM), designated by White, 1975: 321. Synonymy with fatigans by

Edwards, 1913b: 55.

fuscus Taylor, 1914a: 699 (Culicelsa ). Type locality: Townsville, Queensland,

Australia. Type(s) ? non-extant. Synonymy with fatigans by Edwards, 1924: 395.

aseyehae Dyar and Knab, 1915: 112. Type locality: New Providence Island, Bahama

Islands. Lectotype male (NMNH), designated by Stone and Knight, 1957: 43.

Synonymy with fatigans by Edwards, 1932a: 208.

townsvillensis Taylor, 1919: 836. Nomen novum for fuscus Taylor, 1914a: 699, non

Theobald, 1910c: 319.

hensemaeon Dyar, 1920: 178. Type locality: Los Banos, Philippines. Holotype

female (NMNH). Synonymy with fatigans by Edwards, 1922: 278.

nigrirostris Enderlein, 1920: 51 (fatigans var.). Type locality: Tananarive, Madagascar.

Type(s) (female) (ZM).

Harbach: Subgenus Culex in Southwestern Asia 230

APPENDIX D

Complete synonymy of Culex (Culex) sitiens Wiedemann

sitiens Wiedemann, 1828: 542. Type locality: Sumatra, Indonesia. Holotype female

(ZMC).

impellens Walker, 1859: 91. Type locality: Makessar, Celebes. *+Holotype female

(BM). Synonymy with sitiens by Edwards, 1913c: 232.

microannulatus Theobald, 1901a: 353. Type locality: Quilon, Travancore, [Madras

State], India. +Lectotype male (BM), designated by Bram, 1967a: 243. Syno-

hymy with sitiens by Edwards, 1913c: 232.

gnophodes Theobald, 1903a: 163. Type locality: Bruas, Dindings, [Malaysia].

*Holotype female (BM). Synonymy with sitiens by Edwards, 1913c: 232.

somaliensis Neveu-Lemaire, 1906: 254. Type locality: Djibouti, Djibouti. Syntypes

(male; female) non-extant (?). Synonymy with sitiens by Edwards, 1913c: 232.

nigricephala Leicester, 1908: 149. Type locality: Batu Gajah, [Perak], Malaysia.

Syntypes (male; female) non-extant. Synonymy with sitiens by Edwards, 1932a:

204.

Salus Theobald, 1908b: 256. Type locality: Port Sudan, [Kassala], Sudan. +Lectotype

male (BM), designated herein (see treatment of sitiens Wiedemann). Synonymy

with somaliensis by Edwards, 1911: 261.

jepsoni Theobald, 1910b: 158. Type locality: Suva, [Viti Levu], Fiji. +Holotype female

(BM). Synonymy with sitiens by Edwards, 1924: 394.

Saibaii Taylor, 1912: 28. Type locality: Saibai Island, Torres Strait. Type(s) (female)

(US). Synonymy with sitiens by Edwards, 1924: 394.

paludis Taylor, 1913: 56 (Culicelsa). Type locality: Townsville, Queensland, Australia.

Type(s) (female) (US). Synonymy with sitiens by Edwards, 1924: 394.

annulata Taylor, 1914a: 689 (Culicada ). Type locality: Townsville, Queensland,

Australia. Syntypes (male; female) (US). Synonymy with sitiens by Edwards,

1924: 394.

milni Taylor, 1914b: 196 (Culicelsa annulirostris var.). Type locality: Milne Bay, Papua,

New Guinea. Syntypes (male; female) (US). Synonymy with sitiens Edwards,

1924: 394.

Salinus Baisas, 1938: 204. Type locality: Caloocan, Rizal, Luzon, Philippines. Type(s)

(male, female, larva, pupa) non-extant. Synonymy with sitiens by Delfinado,

1966: 152.

236 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

CONSPECTUS OF TAXONOMIC CHANGES

Lectotype Designations

SOS CARAT are A ee rR gee oS a ON en's 82

Peo co ie VG aa AS ea i oll ent cee SO i! MIR Raia Kesey car UR MEL cen aan cae fo SUM 61

te ee er eho 92

Digi These. VOCs e eink). tee AN eon mere l, RE OEE. pole! a pag by. 82

CAMPOTN VECTOR TOO SI 8 EEG IR Vaid CU aia aah er As Cake los ya Sot 31

GOCIIS TT OT Pe OE re RANE Tales REISS MOA RAIS ge eons od CNS 44

Cee Teoma feo la oe ei Ae ee. bleed T eit bioraoe 82

CONGGl TREO FOU One vl Seto RUS elliot ail Gage. 82

FUSCA VECO, TOOT a ee ik ETE ae ite es Moai SMC thi | 8 82

PROT ye Cr rer enigma ies (eee oe Rte Bae SCG | mew OPTUS OW Siar 48

Wigoce ras Gram Io sr et ea Pa Ph, aeieed © eta tice 45

Macagascanensie VENTOn GO om ce ectay Hib wines | AS), abe? prey tiie 92

FUSS TO Wha ate Pe ON Ng oo RI ES UAT SRT Vi, ARERR) FI og ey 44

PUOIS THEODAI TOUS a ee ae Ee a en ome aavared Gane raed 44

TIOOCeSIATs TNC OA TAOS ain ie wire AP ie eats Pole Led cu 44

Oe ar Wee Ue ere Ce Me LaPierre 92

noniennoramiand De Menon, Fa27 “sian Ga RP BO PR We ies he 4D

BANS FOO ARTE oe ey APS a PR ut aaah iy RR Bd ais oO, 87

na og Wo. ih ce aC a Re oR neh nm GLA ae RC ab na 1, 75

DE OTUOO NIG aM FT i EE OR no ANGE Sa RA hl boi NS RO eR 105

New Synonymies

alpha Séguy, 1924, synonymy with theileri Theobald, 1903a ............... 61

calloti Rioux and Pech, 1959, synonymy with pipiens Linnaeus, 1758 ......... 231

erectus \glisch, 1977, synonymy with pipiens Linnaeus, 1758 .............. 231

ethiopicus Edwards, 1912c, synonymy with biaeniorhynchus Giles, 1901a...... 101

mauritanicus Callot, 1940, synonymy with simpsoni Theobald, 1905c .......... 18

torridus l|glisch, 1977, synonymy with pipiens Linnaeus, 1958 .............. 231

Harbach: Subgenus Culex in Southwestern Asia 237

SYSTEMATIC INDEX

Valid taxa are in roman type, valid taxa treated in detail are in bold roman type;

synonyms and invalid taxa are italicized. Boldface page numbers refer to species

treatments; figure numbers and chaetotaxy tables are italicized.

abdominalis (Culiselsa ), 101

acer (Culex), 232

adairi (Culex), 228

Aedes, 5, 6

aegypti (Aedes), 228

aestuans (Culex), 232

ager (Taeniorhynchus ), 101, 105

agilis (Culex), 230

aikenii (Culex), 234

albolineatus (Culex), 233

albovirgatus (Culex), 82, 87, 236

algeriensis (Anopheles), 227

alpha (Culex), 61, 66, 236

anarmostus (Culex), 82, 87

andersoni (Culex), 66

annulata (Culex), 61

annulata (Culicada ), 235

annulata (Culiseta), 71, 228

annulata (Trichopronomyia), 10

annulus (Culex), 87

Anopheles, 5, 6, 98, 100

antennatus (Culex), 7, 11, 12, 15, 19, 20,

29, 47, 48-52, 55, 56, 60, 65, 66, 77,

78, 80, 81, 96; Figs. 13, 14; 191, 2117

anxifer (Culex), 232

apoci (Anopheles), 227

Aporoculex, 10

arabiensis (Anopheles), 78, 81, 227

arbieeni (Culex), 78, 81, 228

aseyehae (Culex), 234

ataeniatus (Culex), 53, 57

auritaenia (Culex), 92, 97, 236

autogenicus (Culex), 230

autumnalis (Culex), 232

azaniae (Anopheles), 227

azoriensis (Culex), 230

barbarus (Culex), 233

barraudi subgroup, 110

Barraudius, 7, 8, 9, 49

berbericus (Culex), 230

berlandi (Aedes), 227

beta (Culex), 81

bicolor (Culex), 23, 229

bifoliata (Culex), 82, 87, 236

bifurcatus (Culex), 229

biroi (Culex), 106

bitaeniorhynchus (Culex) 11, 12,

13, 16, 19, 22, 70, 78, 81, 101-106,

114, 115; Figs. 2, 37, 38; 203,

223, 236

bitaeniorhynchus subgroup, 11, 91

buxtoni (Coquillettidia), 228

caballus (Aedes), 227

Cacoculex, 10

calcitrans (Culex), 229

calloti (Culex), 231, 236

cartroni (Culex), 31, 233, 236

caspius (Aedes), 29, 60, 71, 227

Cellia, 98, 100

chelli (Aedes), 227

christophersii (Culex), 234

cinereus (Anopheles), 78, 227

cingulatus (Culex), 232, 233

claviger (Anopheles), 71, 227

condylodesmus (Culex), 82

consobrinus (Culex) ,229

Coquillettidia, 5, 6

coustani (Anopheles), 227

cretinus (Aedes), 228

creticus (Culex), 61, 67

cubensis (Culex), 232

Culex (Genus), 2, 5, 6, 7, 10

Culex (Subgenus), 2, 3, 7, 8, 9, 10, 11,

13,29,45'51,63,71,61,99, 410; “227

culicifacies (Anopheles), 227

Culicidae, 5, 227

Culiciomyia, 7, 8

Culiseta, 5, 6

debilis (Phalangomyia ), 10

decens (Culex), 11, 12, 15, 18, 20,

44-48, 50, 57, 64, 70, 76; Figs

11,14; 790, 2170, 236

decens subgroup, 11, 47, 51, 77

238 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

demeilloni (Anopheles), 227

deserticola (Culex), 81, 100, 228

detritus (Aedes), 227

didieri (Culex), 233

disjunctus (Culex), 231

dissimilis (Culex), 82, 87, 236

doleschallii (Culex), 233

doliorum (Culex), 230

dolosa (Heteronycha ), 10

domesticus (Culex), 229

dthali (Anopheles), 81, 227

duplex (Aedes), 228

duttoni (Culex), 8, 11, 12, 13, 17, 19, 70,

82-87; Figs. 2, 29, 30; 199, 219,

236

duttoni group, 11, 13

echinus (Aedes), 227

eleuthera (Culex), 10

erectus (Culex), 231, 236

ethiopicus (Culex), 101, 102, 105, 106

236

exilis (Culex), 36, 40

fasciatus (Culex), 229

fatigans (Culex), 23, 31, 232, 233, 234

flavescens (Aedes), 228

fluviatilis (Anopheles), 227

fouchowensis (Culex), 233

fuscus (Culicelsa ), 234

gambiae (Anopheles), 29, 78

gelidus (Culex), 10

geniculatus (Aedes), 227

gnophodes (Culex), 92, 235

goughii (Culex), 234

habibi (Anopheles), 227

habilitator (Culex), 10

haematophagus (Culex), 230

hensemaeon (Culex), 234

Heptaphlebomyia, 10, 52

Heteronycha, 10

hirsutipalpis (Culex), 82, 87, 236

hirsutus (Aedes), 227

hispaniola (Anopheles), 71, 81, 101, 227

hortensis (Culex), 71, 228

hutchinsoni (Culex), 86

hyrcanus (Anopheles), 227

impellens (Culex), 87, 92, 235

impudicus (Culex), 228

infula (Culex), 104, 105

invidiosus (Culex), 47, 48, 49

Jenkinsi, 78

jepsoni (Culex), 92, 235

judaicus (Culex), 228

karatsuensis (Culex), 101

lachrimans (Culex), 234

Laiomyia, 10

Lasioconops, 10

Lasiosiphon, 7, 8, 9

laticinctus (Culex), 11, 12, 13, 14, 15,

17, 20, 23, 48, 64, 66, 67-72, 73,

74, 81, 85, 90; Figs. 27, 22; 195,

215

laurenti (Culex), 48, 49, 52, 236

lepidonotus (Aedes), 228

leucomelas (Aedes), 228

Leucomyia, 10

litwakae (Culex), 47, 51

lividocostalis (Culex) ,45,48, 236

longefurcatus (Culex), 23, 230

longiareolata (Culiseta), 71, 81, 228

luteoannulatus (Culex), 233

luteus (Culex), 229

Lutzia, 7, 8, 93

macleayi (Culex), 232

maculipennis (Anopheles), 29, 227

maculipes (Taeniorhynchus), 92,

maculipesarabiensis (Pseudo-

taeniorhynchus), 101

madagascariensis (Pseudohep-

taphlebomyia ), 10, 92, 97, 236

Maillotia, 7, 8,9

marginalis (Culex), 229

marteri (Anopheles), 227

martinii (Culex), 7, 228

masculus (Culex), 44, 48, 236

mattinglyi (Culex), 11, 12, 15, 17, 20,

64, 66, 68, 69, 71, 72-75, 85, 90;

Figs. 23, 24; 196, 216

mauritanicus (Culex), 75, 78, 236

melanoon (Anopheles), 227

Harbach: Subgenus Culex in Southwestem Asia 239

melanorhinus (Culex), 23, 230

meridionalis (Culex), 229

microannulatus (Culex), 87, 92, 235

milni (Culicelsa ), 235

mimeticus (Culex), 11, 12, 13, 16, 19,

22, 71, 81, 90, 97-101, 109, 113;

Figs. 2, 35, 36; 203) 222

mimeticus subgroup, 11, 71

minor (Culex), 234

minutus (Culex), 44, 48, 236

modestus (Culex), 228

moghulensis (Anopheles), 227

molestus (Culex), 2,23,24,31,229,230,231

monttforti (Heptaphlebomyia ), 52, 57

morsitans (Culiseta), 228

multicolor (Anopheles), 71, 81, 227

natronius (Aedes), 227

neavei (Culex), 53, 56, 57, 226

nebulosus (Culex) ,228

Neoculex, 7, 8, 9

neovishnui (Culex), 111

nigricephala (Culex), 235

nigrirostris (Culex), 234

nigrocanus (Aedes), 228

nigrocostalis (Culex), 44, 48, 236

Oculeomyia, 10

onderstepoortensis (Culex), 61, 67

ornatothoracis (Culex), 47

Orthopodomyia, 5, 6

pallidocephala (Culex), 31, 57, 233

pallipes (Culex), 23, 230

paludis (Culicelsa ), 235

par (Culex), 92, 97

pavlovsky (Culex), 40

penafieli (Culex), 232

perexiguus (Culex), 11, 12, 13, 14, 18,

20,51 ,52,53;55,56, /57-61, 63,065, 66,

71, 75,81; 96 Figs 47, 118:9 93:

213, 226

perfidiosus (Culex), 47

perfuscus (Culex), 47

pettigrewii (Culex), 61, 67

Phalangomyia, 10

pharoensis (Anopheles), 78, 81, 96, 227

phoeniciae (Aedes), 228

phytophagus (Culex), 230

pipiens(Culex),2,3,10,11,12,13,15,16

18,21, 23-31, 32, 34, 35, 36, 38, 39,

40,42,43,44,46, 50, 51, 52,55,60, 66

71,81;Figs3,4; 186,206,229, 230,236

pipiens group,11,13, 14, 17, 20, 65, 86

109, 114

pipiens subgroup, 11, 39, 51, 86

plumbeus (Anopheles), 227

poicilipes (Culex), 10, 11, 12, 16, 19,

22, 52, 70, 91, 92-97, 104, 110,

114; Figs. 2, 33, 34; 201, 221

pretoriensis (Anopheles), 81, 227

Pseudoheptaphlebomyia, 10

pseudomimeticus (Culex), 97

pseudovishnui (Culex), 11, 12, 13,

17, 19, 22, 110, 111-115; Figs.

41, 42; 205, 225

pulcherrimus (Anopheles), 227

pulcripalpis (Orthopodomyia), 228

pulchritarsis (Aedes), 228

pungens (Culex), 232

punctipes (Aporoculex ), 10, 92, 97

pusillus (Culex), 49, 51, 71, 228

pygmaeus (Culex), 31, 233

quasigelidus (Culex), 92, 93, 236

quasilinealis (Culex), 234

quasimodestus (Culex), 230

quasipipiens (Culex), 233

quettensis (Culex), 228

quinquefasciatus (Culex), 3, 11, 12,

13, 15,16, 18, 21.24, 29;:31+35:

39; Figs. 5,6; 187, 207,232, 233.204

raymondii (Culex), 234

reesi (Culex), 233

refiki (Aedes), 228

revocator (Culex), 234

rhodesiensis (Anopheles), 81, 227

richiardii (Coquillettidia), 228

richteri (Culex), 75, 78, 236

rufinus (Culex), 230

rufus (Culex), 229

rusticus (Aedes), 228

sacharovi (Anopheles), 227

Saibaii (Culex), 235

Salinus (Culex), 235

salisburiensis (Culex), 228

Salus (Culex), 87, 92, 235, 236

sarawaki (Oculeomyia ), 10

240 Contrib. Amer. Ent. Inst., vol. 24, no. 1, 1988

seldeslachtsi (Culex), 77

sergentii (Anopheles), 71,78,81, 101, 227

sericeus (Culex), 233

serotinus (Culex), 232

setivalva (Culiseta), 228

siamensis (Culex), 106

simplex (Heptaphlebomyia), 10, 52,

simpsoni (Culex), 11, 12, 14, 19, 21, 53,

55, 65, 66, 75-78, 79, 80, 81, 100:

Figs... 2, 25, 26; 197, 217,236

simpsoni subgroup, 11, 50, 77

sinaiticus (Culex), 11, 12, 13, 14, 19, 21,

53, 55, 65, 71, 77, 78-82, 96, 100;

2, 2f, 28;-196, 218

sitiens (Culex), 11, 12, 13, 16, 19, 22, 70,

87-92, 94, 95, 99, 100, 107, 109,113,

114; Figs. 37,32; 200, 220, 235

sitiens group, 11, 13, 16, 19, 22, 51, 86,

96, 100, 104, 109, 110, 114

sitiens subgroup, 11, 91, 96, 110

skusii (Culex), 232

somaliensis (Culex), 87, 88, 235

splendidus (Anopheles), 227

sp. near infula (Culex), 78

squamosus (Anopheles), 227

stephensi (Anopheles), 227

sternopallidus (Culex), 231

Sternopunctatus (Culex), 231

stigmatosoma (Culex), 10

Stoehri (Culex), 234

Sstriatipes (Culex), 77

subochrea (Culiseta), 228

subpictus (Anopheles), 227

summorosus (Culex), 106, 107

superpictus (Anopheles), 81, 101, 227

taeniorhynchoides (Culex), 92, 97

telesilla (Culex), 47

tenax (Taeniorhynchus), 92

tenebrosus (Anopheles), 227

territans (Culex), 228

terzii (Culex), 77

thalassius (Culex), 78, 88, 91, 107

theileri(Culex),11,12,13,14,18,21, 36,

39,52,55,61-67 ,69,70,71,73,74,77,

78, 81, 90, 100; Figs. 2, 19, 20; 194,

214, 236 |

theileri subgroup, 11, 66, 71, 74

Theobaldiomyia, 10

thoracicus (Culex), 230

tigripes (Culex), 48, 93, 228

tipuliformis (Culex), 36, 40, 62

torrentium (Culex), 11, 12, 13, 14, 16,

18, 21, 29, 36, 39, 40-44, 55; Figs.

2, 9, 10; 189, 209

torridus (Culex), 321, 236

townsvillensis (Culex), 234

Transculicia, 10

Trichopronomyia, 10

trifilatus subgroup, 10, 11, 39, 86

trifoliatus (Culex), 47

trifurcatus (Culex), 229

trilineatus (Culex), 233

tritaeniorhynchus (Culex), 11, 12,

13, 17, 19, 22, 71, 81, 91, 106-

474 313, 104; 11s Figs. GO, 40;

204, 224

turkhudi (Anopheles), 81, 227

unguiculata (Uranotaenia), 71, 81, 228

unistriatus (Culex), 230

univittatus aggregate, 56, 226

univittatus(Culex),11,12,13,14,18,20

51, 52-57, 58, 59, 60, 63, 65, 75,

77; Figs. 2, 15, 16; 192, 212, 226

univittatus subgroup, 11, 50, 51

Uranotaenia, 5, 6

vagans (Culex), 11, 12, 13, 15, 17, 21,

36-40, 42, 43, 55, 63; Figs. 2, 7, 8;

188, 208

varioannulatus (Culex), 230

vexans (Aedes), 227

virgatipes (Culex), 36, 40

vishnui (Culex), 111, 114

vishnui subgroup, 11, 91, 110

vittatus (Aedes), 227

watti (Culex), 86

whitmorei (Culex), 96, 110

zeltneri (Culex), 31, 233

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Contributions

of the

American Entomological Institute

Volume 24, Number 2, 1988

A SYNOPSIS OF THE NORTH AMERICAN

LEPIDOSTOMATIDAE (TRICHOPTERA)

John S. Weaver III

CONTENTS

SNS 6 OCTET MANES Ue tai AE aad Mg arta ce gk Ucar NU ates gly aki ay VO rarely aT ne a eo

EUR Le |e) S20 Oi) UR ROR ae rN meena giant Moma a ae, Mt es ea Er een aa og ae eae

Pe CC eae dia Mie Sg AMUN GMCS Beit yoke be boas 8

CEOS TOMATTOAR UiLWiiIK ey biel 2 Sell BO a ade ees

Bo te ies ions hers ea ea ek ee uO a Gis wa 4

bet Va ee 6 eka He Bana waar Uc Guernica Pci mere bake | ANN Sun Sacre ce On Pra ty ea ee

PAU alae Soe GMachaxats ni picy og BVA Sis Ni aeRO cc TPL “ota MINE Ee osc Aral Beary eS UN ns WOR w lack Sd ds

Rg EE 0 a ea epee ier eae irelanatnntmmepiel OS Say ay GR EY Slee w

LO 1L ERURER BSS Ale a URE IN? UR TRDE NE SA GOGO Tabod CVNNEG ely GPCL tease tes ate Ap GD hat aetna aa ae

AON aU ee ra ta oa ua NDMP trig) Uri pyre ele AG eligi Wwe Mido wie

ee pe tere WRC rye Re Ns FR RES D'y a See es

Keys to North American Lepidostomatidae Genera ..............00 002s

SO urn et aie ie Mane CRIA aR a vale Yar nucini te ater a tera Muny ate Re daria Gee

PTT he ie ie PINE re its MSSM R A Ce Se hae ace

PF seh ye Ng ee AL ig Oh gilda an bes A Smale SG Jet yee a deaiee

Key to Aauets Or 7 HeTODSVCNe Species. oi he ei eA bed ca.

TTC OBEN CS OPO Se ik ee are RE ae Be ae ho

TONG O D2 VGie mOrOn & KORE oy aw ee Ge ame eae ee ON aie ea eee bg

PRTC OE VOTE reo TC HAGe) air au reas Cae Newer he he eas

Te ODS ee es SOW aay bP es Ga ae bee be Ra ed a le ees

PUGRTG eye ATT ON ie ie a tia By Ai ora GS Ralalee ik oa es

Tele vere Pano poosa Haris oe OG Le Wa ee ea

CLONE Ses ee il ens iw eet COR ets NE ee a ay

Key to Larvae of Lepidostoma Species or Species Groups ..........

Key tO Adults) Gf epidostomdg SUBOEIera i. flu ee PEW a Seed eed

Lepidostoma Subgenus Nosopus McLachlan

Key to Adults of L. ({Nosopus) Species

Lepidostoma (Nosopus) Reosum Group

L. (Nosopus) reosum Denning ....

UR oo ie Timer So eh, Umm i et SOs Me Per ena ae RT aie Se gk a YO ME Oe RO a

Oe NOS OL TOS HO ces “Eel he, Ke) 2a) eee el Ce we! ee se le nee én ee

BL SOU 8) VOR Ort KOLGe TOA OHS BON er On el © Ae VO! Men SS Ot ee ce

Wiies tee so.) ‘ere Ses .e RLS eter er ie! OSS er ae ee See eee. 6

Leproostenm TVes opus) AStonedi Grou foe bee IIe Bee ed en

Lepidostoma (Nosopus) astaneum De

RE ee it Oe Nak he ARRON KG

Lepidostoma (Nosopus) goedeni Denning (n. syn.) ..........

Lepidostoma (Nosopus) recinum Den

Lh SRE AU Cra eee ra LG. Frnt ao aan

Lepidostoma (Nosopus) calense Denning (n. syn.) ..........

Lepidostoma (Nosopus) Jewetti Group .

Lepidostoma (Nosopus) jewetti Ross

Lepidostoma (Nosopus) licolum Denn

COOMA esc Ri ars ue Wi? Seat ae Saeko Sekt AM SOME Me Tea SO, A I MR ee ce ek Ue SR

Wilde hie: OO. ee met Ae ee ela e588 6. ele Bg Be 88: Oe es we

OS ee eb Os a ee

Lepidostoma {Nosonus) Americanum Group © 2.6 ee ee ea eas

Lepidostoma (Nosopus) americanum

ee Wes oe a

hepidostona (Nason s) Drvant): (eats) sleek he ak boa e ewe

Lepidostoma wisconsimense. Vochies (Sy. yoo seers eo ean.

Lepidostoma (Nosopus) costale (Ban

ee GUNS Par ea mica te ea us

Lepidostoma (Nosopus) prominens (Banks) co.cc. oe oe

Lepiagostoma (Nosopus}) FPodaoriaan Group 50s. sc cl eo et es heraere

Lepidostoma {Nosopus) podagrum (Mclachlan) .....2)...00.0008.

LERIOCSLOMG LNOSOBUS?) CHEF CINUT ROSS. eee Oe ee ca bes

LepiGosrore JNoOsOpus? (ROG Cine |. oh fcwdeods Oicc cenlna Saeed ewes

Lepidostoma (Nosopus) roafi (Milne)

—

oOODmDuUUNpAL GO eS)

Lepidostoma

Lepidostoma swannanoa Ross, n.

Lepidostoma

Lepidostoma strophe Ross, n.

Lepidostoma

Lepidostoma leechi Denning (syn.)

Lepidostoma

Lepidostoma (Nosopus) Mexicanum Group

Lepidostoma

Atomyiodes bispinosa Ulmer (syn. )

Lepidostoma alexanderi Denning (syn.)

Lepidostoma

Lepidostoma (Nosopus) Hoodi Group

Lepidostoma

Lepidostoma roemhildi Denning, n.

Lepidostome (Nosopus}, incertem Sens S25 DRS Oe RE PRS

Lepidostoma

Lepidostoma plecum Ross (syn.)

Lepidostoma mirum Denning, n. syn.

Lepidostoma

Lepidostoma Subgenus Mormomyia Banks

Key to Adulte-of 2. (Morpmemyla) Soeciog iy ecis ti Sivek r i a

Lepidaostome (Mormonvia} Crleeum Groum oo oie eed ce Se eho leek

Lepidostoma

Lepidostoma (Mormomyia) etnieri Weaver n. sp.

Lepidostoma: (Nosopus) Modestui Grou? oii UG OR Re

(Nosopus }compressum Pinter *t Way ) ei ores.

(Nosepus } tydia Kase oi) Wa ee ee ee ae es

(Nosopus) modestum (Banks)

[Nosopus) ontario Rese re rer ee ae eas

(Nosopus) pictile (Banks)

Be Oe 0 EOS ee ee eee ae 8 ee Se MCE

syn.

(Nosopus) weaveri Harris

©) Oe 6, er 6s eS EL 6: Se ee 8 Ser we eS, LO

Lepidostome: (Nosonus) UnieGler: Group Veer a ae

[Nosopus) DORR) ewe Cree See a SS

(Nosopus) cinereum (Banks)

syn.

(Noso pus) Froste Ee Se ee Srey

(WeSORES Kh age OP at IS

RT eh eee ees le Ue) cer ee Ge we @ Seb ter Me” eee

9 0 ware 8 8 OO gt eee Oe OP ele! 6 6 a Le ere ee

(Nosopus) unicolor (Ban ks) re cee ener

(Nosopius) ecaralgn Denning: (Oo. a WPS eh

(Nosopus} ertectneFiint Bo Buena wo 9 CSO

[Nosopus?) dafila Buens’' © Contrenwe >. 22 eewee...

[Nosepus). elena) Resse OO Dy Pease EOE RS Y

(Nosopus) denningi Weaver n. sp.

[Nosopus) frontale PRaenks) OE eho REPRE .

(Nosopus) heveli Flint & Bueno

(Nosoous) ‘Hachnatane PHAR Ve ie iA ic ce:

(Nosopus) leonilae’ Bueno & Contreras 0.5 Oelias...

(Nosopus) mexicanum (Banks)

CO a ON Ae (ge oe el ate Ger te ee he ede ©

Piste) ee See BTS Se Se) ie Ne 78: ele vet at Veo ee at ig ®

ea es ONE e Ne oe, ove Ab ie) Te De) ete Nee © Seo ce fe

(Nosopus) oaxacense Bueno & Contreras ...........

{(Nosopus) gallo’ Bueno’ t= Padilla oP 4e, Pe aied. ae Pare

(Nosopus) ‘rectangulare: PUNE 206.4502 12s RAE

(Nosopus) reimosert: Flint: s Bwene” .ecier wees Oo 2

(Nosopus) steinhauseri Flint & Bueno 9 Slii i vate..

(Nosopus) talamancense Flint © Bueno Jo. eee... 0.6.

(Nosopus) ¢rimdnde: Weaver ne Sb Por ET een. ek

(Nosapuay Heed? Rese 4 ANT Are Ree Ae,

(Nosopus) spicatum Denning

syn.

(Nosopus) cdscadense® (MiNe) (06000 Rc a EX Bes es

a istite ie 8rie oe 4! 7 6. Celia (eo e- a Soe Fe Slee @) &

(INOSODUS): PRTOOGUNT BOS8 is Cor ks FARE. OAR &

(Nosopus) deceptivum (Banks), nomen dubium

eR eR 8 Oe OE cere ey, 8) 8) ee Ve ye ee ig ke A! we he <e

iMorpmomyla): grineum LBanks)- oon ci Cr wewiain.

(Mormomyia) morsei Weaver n. sp.

OF ee eS Aah ce BAGS Op be ew 8 8

Lepidostoma (Mormonyia}): Serratum Growe ee. 66. io ee ee ca ek s

(MOPMOINYIO) COFPOH! FANE A iieu 544 Ga Gh Rs ek

(Mormomyia) serratum Flint & Wiggins .............

pre! a AT Ey a Ce ge i ge

Lepidestoma (Mormomyia) Libum Groupiiniiu. di Wann).

Lepidostoma (Mormomyia) excavatum Flint & Wiggins ..

Lepidostema (Mormomyia) (fbuni ROSSiiic.. oye aie. «0s

Lepidostoma (Mormomyia) sommermanae Ross .........

Lepidestona (Mormomyla) Vernale Groupies onriniin vo' > ee niAd elas ds

eee ee ec e ee @

C8 we One Sh ee ie

Lepidostoma (Mormomyia) flinti Wallace & Sherberger ...........

Lepidostoma (Mormomyia) glenni Wallace & Sherberger

Lepidostoma (Mormomyia) sackeni (Banks) ...........

Lepidostoma (Mormomyia) vernale (Banks) ...........

Lepidestoma (Mormomyia) Mitchelli Groupe< ia .c.0Ks)+4a%

Lepidostoma (Mormomyia) lobatum Wallace & Sherberger

Lepidostoma (Mormomyia) mitchelli Flint & Wiggins ....

Oy POLeh erie) te erie 6 fe

ee e@ © © © we ee

O26 AO) -0:. 01) 02 eo) et ies Co

@2 ey ee: Tree. ‘lel ‘ee: |e

ee © © © © © ©

Lepidostoma (Mormomyia) styliferum Flint & Wiggins ............

Lepidostoma subgenus Neodinarthrum Weaver n. subgen. ...

Key. to Adults: oF Lo N@Gd Ore Li) \SPRGIES) ern! Mos Gel Maile a oo

Lepidostoma (Neodiaarthrum ji;Cant hum: Group wi sons seus wh ye ae a.

Lepidostoma. (Neodinarthrum)sbaxea Denning Gi eee caWy Mars ais

Lepidostoma (Needinarencium): COMLAUMAROSS i suisse siinenies eae.

Lepidostoma (Neodinerthcuan PluVigle Group ponyionwe)). Se cgheaisieuies..

Lepidostoma (Neodinarthrum) apornum Denning ......

eee © © © © © 8

Lepidostoma.{ Neodinarthrum) .errigenum: Denning: . 2.8.6.4...

Lepidostoma, {Neodinarthetium) JObOR ROSS \ <siierey) s s/s

Lepidostoma (Neodinarthrum) ormeum Ross ...........

Léepidostonid Guerin Denning, MsiiSy Wan wees deeshwekehe d+

Lepidostoma monekum Denning pon wy SY Ne wae sues - ain.

eee © © @ © @ @ @

Lepidostema tNeodinattihrum) pluviale (Mine) on. aun anda oe.

Lepiaostome ui hi no! ROSS) (NenSVne cane Dake wee Pea eld»

hepigostoma /Veled Um) Derniae Muni S Wiad ae nla Qalasigei wills a. es

Lepidostoma UN eodlnarthncum)-FOyVNer! ROS Siiyieudd dicaimei ua eiede vee

Pepi dostemenery OCURT GOED LING pny SY May nena eid igre oa eewienety 6

Lepicgostoma fischer) ‘DENWING i NASYN. haveaeaid s swe

Lepidostoma Subgenus, Lepidostoma -Rambur jvc...) sxe ewies’s kas

OPO Ore Sey Ker ei ceae. o

Key te Adulte, of Nearctic Lepicostoman( 1, 1) Species: wand embinge.’s »

Lepidostoma: {(Lepidostoma)) carolind:( Banks.) sms. aes

Lepidostoma (Lepidostoma) latipenne (Banks) ........

Lepidestomea | Repidestoma). stigmanDBalniksii it siom saya. payee ales

Letmigostonia Mae wltonh) MOssy MN iS yl. (ise ob abi. ae ee ASL

Lepidostoma (Lepidostoma) tibiale (Carpenter) .......

$59 4e)) le Were) ee ie) e. ©

Lee praosteria rile) MARINE TOM MM i56e <a samen Sa G) «doula Layee a) mle

Lepidostoma (Lepidostoma) togatum (Hagen) ..........

Silo. PahiiGias Bias Sy Mois aia mar i whignaeer A .

Pristosilo (Canddensisy Bal her (SViiieiir P aiwincaw inks tes

@) 9) Gor ke ied eye ese

Ee PR aa are ek ela cabal iattiks, Nae alana aliiiiethd ak a RA arb awit olietahiai de aNd Gtidlie @ * ele

Coe heh te ti ekt let ve (ie, ‘ou Lacie (6. Ts \letbienrce (ke en cen e4 (6 len) oe] Gmeeee. seller Tet Onuleneeiiel LOULe Ler e Paar iene PORni@s Ui Sy Si) MC. ONO? CuO, LON Olea VO Tel: OES Be ee

Oey ob OL iecched Cer leks eet eaelel ielite Selina) <6) eee: Yelle! 6@ wh reuieh, 6 he rrelena'e)) ed Kener ie lee Ue) Te. A8\)) OYLOL MSG Suet Re ey eee CS) Oh [en Oy NOL en NOU Orie). Rh Bi. St (ea! Oe

A SYNOPSIS OF THE NORTH AMERICAN

LEPIDOSTOMATIDAE (TRICHOPTERA) ?- ?

John’ S. Weaver’ Til?

ABSTRACT

A synopsis of the North American Lepidostomatidae (79 spp.) is provided,

including Theliopsyche ‘(6 spp.) and Lepidostoma (73 “spp: ). The genus

Lepidostoma is divided into 4 subgenera: Nosopus (45 spp.), Mormomyia (15

spp.), Neodinarthrum, n. subgen. (8 spp.), and Lepidostoma (5 North American

spp i), In the genus Lepidostoma 4 new species are described, denningi,

ermanae, morsei, and etnieri; and 13 new junior synonyms are recognized, goedeni

Denning of astaneum Denning, calense Denning of recinum Denning, swannanoa

Ross of pictile (Banks), strophe Ross of cinereum (Banks), roemhildi Denning

of spicatum Denning, mirum Denning of cascadense (Milne), quer/um Denning and

monekum Denning of ormeum Ross, rhino Ross and veledum Denning of p/uviale

(Milne), cryptum Denning and fischeri Denning of rayneri Ross, and knowltoni

Ross of stigma Banks. Three species previously known only from female types

are herein associated with males: pjictile previously considered a junior synonym

of americanum (Banks) is a senior synonym of swannanoa, and cinereum and

stigma are senior synonyms of strophe and knowltoni, respectively; deceptivum

(Banks), nomen dubium, remains the only North American species of Lepidostoma

known solely from its female. Keys are provided for males, females and larvae

to genus and subgenus, males to species, females of Lepidostoma (L.) to species,

and larvae of Lepidostoma to species or species groups.

INTRODUCTION

Sexual dimorphism prevails throughout the Lepidostomatidae with variable

secondary sexual characters so bizarre that McLachlan (18/76) referred to this

group (section IV of Sericostomatidae) as the "curiosity shop" of the Trichoptera.

Many trichopterists have viewed these odd characteristic differences as a basis

of recognizing new genera. As a result, 20 genera in the family are monotypic

or contain only two or three species, reflecting a general indecisiveness regarding

generic relationships within the family. Furthermore, this problem is evident in

certain monophyletic species groups within the genus Lepidostoma which appear

to include members of other "genera" from the Old World.

*Completed in partial fulfillment of the requirements for the degree of Doctor of

Philosophy in Entomology from Clemson University.

This is Technical” Contribution’ No. 2832 of the South Carolina’ Agricultural

Experiment Station, Clemson University.

*Department of Entomology, University of New Hampshire, Nesmith Hall, Durham,

Ne 03624) USA.

2 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

Ross (1938b) found it necessary to define the genus Lepidostoma very

broadly and mentioned that a revision of the entire world fauna of the family would

be necessary to correctly evaluate the genus. Consequently, Ross (1944, 1946)

decided that the system of classification in use was artificial and presented a

contrasting treatment for the Nearctic Lepidostomatidae. Rejecting the tradition

of previous workers, he concluded that the secondary sexual characters of males

were not important generic-level indicators. Accordingly, Ross (1944) included

nearly all of the Nearctic genera in the synonomy of Lepidostoma. However,

Schmid (1980) remarked that this solution was too radical and suggested that

eventually the secondary sexual characters as well as others must be considered

to provide a much desired revision of the world Lepidostomatidae.

Herein, Lepidostoma sens. fat. is recognized with four subgenera

(Lepidostoma, Mormomyia, Nosopus, and Neodinarthrum) and 19 species groups.

This position seems to offer more nomenclatural stability than the more subjective

treatments of this group used in the past. However, this classification certainly

is not intended to be final, rather it is offered as an improvement to the current

treatment of the North American fauna. This scheme is a compromise between two

general schools of thought regarding generic determination in Lepidostomatidae,

that of the Old World which has been overly generous and that of the New World

which has been too conservative. Presently, | refrain from splitting Lepidostoma

into other genera until a more definitive revision of the world lepidostomatid

genera can be provided.

In the bibliographies of each taxon that follow, original descriptions, junior

synonyms, and pertinent works that have become available after 1960 are included

(brackets are mine). Other earlier citations of the Lepidostomatidae are listed

in the Trichopterorum Catalogus by Fischer (1970, 1973).

In the material examined, the known collectors correspond with their initials

as: follows: . CPA, sC.. Ps. Alexander: NHA, Whe Anderson: JHB,..J.. H:.. Baker:

WEG, .W. Fs Barre AWE RoW Baumann: 26a)... Bueno. Sdria:..MRB, Mau R.

Butcher: PHC, ePo te GCerigon AD A. DammpkhsoCbbD, «GO. .8): Danns..DAE . De. A.

Biners GME. Cs Vio Plt OSE, OucnS a. Flint. cre Bee. 1. Prison: THF: dx;

J. Beobisong. Je ARG Ast Rico Gautine -DRGy JD. Re Givens KG; Ka Goeden;

Di, a bs Gostateans tow S. Wo Hamilton: SCH, SiC. Harriss DDH, . Du. Da

Herlong; RWH, R. W. Holzenthal; SGJ, S. G. Jewett; HDK, H. D. Kennedy; JK,

J, Kitresolvers -Hek, ft. abe beech; “EVM, E. “Mic wMeEwan. Hack: ECM, EW ©.

Masteller; JCM, J. CG. Morse: APN, Av Buc Nimmes DEP; D.. Las Peck; JP, Jd. Peck:

SP, Si-Prichard: WEB. Wo £. Ricker: HAR, HoH. Ross: JAR, J. As Ross: MWS,

Mo OW GanGgereeme Les: so es Serpe PiS. Peogu: Steburth: SDS,.S:..D. Smith;

WPS...W Pa Starke RES) Rees Surdieks: JLG..9.. d. Sykorak. KWS ck. Wi Stewart:

Hoe MT, Ai sbiMa> Townes; Jew od: Wei Weaver iit: RLW) <Re.Le Wescott; TRW,

Toc RR. White: .. GBW, °G.. .B. Wiggine)..RWW, RR. Wi Wisseman;.: JRW,..Ja.:R.) Woods

TY, T. Yamamoto. Also, in the material examined following the collector's initials,

the institution or person currently holding the specimens is indicated in

parentheses and abbreviated as_ follows: BYUC, Brigham Young University;

CASC, California Academy of Sciences; CNCI, Canadian National Collection;

CUCC, Clemson University; DGD, Donald G. Denning's collection; SCH, Steven

C. Harris's collection; INHS, Illinois Natural History Survey; MCZC, Museum of

Comparative Zoology, Harvard University; OSUO, Oregon State University; PURC,

Purdue University; ROME, Royal Ontario Museum; LES, Larry E. Serpa's col-

lection; SDS, Stamford D. Smith's collection; USNM, Smithsonian Institution,

National Museum of Natural History; UASM, University of Alberta, Entomology

Department; UICM, University of Idaho; ECUT, University of Tennessee; JSW,

author's collection.

Weaver: North American Lepidostomatidae 3

ACKNOWLEDGMENTS

This work would not have been possible without the assistance of many

people. | am very thankful to John C. Morse, who has supervised the progress

of this investigation from its commencement to its completion; his knowledge of

phylogenetic systematics and caddisflies has been invaluable. Also, | wish to

thank Jewel A. Jordan and Ada Lou Steirer for reviewing the manuscript and

Dianne H. Haselton for editing the final text. | am grateful to the following people

who willingly provided many specimens and data from either their personal col-

lections or resident institutions: William F. Barr, Richard W. Baumann, Peter C.

Barnard, Joaquin Bueno Soria, Donald G. Denning, Nancy A. Erman, David A.

Etnier, Oliver S. Flint, Jr., Daniel L. Gustafson, Eileen M. McEwan Hack, Steven

W. Hamilton, Steven C. Harris, Ralph W. Holzenthal, Kenneth L. Manuel, John

C. Morse, Alfred F. Newton, Andrew P. Nimmo, Wojciech J. Pulawski, Milton W.

Sanderson, Fernand Schmid, Larry E. Serpa, Scott R. Shaw, Stamford D. Smith,

William P. Stark, Jeffery C. Steven, Rebecca F. Surdick, Bernard G. Swegman,

Jan L. Sykora, John D. Unzicker, Robert D. Waltz, Glenn B. Wiggins, Robert

W. Wisseman, and John S. Wojtowicz.

Finally, | am appreciative to all former trichopterists (especially Herbert H.

Ross) who, in gathering systematic information, laid the foundation upon which

this work was constructed. | am indebted to them all!

LEPIDOSTOMATIDAE ULMER

Ulmer 1903:89 (Lepidostomatinae); Ross 1964:214, f. 6; Lepneva 1966:6, 14, 62,

349; Ross. 1967;174, 178, £2. 1;>Unzieker/1968:4 - 14; '19+20;, 30;°31; 48-51; Fischer

1940335) Panis: 19742452485). Fischer :19732162;:Macan ? 4973: 7) 12 P4909 992 » 93:

Hilsenhoff 1975:24; Anderson 1976:116; Wiggins 1977:13, 16, 19, 21, 31, 154, 155;

Solomon and -Leiser. 1977:32,' 1814: 4668, 472, 1%8y 183; Ress 197802 (63, fe lo Wiegins

19/8:147,.. 158, .163,. 173,179. Pennak- 1976: 601,, 616; Schmid 1980:24, 347+149;

McCafferty 1981:241, 243, 264; LaFontaine 1981:157, 247-253; Hilsenhoff 1981:28;

Unzicker et aly :198279.4; 9:6; :°9.6A; 9:9,°.9.15, 9.19, 9:24, -9/108, 91104-9. 107:

Wiggins. 196412727) 275, 277 ,- 2644 /291 , , 294; | 306) F046 2 995. 16.76," 164 108: 16: 154:

The Lepidostomatidae consist of approximately 285 extant species known.

The family is predominately Holarctic, Afrotropical, and Oriental in distribution,

with however a few species occurring in the northern fringes of the Australian

Region (New Guinea) (Neboiss 1986) and the Neotropical Region (Panama to Mex-

ico) (Flint and Bueno 1977). They were first recognized as a subfamily of Seri-

costomatidae by Ulmer (1903) and later elevated to full family status by Ross

(1944). ;

The lepidostomatids made available as of 1960 inclusively, were cataloged by

Fischer (1970, 1973). Since 1960 the status of a few genera has changed. Ina

review of the Japanese lepidostomatids, Tani (1971) presented two new genera,

Yamatopsyche and Paradinarthrodes. The genus Trichovespula Schmid (1955) of

Chile, originally considered to be a lepidostomatid, was reviewed by Flint (1967b)

and subsequently placed in the Tasimiidae (Flint 1974). Two monotypic genera

Atomyiodes Banks and Eremopsyche Banks from Central America joined the long

list of junior synonyms of Lepidostoma (Denning 1962, Flint 1967a, Flint and Bueno

1977). Ito (1984) recognized Yamatopsyche Tani as a new junior synonym of

Goerodes Ulmer.

A comprehensive review of the Nearctic Lepidostomatidae was presented by

Ross (1946) who listed 4 species of Theliopsyche and 34 species and 6 species

groups of Lepidostoma. Other major works on North American lepidostomatids

followed by Denning (1956), Flint and Wiggins (1961), and Flint and Bueno (1977).

Since Ross's (1946) review the number of North American species has more than

doubled to 6 species of Theliopsyche and 73 species of Lepidostoma.

4 Contrib. Amer: Ent. lnst., Volume 24, Number 2, 1988

Egg and Oviposition

Observations on Trichoptera eggs and oviposition are not abundant. How-

ever, works on this subject by Anderson and Wallace (1984), Hinton (1981),

Silfvenius (1906), LaFontaine (1981), and Wiggins (1973) are highly informative.

In Lepidostomatidae, egg extrusion occurs prior to the act of oviposition,

long before the female releases her spawn. To carry the eggmass, the egg pouch

becomes concave by the contraction of sternites VII and VIII, and segment IX

remains unaltered, overhanging the eggmass like a hood.

There are few reports on the oviposition of Lepidostomatidae. Milne (1943)

observed many females of Lepidostoma (L.) togatum in July "ready for oviposition,

as each had a spherical mass of light green eggs hanging from the tir of the

abdomen." Later he found several such eggmasses in the water. Also, in July,

Roos (1957) netted 20 females of L. (L.) hirtum, eight of which were carrying

eggmasses. Anderson (1976) found many eggs of Lepidostoma in the water. Ross

(1944) states that lepidostomatids deposit eggs in or near the water. Solomon

and Leiser (1977) report that eggmasses are dropped on damp areas near the water

or actually in the water. These reports have been corroborated by R. W.

Wisseman (pers. comm.), who observed that L. (Nosopus) podagrum and L. (N.)

quercinum swarm at dusk and drop their eggmasses directly into the stream during

flight; the eggmasses are very sticky and tend to adhere to benthic substrates.

He also saw females of L. (N.) jewetti oviposit eggmasses directly on damp wood

in seepage areas.

| have examined eggmasses attached to specimens of Lepidostoma pinned dry

and preserved in alcohol. A greater percentage of pinned females have eggmasses

retained than those of alcohol collections. Apparently eggmasses are often dis-

lodged and separated from insects collected and stored in fluids, or they may be

discarded since it is difficult to associate eggmasses with corresponding females.

Larva

Review. Larval descriptions of lepidostomatids have been made available by

Hickin (1967), Ito (1978), Lepneva (1966), Morse and Holzenthal (1984), Nielsen

(1942): “Ross (1944), Ulmer CI903, 1955) and Wiggins (1977, 1984).

Body... Eruciferm:

Head. Hypognathous, distinct from all other caddisflies (except Oeconesidae

and Calocidae) by having antenna adjacent to eye. Apical portion of antenna

seated on small basal lobe. Genae almost meeting near ventral ecdysial line.

Labrum elliptical, its maximum width about a third that of head and about half

as long as wide, usually bearing small membranous anterolateral fringe. Eyes

dorsolateral, a distance of about a third length of head posteriad of mandibles.

Known larvae of Theliopsyche and some species of Lepidostoma (Nosopus) from

Central America have cranial carinae.

Mouthparts. Similar to those of most other phytophagous caddis larvae and

lepidopteran caterpillars (Snodgrass 1935). Maxilla, labium, and hypopharynx

united to form composite structure, with maxillolabial-hypopharyngeal complex

which projects forward like lower lip. Mandibles small and triangular with scra-

perlike ventromesal edge. Maxilla with cardo, stipes, and palpifer, supporting

four-segmented palps and rounded lobelike galea bearing several sensilla. Labium

with triangular postmentum and rectangular mentum and prementum; these col-

lectively supporting pair of rodlike palpigers, two small lateral palps with apical

sensilla, and single apical spinneret.

Thorax. Nota predominantly sclerotized. Sternum with few. sclerites.

Pronotum completely sclerotized by two large plates separated narrowly by mid-

dorsal ecdysial line. Prosternal horn present. Mesonotum almost completely

covered by pair of heavily sclerotized square plates, separated narrowly by

middorsal ecdysial line. Metanotum mostly membranous, usually with three pairs

Weaver: North American Lepidostomatidae 3

of sclerites: lateral pair, setal areas 1, anteromesal pair, sa2, and posteriomesal

pair, sa3. Anterior leg broadest and shortest, with small triangular trochantin.

Middle leg approximately 1.8 times longer than foreleg. Posterior leg 1.5 times

longer than foreleg. Posterior coxa with dense patch of branched or plumose

setae.

Abdomen. Round sagittally and usually with dorsolateral and ventrolateral

row of single filamentous gills on segments II-VII, with varying numbers, some-

times none. Segment | with pair of lateral humps, but middorsal hump absent.

Lateral fringe sparse or absent, and lateral tubercles usually present on segments

MeV aT: Segment VIII with pair of lateral lobes similar to those of Micrasema.

Segment |X with small tergite bearing four major setae. Anal prolegs small and

similar to those of most limnephilids. Anal papillae usually present.

Pupa

Review. The key to pupae of the North American Trichoptera families pro-

vided by Wiggins (1984) is highly recommended for identification purposes. Other

pupal descriptions of lepidostomatids have been made available by Hickin (1949,

1967), Lepneva (1966), Ross (1944), and Ulmer (1909, 1955).

Head. Antennae not excessively long, extending only short distance poste-

riorly of abdominal segment VIII. Labrum semicircular. Mandibles triangular,

stout and pointed apically, being similar to those of most limnephilids. Maxillary

palps two-segmented or apparently one-segmented in male pupae and five-seg-

mented in female pupae. Labial palps always three-segmented.

Thorax. Wingpads appressed against side of abdomen. Middle tarsae mod-

ified for swimming with many fine lateral setae.

Abdomen. Posteriolateral corners of tergite | with dentate ridge. Terga

of segments III-VII each with an anterior pair of hooked-plates, each bearing four

or five short spines pointed posteriorly. Segment V with posterior pair of hooked

plates each bearing five short spines pointed anteriorly. Lateral fringe of

abdomen abbreviated. Abdominal gills single, filamentous, present on segments

II-VI and apparently corresponding in number and position with those of larva.

Anal processes, short stout lobes, about twice as long as wide and _ usually

obliquely truncated and with four long major apical setae. Hickin (1949, 1967)

indicated that the anal processes of Lepidostoma hirtum are long and slender, but

apparently his figures of this species and Goera pilosa have been mismatched.

Case

Lepidostomatid larvae construct many different types of cases (f. 1-6), and

those of some species groups are distinctive. The larva of Lepidostoma (Neodi-

narthrum) constructs a case of fine sand grains (f. 1) which is very slender and

tapered posteriorly. The larva of Theliopsyche, the libum group of Lepidostoma

(Mormomyia) and the mexicanum and hoodi groups of Lepidostoma (Nosopus)

predominantly construct stouter cases of small stones (f. 2, 3). Also, a western

species of Lepidostoma, possibly L. (N.) verodum, constructs a unique spiral case

(Wiggins TO7TTr F.8y bh G)):

Early instar larvae of most lepidostomatids appear to build cases of stones

and the later instars of several species adhere to this primary case construction

plan (f. 1-3). However, later instars of many species switch to a secondary

construction plan in which the larvae make cases of stems, leaves, and other

phytogenic debris that they commonly use to construct either a panel case (f.

4) or a turret case (f. 5, 6). Construction of the panel case was documented

by Sattler (1957) for the larva of Crunoecia irrorata; the larva cuts an oval piece

out of a leaf and then chews four small pieces from the two lateral sides, forming

a hexagonal-shaped panel. Panels are fitted together tightly and arranged in a

6 Contribs Amer. Ent. 4nst«,;: Vohume’24, Number 2, 1988

staggered pattern, positioned opposite the transverse seams of adjacent sides.

This probably reduces the amount of silk needed to glue seams together and

provides longitudinal rigidity. Individual leaf panels are slightly flexible, so that

cases predominantly composed of leaf material are usually somewhat round sagit-

tally, resembling a cigar (f. 4) as in the modestum group of L. (Nosopus).

Occasionally, paneled cases are made from bark or wood and are square sagittally

(Wiggins 1977: f. 8.1 C). Small stones may be incorporated intermittently in

cases, especially cases which are constructed during the transitional construction

phase.

The turret case (f. 5, 6), resembling the turret of a wolf spider, consists

of a whorl of small stems or other long slender plant fragments arranged tran-

sversely and is typical of the unicofor group of L. (Nosopus). The length of

transverse pieces. may vary according to species, and the lengths of stems

incorporated into an individual case may be similar or variable.

Adult

Review. Major contributions to lepidostomatid adult taxonomy and morphology

have been made available by Crichton (1957), Denning (1956), Flint and Bueno

(1977), Flint and Wiggins (1961), McLachlan (1876), Mosely (1939a, b), Ross

(1944, 1946), Schmid (1980), Tani (1971), Ulmer (1907, 1909, 1951) and Wiggins

(1984).

Body. Adult lepidostomatids are medium-sized (forewing length 4-14 mm),

usually dull brown in color, and with elliptical wings. Males exhibit many

extraordinary secondary sexual characters in their wings, thorax, and appendages

of the head. These characters vary greatly and may be involved with dispersion

of pheromones, and thus compensating for the lack of the abdominal scent glands

of segment V. Moretti and Bicchierai (1981) discerned that stout cylindrical setae

in the basal pocket of the forewing's anterior margin of Lasiocephala basalis are

androconia. Furthermore they hypothesized that a secretion is discharged into

the bristle lumen and subsequently dispersed through many small exterior bristle

holes. Similar stout setae (probably androconia) also occur on specific areas of

the scape, head, thorax, and wings (f. 10-13) of many lepidostomatid males.

Frequently setae are modified as scales which may be adjacent to stout setae or

segregated in areas of the wings and maxillary palps. However, Lepidostoma (N.)

podagrum is a curious exception in having modified setae on the anterior baso-

tarsus (f. 29). Modified setae that occur on appendages may be petiolate. Size

of setose warts are sometimes dissimilar in sexes; for example, in males, warts

of the mesoscutellum are larger in L. (Mormomyia) griseum, smaller in L. (N.)

cascadense, and absent in L. (L.) latipenne.

Head. Head short, with eyes large, apparently glabrous. Ocelli absent.

Lateral, frontal and three dorsal pairs of setose warts usually present. Males

often with modified secondary sexual characters in scape and maxillary palps.

Antennae extending posteriorly about as far as apices of forewings in their folded

position. Length of scape variable, often longer than head, but with several

exceptions among males. In New World lepidostomatids, scape often shorter in

males and longer in females, often vice versa in Old World species. For example,

in Dinarthrum the male scape often is 4-9 times as long as the head; in Dinar-

throdes the scape is sometimes longer and is usually subdivided with a false joint.

A great amount of intraspecific variation has been observed in the male scapes

of Dinarthrum iranicum by Schmid (1959) and of Neoseverinia crassicornis by I|to

(1983). Pedicel usually no longer than first segment of flagellum, constricted and

pivoted basally, as in most caddisflies, but sometimes longer or modified as in

Lepidostoma (Nosopus) ermanae, spicatum, verodum and hoodi. Flagellum with

about 30-40 segments, each simple and cylindrical. Tentorium complete (f. 14),

in lateral view shaped like inverted T. Anterior and dorsal arms of tentorium

diverging from posterior arms mesad. Dorsal arms slender, perpendicular to

Weaver: North American Lepidostomatidae q

posterior and anterior arms, and fused with head capsule near antennal bases.

Anterior arms are shorter and in line with posterior arms. Labrum in lateral view

shaped like small beak, about three times as long as broad with slightly bulbous

base and pointed apex. Mandibles small inconspicuous lobes. Maxillary palps

five-segmented in females and two-segmented or apparently one-segmented in

males. Haustellum anterior surface with system of converging channels through

which liquids are imbibed.

Thorax. Pronotum with two pairs of setose warts, mesoscutum and mesos-

cutellum each usually with one pair of warts, and metanotum without warts. Warts

of pleuron variable in number. In Lepidostoma, propleuron with one pair,

mesopleuron with two pairs and metapleuron with one pair. In. Theliopsyche,

setose warts smaller and fewer in number. Legs with few spines but densely

covered with small setae. Spurs also very setose (f. 11). Tibial spur count

always 2, 4, 4 (f. 15) on the fore, middle and hind legs, respectively.

Wings. Herein the Comstock-Needham wing venation system is employed.

Lepidostomatid wings are elliptical because anal regions are reduced and apices

are rounded. In Trichoptera, unique differences in wing venation often have

been regarded as important generic-level indicators. However, because sexual

dimorphism causes excessive variation in the venation of male lepidostomatids,

Mosely (1939a) suggested that "the practice should be reversed and that where

species can be conveniently grouped on a similarity of genitalia, this character

should be considered as pertaining to the genus and that the species should be

separated on neuration." In most lepidostomatid females wing venation is con-

sistent and more complete than in males. However, in Theliopsyche the wing

venation of males and females is very similar.

Anterior wing: Costa (C) normally straight, but in males sometimes curved

anteriorly, forming marginal costal cell area of various size folding over the wing

or sometimes forming small basal pocket. Many modified setae often in this

anterior marginal fold. Subcosta (Sc) heavy, straight, usually parallel with

radius (R). Discal cell (d) delimited by first fork of radial sectorial veins and

sectorial crossvein (r), long in forewing, and much smaller in hindwing. Radial

sector (Rs) branched into four apical veins (R2, R3, R4, R5); fork of R2+3

defining « first “fork: CF) and fork of (R445 defining» 2. Media (M) rarely

unbranched; usually branched into M1+2 (Ma) and M3+t4 (Mp); Ma sometimes

branched, forming F3, but Mp never separating, such that F4 always absent in

Lepidostomatidae. Thyridial cell (th) delimited by bases of median and cubital

veins and mediocubital crossvein (m-cu), usually long and containing a thyridium

(a small transparent spot, similar to the nigma) in females, but usually absent

in males. Cubitus (Cul) thick and usually branched into Cula and Culb,

delimiting F5. Postcubitus (Cu2) unbranched, normally converging with crossvein

(cu-a) and anal veins (1A, 2A, 3A) in distal submarginal hyaline area, the arculus

(ar). However, these veins often confounding due to secondary modifications of

males. Usually two anal cells (anal loops) present, anterior cell (1a) and posterior

cell (2a), delimited by 1A, 2A and 3A. These veins diverging basad and then

fusing distad. However, sometimes one or both anal cells absent. Anterior cell

sometimes open with 1A terminating in posterior margin independently. When both

cells absent, 2A and 3A apparently fused with 1A terminating in hind margin

independently.

Posterior wing: Sc and R often thick and closely appressed. In males Fl

present and F2 usually absent, R4 apparently fused with R5. In Trichoptera it

is difficult to determine the location of R5 when it is not isolated.? In

Lepidostomatidae it usually appears to be fused with R4 above the nigma. This

*However, Betten (1934) and Morse (1975) have taken the position that the nigma

always identifies F2; hence, when the nigma is evident, R5 is always posterior

of it!

8 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

is suggested by the observation that R4 and R5 are obviously fused in male hind

wings of Lepidostoma (M.) vernale (f. 126) and in L. (N.) unicolor and L. (N.)

acarolum (f. 47, 50). M usually with Ma and Mp each unbranched and F3 and

F4 absent. In male Cul usually unbranched, hence without F5; 1A, 2A and 3A

usually terminating independently in posterior margin, but 1A and 2A often

partially fused and then diverging distad, or they may be completely fused.

Abdomen. In lepidostomatids, abdominal hemobrachial apparatus well devel-

oped and composed of simple filaments about as long as segment | (Schmid 1980).

The scent glands of sternite V are absent in both sexes, being one of the most

outstanding derived characters of the Lepidostomatidae.

Male Genitalia

Review. Important contributions on homologies of Trichoptera male genitalia

have been provided by Morse (1975), Mosely (1939a), Nielsen (1957), Ross (1944,

1996) ,cand Schmid. (1970; 1980). However, because all of these workers use

different systems of morphological nomenclature, it is necessary to clarify the

terms used herein.

Segment VII. Theliopsyche with depressed rounded midventral process (f.

19, 23). Lepidostoma (Mormomyia) males with pair of dorsal setose warts.

Segment VIII. Lepidostomatids without sternal process. Lepidostoma

(Mormomyia) males with pair of dorsal patches of major setae.

Segment IX. Completely sclerotized ring. In Lepidostoma (L.) with pair

of sclerotized straps of phallocrypt extending from posterior lateral margins near

base of segment X to articulate with base of phallus and basal plate of inferior

appendages (f. 176). Lepidostoma (Mormomyia) males (f. 127) and females (f.

142) with pair of dorsal patches of major setae.

Segment X. Fused with dorsum of segment IX. Preanal appendages absent

or expressed only as small dorsal setose lobes. Main body consisting of paired

dorsal superior processes, usually fused together and sometimes with a pair of

lateral processes. Intermediate appendages apparently absent, but possibly

present as lateral processes in Theliopsyche (f. 19), several species of

Lepidostoma (L.) (f. 179), and the canthum group of LL. (Neodinarthrum).. Anus

directly below segment X, in L. (Mormomyia) opens from large membranous lobe.

Phallus. Above inferior appendages, with its base situated in membranous

phallocrypt. Phallocrypt outer circumference attached to posterior edges of

segment IX, base of segment X, and basal plate of inferior appendages.

Phallocrypt near its center united with circumference of cylindrical phallus.

Resulting ental portion of phallus defines phallic apodeme. The portion of phallus

inside the phallocrypt is the phallobase. Sclerotized phallocrypt immediately

surrounding phallus defines phallic shield. In Theliopsyche phallic shield heavily

sclerotized obscuring phallobase from view (f. 19), in L. (Mormomyia) semi-

spherical (f. 127), in L. (Neodinarthrum) spoon-shaped (f. 160), in L. (Nosopus)

collar-shaped with extended dorsum (f. 60), and in Lepidostoma (L.) ring-shaped

and attached to lateral sclerotized strips of phallic shield (f. 176). Membranous

endotheca attached to apical portion of phallobase. Paired parameres superior,

usually heavily sclerotized spiniform rods (f. 19, 160), but occasionally short (f.

598, 129) or apparently reduced to flexible membranous lobes (f. 71). However,

in Lepidostoma (L.) and Theliopsyche (except T. corona) parameres absent.

In L. (Mormomyia) parameres and phalicata often asymmetrical, and base of heavily

sclerotized phalicata situated in bulbous endotheca. Phalicata short and straight

in Theliopsyche, but in Lepidostoma strongly curved at base, being somewhat

semicircular. Ejaculatory duct entering phallus through phallic apodeme,

extending along its axis, and terminating in phallotremal sclerite in dorsomesal

subapical concavity of endophallic membranes.

Inferior appendages. Very setose, usually short and_= stout in

Theliopsyche, but more long and slender in Lepidostoma. First article with four

Weaver: North American Lepidostomatidae 9

distinct processes, some of which may be absent or reduced: (1) second article

Inconspicueus lobe -({. »129) er obvious, «subapical: mesal process -(f.. 179); 02)

basodorsal process usually vertical at basal position on first article; in

Theliopsyche usually short and thumblike (f. 19); but in 7. grisea and tallapoosa

long (f. 21); in Lepidostoma often capitate, usually long, slender (f. 179) or

sometimes: reduced -(f..68); in Lb. (Nosopyus) medestam: group (@. 71) slender,

capitate, horizontal and about as long as first article; (3) horizontal process in

Lepidostoma (L.) (f. 179) a basomesal process, apparently functioning as phallic

guide;in: Ls €(Mormomyia) «(f.. 127). ands most) species of .L. (Nesopus)-(. 66) a

middorsal process of first article, apparently aiding in clasping female, but absent

in Theliopsyche, the modestum group of L. (Nosopus), and L. (Neodinarthrum);

(4) ventromesal ridge broad with apical angle in Theliopsyche (f. 19) and several

species of Lepidostoma (f. 158, 179).

Female Genitalia

Review. Detailed investigations of female Trichoptera morphology have been

made by Nielsen (1980) and Unzicker (1968). Other important points regarding

caddisfly. female genitatia have been mentioned by Schmid (1980) and Matsuda

(1976).

General characters. In Lepidostomatidae the female genitalia are typical of

Integripalpia being blunt, without the "functional ovipositor"” (tubulus) typical

of Annulipalpia, without long slender anterior apophysis of segments VIII and IX,

cerci absent, rectum and common oviduct separate (hence cloaca absent), common

oviduct opening between segments VIII and IX, rectum above common oviduct,

opening within segments |IX+X.

Segment VI and VII. Circular sagittally, without sternal processes, pleura

unsclerotized.

Segment VIII. Tergite curved sagittally, well developed, extending over

pleuron. Sternite (ventral plate) flattened, slightly reduced, forming concave

surface (egg pouch) for temporary retention of eggmass; in Theliopsyche with

posterior margin bearing medial and lateral pair of lobes, in L. (Neodinarthrum)

(f. 167) chalice-shaped, in L. (Nosopus) modestum group U-shaped with posterior

margin emarginated, and in L. (Nosopus) and L. (Mormomyia) with several lon-

gitudinal striations indicating flexibility.

Segment |XX. Tectiform (roof-shaped) with two blunt anterolateral

apodemes. Segment X inconspicuous, reduced to posterior pair of small broad

lobes, almost completely integrated with segment IX.

Spermathecal sclerite. Also referred to as "appareil vaginal" (Schmid 1980)

or "processus spermathecae” (Nielsen 1980), heavily sclerotized internal process,

constricting posterior end of spermathecal duct, between spermatheca and common

oviduct, possibly regulating sperm discharge from spermatheca or sperm intake

from male phallus via common oviduct. In ventral view somewhat oval, bell-

shaped, triangular, or pentagonal; in lateral view trapezoidal. With lateral pair

of bands fusing ventrally, in ventral view sometimes with lateral lobes. Also with

anterodorsal process and posteroventral process. In ventral view anterodorsal

process broad and posteroventral process narrow, long, medial, keyhole-shaped,

bearing spermathecal duct opening posteriorly. In lateral view anterodorsal

process cup-shaped and posteroventral process forming ventral keel with lateral

bands extending below.

Spermatheca. Spermatheca of Lepidostoma (L.) bearing many conspicuous

microtrichia which usually occur in small clusters anteriorly, forming reticulate

anterior pattern and being a good diagnostic character of species. Also, in this

subgenus opening of spermathecal sclerite into posterior duct, often with valve

bearing several spiniform processes. In other North American lepidostomatids

spermatheca bearing less conspicuous microtrichia.

10 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

Keys to North American Lepidostomatidae Genera

Larva

la. Length of postmentum (ventral apotome) less than half that of midventral

ecdysial line of adjacent margins of genae (f. 7); case made of stones (f.3);

PATO Gt a a eae yee ac Theliopsyche, p. 10.

lb. Length of postmentum subequal to or greater than that of midventral ecdysial

line of genae (f. 8, 9); case made of stones or of various plant fragments

Phi heG) te whdeseread.s 6 sG.4 Gena wc ee Lepidoestomad).ps 15:

Adult

la. Head dorsum with pair of setose warts oval, aligned with occipital margin,

and small (f. 15); male with sternal process on abdominal segment VII (Cf.

19); female posterior margin of sternite VII! with medial and lateral pair of

gmail tubercles: Cf. 124): eastemhe ane. iGuimeree es eee ws Theliopsyche, p. 10.

1b. Setose warts somewhat teardrop-shaped or oval, not aligned with occipital

margin, large, occupying most of area between the eyes and occipital margin

(f. 28, 124, 151, 169); male without a sternal process on abdominal segment

Vil; female without two pairs of tubercles on the posterior margin of sternite

VLE eho: idee reed acaba sins duuwiee cae Lepidostoma, p. 15.

Theliopsyche Banks

Theliopsyche Banks 1911:356, type species: parva, monobasic; Unzicker 1968:

by 44, 19-20,0 32, 48, 49,0 fs 22.(femaled: Fischer: 1970:92-93; Wiggins

19th; 4045-156, fin 8. 2A-E (larva); »Pennak :19078:610;° Wiggins 1978: 179

(larvay pupady tnzieker et aly. 196279, 19,<9).104-9.107,-f 8.257 (larva);

Wiggins 1984:306; Morse and Holzenthal 1984:324, f. 17.104.

Quisilo Milne 1936:118, 124, type species: Silo griseus Hagen, original desig-

nation.

Aopsyche Ross 1938a:174, type species: corona, original designation.

Etymology: Greek, thelion, little nipple; psyche, butterfly; gender feminine.

Body. Smaller than the majority of lepidostomatids, forewing approximately

9-6 mm long, color dark brown.

Head. Elongate pair of oval posterior warts aligned with occipital margin

(f. 14, 15). Antennal scape longer than head, not highly modified, cylindrical

and occasionally bearing several mesal bristles. Male maxillary palps always

two-segmented with each segment distinct and cylindrical. Female maxillary palps

five-segmented.

Wings. Little sexual dimorphism in wings (f. 16-18). Venation more complete

than that of most other lepidostomatids. Forewings with Ma branched in both

sexes and arculus with normal distal configuration of veins. Slight sexual

dimorphism exhibited in forewing of 7. corona and in the hindwing of T. melas.

Venation usually differing between sexes only in posterior wing in which R4*5

unbranched (F2 absent) in males, but branched in females (typical of

Lepidostomatidae). Posterior wing with discal cell open in both sexes.

Legs. Tibial spur count 2, 4, 4 on front, middle and hind legs, respectively

ffs Op

Abdomen. Males distinguishable by having paddle-shaped sternal process

on abdominal segment VII (f. 19, 23) (not on segment VIII as reported by Ross

[1944, 1946] and Schmid [1980]).

Weaver: North American Lepidostomatidae 11

Male genitalia. Segment IX (f. 19) long in lateral view with anterior margin

of pleuron projecting cephalad and posterior margin nearly straight, pleuron at

least two times as long as venter, dorsum reduced. Segment X usually about as

long as inferior appendages, lateral processes usually prominent and spiniform

(reduced in 7. grisea). Preanal and intermediate appendages apparently absent.

Phallus straight, phallotremal sclerite large, parameres present only in one spe-

cies, T. corona; phallobase obscured by heavily sclerotized phallic shield. Each

inferior appendage with (1) first article usually longer than basodorsal process

(except in 7. grisea and tallapoosa); (2) small apicomesal point of first article

possibly rudiment of second article (The first article is a midprocess, recognized

by having an apicomesal toothlike process.); (3) basodorsal process usually short

and thumblike, but sometimes long and _ fingerlike or acuminate; and (4)

ventromesal ridge usually extending along basal half of main article.

Female genitalia. Posterior margin of sternite VII with two pairs of short

points (f. 24). Spermathecal sclerite with long pointed anterior process in ventral

view.

Immatures, case and habitat. Larva with postmentum about half as long as

midventral ecdysial line. Case cylindrical and made of stones. | have collected

adults of T. grisea twice, on both occasions near a spring. Wiggins (1977)

mentioned that the larvae of T. melas were found in a small spring, also. It seems

likely that most species of this genus are endemic to mountain springs, but Harris

(1986) has collected adults of T. tallapoosa near a stream.

Distribution. This genus is distributed throughout the Appalachian Moun-

tains from Quebec to Alabama. Few records are available, indicating that most

species are somewhat uncommon.

Key to Adults of Theliopsyche Species

(males)

la. Parameres present (f. 19); Mountains of North Carolina and Tennessee

ee EMO Maa U CR V URE tren Brit. tice? Sie sar Aan 4 Emi Gs.? Oe: Beh de PE corofia’ Ross,©p.. 12.

to. Parameret: ebeent (horGOyeiy nC Barely OA ee, a ee ot 2

2a. Dorsal process of each inferior appendage greatly extended posteriorly

peyond maincboudy: (6) ieiiien, Phe eae Bie) seme ARES ia) 3

2b. Dorsal process of each inferior appendage short, not extended posteriorly

bevyorid 106 mal Hodes ZOOPLA PO oe Le ls ee ee ee 4

3a. Lateral processes of segment X shorter than main body; dorsal process of

inferior appendages fingerlike (f. 21); mountains of New Hampshire to South

Car es er i PE ce er Pe Oe ea alga grisea (Hagen), p. 13.

3b. Lateral processes of segment X longer than main body; dorsal process of

interior appendage acuminate (Harris 1986: f. 3); Alabama ................

SF aig ip Mea alae ae GR RRR HORS A AN cs Al Rn ate oR TC tallapoosa Harris, p. 15.

4a. Lateral processes of abdominal segment X sinuate (f. 22); posterior wing

broad, with many stout setae (f. 18); mountains of Kentucky, Tennessee,

ard: Mig aie AL balay pee. eee iad | Lanes melas Edwards, p. 14.

4b. Lateral processes of segment X straight (f. 20); posterior wings oval, with

normal: setNGis os parole ere EASED BOATS 2 SO eS PH EET OS 5

Sa. Mesal processes of segment X almost completely fused and long, each about

six times as long as basal width; ventral process of each inferior appendage

longer than basodorsal process (f. 20); mountains of North Carolina and

PPAR RO 6c le SA re OS Ce yo we ae epilonis Ross, p. 12.

12 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

5b. Mesal processes of segment X fused together about 0.6 times their length

and short, each about 2.5 times as long as basal width; ventral process of

each inferior appendage shorter than basodorsal process (f. 23);

Massachusetts: New York, and »Quebee (ioc)... cee. parva Banks, p. 14.

Theliopsyche corona Ross

(#10, 14-16,-19 24)

Theliopsyche corona Ross 1938a:174, f. 116 (male); Wray 1967:57; Fischer

1970:94; Etnier and Schuster 1979:20; Webb 1980:98; Unzicker et al.

1982:9.106, 9.107.

Male. Head (f. 14, 15). Scape, cylindrical, approximately 0.5 mm long,

with several long mesal setae. Maxillary palps, with two distinct cylindrical

segments, approximately 1.1 mm long (about twice as long as in other species of

the genus). Wings (f. 16). Anterior wing with specialized region between C and

R bearing many modified setae (f. 10); C very thick, Sc apparently branched

distad, Rl atrophied before reaching the margin, basal stems of R2, R3, R4 close

together. Hindwing normal for the genus. Genitalia (f. 19). Segment |X with

unusual sclerotized pattern on dorsum. Segment X with rounded dorsum and

lateral processes spiniform and long. Phallus with parameres present (absent in

all other species of the genus). Inferior appendage with basodorsal process short

and thumblike, main article extended about as far as lateral processes of segment

xX, ventromesal ridge angled with blunt apex.

Female. Antennal scape about 0.5 mm long. Wings (f. 16) normal for the

family. Anterior wing with nigma and thyridium, length approximately 6 mm.

Genitalia (f. 24) having spermathecal sclerite short and posterior margin of

sternite VIII with mesal points separated by wide square notch.

Immatures, case and habitat. Unknown.

Life history. Recorded flight dates: 2/7 May - 2 July.

Distribution. Only Known from the Great Smoky Mountains National Park of

North Carolina and Tennessee.

Type data. Ross (1938a): “Holotype, male. - Gatlinburg, Tennessee: May

27, 1934, along branch of Little Pigeon River, T. H. Frison." Holotype at the

INHS (Webb 1980). See type material examined below,

Material examined. Allotype female, (INHS). NORTH CAROLINA: ~ Great

Smoky Mts Natl Pk, Wild Cherry Branch, male, 8 VI 1949, WER, (INHS); Andrews

Bald,-“@ «males females 1e°WP1647, “CPAP (UGNM) CO TENNESSEE: | Gatlinburg,

Great Smoky Mts Natl Pk, Peach Gap, male, 2 VII 1947, RH Whittaker, (USNM).

Theliopsyche epilonis Ross

(FL! 20225)

Theliopsyche epilone Ross 1938a:173-174, f. 115 (male); Wray 1967:57; Fischer

1970:93; Etnier and Schuster 1979:20; Webb 1980:98; Unzicker et al.

4982: 9.106; 98107,

Male. Scape 0.6 mm long, cylindrical. Maxillary palps 0.8 mm long. Wings

similar to those of T. parva. Genitalia (f. 20) with segment X having mesal

process in dorsal view straight, slightly constricted basally, about six times as

long a basal width, with two short apical humps; and lateral process spiniform,

nearly straight, almost as long as mesal process. Phallus straight, without

parameres, phalicata subequal to length of segment X. Inferior appendages with

basodorsal process in lateral view small, thumblike, about half as long as main

article which is slightly curved.

Weaver: North American Lepidostomatidae 13

Female. Genitalia (f. 25) with spermathecal sclerite in ventral view similar

to that of T. parva and corona, but unique in having keyhole-shaped process

shorter, about half as long as its length.

Immatures, case and habitat. Unknown.

Life history. Recorded flight date: 13 June.

Distribution. Great Smoky Mountains National Park, North Carolina and

Tennessee.

Type data. Ross (1938a): "Holotype, male. - Newfound Gap, North

Carolina: June 13, 1935, along Little Pigeon River, H. H. Ross. Paratypes. -

North Carolina. - Same data as for holotype, 3 males." Holotype at the INHS

(Webb 1980). See type material examined below.

Material examined. Holotype male and allotype female, (INHS).

Theliopsyche grisea (Hagen)

(ft 2a)

Silo griseus Hagen 1861:293 (male).

Theliopsyche grisea: McConnochie and Likens 1969:150; Fischer 1970:93; Schmid

1980:150, f. 541, 544-550 (male, female); Parker and Voshell 1981:6;

Unzicker et al. 1982:9.106, 9.107.

Male. Scape 0.5 mm long. Maxillary palp 0.5 mm long. Wings similar to

T. parva. Genitalia (f. 21). Segment IX in lateral view similar to 7. corona,

but without dorsal pattern. Segment X reduced with slender lateral processes

(sometimes reduced or apparently broken). Phallus without parameres; phalicata

about as long as segment X, with phallotremal sclerite large, about 0.7 times as

long as phalicata. Inferior appendages each with dorsal process long, about twice

as long as segment X and extended far beyond other processes of main article.

This species and T. tallapoosa are the only species in the genus for which the

dorsal process is the longest process of the inferior appendage.

Female. Genitalia (Schmid 1980: f. 550) similar to T. corona and parva in

having a short triangular spermathecal sclerite. Posterior margin of sternite VIII

with mesal points separated by a wide, square notch wider than the length of one

point.

Immatures, case and habitat. Unknown.

Life history. Recorded flight dates: 19 May - 14 August.

Distribution. Known from the Appalachian Mountains of New Hampshire, New

Jersey, New York, North Carolina, Pennsylvania, South Carolina, Tennessee, and

Virginia.

Type data. Hagen (1861): "Hab. Trenton Falls (Osten Sacken)."

Silo griseus Hagen, New Lectotype, male: Trenton Falls, New Jersey, 1854,

O. Sacken. MCZC Type 10951. See type material examined below.

Material examined. Lectotype male, (MCZC). NEW HAMPSHIRE: Grafton

Co: West Thornton, Hubbard Brook, 4 males, K McConnochie, (USNM). NORTH

CAROLINA: Swaine Co: Great Smoky Mts Natl Pk, Juneywhank Falls, Deep Cr

Cpg, male, 21 V 1970, OSF, (USNM). SOUTH CAROLINA: Oconee Co: Sumter

Natl Forest, Yellow Branch, 2 _ males, Ws wh BOF OSFy) v) CUS:

. PENNSYLVANIA: Cumberland Co: Mount Holly Spring, Whisky Spring, 1 VI

1977, JSW, (JSW). TENNESSEE: Blount Co: Great Smoky Mts Natl Pk, small

spring at camp 13 near Sheep-pen Gap, male, 13-14 VIII 1982, JSW & PJS, (JSW).

VIRGINIA: Smyth Co: small stream N of White Top Mt, at Hwy 600, male, 10

Vit. 1980, SWH) CUCEi

14 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

Theliopsyche melas Edwards

Cheep 918) 22)

Theliopsyche melas Edwards 1956:16-17, pl. 1 f. 4-5 (male, female); Fischer

19/0793; Resh 1975:15; “Wiagins: 1977) 1585" ft; -8.2A-E* Clarvay® Etnier and

Schuster 1979:20; Parker and Voshell 1981:6; Unzicker et al. 1982:9.107.

Male. Scape cylindrical, 0.6 mm long. Maxillary palps 0.8 mm long.

Anterior wing similar to that of 7. parva. Posterior wing (f. 18) slightly trun-

cated apically, broad, with many modified, stout setae. Genitalia (f. 22). Seg-

ment X with mesal processes about 2.5 times as long as broad in dorsal view,

apices attenuated and widely separated by long pointed notch; lateral processes

sinuate, spiniform, and long, extended posteriorly as far as mesal processes.

Phallus long, extended posteriorly as far as segment X, without parameres.

Inferior appendages in lateral view each with short thumblike basodorsal process,

main article extension about two times as long as basodorsal process, ventromesal

ridge broad, freely extended posteriorly.

Female. Genitalia (Edwards 1956: f. 5) having spermathecal sclerite long

and narrow, similar to that of 7. epilonis, but apparently with longer keyhole-

shaped medial process, greater than half length of sclerite.

Immatures, case and habitat. Larva (Wiggins 1977: f. 8.2), inhabiting a

small spring; head ventral (f. 7).

Life history. Larvae collected 14-15 May. Recorded flight dates: 19 April

- 28 June.

Distribution. Kentucky, Tennessee, and Virginia.

Type data. Edwards (1956): "Holotype, male. - Wellington Mills, four miles

northwest of Winchester, Franklin County, Tennessee, on state highway 50, and

one and one-half miles north of this highway on a rural road. April 19, 1955.

Preserved in alcohol. S. W. Edwards. Allotype female. - Same data as holotype.

Paratypes. - Same data as holotype. Four males, three females. Disposition of

the type material. - Holotype and allotype deposited in the U.S. National Museum;

paratypes to Illinois Natural History Survey, Vanderbilt University Entomological

Collection, and the author's collection." See type material examined below.

Material examined. Paratypes, two males, (INHS). TENNESSEE: Franklin

Co: 4 mi W of Wellington Mills, Hwy 50, 1.5 mi N of Winchester, 3 larvae, 14-15

V°1970; GBW Ss: TY,2(ROME)> Putnam Cor’ tst spring outflow;* Hwy? 84,1 mi. S

of Hwy 7ON,» 3: males 6 Vi 1972, DAE: & OC Saylor, CUSNM).

Theliopsyche parva Banks

CR RTS: 239 28)

Theliopsyche parva Banks 1911:356, pl. 12 f. 16 (male); Fischer 1970:93-94; Roy

| and Harper 1975:1085; Roy and Harper 1979:163; Schmid 1980:150.

Male. Scape 0.4 mm long, cylindrical. Maxillary palps 0.3 mm long, two-

segmented, each segment cylindrical. Wings (f. 17). Anterior wing 4.2 mm long,

venation typical for the genus, posterior wing also typical. Genitalia (f. 23).

Segment X having mesal process somewhat rectangular with Y-shaped mesal notch

in dorsal view, with dorsum and venter curved and with obtuse mesal notch and

blunt subapical process in lateral view; lateral processes straight, spiniform,

extending beyond apex of mesal process. Phallus straight, without parameres,

phalicata about as long as lateral processes of segment X. Inferior appendages

each with basodorsal process thumblike, short, less than half as long as extended

main article; ventromesal ridge crooked, with small apical point.

Female. Forewing with venation similar to that of male. Hindwing (f. 17)

with more complete venation than in male. Genitalia (f. 26) similar to those of

T. corona and grisea with short triangular spermathecal sclerite in ventral view;

Weaver: North American Lepidostomatidae 15

however, posterior margin of sternite VIII with mesal points close together with

narrow V-shaped mesal notch.

Immatures, case and habitat. Unknown.

Life history. Recorded flight dates: 1 June - 24 July.

Distribution. Massachusetts, New York, and Quebec.

Type data. Banks (1911): ‘From Woodworth's Lake, Fulton County, New

York, 23 June (Alexander)."

Theliopsyche parva Banks, New Lectotypes: There are four syntypes at the

MCZC, Type 11710, of which one has been selected as Lectotype male and another

as Lectoallotype female by Ross in 1937, but unpublished (Ross 1938b). Both of

these specimens bear Ross's yellow lectotype labels and | herein recognize them

as he intended. See type material examined below.

Material examined. Lectotype male and lectoallotype female, (MCZC).

MASSACHUSETTS: Franklin Co: Sunderland Fish Hatchery, male, 2 females, 1

Vi.1954,. OSF~.CINHS)-

Theliopsyche tallapoosa Harris

Theliopsyche tallapoosa Harris 1986:34-36, f. 3A-G (male, female).

This species and 7. grisea are the only species in the genus having male

inferior appendages with dorsal process longer than the medial extension of the

first article. It differs from T. grisea by having male, basodorsal process of the

inferior appendage acuminate, rather than. being fingerlike, and segment X with

lateral processes longer than its main body. The following diagnosis is based on

the original description by Harris (1986).

Male. Length 5.2 mm long. Color brown. Head with scape 0.4 mm long,

maxillary palp 0.5 mm long. Genitalia: Segment X about half as long as inferior

appendages, bearing lateral pair of spiniform processes longer than remainder of

segment X and each process broadest mesad; main body comprising two trapezoidal

plates, their basomesal halves fused together, and with blunt, lateral apices

separated by acute mesal notch. Inferior appendages with dorsal process

acuminate, and longer and broader than medial extension of first article. Phallus

cylindrical and without parameres.

Female. Posterior margin of sternite VIII with mesal points broadly sepa-

rated. Spermathecal sclerite fork-shaped having central keyhole-shaped process

as long as lateral flanges which diverge caudad from its anterior at acute angles.

Immatures, case and habitat. Immatures unknown. Harris (1986) collected

this species near a swift (6 m wide) stream, with rocky bottom.

Life history. Adults collected 19 May.

Distribution. Piedmont region of Alabama. Only known from the type

locality.

Type data. Harris (1986): "Holotype male. - Alabama, Tallapoosa County,

Timbergut Creek, downstream crossing Hwy. 22, 19 May 1984, S. Harris and P.

Lago (USNM). Paratypes. - Alabama, same as above, 1 male, 1 female (USNM)."

Lepidostoma Rambur

Mormonia Curtis 1834:215 (preoccupied), type species: gracilicornis Curtis, syn.

of Phryganea hirta Fabricius (selected by Fischer 1970).

Lepidostoma Rambur 1842:493, type species: squamulosum- Rambur, syn. of

Phryganea hirta Fabricius (selected by Ross 1944); Flint and Wiggins

1961:279-281 (vernalis group); Denning 1962:36-37, 39 (Atomyiodes

syn.);. Flint),1967a123-24:.Unzieker 1968:11,° I4..19>. 32, 50, 51;~Fiseher

1970:33-37; Wallace and Sherberger 1972:222, 226-228 (vernalis group);

Fischer 1973:162;.. Hilsenhoff 1975:24, ££. 82,.c83<¢ Clarva);. Anderson

16 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

19/6:116-118, f. 19 (larva); Solomon and Leiser 1977:166, 172, 178: Flint

and “Bueno. 19%7:315-376>°: Wiggins: “1970: 13 “S48 156, $158) f. 8 TA=H,

(larva); Wiggins 1978:179; Pennak 1978:610; Schmid 1980:147, 148, 149,

100; “MeCatieriv. > 19810264" Hilsenhoff: “I98hi26ywt: -83,°%64 * (larva);

Lafontaine: 198hcs¥, 147) 2180," 195).:° 20830 248-252, 327; “Dudley and

Anderson. 1982°9,. 102 Wnztekér etial.:'196279 6) °9.6A, 9.99.10; 9.19,

Di 28y Bete Os 104 Oe FOU oo Feo O26 “Clakvay ih Walte < ard MeCafferty

1983b:16; Wiggins 1984:306; Morse and Holzenthal 1984:324, f. 17.103.

Pristosilo Banks 1899:212, type species: canadensis Banks, syn. of Mormomyia

togata Hagen (monobasic).

Notiopsyche Banks 1905:216, type species: /atipennis Banks (original desig-

nation).

Oligopsyche Carpenter 1933:36, type species: Notiopsyche carolina Banks (original

designation).

Neuropsyche Carpenter 1933:38, (preoccupied), type species: tibialis Carpenter

(original designation).

Flura Milne 1936:118, 124 (new name for Neuropsyche Carpenter 1933), type

species: Neuropsyche tibialis Carpenter (replacement).

Etymology: Greek, lepis, scale; stoma, mouth; gender neuter.

Similar to Ross (1938b, 1944) | define the genus Lepidostoma rather broadly.

In respect to the Old World genera, Lepidostoma does not appear to be

monophyletic. Only a revision of the world Lepidostomatidae will provide an

accurate assessment of this genus.

Key to Larvae of Lepidostoma Species or Species Groups?’

(final instars only)

la. Case made of small stones (f. 1-3), or head sometimes with carina .... 2

lb. Case made of various plant fragments (f. 4-6), head without carina .. 13

2a. Case linear (f. 2), head without carina; eastern North America; L.

EMOPTRONIIIE i OI GUUS Sie a a De BES he 3

2b. Case curved slightly posteriad (f. 1, 3), or head with carina; western ..

MRE Mares e ie unite CONN 1 bc Lae Foner ota SE Sie ee Ae a OM 0 oot 2 aaa 5

Sat PE HOMEES WISEORISiA Mannie vols iy, aR i tM) ei bum I RE6SSs fp. 68,

ob.” “Appalachian aMoontairs Ohio atieeiiiciaiee . ch ol. Soke he DIS AL ee. 4

CEPR MTA UR ails SIRE Tice eu L. (M.) excavatum Flint & Wiggins, p. 6/7.

4 hahenaatoiMatsachueerts forvitrginian war. PaGe ees. Ua PN ee...

9a. Case slender, tapered posteriad and made of fine sand grains, posterior

wicta: tess'(thaaih, Onn t ado ehead avithowrt carina i .iseiuscbede oh on.

6 I RR WPS WE RATS AAD cath eats AMET ISU) a an ie ae ee LE. (Neodinorthrum); pi. 15.

Sb. Case more robust, not so markedly tapered posteriorly and usually made

of larger stones, posterior width greater than 1.0 mm (f. 3), or head with

Bay HRT MRA SEO SLVR eich oe ob AMEN te a ei aR RN ADD RS ca ue eS 6

*Larvae of L. (N.}) reosum (reosum group) and L. (N.) deceptivum (nomen

dubium) are unknown. Also, larval cases of L. (N.) cinereum and L. (M.)

libum are Known to vary.

6a.

6b.

dae

ib.

8a.

8b.

9a.

Ob.

Yom

10a.

10b.

lla.

11b.

12a.

12).

13a

13b.,

14a.

14b.

laa.

1b:

16a.

16b.

1das

1/7b.

Weaver: North American Lepidostomatidae 17

Rocky Mountains: and-Racifieveoastal-recions' wali iwoeyl aw iule i. | ri

Arizona and southern Colorado to Panama; L. (Nosopus) mexicanum group

Ee Re ee a ate Sam ee eT Pen Neat rie Par hom po tr ie Lc nis Gal fy een mea 19)

Withuebdominal ills: sy Wwyesneanien. « dmand el bea Alt eed amie be ubiey hdl cia Ne os 8

Without abdominal@ils; -GeuiNascpus) \hood bignotum anti acti ine ion 9

Metanotal sal and sa2 sclerites large, oval, each with 15-20 setae ........

aiscituete OE oy he Ri te RRS te 2 ak a ek tl a ae L. (N.) cascadense (Milne), p. 57.

Mesonotal, saland sq2. sclerites):smalil, -round,,each iwithc1-2) setagww..'. ...'.

nent Ge ake 9 ga SN, merce aaa Ae L. (Nosopus) astaneum group, p. 25.

Northwestern, Oregon) to Binitish: <Golumlbbiaies ax voniueaieiie ware eves epi 8 oiled

Boe enn ae ey ante Pate eh OMnoR Tate ee cura oma Wen toot NedePOegl Ress. . 56.

North, centwal. Recky Mountains: .Albentarte: Otatiin ithe ee ai Meee Bi Ll

OYE ERE hres ae ML Te ae Pm Ns | Ln) (Nas picatum) Denning;.p:...06:

Northern Sterra: Nevada: Mountains sof -Galitormias accdaweeg obs ww eit oo

site ach 8 CRhSba 6 ugh Sa beg Ae Te aie SO ses Me Ten li Ue L. (N.) ermanae Weaver p. 55

Head with weak carina; southern Colorado and Arizona to Panama ........

Sn. a Sean Pear ea tae Darran Bi sa 5 dca i le a Ral Linco IN oo mexticantiumci® . SA.

Head with distinct carina or none at all; Central America to Arizona .....

eT ee ee ee ee ee eee Ne Ree ree hear eerie ART OR 3 CURA anew cr nt 11

eo pe CA ae OCL Sal RE ee CRE Ch Mr oem ren ova MMT ES. ie tien ewe Spoons MN c Meme re Gott: am 12

Head without carina L. (Nosopus) mexicanum group in part, p. 46

Head evlingrieal, (mere flats wee aed o Milioe Wicihouiiiaeeraned aa: ANe cic Clie Sia a i

si pedis ga te in ie I Nay le a a L. (N.) acarolum Denning, p. 47.

Head shaped like WW | army helmet, vertex convex; Costa Rica ..........

hide alti: a Bi tdederolnaab Achaea tau ie WE aig L. (N.) talamancense Flint & Bueno, p. 54.

Case, spiral type (Wiggins 1977: f. 8.1 G); northern California to Oregon

Ke aneicalabe date a + peta aelai cit, i Mees netball AON ea Semel baths} verodum? Rossi... oo,

Case, turret. or panel type LE A> 8). Giese mh les di tun ll a lbiniuemy 14

Case; itunret)tyee (ti. 6)ivbw i Nesonus): umieoler aroupe|,Jjiguliese aul. 15

Cas Gu pane batay Pe Cau eM lied aus hese GL iih clin yeataad oad tlhe sem aM iw clad da atl gs acl atu eb Dutra 19

Head light brown, vertex with pattern of darker spots; eastern, Nova Scotia

and. Quebec. £o,Neorih Ala roll ie eri Gln OI eS BION iste, 5 scheree alls cll eA, Mid iment iM oc gg

ogc pS Sense ah tat) BOO athe inal bo Sta tle a Ua bo (Nw jyifresth AMilive) pip. 42.

Head brown, vertex unicolorous or with lighter spots; widespread, Central

American, western and northeastern Alaska to Maine and Newfoundland

Sin) Dans deh llc ial i a tin SANA rel Aaa et teats AAR Um gs ok 1 a a i iad 16

Mesonotal. .scleritas: with) dark) poasterion amare tinge iid vk dik Mies J gabe bn esea gies i

Mesonotal scleriteswithout.dank. posterior marge «ian earrdiia i wie. 18

Case with several long freely extended transverse plant fragments (f. 5);

Northern Mexied): Arizana.. and Mew MaMa ie bu ddl ea iuumigian burma weal Nr

a iin wal hate Wy Bac Mena ae a alli ota dna a a lla AR Lp t(Nadoknulli Ross, «po. 44.

Case with only short transverse plant fragments (f. 7); Guatemala, Mexico,

Arizona, and ‘southern Colorado) .cdodxuiee Low tN ibeakert: blinks po. 40.

18a.

18b.

19a.

19b.

20a.

20b.

2la.

210:

la.

TBs

2a.

20:

3a.

3b:

Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

Head and notal sclerites light brown, mesonotal sal and sa2 sclerites do

not contrast much with membranous background area; midtibia light brown

with dark distal apex; California to Alaska, Rocky Mountains, Michigan to

Maine and Newfoundland ............. L. (N.) cinereum (Banks), p. 41.

Head and notal sclerites dark brown, mesonotal sal and sa2 sclerites con-

trast well with membranous background area; midtibia unicolorous;

California to British Columbia, Rocky Mountains, South Dakota to Quebec

ERO GaN See Va Vy L. (N.) unicolor (Banks), p. 44.

Length of postmentum subequal to ventral ecdysial line of adjacent genae

(1.28) 2 case Somewnat“round sacittally: eastern... se es

PET Hoy ek ar UPD Celine he ere wea L..(N.) modestum group, p. 35.

Length of postmentum greater than ecdysial line (f. 9); case, round or

SGGTe Sagan y WIiGeSOredd. 6. sides oe eG eee ek ei eee we eee ee 20

Case usually tapering posteriorly and sagittally square; USA and Canada

exclusive of Southwest; endemic to coastal streams of Southeast, lakes and

large streams of North, but exclusive of small springs ...................

SS MELBOL Ware peak Ginn ate Why ace) ca cite We aah u k Gre ROU. a calor ges Lepidostoma (L.), p. 84.

Case usually with lateral sides parallel, round or square sagittally; wide-

spread Nearctic; endemic to springs and small mountain streams ..... 21

Vy OST RR NE AREiG si fool ee Ore ih Re LS GURUS Want SERA D sR OS

Pee aks L. (Nosopus) jewetti, podagrum, roafi groups, p. 27, 31, 33.

Eastern Nearctic, including Georgia to Newfoundland, and Great Lakes

POGIONS Fu Se OR UA Se Lak ee el DOU ES L. (Nosopus)

americanum group and L. (Mormomyia) (except /ibum group), p. 28, 60.

Key to Adults of Lepidostoma Subgenera

Male and female abdominal dorsum |X with paired setose warts; (f. 127, 142);

Sas terhies usally WiOntene Gl Vs Ae Wr 85 Mormomyia, p. 60.

Male and female with normal setae on abdominal dorsum |X; widespread

Male phallus without parameres, phallocrypt with lateral straps (f. 175-179);

female spermatheca with many conspicuous microtrichia, segregated anteriorly

in small clusters, sometimes forming a reticulate pattern (f. 180-184); wide-

spread in eastern coastal plains and northern lake regions but exclusive of

Southwest USA and Central America, occasionally montane ................

PU ae ee Ce Ay ch oe ie ene ye ee ees Lepidostoma, p. 84.

Male phallus with parameres present but sometimes reduced to small lobes

(f. 58, 71), phallocrypt without lateral straps (f. 60, 160); female

spermatheca with inconspicuous microtrichia, not segregated in reticulate

clusters; widespread, usually montane ......... Ri rgeieaal sabes Gi Mo iChat Nea abe ge 3

Male forewing with reflexed anterior margin and unique fold in anal region

which obscures Cu2 and 1A adjacent to arculus (f. 152, 153); female sternite

VIII heavily’ sclerotized with chalice-shaped area (f. 167); western ........

PURINA Whe PA Ee MUR oe ae NP aay SPOR WUT QR NI HURON Glby tiene atl! 8 Neodinarthrum, p. 75.

Male forewing usually without reflexed anterior margin and without such fold

near arculus, Cu2 atrophied (f. 41-53), except in reosum (f. 40), sometimes

with anal groove or medial fold (f. 54-56); female sternite VIII without

sclerotized chalice-shaped area; widespread ............. Nosopus; 2. 12.

Weaver: North American Lepidostomatidae 19

Lepidostoma Subgenus Nosopus McLachlan

Nosopus McLachlan 1871:114-116, type species: podager McLachlan (monobasic).

Olemira Banks 1897:29, type species: americana Banks (monobasic); Flint

1967a:24.

Eremopsyche Banks 1901:367, type species: frontalis Banks (original designation);

Flint 1967a:23 (as syn. of Lepidostoma); Fischer 1970:83.

Atomyia Banks 1905:217, type species: modesta Banks (original designation).

Alepomyia Banks 1908:64, type species: bryanti Banks (original designation).

Atomyiodes Ulmer 1911:23, f. 10-11, type species: bispinosa Ulmer (monobasic);

Fischer 1970:70; Flint and Bueno 1977:382, Atomyiodes bispinosus syn.

of Olemira mexicana Banks.

Alepomyiodes Sibley 1926:106, 190, type species: Lepidostoma wisconsinensis

Vorhies syn. of Alepomyia bryanti Banks (original designation).

Arcadopsyche Banks 1930:129, type species: prominens Banks (monobasic).

Jenortha Milne 1936:119, 124 type species: cascadensis Milne (original desig-

nation).

Etymology: Greek, nosos, disease; pous, foot.

This is the most diverse subgenus of Lepidostoma, presently accounting for

45 species. The majority are restricted to the West, but nine species occur in

the East of which two species L. (N.) cinereum and unicolor are transcontinental.

Male. Head with scape sometimes modified, but often simple and cylindrical.

Maxillary palps often modified with segmentation usually obscured; generally apical

segment spatulate, mesal surface concave bearing several modified petiolate scales;

basal segment cylindrical. Wings with Cu2 of anterior wing usually atrophied (f.

41, 43-53), sometimes joining Cul basally (f. 42), or modified with setose anal

groove or medial fold (f. 54-56), but complete in L. (N.}) reosum (f. 40). FI,

F2, and FS (in L. [N.] jewetti F3) present. Arculus usually modified, distal

submarginal convergence of anal veins (2A, 3A sometimes atrophied) in typical

distal position only in L. (N.) reosum, astaneum and jewetti groups. In the other

species groups of L. (Nosopus), 1A, 2A, 3A converge with crossvein cu-a at

arculus, forming distinct Z-shaped configuration, more basad, usually in middle

of anal region. Number of anal loops variable, sometimes two present, but often

anterior loop absent. Discal cell usually about as long as Fl and thyridial cell

about 1.5 times as long as Fl. Costal cell folding over wing, which varies in size,

exhibited by some species of L. (N.) astaneum, jewetti, americanum and

mexicanum groups. However, many species without costal cell. Hindwing usually

not highly modified. Fl always present, F2 present only in most species of L.

(N.) mexicanum group, F3, F4, and F5 absent. Unique submarginal setose groove

exhibited by L. (N.) jewetti. Genitalia (f. 57-96) having segment X usually about

as high as long in lateral view, bearing one or two apicolateral points. Phallus

symmetrical, phallobase short. Parameres present but variable in size and shape.

Inferior appendages generally similar to those of L. (Mormomyia); first article in

lateral view usually triangular and acuminate, but sometimes long and uniform in

width; second article and horizontal middorsal process of first article sometimes

absent as in L. (N.) jewetti group; basodorsal process variable, usually clavate,

directed dorsally except in L. (N.) modestum group directed posteriorly; hori-

zontal middorsal process parallel to first article, but absent in L. (N.) modestum

group.

Female. Wings typical of female Lepidostomatidae. Genitalia similar to those

of L. (Mormomyia), but dorsum |X lacks setose warts. Spermatheca without

conspicuous microtrichia. Sternite VII! rectangular with posterior margin curved

outward, with several longitudinal striations indicating flexibility and position of

egg pouch. Spermathecal sclerite usually oval in ventral view.

Immatures, case and habitat. Early instar larvae appear to construct stone

cases only, but later instars make panel, turret, spiral, or stone cases (f. 3-6).

Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

Most species of this subgenus are endemic to mountain streams and springs, but

some species also occur in lentic environments.

Distribution. Widespread throughout North America, Panama to Alaska to

Newfoundland to Georgia, especially common in mountainous regions.

la.

Das

2b.

oa.

3b.

Aa.

Ab.

Da.

ob.

6a.

6b.

Tai

Wook

8a.

Key to Adults of L. (Nosopus) Species

(males)

Forewing with Cu2 complete, fused with 1A distally, (f. 40), small, about

4-5 mm long; Maine and New Brunswick to Massachusetts ................

SIRUAD Ns ER Ue MOL A ot Al sa Fa gil Caran Ns kN Ra reosum Denning, p. 24.

Forewing with Cu2 atrophied or apparently fused with Cul or 1A basally

(f. 41-53), or with setose anal groove or medial fold (f. 54-56), about 6-11

UT Gir AACS OTSA Ah Ty, MOO TL, IY cee etait MMe Sie TOMO W ME 2

Forewing with R5 and Ma forming petiolate apical fork and with long setose

groove traversing veins in anal region (f. 55); scape with long fingerlike

mesal process (f.\ 36);. Galifofinia tov Yukon ito: Colorado to’ Montana ov...

SA GOR i Lae eh te Coley iM alten ene Re ese lem an nn RN! cascadense (Milne), p. 5/7.

Forewing never with R5 and Ma forming petiolate apical fork and without

setose anal groove (f. 41-53), but sometimes with short anal groove (f. 56)

or with large fold along M (f. 54); scape without long fingerlike mesal

process, but sometimes modified otherwise (f. 33, 38); widespread .... 3

Pedicel at least twice as long as first segment of flagellum (f. 37-39);

Calitormlaitoveritish Columbia to Montana ie yy Meee yy eee as os 4

Pedicel approximately as long as first segment of flagellum; widespread

Forewing without deep setose fold along M (f. 56); scape bulbous (f. 39);

Calliorniatonwesiiaarom: hele ee ees lial verodum Ross, p. 59.

Forewing with deep setose fold along M (f. 54); scape not bulbous, either

elbow-shaped in lateral view (f. 37) or rectangular in dorsal view (f. 38);

ge clep BC Rs) Tote. MNES UT A RN eee ase eu em oe an 5

Scape elbow-shaped, pedicel without mesal process (f. 37); Oregon to

og) bales Cok UO) VLEs) Cu) Suk OMe DCE Aa am Pe STAM ae SLAP na Hoodl “Ross. ps" 56:

Scape rectangular in dorsal view with expandable mesal bulb, pedicel with

SI Oeh Wiesel DOr OC eee MIN Cy PMR NIM RMN OIE RUE EES eek 6

Segment X in lateral view with length subequal to its height (f. 94); Alberta

LSS UU Cl ohiGc) Uo MPR Ee. UCT SUAS SRL aA ar Ua err cee RRMA Se spicatum Denning, p. 956.

Segment X in lateral view about twice as long as its height (f. 92); northern

ee) VCC. ON Re GEN LALOR a GbE ans Su nea a Pa aga CAR abe SRC ermanae Weaver, p. 59.

Forewing with arculus in its typical distal submarginal position, having anal

veins joining crossvein cu-a adjacent to hind margin, sometimes 2A and 3A

FUSES Vain achem mee T ey Aiea MA ie ge a SER ae es eI SO, 8

Forewing with arculus modified, slightly more basad of its typical distal

position, having anal veins joining crossvein cu-a in middle of anal region

FED Sha PERU EARLE RSAOe EAU RMT ee OD GUISE ROI Sra! aT I SRN asia ES i 11

Forewing without any anal loops, 2A and 3A apparently fused, having 1A

atrophied, hence appearing to have only one complete vein in anal field

reaching arculus (f. 41); parameres blunt, short, about a fourth as long

asvonaticata (ti Ge) SO)e Nestaneunigroup Mee R Ae a 9

8b.

9a.

9b.

10a.

10b.

1 ta:

ITS.

2a.

126.

13a:

Tae.

14a.

14b.

La,

Toe.

16a.

16b.

I bs

Weaver: North American Lepidostomatidae 21

Forewing with anal veins forming one large anal loop by fusing just before

arculus (f. 42, 43); parameres long, about 0.7 times as long as phalicata

CF 60, (84): JewreGet , Guroadee tS BW iid amet dint dietician el | ese eI el Sid a 10

Segment X in lateral view with posterolateral process bearing three dorsal

LO cfs ANGI: CAR ee. Ta ee ane SRE CU ANE EMOTE EI. Cd recinum Denning, p. 26.

Segment X in lateral view with posterolateral rounded apical lobe (f. 58)

Gi IIE Gt oi at a aR NU HEN i a ke ce I astaneum Denning, p. 25.

Inferior appendage in lateral view with long slender horizontal process (f.

60); hindwing with setose submarginal anal groove (f. 42); California to

British Gotu ce ak OG Baie eins ese Lema eres oy a lla jewetti Ross, p. 27.

Inferior appendage in lateral view with short, stout, horizontal process (f.

6i)y hindwing Without setose groove Ci Agi Callitorn ian ali) Powe Ue e

Se eT ee a ee a am Re MeL wee eat hie, stn licolum Denning, p. 27.

Maxillary palps with flexible apical segment bearing many long petiolate

scales; frons concave and vertex projected forward like visor (f. 31);

eastern: IMeriEC QU | Grouisy (eeu Gane. Bian eich UN a ae Ma eat 12

Maxillary palps not flexible, usually spatulate or triangulate with small scales

mesad, frons not concave and vertex not extended forward like visor;

WIGESD MEAG aidiawen hiss Lee Hee ad einai a nani Repl a rN aati aly eA aa nae 2 15

Forewing with broad costal fold, when folded concealing Sc, R, Rs medially,

and Bearinen mah yshortiecatese bu Ua): dnl wlabiens inn aie uu aimiiliieli chet ial oaly 13

Forewing without: broadvcostal fold) (fi Abode n Wien kbs lie duly Bs. 4 14

Segment X in lateral view with two long terminal points, both directed

posténedorsad (.(f: 162) Geer gianitoiQuebecianiunuia win mie wean Wbinliins Gime). CN.

ae aN amr a REPOS EDA DRT a CMM Vil me ty 8) Get np lad eum americanum (Banks), p. 28.

Segment X in lateral view with two short terminal points, dorsal point

directed anterodorsad and ventral point directed posterodorsad (f. 64);

Virginian te- Quebec tor Mimmesotay isi uw widen, costale (Banks), p. 30.

Phalicata bulbous mesially; segment X in lateral view with long curved

ventral process (f. 65); Massachusetts to Quebec to Newfoundland .......

idole say ty yl lRR Renn hc NR le a a eae prominens (Banks), p. 31.

Phalicata not bulbous mesially; segment X in lateral view without long curved

ventral process (f. 63); Georgia and Tennessee to Newfoundland and

Mie sOtae yada dint ave ph hee aha aati lat a eae Ll fee a bryanti (Banks), p. 29.

Forewing with two anal loops (f. 45); western; podagrum group ..... 16

Forewing. withione anal loop. (ft. 46-53) :widesp read. iis is Ii6 eswk oes os ‘7?

Foreleg basotarsus broad, modified, bearing many long petiolate setae

anteriad; labial palps also broadened (f. 29); California to Alberta .......

eh PA ear IN OR WIR is Phase bie Nae mite Ele eat ata Oi HAIN podagrum McLachlan, p. 31.

Foreleg and labial palps unmodified, normal (f. 30); Oregon to British

Co lertribatay Bee Gia eid) at aety mye ln Agata aM bie elect dig c Lh guercinum Ross, p. 31.

[Inferior appendage in lateral view with capitate, basodorsal process at least

as long as and parallel to main article, without horizontal middorsal process;

parameres membranous, lobiform (f. 69-73); eastern; modestum group ...

Inferior appendage in lateral view with basodorsal process directed verti-

cally, shorter than and not parallel to main article, with horizontal middorsal

process; parameres sclerotized, spiniform (f. 74-91); widespread ........

18a.

18b.

19a.

19b.

20a.

20b.

2la.

Z1p..

22a.

226%

23a.

Zan.

Zao:

24b.

23a.

Zob..

26a.

26b.

2TR

2S,

Contribs (Amer. Enteolnst.: Volume 24>: Number 2, 1988

Segment X with long lateral processes (f. 72); South Carolina to

Nir ruled eh Se ee A SEB ey ontario Ross, p. 37

Segment X without lateral processes (f. 71)

Segment X in dorsal view with 3/4 of mesal processes fused basad (f. /0);

Ceara ts ew POunalene a ai wei an EU Tok a lydia Ress,p. 36:

Segment X in dorsal view with half or less of mesal processes fused basad

Fee OU TA) Gary gt chehaah eamnnadaed nay ate mk locale kahuatou Mendhubudlek tn eeaes ne MEU anodes, a 20

Segment X in lateral view tapering gradually to blunt apical process (f. 73);

North Carolina to Newfoundland: .iiir ous ase pictile (Banks), p. 38.

Segment X in lateral view tapering abruptly to slender apical processes (f.

BOF) uid eam Bee Sapte nN RNIN Re ING RUM Sk a ME BU Saf RR hal Big WOES ae, 21

Segment X in dorsal view with apical arms slender, parallel, straight and

directed posteriad (f. 71); North Carolina and Tennessee to Quebec .....

PiAeerc hae See eee EAT AMEN Y See is WMS Seton: LMU ar een hang cea Gage a modestum (Banks), p. 37.

Segment X in dorsal view with apical arms not slender, parallel nor straight,

butibernt with apices. directed. posteriomesadi (iy BOJUR ries, Bok eee. . ye

Segment X in dorsal view with lateral margins of distal arms obliquely angled

mesad; apices pointed and almost touching (f. 69); Tennessee oi. ..0......

eg re ene Ser ner ct aH Ree CN ik Sern. ea ee compressum Etnier & Way, p. 36.

Segment X in dorsal view with lateral margins of distal arms gradually

curved mesad; apices rounded and widely separated (Harris 1986: f. 4);

PABA iii cipal WA di es WO oe ee OE weaveri Harris, p. 39.

Inferior appendage in lateral view with robust thumblike basodorsal process

extended posteriad about as far as main process (f. 74-78); widespread;

a iene Wid Oi eR nerd OP RW AW ene nee wae ek OREO 24

Inferior appendage in lateral view with less robust basodorsal process, not

extended posteriad as far as main process (f. 68, 79-91); western;

SS A FAO CONS). 4 viewers PRES UR WD ik ER Ie 28

Scape with small mesal process, bearing retractable ball within membranous

lobe (f. 33); posterior margin of segment X in lateral view without acute

War, Semana 7a ee ON Geet Ur Geena xh arti Waa WED Soy 8b mcg ckle wee 26

Scape without mesal process (f. 34); posterior margin of segment X in lateral

Via Wat aeU terrae HEC 76 FS he ah. ele OR Re, 2

Posterior margin of segment X in lateral view with acute notch (f. 7/8);

Californiante British Columbia: to: Minnesota: to Quebecwl boo OiWeli es. ce een

Sik He acest Daal bel gig Ae ce SIRS ce Getty EAS RNS Be unicolor (Banks), p. 44.

Posterior margin of segment X in lateral view with square notch (f. 16);

Nerti Carolma te! Nove Seotiacs 480200) wh VG frosti (Milne), p. 43.

Inferior appendage with pointed apical process (f. 74); Colorado to Mexico

COO EN el ek ee ee a ak om bakeri Flint, p. 40.

Inferior appendage with two aligned disk-shaped apical processes (f. 75,

PPB ee kes eRe) dhe dndlanee WONA ARE Ca MUMS CubueMa IK Bye alah aay ian Bias Map ennee fea 27

Segment X in lateral view with oblique dorsal margin, an acute apex and

with many large tubercles (f. 77); northern Mexico, Arizona, and New

COST ESE ARE aa ROMP ne May ON Ome RN EU eae AYE TS ee Varay oh: aOR OY knulli Ross, p. 44.

Segment X in lateral view with curved dorsal margin, small rounded apical

lobe and with few small tubercles (f. 75); California to Alaska to Maine and

HAP RO ies Cae RR ER pee DOO cinereum (Banks), p. 41.

28a.

28b.

29a.

29b.

30a.

ae.”

aoe:

Bib:

Soe:

32b.

S38:

34a.

34b.

35a.

36a.

36b.

3/a.

37b.

38a.

38b.

Weaver: North American Lepidostomatidae 23

Hindwing=with: Ro four-brarched " CFs 49-50) acd veek ORK Sey 29

Hindwing with R five-branched (f. 51-53); mexicanum group (in part) ...

ONE A Cre Umar. Lie maa Nene Ea PT MOEA Ty Tee 251 Syme REN AOY BemEe ANE Da AM et see 33

Segment X in lateral view with two long posterior projections ........ 30

SSNOAT tO ae ee I eae ek age ee Pe I ak yaaa nega 31

Inferior appendage without basodorsal process (Bueno and Contreras 1986:

C...7)) Gakees i aisae Deb od. eae ra dafila Bueno & Contreras, p. 48.

Inferior appendage with basodorsal process (Bueno and Contreras 1986: f.

135 OBMeGe SEP Coe ea oaxacense Bueno & Contreras, p. 52.

Segment X in lateral view with two blunt posterior projections (f. 81);

PRS OS, Pe EA Ee eg te a Rie kaa ri delongi Ross, p. 48.

Segment X in lateral view with only one acute posterior projection (f. 68,

PDI ess ails. Vs amet Dae GP sh Cn ee eee Bs MEE ho ah acer ee ee A 32

Segment X in lateral view with medial apex (f. 68); New Mexico to Alaska

Montana i6 Comradovnumine ean, s Sie he eae roafi (Milne), p. 34.

Segment X in lateral view with ventral apex (f. 79); Arizona ............

CPS RG Fe a ee a ee IE oe es acarolum Denning, p. 47.

Inferior appendage in lateral view with venter truncated apically and narrow

apicodorsal ‘process extending posteriorly (f: 83; 66)> oi. Yoelsieaa. : 34

Inferior appendage in lateral view without venter truncated apically and

narrow apicodorsal process extending posteriorly (f. 82, 85, 87-91) ......

Forewing with narrow anterior margin reflexed over apices of Sc, R1, R2

anc AS f.: Si sinaxillary pales spatelata: Weracrus 162s. oir ake

SiS ORE EE A We a ee ES ee oa eae frontale (Banks), p. 49.

Forewing without reflexed anterior margin; maxillary palps triangular (f.

35); Panama to Arizona anc southwestern: Colorade iin tsar hoods be ee, &.

Inferior appendage in lateral view with serrated subapical ectal flange,

bearing. 2-23 teeth 050, Gal aa Bee BO) icen ued Sow eos kk Be 36

Inferior appendage in lateral view without subapical ectal flange, or with

fiange ret serrated Ch) 825 GP) Saye hea Re Pee ew lee. 40

Subapical flange of inferior appendage semicircular with 6-25 teeth (f. 84,

is EL) Se Perc ENE A Eee ne ie Okan Wms 2 oor et te En Cre merer ee yee ye ee 37

Subapical flange of inferior appendage not semicircular with two or three

(eer its Be Sa aes PO ee Pe ae Ba rete Ried Cee 39

Subapical flange of inferior appendage with six big teeth (f. 85); anterior

wing without costal cell basad (f. 53); Arizona to Durango and Sinaloa

Os PAs Tee eo a eres ear hea OD we: Gat aie Pak ea lacinatum Flint» p. 50.

Subapical flange of inferior appendage with 10-25 small teeth (43084, . 90);

anterior wing witt: Gestal dallvhased 06) B2)025 oss cae es eh logetoor.: 38

Segment X in lateral view small portion of main process hangs below

posteroventral:p rocass (6) BA) «Guatemala «cosine (oe eihawaeih: OBA w oe.

RUN es Ban am Ae Oe ye Monee ete bemte Grol viantet Poe heveli Flint & Bueno, p. 49.

Segment X in lateral view no portion of main process hanging below

posteroventral procese (8) Go)» El Satvetio’ eG te Gide: ewe chi

24 Contrib. Amer, ‘Ent. Iinst:);) Volume’24, Number 2, 1988

39a. Segment X with two dorsal processes in lateral view; inferior appendage

with ‘hatchet-shaped apex (f. 80); Morelos “and Veracruz is... eer. on. Le,

RG LOS AEC SGU COE CR Uiy WERT ac ralne VIAN carci: B Sot MOF ae CO a aztecum Flint & Bueno, p. 47.

39b. Segment X without two dorsal processes in lateral view; inferior appendage

with trumeatedmMapex (fo68) 3 Durargé. sao. rectangulare Flint, p. 53.

40a. Segment X in lateral view with three broad posterior lobes (f. 91); Costa

iCal eh See ed aay talamancense Flint & Bueno, p. 54.

40b. Segment X in lateral view without three rounded posterior lobes (f. 82, 87,

CO VR UC 5 0h eae Na the | sii a ht a ie ps ha etna he Al

4la. Segment: X.with posterionm margin serrated (f:)082)> shoe. geod. eee 42

41b. | Segnient “xX owith posterioromargin mot serrated (f ) 67), 89) oo weiss. 43

42a. Inferior appendage with semicircular subapical ectal flange, parameres broad

STOR) nia as Fe a Sa Le a a denningi Weaver, p. 49.

42b. Inferior appendage without subapical ectal flange, parameres slender (Bueno

and’ Con theras (1966 i 4-6)e Nuevo eons it ee ee ee ee ee

CORI) ESET Sn ahaa Pl a Rudi, leonilae Bueno & Contreras, p. 50.

43a. Segment X with many tubercles; inferior appendage without ventral subapical

flange: Ghr60) s@osta Rica Hes leona reimoseri Flint & Bueno, p. 53.

43b. Segment X with only two or three apical tubercles in lateral view; inferior

appendage with distinct ventral subapical flange in lateral view (f. 87);

Mex O mS Pate Po et Rea tata oc ola ie oa cuifa Baeno 6 Padilla, ip: 52.

Lepidostoma (Nosopus) Reosum Group

This group consists of a single species, L. (N.) reosum, of which only the

adult male is known. Male wing venation unique (f. 40). Forewing with Cu2

running parallel to 1A and both veins fusing distad, forming long narrow cell;

2A and 3A fused medially forming only one anal cell and terminating in posterior

margin together; distal convergence of veins at arculus. similar to that of

Theliopsyche. Hindwing with Fl and F5, M unbranched; 1A and 2A fused basally,

forming small anal cell, and then terminating in posterior margin together.

Lepidostoma (Nosopus) reosum Denning

Che 40). 8x9

Lepidostoma reosa Denning 1954:192, f. 6GA-C (male); Fischer 1970:63; Neves

19 7Sea 74:

Male. Scape not highly modified, cylindrical, 0.3 mm long. Maxillary palp

fingerlike, 0.2 mm long. Wings (f. 40) with anterior wing 4.3 mm long, brown.

Genitalia (f. 57) with segment IX short, dorsum slightly longer than venter in

lateral view. Segment X in lateral view lobe-shaped, basal height about twice its

length, apex slightly extended posteriorly beyond inferior appendages, with three

stout dorsoapical spines; in dorsal view with two straight fingerlike processes

half its length, and pair of short lateral processes basally. Inferior appendages

in lateral view with first article triangular; basodorsal process long, directed

vertically fingerlike; middorsal process horizontal, half as long as basodorsal

process; second article short, subapical mesal process.

Female, immatures, case and habitat. Unknown. However, larvae may

inhabit bog areas where adults have been collected.

Life history. Recorded flight dates: 22 May - 14 June.

Distribution. Maine, Massachusetts, New Brunswick.

Weaver: North American Lepidostomatidae 25

Type data. Denning (1954): "Holotype male, Belchertown, Massachusetts,

May 21, 1938, (from University of Massachusetts Collection)." See type material

examined below.

Material examined. Holotype male, (DGD). MAINE: Kennebec Co: Mt.

Vernon, 2 males, 7 VI 1963, AE Brower, "Flitting like Micropteryx over tiny bog

pools,’ (USNM); Augusta, 2 males, 22 V 1941, AE Brower, (USNM). NEW

BRUNSWICK: . Kent Co: Kouchibouguac Nat! Pk, male, 14 Vi 1978. DB Lyons,

(GNCI).

Lepidostoma (Nosopus) Astaneum Group

This group comprises two species, L. (N.) astaneum and recinum.

Male. Head (f. 27) with scape cylindrical, unmodified. Maxillary palps

apparently two-segmented, segmentation obscured, apical segment slightly longer

than basal segment. Wings (f. 41) having anterior wing with small costal cell;

Fl, \F2,. and F3S present; discal cell long and. narrow; thyridial cell long; 1A and

Cu2 atrophied basally; 2A and 3A apparently fused completely and merging with

crossvein cu-a near hind margin, distal submarginal configuration of veins at

arculus present. Posterior wing with Fl and F5; M unbranched: 1A and 2A

terminating in hind margin separately. Genitalia (f. 58, 59) generally similar to

those of other species of L. (Nosopus). Phallus with parameres short and blunt,

about a fourth as long as phalicata, phallotremal sclerite relatively large. Inferior

appendage in lateral view with first article triangular and acuminate; second

article, subapical mesal lobe pointed posteriorly; basodorsal process directed

vertically, slightly shorter than middorsal process; horizontal middorsal process

parallel to first article.

Female. Genitalia similar to those of many species of L. (Nosopus).

Immatures, case and habitat. The pupae and probable larvae of recinum

were examined. These specimens were collected from spring and temporary pool

areas. Larval cases cylindrical and composed of small stones.

Distribution. Species of this group are known to occur in northern California

and Oregon.

Lepidostoma (Nosopus) astaneum Denning

(hs (Ab, (oe)

Lepidostoma astanea Denning 1954:190-192, f. 5A (male); Fischer 1970:38, 39.

Lepidostoma goedeni Denning 1971:208, f. 6-6B (male) NEW SYNONYM; Anderson

1D (oe 42.

Male. Scape cylindrical, 0.8 mm long. Maxillary palps 0.6 mm long, with

many scales on mesal surface. Wings (f. 41) brown. Anterior wing 6.3 mm long.

Genitalia (f. 58) similar to those of L. (N.) recinum, but having segment X with

posterolateral process a rounded apical lobe in lateral view.

Female, immatures, case and habitat. Unknown.

Life history. Recorded flight dates: 14 June - 24 August.

Distribution. Northwestern California and western Oregon.

Type data. Lepidostoma astanea Denning (1954): "Holotype male, Patrick

Creek, Del Norte County, California, June 24/1951, D. G. Denning.”. ‘See type

material examined below.

Lepidostoma goedeni Denning (1971): "Holotype male, Beverly Beach,

Lincoln County, Oregon, 2 August 1969, blacklight trap, Kenneth Goeden." See

type material examined below.

Material examined. Holotypes of astaneum and goedeni (DGD). OREGON:

Lincoln Co: Flynn Cr, 2)males, 19°VIl 1982%mate, Gy Wid) .1982: 3 malas. 180V1i1

26 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

1982, RWW, (OSUO); Big Elk River, 1 mi N of Harlan, male, 24 VIII 1982, RWW,

(OSUO).

Lepidostoma (Nosopus) recinum Denning

Cr 2b, os, oe

Lepidostoma recina Denning 1954:188-189, ff. 2A-B (male); Fischer 1970:63;

Anderson 19/76:122; Anderson et al. 1982:14.

Lepidostoma calensis Denning 1968:68, f. 6-6B (male) NEW SYNONYM; Anderson

1976:118; Wisseman and Anderson 1984:459.

Male. Head (f. 27) with scape cylindrical, 1.1 mm long, and maxillary palp

0.5 mm long. Anterior wing 6.3 mm long, brown. Genitalia (f. 59) similar to

those of L. (N.) astaneum, but having segment X with posterolateral process

bearing three dorsal points in lateral view.

Female. Genitalia illustrations (f. 97) based on examination of developed

pupa.

immatures, case and habitat. Several pupal cases examined indicate that late

instar larvae construct cylindrical cases of stones approximately 9.5 mm long, 2.0

mm wide and slightly curved. The collection label of pupae examined reads:

"clogging water supply from covered spring." Also, larvae inhabit temporary

pools.

Life history. Larvae collected 19 January to 27 April, prepupae late April.

Recorded flight dates: 29 June - 24 August. |

Distribution. Central California to Oregon.

Type data. Lepidostoma recina Denning (1954): "Holotype male, Peavine

Ridge, McMinnville, Oregon, June 29, 1950, K. M. Fender." See type material

examined below.

Lepidostoma calensis Denning (1968): "Holotype. Male. Strawberry,

Tuolumne County, California, 13 August 1960, D. Q. Cavagnaro. Type to be

deposited in the Academy of Sciences, Golden Gate Park, San Francisco,

California."" See type material examined below.

Material examined. Holotypes of recinum and_ calensis (DGD).

CALIFORNIA: Siskiyou Co: Preston Peak, 2000', camp at bridge, male, AB

Gurney, (USNM). OREGON: Benton Co: Marys Peak, road 12-7-30.1, temporary

pool, larvae & prepupae 27 IV 1983 (7 males & 11 females reared, emerged late

V-early VI), RWW, (OSUO); Lincoln Co: Flynn Cr, larvae? 19 | 1982 (male pupa

reared); male, 19 Vil 1982; 2 males, 24 VIII 1982, RWW, (OSUO); Polk Co: Falls

City, 21 VIII 1967, 8 male pupae, female pupa, (ROME).

Lepidostoma (Nosopus) Jewetti Group

This group contains only two species, L. (N.) jewetti and licolum.

Male. Head (f. 28) with scape cylindrical. Maxillary palps slender and

fingerlike. Wings (f. 42, 43) having anterior wing with distal arculus and small

costal cell basad, only one very long anal cell present. Posterior wing with F1

only, anal veins separate. Genitalia (f. 60, 61) having sternite VIII with a

conspicuous patch of setae. Segment X in lateral view about twice as long as

high, and about as long as inferior appendage. Phallus heavily sclerotized about

0.7 times as long as phalicata. Inferior appendage long with first article long

and fingerlike; second article absent; dorsal process vertical and slender; hori-

zontal middorsal process, parallel to first article. |

Female. Female of only L. (N.) jewetti known, genitalia (f. 98).

Immatures, case and habitat. Final instars of L. (N.) jewetti build panel

cases; immatures collected at springs.

Weaver: North American Lepidostomatidae ay

Distribution. Species of this group are indigenous to the Pacific Coastal

region.

Lepidostoma (Nosopus) jewetti Ross

(f{. 429 60,98)

Lepidostoma jewetti Ross 1946:269, 285-286, f. 16, 16C (male); Fischer 1970:58;

Anderson 1976:119; Nimmo and Scudder 1978:120; Webb 1980:97; Nimmo

and Scudder 1983:80.

Male. Scape cylindrical, 0.9 mm long. Maxillary palp fingerlike, 0.9 mm

long. Wings (f. 42) brown. Anterior wing about 9.3 mm long; venation unique,

being more complete than most male lepidostomatids, with Fl, F2, F3 and F5, but

only one anal cell, Cu2 fused with Cul basad, and 1A atrophied basad. Posterior

wing with Fl only, and with unique subapical marginal setose groove. Genitalia

(f. 60) having inferior appendage with long, slender, horizontal, middorsal

process and basodorsal process half as long.

Female. Genitalia (f. 98) having segment |X with short blunt anterolateral

apodeme and dorsum with several transverse grooves. Spermathecal sclerite oval

in ventral view and narrow in lateral view.

Immatures, case and habitat. Glenn Wiggins has provided several identified

immature specimens of L. (N.) jewetti, which show that this species may be

indigenous to spring habitats and that final larval instars construct panel cases.

Life history. Larvae collected 14 April, pupae 22 May. Recorded flight

dates: 15 May - 25 September.

Distribution. California, British Columbia, Oregon, and Washington.

Type data. Ross (1946): "Holotype, male. - Ellsworth, Washington, May

15, 1941, S. Jewett, Jr." Holotype at the INHS (Webb 1980). See type material

examined below.

Material examined. Holotype male, (INHS). CALIFORNIA: Tehama Co:

Dead Mule Spring, by Covelo Road 2 mi N of Pashenta, 5150’, 4 males, 6 females,

28 Vill 1972, HBL, (CASC). OREGON: Benton Co: Marys Peak, spring along

road at Chintimini Cr, larvae, 14 April 1964, (male and 2 females reared), NHA

& GBW, (ROME); Lincoln Co: Newport, male, 12 Vil 1968, B Brown, (OSUO);

Fiynn Cre-male pupa, 22:V 1982; maley14:1™ 1982, RWW, LOSUO).

Lepidostoma (Nosopus) licolum Denning

Cf G26, 439) 632

Lepidostoma licola Denning 1975:325-326, f. 7-7D (male).

Male. Head (f. 28) with scape cylindrical, 1.2 mm long. Maxillary palps

slender fingerlike, 0.5 mm long. Wings (f. 43) brown. Anterior wing, about

7.0 mm long, venation less complete than that of L. (N.) jewetti, with Fl, F2,

and FS, and Cu2 atrophied basally. Genitalia (f. 61) having segment X in lateral

view with small midventral notch adjacent to small protrusion. Phallus with

phalicata gradually curved. Inferior appendage similar to that of L. (N.)

jewetti, but with basodorsal process shorter.

Female, immatures, case and habitat. Unknown.

Life history. Recorded flight date: 9 July.

Distribution. Northeastern California. Only known from the type locality.

Type data. Denning (1975): "Holotype male. - Sierra County, California,

Yuba Pass, 6700', 9 July 1967. Paratypes, three males, same data as for holotype.

Types to be deposited in Academy of Sciences, Golden Gate Park, San Francisco,

California." See type material examined below.

28 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

Material examined. Three male paratypes (DGD), (of which a paratype was

kindly given to me by Dr. Denning).

Lepidostoma (Nosopus) Americanum Group

This species comprises four eastern species, L. (N.) americanum, costale,

bryanti, and prominens.

Male. Head unique (f. 31) with vertex projected anteriorly between eyes

like small visor, forming concave frontal area. Scape cylindrical, slightly longer

than head. Maxillary palp having flexible apical segment with many petiolate

scales and held before concave area of frons. Wings (f. 44) with venation as in

iL. {N.): podagrum. Anterior wing with Fl, F2, and F5 and usually both anal

cells, veins of arculus converging mesad, sometimes modified with large costal cell

as in L. (N.) americanum or costale, or with small costal cell basad as in L. (N.)

prominens. Posterior wing similar to that of L. (N.) podagrum with F1 present,

1A and 2A apparently completely fused. Genitalia (f. 62-65) with segment IX

relatively shorter than that of L. (N.) jewetti group. Segment X about as long

as high with two apical points in lateral view. Phallus short, except in L. (N.)

prominens. Inferior appendage with first article triangular and acuminate.

Female. Genitalia (f. 99-102) of L. (N.}) americanum and bryanti similar to

those of females of L. (Nosopus) unicolor group. Spermathecal sclerite broad in

lateral view and oval in ventral view, but in L. (N.) costale and prominens with

oval, lateral flanges in ventral view.

Immatures, case and habitat. Final instars construct panel cases, square

in cross section. Immatures show preference to inhabit small streams, rather than

springs.

Distribution. The four species of the L. (Nosopus) americanum group are

indigenous to eastern North America.

Lepidostoma (Nosopus) americanum (Banks)

Che 4A G2 99)

Olemira americana Banks 1897:29 (male).

Lepidostoma americanum: Flint and Wiggins 1961:295, f. 13A-B (female); Blickle

and Morse ,196Go°11;. Fisener 1970.28: Hilsennott. 1979:25; Wray .1967:56;

Roy and Harper 1975:1084; Roy and Harper 1979:162; Etnier and Schuster

1979:19; Neves 1979:171; Mingo and Gibbs 1980:45; Parker and Voshell

198-6: Ungieker ef al. 1962:9. 706) .9.3077 Lake 1984:221.

Lepidostoma americana: Seward and Swegman 1979:24.

[Mormonia pictilis Banks removed from this synonymy. |

Male. Head (f. 31). Scape cylindrical, 1.0 mm long. Maxillary palps flexible

bearing many petiolate scales and with segmentation obscured, 1.5-2.0 mm long.

Wings (f. 44) brown. Anterior wing similar to that of L. (N.) costale with large

costal cell bearing many scales (f. 12). Genitalia (f. 62). Similar to those of

L. (N.) costale having segment X in lateral view with long narrow dorsal point

parallel to shorter ventral point, both directed posterodorsally.

Female. Genitalia (f. 99) having spermathecal sclerite somewhat rectangular,

with posterior lobe, in lateral view; oval and slightly acuminate anteriorly, without

lateral flanges, in ventral view .

Immatures, case and habitat. Late instars construct panel cases with

hexagonal pieces of bark and wood. Pupae collected from small streams.

Life history. Pupae collected in late July. Recorded flight dates: 15 June

“i 13 August.

Weaver: North American Lepidostomatidae 29

Distribution. Delaware, Georgia, Maine, Massachusetts, New Hampshire,

New York, North Carolina, Pennsylvania, Quebec, South Carolina, and Virginia.

Report from Manitoba (Ross 1946) by Banks is in error (Flint and Wiggins 1961).

Type data. Banks (1897): "One male, several females, Sea Cliff, N.Y.,

Julyie

Olemira americana Banks, New Lectotype, male: Sea Cliff, L.Il., New York.

MCZC. Type 11700.

Material examined. GEORGIA: Union Co: Wolf Cr, 3.5 mi N of Neel Gap,

male; «. 0: Vil. A940, .. PW Fattios- CINHS)., MASSACHUSETTS: Hampshire Co:

Williamsburg, E branch near Ashfield Road, male pupa, 25 VII 1967, Merrill,

(ROME). NEW HAMPSHIRE: Coos. Cox Fabayan, male, 3 VIII 1958, OSF,

(USNM). NORTH CAROLINA: Jackson Co: 5 km SE of Balsam, 4 males, 3

females, 9-12 VII 1974, (USNM); Whitewater Falls, male, 10 females, 14 VII 1979,

JCM, (CUCC)... PENNSYLVANIA: Fayette €o:,- Bear Runs male, 6 VII 1978. BG

Swegman,: (CUSNM); Fulton Co: »-Crystal Spring. Chestnut Bridge Hollow, male,

t VIP VVSTT, SSW (JBW)s Aughwick Cr, male, femate, 30 Wb 477; JSw, (JSW);

Westmoreland Co: Rector, Powdermill Nature Reserve, Maul Spring, 18 males,

140 females, 16 VII-13 VIII 1975, JSW & JLS, (JSW). SOUTH CAROLINA: Pickens

Co: Carrick .Cresk;« Table Rock: St “Pk, male 5. Vib 1968) PHOr-(CUCC).

VIRGINIA: Fauquier Co: Burtons Br., Jackson Hollow, 7 males, female, OSF,

(USNM); Spotsylvania Co: 2 males, 15 VI 1974, B Strickler, (USNM).

Lepidostoma (Nosopus) bryanti (Banks)

Ga, 803

Alepomyia; bryanti Banks: 1908:65;. bla12,° 4. 12,13: Omete).

Lepidostoma wisconsinensis Vorhies 1909:685-687, f. 9, pl. 52 f. 8, pl 57 f. 1-10.

Lepidostoma bryanti: Ellts 1962:5, 14: Etnier 1965-151: Blickle and Morse

1966:11; Fischer 1970:39; Hilsenhoff et al. 1972:22, 28; Longridge and

Hilsenhoff 1973:181; Hilsenhoff 1975:25; Karl and Hilsenhoff Tere: 33}

39-40; Neves 1979:171; Roy and Harper 1979:162: Hilsenhoff IISA;

Parker and Voshell 1981:6; LaFontaine 1981:249, 251; Marshall and Larson

1982:78249, of. 15524 Krueger and Cook. 1984: 1482+: Singh ebialy 1984:296:

Steven and Hilsenhoff 1984:165.

Lepidostoma wisconsinense: Fischer 1970:69.

Male. Head and its appendages similar to those of L. (N.) americanum.

Scape 0.6 mm long. Anterior wing brown, about 7.3 mm long, without costal cell,

venation same as that of L. (N.) podagrum. Genitalia (f. 63) having segment X

in lateral view with two small subequal apical hooks directed dorsad.

Female. Genitalia (f. 90). Spermathecal sclerite similar to that of L. (N.)

americanum, in ventral and lateral view somewhat oval.

Immatures, case and habitat. Karl and Hilsenhoff (1979) observed many

larvae inhabiting Glen Creek, Wisconsin. They also remarked that the larvae

construct panel cases similar to those of L. (Mormomyia) griseum.

Life history. Larvae collected from November to late June: pupae from 5

May to 1 June. Recorded flight dates: 20 May - 7 September.

Distribution. Georgia, Maine, Massachusetts, Michigan, Minnesota, New

Hampshire, Newfoundland, New York, North Carolina, Nova Scotia, Ontario,

Pennsylvania, Quebec, Rhode Island, Tennessee, Virginia, and Wisconsin.

Type data. A/epomyia bryanti Banks (1908): "Four males from Grand Lake,

Newfoundland.” Ross (1938b): "Lectotype, male. - Grand Lake Newfoundland,

July 28,. .1906,.0,.. Bryant. “No. 11709.": 1 havecseen thatthe lectotype and one

other male syntype are at the MCZC.

Lepidostoma wisconsinensis Vorhies (1909): “These larvae are quite plentiful

in the streams at Devil's Lake and Parfrey's Glen, where they inhabit the pools.

The pupae were abundant at the Glen on June 1. They could be found only among

30 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

roots and grasses where these hung in the stream, as, for example, in a bit of

overhanging sod. | have not taken the imagos in their natural habitat. Those

reared in the laboratory appeared in the first days of June." New Lectotype,

Dane Cotnty,. Wisconsin, 7 Wi 1907/7, Co Pee Vorhies. MCZC Type 11688.

Material examined. GEORGIA: Union Co: Vogel St Pk, Wolf Cr, male, 6

females, :30-31 V 1980, JCM, €CUCC)u “MAINES Aroostook Ce: "Oxbow, 8-9 VII

1961, A Brower, (USNM). NORTH CAROLINA: Macon Co: Highlands, 5 males,

8 females, 3-8 VI 1961, OSF, (USNM). PENNSYLVANIA: Forest Co: Kellettville,

Ross Run, 18-19 VI 1976, JSW, (JSW); Lancaster Co: Welsh Mountains, New

Holland Watershed Area, male, female, 4-5 VI 1977, JSW, (JSW); Sommerset Co:

Kooser St Pk, Kooser Run, 20 males, 56 females, 7-8 VI 1976, JSW & JLS, (JSW);

Westmoreland Co: Rector, Powdermill Nature Reserve, Maul Spring, male, 20 VII

1877; JUS “6 JSW, (ISW).@ RHODE. “ISLAND: Washington Co: Richmond, ‘male,

20. Vi Toys AG. beavelleen (CUCL): “VIRGINIA: (Grayson Co: Fox Cry). & mi W

of Troutdale; male; 2 females, CM: 8° OSF, CUSINM)’.

Lepidostoma (Nosopus) costale (Banks)

Ch. 64°81)

Olemira costalis Banks 1914:265, pl. 10 f. 34 (male).

Lepidostoma_ costalis: Btls WS6235 °° 14>. Etnrer 1966: 151; Blickle: ‘and Morse

1966:11; Longridge and Hilsenhoff 1973:180; Hilsenhoff 1975:25; Karl and

1979:171; Mingo and Gibbs 1980:46; Hilsenhoff 1981:31; LaFontaine

1981:249, 251; Lake 1984:221; Steven and Hilsenhoff 1984:165.

Lepidostoma costale: Fischer 19/70:41; Roy and Harper 1975:1084; Roy and Harper

1979:162; Swegman et al. 1981:131, 144.

Male. Head and its appendages like those of L. (N.) americanum. Scape

1.0 mm long. Maxillary palp 0.7-0.8 mm long. Anterior wing brown, with large

costal cell, like L. (N.}) americanum. Genitalia (f. 64) similar to those of L. (N.)

americanum, but segment X in lateral view with two short apical points perpen-

dicular and subequal to each other, dorsal point directed anterodorsad, ventral

point directed posterodorsad.

Female. Genitalia (f. 91) with spermathecal sclerite similar to that of L. (N.)

prominens being nearly round and with pair of round lateral flanges in ventral

view.

Immatures, case and habitat. Larva and case undescribed, but Karl and

Hilsenhoff (1979) reported that many larvae and pupae were found at Glen Creek,

Wisconsin.

Life history. Larvae collected from mid-August to late June; pupae from

24 July to 5 August. Recorded flight dates: 26 May - 9 September.

Distribution. Connecticut, Delaware, Maine, Massachusetts, Michigan,

Minnesota, New Hampshire, New York, Ontario, Pennsylvania, Quebec, Virginia,

and Wisconsin.

Type data. Banks (1914): “From Woodworth's Lake, Fulton Co., N.Y., 19

August (Alexander)." Ross (1938b): ‘Lectotype, male. - Woodworth's Lake,

Fulton Co., New York, August 19, 1909. No. 11/01. Allotype, female. - Same

data." | have seen these specimens at the MCZC; only the lectotype bears a MCZC

type-label. :

Material examined. CONNECTICUT: Litchfield Co: Canaan, Music Mt, male,

31 VIII 1962, SW Hitchock, (USNM). PENNSYLVANIA: Crawford Co: Linesville

Cr, 2 males, 6 VIII 1973, Walker, (USNM); Forest Co: Pigeon, Marienville Rod

& Gun Club Poomales 028-20: Wake JSW, -Swh

Weaver: North American Lepidostomatidae Re

Lepidostoma (Nosopus) prominens (Banks)

($265,924

Arcadopsyche prominens Banks 1930:129, pl. 7 f. 1, 8, 10 (male)

Lepidostoma prominens: Fischer 1970:62; Roy and Harper 1975:1085; Williams and

Williams 1979:2407; Roy and Harper 1979:163; Neves 1979:171.

Male. Head and its appendages similar to those of L. (N.) americanum.

Scape 1.1 mm long. Maxillary palps 1.4 mm long. Anterior wing 9 mm _ long,

venation similar to that of L. (N.) bryanti, but with only one anal cell and small

costal cell basad. Genitalia. 1.;.65) similarntoy those-of: ..1N.) beyanti, “out

segment X in lateral view with long curved ventral point and shorter apical point.

Female. Genitalia (f. 92) with spermathecal sclerite in ventral view oval,

similar to that of -L: (N.)-costeale.

Immatures, case and habitat. Unknown.

Life history. Recorded flight dates: 3 June - 19 August.

Distribution. Massachusetts, New Hampshire, Newfoundland (Labrador),

Nova Scotia, and Quebec.

Type data. Banks (1930): "From Cape North, Cape Breton, Nova Scotia,

7 August." Ross (1938b): "Lectotype, male. - Cape North, Cape Berton Island,

Nova Scotia, August 7, 1928. No. 16321. Lectoallotype, female. - Same data."

| have seen these lectotypes and three additional male syntypes at the MCZC.

One male syntype is at the INHS (pers. comm. J. D. Unzicker).

Material examined. NEWFOUNDLAND: Labrador, Goose Bay, 4 males, 2

females, 34 July 1950; :Jd. Tibbles,; (GNChs: «NEW HAMPSHIRE::() Goes Gov Mt

Washington, Cutler ‘River,:«male; 4-5. Vi1l- 1958, OSF, CUSNM).:.. QUEBEC: - Parc

des Laurentides, Grand Lac Jacques Cartier, male, female, 8 VII| 1938, Vladyhoff,

(INHS).

Lepidostoma (Nosopus) Podagrum Group

This group comprises two species, L. (N.) podagrum and quercinum.

Male. Scape cylindrical unmodified. Maxillary palps slender and fingerlike.

The two species of this group have nearly identical male genitalia, and are best

distinguished on the bases of unique secondary sexual modifications of the labial

palps and anterior leg, exhibited only in L. (N.)}) podagrum (f. 29). Wings (Cf.

45) with anterior | Wing ‘similar’ to ‘thatiot Lich.) bryanti.. with Fiche and,

Cu2 atrophied, two anal cells present, without costal cell or other modifications.

Posterior wing with Fl; 1A and 2A completely fused. Genitalia (f. 66, 67) with

segment X about as long-as high, triangular, in lateral view. Phallus short,

similar to that: of miaolN. Jhbryenti,

Female. Genitalia (f. 93, 94) having segment IX with blunt dorsal subapical

lobe in lateral view. Spermathecal sclerite in ventral view oval, slightly pointed

anteriorly and posteriorly, about twice as long as wide.

Immatures, case and habitat. Final instar larvae construct panel cases,

square sagittally. Immatures inhabit mountain streams.

Distribution. The two species of the L. (N.)}) podagrum group are endemic

to the Pacific Northwest.

Lepidostoma (Nosopus) podagrum (McLachlan)

(ho 99) AG. (66, 92)

Nosopus podager McLachlan 1871:116, pl. 2 f. 6 (male).

Lepidostoma podager: Fischer 1970:61-62; Newell and Potter 1973:20; Anderson

1976:120:, Baumann: and Unzicker 1981:27:). Anderson et al. .1962:14:

Roemhild 1982:11; Ruiter and Lavigne 1985:43.

32 Contribs: Amer. Ent: Inst., Volume 24, Number 2, 1988

This was the first Lepidostomatid discovered in the New World. Contrary

to the general rule that lepidostomatid males have modified antennal scapes and

maxillary palps (or they have no modified appendages), L. (N.) podagrum is an

exception, having modified labial palps and anterior legs instead. This may

possibly be the result of a homeotic mutation, as the expression of the genes which

usually cause modifications of the antennal and maxillary somites of the head has

shifted posteriorly.

Male. Head and prothorax as in (f. 29): Scape 1.1 mm long, cylindrical.

Maxillary palp 0.8 mm long, fingerlike, slender. Labial palp 1.8 mm long, more

robust than that of female. Anterior leg with enlarged tibia, dilated basotarsus

bearing many petiolate scales. Wings (f. 45) brown. Anterior wing 8.5 mm long.

Genitalia (f. 66) similar to those of L. (N.) quercinum.

Female. Head and prothorax' (4/729). “Wings: (f. 49). (Genitaha (fF. 93)

similar to those of L. (N.) quercinum.

Immatures, case and habitat. Final instar larvae construct panel cases

square in cross section with irregular square pieces of bark and wood. They

inhabit small streams in pool areas. Pupae found in decayed wood.

Life history. Larvae collected 8 to 11 November, pupae 20 March. Recorded

flight dates: 4 February - 1/ September.

Distribution. Alberta, California, Idaho, Montana, Oregon, Utah and

Wyoming.

Type data. McLachlan (1871) as cited by Kimmins and Denning (1951):

"California: (H. Edwards) 1 male. Type mounted as a microscope preparation."

Dr. P. C. Barnard (pers. comm.) has confirmed that the unique type is in the

British Museum (Nat. Hist.). | have seen that there is a male at the MCZC labeled

"Type 10941 Sierra Nev Cal" and there is another male labeled as a type at the

INHS (pers. comm. J. D. Unzicker), however neither specimen has type-status.

Material examined. ALBERTA: Carrot Cr, old Hwy 16, 92 males, 15 females,

4 Vl 1976, APN, (UASM);: Chickadee Cr, Hwy 43, W. of Whitecourt, male, 1 VI!

1967, APN, (UASM); House River, Hwy 63, 2 males, 32 females, 29 VI 1978, APN,

(UASM); Lobstick River, Hwy 16, Niton Junction, 5 males, 3 females, 4 VI 19/6,

APN; (UASM):: Rourke Cr, Hwy 44, S of Smith; 5 males, 9 females, G VI 1976,

APN, (UASM); Smulland Cr, Forestry Trunk Road, 4/7 mi S of Goodwin, male,

26 females, 2 VII 1967, APN, (UASM); Two Cr, Hwy 43, W of Whitecourt, 7 males,

Tle Mil 4967, APN, GUASM) .© CALIFORNIA: (Contra Costa Co: “Mt.: Diavlo,: male,

20 Il 1954, SW Hitchock, (USNM); El Dorado Co: Fallen Leaf Lake, 4 males, 20

Vi 1961, 6 males, 20 VI 1961, SDS, (UICM); River between Fallen Leaf & Tohoe

Lakes 3 males, 3° females) 22° V) 1961, SPS; (UICM): Marini Gos: Lageonitas Cr,

0.5 mi W of Jewell, 40’, larvae in pools on vegetation & roots 11 XI 1982 (adults

reared), LES, (LES); Monterey Co: Big Sur River, 2 males, 2 females, 1 VI 1961,

SDS. (UCM? SantasGlara’ Co: “Guadaluse Cr,.-male, s20ch4 1960; SDS, CUCM):

Coyote Cr, 2 males, 2 females, 4.11 °1961, SDS,°(WICM); Uvus Cr, male,°201V

1960;:SDS\« CUSNM):.. IDAHO: Teton: Co: 2 mic Woof -Tetonia, Teton River, 17

males, female, 14 VI 1969, SDS, (JSW). MONTANA: — Gallatin Co: Bozeman,

Bozeman Cr, 6 mates, 2 females. 16° Vi 1986, DLG,: (JSW): > OREGON: ‘Harriey

Cot “Pueblo Mt, Cottonwoed Cr, male, 29 V 1925; SGJ>°CUSNM);. dosephine Co:

Grayback St Pk, 12 mi E of Cave Junction, 3 males, 2 females, 13 VI 1963, NHA,

(OSU0): «Lane Co: — Lockout Cr, 6 mi NE of Blue River. ‘final instars 8° xX! 19/6

(preserved as pupae 11 | 1977), NHA & Frost, (ROME); 12 mi SE of Eugene, male,

17 1X 1966, KG, (OSUO); Lincoln Co: Flynn Cr, pupae burrowed in punky wood,

20 Ill 1981 (male & 4 females emerged 13 IV); 9 males, female, 19 V 1982, RWW,

COSUG):.

Weaver: North American Lepidostomatidae 33

Lepidostoma (Nosopus) quercinum Ross

(fie B07 68, 94)

Lepidostoma gquercina Ross 1938a:1/76-177, f. 119 (female); Anderson and Wold

1972:192, 19%;%. Anderson 1976: 120-1215 = Grafius: ‘and Andereon < 1979:

433-441; Nimmo and Scudder 1978:120; Webb 1980:98; LaFontaine

1981:249; Dudley and Anderson 1982:10; Roemhild 1982:11; Nimmo and

Scudder 1983:80.

Lepidostoma quercinum: Fischer 1970:62.

Male. Head (f. 30) with scape cylindrical, 1.0 mm long. Maxillary palp 0.8

mm long, fingerlike. Labial palp 1.5 mm long, about as slender as maxillary palp.

Anterior leg unmodified. Wings brown, venation similar to that of L. (N.)

podagrum. Anterior wing 8.3 mm long. Genitalia (f. 67) indistinguishable from

those of L. (N.) podagrum.

Female. Genitalia (f. 94) similar to those of L. (N.) podagrum.

Immatures Cases and _ Habitat. Contributions .to” the bionomics of. L.

quercinum have been presented by Anderson and Wold (1972), Anderson (1976),

and Grafius and Anderson (1979). These studies revea! that larvae of this species

inhabit small streams and that final instars construct panel cases similar to those

of dt: (N.).podagrum.

Life history. Larvae were collected from August to February; pupae from

January to February. Recorded flight dates: 21 March - 2 September.

Distribution. British Columbia, Idaho, Montana, Oregon and Washington.

Type data. Ross (1938a): "Holotype, male. - Corvallis, Oregon: April 2,

1935, Oak Creek." Holotype at the INHS (Webb 1980).

Material examined. IDAHO: Shoshone Co: Emeral Cr, male, 26 V 1962,

SD$, (SDS)... OREGON::. Benton.Co: Camp Adair, male, 15-1965, SGJ, (USNM);

9 mt°N of Corvallis, Berry Cr, 5 mates;. 19° IV 1967,/ED Evans, (USNM)< ibid, 3

males, 14.4Y 1973, (NHA,<€OSUO)?; > 3< males: 12:4M. 1980; 6 males, 3 females,’ -20

Vi 1982, RWW, (OSUO); Corvallis, Oregon’St Univ, Oak Cry''5 malés,> 4 females,

We22 AM 1969, Oo Kerstys GUSNM)s diobster Cry camels mt SW of (Alsea.- larva,

pupa, 4 VIII 1974 (emerged 15-21 Vill), NHA, (ROME); 5.5 mi’S of Philomath,

Rock Cr, Hwy 34,02 males,.21 V-1970, “E’ Evans; (OSUO); Clatsop Cor €& Humbug

Cr, male, 9 V 1965, SGJ, (USNM); Nehalem R at Red Bluff Rd, 4 mi NE of Elsie,

male; 8 V. 1965; 4male; 23) 1W'4966; SGJ. CUSNM)>Linceln Ces: Deer’ Cr)’ 2 males.

24-28 V. 1974, RWW, (OSU0)::FlynnGr,; 3: males, female, 4-5°V 1982: 4 males.

2M 1962, RWW; {OSUO)}) 1: mi.N Marlan, Big Elk Cr, male,.24 Wilt 1982° 3 males;

21% 1982, RWW (OSVUO)s Union. Co} .20-mi SW ofvhe Grande, Ladd Cr. 3 males.

13921 Vb 1978;.SD8, (8D0$)> male. 20-44 1978) SDS) 24dSW):- WASHINGTON: Grays

Harbor Core 25 mpcN’ Hoquiam, Donkey ‘Or, male, 18 Vo 1973, SDS; (SDS):

Lepidostoma (Nosopus) Roafi Group

This group comprises only one species, L. (N.) roafi.

Male. Wings (f. 48) similar to those of L. (N.) unicolor. Genitalia (f. 68)

with segment X short. Phalicata and parameres short. Inferior appendages with

first article acuminate, basodorsal process short; second article acuminate,

directed posteriorly.

Female. Genitalia of L. (N.) roafi (f. 95) with spermathecal sclerite similar

to that of L. (N.) cascadense in lateral view.

Immatures, case and habitat. The final instar larvae of L. (N.) roafi con-

struct panel cases.

Distribution. Western North America.

34 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

Lepidostoma (Nosopus) roafi (Milne)

| (62:48;) 68°95)

Atomyia roafi Milne 1936:120, 121, 124 (male).

Lepidostoma_ roafi: Fischer 1970:63-64; Anderson and Wold 1972:192, 197-198;

Newell and Potter 1973:20; Anderson 1976:122; Ellis 1978:203, 204; Nimmo

and Scudder 1978:120; LaFontaine 1981:251; Roemhild 1982:11; Baumann

and Unzicker 1981:27; Anderson et al. 1982:14: Nimmo and Scudder

1983:80; Anderson et al. 1984:15, 17, 18; Nimmo and Wickstrom 1984:4;

Ruiter and Lavigne 1985:43; Herrmann et al. 1986:435; Nimmo 1986a:2.

Male. Scape 0.8 mm long, not highly modified. Maxillary palps slender,

spatulate. Wings (f. 48) with venation similar to that of L. (N.) unicolor.

Anterior wing about 7.7 mm long, not highly modified. Genitalia (f. 68) similar

to those of L. (N.) verodum. Segment X about 2/3 as long as inferior appendage,

dorsum and venter curved together forming blunt posterior apex in lateral view.

Phalicata short, about half as long as parameres. Inferior appendage with vertical

basodorsal process about half as long as horizontal middorsal process, first article

acuminate, middorsal process extending to base of second article.

Female. Genitalia (f. 95) having spermathecal sclerite with height greater

than length in lateral view. |

Immatures, case and habitat. Final instar larvae construct panel cases.

Larvae inhabit small streams in pool areas, pupae found attached to wood.

Life history. Larvae collected 8 July to 20 August; pupae early July to 15

September. Recorded flight dates: 28 May - 19 October.

Distribution. Alaska, Alberta, British Columbia, California, Colorado,

Idaho, Montana, Nevada, New Mexico, Oregon, Utah, Washington, Wyoming, and

Yukon.

Type data. Milne (1936): “Holotype male 'Eagle Co. (Sheephorn, 7900'),

T-Vill, eG: Ff. Clage ,. 1: male paratype Horseshoe: Ro section: (Grand ‘Co., 8500’),

Colo;G<7G5Vill ain theo M.C. 2.3: fourteen other males<from BoC:, Ores, 28-V- to

TOaXy 3 tmoethe: Cans: Nat. collection, 11 otn. the author's. collections ":o b have ‘seen

the holotype and three paratypes, one male and two females at the MCZC. A male

and female paratype are at the INHS (pers. comm. J. D. Unzicker).

Material examined. ALASKA: Prince of Wales Isl, Red Lake, male, 13 IX

1951,°8 Malkiny SOASC)) Prepper Cr. Hwy 3, 2 males; 2 temates, 2° Vib: 1979;

APN, (UASM)) “ALBERTA: Cold Cri Hwy..J6,. Nojack): 2>malesys female, .24°Vi4

1977, APN, (UASM); Ernst Cr, Maycroft-Gap Rd, 3 males, female, 31 VII 1976,

APNG (OASM)s\ Lusk Gr Forestry: Trunk Road, “Kananaskis, male, 1 VII 1976,

APN, (UICM); Moose Cr, Forestry Trunk Road, S of Clearwater River, 3 males,

2 females, 17 VII 1976, APN, (UASM); Red Deer River, Crossing, Forestry Trunk

road, 2: males, 6 MIN? 18ie, oA PN, CUASM). BRITISH COLUMBIA: UBC. Exp

Perest, Spring Cry. 2 males; 19 Vill 19al, K. Van Frankenhuyzen,. COSUO): 4bid,

‘Maythy Gr, near Haney, 8 males, 13-15 (Vili 1986,. JS. Richardson, (JSW).

CALIFORNIA: Fresno Co: Dinkey Cr, 2 males, 10 IX 1983, RD Haines, (DGD);

Sonoma Co: Fairfield, Osborn Preserve, Copeland Cr, 1540', larvae in pools 20

Vill 1980: (adults réared), LES;) (LES)... IDAHO: © Blaine: Co: Ketchum, °2:males,

2F WINS 1946) BES) (2SW): Butte Co: ooCraters’ Moon Nat! Monument; «2: malesy3

females, 12 Vilh.l965,; DS Horhingy: (SD5)>.ranktin Co: . Gub River Canyon, male

pupa; 25 Vill 19/73, GF Knowlton, CUSNM)>. Shoshone Co: Mullan; Willow: Cr,

male, 22 VIII 1950, JAR & HHR, (USNM). MONTANA: Lake Co: Yellow Bay

State Pk, Yellow Bay Cr, mate, sods W6e.. RWBy CUSNM):) Missoula: Ce:

Rattlesnake Camp, male, 16 IX 1968, RWB, (USNM). NEVADA: Washoe Co:

Galena Cr, male, 31 VII 1968, RWB, (USNM). OREGON: Baker Co: Spring Cr,

male, female, 25 IX 1967, JHB, (USNM); Benton Co: 7 mi NW of Corvallis, male,

» femaies, 3-10 1X 1969. CD: Kerst,. CUSNM)> Clatsop Co: Vie Gronne!l Rd, 2 mi

E of Biste, mate, 1-14 Vil 1e60, SG), CUSNM);. Crook Co: Ocheco Mts, Hwy 26,

near Summit, 2 males, 31 VII 1965, SGJ, (USNM); Lane Co: Mack Cr, HJ Andrews

Weaver: North American Lepidostomatidae BO

Forest, 11 mi NE of Blue River, 4 larvae, 8 VII 1976 (reared to male, 2 females,

male pupa preserved 19 VIII), NHA, (ROME); HJ Andrews Exp For, Mack Cr,

male emerged, 28 VIII 1973, (OSUO); ibid, 2 male pupae, 15 IX 1977, L Roberts,

(OSUO): “ibid, dookout: Cr, male female, (25° Vidb 1982, RWW (OSU0l> aan Co:

Quartzville Rd, Loan Star Bridge, 8 larvae?, 3 VIII 1974 (plus 4 adults, emerged

5-15 VIII), NHA, (OSUO); Quartzville Rd, 2.5 mi E Rocky Top Bridge, immatures,

26 VI 1973 (male pupa & female, emerged IX), NHA, (OSUO); Lincoln Co: Flynn

Cr, male, female, 30 VIII-6 IX 1962, (OSUO): ibid, 5th instar larvae (2 pupae,

4 males, 4 females emerged early VII 1981), RWW, (OSUO); ibid, attached to wood,

prepupae & pupae 19 VIII 1981 (male & 8 females emerged 23 Vill-mid IX), RWW,

(OSUO); Union Co: Big Cr, 7 mi E of Medical Springs, males & females, 15 VIII-5

Ix 1975, SDS, (SDS); Goose: Gry:1d mich cot: Medical: Springs 2 ¢males? 22 Vill

1975, SDS, (SDS); Lick Cr,:6 meue Of Medical’Springs males © females, 15 Vit-5

[xX 19/5, SDS, (SDS)9 Velvet’ Cr,’ S mE of Medical Springs; males? 6°terales, 22

Vil-16:.1X 1913;.SD8, €SDS)> Jordan Cr, 20 mi SW of-la Grarige, males © females,

Vo W111 Te 1X: 89 /5)-8DS -4$D$). 0 UTAH < Summie Co: Praihieke, rail Camo,

male, 20 VIII 1973, RWB, (USNM); Uinta Mts, Hayder Fork of Bear R., male, 4

Vit 1978, RES» (JSW) eo: WASHINGTON: King Go: Hull Cr, G@ mieAk of North Bend,

male, 2. females p11 4% 21960,,°SDS, “(S08)¢0 Kittitas Go: 2 mi NE of ENesbutg;, 15

males, 4 females, 19 X 1968, SDS, (SDS); Gale Cr, Kachess, male, 25 VIII 1970,

SDS, (SDS); 2 mi N of Little Kachese: bakes 2 mates, 2¢femates, LO Wil POF,

SDS, (JSW); 17 mi N of Roslyn, Cla Elum River; 4 males, @ 1x<'9968, SDS, (SDS);

Lewis’ Co: Tt mi 'S) of Randle, 1200, Camo Cr. 2 males: SB 104% 180, DRGs

SDS, (8D8)r Whatcom Cot Glacier, Gallup Cr, 2 4males, 6 PX’ 1970, SOS (8 0s).

WYOMING: Yellowstone Natl Pk, Virginia Meadows, Gibbons River Br, male, 31

Vb 1940, THE: & THF Jr yoCUSNM)? Shoshone Natl Forest, “Crazy Cre? males; 2

females, 25 VIII 1955, KM Fender, (USNM). YUKON: Judas Cr, Alaska Hwy,

110.4 km W of Teslin Bridge, male, female, 24 VII 1979, APN, (UASM).

Lepidostoma (Nosopus) Modestum Group

This group comprises six species, L. (N.) compressum, lydia, modestum,

ontario, pictile, and weaveri. The adults and immatures appear to be slightly

smaller than most other eastern species of Lepidostoma.

Male. Head (f. 32) with scape not highly modified, cylindrical. Maxillary

palp with apical segment broader than basal segment, mesal surface concave

bearing several long scales, segmentation obscured. Wings (f. 46) with venation

similar to that of L. (N.) podagrum group, but anterior wing with only one anal

cell and hindwing with 1A and 2A not completely fused. Genitalia (f. 69-73) with

segment X usually long and narrow, about 0.75 times as long as_ inferior

appendages. Phallus short. Parameres, lightly sclerotized flexible lobes. Infe-

rior appendage, without middorsal process; first article, long and fingerlike;

second article, subapical mesal lobe; basodorsal process capitate, parallel to and

not shorter than first article.

Female. Wings (f. 46) with venation typical of female lepidostomatids.

Genitalia (f. 96-99) having sternite VIII with anterior lateral margins heavily

sclerotized and mesoposterior region lightly sclerotized, forming genital pouch.

Spermathecal sclerite in ventral view bell-shaped, being widest anteriorly.

Immatures, case and habitat. Larval head dark brown with length of

postmentum about equal to that of midventral ecdysial line (f. 8), being shorter

than that of other known larvae of Lepidostoma. Final instars construct cigar-

shaped, panel cases (f. 4), with hexagonal pieces of leaf. Identified larvae of

L. (N.) lydia and ontario found in mountain springs suggest that other species

of this group may be endemic to similar habitats.

Distribution. The species of the L. (N.) modestum group are indigenous

to eastern North America, being particularly common throughout the Appalachian

Mountains.

36 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

Lepidostoma (Nosopus) compressum Etnier & Way

th. Go)

Lepidostoma compressa Etnier and Way 1973:428-429, f. 5A-C (male); Etnier and

Schuster 1979: 29. |

Lepidostoma compressum: Unzicker et al. 1982:9.107.

Male. Scape, maxillary palps ana wings similar: to. those of Lb. - (N;)

modestum. Genitalia (f. 69) with segment X in lateral view similar to L. (N.)

modestum, but in dorsal view similar to L. (N.) pictile and weaveri. Apex of

phalicata curved ventrad.

Female, immatures, case and habitat. Unknown.

Life history. Recorded flight date: 23 May.

Distribution. Tennessee.

Type data. Etnier and Way (1973): ‘Holotype, male. - Plateau Experiment

station, Cumberland: Co., Tenn: , 23 May 1970, Univ. Tenn. Dept. Ag. Biol. staff.

Paratypes. - Same data as for holotype, 28 April 1967, 2 males. The holotype

has been deposited at the U.S. National Museum. A paratype has been deposited

at the University of Georgia. The remaining paratype is in the senior author's

collection." The first mentioned paratype is now at the INHS (pers. comm. J.

D. Unzicker), the other is mentioned below.

Material examined. Paratype male, (ECUT).

Lepidostoma (Nosopus) lydia Ross

ho 10, 9G)

Lepidostoma lydia Ross 1939:/70, pl. 10, f. 9-9B (male, female); McConnochie and

Likens 1969:150; Fischer 19/0:59: Roy and Harper 19/75:1085: Etnier and

Schuster “19/79: 19; Roy and Harper 19/97162;° Neves 19/9:171; Webb

1980:97; Parker and Voshell 1981:6; Unzicker et al. toon. 2. 106, Oo 107.

Male. Scape, maxillary palps and wings similar to those of L. (N.)

modestum. Genitalia (f. 70) with segment X in lateral view similar to that of L.

(N.) pictile, but with shallow mesoapical notch in dorsal view. Phallus with apex

of phalicata curved ventrad.

Female. Genitalia (f. 96) having segment |X with short blunt apodeme.

Spermathecal sclerite in ventral view similar to that of L. (N.) ontario; in lateral

view similar to that of L. (N.}) modestum, but somewhat larger and without

membranous-like dorsal mass.

Immatures, case and_ habitat. Immatures appear to be endemic to

springbrooks. Final instars construct cigar-shaped panel cases, which are round

sagittally and composed of hexagonal pieces of leaf similar to that of L. (N.)

ontario.

Life history. Larvae collected February and August; pupae 23 February to

19 June. Recorded flight dates: 20 April - 6 September.

Distribution. Georgia, Massachusetts, Newfoundland, New Hampshire, New

York, North Carolina, Nova Scotia, Pennsylvania, Quebec, South Carolina,

Tennessee, and Virginia.

ype data. «Ross (1939). Holotype, males - Lydia, Va, April 20;' 1938,

Ross. & Burks, Allotype, female. - Same data. Paratypes. - Same data, 18

males." Holotype at the INHS (Webb 1980). Other type material at the INHS

(pers, comm. do D. Unzieker):

Material examined. GEORGIA: Union Co: Vogel St Pk, Wolf Cr, 4 males,

22 females, 30-31 V 1980, JCM, (CUCC). NEWFOUNDLAND: Big Falls, male, 28

Vi 1966, DR Smith, (USNM). NEW HAMPSHIRE: Co00sCo: Mt Washington,

Jefferson Notch Rd, male, 7-8 VII 1977, JSW, (JSW); ibid, Pinkhams Notch, Cutler

River, 15 males, 8-9 VII 1977, JSW, (JSW). NORTH CAROLINA: Macon Co:

Weaver: North American Lepidostomatidae at

Standing Indian Cpg,-Kimsey ECriy4amales,2 temales, 13 V¥1971, JCMeehOCUCEe):

Wayah Bald.at hairpin turn; 20: V. 19/0;-Q0S5F, (USNM)> Yancey Go: Mt. Mitchell,

Camp Alice, male, 9 VI 1961, OSF, (USNM). PENNSYLVANIA: Westmoreland

Co: Rector, Powdermill Nature Reserve, Maul Spring, 3 larva, male pupa, 23 II

1976; 3-male pupae, ¢ pupae, 6.V 1975: pupa, 19 Vl 19/5; 2 Jarvae, 14 Vill 4975,

JSW S/S, (05W)..> PENNESSEE: Sevier Co: 10.2 mi E of Gatlinburg, 7 males;

female, 20 V 1970, OSF, (USNM). VIRGINIA: Shenandoah Natl Pk, Skyline

Drive; mile post 79.5, male,..20 V.1961, OSE, LUSNM)> Shenandoah Nath Pk,

Hogcamp .Brk, male,..22-23..V 1940,..0OSF, LUSNM):> Madison; Co: tributary of

Hughes River, N of Nethers, male, 6 V 1973, OSF, (USNM); Rockingham Co:

Hone Quarry Cpg, male pupa, 5 larvae, 29 IV 1967, OSF, (USNM).

Lepidostoma (Nosopus) modestum (Banks)

(fe 32, 46; hl 8T)

Atomyia modesta Banks 1905:217.

Lepidostoma modestum: Wray § 1967:56%; Fischer - 1970760; Etnier -and< Way

1973:429, f. 6A-C (male); Roy and Harper 1975:1085; Etnier and Schuster

1919:19; Roy and Harper -19/9:162. Unzicker et al. 1962:9.106, 9.107,

Male. Head (f. 32) with scape cylindrical, unmodified, 0.4 mm _ long.

Maxillary palps 0.4 mm long, fingerlike with mesoapical portion concave and

bearing many petiolate scales. Wings (f. 46) dark brown. Forewing without costal

cell or setose grooves, venation similar to that of L. (N.) unicofor group.

Genitalia (f. 71) with segment X similar to that of L. (N.) compressum in lateral

view, but with long narrow pair of fingerlike mesoapical processes, in dorsal view.

Female. Wings (f. 46) typical of lepidostomatid females. Genitalia (f. 97)

with spermathecal sclerite in ventral view similar to that of L. (N.) ontario, in

lateral view somewhat like that of L. (N.) lydia, but with more massive, irregular

dorsum.

Immatures, case and habitat. Unknown.

Life history. Recorded flight dates: 13 May - 18 August.

Distribution. New Hampshire, North Carolina, Quebec, Tennessee, and

Virginia.

Type data. Banks (1905): "Many specimens from Black Mountains, North

Carolina, May and June (Beutenmuller)."” Ross (1938b): "Lectotype, male. -

Black Mts., North Carolina, May. No. 11702." | have seen the lectotype and

one other male syntype at the MCZC. A male syntype is at the INHS (pers. comm.

Jiod Uazieker)-.

Material examined. NORTH CAROLINA: Macon Co: Standing Indian Cpg,

Nantahala» River, 26 males, 35: females;:13.V 1977) JCM, (CUCC). VIRGINIA:

Rappahannock Co: Shenandoah Natl Pk, Piney River, male, 28 V 1977, OSF,

(USNM).

Lepidostoma (Nosopus) ontario Ross

Che) 428) 72298)

Lepidostoma ontario Ross 1941:119-120, pl. 13 f. 98-98A (male, female); Blickle

and. .Morse 1966:.11; Wray: 1967:56: Fischer 1970:60-61;\.Roy and Harper

1975:1085; Etnier and Schuster 1979:19; Roy and Harper 1979:163; Neves

1979:.171;., Webb: 1980: 98; Unzieker et al. 198229. 106, 9107.

Male. Scape, maxillary palp and wings similar to those of L. (€N.)

modestum. Genitalia (f. 72) having segment X unique with pair of ventrolateral

processes which extend posteriorly beyond mesal processes.

38 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

Female. Genitalia (f. 98) having segment IX in lateral view with long

attenuated apodeme. Spermathecal sclerite in ventral view similar to that of L.

(N.) pictile.

Immatures, case and habitat. Larval head venter (f. 8). Larvae appear

to be endemic to mountain springs and final instars construct cigar-shaped, panel

caseeu( fa 4);

Life history. Larvae and pupae collected 18 April 1981. Recorded flight

dates: 18 April - 24 August.

Distribution. Maine, Massachusetts, Newfoundland, New Hampshire, North

Carolina, Nova Scotia, Ontario, Quebec, South Carolina, Tennessee, and Virginia.

Type data. Ross (1941): “Holotype. - Male; Costello Lake, Algonquin Park,

Ontario, Ontario Fisheries Research Laboratory; May 29, 1938; (W. M. Sprules;

station #2). Allotype. - female; same data as for holotype; [INHS]. Paratypes.

- 2 males, 2 females; Randolph, New Hampshire; June 25, 1930. 3 males; same

data as for holotype, but May 30, 1938: 1 female; May 31: 1 male; May 26.

Paratypes are in the collections of the Illinois Natural History Survey, the Museum

of Comparative Zoology, and The Academy of Natural Sciences of Philadelphia."

Holotype at the INHS (Webb 1980). | have seen 3 paratypes, 1 male and 2 females,

at the MCZC; Type: 25863.

Material examined. NEWFOUNDLAND: Lewisporte near Notre Dame Bay, 2

males, 12 VII 1961, CPA, (USNM). NEW HAMPSHIRE: Coos Co: Mt. Washington,

Jefferson Notch Road, Mt. Pleasant Brook, male, 4 females, 9 VII 1977, JSW,

(JSW); ibid, Abenaki Brook, 3 males, female, 7-8 VII 1977, JSW, (JSW). NORTH

CAROLINA: Blue Ridge Parkway, Crabtree Meadows, male, 5-6 VI 1962, EG

MacLeod, (USNM); Macon Co: Highlands, male, 3-8 VI 1961, OSF, (USNM); Wayah

Bald, male pupae, 8 VI 1961, OSF, (USNM); Wayah Bald, stream at hairpin turn,

2 males, 8 females, 20 V 1970, OSF, (USNM); Wayah Crest Cpg, 3 males, 30-31

V 1981, JSW & RWH, (JSW); Wayah Bald, Rattlesnake Spring, 5 larvae, 4 pupae,

18 IV 1981 (4 males, female emerged same day); 3 males, 3 females, 30-31 V 1981,

JSW & RWH, (JSW); Jackson Co: Whitewater Falls, Whitewater River, male, 6 V

1975, PHOY CCUCC): s VYaneey Co: > Black: Mts. 4/60:,.. Mamphill Spring, .2 males,

female, 1-2 VII 1979, JSW, (JSW). SOUTH CAROLINA: Oconee Co: Wash Branch

above Crane Cr, Tamassee Rd 0.4 mi E of Hwy 107, male, 3 females, 24 IV-15 V

1983, JSWoe sPJS.. WeSW); Pickens Co: Wildcat Cro (S mi NW ‘of: Clemson) 4 males,

7 females, 17-21 IV 1968, PHC, (CUCC). VIRGINIA: Shenandoah Natl Pk, White

Oak Canyon, male, 27 V 1962, OSF, (USNM).

Lepidostoma (Nosopus) pictile (Banks)

Of Fa 8)

Mormonia pictilis Banks 1899:211 [removed from synonymy of Lepidostoma

americanum (Banks) ].

Lepidostoma swannanoa Ross 1939:69-70, pl. 10 f. 11-11A (male, female) NEW

SYNONYM; Wiggins 1961:/02; Wray 1967:56, 57; McConnochie and Likens

1969:150; Fischer 1970:65; Roy and Harper 1975:1085; Etnier and Schuster

1979:20; Williams and Williams 1979:2407, 2410; Neves 1979:171; Roy and

Harper 1979:163; Mingo and Gibbs 1980:46; Webb 1980:98; Parker and

Voshell 17981:6; Unzicker et: al..1982:9.106, 9.107; Marshall and. Larson

1BGA 78). Fh kes :

New male and female associations of several species of the L. (N.) modestum

group revealed that L. (N.) pictile (previously Known only from the female

holotype) is the senior synonym of L. swannanoa (which was based on the unique

characteristics: of the male).

Male. Scape, maxillary palp and wings similar to those of L. (N.)

modestum. Genitalia (f. 73) with segment X similar to that of L. (N.) lydia in

Weaver: North American Lepidostomatidae 39

lateral view; but with two acuminate processes bearing serrate lateral margins,

separated by long narrow gap, in dorsal view.

Female. Genitalia (f. 99) having segment IX with short blunt apodeme in.

lateral view. Spermathecal sclerite in lateral view similar to that of L. (N.)

ontario, being short in length and elongated in height.

Immatures, case and habitat. Unknown.

Life history. Recorded flight dates: 21 April - 1 August.

Distribution. Maine, Massachusetts, New Hampshire, Newfoundland, New

York, North Carolina, Nova Scotia, Quebec, Tennessee, Virginia, and West

Virginia.

Type data. Mormonia pictilis Banks (1899): "Franconia, New Hampshire

(Mrs. A. T. Slosson)." New Lectotype, female: Franconia, New Hampshire,

MCZC Type 11699.

Lepidostoma swannanoa Ross (1939): "Holotype, male. - Black Mountain,

N.C., April 24, 1938, North Fork Swannanoa River, Ross & Burks. Allotype,

female. - Same data. Paratypes. - Same data, 2 males." Holotype at the INHS

(Webb 1980). Allotype and paratype at the INHS (pers. comm. J. D. Unzicker).

See type material examined below.

Material examined. Lectotype female of M. pictilis, (MCZC); allotype female

of L. swannanoa, (INHS). NEWFOUNDLAND: Big Falls, 26 VI 1966, DR Smith,

(USNM); Aspen Brk Cag, male, ies voaly 1961, GPA, (USNM).

28 NEW HAMPSHIRE: Coos Co: Groveton, male, 9 VI 1973, WJ Kief, (USNM).

NEW YORK: Allegany St Pk, 7 males/females, V-3 VI 1963, WW Wirth, (USNM).

NORTH CAROLINA: Swain Co: Great Smoky Mts Natl Pk, Smokemont Cpg, 3

males, 3 females, 12 V 1970, OSF, (USNM). MASSACHUSETTS: Hampshire Co:

Cummington, woodland spring, 3 males, 4 VI 1973, AB Gurney, (USNM). RHODE

ISLAND: Washington Co: Richmond, male, 20 VI 1971, AC Lavallee, (CUCC).

VIRGINIA: «Bath Go: Blowing Springs, Black River, male, 16 V 1975, OSF,

(USNM); Blowing Springs, camp 8 mi W of Warm Springs, 57 males, 69 females,

18-29 V 1963, Field & OSF, (USNM); Douthat St Pk, 2 males, 3 females (immatures

in Il) emerged 15 IV 1973, B. Stieckler, (USNM). WEST VIRGINIA: Tucker

Co: Red Cr, near Laneville, male, 26 V 1973, OSF, (USNM).

Lepidostoma (Nosopus) weaveri Harris

Lepidostoma weaveri Harris 1986:36-38, f. 4A-G (male, female).

This species is closely related to L. (N.) pictile and especially L. (N:)

compressum. I\t differs from L. (N.) pictile by having male segment X in lateral

view tapering abruptly into a slender apical arm. Also, it differs from L. (N.)

compressum by having segment X in dorsal view with lateral margins of distal arms

gradually curved mesad, with their apices rounded and widely separated.

Male. Color brown. Anterior wing 7.0 mm long. Venation similar to L. (N.)

modestum. Scape cylindrical, 0.5 mm long, and maxillary palp, 0.7 mm long, both

similarly modified in L. (N.) modestum. Genitalia: Segment X in lateral view

with dorsum curved ventrad toward subapex, venter almost horizontal, and

slender apical arm curved slightly and directed posteriodorsad; in dorsal view

comprising two mesal processes, having basal third fused together, lateral margins

parallel and straight, and apical arms acuminate and curved slightly mesad.

Inferior appendage similar to that in L. (N.) compressum, but having apex of

main ventral process broader and more rounded.

Female. Color brown. Anterior wing 7.0 mm long. Scape cylindrical 0.6

mm long. Genitalia having dorsum of segment IX and spermathecal sclerite similar

to that of £. (Nv) pletile:

Immatures, case and habitat. According to Harris (1986) the larva has

postmentum and larval case characteristic of the L. (N.) modestum group, and

inhabit several small, spring-flowing, temporary streams.

40 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

Life history. Final instar larvae collected in February and March. Recorded

flight dates: 13-19 March.

Distribution. The Cumberland Plateau of Alabama. Only known from the

type localities.

Type data. Harris (1986): "Holotype male. - Alabama, Tuscaloosa County,

unnamed tributary to Tyro Creek at bridge 303, 4.5 miles east New Lexington

(RIOW, T17S;7 S15), c12¢March: 1984," S:- Harris’ (USNM). Paratypes. - Alabama,

same as above, / males, 2 females (USNM, INHS, UA), same, but 14 March 1984,

2 males, 4 females (USNM, INHS, FSCA), same, but 15 March 1984, 3 males, 10

females (USNM, INHS, UA, ROM), same, but 13 March 1984, 3 males, 2 females,

same but 19 March 1982, 1 male, unnamed tributary to Wallace Branch, 5 miles

southeast Berry (RIOW) 7117S, $2); 15 March 1984, S$. Harris; 3 males,.1>féemale

(USNM, INHS)." See type material examined below.

Material examined. Paratypes: male, 19 III 1982, SCH, (SCH); male and

female, 13 March 1984, SCH, (JSW). | wish to thank Steven C. Harris for kindly

giving me the latter paratypes.

Lepidostoma (Nosopus) Unicolor Group

This group comprises five known species, L. (N.) bakeri, cinereum, frosti,

knulli, and unicolor.

Male. Head: (F234). Scape not highly modified ino ts (N.) frosti and

unicolor, but in L. (N.) bakeri, cinereum and knulli having retractable spherical

mesal- process: (f. 33). Maxillary palps and wings (f. 47) similar to those of

(N.}) modestum group. Genitalia (f. 74-78) having inferior appendage unique,

with large thumblike basodorsal process extending posteriorly about as far as apex

of first article, first article which is short and broad in lateral view, second

article small subapical rounded lobe, horizontal middorsal process present.

Female. Genitalia (f. 110-114) similar to those of several other members of

L. (Nosopus), such as bryanti and americanum. Wings (f. 47). Spermathecal

sclerite in ventral view oval, elongated longitudinally.

Immatures, case and habitat. The larvae of all five species of this group

have been observed. Final instar larvae of the L. (N.) unicolor group construct

a turret case-(f. 5, 6). > However, oneseries of final instar larvae’ of L. (N-)

cinereum with stone cases only. The turret case consists of several transverse

plant fragments arranged in a random whorl about the exterior, resembling the

turret of a lycosid spiders. Anderson (1976) referred to this larval case as the

"log-cabin type.”

Distribution. Two species, L. (N.)}) cinereum and unjcolor have transconti-

nental distributions, occurring throughout western and northeastern North

America; L. (N.) frosti is strictly eastern; L. (N.) bakeri and knulli are endemic

to the Southwest.

Lepidostoma (Nosopus) bakeri Flint

(fy FAs FIO)

Lepraostome baker! Flint 1965:175; f2.10A-B» (male)s° Flint 1967e:1752 Denning

166269 -C9p; ts) -7C;. Plintcand: Buena. 1977. sp. 1377, 378=380.

Male. Scape with small spherical retractable mesal process, similar to that

of L. (N.) cinereum. Maxillary palp, 1.1 mm long. Wings brown. Anterior wing

about 8.0 mm long. Genitalia (f. 74) with segment X similar to those of L. (N.)

cinereum and knulli, but first article of inferior appendage with small hooked

apex.

Female. Genitalia (f. 110)' similar to those of other «species of L. °(N.)

unicolor group. Segment IX in lateral view with attenuated anterior apodeme

Weaver: North American Lepidostomatidae 4]

curved ventrad. Spermathecal sclerite oval in ventral view, elongated

longitudinally.

Immatures, case and habitat. Final instar larva head light brown with few.

lateral muscle scars. Mesonotum also light brown with posterior margin black,

similar to that of L. (N.) knulli. Hind leg with many long setae. Case consists

of small transverse plant fragments, similar to that of L. (N.) frosti and

unicolor.

Life history. Larvae and pupae collected in July. Recorded flight dates:

11 April - 8 September.

Distribution. Arizona, Colorado, Mexico, and Guatemala.

: Type data. Flint: 41965) : "Holotype, male: Arizona, Portal, Southwest

Research Station, 16 June 1963, J. H. Baker. United States National Museum type

6741/7. Paratype, male: same data.”

Material examined. ARIZONA: Cochise Co: Southwestern Research Station,

2 mi SW. of Portal, male paratype, 16 V1 1963, JHB .<CUSNM): mates 6 females.

hE AN=e2T Vile 19965, (CUSNM) 6 males; 10 females: 158 1X%:1974. “He M7, CCveC).

COLORADO: Jackson ‘Co? Gould; 2 mates, 2 demales, 4-Vill 1974. H 6 MT,

(CUCC). MEXICO: La Marquesa, La Cruces Natl Pk, many larvae, 5-9 VII, OSF

& Ortiz, (USNM); VERACRUZ: La Joya Rt. 140, Km 307, male pupa, many larvae,

21: Wit 1965; OSEs Grtiz.. (USNM):

_Lepidostoma (Nosopus) cinereum (Banks)

Gee ee opera a

Silo cinereus Banks 1899:210-211.

Lepidostoma cinereum: Fischer 1970:40-41.

Lepidostoma strophis Ross 1938a:177, f. 120 (male) NEW SYNONYM; Ellis 1962:5,

14;. Fischer. .1970:65; Newell and Potter 1973220; Hilsenhoff 1975:25;

Anderson 1976:122; Nimmo and Scudder 1978:120; Roy and Harper 1979.

‘p. 163; Webb 1980:98; Hilsenhoff 1981:31; LaFontaine 1981:251; Baumann

and Unzicker 1981:27; Roemhild 1982:11; Marshall and Larson 1982:79,

f. 156; Nimmo and Scudder 1983:80; Nimmo and Wickstrom 1984:4;

Herrmann et al. 1986:435; Nimmo 1986a:2; Nimmo 1986b:96; McElravy and

Resh 1987:85: °88.

Lepidostoma strophe: Unzicker et al. 1982:9.107.

The male and female associations of this species and its known distribution,

suggests that L. strophe (based on the male type) is a junior synonym of L. (N.)

cinereum (previously \known only from the female holotype).

Male. Scape (f. 33) with retractable spherical mesal process similar to those

of L. (N.) bakeri and knulli, 0.4 mm long. Maxillary palp 0.7 mm long. Anterior

wing, /.5 mm long. Genitalia (f. 75) having segment X in lateral view smoothly

curved with venter linear, being similar to that of L. (N.) bakeri. Inferior

appendage with incised rounded apex, similar to that of L. (N.) knulli.

Female. Genitalia (f. 111) with spermathecal sclerite in lateral view oval,

elongated longitudinally, similar to those of L. (N.) bakeri and knulli.

Immatures, case and habitat. Larval head light brown. Mesonotum light

brown, without black posterior margin as in L. (N.) knulli and bakeri. Legs light

brown, midleg tibia with distal apex dark brown. Case of final instar variable,

usually turret type, similar to that of L. (N.) knulli with fewer long transverse

pieces, but sometimes made only of stones. Several larvae collected from a pond

with turret cases, few larvae from a stream with stone cases.

Life history. Larvae collected 12 May to 20 August, some reared to pupae

and adults in June. Recorded flight dates: 22 March - 9 November.

Distribution. Alaska, Alberta, British Columbia, California, Colorado,

Idaho, Maine, Michigan, Montana, Newfoundland, Northwestern Territories,

Oregon, Quebec, Saskatchewan, Utah, Washington, Wyoming, and Yukon. Record

42 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

from Georgia (Unzicker et al. 1982) is a female specimen which is probably L.

(N.}) americanum.

Type data. Silo cinereus Banks (1899): "Gazelle, California, September

(Morse)." New Lectotype, female: Gazelle, California, September 1897. MCZC

Type 11691. See type material examined below.

Lepidostoma strophis Ross (1938a): "Holotype, male. - Beulah, Michigan:

Sept, 16; 1936, Ross € Burks. ““Pardtypes: - British: Coltimbia.+ Cultus’ Lake:

June 5, 1927, H. H. Ross, 2 males. Michigan - Same data as for holotype, 1

male." Holotype at the INHS (Webb 1980). Paratype male at the INHS (pers.

comms J bcUnzioker):

Material examined. Lectotype female of S. cinereus, (MCZC). ALASKA:

Prince. of Wales Island, N Fork Stoney Cr, 7 males, 12-18: Vill, DGD, (DGD); 9

mit Evof Kefai, male, 8. Vill 193%). WM Bryant; (USN): ALBERTA: 2.5 mi: £ of

Nordegg, Shunda Cr Cpg, 22 males, female, 13 VIII 1968, 5 males, 8 VIII 1965;

35 males, 3 females, 8 VIII 1965, APN, (UASM); Cold Cr, Hwy 16, Nojack, male,

3 females, 9 VIII 1967; 2 males, 3 females, 22 VIII| 1967, APN, (UASM). BRITISH

COLUMBIA: 22.6 km E of Princeton, Wolfe Cr, Hwy 3, male, female, 14 IX 1979,

APN, CONG)... GALIFORNIA:” Eb Dorado: Cot. Fallen Leaf Lake; male, 20. Vi 1961,

SDS» (UICM);. Los Angeles Cor. Topanga Cyn, 300. larva;) 20 VII 1965, WP Vann;

(INHS):>~ Plumas: Co: Quincy, Spanish Cr, male, -# females, 26> Vi 1973; DRGs

SDS, (SD8)s (Santa Clara Cos, Alum Rock (Park, mate. Pow I1960,.SDS, (UCM);

Siskiyou Co: Shasta River, Weed, 6 males, 29 V 1967, SGJ, (USNM). Sonoma

Co: Fairfield, Osborn Preserve, Copeland Cr, 1360', larvae in pools on rocks

with stone cases only 20 VIII 1980 (adults reared), LES, (LES). IDAHO: Adams

Co: New Meadows, Little Salmon River, 5 males, 2 females, 10 IX 1964, SDS §&

WFBpfUICM). (SDS). Bonner: Co: «Hope, Lighting Cr, male, 16 1% 19699¢SDS ‘6

WFB, (UICM); Boundary .Co: 5 mi SW of Bonners Ferry, Deep Cr, male, 16 1X

1969,-SDS: 8 WEB, (OICM): Butte: Coc. Craters: Moon Nath’ Monument,;'°S males: 2

females, 11 VIII 1965, SDS, (UICM); Gooding Co: Thousand Springs, Riley Cr,

mates sided Vs 1865, “SDS {CCUICM) > “tdahe “Cot Fishes Gr). mate. Ocho 1064..: WEB,

(UICM); Shoshone Co: 3 mi E of Kellogg, Moon Cr, 8 males, female, 16 IX 1964,

SOS ACU Gis weer alts Coe ol mie WoofeBulls. male, ? T2014 Vereesa. SDS}. CUICM).

MICHIGAN: Montmorency Co: Hunt Cr, 38 males, 30 VIII-3 IX, JW Leonard,

(INHS). MONTANA: Flathead Co: Glacier Natl Pk, McDonald Cr below McDonald

Lake, male, 10 IX 1968, RWB, (USNM). NORTHWEST TERRITORIES: = Simpson

Island, Great Slave Lake, male, 20 VIII 1964, D Larson, (UASM). OREGON:

Benton Co: MacFadden Pond, 12 mi S of Corvallis on Hwy 99, 5 larvae, 5 pupae,

12 V 1971 (some adults emerged V!), ED Evans, (ROME); Corvallis, 35th Street,

Oak: Cr, male, 13 Vi 19861, D Walters, (QSUO); Harney Co: > 8 mi‘N of Burns;

male, 15 VI! 1965, KG, (OSUO); Lane Co: Springfield, 1960, 4:males, 8 females,

(USNM);°12 mi: SEvof Eugene, male; 121%: 1966, KG), (OSUO); .kinn Co: Albany,

Western Kraft Streams, 5 males, 3 females, 10 V 1972, D Craven, (OSUO); Marion

Connmumevitle, mate, 6 VI oM6ys KG: €QSU0); Niagra, male, 37°V 1970, E Evans;

(OSUO); 12 mi S of Salem, 2 males, female, 4 X 1968, KG, (OSUO); Josephine

Co: Grants Pass, male, 24 V 1964, KG, (OSUO). WASHINGTON: Cedar Grove,

Cedar Rinvery \\2)cmales;> 3 xkIn1972) ARG) Malick, Stansbury, SDS, (SDS): ‘Chelan

CO! Lake Wenatchee, 9 males, female, 27 X 1968, SDS, (SDS); Grant Co:

Vantage, Koleche Lake, male, 6 V 1969, J Gooding, SDS, (SDS); Kittitas Co: 4

mi W of Ellensburg, Yakima River, male, 2 V 1969; male, 7 IV 1969; 2 males, 2

females, 14 X 1970, SDS, (SDS); Ellensburg, Carey Pond, 2 males, 2 females,

Zoe HOO, (SOS oWSW yr Eilensburnay: golf ehurse, 09 males, FOYIK® 1969, SDS;

(JSW); Squaw Cr & Yahima River, male, female, 4 VLII 1977, SDS, (SDS); Pierce

Co: Clover Cr, Hwy 7, near Spanaway Lake, 2 males, 14 VI 1969, RWB, (USNM).

WYOMING: Yellowstone Natl Park, Madison River, male, female, 29 VIII 1963,

JR Heaton, (USNM).

Weaver: North American Lepidostomatidae 43

Lepidostoma (Nosopus) frosti (Milne)

Cro ye fo 192)

Atomyia frosti Milne 1936:119-120, 122, 124 (male).

Lepidostoma frosti: Flint & Wiggins 1961:295-296, f. 12A-B (female); Blickle and

Morse 1966:11; Wray 1967:57; McConnochie and Likens 1969:150; Fischer

1970:42-43; Roy and Harper 1975:1084; Etnier and Schuster 1979:19; Roy

and Harper 1979:162; Neves 1979:171; Unzicker et al. 1982:9.106, 9.107.

Male. Scape unmodified, cylindrical, 0.4 mm long. Maxillary palps 0.7 mm

long. Anterior wing about 7.0 mm long. Genitalia (f. 76) similar to those of L.

(N.) unicolor, but segment X in lateral view with slender posterior apex, square

notch in posteroventral margin.

Female. Genitalia (f. 112) similar to those of L. (N.) americanum.

Spermathecal sclerite in ventral view oval elongated longitudinally.

Immatures, case and habitat. Larval head light brown with mesal pattern

of dark spots, venter (f. 9). First and second larval instars construct cases of

fine sand grains. Later instars make a turret case (f. 6) composed of short

transverse plant fragments similar to that of L. (N.) unicolfor. Larvae inhabit

springs.

Life history. A concentrated population of L. (N.) frosti at Maul Spring,

a constant temperature spring in Pennsylvania, was sampled monthly with three

1 ft? benthic samples and light-trap collections. The first instars occurred 14

August to 5 November, second instars 14 August to 27 March, third instars 11

October to 27 March, fourth instars 3 December to 19 June, fifth instars 28

February to 16 July, pupae 27 May to 14 August. Recorded flight dates: 20

May - 21 September.

Distribution. Maine, Massachusetts, New Hampshire, New York, North

Carolina, Nova Scotia, Pennsylvania, Quebec, Rhode Island, and Tennessee.

Type data. Milne (1936): "Holotype male ‘Catskills, N.Y., VII-1874, O.

Sacken', 1 male paratype ‘White Mts., N.H., VI, Morrison’, 1 no data but

21-1X=1900) in the MeO Zs. NineJether males tram: P.O, Nevievand MOWe-29-VI

te :26-VIl, 2) im the Can: Nat. collection; 1 ine the Ameri: Mus.\Nat. Hist. 6 in

the author's collection." Male and female syntype at the INHS (pers. comm. J.

D. Unzicker). | have seen the holotype and 6 paratypes, 2 males and 4 females,

at the MCZC, Type 22276. However, the labeled holotype is from New Hampshire,

Pinkham, 26 VII 1930, C. A. Frost, differing from locality data given by Milne

(1936).

Material examined. MASSACHUSETTS: Hampshire Co: Westhampton, larva,

20 Me, TED Rie ae 2B IY POR kl eat AL eA NEW HAMPSHIRE:

Pemigewasset River, 22 VI 1941, 8 larvae, THF & HHR, (INHS); Coos Co: Mt

Washington, Jefferson Notch Road, Abenaki Brook, male, 5 females, 7-8 VII 1977,

JSW, (JSW); ibid, Mt Pleasant Brook, male, 9-10 VII 1977, JSW, (JSW). NORTH

CAROLINA: Macon Co: Wayah Crest Cpg, larva, 30 V 1981, JSW & RWH, (JSW);

Yancey Co: Black Mts, S Toe River, male, 1 VII 1979, JSW, (JSW); Mt. Mitchell,

Camp Alice, 4 larvae, 9 VI 1961, OSF, (USNM). RHODE ISLAND: Washington

Ce: Richmond, male, female, 30 VI’ 1971;°2 males, 20. 'V1 1971, AG Lavalle,

(OCG). PENNSYLVANIA: Westmoreland Co: Rector, Powdermill Nature

Reserve, Maul Spring, 2 males, 3 females, 20-21 V 1977; 5 males, 7 females, 31

V 1975; 22 males, 40 females, 19-20 VI 1975; 3 males, 2 females, 22 VI 1977; 2

male pupae, 2 female pupae, 6 VI 1975; 23 males, 31 females, 16-17 VII 1975: 5

males, 20 VII 1977; 7 males, 19 females, 13-14 VIII 1975; larvae, 28 1-22 V 1975:

pupae 19 Vied4o Wi TERS: tape) IO LK) POF arait. MOP ey SMe Ee CISWY.

TENNESSEE: Sevier Co: Great Smoky Mts Natl Pk, Walker Prong, W Prong Little

Pigeon River, 4500’, male, female, 4 Vil 1979, JSW, (JSW); Great Smoky Mts Natl

Pk, 10.2 mi -& of ‘Gatlinourg, larvae, 20 V: 1970; OSF,(USNM).

44 Contribs Amer Ent>inst.,° Volume 24, Number 2, 1988

Lepidostoma (Nosopus) kKnulli Ross

(fh ries 743)

Lepidostoma knulli Ross 1946:269, 280, f. 10A-C (male); Flint 1967c:175; Fischer

1070: 50 First and Bueno 19772377... 361-362; Webb. 1960°97;: Waltz: and

McCafferty 1983a:414.

Lepidostoma leechi Denning 1962:37-38, f. 5SA-C (male); Flint and Bueno

12772381 (as syn.of Ly knulli).

Male. Scape 0.8 mm long, modified with retractable spherical mesal process

similar to that of L. (N.) cinereum and bakeri. Maxillary palps 0.9 mm _ long.

Anterior wing 8.2 mm long. Genitalia (f. 77) having segment X similar to that

of bakeri with many dorsal tubercles, and inferior appendage similar to that of

L. (N.) cinereum with rounded apex.

Female. Genitalia (f. 113) similar to those of L. (N.) bakeri. Spermathecal

sclerite in lateral view elongated dorsoventrad, in ventral view oval and elongated

longitudinally. Segment |X with short blunt apodeme.

Immatures, case and habitat. Larva having mesonotum light brown with dark

posterior margin similar to that of L. (N.) bakeri. Turret case (f. 5) with many

long transverse plant fragments. Based on collection data immatures appear to

inhabit small streams.

Life history. Several larvae and pupae were taken 8 to 18 June. Recorded

flight dates: 5 May - 18 December.

Distribution. Arizona, Durango, Michoacan, Mexico State, and New Mexico.

Type data. Lepidostoma knulli Ross (1946): "Holotype, male. - Oak Creek

Canyon, Arizeda, Aug. 15, 1988) D..3. 6 doN. Knull. Holotype at the INHS

(Webb 1980). See type material examined below.

Lepidostoma leechi Denning (1962): "Holotype male, one mile west of La

Marquesa, Mexico, approximately 9200 feet elevation (34 kilometers west of Mexico,

D.F.), December 8, 1948, H. B. Leech. Type deposited in the collection of the

California Academy of Sciences, San Francisco, California."

Material examined. Holotype male (INHS). ARIZONA: Cochise Co:

Chiricahua Mts, Southwestern Research Station, 5 mi SW of Portal, Cave Cr, male,

9 VI 1961, JK, (INHS); Chiricahua Mts, Rucker Canyon, male pupa, 7 larvae, 8

Vi1968, Menke. €OSF, GJSNM): Oak Cr Canyon; Halfway Camp; 9 larvae, 18

V |: 1968; OSE. 6 Menke,.<GuSNM): -Cocohine’ Co: W Fork Oak=Cr, 5: females, 19

Is 1979: G-males. female,.49 xX -19/8, MWS... (JSW).. NEW MEXICO:-. Catron Co:

Gila Natl Forest, Scorpion Camp, W Fork. Gila River, male, 5-6 V 1981, WP

McCafferty & AV Provonsha, (PURC).

Lepidostoma (Nosopus) unicolor (Banks)

CP iB4, 48 dA)

Mormomyia unicolor Banks 1911:357, pl. 12 f. 21 (male).

Lepidostome unhicolor: “Etrier 1965:151;. Anderson. .1967:517, (case): Fischer

1970:6/-68; .. Anderson: (and. Wold: 1972: 792, 1972198: Hilsenhoff 1975: 25;

Anderson: 19/76:123:. Anderson: et aly 619078: 71,.. 727 Nimmo. and .Scudder

1978::120;:Resh and Song 1978: 396; ‘Roy and Harper 19/9:163%. 'Grafius and

Anderson 1980:808-816; -Hilsenhoff 1981:31; LaFontaine 1981:249, 251;

Bauman: “ands -Unzieker..1961:27::. Roemhild .19822115 » Anderson” et al:

1962: 14> Nimme and Scudder .1983:74.. 80; Anderson. et.al. 1984: 15, 183

Ruiter and Lavigne 1985:43; Herrmann et al. 1986:435.

Male. Head (f. 34) with scape cylindrical and unmodified, 0.5 mm long and

maxillary palps 0.7 mm long. Wings (f. 47) with venation similar to that of L.

(N.}) modestum. Anterior wing 9.0 mm long. Genitalia (f. 78) similar to those

Weaver: North American Lepidostomatidae 45

of L. (N.) frosti, but segment X with acute angle in ventrolateral margin in lateral

view.

Female. Wings (f. 47). Genitalia (f. 114) similar to those of other species

of unicolor group. Spermathecal sclerite somewhat triangular in ventral view.

Immatures, case and habitat. Larva mesonotum brown without black posterior

margin, setae of hind leg not as numerous as in L. (N.) bakeri. Contributions

to the bionomics of L. (N.) unicolor have been documented by Anderson (1967),

Anderson and Wold (1972), Anderson (1976), Anderson et al. (1978), and Resh

and Sorg (1978). Larvae inhabit mountain streams. Final instar larvae construct

turret cases similar to those of L. (N.) knulli, but sometimes with fewer long

transverse pieces.

Life history. Larvae occur from October to July, pupae April to 3 Septem-

ber. Recorded flight dates: 9 April - 27 September.

Distribution. Alberta, Arizona, British Columbia, California, Colorado,

Idaho, Minnesota, Montana, New Mexico, Oregon, Quebec, Saskatchewan, South

Dakota, Utah, and Wyoming.

Type data. Banks (1911): "From Switzer's Camp, San Gabriel Mountains,

California, June (Grinnell coll.)." A female syntype is at the INHS (pers. comm.

Jo2D. Unzieker):

Mormomyia unicolor Banks, New Lectotype, Switzer's Camp, San Gabriel

Mountains, California, June, MCZC Type 11686.

Material examined. ALBERTA: Carrot Cr, old Hwy 16, 3 males, 1 female,

29 Vil 1976,, APN, CUASM)s: Cold Gr} Hwy-16, Nojack» male, female ;-25 wll 1976;

APN, (UASM); Waterton R at bridge, Hwy 5, Waterton Lakes Natl Pk, male,

female, 31 VII 1976, APN, (US). ARIZONA: Cochise Co: Herb Martyr Cea. 7

mi W of Portal, 4 larvae, male & female pupae, 23 VI 1966, GBW, (ROME);

Southwestern Research Station, 5 mi SW of Portal, Cave Cr, male pupa, 7-10 VI

1968, OSF & Menke, (USNM); Chiricahua Mts, E Turkey Cr, many larvae, 9 VI

1968, OSF & Menke, (USNM). BRITISH COLUMBIA: U BC Exp Forest, Spring

Cr, male, 28 VII 1981, K Van Frankenhuyzen, (OSUQ); Vancouver Island, Tsable

R, Hwy 19, S of Courtney, male, 8 VII 1979, APN, (UASM); Vancouver Island,

28.2 km W of Caysuse, male, 7 VII 1979, APN, (UASM); Shannon Cr, Hwy 99,

2.4 km, S of Squamish, male, 12 VII 1979, APN, (UASM); Hwy 97, 9 km N of

Lac la Hache, male, 14 VII 1979, APN, (CNC). CALIFORNIA: Modoc Co: 5 mi

Evof Likely, S Ferk. of Pit River, 1/+males; 13° females, S<1xX 1069, AB Gurney,

(USNM); Mono Co: Barnes Lake, 15 mi W of Bridgeport, male, female, 20 VII

1979, SDS, 4$D8d; ‘San Diego Go: -Petreros, 5 maless 12: females,’ 9 Hyo1974, #H

& MT, (CUCC); Santa Clara Co; Guadalupe Cr, male, 6 VIII 1960, SDS, (UICM);

Alum Rock Park, male, VI 1960, SDS, (UICM); Sonoma Co: Fairfield, Osborn

Preserve, Copeland Cr, 1540', larvae in pools 27 IV 1980 (adults reared), LES,

(LES). COLORADO: Routt Co:»:Steamboat Springs Fish Cry males 11 VIM 1973.

DLP € SDS, (SDS). 'IDAHO: Custer Co: 7 mi-S Stanley, Red) Fish take: 2 males:

3 females ¢:2.Milb. 4964, SDS, (JSWiee ibid; male, 223 Vil. 19657-SDS: CUICM)* Idaho

Co: Grave Park, Selway-Bitterroot Wilderness Area, male, 21 VII 1969, LM §&

WB Potter, (USNM); ibid, Wind Lake outlet stream, 3 males, 25 females, 4 IX 1980,

SDS, (JSW); Kootenai Co: 3 mi NE Coeur d'Alene, male, 22 VI 1961, RE Stecker,

(UICM); ValleywCo:- Sof. Yellow Pine, male, female, 3 VItl 1964, SDS & RULW;

(SDS ).-4:MONTANA®?. Lineeln) Cot «vLibby,p «Libley Cr) So males. 12 femates> 1901

1986, DLG, (JSW). -OREGON: Baker: Cor. Blue Mt., Spring Cr, mate; 23-1 1974;

maley, 16:Vbbb 1969, JHB, (USNM);>Benten: Con: 7. ami NW: of: Comvaltie) male, 27

Vili-3 1X 1968, CD Kerst, (USNM); Crook Co: —-Ochoco Mts near summit, Hwy

26, male, 31 VII 1965, SGJ, (USNM);. Lane Co: HJ Andrews Exp For, Mack Cr,

2 males & 3 females reared, 15 VI 1973, NHA, (OSUO); Lineoln Co: Flynn Cr,

pupa in soft wood 19 VIII 1981 (male emerged 23 VIII), RWW, (OSUO); ibid,

larvae? prepupae? male pupa, 22 VI 1982, SB & AL Flotsam; ibid, 10 mi S of

Toledo, male pupa from decaying wood, 12 VII 1977, TL Dudley, (OSUO); ibid,

Oak Creek Drippery, reared from eggs, male pupa & male emerged 15 VI 1974,

(OSUO); Needle Branch Cr, 10 mi 5 of: Toledo, larvae 24 tV 1974, (male puna,

46 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

7 males, 6 females, emerged VI-VII), NHA, (OSUO); Jefferson Co: Metolius

River, 1 mi S of Camp Sherman, male & female pupae, 3 IX 1964, NHA, (OSUO);

Union Co: Goose Cr, 15 mi E of Medical Springs, male, female, 22 VIII 1975;

male, 5 (3078, MRB, > €SDS)iatick Cr, 6 ombE of: Medical: Springs, males €

females, 29 VII-5 IX 1975, MRB, (SDS); Jordan Cr, 20 mi SW La Grande, males

& females, 25 VII-18 VIII 1975, MRB, (SDS); Velvet Cr, 8 mi E of Medical Springs,

males & females, 22 VII-19 VIII 1975, MRB, (SDS). SOUTH DAKOTA: Spearfish

Cr, male, 11 females, 7 VIl 1965, RW Hodges, (USNM). UTAH: Salt Lake Co:

Lhevsand Springss:Mub oCr «Canyon; males 2orWilloe 197, RWB, “CUSNM):

WASHINGTON: Kittitas Co: Dickey, Tearway Cr, male, 7 |X 1968, SDS, (SDS);

6.5 mi N of Ellensburg, Wilson Cr, male, 29 VII 1980, DRG, (SDS); Lewis_Co:

5 mi E Randle, Davis Cr, male, 24 VII 1980, SDS, (SDS). WYOMING: Yellowstone

Natl Park, Madison River Junction, male, 2 females, 21 VII] 1955, KM Fender,

(USNM); Teton Co: near Moose, Grand Teton Natl Pk, male, 4 VIII 1973, DLP

SiS Po CSDS) .

Lepidostoma (Nosopus) Mexicanum Group

This group comprises L. (N.) acarolum, aztecum, dafila, delongi, denningi,

frontale, facinatum, leonilae, mexicanum, oaxacense, quila, rectangulare,

reimoseri, steinhauseri, and talamancense. However, L. (N.) acarolum and

delongi are only tentatively included within this group.

Male. Head (f. 35) with scape cylindrical, not highly modified. Maxillary

palps with segmentation obscure, mesal surface bearing scales, apical portion

usually spatulate, but triangular in L. (N.) mexicanum. Wings (f. 49-53) having

forewing with venation similar to that of L. (N.) unicolor group, anterior costal

cell usually absent (f. 53), but sometimes with small basal pocket (f. 52); however

L. (N.) frontale with long narrow anterior marginal fold (f. 51). Hindwing

unique, having both Fl and F2, but in L. (N.) acarolum (f. 50) and delongi (f.

49) F2 absent; 1A and 2A not fused together completely (except in L. [N.])

talamancense). Genitalia (f. 79-91) with inferior appendages variable; L. (N.)

mexicanum and frontale with first article broad basally, venter truncated, short,

with slender horizontal dorsal process extended posteriorly, and with apex usually

tricuspid in dorsal view; most other species of L. (N.}) mexicanum group with

serrate subapical lateral flange.

Female. Only females of L. (N.) acarolum, lacinatum, and mexicanum known.

Genitalia (f. 115, 116) having spermathecal sclerite similar to that of L. (N.)

unicolor group, but longer, with ventral keyhole-shaped mesal process about 2/3

as long as sclerite in ventral view. Several unidentified females from Central

America, (probably of the L. [N.] mexicanum group) with similar long

spermathecal sclerite.

Immatures, case and habitat. Larval head with distinct carina in L. (N.)

acarolum and talamancense, weak carina present in L. (N.) mexicanum. Larvae

of other species unknown. Some unidentified larvae from Central America (and

probably of this group) with metanotal sa2 sclerite bearing three or more setae,

sometimes without carina on head. Larva of L. (N.) acarolum, mexicanum, and

talamancense construct cylindrical cases of small stones (f. 3). Most other uni-

dentified larvae of the L. (N.) mexicanum group will probably key to couplet 11

in the larval key. of Lepidostoma species herein. However, | have examined

several other unidentified larvae from Central America which construct cases of

plant fragments.

Distribution. The majority of species in the L. (N.) mexicanum group occur

in Central America. However, a few species, such as L. (N.) mexicanum and

delongi have ranges extending into the southwestern United States, and one

species, L. (N.) acarolum, is known only from Arizona and New Mexico.

Weaver: North American Lepidostomatidae 47

Lepidostoma (Nosopus) acarolum Denning

Lf. By 26 ee 145)

Lepidostoma acarola Denning 1962:38-39, f. 6A-B (male).

This species is tentatively included in the L. (N.) mexicanum group.

Male. Scape not highly modified, cylindrical, 0.7 mm long. Maxillary palps

slender and spatulate, 0.7 mm long. Wings (f. 50) brown, venation similar to

that of L. (N.) unicolor. Anterior wing about 7.5 mm long, not highly modified.

Hindwing without F2. Genitalia (f. 79) having segment X in lateral view with

dorsum short and curved caudad into narrow upturned ventroapical point, venter

sinuate and longer than dorsum. Parameres rather broad in lateral view. Inferior

appendage similar to that of L. (N.) bakeri, with hooked apex, but with shorter

basodorsal process.

Female. Genitalia (f. 115) with spermathecal sclerite in lateral view elongated

longitudinally; in ventral view, posterior end truncated and anterior end tapered.

Immatures, case and habitat. Larval head brown, cylindrical, vertex flat

with circular carina. Case (f. 3) cylindrical, slightly curved posteriorly and

composed of small stones. Case of final instar with anterior width 2.1-2.3 mm,

posterior width 1.2-2.0 mm, and length 9.0-11.0 mm.

Life history. Several larvae and pupae collected in June. Recorded flight

dates: 13 - 25 August.

Distribution. Arizona and New Mexico.

Type data. Denning (1962): "Holotype male, Cochise County, Arizona,

Southwestern Research Station, five miles northwest of Portal, August 25, 1960,

D. C. Rentz. Allotype. - female, same data as for holotype. Paratypes (1 male,

1 female) same data as the holotype. Holotype and allotype deposited in the

collection of the Academy of Sciences, San Francisco, California."

Material examined. ARIZONA: Cochise Co: Southwestern Research Station,

29 mi SW of Portal, seep, male pupa, 50 larvae, 7-8 VI 1998, -OSF -& Menke,

(USNM). NEW MEXICO: Catron Co: Glenwood, 2 males, 5 females, 13-14 VIII

1978, MWS, (JSW); | wish to thank Milton W. Sanderson for giving me these

specimens.

Lepidostoma (Nosopus) aztecum Flint & Bueno

Chi 80}

Lepidostoma aztecum Flint and Bueno 1977:376, 378, f. 2-3 (male): Bueno and

Padilla 1981:391.

Male. Scape cylindrical, 0.6 mm long. Maxillary palp similar to that of L.

(N.) modestum, 0.6 mm long. Wings with venation similar to that of L. (N.)

lacinatum. Anterior wing 10.7 mm long. Genitalia (f. 80) similar to those of L.

(N.) lacinatum. Segment X in lateral view with two short dorsal processes.

Inferior appendage main article in lateral view with hatchet-shaped apex and

pointed bifid subapical lateral flange.

Female. Unknown.

Immatures, case and habitat. A male pupa close to L. (N.) aztecum was

examined with a cylindrical case of small stones similar to that of L. (N.)

mexicanum.

Life history. Pupa (possible) collected in August. Recorded flight dates:

2 June - 7 November.

Distribution. Morelos and Veracruz.

Type data. Flint and Bueno (1977): "Holotype, male: Mexico, Morelos,

Lagunas Zempoala, 2 June 1976, J. Bueno-Soria. USNM Type /4073. Paratype:

Same data, 1 male." See type material examined below.

48 Contrib. Amero Ent. Inst... Velume 24, Number 2, 1988

Material examined. Holotype male, (USNM). VERACRUZ: Perote, male, /

IX 1977, JB, CUSNM); La Joya, Rt 140, km 307, (probable) male pupa, 4 pupae,

21-Wit- 1965, °OSF © Ortiz, (USNM).

Lepidostoma (Nosopus) dafila Bueno & Contreras

Lepidostoma dafila Bueno and Contreras 1986:209-210, f. 7-9 (male).

The following is based on the original species description. This species ts

closely related to L. (N.) heveli, having similar male segment X. It differs from

all Known members of the L. (N.) mexicanum group by lacking a_ basodorsal

process of the inferior appendage.

Male. Scape cylindrical. Maxillary palp unsegmented with basal portion

narrow and cylindrical and apical portion broader and ellipsoidal. Anterior wing

8 mm long. Posterior wing without F2 (pers. comm. Bueno). Genitalia: Segment

xX in lateral view having two linear processes nearly parallel to each other, sub-

equal in length, directed posteriad, and separated widely by U-shaped notch;

dorsal process with pointed apex and more slender than ventral process; ventral

process bearing subapical trapezoidal extension ventrad and with fingerlike apex.

Phallus with slender parameres about half as long as_ phalicata. Inferior

appendage in lateral view having main article broad and rectangular, dorsum with

short extension posteriad, without basodorsal process, horizontal middorsal

process broad and acuminate.

Female, immatures, cases and habitat. Unknown.

Life history. Holotype collected 17 June.

Distribution. Oaxaca. Only Known from the type locality.

Type data. Bueno and Contreras (1986): "Holotipo macho. Mexico:

Oaxaca, Finca Pacifica Mpio. de Pluma Hidalgo, recolectado el 1/7 de junio de 1982

por E. Barrerra. Depositado en el IBUNAM."

Lepidostoma (Nosopus) delongi Ross

(ft: 49, 31)

Lepidostoma delongi Ross 1946:270, 283, f. 12A-B (male); Fischer 1970:42; Flint

and Bueno 197/7:377, 380; Webb 1980797: Bueno and Padilla 1981:39T.

This species is tentatively included in the L. (N.) mexicanum group.

Male. Scape, cylindrical, not highly modified, 1.0 mm long. Maxillary palps

fingerlike, concave mesially, 1.2 mm long. Wings (f. 49) with venation similar

to that of L. (N.) roafi and unicolor. Anterior wing about 7.0 mm long. Genitalia

(f. 81) having segment X in lateral view with dorsum slightly curved, bearing

two or three small tubercles, posterior with obtuse apical notch, venter nearly

linear with a square apical angle. Parameres spinous. Inferior appendage narrow

with apex bifid in dorsal view.

Female, immatures, case and habitat. Unknown.

Life history. Recorded flight dates: 7 February - 10 December.

Distribution. Chiapas, Michoacan, and Morelos.

Type data. Ross (1946): "Holotype, male. - Zitacuara, Mexico, at light,

1941, D. M. Delong.” Holotype at the INHS (Webb 1980).

Type material. CHIAPAS: Finca Esperanza, 5 males, 12 V-5 VII 1938; male,

6 Ki 1636s. male, 10 Xile-1938; AD: <CINHS): Finca Germaniay: mate,” 2° V1 1935,

AD, GINHS)3¢Finca Vergel,. male, 12 44935 male, 23:V°1935;.. male; 28° V.1935;

2mates (7: Vii 4985). AD: CINHS) > Finca: Victoria, «male; i2° Vie 1935). AD)GINHS).

MORELOS: Park Nac. Zempoala, male reared, 7 II 1977, JB, (USNM).

Weaver: North American Lepidostomatidae 49

Lepidostoma (Nosopus) denningi Weaver n. sp.

(i582)

This species is closely related to L. (N.) rectangulare, but differs by having

male inferior appendage with apex of main article rounded, rather than rectan-

gular.

Male. Scape simple, cylindrical, 0.6 mm long. Maxillary palps spatulate,

0.6 mm long. Wings with venation similar to that of L. (N.) lacinatum. Anterior

wing 8.5 mm long, with small basal anterior marginal fold. Genitalia (f. 82) similar

to those of L. (N.) rectangulare. Segment X in lateral view with posterior margin

serrate. Parameres at least as broad as phalicata. Inferior appendage in lateral

view with subapical lateral flange oval, apex of main article rounded.

Female, immatures, case and habitat. Unknown.

Life history. Recorded flight date: 21 June.

Type material. Holotype: CHIAPAS: San Cristobal, (at light), male, 21 June

1926, AD, (USNM).

_ Etymology. This species is named after Donald G. Denning in honor of his

contributions to the taxonomy of Trichoptera.

Lepidostoma (Nosopus) frontale (Banks)

(fo S488)

Eremopsyche frontalis Banks 1901:367, pl. 7 f. 13, 14 (male); Fischer 1970: 83;

Plant: 967a223.

Lepidostoma frontaliss: Flint: 19G7as 24) fii F248 125 (male).

Lepidostoma frontale: Flint and Bueno 1977:375, 380+381; Bueno?’’and Padilla

1981:39%:

Male. Scape cylindrical, but bent near base, 1.2 mm long. Maxillary palp

about 1.2 mm long, apex spatulate with many black setae. Wings (f. 51): Anterior

wing with long narrow anterior marginal fold, 9.0 mm long. Genitalia (f. 83)

identical to some variants of L. (N.) mexicanum, but characters of the maxillary

palps and wings are diagnostic.

Female, immatures, case and habitat. Unknown.

Life history. Recorded flight dates: 1 August - 10 December.

Distribution. Hidalgo and Veracruz.

Type data. Banks (1901): "One specimen from Jalapa, Vera Cruz, Sep-

tember."

Eremopsyche frontalis Banks, New Lectotype, male, Jalapa V.C. sept y "MCZC

Type 11825. See type material examined below.

Material examined. Lectotype male, (MCZC). VERACRUZ: La Joya, Hwy

140, koe O07,” male, h Vides OSPR & Ortiz, (USNM); Rio Jamapa, 6 km N of

Coscomatepec, male, 2 V 1981, CMF & OSF, (USNM).

Lepidostoma (Nosopus) heveli Flint & Bueno

(42 ce)

Lepidostoma heveli Flint and Bueno 1977:376, 381, f. 4-5 (male).

Male. Scape not highly modified, cylindrical, 0.6 mm long. Maxillary palp

capitate; 0:6 mm: déng, Wing) venation’, (f,°.52). “similar to that of 1h) PNR)

lacinatum, but anterior wing, about 10 mm long, with small basal anterior marginal

fold. Genitalia (f. 84) similar to those of L. (N.) steinhauseri. Segment X in

dorsal view with narrow mesal and lateral processes. Parameres in lateral view

not quite as broad as middle of phalicata.

Female, immatures, case and habitat. Unknown.

50 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

Life history. Recorded flight date: 28 May.

Distribution. Guatemala. Only known from the type locality.

Type data. Flint and Bueno (1977): "Holotype male: Guatemala, Quiche,

El Quiche, 7.3 km S Chichicastenango (14°54'N, 91°0O7'W), 2400 mts., 28 May 1973,

Erwin & Hevel. USNM Type 74074." See type material examined below.

Material examined. Holotype male, (USNM).

Lepidostoma (Nosopus) lacinatum Flint

(F< 158) 85)

Lepidostoma lacinatum Flint 196/c:1/75-1/76, f. 24-25 (male); Denning 1973:142, f.

12 (male); Flint and Bueno 1977:377, 382.

Male. Scape simple, cylindrical, 0.7 mm long. Maxillary palp spatulate, 0.7

mm long. Wings (f. 53) anterior wing about 9.3 mm long, without anterior mar-

ginal fold. Posterior wing with Fl and F2. Genitalia (f. 85) similar to those of

L. (N.) aztecum, but segment X in dorsal view with shorter dorsal process.

Inferior appendage in lateral view with main article broad somewhat oval and

tapered into blunt apex, subapical lateral flange serrated with about six teeth,

mesal process in dorsal view narrow and acuminate with apex bent mesad.

Female. Scape 1.0 mm long. Forewing brown, 12.0 mm long. Genitalia with

spermathecal sclerite similar to that of L. (N.) mexicanum, but more oval in

ventral view.

Immatures, case and habitat. Unknown.

Life history. Recorded flight dates: 24 May - 31 October.

Distribution. Arizona, Durango, and Sinaloa.

Type data. Flint (1967c): ‘Holotype, male. Mexico (Durango), 10 miles

west of El Salto, 9000, 2 July 1964, J: ba Ho Martin’ Paratypes. Same ‘data,

3 males; 6 June 1964, 1 male; 8 June 1964, 2 males; 13 June 1964, 1 male; 17 June

1964, 3 males; 27 June 1964, 1 male; 7 July 1964, 1 male; 15 July, 1964, 1 male;

29 July 1964, 1 male. Mexico (Sinaloa), El Palmito, 15 July 1964, J. F. McAlpine,

1 male." See type material examined below.

Material examined. Male paratype, (USNM). ARIZONA: Cochise Co: 15

mS of Sierra Vista, Ramsey iGynys 6000, 2 males, 2iifemales:: 24 V°196/7s°7 males,

11 females, 20 VII 1967; 16 males, 9 females, 1-31 X 1967, 9 males, 17 females,

XI 1967s RE Sternitzky; CONC!) -

Lepidostoma (Nosopus) leonilae Bueno & Contreras

Lepidostoma leonilae Bueno and Contreras 1986:209, f. 4-6 (male).

The following is based on the original species description. It is not clearly

apparent that this species is closely related to any other particular known species

of the L. (N.) mexicanum group, but perhaps it is close to L. (N.) reimoseri.

It has male segment X similar to L. (N.) bryanti and some forms of L. (N.)

frontale. However, its male inferior appendage is more like that of L. (N.)

reimoseri, from which it differs by having main article more acuminate in lateral

view and lacking a‘subapical ectal point.

Male. Maxillary palp unsegmented and triangular in lateral view, perhaps

similar to that of L. (N.}) mexicanum. Anterior wing 9 mm long. Posterior wing

with R five-branched (pers. comm. Bueno). Genitalia: Segment X pentagonal

in lateral view, having an obtuse angle, anterior dorsum with about four

tubercles, posterior dorsum and venter smooth; posterior margin bearing two

prominent points (one ventral point and one mesal point) and two smaller dorsal

points, all points directed dorsad. Phallus with slender parameres about 3/4 as

long as phalicata. Inferior appendage in lateral view having main article slender

Weaver: North American Lepidostomatidae 31

and acuminate, bearing basodorsal process prominent and thumblike, and hori-

zontal middorsal process lobiform and about half as long as basodorsal process,

but without subapical ectal point or flange.

Female, immatures, cases and habitat. Unknown.

Life history. Holotype collected 16 March.

Distribution. Nuevo Ledn. Only known from the type locality.

Type data. Bueno and Contreras (1986): "Ho/lotipo macho. Mexico: Nuevo

Leon: Santiago Potero Redondo, recolectado el 16 de marzo de 1984 por A.

Contreras. Depositado en IBUNAM."

Lepidostoma (Nosopus) mexicanum (Banks)

CF) 35,08, 016)

Olemira mexicana Banks 1901:367; Flint 1967a:24.

Lepidostoma mexicana: Flint 1967a:24, f. 126, 127 (female holotype).

Lepidostoma mexicanum: Fischer 1970:59-60; Flint and Bueno 19772375, 382-383.

Atomyiodes bispinosa Ulmer 1911:25-26, f. 12-13 (male); Flint and Bueno

1977:382 (A. bispinosus as syn. of L. mexicanum).

Lepidostoma bispinosa: Denning 1962:39, f. 7A-C (male); Flint 1967c:175.

Lepidostoma alexanderi Denning 1962:37, f. 4A-C (male, female); Flint 1967c:175

(as syn. of L. bispinosa).

Some variants of male genitalia of L. (N.) mexicanum are virtually identical

to those of L. (N.) frontale. However, these species are easily distinguished

by having unique secondary sexual characters expressed in the male antennal

scape and forewing.

Male. Head (f. 35) with scape cylindrical, 0.9 mm long. Maxillary palps

triangular, 1.0 mm _ long. Wings with venation similar to that of L. (N.)

lacinatum. Anterior wing brown, 7.5 mm long, without anterior marginal fold.

Genitalia (f. 86) often very similar to those of L. (N.) frontale.

Female. Genitalia (f. 116) with spermathecal sclerite in ventral view oval

and elongated longitudinally with keyhole-shaped process 2/3 as long as sclerite.

The genitalia of the female holotype are illustrated by Flint (1967a).

Immatures, case and habitat. Larval head brown with weak carina around

posterior dorsal perimeter. Case cylindrical, slightly curved and composed of

small stones, similar to that of L. (N.) acarolum. Several larvae and pupae col-

lected in June.

Life history. Recorded flight dates: 7 February - November.

Distribution. Widespread throughout Central America and southwestern

United States including Arizona, Colorado, Costa Rica, Guatemala, Mexico, and

Panama.

Type data. Ofemira mexicana Banks (1901): "One specimen from Tacubaya,

November."

Olemira mexicanam Banks, New Lectotype, female, Tacubaya, Mex., MCZC

Type 11824. See material examined below.

Atomyiodes bispinosa Ulmer (1911): "1 male, Costa Rica, San José, Stieler

leg., im Berliner Museum." One possible syntype at the INHS (J. D. Unzicker).

Lepidostoma alexanderi Denning (1962): "Holotype male, Southwestern

Research Station, Chiricahua Mountains, Cochise County, Arizona, August 7-12,

1957, C. P. Alexander. Allotype. - female, same data as for holotype. Paratypes

(4 males, 19 females) same data as the holotype." See type material examined

below.

Material examined. Lectotype female (MCZC) of O. mexicanam. ARIZONA:

Cochise Co: Southwestern Research Station, 5 mi SW of Portal, Cave Cr, 3 males,

3 females, 23 V 1967, PJ Spangler, (USNM); 3 males, 2 females, 11 VII 1963, CPA,

(USNM); male (paratype of alexanderi), 7-12 VIII 1957, CPA, (USNM); 11 males,

6 females, 12 VIII 1974; 9 males, 1-8 IX 1974, H & MT, (CUCC). COLORADO:

52 Contrib. Amer. Ent. inst. Volume 24, Number 2, 1988

Jacksom GCos« Gould: 4-Vill 1974. Samales, 2 females, H.& M1). (CUCC)«. COSTA

RICA: Cartago, 2 male pupae, 24 VI 1967, OSF & Ortiz, (USNM); Heredia, Los

Cartagos, male, 24 VI 1967, PJ Spangler, (USNM); Vara Blanca, 4 larvae, male

pupa, 1.VIi 1967, OSPF & Ortiz, GUSNM).> MEXICO: , DISTRITO FEDERAL: male,

30... V1. 1938.5 AD CINHS); MORELGS: Cuernevacas: male, 23lhlhul932> AD €

Plummer, (INHS); Park Nac Zempoala, male, 7 11 1977, JB, (USNM). PANAMA:

Chiriqui, El Volcan, Chiriqui Viejo River, male, 22 VII 1964, A Broce, (USNM).

Lepidostoma (Nosopus) oaxacense Bueno & Contreras

Lepidostoma oaxacensis Bueno and Contreras 1986:208-209, f. 1-3 (male).

The following is based on the original species description. This species is

apparently closely related to L (N.) quila by having similar apex of male inferior

appendage in dorsal view, but it differs by having segment X in left lateral view

C-shaped. :

Male. Maxillary palp unsegmented and cylindrical. Anterior wing 6 mm long.

Posterior wing R only four-branched (pers. comm. Bueno). Genitalia: Segment

x in left lateral view C-shaped, having ventral process acuminate and curved

dorsoposteriad, and dorsal process more blunt and linear, and bearing about three

tubercles. Phallus with slender parameres about 3/4 as long as_ phalicata.

Inferior appendage, in lateral view main article broad and rectangular, having

apex in dorsal view similar to that of L (N.) quila with broad mesal lobe and

posterior point, but without subapical extal point; basodorsal process slender and

fingerlike; horizontal middorsal process slender acuminate and subequal in length

to basodorsal process.

Female, immatures, cases and habitate. Unknown.

Life history. Holotype collected in December.

Distribution. Oaxaca. Only known from the type locality.

Type data. Bueno and Contreras (1986): "Holotipo macho. Mexico:

Oaxaca, Carretera No. 175 Portillo del Rayo a 1540 m.s.n.m., recolectado el lo

de diciembre de 1982 por A. Ibarra y M. Garcfa. Depositado en IBUNAM."”

Lepidostoma (Nosopus) quila Bueno & Padilla

CF. 87)

Lepidostoma quila Bueno and Padilla 1981:389-391, f. 1-2 (male).

This species is similar to L. (N.) talamancense, delongi and denningi by

having male inferior appendage with a subapical flange which is not serrated.

However, it is distinct, having inferior appendage with acuminate subapical flange

projecting from position about a third length from apex of first article.

Male. Forewing 11 mm long. Hindwing with AR5, Fl and F2 present.

Genitalia (f. 87) with segment X similar to that of L. (N.) aztecum, but inferior

appendage in lateral view with first article acuminate. —

Female, immatures, case and habitat. Unknown.

Life history. Recorded flight dates: 3-18 August and 2 December.

Distribution. -Mexico State and Morelos.

Type data. Buenos and. Padilla. (1981): “Holotipo macho. Lugunas de

Zempoala, Estado de México. 3-VIII-78. Colect6: J. Bueno. Paratipos 9 machos

18-VIII-79. Colectados por J. Padilla y M. Rivera. Holotipo y paratipos estan

depositados en la Colleci6n Entomolégica del Instituto de Biologia, UNAM." See

type material examined below.

Material examined. | have examined one male paratype kindly offered as a

gift by Joaquin Bueno Soria; however, its collection data were not listed in the

original description: MORELOS: Zempoala laguna, 2 XII 1978, J. Padilla, (JSW).

Weaver: North American Lepidostomatidae 53

Lepidostoma (Nosopus) rectangulare Flint

(fF 286)

Lepidostoma rectangulare Flint 1967c:176, f. 26-27 (male); Flint and Bueno

TOTS 371 | Ses,

Male. Scape not highly modified, cylindrical, 0.6 mm long, similar to L. (N.)

mexicanum. Maxillary palp 0.6 mm long, spatulate. Wings with venation similar

to that of L. (N.) lacinatum. Anterior wing without anterior marginal fold, 9.0

mm long. Genitalia (f. 88) similar to those of L. (N.) denningi. Inferior

appendage distinctive in lateral view, main article truncated apically, serrated

subapical lateral flange with three teeth. Parameres in lateral view with apices

hooked dorsad.

Female, immatures, case and habitat. Unknown.

Life history. Recorded flight date: 25 July.

Distribution. Durango. Only known from the type locality.

Type data. Flint (1967c): "Holotype, male. Mexico (Durango), 10 miles

west of El Salto, 9000° 25 July 1964, W. C. McGuffin." See type material examined

below.

Material examined. Holotype male, (CNCI).

Lepidostoma (Nosopus) reimoseri Flint & Bueno

Ch. eo)

Lepidostoma reimoseri Flint and Bueno 1977:377, 383-384, f. 10-11 (male).

Male. Scape not highly modified, cylindrical, 1.0 mm long. Maxillary palp

short, 0.5 mm long, slender fingerlike. Wings similar to those of L. (N.)

lacinatum. Anterior wing 9.5 mm long without costal cell. Genitalia (f. 89) having

segment X with many dorsal tubercles. Inferior appendage with subapical lateral

flange inconspicuous in lateral view, but small apicolateral point in dorsal view.

Female, immatures, case and habitat. Unknown.

Life history. Recorded flight dates: 21 - 28 May.

Distribution. Costa Rica. Only known from the type locality.

Type data. Flint and Bueno (1977): "Holotype, male: Costa Rica

[Cartago], Volcan lIrazu, 2200-2500 m., 21-28 May 1930, Reimoser (Vienna

Museum). Paratype: Same data, 1 male (U.S.N.M.)." See type material examined

below.

Material examined. Paratype male, (USNM).

Lepidostoma (Nosopus) steinhauseri Flint & Bueno

(fF, 90)

Lepidostoma steinhauseri Flint and Bueno 1977:376, 384-386, f. 6-7 (male).

Male. Scape not greatly modified, cylindrical, 0.7 mm long. Maxillary palp

fingerlike, 0.5 mm_ long. Wings with venation similar to that of L. (N.)

lacinatum. Anterior wing about 11 mm long, with small basal costal cell. Genitalia

(f. 90) similar to L. (N.) heveli; but segment X in dorsal view with mesal process

broader than lateral process. Parameres at least as broad as midsection of

phalicata in lateral view.

Female, immatures, case and habitat. Unknown.

Life history. Recorded flight date: 23 January.

Distribution. El Salvador. Only known from the type locality.

54 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

Type data. Flint and Bueno (1977): "Holotype male: El Salvador, Santa

Anas: Cerro: Miramundo, ‘2300 m., 26°Uan) 1971, S. Steinhauser. USNM Type

74075." See type material examined below.

Material examined. Holotype male, (USNM).

Lepidostoma (Nosopus) talamancense Flint & Bueno

Cf. 9)

Lepidostoma talamancense Flint and Bueno 1977:376, 386, f. 8-9 (male).

Male. Scape not highly modified, cylindrical, 0.6 mm long. Maxillary palps

spatulate, 0.5 mm _ long. Wings with venation similar to that of L. (N.)

lacinatum, except posterior wing with 1A and 2A completely fused. Anterior wing

9-13 mm long, without costal cell. Genitalia (f. 91) having segment X in lateral

view with three undulate lobes, middle lobe largest. Parameres sinuate at base.

Inferior appendages with subapical lateral flange located near apex and only

conspicuous in dorsoventral aspect. |

Female. Unknown.

Immatures, case and habitat. Larval head, chestnut brown, vertex convex

and with carina extended laterally, defining perimeter, shaped like a USA World

War | Army helmet. Cases of late and early instars cylindrical and composed of

small stones. Case of final instar with anterior width 2.0 mm, posterior width

1.5-1.8 mm, and length 9.0-10.0 mm. | have examined several larvae and pupae

collected in June at the type locality, Ojo de Agua (eye of water), a very large

spring.

Life history. Several different instars of larvae and mature pupae collected

30 June. Recorded flight dates: 20 - 30 June.

Distribution. Costa Rica. Only known from the type locality.

Type data. Flint and Bueno (1977): “Holotype, male: Costa Rica, Cartago,

Ojo. ge Aguay routes? Kkmis/S, 30 June 1967). Flint and (Ortiz: “USNM: Type 7076:

Paratype: Same data, 1 male, 2 females; same, but 20 July 1967, 3 males, 2

females."

Material examined. COSTA RICA: Cartago, Ojo de Agua, Rt. 2, km 75,

larvae & pupae, 2 male pupae, male, 30 VI 1967, OSF & Ortiz, (USNM).

Lepidostoma (Nosopus) Hoodi Group

This greup comprises three “species; al. (Ni) enmanae, <hoodi, and

spicatum. The relationship of this species group to others remains elusive because

of modifications in the male anterior wing. Interestingly, similar modifications in

the male wing are exhibited by Dinarthrella. The male genitalia are similar to

those of other species of L. (Nosopus), to which this group is tentatively

assigned.

Male. Antenna, scape modified, with mesal process or bend near its base;

pedicel much longer than individual segments of flagellum. Anterior wing modi-

fied, having Cu2 apparently atrophied, thyridial cell 4/5 as long as wing, venation

obscured by medial fold (f. 54). Posterior wing with Fl only, 1A and 2A fused.

Genitalia (f. 92-94) similar to those of other species of L. (Nosopus). Segment

xX tuberculate. Phallus symmetrical. Inferior appendage with first article

acuminate, triangular; second article, small subapical mesal process; dorsal and

dorsomesal process present. :

Female. Female of L. (N.) spicatum unknown. Wings of L. (N.) hoodi

typical of the family. Spermathecal sclerite in L. (N.}) ermanae and hoodi narrow

posteriorly and wider anteriorly, bell-shaped in ventral view (f. 119).

Immatures, case and habitat. The pupal cases of L. ({N.) hoodi and spicatum

in this group are known to be similar, cylindrical and composed of small stones,

Weaver: North American Lepidostomatidae SS

suggesting that their larvae construct stone cases only. Other than L. (N.) hoodi

| have observed North American lepidostomatid larvae without abdominal gills only

from the type locality of L. (N.}) ermanae and a watershed where L. (N.) spicatum

occurs; thus | predict that these larvae are of these two species which are closely

related to L. (N.) hoodi. The presumed larvae of L. (N.) ermanae and spicatum

inhabit springs; the larvae of L. (N.) hoodi dwell in streams and springs.

Distribution. Northwestern North America.

Lepidostoma (Nosopus) ermanae Weaver n. sp.

82. Vea

This species is included in the L. (N.) hoodi group of which it is more

closely related to L. (N.) spicatum, having similar modifications of the male scape

and pedicel. However, it differs by having male, segment X more slender, almost

twice as long as its height in lateral view.

Male. Color light brown. Head vertex with median protuberance between

antennae about a fourth as long as scape. Scape and pedicel similar to those of

L. (N.) spicatum (f. 38); scape 0.4 mm long, stout, somewhat rectangular in

dorsal view with mesal surface expanded and slightly bulbous; pedicel 0.3 mm

long, with basomesal process, an extension of a mesal C-shaped cavity which bears

many small setae; first segment of flagellum about half as long as pedicel.

Maxillary palp apparently one-segmented and 0.6 mm long. Labial palp 0.95 mm

long. Anterior wing 7.0 mm long, venation similar to that of L. (N.}) hoodi with

long setose fold along M. Posterior wing with venation different from that of L.

(N.) hoodi, having M unbranched and F5 present. Genitalia (f. 92): Segment

X in lateral view almost twice as long as basal height, dorsum curved toward

triangular apex, venter more nearly straight with rounded, shallow, subapical

emargination; in dorsal view having a pair of acuminate processes with their basal

halves fused and distal halves separate, each separated by narrow V-shaped

notch, and apices slightly curved and pointed laterally. Phallus with parameres

more slender than in L. (N.) spicatum and more curved than in L. (N.}) hoodi

or spicatum.

Female. Genitalia (f. 118): Segment IX with anterior apodeme longer than

in L. (N.) hoodi. Spermathecal sclerite similar to that of L. GINS)! Rood! “upwe

having length greater than its height in lateral view.

Immatures, case and habitat. One kind of larva was collected in abundance

in a spring at the type locality and it is very probably of this species. Larva

head and pronotum brown;.pronotum in dorsal view with lateral margins curved

and slightly broader than head; abdomen without gills. Case cylindrical and made

of stones. Nancy Erman (pers. comm.) is conducting an ecological study of the

spring habitat and has observed its water temperature 3.5-4.0°C. |

Life history. A few early instar larvae (probably third of fourth instars)

were collected 9 October. Recorded flight date: 24 July.

Distribution. Known only from the type locality in the northern Sierra

Nevada Mountains of California.

Type material. Holotype: CALIFORNIA: Nevada Co: spring, northern

slope of Sagehen Creek basin, 2000 m, emergence trap, male, 24 July 1984, NA

Erman, (USNM). Paratypes: ibid, 6 males, 8 females, NA Erman: 3 males and

4 females, (USNM), 3 males and 4 females, (JSW).

Other material examined. Ibid, 9 October 1982, 7 larvae (probably third

or fourth instars), NA Erman, (JSW).

Etymology. This species is named after its collector Nancy A. Erman in

honor of her remarkable discoveries in trichopteran behavior and ecology.

56 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

Lepidostoma (Nosopus) hoodi Ross

(te: 3h ody? B2) 119)

Lepidostoma hoodi Ross 1948:152-153, pl. 14 f. 2-2A (male); Fischer 1970:57;

Anderson and Wold 1972:192, 198; Anderson 1976:119; Nimmo and Scudder

1978:119; Webb 1980:97; Anderson et al. 1982:14; Nimmo and Scudder

1983:80; Anderson et al. 1984:15.

Male. Head (f. 37) with scape 1.0 mm long, bent mesad near base, setose.

Maxillary palp fingerlike, apparently one-segmented, 1.2 mm long, setose. Wings

(f. 54) brown. Anterior wing 8.2 mm long, highly modified with deep expansible

medial fold. Genitalia (f. 93) with segment X in lateral view narrow and long,

about twice as long as high. Phallus symmetrical.

Female. Wings (f. 54) with normal venation for lepidostomatid female.

Genitalia (f. 119) with spermathecal sclerite unique, in lateral view height about

twice as long as length; in ventral view anterior portion about three times wider

than posterior. 3

Immatures, case and habitat. Larvae:and pupae found in mountain stream

and spring habitats. Case similar to that of L. (N.) cascadense, cylindrical and

made of small stones.

Life history. Larvae collected 4 May to 20 July, 14 September; pupae 20

June to 14-23 August). Recorded flight dates: 21 May - 15 September.

Distribution. British Columbia, Oregon, and Washington.

Type data. Ross (1948): "Holotype, male. - Government Camp, 4000' elev.,

on south slope of Mt. Hood, Oregon, July 23, 1946,. H. H.: Ross. < Paratype.: -

Hood River Meadows, Mt. Hood, Oregon, July 17, 1947, Kenneth M. Fender."

Holotype at the INHS (Webb 1980). Paratype also at the INHS (pers. comm. J.

D. Unzicker).

Material examined. OREGON: MEL Hood: *NoFork lron Cry Rt: 35; near

Government Camp, 8 pupae, 20 VI 1968 (2 emerged 23 VIII, preserved 26 VIII),

GBW. T¥> SDS, (ROME); Benton Co: (6</75ant 'N of Corvallis, Oak Cr, 2. males;

Zé Villa 1X.1968,°CD Kent, (USNM):cabid) trap at right branch of:Oak Cr E

side of Patterson Rd, 1.5 mi N of Fisheries Lab, 700’, male, 4-8 |X 1970, C Kerst,

(OSUO); Marys Peak, Parker. Cr)» at Summit Cpg,/ 2. larvae, 14°1K 19819: larva,

SO: V 1862,. RWW. (OSU0)) Zahn: Cr: ‘headwaters: et: hS road, of sleat: larva, 20

Vib la8i, RWW, (OSU0); Linceln Cont FlynncGr, weir*strspringhead «in seep;

NM darvae,: 22 VstI982, RWW, (OSUO):: ibid, gate trib, 3 larvae, 6 Vil 1982) RWW,

(OSU0O); ibid, gauge trib, larva, 6VIl 1982: Jjarva;:4:V 1982° male,’ 21222 VY 1982,

RWW; ( OSUO);) Linn’ Cor’ 1 mi W of Marion’ Forks,;.N Santiam Hwy, 3 males; 9

Vill 1952, K Fender, (INHS); Tombstone, male pupa, 14 VIII 1982, RWW, (OSUO);

wnion Gere” in oN ot: Biggin Gordon Cr, -aAvdervaeiol2 Vi°o1974. (NHA, ~COSUO):

WASHINGTON: Chelan Co: 11 mi W of Wenatchee Lake, Minotaur Cr, males &

females, 1-15 IX 1976, JRW (SDS).

Lepidostoma (Nosopus) spicatum Denning

(f.° 36, 94)

Lepidostoma spicata Denning 1954:190, f. 4A-C (male); Newell and Potter

1973:20:.. Roembhite:. 1962: 1%.

Lepidostoma spicatum: Fischer 1970:64.

Lepidostoma roemhildi Denning 1982:214-215, f. 13, 13A-D (male) NEW SYNONYM.

Male. Head (f. 38) having scape about 0.8 mm long, modified with slender

curved mesal process (Denning 1954: f. 4C). Maxillary palps slender and

spatulate, 1.1 mm long. Anterior wing 7.6 mm long, similar to L. (N.) hoodi.

Genitalia (f. 94) with segment X in lateral view having height about equal to

length, with several pointed tubercles.

Weaver: North American Lepidostomatidae of

Female. Unknown.

Immatures, case and habitat. A few specimens examined were collected from

a spring habitat. The case is similar to those of L. (N.) hoodi and cascadense.

Life history. Larvae collected 14 to 22 June, pupa preserved 12 July.

Recorded flight date: 21 - 29 July.

Distribution. Alberta, Idaho, Montana, and Utah.

Type data. Lepidostoma spicata Denning (1954): "Holotype, male, Twin

Creek Camp, Bitterroot Mountains, Salmon, Odaho, 5000' elevation, July 25, 1952,

Borys Malkin. Paratype, male, same data as for holotype." See type material

examined below.

Lepidostoma roemhildi Denning (1983): "Holotype, male. - Montana: Sanders

County, 10 miles NE Thompson Falls, West Fork Thompson River, 25 July 1975,

George Roemhild. Paratype, male. - Glacier County, Montana, Many Glacier Area,

small seeps along Iceberg Lake Trail, 21 July 1979, R. W. Baumann of Brigham

Young University, Provo, Utah."

Material examined. ALBERTA: Banff roadside spring on Sundance Canyon

Rd, 2 immatures, 22 VI 1962 (reared male pupa, preserved 12 VI1-29 VIII), GBW,

(ROME). IDAHO: Lemhi Co: Salmon, Bitterroot Mountains, Twin Creek Camp,

5000', male (paratype of spicatum, 25 VII 1952, B Malkin, (DGD)... MONTANA:

Gallatin Co: West Cr, 2 larvae, 14 IV 1985, DLG, (JSW); Bozeman Cr, male, 29

W1A1986, ‘DLG,+(JISW).

Lepidostoma (Nosopus), Incertae Sedis

The species group positions of the following three species of L. (Nosopus)

are uncertain.

Lepidostoma (Nosopus) cascadense (Milne)

Chie BG BS)" OSe a

Jenortha cascadensis Milne 1936:119, 121, 124 (male); Webb 1980:97.

Lepidostoma pleca Ross 1938a:1/75-176, f. 118 (male); Webb 1980:98.

Lepidostoma cascadense: Fischer 1970:40; Anderson 1976:118; Schmid 1980:150,

ff: 330: O40, . S434 slmrate} , wicoeced( sim 0206) Gratis and. Anderson

1980:808-816; LaFontaine 1981:251; Roemhild 1982:11; Baumann and

Unzicker 1981:27; Anderson et.al. °1982:14; Anderson et aly’ 1984: 15: -17,

18; Nimmo and Wickstrom 1984:4; Ruiter and Lavigne 1985:41; Herrmann

et al. 1986:434; Nimmo 1986a:2.

Lepidostoma cascadensis: Newell and Potter 1973:20; Nimmo and Scudder 1978:

119; Nimmo and Scudder 1983:80.

Lepidostoma mira Denning 1954:189-190, f. 3A-D (male, female) NEW SYNONYM;

Smith 1969:52;) Fischer 1970760) “Anderson.’'1976: 119; * Bauman: -and

Unzicker 1981:27; Anderson et al. 1982:14; Ruiter and Lavigne 1985:42;

Herrmann et al. 1986:434.

This is the only North American lepidostomatid of which both the male and

female can be recognized on the bases of wing venation alone.

Male. Scape 0.8 mm long, with mesal lobe bearing many stout setae;

maxillary palp 1.0 mm _ long, with apical segment spatulate and_ flexible;

mesoscutellum with small setose warts (f. 36). Wings (f. 55) light brown and

very narrow. Anterior wing, 8.0 mm long, venation unique, with submarginal

anal grove transecting Cul, 1A, 2A, 3A and each vein terminating in apical margin

independently; RS and Ma forming petiolate apical fork; with many modified setae

(f. 13). Genitalia (f. 95) having segment X in lateral view with dorsum short

and curved ventrad, venter longer and somewhat linear, apex pointed and bearing

58 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

many dorsal tubercles. Phallus symmetrical. Parameres with several basodorsal

tubercles. .

Female. Anterior wing venation unique, with Fl, F2, F3, and F5; R5 and

M1 forming petiolate fork (similar configuration as in male). Genitalia (f. 117)

in lateral view similar to those of L. (N.) roafi. Segment |X with short posterior

apodeme and curved ventroapical notch. Spermathecal sclerite in lateral view with

height greater than length, in ventral view oval, with length greater that width.

Immatures, case and habitat. Larva unique, metanotal sal and sa?2 sclerites

large, oval, each bearing 15-20 setae. Contributions to the life history of L. (N.)

cascadense have been documented by Anderson (1976), Grafius and Anderson

(1980). Cases of all larval instars cylindrical and composed of small stones.

Larvae inhabit mountain streams and rivers.

Life history. Larvae have been collected from September to the end of July,

pupae from mid-June to the end of July. Recorded flight dates: 14 May - 10

October.

Distribution. Alberta, Alaska, British Columbia, California, Colorado,

Idaho, Montana, Nevada, Oregon, Utah, Washington, Wyoming, and Yukon.

Type data. Jenortha cascadensis Milne (1936): "Holotype, male, 1 male

parotyoe Cultus bo, Ba Gon.8-Vie1934,. Wo EooRicker, <1 male same data. but

28-V-1935, all in the author's collection.” Holotype at the INHS (Webb 1980).

See type material examined below.

Lepidostoma pleca Ross (1938a): "Holotype, male. - Ingles Creek (near

Bluett Pass), Washington: July.10, 1936, H. Hs Ross.”...Holotype at the INHS

(Webb 1980). See type material examined below.

Lepidostoma mira Denning (1954): "Holotype, male, Strawberry, Tuolumne

County, California, July 8, 1951, E. L. Silver. Allotype. - female, same data

as for holotype. Paratype, males, 1 male, same data as for holotype; 1 male,

Castle Lake, Siskiyou County, California, Harry Chandler. Holotype deposited

in the collection of the University of California, Berkeley, California." See type

material examined below.

Material examined. Holotype male, paratypes male and female of J.

cascadensis, (INHS); paratypes 2 males of L. mira, (DGD); holotype male of L.

pleca, (INHS). ALBERTA: Bread Cr, Forestry Trunk Road, N of Coleman, male,

2 females, 17 VII 1976, APN, (UASM); Cameron Cr, WNP, 5400', male, 26 VII 1975,

DB: Donaid; CUASM). Cameron. Ce afalls,. Watertona: mate, «2oWht 1968:.. APN,

(UASMIJs > Crowsnest) RK, Erank, imate, 42: Vill 196%, APN: (WASM) Ernst. Cr,

Maycroft, Gap. Rd» male, 5: 1X 19/6, APN, CUASM): Galway Bk, Hwy 6, S:. of

Pincher Cr, male, 31 VII 1976, APN, C(UASM); McLeod R, Forestry Trunk Road,

S of Hinton, male, 25 VII 1976, APN, (UASM); Shunda Cr, 2-3 mi E of Nordegg,

male, 8 VIII 1965, APN, (UASM). BRITISH COLUMBIA: Bella Coola R, Hwy 20,

W of Anahim Lake, male, 15 VIl 1976, APN, €CNCI); 44.6 mi W of Burns. Lake,

Hwy 16,:.male, 3 females, 18 VII.1979, APN, (€CNCI);: W of: Endako, Hwy 16, 2

males, female, 18 VII 1979, APN, (CNCI); 13 km E of Hope, Hwy 3, male, 13 VII

1979, APN, (CNCI); W of Keremeos, Sixteen-mile Cr, Hwy 3, 2 males, female, 5

Vill 1979, APN, -CONCI2:: Rock: Creek, Hwyi3) Rock: Cr, male,:3 VUb) 19879, APN,

CONCT): 3.3 Kin Sor Squamish, «Hwy 98, male iT, VE IOV wAPNe fGNCI):.5.7

km 2. of Squamish, hwy 9o,; 3 males, - 12 Vail 1e79; APN,. (CNCI): Stone Creek,

Hwy Gfo0Steme Cry male. 1% Vil 1979, APN, CONCI); S&S ke Woof Topley, «Hwy 16,

2 males,.5 females, 18: Vil.1979, APN;. €GNCI):> Vancover island: 74.9<km Sof

Beaver Cove, Hwy 19, 2 males, 2 females, 10 VII 1979, APN, (JSW) (CNCI); 24.1

km S of Beaver Cove, E side of Nimpkish Lake, Hwy 19, 10 VII 1979, male, APN,

(CNCI); 24 km SW of Franklin R, Bamfield Rd, male, 7 VII 1979, APN, (CNCI);

28 $km..N of Gold River, male, female, 9 VI! 1979, APN, (CNCI); 10:km W of

Honeymoon Bay, male, 7 VIII 1979, APN, (CNCI); S of Courtney, Hwy 19, Tsable

Ry imeles ceo Wil 1979, APN. CONGIe. Kelsay Bay, \2omatesy.: O2VbLb: 1979). APNy

(CNCI)w. CALIFORNIA: (Calaveras Coi-i male, female, 17.Vil. 1973) ‘(DRG ¢°SDS;

(SDS); El Dorado:.Go:.. Pyramid. Ste, chiwy 50,.0nale, 16 Wh. 1973. DRG.-6 SDS,

(SDS); Fresno Co: Dinkey Cr, male, 10 IX 1983, RD Haines, (DGD); Mono Co:

Weaver: North American Lepidostomatidae 99

Convict Cr, male, 17 VII 1963, HDK, (USNM); Yosemite Natl Pk, Mosquito Cr,

male pupa, 17 VI 1961, GBW, (ROME); Tuolumne Co: Crabtree Cpe, 700011

mink of: Pinecrest; Ball Op 12) mabespul ie Wil 1S7Sp DRG vos SDS) (SS)

COLORADO: Boulder Co: Middle Boulder Cr, Hwy 119, Barker Res, 4 males,

6 VIII 1973, RSB & WPS, (USNM); 12 mi W of Boulder Cr, male, female, 10 VIII

1973) DLP \@ vSDSy (SBS); s@hatfee Cor Buena’ Vistal(2 meles) 9: 4 1973, DLP

& SDS, (SDS); Clear Creek Co: Idaho Springs, Clear Cr, male, female, 5 VIII

1973, RWB & WPS, (USNM); Gunnison Co: 14 mi W of Crested Butte, male, 7

Vill 1972, KWS, (USNM); Larimer Co: Rocky Mts Natl Pk, Hwy 34, Fall River,

male, 24 VII 1960, ARG, (USNM); ibid, Roaring R, 3 males, 2 females, 11 VIII

1973, DLP &€ SDS, (JSW). IDAHO: 1 mi SE of Lolo Summit,” Pack oCr,’ male, “24

MID’ 1963, SDS; (ULCM); “Boundary Go: Lowe rCanyon? Or) mater v2 1965,

WEB) COICM); 0tdaho Cor: oN Riggitis, John "Day Cer males“ ovii 1964, * SDS,

(UICM); Latah Co: Moscow, Chmarine Cr, male, 14 VII 1963""SDS8)) COFEM))

MONTANA: Glacier Natl Pk, Trick Falls: Cr,: 2 males, 3°Vit 1940, JAR. & HHR,

(INHS); Gallatin Co: West Cr, 2 larvae, 14 IV 1985, DLG, (JSW); Madison Co:

S Willow Cr, 33 larvae, 6 IV 1986, DLG, (JSW). NEVADA: Washoe Co: Galena

Cr, Whochoe Co Recreation Area, Hwy 27, 2 males, 31 VII 1968, RWB, (USNM).

OREGON: Mt. Hood, Still Cr, Forest Camp, male pupa, 17 VII 1963 (preserved

6-MI1t), GBW, (ROME); Baker Co? Spring Cr, male,» female.’ 7 Vi 1963; male, 2

females, 16 VIII 1969; male, female, 6 IX 1968, JHB, (USNM); Big Lookout Mt.,

13 VII 1974, JHB, (USNM); Lane Co: Grasshopper Cr, larva, 23 V 1982, RWW,

(OSU0O)} ‘Lincoln, Cot Flynn Cre)? farwad; 29 1 1982; 4 larvae, 7 III 1982; ibid,

immatures on cobble below bedrock collected 5 V 1982 female emerged 14 V), RWW,

(OSUO). UTAH: Summit Co: Smith & Morehouse Cr, S fork Grand Station,

male, 19 VII 1961, ARG, (USNM); Utah Co: Timpanagos St Pk, male, 7 females,

14 Vil 1972, CED, (CUCC). WASHINGTON: Chelan Co: Little Wenatchee River,

29 mi W of Leavenworth, 2 males, 24 VIII 1980, SDS, (SDS); Clallam Co: Elwah

River, Olympic Natl Pk, male, 4 females, 9 VIII 1969, SDS, (SDS) 2" Kittitas’ ‘Co:

Kachess Cp, Gale Cr, male, 25 VIII 1970, SDS, (SDS); 2 mi N of Little Kachess

Lake, 2 males, 10 VIII 1971, SDS, (SDS); Lewis Co: 10 mi S of Randle, Camp

Cr, male, 2 VII 1980, SDS, (SDS). YUKON: 32.6 km W of Teslin Bridge, Alaska

Hwy, Seaforth Cr, male, 24 VII 1979, APN, (UASM).

Lepidostoma (Nosopus) verodum Ross

CF BO BS; OB, 120)

Lepidostoma veroda Ross 1948:151-152, pl. 14 f. 1-1A (male); Fischer 1970:69:

Anderson 1976:124; Webb 1980:98; Anderson et al. 1982:14: Anderson et

al P964:15.

Male. Head (f. 39) with appendages highly modified. Scape bulbous, about

1.0 mm long. Maxillary palps, two-segmented, base bulbous and bearing long tuft

of setae dorsad, apex broad and fingerlike, 1.5 mm long. Wings (f. 56) brown.

Anterior wing 7 mm long, venation similar to L. (N.) unicofor, but with small anal

setose groove. Posterior wing similar to that of L. (N.) roafi, but fork of M

longer, about 2/3 length of wing. Genitalia (f. 96) similar to those of L. (N.)

roafi. Segment X in lateral view, having dorsum with oblique posterior angle and

ventral apex truncated. Parameres about half as long as phalicata. Inferior

appendage short, acuminate, about as long as segment X, with dorsal process

shorter than mesal process.

Female. Genitalia (f. 120) similar to those of several other species of L.

(Nosopus). Spermathecal sclerite oval in ventral view.

Immatures, case and habitat. The adult-larval association of this species is

not certain. However, based on the process of elimination of larvae known from

other Lepidostoma species groups in the West and specimens provided by R. W.

Wisseman, the larvae is tentatively identified as the one which makes a spiral case

60 Contrib. Amer. -Ent.insts, Velame 24; Number 2, 1988

as depicted by Wiggins (1977: f. 8.1 G). A few such larvae were found inhabiting

small mountain streams on wood debris.

Life history. Probable larvae collected 4 April to 10 June. Recorded flight

dates: 1/7 July - 13 September.

Distribution. Northern California, Oregon, and Washington.

Type data. Ross (1948): "Holotype, male; Hood River Meadows, Mt. Hood,

Oregon, July 17, 1947, Kenneth M. Fender." Holotype at the INHS (Webb 1980).

Material examined. CALIFORNIA: Nevada Co: Sagehen Cr at spring trib-

utary, 6400', 2 males, female, 20 VIII 1965, A Sheldon, (JSW); 3 males, 8 females,

111i Vill 1965. -(JSW)> .11 -males,.-7 -fémales, 13. 1X 1965: A Sheldon, (USNM).

OREGON: Benton Co: Marys Peak, in drainage area of road cut near Parker

Creek Falls’ near. summit, .3-larvae?, 4 1V 1981. RWW, (OSUO); Jefferson Co:

stream going into Metolius branch of Billy Chinook reserv, on wood, larva?, 10

Vi 1978, <1: Dudley, -(QSUO);. Lane Co:.. Hd: Andrews: Exp Forest; Mack Cr, 2

mates; 20 Vil 1982: male 28 VII. 1982,. RWW; COSUO).

Lepidostoma (Nosopus) deceptivum (Banks), nomen dubium

Cfesl2i]

Thremma deceptiva Banks 1907:125, pl. 8 f. 1, (female).

Lepidostoma deceptivum: Fischer 19/0:41.

The male of this species remains unassociated with its female. Additional

species associations of male and female lepidostomatids from the Southwest are

necessary to reveal the unknown male of L. (N.}) deceptivum. | have examined

the female holotype, clearing and illustrating its genitalia for the first time.

Female. Genitalia (f. 121) somewhat similar to those of L. (N.) bakeri and

knulli, but connective tube between spermatheca and spermathecal sclerite more

narrow and not as bulbous as in these species. However, this may or may not

be a reliable distinction.

Male, immatures, case, habitat, and life history. Unknown.

Distribution. New Mexico. Only known from the type locality.

Type data. Banks (1907): "One female from Beulah, New Mexico, 8,000 feet

(Cockerell) ."

Thremma deceptiva Banks, New Lectotype, male; Beulah, New Mexico, 8,000

feet, MCZC “Type 11706."

Material examined. Lectotype female, no date on label, (MCZC).

Lepidostoma Subgenus Mormomyia Banks

Mormomyia Banks 1907:127, type species: Mormonia vernalis Banks (monobasic).

Phanopsyche Banks 1911:357, type species: grisea Banks (monobasic).

Etymology: Greek, mormo, bugbear, hobgoblin; my/a, fly.

This subgenus comprises 15 Known species, including species formerly of the

L. vernalis group sensu Ross (1946). Adults of both sexes are distinguished

by having abdominal segment IX with a dorsal pair of setose warts (f. 125-148).

Flint and Wiggins (1961) and Wallace and Sherberger (1972) have made contrib-

utions to the systematics of this group.

Male. Head (f. 122, 124) with scape 0.5-0.6 mm long, shorter than head,

not highly modified, cylindrical. Maxillary palp 0.5-0.8 mm long, segmentation

reduced, apparently with only one segment, fingerlike, slender and somewhat

linear, except in L. (M.) griseum group angled mesad (f. 122). Mesonotum of

L. (M.) griseum bears several stout setae (f. 11). Wing venation different from

that of female (f. 125, 126). Anterior wing unique, with thyridial cell petiolate,

Weaver: North American Lepidostomatidae 61

being very narrow basad and broad distad, with F5 and each of its adjacent open

cells subequal in length; both anal cells usually present, except in modified

anterior wing of L. (M.) griseum group with long setose submarginal anal groove.

Posterior wing with Fl and sometimes F2 present, M and Cul unbranched, 1A and

2A completely fused. Hindwing of L. (M.) griseum group with submarginal anal

groove aligned with posterior margin of anterior wing in lateral position. Genitalia

(f. 127-141) having segment IX with pair of dorsal setose warts. Terga of other

posterior abdominal segments bearing many long setae. Unique massive egg-

shaped dorsum of abdominal segment IX in L. (M.) sackeni, possibly enlarged

modification of dorsal setose warts. Segment X dorsum large, somewhat circular,

lobelike process only in L. (M.) griseum group; in other groups dorsum absent

and with only ventrolateral processes (usually bifid). Anus opens from large

membranous lobe below abdominal segment X. Phallus asymmetrical (except in

L. [M.] vernale group), endothecal membrane and phallocrypt large and bulbous.

Inferior appendage similar to that of L. (Nosopus) species, having vertical

basodorsal and horizontal middorsal processes; second article in subapical mesal

position, but first article acuminate and triangular in lateral view,

Female. Wings typical (f. 125, 126). Genitalia (f. 142-150) having segment

[IX with pair of dorsal setose warts. Spermathecal sclerite sometimes pointed

posteriorly. Ventral sclerite of oviduct with posterior margin crenate or with

several apical ridges. Key for females of several species provided by Flint and

Wiggins (1961:281).

Immatures, case and habitat. Larvae of most species are endemic to springs,

but some occur in small streams. All known immatures apparently construct panel

cases (f. 4) during final instar, except species of the L. (M.) /ibum group which

predominantly construct stone cases (f. 2).

Distribution. L. (Mormomyia) occurs throughout most of eastern North

America and is especially diverse in the Appalachian Mountains.

Key to Adults of L. (Mormomyia) Species

(males)

la. Dorsum of segment IX ovoid, extended posteriorly beyond segment X (f.

137); West. Virginia to (Newfoundland te Wiseorsli W) HaolRe i 8k. RO ee.

Oe Ao... ae eer A)? ONS? Mattie Merman Mra Ce ee, une RR 8 Se koe. ea 2

2a. Maxillary palp angled (f. 122); warts of segment IX oval and aligned

transversely; (ty 120.) 4B8) shel euann eprodireds.. Yiu ohana ma a oe 3

2b. Maxillary palp straight (f. 124); warts on dorsum of segment IX round (f.

P04 ae cas 28 ey BM i, pre ee Nh aed stat Re ew, 4

3a. Mesonotal warts bearing many stout androconia (f. 122); posterior ventral

process of segment X short, not extended far beyond hind margin in lateral

view (f. 127); Georgia to Nova Scotia, Tennessee to Wisconsin ...........

£5 DE HE Sah A i LE Ltt cy ee ea gthicd griseum (Banks), p. 63.

3b. Mesonotal warts with normal setae (f. 123); posterior ventral process of

segment X long, extended far beyond hind margin in lateral view (f. 128);

Fiona 40. Miesiaaotoy ier sueiietiien cg wig ately f Calle morsei Weaver, p. 64.

4a. Segment X with dorsomesal hump-shaped flange extended posteriorly over

large membranous: anal lobe in. dorsal view (Ff. 130); 18V) aie... o... 5

4b. Segment X without dorsomesal flange extended over anal lobe in dorsal view

(Fx: 12S 12 4 BAe a a I i i pn si ke a 6

Dai;

20:

6a.

6b.

fa.

fb.

8a.

8b.

9a.

Sb:

10a.

106.

The

Tbe

Wa:

Tbe

13a.

Tob,

Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

Phallic parameres spiniform, subequal in length (f. 130); Louisiana, North

Carolinaend Gonmectioul esse sce. os serratum Flint & Wiggins; p. 66:

Phallic parameres broad, not spiniform, unequal in length, right paramere

aboud twice as jong as: left ones(t. (181) Tennessee oF. e ewer. eas

RPE Rh Sei Ee SuC pena ew UMO St Cle nan egt RP AME Reals Maree ete Ug etnieri Weaver, p. 6/7.

Right paramere spade-shaped in dorsal view, left paramere small and

obscure; segment X in dorsal view with two short, straight, slender, mesal

processes, without long lateral processes (f. 129); South Carolina to Maine

BAS a Ret) WER arae SNR tn Ok OS ain rata a ee igh as re ie corroll Flint? p : 65:

Right paramere in dorsal view not spade-shaped, subequal to left paramere;

segment X in dorsal view with curved lateral processes or with long,

cunved: (mesalewprocessesn(t.:. lG22I4hhisie wid a es Be ee eee e

Segment IX with large setose warts, covering most of dorsum (f. 135, 136,

Tao saweriele nercup: Giana Ws se Ouest si dd ea IN Pee ae bie, ia yy 8

Segment IX with small setose warts not covering most of dorsum (f. 132-134,

Peel eo U ek eee CEN ca tee AE ee Ras Page tie oh Pe a 10

Segment X in lateral view with short dorsolateral process and longer curved

ventrolateral rod (f. 138); Virginia to Newfoundland to Wisconsin ........

ADORE S Gh aN aiaiiere SU TORN e a re RE ea ee cae Uk vernale (Banks), p. 72.

Segment X in lateral view with long dorsolateral process and shorter

Venirolaterns) process fi Tani: hung e A Ns Rl hs L 9g

Basodorsal process of inferior appendage long, extending posteriorly as far

as apex of middorsal process (f. 135); mountains of North and South

Caroline. Giese. Io eel: ... flinti Wallace & Sherberger, p. 70.

Basodorsal process of inferior appendage short extending posteriorly as far

as bese tot middorsal.orocess (ies ioG)), Geergian wwii yew. i Sew. os

ee moretat NRE SES Gea rye en eR glenni Wallace & Sherberger, p. 71.

Inferior appendages in dorsal aspect with small pair of mesal hooks near

V-shaped notch of posterior margin, in lateral aspect without subapical

vertical process (Fs MO )s shite eih earoUu bias. Gye. CR 1]

Inferior appendages in dorsal aspect without small pair of mesal hooks near

notch of posterior margin, in lateral aspect with subapical vertical process

Oe ere OR i atite Pa? 13

Dorsal aspect of inferior appendages with broad squarely truncated apices,

phallus symmetrical (f. 140); mountains of North Carolina and Tennessee

PEA CME ce TN eMC Coes rset ate MEN aN ease ere ~mitchelli Flint & Wiggins, p. 74.

Dorsal aspect of inferior appendages with oblique apices; phallus asymmet-

Tr Ag SOR REGS SSR [Sb SARRIO REONSEUEN ARS UME ge ter POR SMM I RTS ICy Gar Sgt] ONE 0 Dana SONS a Cae et ws

Main article of inferior appendage broad, obscuring most of middorsal

process in lateral view (f. 141); Appalachian Mountains of North and South

CAROUSEL. Siw e tenner Li. eae ai styliferum Flint & Wiggins, p. 74.

Main article of inferior appendage slender, not obscuring middorsal process |

in lateral view (fi Gabo AppalachianvMountauys. of «Georgia: i.e. cs)

BA RN Ue ul ag A TANG shaggy tl HOG wu Wobetum: Wallace: é Sherberger,! po. (73:

Lateral aspect of inferior appendage with large subapical vertical process

io (eo mountains: 6h Nort Canrotina: amd Tah neseeawe eho wna evi es

ASN Ry ae ARN Sh acuta ei Rr ia tL excavatum Flint & Wiggins, BUNGE:

Lateral aspect of inferior appendage with small subapical vertical process

Gee CSCI 0,72 ESD Ce PCcU anon ee Sse cg St US OR GA MINS a ASN vent eae Sea ch rT 14

Weaver: North American Lepidostomatidae 63

14a. Inferior appendage with short curved basodorsal process, about a fourth

as long as main process (f. 134); Virginia to New Hampshire to Indiana ..

bh ala ts geile QBN aR ING a's Ce ats Mae Sal a FACE ae ote sommermanae Ross, p. 69.

14b. Inferior appendage with long curved basodorsal process, about half as long

as) rain serocess 175s 1338 )3) \ineie to Wisconsiv eee oe ee.

Lepidostoma (Mormomyia) Griseum Group

This group comprises two species, L. (M.) griseum and morse.

Male. Maxillary palps angled in front of face (f. 122). Wings (f. 125) having

anterior wing, highly specialized with posterobasal setose pocket. Posterior wing

modified with posterior submarginal setose groove. These modified sites overlap

each other when both wings are spread laterally and probably function in

pheromone dispersion. Genitalia (f. 127) having segment IX with pair of elongate

oval, dorsal setose warts. Segment X in lateral view with large circular dorsum

and two ventrolateral processes. Phallus asymmetrical, parameres diverging to

left side of phalicata. Inferior appendages with first article acuminate in lateral

view and with apex curved mesad in ventral view; basodorsal process shorter than

middorsal process; second article, subapical mesal point.

Female. Genitalia (f. 142) having segment |X with small setose wart:

spermathecal sclerite in ventral view broad posteriorly and curved anteriorly.

Immatures, case and habitat. Final instars of L. (M.) griseum construct

panel cases.

Distribution. Widespread, eastern North America.

Lepidostoma (Mormomyia) griseum (Banks)

CPG F220 D2 Pees AD)

Phahopsyche. grisea. Banks 19117°357-358 ph )'12 2° 0791922 Crate?

Lepidostoma griseum: Flint and Wiggins 1961:280, 281, 290-292, f. 9A-B, 19 (male,

female); Ellis 1962:5, 14; Blickle and Morse 1966:11; Wray’ 1967 >" p56)

McConnochie and Likens 1969:150; Fischer 1970:43: Hilsenhoff et al.

1972:22, 28; Wallace and Sherberger 1972:227; Longridge and Hilsenhoff

1973:180; Resh 1975:15; Roy and Harper 1975: 1084; Hilsenhoff 1975:25:

Seward and Swegman 1978:24; Neves 1979:171; Roy and _ Harper

1979:162; Etnier and Schuster 1979:19; Tarter and Hill 1979:206; Masteller

and Flint 1979:169, 174; Karl and Hilsenhoff 1979:33, 40-41; Masteller and

Flint 1980a:18, 19,\ 21; Masteler and Flint 1980b:202,° 207: Swegman et

al. 1981:131, 144; LaFontaine 1981:251; Unzicker et al. 1982:9.106, o¢ POT

Huryn and Foote 1983:792+ Lake. 1984+221; ‘Singh et aly. 1984:225>° Steven

and Hilsenhoff 1984:165.

Lepidostoma grisea: Herrmann et al. 1986:434.

Male. Head (f. 122) with scape 0.5 mm long, simple. Maxillary palps angled

mesad, 0.8-1.0 mm long. Mesonotum (f. 11) modified with many stout setae,

mesoscutum with setose warts larger than those of L. (M.) vernale. Tibial spurs

(f. 11). Wings (f. 125). Genitalia (f. 127) similar to 2. (M.) morsei. °° Tergum

IX with oval, transverse, setose warts. Segment X in lateral view with circular

dorsum and two short ventroapical processes. Phallus asymmetrical, right

paramere sinuate, endothecal membrane large bulbous. Inferior appendage typical

of L. (M.) griseum group.

Female. Wines Ch) nT2a): Genitalia (f. 142) similar to those of L. (M.)

sackeni; however, segment |X with smaller setose wart. Spermathecal sclerite in

lateral view with small dorsal lobe; in ventral view oval, posteriorly elongated

laterally. Ventral sclerite of vulva with several apical striations.

64 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

Immatures, case and habitat. Final instar larvae construct panel case square

in cross-section composed of hexagonal pieces of bark. Flint and Wiggins (1961)

and Karl and Hilsenhoff (1979) reported on the life history of L. (M.) griseum.

Larvae appear to be endemic to small second order streams.

Life history. Larvae collected 6 October to 28 July, pupae 14 July to 6

September. Recorded flight dates: 20 June - 27 September.

Distribution. Delaware, Georgia, Kentucky, Maine, Massachusetts, Michigan,

New Hampshire, New Jersey, New York, North Carolina, Nova Scotia, Ohio,

Ontario, Pennsylvania, Quebec, South Carolina, Tennessee, Virginia, West

Virginia, and Wisconsin. (A record from Colorado [Herrmann et al. 1986] is

probably in error pers. comm. Ds Es Ruiter).)

Type data. Banks (1911): “From Woodworth's Lake, Fulton County, New

York,..22. August: and .14...September. (Alexander .< coll.) ..” Ross .(1938b):

"Lectotype, male, Woodworth's Lake, Fulton Co., New York, August 22, No.

11693." | have seen the lectotype and one syntype at the MCZC.

Material examined. MICHIGAN: Cheboygan Co: Niggor Cr at Indian Trail,

male pupa (reared), 14 VII 1968, D Merrill, (ROME). NEW YORK: Tompkins

Co: Dryden, Ellis Hollow, male, 6 IX 1956, OSF, (USNM). NORTH CAROLINA:

Haywood Co: Great Smoky Mts Natl Pk, Chestnut Branch at Big Creek Cpg, 12

males, 9 females, 21-22 VII. 1982, JSW, (JSW); Macon -Co:. Nantahala Natl Forest,

Standing Indian Cog, male plipa,.4 larvae, 28 Vil 1979, JCM, (CUCC): 35° 5°.N

83° 14° 30" W, 2nd order stream, 2 male pupae, female pupa, RD Davie, (JSW);

Swain Co: Great Smoky Mts Natl Pk, Mingus Cr, male, 9-10 IX 1983, JSW, (JSW);

Yancey Co: Black Mts, 4760', Hemphill spring, male, 8 females, 2-3 IX 1978, JSW,

(JSW). “NOVA SCOTIA: Lunenburg Co: Petite River, New Cumberland Road,

9. males, 41% 91956, male; «female, 5 4x -4956,..DCG. Ferguson) ~(USNM).

PENNSYLVANIA: Erie Co: Behrend College, male, 20 IX 1976; male, female, 29

Vill -o ee 77, ECM, CUSNM); Six Mile Cr, 2 males, female, 26oVbb1979,-ECM,

(USNM); Forest Co: Pigeon, Marienville Rod & Gun Club, 2 males, 28-29 VII 1976,

JSW, (JSW);:Semmerset Co: laurel Hill St Pk, Jones Mill Run, male, 20-21 VII!

1976, JLS & JSW, (JSW); Westmoreland Co: Rector, Powdermill Nature Reserve,

Maul Spring, 6 males, 10 females, 10-11 1X 1975, JSW & JLS, (JSW); Laughington,

Laurel’ SummiteRd, Spruce Run, male; °21-22 VWkho1976;. male, 20-24.-Viddi1976;.. 13

males, (females. 20-26. 1: 1976,. JSW,. (WSW)s Linn Rune St PK? Linn Run; male,

29226 JO 4976, JEW, (ISW). SOUTH CAROLINA». Geonee: Cos: «£ fork Chattooga

Riven. U,Soo Fish Hasehery male, dg Wile (969. GEM. fCUCC)* Pickens Co:

Wildcat Cr, 5 mi NW of Clemson, male, 21-22 X 1979, JSW, (JSW); 4 males, 26-27

IX O68, JCM, (CUCECIs laile: Rock St Pk, ° CarrieksCr,anateg<9 1X:1969, JCM,

(CUCC), . TENNESSEE: .. Blount ‘Cor’ Great Smoky Mts Natl: Pk: cRabbit Cr, 11

males, 9 females, 22-23 VIII 1982, JSW & PUS, (JSW). VIRGINIA: Prince William

Co:...Prince William Forest Park, male, 14 1X 1973, RWB, (USNM).

Lepidostoma (Mormomyia) morsei Weaver n. sp.

(fo Se 128)

This species is closely related to L. (M.) griseum, but its male is distin-

guishable, having mesonotum not bearing stout, modified setae and segment X

with longer ventrolateral processes in lateral view. Presently, this is the only

lepidostomatid Known from Florida.

Male. Scape and maxillary palps similar to those of L. (M.) griseum.

Mesonotal warts,.(f, 123). not bearing -modified;.stewt setae as: inv os (M.)

griseum. Wings modified similarly as those of L. (M.) griseum. Forewing 7.0

mm long. Genitalia (f. 128) similar to those of L. (M.) griseum. Tergum |X with

oval setose warts. Segment X dorsum circular and smaller than that of L. (M.)

griseum, with two ventrolateral processes in lateral view more than half as long

as the dorsum and longer than those of L. (M.) griseum. Phallus asymmetrical,

Weaver: North American Lepidostomatidae 65

with broad parameres. Inferior appendage similar to that of L. (M.) griseum,

but with more slender basodorsal process.

Female, immatures, case and habitat. Unknown.

Life history. Recorded flight dates: 18 - 19 October.

Distribution. Florida to Mississippi.

hype material, - Holotype: “MISSISSIPPI:.° Stone-Cos-U:S. Forest Lands,

male, 19 October 1984, PK Lago, (USNM). Paratype: FLORIDA: Walton Co:

Portland, . Little Alaqua Cr,.. male,’ 18 Octobér -1970, PHC, (SW): (some: of the

locality information as labeled is questionable [pers. comm. K. L. Manuel]).

Etymology. This species is named after John C. Morse in honor of his

guidance, encouragement, and support in the completion of this work.

Lepidostoma (Mormomyia) Serratum Group

This group comprises three species, L. (M.) carrolli, serratum, and

etnieri.

Male. Maxillary palp slender, fingerlike, linear. Wings without secondary

modification as in L. (M.) griseum; venation similar to that of L. (M.) vernale.

Genitalia (f. 129-131) having tergum IX with round dorsal warts. Segment X in

dorsal view with short dorsomesal projection (except absent in L. [M.] carrolli)

and lateral processes not bifid. Phallus asymmetrical. Parameres separate, not

fused basally, each markedly dissimilar in size with right paramere more than twice

as large as left one in L. (M.) carrolli and etnieri.

Female. Females of L. (M.) carrolli and serratum known (f. 143).

Immatures, case and habitat. Only larvae of L. (M.) serratum known whose

final instars construct a cigar-shaped, panel case and inhabit springs.

Distribution. Louisiana to South Carolina to Tennessee to Connecticut.

Lepidostoma (Mormomyia) carrolli Flint

(fe 5429)

Lepidostoma carrolli Flint 1958:22-24, £. 4A-F (male, female); Flint and Wiggins

1961.:279;..2814,. 292-293, f. 8A-B,;-8D)15 4male.. femated« Blickle:. and: Morse

1966: 114:..Fischer -1970:40: Wallace and Sherberger 1972422732 Etnier

1973:274; Etnier and Schuster 1979:19; Masteller and Flint 1979:169; Morse

et. -al..,.1980:914. Unzieker et.ak, °198229),106)): 9.107%: .\Huryn..and “Foote

19335702.

Male. Scape 0.6 mm long. Maxillary palp 0.7 mm long. Wings similar to

those of L. (M.) vernale. Anterior wing 6.5 mm long. Genitalia (f. 129) having

tergum IX with small pair of round setose warts. Segment X in lateral view, with

dorsum extending posteriorly, venter short, irregularly shaped and _ slanted

obliquely towards dorsal apex. Parameres in dorsal view with right paramere

spade-shaped, left paramere an obscure small point. Inferior appendage in lateral

view with basodorsal process short, broad and pointed apically, and seated on

long horizontal middorsal process; first article slender and extending posteriorly

just beyond horizontal middorsal process.

Female. Genitalia (Flint and Wiggins 1961: f. 15) with ventral sclerite of

oviduct in ventral view extending beyond sternite VIII and bilobed.

Immatures, case and habitat. Unknown.

Life history. Recorded flight dates: 28 July - 16 October.

Distribution. Maine, Maryland, New Jersey, Ohio, Pennsylvania, South

Carolina, and Tennessee.

Type data. Flint (1958): "Holotype, male, Lakehurst, New Jersey, Sep-

tember 4, 1956, C. B. Knowlton, Jr. and J. G. Franclemont; allotopotype, female,

September 1, 1956; deposited in the Cornell University — collection.

66 Contrib. Amer: Ent.inst:, “Volime 24 Number 2, 1988

Paratopotypes: Aug. 25, 1 male; Aug. 26, 4 males; Aug. 30, 1 male, 1 female;

Sept. 1, 1 male; deposited in the collections of U.S. National Museum, Illinois

Natural History Survey, Dr. D. G. Denning and author." See type material

examined below.

Material examined. MARYLAND: Prince Georges Co: Beltsville, 2 males,

1°30 1% 2 1964" PHO Arnaud: GOASC): NEW JERSEY: Oégean “Co: Lakehurst,

paratype! omale,o Po 1X: 1806>> CB Knowlton “€ - JG Francléemont,(USNM).

PENNSYLVANIA: Erie Co: West Springfield, male, 5 IX 1977, ECM, (USNM);

Forest Co: Pigeon, Marienville Rod € Gun Club, male, 28-29 July 1976, JSW,

(JSW).. SOUTH CAROLINA: Aiken Co: “Savannah River Plant, Tinker’ Cr, male,

1o416 1976, “DDE SS SP, 3(CUCe):

Lepidostoma (Mormomyia) serratum Flint & Wiggins

Chi TSO e743)

Lepidostoma serratum Flint and Wiggins 1961:281, 292, f. 7A-C, 17 (male, female);

Wray 1967:56; Wallace and Sherberger 1972:227; Etnier and Schuster

bO79: 19-20 Wngicker et eb. 1962s92106; 1 Scr.

Male. Scape simple, 0.5 mm long. Maxillary palp simple linear, 0.5 mm long.

Wings similar to L. (M.) vernale. Anterior wing 8.0 mm long. Genitalia (f. 130)

having segment X in dorsal view with blunt mesal process; ventrolateral processes

about twice as long as mesal processes, slightly curved mesad and acuminate.

Parameres long, slender, asymmetrical and not fused mesially or basally (a series

of males collected at Louisiana, Schoolhouse Spring, exhibit an interesting vari-

ation, sometimes having one, both, or neither parameres bifid apically). Inferior

appendage with short, broad basodorsal process.

Female. Genitalia (f. 143) segment IX in lateral view with deep posterior

groove. Spermathecal sclerite similar to that of L. (M.) griseum, but in lateral

view without dorsal lobe.

Immatures, case and habitat. Final instar larva builds panel case, usually

cigar-shaped, but sometimes square in cross-section, consisting of hexagonal

pieces of leaf or bark. Several immatures collected at Louisiana, Schoolhouse

Spring (water temperature 19°C), indicating that this species may be endemic to

spring habitats.

Life history. Larvae collected 30 March to 24 August, pupae 30 March to 2

September. Recorded flight dates: 30 March - 20 September.

Distribution. Louisiana, Connecticut, and North Carolina.

Type data. Flint and Wiggins (1961): "Holotype, male and allotype,

female: Crabtree Meadows Camp Ground, Blue Ridge Parkway, North Carolina,

[X-16-58 (Flint and Wiggins). Paratypes: Same locality, but IX-3-59, 1 female;

[X-2-59, 1 male pupa (Flint and Wiggins)." See type material examined below.

Material examined. Holotype male and _ allotype female, (USNM).

CONNECTICUT: Windham’ Co: Putnam: mate; 1-20 1X<1965>° AB “Klots, (USNM).

LOUISIANA: > Ouachita Coz: 6 mie SW ef Calhoun,’ Jackson ‘Parish, Schoolhouse

Spring, male, 2 females, 2 male pupae, /¢ pupae, 40 larvae, 30 III 1973, HHR et

alsa CGUCe’ poniale ANMHSd es Qanetes Stemales, "23: 1451982, RWH Ss SWHS (CUCC);

male, Ohiefemaless (2O larvae; (260M 1ex3y JOM Se: CED, hetton, (CUCC) «50: larvae,

pupa. CO Vintoba, INeS: mate? larva, 9° 1999; HAR = Stannard, (CUCC); 25

larvae, 03°. Vi We WIC Se Motions | CCUCO): So prepupae: 24 Vili 19738, JCM ss

Louton, (CUCC). NORTH CAROLINA: Swain Co: Great Smoky Mts Natl Pk,

Mingus Cr, 2 males, 20 females, 9-10 IX 1983, JSW, (JSW).

Weaver: North American Lepidostomatidae 67

Lepidostoma (Mormomyia) etnieri Weaver n. sp.

fa e334)

This species is closely related to L. (M.) carrolli by having similar para-

meres, but its male is distinguishable by having a unique C-shaped segment X

in lateral view.

Male. Scape and maxillary palp similar to those of L. (M.) vernale. Wings

extraordinarily broad. Anterior wing about 8.0 mm long and 3.0 mm broad. (In

type, apices of both fore wings torn and missing.) Posterior wing 3.0 mm broad

and truncate. Genitalia (f. 131) having tergum |X with pair of small round setose

warts. Segment X with short dorsomesal process in dorsal view and with large

curved hook and smaller ventral process pointed ventrad, forming C-shaped

process in lateral view. Phallus asymmetrical. Parameres with right paramere

spade-shaped and twice as large as left one. Inferior appendage with basodorsal

and middorsal processes subequal in length, first article with one or two

apicolateral points curved mesad and with basodorsal bulbous hump in dorsal view.

Female, immature, case and habitat. Unknown.

Life history. Recorded flight date: 30 April.

Type material. Holotype: TENNESSEE: Knox Co: Knoxville, stream at

Paper Mill Rd. and Weisgarber Ave., male, 30 IV 1966, DAE, (USNM).

Etymology This species is named in honor of its collector David A. Etnier.

Lepidostoma (Mormomyia) Libum Group

This group comprises three species, L. (M.}) excavatum, libum, and

sommermanae.

Male. Maxillary palp slender, fingerlike and linear. Wing venation similar

to. -thatinet«.2.. (Mey) vernale: Genitalia (f. 132-134) having tergum IX with

medium-sized setose warts. Segment X with pair of fingerlike setose lateral

processes. Phallus with left basal side of phalicata enlarged in dorsal view.

Parameres with basal halves fused. Inferior appendage having first article with

dorsal subapical emargination in lateral view, second article apparently in dorsal

subapical position.

Female. Genitalia (f. 144-146) having spermathecal sclerite similar to that

of: Lew AND mitchell,

Immatures, case and habitat. Larvae of L. (M.) libum, excavatum and

sommermanae have been identified. Both early and late instars of these species

usually construct stone cases only (f. 2), they also appear to be endemic to small

springs. :

Distribution. Wisconsin to Quebec to North Carolina.

Lepidostoma (Mormomyia) excavatum Flint & Wiggins

ChE ABD, 4A)

Lepidostoma excavatum Flint and Wiggins 1961:281, 289-290, f. 6, 22 (male,

female); Wray 1967:56; Wallace and Sherberger 1972:227; Etnier and

Schuster, 19792193 4) Parker “rand “Weshelh. 196176) °° Ungeker if et.al.

1982:9:..306, 9.107.

Male. Scape, maxillary palp and wing venation similar to that of L. (M.)

vernale. Anterior wing about 8.0 mm long. Genitalia (f. 132) similar to those

of L. (M.) sommermanae, but inferior appendage with second article acuminate

and longer than middorsal process.

Female. Genitalia (f. 144) having spermathecal sclerite almost rectangular

in lateral view. Ventral sclerite of oviduct crenate apically, with about three small

lobes.

68 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

Immatures, case and habitat. Like other species of the L. (M.) libum group,

larva of this species are endemic to small mountain springs and construct cylin-

drical cases of small stones only (f. 2). Case of final instar with anterior width

2.0=2.2 mm, posterior width 1.5-1.0 mm, and length 9.5-10.0 mm.

Life history. Larvae collected 30 May to 15 August, pupae 8 June to 15

August. Recorded flight dates: 13 August - 13 September.

Distribution. North Carolina and Tennessee.

Type data. Flint and Wiggins (1961): "Holotype, male and allotype,

female: Indian Gap, about 5,000 ft., Great Smoky Mountains National Park,

Tennessee, IX-13-58 (Flint and Wiggins). Paratypes: Same data, 2 males, 3

females." Paratype male at the INHS (J. D. Unzicker). See type material

examined below.

Material examined. Holotype male, allotype female, (USNM). NORTH

CAROLINA: Macon Co: Wayah Crest Cpg, 10 larvae, 30 V 1981, JSW & RWH,

(JSW)> Wayah Bald, male pupa, 10 larvae, 8 VI- 196), OSF, (CUSNM):* ibid,

Rattlesnake Spring, prepupa, 2 pupae?, 26 V 1981; 12 males, 4 females, 22-23

Vil 1983, JSE, (JSW); Swain Co: Great Smoky Mts Nat! Pk, Moore Spring, 4650',

male, male pupa, 10 larvae, 14-15 VIII 1982, JSW & PJS, (JSW); ibid, spring at

buftee “Gao, 5000. 3 males; 68-9: |xX°1983. JSw, (SW): TENNESSEE: ~ Blount

Co: Great Smoky Mts Natl Pk, Gregory Bald, small spring near Sheep-pen Gap,

Cpg 13, 13 mates, 4 females, 13-14 Vii] 1982, JSW°e PIS, CSW).

Lepidostoma (Mormomyia) libum Ross

C62, “Tas; 145)

Lepidostoma liba Ross 1941:120-121, pl. 13 f. 97-97D (male, female); Flint and

Wiggins 1961:280, 281, 284-286, f. 1A-B, 8 (male, female); Weidner

1964:83; Edwards 1967:126; Fischer 1970:59; Wallace and Sherberger

19/2:226+: S6lomon and Leiser 19777166; Webb ~1980:97;. Steven and

Hilsenhoff 1984:165.

Male. Scape, maxillary palp and wing venation similar to those of L. (M.)

vernale. Genitalia (f. 133) similar to those of L. (M.) sommermanae, but inferior

appendage with longer basodorsal process, about half as long as first article.

Female. Genitalia (f. 145) with spermathecal sclerite similar to that of L.

(M.) griseum, but without dorsal lobe in lateral view.

Immatures, case and habitat. Larvae inhabit small springs. Ross (1944)

reported that larvae of this species construct both stone and panel cases. | have

observed specimens which substantiate this, but there seems to be a much lower

frequency of larvae with panel cases. In a series of 42 pupae, | observed only

2 specimens with panel cases and all others with stone cases (f. 2). Hence, the

typical larval behavioral development to convert to panel type construction

apparently is suppressed strongly and is not obligatory in L. (M.) libum. Stone

case of final instar with anterior width 1.3-1.6 mm, posterior width 2.0 mm and

length 8.2-8.5 mm.

Life history. Larvae collected year-round, pupae 16 April to 15 September.

Recorded flight dates: 12 May - 15 September.

Distribution. Hlineis and Wisconsin.” “Edwards “C1967). record’ of L. (M.)

libum from Tennessee is doubtful and is more probably L. (M.) excavatum.

Justly, Etnier and Schuster (1979) did not include L. (M.) libum in their species

list of Tennessee caddisflies.

Type data. Ross (1941): "Holotype. - male, stream in Botanical Garden,

Elgin, Illinois; June 6, 1939 (Burks & Riegel). Allotype. - female, same data as

for holotype [INHS]. Paratypes. - 1 female, same data as for holotype, but June

13, 1939 (Frison & Ross); 1 female, same data as for allotype; 3 males, 3/ females,

Quincy, Cave Springs, Illinois, September 15, 1939 (Ross & Riegel); 1 female,

spring at MeCann school, LaRue, Hliinols, May 12, 19388 (Burks €. Riegel).

Weaver: North American Lepidostomatidae 69

Paratypes are in the collections of the Illinois Natural History Survey and The

Academy of Natural Sciences of Philadelphia." Holotype at the INHS (Webb 1980).

One paratype in the Zoologisches Museums Hamburg (Weidner 1964). | have seen

one female paratype at the MCZC, "M.C.Z. Paratype 26848." See type material

examined below.

Material examined. Holotype male, allotype female, (INHS). ILLINOIS:

Aésculus Glen, Wolf Lake,-6 larvae. 30.1V 1953, L Stannard, HHR, UNHS); Elgin,

Botanical Gardens, larva, 9.1930, BHR eG Burkes: (NMS): Adams Co: Ouincy,

Cave Spring, 36 pupae, 4 male pupae, 15 |X 1939, HHR & Mohr, (INHS); La Salle

Co: .4ani:W of: Utica at Split: Bock 410 darvae, 77H) 961 HAR suGPA, CINHS).

WISCONSIN: Sauk Co: Otter Cr, 16 larvae: 2 :1V 1980; 2 male pupae, 16 IVe1980;

2-males, 2. females, .4,V1 1980,.JG: Steven, (JSW).

Lepidostoma (Mormomyia}) sommermanae Ross

Cf nod SA cl46)

Lepidostoma sommermanae Ross 1946:267, 268, 270, 286-287, f. 18A, 34A-B (male,

female); Flint .and-Wiegins-.1961: 281, 282, 288-289. 4, SA-G;;521 (male,

female); McConnochie and Likens 1969:150; Fischer 1970:64; Wallace and

Sherberger 1972:227; Roy and Harper 19/79:163; Masteller and Flint

1979:169, 174; Neves 1979:171; Webb 1980:98; Swegman et al. 1981:131,

144; Huryn and Foote 1983:792; Waltz and McCafferty. 1983b:16; Lake

1984:221.,

Male. Scape, maxillary palp and wings similar to those of L. (M.) vernale.

Genitalia (f. 134) with inferior appendage similar to that of L. (M.) /libum, but

basodorsal process shorter, stout and about 0.25 times as long as first article.

Female. Genitalia (f. 146) similar to those. of LL. {M.) mitehelli,.. out

spermathecal sclerite with height and length subequal in lateral view and ventral

sclerite of oviduct crenate apically in ventral view.

Immatures, case and _ habitat. Ross (1946) and Flint and Wiggins (1961)

reported that the larvae of L. (M.}) sommermanae construct cases of small stones

and were found in springs; my observations agree with theirs. Final instar case

anterior width 1.9-2.0 mm, posterior width 1.5 mm, and length 8.5-9.0 mm.

Life history. Larvae collected 27 March to 19 June, pupae 26 May to 28

August. Recorded flight dates: 15 May - 28 October.

Distribution. Connecticut, Delaware, Indiana, Massachusetts, New

Hampshire, New York, Ohio, Pennsylvania, Quebec, and Virginia.

Type data. Ross (1946): "Holotype. - male, Mt. Carmel, Connecticut,

Woodruff's Pond, May 15, 1944, Mrs. A. H. Sommerman. Allotype. - female, Mt.

Carmel, Connecticut, September 4, 1944, Mrs. A. H. Sommerman. Paratypes. -

Connecticut, Mt. Carmel, Spruce Bank Spring, June 26, 1944, Mrs. A. H.

Sommerman, 1 male; Sept. 4, 1944, 1 female. Pennsylvania - Roxboro, June 28,

1 male; Aug. 15, 1909, Haimbach, 1 male (ANSP). Holotype at the INHS (Webb

1980). Allotype and paratypes at the INHS (pers. comm. J. C. Unzicker).

Material examined. INDIANA: Tippecanoe Co: West Lafayette, 2 males,

241%. 1978: male,..24 X. 1978; mate... female, O° X..1979; omale,' femates.28 ¢..) 1979,

(PURC);. Wayne Co: Richmond; Haves R.. male, 23 Vill e0, EPURC)~ “NEW

YORK: Tompkins Co:. Dryden, Ellis Hollow, male, female; 6G.1X 1956,: OSF,

(USNM). OHIO: Wayne Co: Wooster, male, 8 VI 1959, CA Triplehorn, (USNM).

PENNSYLVANIA: Chester Co: Atglen, Glen Run, 2 males, 1/7-18 V 1977; male,

31..V.H 19762 male,«.2230¥ biA9RT sy d8Wee (USWde Elk Ce: Ridgway, spring near

Clarion River, 2 males, 29-30 V 1977, JSW, (JSW); Erie Co: Behrend College,

5. males, 23: Vl-12 Vili 1976: mate, 18 Vili 1978s ECM, CUSNM): Pultem Co::) Crystal

Spring, Chestnut Bridge Hollow 3 male pupae, 15 larvae, 29 VI 1976; male, female,

23724 VII 1978;.. male pupa, female pupa,.w1S> larvads «20 4)? 1977:) -19malesi;« a7

females, 31 VII 1976, JSW, (JSW); Sommerset Co: Laurel Hill St Pk, female, male

10 Contrib: “Amer Ent. lastiy Volume 24; Number 2, 1988

pupa, 26 V1974,.RFS,°(USNM); Tioga Coe Ansonia, Colton’ Point, ‘Right Branch

Run, male, 30 VI 1978, JSW, (JSW); Washington Co: Amity, spring near Ten

Mile Run, 2 males, 2 females, 15 VIII 1976, B Owen, (JSW); Westmoreland Co:

Rector, Powdermill Nature Reserve, Maul Spring, male, 10-11 IX 1975, JSW & JLS,

(JSW).

Lepidostoma (Mormomyia) Vernale Group

This group comprises four species, L. (M.) flinti, glenni, sackeni, and

vernale.

Male. Maxillary palps (f. 124) slender, fingerlike, linear, held anterior of

frons and apparently one-segmented. Wings with venation like that of L. (M.)

vernale (f. 126). Genitalia (f. 135-138) having segment IX with large dorsum

and with large setose warts comprising most of dorsum (the highly specialized and

enlarged ovoid dorsum of L. [M.] sackeni may be an extreme modification of this

character). Segment X with lateral pair of slender processes sometimes bifid.

Phallus symmetrical (except for L. [M.] sackeni). Parameres with basal halves

fused. Inferior appendages with basal ventromesal margins of first articles

broadly fused at least half their lengths in ventral view.

Female. Genitalia (f. 147-149) having spermathecal sclerite of L. (M.) flinti

and vernale pointed posteriorly, but that of L. (M.) sackeni more similar to that

of L. (M.) griseum.

Immatures, case and habitat. Observation of pupal specimens of L. (M.)

flinti and vernale suggests that final larval instars of this group construct panel

cases.

Distribution. Eastern North America including Georgia to Newfoundland to

Wisconsin.

Lepidostoma (Mormomyia) flinti Wallace & Sherberger

Ch Toa TAL)

Lepidostoma flinti Wallace and Sherberger 1972:224-225, 227, f. 5-7 (male);

Unzieker at iahoii862.9: 106; ° Obey.

Male. Scape, maxillary palp and wings similar to those of L. (M.) vernale.

Anterior wing about 8.8 mm long. Genitalia (f. 135) having tergum |X large with

setose warts comprising most of dorsum. Segment X with bifid lateral processes.

Phallus symmetrical, basal half of parameres fused. Inferior appendages with

basodorsal process spiniform and approximately as long as first article.

Female. Genitalia (f. 147) similar to those of L. (M.) vernale, having

spermathecal sclerite pointed posteriorly in ventral view, but in lateral view

somewhat circular with small anterior lobe.

Immatures, case and habitat. Larvae appear to be endemic to mountain

springs and late instars build panel cases which are square in cross-section,

composed of hexagonal pieces of leaf and bark.

Life history. Larvae and pupae collected 11 May to 15 August. Recorded

flight dates: 3 April - 7 June.

Distribution. Appalachian Mountains of North Carolina and South Carolina.

Type data. Wallace & Sherberger (1972): "Holotype, male, Highlands, Macon

Cow NortoGanclina ivApMh ISO) By Oc Se Flint, Jeo. Cavs. National Museum,

type no. 71978). Paratypes: - 1 male, Smokemont Camp Ground, Great Smoky

Mi Netior hii North Carolina wt hays O70 by KOS selene CUS National

Museum); 1 male, 4.0 mi. south of summit of Mt. Pisgah, Transylvania Co., North

Carolina, 7¢ June, 1971, by J. B. Wallace, et al." Male paratype in the INHS

(pers. comm. J. D. Unzicker). See type material examined below.

Weaver: North American Lepidostomatidae Tl

Material examined. NORTH CAROLINA: Macon Co: Wayah Bald, Rattlesnake

Spring, 4 males, 3 females, 30-31 V 1981, JSW & RWH, (JSW); ibid, Wayah Crest

Cpg, male, 30-31 V 1981, JSW & RWH, (CUCC); Swain Co: Great Smoky Mts Natl

Pk, Moore Spring, male pupa, 8 larvae, JSW & PJS, (JSW); Great Smoky Mts Natl

Pk, Smokemont Cpg, male paratype, male pupa, 5 pupae, 11-14 V 1970, OSF,

(USNM). SOUTH CAROLINA: Oconee Co: Crane Cr, 5 mi NW of Tamassee, male,

26-20. V1 1930, JSW SRW (ISW):

Lepidostoma (Mormomyia) glenni Wallace & Sherberger

(64136)

Lepidostoma glenni Wallace and Sherberger 1972:225-226, 227, f. 8-9 (male);

Unzicker et al. 1982:9.107.

Male. Scape, maxillary palp and wings similar to those of L. (M.) vernale.

Genitalia (f. 136) similar to those of L. (M.) flinti, but distinguished by having

inferior appendage with basodorsal process much shorter than first article,

slightly shorter than middorsal process.

Female, immatures, case and habitat. Unknown.

Life history. Recorded flight dates: 30 May - 2 June.

Distribution. Only known from Union County, Georgia.

Type data. Wallace & Sherberger (1972): "Holotype. - male, extreme

southwestern Union Co., Georgia, 2 June 1971, taken by a small spring-fed stream

at 2000’ by F. F. Sherberger et al."' See type material examined below.

Material examined. Holotype male, (INHS). GEORGIA: Union Co: Vogel

StiPky Wolf Cr; mate <-30531-M 1966). JCM (GuUCC).

Lepidostoma (Mormomyia) sackeni (Banks)

Ch) Gaiy 148)

Mormomyia sackeni Banks 1936:267-268, f. 9, (male).

Lepidostoma~sackeni. Flint..and: Wiggins ,1961¢280,..281, 282-284,. 10A-B,, 14A-B

(male, female); Ellis 1962:5, 14; Blickle and Morse 1966:11; McConnochie

and Likens 1969:150; Fischer 1970:64; Wallace and Sherberger 1972:226;

Hilsenhoff et al. 1972:22, 28; Longridge and Hilsenhoff 1973:180; Roy and

Harper 1975:1085; Hilsenhoff 1975:25; Roy and Harper 1979:163; Tarter

and Hill 1979:206; Masteller and Flint 1979:169, 174; Neves 1979:171;

Hilsenhoff 1981:31; Swegman et al. 1981:131, 144; Marshall and Larson

1982:78; Huryn and Foote 1983:792; Steven and Hilsenhoff 1984:166.

Male. Scape, maxillary palp and wings similar to L. (M.) vernale. Anterior

wing 7-8 mm long. Genitalia (f. 137) having tergum |X large, setose, egg-shaped,

extending posteriorly beyond other genital appendages. Phallus slightly asym-

metrical, with basal half of parameres fused. Inferior appendages short, in

ventral view about twice as wide as long.

Female. Genitalia (f. 148) with spermathecal sclerite similar to that of L.

(M.) griseum, but posterior margin of sternite VIII with long bilobed mesal

process (Flint and Wiggins 1961: f. 14A).

Immatures, case and habitat. Unknown.

Life history. Recorded flight dates: 22 July - 17 September.

Distribution. Maine, Massachusetts, Michigan, Newfoundland, New

Hampshire, New York, Nova Scotia, Ohio, Ontario, Pennsylvania, Quebec, West

Virginia, and Wisconsin.

Type data. Banks (1936): "From Dubois, Pa., 5 Sept. (Melander), North

Conway, N.H., 16 Aug. (Osten Sacken) and Trenton Falls, N.Y: (Osten

Sacken). Types in Deutsches Entomologisches Institut, Berlin-Dahlem, and

2 Contrib: Amer. Entclnst.; Volume 24, Number 2, 1988

Museum of Comparative Zoology, Cambridge, Mass." Male paratype in the INHS

(pers comm. J. Do Unzicker) .

Mormomyia sackeni Banks, New Lectotype. - male, North Conway, N.H.,

Vill 16, O. Sacken, "M.C.Z. Type 22667" MCZC. Also, one male paratype at the

MEZC, Trentom Falls, 2Nod.5 1860250. Seckena MG Z. “Paratype 22667 -MC7C:

Material examined. MAINE: Aroostook Co: Oxbow, male, 22-23 VII 1961,

A Brower, (USNM). MICHIGAN: Presque Isle Co: Ocqueoc Lake, male, female,

21-31 Vil 1974, RW ©€7ER Hodges, CUSNM). NOVA SCOTIA: Lunenburg Co:

Green Bay, male, female, 5 IX 1956, DC Ferguson, (USNM). PENNSYLVANIA:

Forest Co: Pigeon, Marienville Rod & Gun Club, Hatchery Spring, 13 males, 28-29

Viteis76, JSw, (JSW): Sommerset Co: Laurel Hill St Pk, Jones Mill Run; male,

20-2 Vill 1976, JLS © JSW.. CSW)? Westmoreland Co: ~“binn Run: St Pk, Linn

Run, male, 25-26 [X 19/6, JSW, (JSW); Laughington, Laurel Summit Rd, Spruce

Run, 20-21 VIII 1976, JSW, (JSW); Rector, Powdermill Nature Reserve, Maul

Spring, mate, female, 10-11 1X 1975, JUSW'€ JLS, (JSW).

Lepidostoma (Mormomyia) vernale (Banks)

(f 0124, 126.7 188, 143)

Mormonia vernalis Banks 1897:29 (male, female).

Lepidostoma vernalis: Flint and Wiggins 1961:181-182, f. 4A-B, 20 (male, female);

Wiggins 1961:702; Wallace and Sherberger 1972:227; Etnier and Schuster

19812251 Marshall” and “larson 1982.76, oF." 152; ..Huryn and - Foote

1983:792; Steven and Hilsenhoff 1984:166.

Lepidostoma vernale: Fischer 1970:68; Roy and Harper 1975:1085; Roy and Harper

1979:163; Swegman et al. 1981:131, 144.

Male. Head (f. 124) with scape cylindrical, 0.5 mm long. Maxillary palp

apparently one-segmented, fingerlike, slender, held anteriorly. Wings (f. 126)

brown. Anterior wing 8-9 mm long. Genitalia (f. 138) having segment |X large

with large dorsal setose warts covering most of tergum. Segment X in dorsal

view, with long pair of curved spiniform processes, originating close together

mesially and diverging laterally, and with lateral pair of much shorter processes.

Phallus symmetrical. Parameres with basal halves fused. Inferior appendage with

basodorsal process bifurcate or trifurcate and shorter than middorsal process.

Female. Wings (f. 126). Genitalia (f. 149) similar to those of L. (M.) flinti

by having spermathecal sclerite pointing posteriorly, but not angled as acutely.

Immatures, case and habitat. Several pupae with panel cases similar to those

of L. (M.) flinti were collected from springs.

Life history. Pupae collected in June. Recorded flight dates: 28 March -

30 wuly.

Distribution. Connecticut, Maine, Maryland, Massachusetts, Michigan,

Newfoundland, New Hampshire, New York, Nova Scotia, Ohio, Ontario,

Pennsylvania, Quebec, Virginia, West Virginia, and Wisconsin. Previous records

from North Carolina originally reported by Banks (1914) and subsequently

referred to by Ross (1946) and Etnier and Schuster (1979) are based on specimens

confirmed to be L. (M.) flinti (Flint and Wiggins 1961, Wallace and Sherberger

1Gf2 y=

Type data. Banks (1897): "Sea Cliff, N.Y.; common near streams, March,

April and May." Ross (1938b): Lectotype, male - Sea Cliff, Long Island, N.Y.,

March 28, No. 11687. Lectoallotype, female. Same data." | have seen these two

lectotypes and three other syntypes at the MCZC. Male syntype at the INHS

(pers? comm. «J. D. Unzicker).

Material examined. CONNECTICUT: Sleeping Giant Pk, tributary of Mill

Cr, male pupa, male, 2 females, 15 V 1962, SW Hitchock, (USNM). MARYLAND:

Prince Georges Co: Oxon Hill, male, 11 V 1972, (USNM). MICHIGAN: Emmet

Weaver: North American Lepidostomatidae 43

Co: Pellston, spring at base of Maple River Dam, male pupa, 12-13 VI 1972,

preserved 30 VI, DW Barr & IM Smith, (ROME). NEW HAMPSHIRE: Coos Co:

Jefferson Notch Road, Seboisis Brook, male, 7-8 VII 1977, JSW, (JSW); Pickham's.

Notch, Cutler River, male, 8-9 VII 1977, JSW, (JSW). PENNSYLVANIA: Chester

Co: Atglen, Glen Run, 25 males, female, 24 IV 1977; male, 11 IV 1977; 4 males,

2-3 MI-49917;-J38W. CISW) ev Elk Cor: «S Branch: Island Run, 2 ai. Sof Ridgway,

male, 29-30 V 1977, JSW, (JSW); Forest Co: Pigeon, Marienville Rod & Gun Club,

male; 28-29 VII. 1976, JSW, (JSW); Fulton Co: Crystal Springs, Chestnut Bridge

Hollow, 3 males, female, 29 V 1977, JSW, (JSW); Lancaster Co: Welsh Mts, New

Holland Watershed Area, male, 23 WH 1978,> JSW.> (SW) 2 Westmoreland Ca:

Rector, Powdermill Nature Reserve, Maul Spring, 16 males, 7 females, 20-21 V

1977; male, 31 V 1975, JSW, (JSW). VIRGINIA: Shenandoah Natl Pk, White Oak

Canyon, male, 8 V 1965, OSF, (USNM); Rappahannock Co: Shenandoah Natl Pk,

Range View Cabin, 13 males, 5 females, 23-24 V 1980, OSF, (USNM). WEST

VIRGINIA: Jefferson Co: Harpers Ferry, male, 1 IV 1979, RFS, (JSW).

Lepidostoma (Mormomyia) Mitchelli Group

Ehis group < ¢omprises« three’ species; ob. -(M.) -lobatum; ° mitchelli, oand

styliferum.

Male. Scape, maxillary palp and wings similar to those of L. (M.) vernale.

Genitalia (f. 139-141) generally similar to those of L. (M.) vernale; but abdominal

segment IX smaller in length with dorsal setose warts smaller, not occupying

majority of dorsum. Segment X with lateral pair of bifid arms. Phallus symmet-

rical-in L.(M.) mitchelli, but» not in Ls (M.). lobatum ‘and styliferum. ° Parameres

similar to those of L. (M.) vernale group, with basal halves fused. Inferior

appendage also similar to that of L. (M.) vernale group, but with additional small

dorsomesal hook.

Female. Genitalia (f. 150) with spermathecal sclerite not pointed posteriorly

as in L. (M.) vernale and flinti, and more similar to that of L. (M.) libum group.

Immatures, case and habitat. Final instar larvae construct panel cases.

Distribution. Southern Appalachian Mountains of Georgia, North Carolina,

South Carolina, and Tennessee.

Lepidostoma (Mormomyia) lobatum Wallace & Sherberger

(ee 139)

Lepidostoma lobatum icles and Sherberger 19/2:222-224, 227, f. 1-3, 3A (male);

Unzicker et ‘als 196209: 106; 9.10%

Male. Scape, maxillary palp and wings similar to those of L. (M.) vernale.

Genitalia (f. 139) very similar to those of L. (M.) styliferum, having asymmetrical

phallus, but with main article of inferior appendage not obscuring middorsal

process in lateral view.

Female, immatures, case and habitat. Unknown.

Life history. Recorded flight dates: 30 May - 23 June.

Distribution. Appalachian Mountains of Georgia.

Type data. Wallace & Sherberger (1972): "Holotype. - male, Coweeta

Hydrologic Laboratory, Macon Co., North Carolina, 11 June 1970. Paratype. -

1 male, extreme southwestern Union Co., Georgia, 23 June 1969. Both specimens

collected at light beside small spring-fed streams at 2000-2500 ft. elevation by

J. B. Wallace and F. F. Sherberger." See type material examined below.

Material examined. Holotype male, (INHS). GEORGIA: Union Co: Vogel

StuPk, Wolf Crpimale, 30-31 Wi 1980;5 ICM): fCUCCy

14 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

Lepidostoma (Mormomyia) mitchelli Flint & Wiggins

Gh 40,, 3tS0)

Lepidostoma mitchelli Flint and Wiggins 1961:281, 286-288, f. 2A-B, 16 (male,

female); Wray 1967:56; Wallace and Sherberger 1972:224, 227, f. 4 (male);

Etnier and Schuster 1979:19; Parker and Voshell 1981:6; Unzicker et al.

198239. 106) 9: 107;

Male. Scape, maxillary palp and wings similar to those of L. (M.) vernale.

Anterior wing about 8.0 mm long. Genitalia (f. 140) similar to those of L. (M.)

flinti and styliferum. Segment |X with small pair of dorsal warts. Phallus almost

symmetrical, basal halves of parameres fused. Inferior appendages with small

pair of dorsomesal hooks.

Female. Genitalia (f. 150) similar to those of L. (M.) excavatum; but

spermathecal sclerite longer in lateral view, ventral sclerite of oviduct crenate

apically in ventral view.

Immatures, case and habitat. Flint and Wiggins (1961) reported that larvae

and pupae dwell in small mountain springs and final instars build panel cases like

those of L. (M.) griseum.

Life history. Larvae and pupae collected in early September. Recorded

flight dates: 20 March - 14 August.

Distribution. North Carolina and Tennessee.

Type data. Fline & Wiggins (1961): "Holotype. - male and allotype, female,

Camp Alice, 5/796 ft., Mt. Mitchell, North Carolina, IX-16-58 (Flint & Wiggins).

Paratypes. - Same locality but 1 female pupa collected |X-1-59, emerged |X-2-59;

1 male, 2 females, pupae collected IX-3-59, emerged |IX-7-59, 1 pupa preserved

(Flint & Wiggins). Stream crossing Route 128, Mt. Mitchell, North Carolina, 1

male pupa collected IX-1-59, emerged |IX-59 (Flint & Wiggins). Highlands

Biological. Station, Highlands, North Carolina, 1 female, |X-4-58 (Flint)

Wiggins)." "This species is Known from North Carolina, with two localities at

elevations of 4000 ft. or more." Male paratype at the INHS (pers. comm. J. D.

Unzicker). See type material examined below.

Material examined. Holotype male, allotype female, (USNM); paratype female,

C(INHS). NORTH CAROLINA: Yancey Co: Black Mts, 4760', Hemphill Spring,

male. t-2 Vil lOO. male, 2-2: IX 1978... JSwW. CISwW).

Lepidostoma (Mormomyia) styliferum Flint & Wiggins

Cf PAL

Lepidostoma stylifer Flint and Wiggins 1961:281, 286, f. 3A-C (male); Wray

1967:56; Wallace and Sherberger 1972:227; Etnier and Schuster 1979:20;

Unzicker et al. 1982:9.106, 9.107.

This species is very close to L. (M.) lobatum, and in fact these species may

be synonymous.

Male. Scape, maxillary palp and wings similar to those of L. (M.) vernale.

Genitalia (f. 141) very similar to those of L. (M.) lobatum by having phallus

asymmetrical and parameres short, about a third as long as phalicata, but dis-

tinguishable by having more slender inferior appendage in lateral view with

middorsal process not greatly obscured by the main article.

Female, immatures, case and habitat. Unknown.

Life history. Recorded flight dates: 2 June - 1 July.

Distribution. South Carolina and Indian Gap, Tennessee (near the North

Carolina state line).

Type data. Flint & Wiggins (1961): "Holotype. - male, Indian Gap, about

9000 ft., Great Smoky Mountains National Park, North Carolina, VII-1-58 (Flint

& J. F. Hanson). See type material examined below.

Weaver: North American Lepidostomatidae "5

Material examined. Holotype male, (USNM). SOUTH CAROLINA: Oconee

Co, spring above Wash Branch, 5 mi NW of Tamassee, male, 2-3 VI 1980, JSW &

EMM, (JSW).

Lepidostoma Subgenus Neodinarthrum Weaver n. subgen.

Type species: Olemira pluvialis Milne.

Etymology: Greek, Neos, new; dejinos, terrible; arthron, joint.

This subgenus comprises the L. (Neo.) canthum (two spp.) and the L.

(Neo.) pluviale groups (six spp.). Species of this subgenus share with members

of the Eurasian genus Dinarthrum the unfortunate characteristic of being difficult

to distinguish from each other due to considerable variation of the male genitalia,

especially in segment X.

Male. Head and its appendages generally resemble those of L. (Neo.)

plavigle lis Tale) Scape longer than head. Maxillary palps distinctly two-

segmented, length subequal to scape, each segment cylindrical, apical segment

with many stout apical setae. Wings (f. 152, 153) more narrow than in other

subgenera of Lepidostoma, having anterior wing with broad or narrow anterior

costal cell, bearing many dark scalelike setae; venation with Cul unbranched,

Cu2 and 1A interrupted by small fold adjacent to 2A just basad of arculus, 3A

about 2/3 as long as 2A and terminating in hind margin. Hindwing with Cul

unbranched, 1A and 2A apparently fused and only F1 present. Genitalia (f.

154-162) with tergum IX broad in lateral view. Segment X with dorsomesal lobe

bearing lateral processes (possibly intermediate appendages), being prominent and

horn-shaped in L. (Neo.) canthum group, but each reduced to a pair of lateral

bifarious points in L. (Neo.) pluviale group. Phallus with phallic apodeme short,

phallic shield spoon-shaped with posterior "handle" articulating with basal plate

of inferior appendages; parameres spiniform, slender, slightly shorter than

phalicata; phalicata curved near its base; phallotremal sclerite small. Inferior

appendages with first article long, narrow; second article small, acuminate, sub-

apical process directed posteriorly in L. (Neo.) pluviale group, but in L. (Neo.)

canthum group reduced; basodorsal process slender, directed dorsoposteriorly;

ventromesal ridge broad, narrow or reduced; without horizontal middorsal process

as in L. (Nosopus) and L. (Mormomyia) species.

Female. Head with scape slightly longer than in male. Wings (f. 152, 153)

typical of genus, hindwing with 1A and 2A apparently fused. Genitalia (f.

163-168) having segment VIII with chalice-shaped sclerotized pattern on sternum

(f. 167) indicating position of concave egg pouch and retained spherical eggmass.

Spermathecal sclerite usually pentagonal in ventral view, but sometimes triangular

or U-shaped.

Immatures, case and habitat. Immatures have been identified for four species

of the subgenus. Found in mountain and coastal streams. Final instar larva

constructs very smooth, slightly curved case composed of small sand grains,

posterior width less than 1.0 mm (f. 1).

Distribution. This subgenus is endemic to the Pacific Coastal and Rocky

Mountain regions, being known from Baja California and New Mexico to British

Columbia and South Dakota.

16 Contrib, Amer. Ent. Inst., Volume 24, Number 2, 1988

Key to Adults of L. (Neodinarthrum) Species

(males)

la. Segment X with long, hornlike, heavily sclerotized lateral process (f. 154,

155), -anterior. wing with nerrow costal cell (f;.153); canthum group ..:>..

1b. Segment X with lateral process reduced to bifarious hooks (f. 156-162),

anterior wing with-broad costal cell (f,. 152); » pluviale group cae. &: 3

2a. Lateral processes of segment X each with apex strongly curved dorsad in

lateral view, not concealed by dorsomesal processes of segment X in dorsal

view; apicomesal processes of segment X with wide V-shaped notch (f. 155);

CANTOR ie ee ee ee ae baxea Denning, p. 77.

2b. Lateral processes of segment X each with apex only slightly curved dorsad

in lateral view, mostly concealed by dorsomesal processes of segment X in

dorsal view; apicomesal processes of segment X with narrow V-shaped notch

(fe 2540s) Califorara ieee lo pee, Dee eacoes canthum Ross;>p: 77.

3a. Segment X with apicomesal lobe extending posteriorly beyond lateral bifarious

MeOKe nt 190. 150d A a ee ee eee Ce ve 4

3b. Segment X with apicomesal process not extended beyond lateral hooks (f.

$eS 8/189, G2 eee ee es OS ee. fe)

4a. Segment X with apicomesal process pointed; lateral hooks with broad bases;

inferior appendage with broad ventromesal ridge (f. 158); California ......

Syd aan se ROE eR RP earr Dhara Ue Talis Cet ort y jotor Ross, 0: 30.

4b. Segment X with apicomesal process curved; lateral hooks narrow basally;

inferior appendage with smaller ventromesal ridge (f. 160); Baja California

Norte to British Columbia and South Dakota ...... pluviale (Milne), p. 81.

oa. Segment X with widely separated bifarious lateral hooks, usually with some

adjacent mesa tubercles. (f..:156, 157-162). 8 Pros. ae. Ae. 6

Sb. Segment X with appressed bifarious lateral hooks, without mesal tubercles

Ct. 150) = Arizona Fontan ake Oar Oa ee I ee ormeum Ross, p. 80.

6a. Segment X with large dorsolateral hook, extended dorsally as far as top of

tergum IM (fs 1o0) Calivrormmiat ses teres | errigenum Denning, p. 79.

6b. Segment X with small dorsolateral hook, not extended dorsally as far as

terount 1 Che teOe eG. Boa oe) Ae es ee i

7a. Ventromesal ridge of inferior appendage broad and squarely angled (f. 162);

Galitornia tor Britisn Goliiibla ceo eo i wees ye rayneri Ross, p. 83.

fb. Ventromesal ridge of inferior appendage reduced, not strongly angled (f.

156). Wiha to Nanbanaec) Skat Sacco Sere eis apornum Denning; p. 79.

Lepidostoma (Neodinarthrum) Canthum Group

Closely related to the L. (N.) pluviale group, this group comprises only two

species, L. (Neo.) baxea and canthum.

Male. Head and its appendages similar to those of L. (Neo.) pluviale. Wings

brown, having anterior wing (f. 153) similar to that of L. (Neo.) pluviale group,

but with narrow costal cell and crossvein sc-r; hindwing similar to that of L.

(Neo.) pluviale. Genitalia (f. 154, 155) with segment IX short and straight and

with broad dorsum in lateral view. Segment X with dorsal process long, extending

beyond lateral processes; lateral processes prominent, each shaped like cow's horn

in lateral view. Phallus shorter than in L. (Neo.) pluviale group, but endothecal

membrane not reduced as much; parameres straight, about half as long as inferior

Weaver: North American Lepidostomatidae ai

appendages in lateral view; phalicata curved at its base. Inferior appendages

with first article long, narrow; second article and ventromesal ridge reduced (one

of which probably represented by small subapical point).

Female. Head and its appendages similar to those of L. (Neo.) pluviale.

Wings brown, having anterior wing (f. 153) with crossvein sc-r present, hindwing

like that of L. (Neo.) pluviale. Genitalia (f. 163, 164) with spermathecal sclerite

triangular or U-shaped in ventral view and with two long narrow dorsal lobes in

lateral view.

Immatures, case and habitat. The larvae of L. (Neo.) canthum are similar

to those of L. (Neo.) pluviale, constructing cases of fine sand grains (f. 1) and

inhabiting coastal streams.

Distribution. California.

_Lepidostoma (Neodinarthrum) baxea Denning

CF 195° 163)

Lepidostoma baxea Denning 1958:97-98, f. 6A-B (male); Fischer 1970:39.

Similar to L. (Neo.) canthum, but this species is distinguished in having

lateral processes of segment X with upturned apices in lateral view. Apparently

the epithet baxea is a noun in apposition derived from Greek meaning “a woven

shoe worn on the comic stage or by philosophers."

Male. Head and its appendages similar to those of L. (Neo.) pluviale. Wings

brown, anterior wing 7.3-8 mm long with venation and form like L. (Neo.)

canthum. Genitalia (f. 155) with segment X having lateral processes, each shaped

like cow's horn with its apex pointing dorsad in lateral view. Phallus and inferior

appendages similar to those of L. (Neo.) canthum group.

Female. Head like that of L. (Neo.) pluviale. Wings brown, venation similar

to that of L. (Neo.) canthum. _ Anterior wing 7.5 mm: long. Genitalia (f. 163)

with spermathecal sclerite triangular in ventral view and with two narrow dorsal

lobes in lateral view.

Immatures, case and habitat. Larva unknown. However, Denning (1958)

mentioned that immatures may inhabit Dry Creek, a small intermittent stream in

Fresne Co: California.

Life history. Recorded flight dates: 1 May - 2 June.

Distribution. California.

Type data. Denning (1958): "Holotype. - male, Dry Creek, Fresno Co.,

California, May 1, 1954, D.-1. Abell, deposited in the collections of California

Academy of Sciences." See type material examined below.

Material examined. Holotype male, (DGD). CALIFORNIA: EI Dorado Co:

El Dorado Nat! Forest, Bridal Veil Picnic Area, 11 mi. W of Kyburz, 3120 ; -meale,

female, 2 VI 1979, JK Gelhaus, (CUCC).

Lepidostoma (Neodinarthrum) canthum Ross

4 153.0 1040164)

Lepidostoma cantha Ross 1941:118-119, pl. 13 f. 94-94A (male female); Fischer

19/70:39; Roy and Harper 1975:1085 [doubtful record]; Webb 1980:97;

McElravy and Resh 1987:80, 88.

Similar to L. (Neo.) baxea, but with apices of lateral processes of segment

xX pointed posteriorly in lateral view.

Male. Head and its appendages like those of L. (Neo.) pluviale. Wings

brown, anterior wing (f. 153) approximately 7.5 mm lone. Genitalia, (¢.. S47:

Segment X with mesal process lobelike, extending beyond lateral processes; lateral

18 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

processes acuminate with apices pointed posteriorly. Phallus and_ inferior

appendages similar to those of L. (Neo.) baxea.

Female. Head and its appendages similar to those of L. (Neo.) pluviale.

Anterior wing (f. 153) approximately 7.5 mm long, crossvein sc-r_ present.

Genitalia (f. 164) having spermathecal sclerite with long narrow dorsal processes

in lateral view, U-shaped in ventral view.

Immatures, case and habitat. Larvae construct slender cases of small stones

and inhabit coastal streams.

Life history. Larvae collected 28 April. Recorded flight dates: 9 April

=e Jane:

Distribution. California.

Type data. Ross (19417: "Holotype. - male, near small, slow stream,

Hastings Natural History Reservation, Monterey Co., California, May 24, 1938 (C.

D. Michener, at light). A/lotype. - female, same data as for holotype [INHS].

Paratypes. - 16 males, 16 females, same data as for holotype. Paratypes are in

the collections of the Illinois Natural History Survey, the California Academy of

Sciences, and The Academy of Natural Sciences of Philadelphia." Holotype at the

INHS (Webb 1980).

Material examined. CALIFORNIA: Los Angeles Co: Santa Monica Mts, West

Stunt Range; Stunt Road): 2 males, «29° V.1978;*K- Dobry; “(JSW): ‘San Diego Co:

Potreros, 5 males; ¢ females, 9: 1V 1974, H°s MT, (CUCC):: Santa Clara ‘Co: Alfum

Rock Pk, 3) males) tt Vi. 1960> SDS? “CUSNM) 26: males? 1 VP 1960" "SDS, €CASC):

Sonoma Co: E of Petacoma Hill Blvd, Copeland Creek, 200', larvae collected 28

Ive T9ST Cadults| reared), ‘LES, (LES):

Lepidostoma (Neodinarthrum) Pluviale Group

This is the sister group of the L. (Neo.) canthum group, comprising six

species. Many of these species vary only in slight detail and are difficult to

distinguish, especially in view of the natural variation expressed in the male

abdominal segment X, which often has been regarded as an important diagnostic

character.

Male. Head and its appendages as in L. (Neo.) pluviale (f. 151). Wings

(f. 152) brown. Anterior wing with broad anterior costal cell, folding dorsally

over wing, crossvein sc-r absent. Genitalia (f. 156-162) having segment IX short

with posterior margin slightly curved and dorsum broader in lateral view. Seg-

ment X dorsomesal process short, usually not extending posteriorly beyond lateral

bifarious points, but sometimes extending just beyond them; lateral processes each

reduced to pair of bifarious points. Phallus longer than in L. (Neo.) canthum

group, extending anteriorly beyond anterior margin of segment |X in lateral view,

phallic shield spoon-shaped with long posterior extension articulating with basal

plate of inferior appendage; endothecal membrane reduced; parameres spiniform,

long, curved near their bases and extending just short of phalicata’s apex;

phalicata semicircular, extending dorsad from its base and then curving

posteriorly near slight constriction at its midlength; both parameres and phalicata

similar in curvature with their lengths subequal to that of inferior appendages.

Female. Wing venation as inl. (Neo.) plaviale (f. 152). Genitalia (Ff.

165-168) with spermathecal sclerite pentagonal in ventral view.

Immatures, case and habitat. Often found in mountain streams. Larvae have

been identified for only a few species of this group. They are easily recognized

by the type of case that they construct (f. 1) which is very slender, posterior

width less than 1.0 mm, slightly curved, and made of small sand grains.

Distribution. This group occurs throughout regions west of and including

the Rocky Mountains.

Weaver: North American Lepidostomatidae #9

Lepidostoma (Neodinarthrum) apornum Denning

Ch. (4G)

Lepidostoma aporna Denning 1949:117-119, f. 10-10B (male, female); Fischer

1970:38; Roemhild 1982:11; Baumann and Unzicker 1981:27.

Male. Head like that of L. (Neo.) pluviale. Wings brown, venation and form

like those of L. (Neo.) pluviale. Genitalia (f. 156) having segment X with

apicomesal processes reduced, lateral processes each reduced to pair of bifarious

points, one point directed dorsad, another point directed ventroposteriorly and

usually two minute points on posterior face in lateral view. Phallus like that of

L. (Neo.) pluviale, apex bulbous. Inferior appendage with reduced ventromesal

ridge.

Female. | have not examined any females of this species; however the female

allotype described by Denning (1949) appears to be close to L. (Neo.) rayneri.

Immatures, case and habitat. Unknown.

Life history. Recorded flight dates: 26 May - 22 July.

Distribution. Utah and Montana.

Type data. Denning (1949): "Holotype. - male, Beaver, Utah, 8000', June

26, 1942 (C. P. Alexander). Allotype. - female, same data as for holotype."

See type material examined below.

Material examined. Holotype male, (DGD). UTAH: Beaver Co: Birch Cr,

3 males, 4 females, 22 VII 1976, RWB; Tooele Co: Clover Cr, 6 males, 6 females,

16.V1.1994,; 0oARG,) (INHS): ibid. males. 18 ViclG¥15 unales3 femates, 26. VEIL 7974,

BR Oblad, (USNM); Utah Co: Payson Can, male, 14 VII 1977, RFS, (JSW); Vivian

Park, spring, male, 16 VII 1971, BR Oblad, (JSW); Washington Co: spring above

North Juniper Cpg, male, 26 V 1976.

Lepidostoma (Neodinarthrum) errigenum Denning

bbe AD Lig BBO

Lepidostoma errigena Denning 1954:188, f. 1A-B (male); Fischer 1970:42.

Similar to L. (Neo.) rayneri and L. (Neo.) apornum this species is distin-

guished by having the lateral bifarious points of segment X with dorsal point much

larger than ventral point.

Male. Head and its appendages as in L. (Neo.) pluviale. Wings brown,

venation like that of L. (Neo.) pluviale. Anterior wing approximately 6.3. mm

long. Genitalia (f. 157) having segment X with short apicomesal processes not

extending beyond lateral bifarious points in lateral view. Lateral processes each

reduced to pair of bifarious points, dorsal point very large with its apex

extending dorsad about as far as dorsal margin of segment |X, ventral point

minute. Phallus like that of L. (Neo.) pluviale. Inferior appendage ventromesal

ridge broad, not extending very far from base, with its apex slightly angled.

Female. Head and wings like those of L. (Neo.) pluviale. Anterior wing

approximately 7.1 mm long. Genitalia (f. 165) with spermathecal sclerite slightly

broader than long in ventral view, dorsal margin with deep curved emargination

in lateral view. i .

Immatures, case and habitat. Larvae make cases similar to those of L. (Neo.)

pluviale and inhabit mountain streams.

Life history. Larvae collected 4 May. Recorded flight dates: 21 June -

28 July.

Distribution. Southeastern California.

Type data. Denning (1954): "Holotype. - male, Seven Oaks, Santa Ana

River, San Bernardino Mts., San Bernardino County, California, July 8, 1950,

John Belkin. Paratypes, two males, same data as for holotype; two males, Palomar

Mt., San Diego County, July 12, 1946, 4700', C. P. Alexander. Holotype and

80 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

one paratype deposited in the collection of the Academy of Sciences, San

Francisco, California." See type material examined below.

Material examined. Holotype male, (DGD). CALIFORNIA: San Bernardino

Co: Seven Oaks, Santa Ana River, 2 males, 22 VI 1977,. RWB, (BYUC):: San

Gogronio Mt, scenic stop just downhill from Forest Falls turnoff, 4000’, larvae,

4 V 1983 (some reared to adults), LES, (LES); Mono Co: E Walker River, male,

female, 28 Vi! 1983, RF Gill, (DGD); Riverside Co: North Fork of San Jacinto

River, 6 mi below Pine Cove, 3 males, female, 21 VI 1977, RWB, (BYUC).

Lepidostoma (Neodinarthrum) lotor Ross

(for ras)

Lepidostoma lotor Ross 1946:268, 275-276, f. 5A-C (male); Fischer 1970:59; Webb

1980:97.

This species is very distinctive, having lateral processes each reduced to a

lateral pair of broad triangular bifarious points and segment X dorsomesal process

with apex triangular and extending posteriorly beyond the bifarious process in

lateral view.

Male. Head and its appendages like those of L. (Neo.) pluviale. Wings

brown, venation and form like those of L. (Neo.) pluviale. Genitalia (f. 158)

having segment X with triangular apex extending beyond two lateral triangular

points in lateral view. Phallus like that of other species of the L. (Neo.) pluviale

group. Inferior appendage ventromesal ridge very broad with sharp apical angle

extending halfway to apex.

Female, immatures, case and habitat. Unknown.

Life history. Recorded flight dates: 15 May - 1 June.

Distribution. California. Only Known from localities listed below.

Type data. Ross (1946): "Holotype. - male, Wadell Creek, California, May

15, 1937." The United States Geological Survey, Gazetteer of California does not

list: 'Wadell Creek, ”.but docs list’ “Waddell ‘Creek ™~ at«37°05'338' NN, 122°16'32"'W,

Santa Cruz Co. Holotype at the INHS (Webb 1980). See type material examined

below.

Material examined. Holotype male, (INHS). CALIFORNIA: Santa Clara

Goce Alum Roekwer, Ss males, |} Vicl960, -SDS - (ISW)> bh owish*to: thank SS. BD.

Smith for giving me these specimens.

Lepidostoma (Neodinarthrum) ormeum Ross

(Ff, 1306166)

Lepidostoma ormea Ross 1946:268 (ormae), 275, f. 6A (male); Fischer 1970:61;

: Webb 1980:98; Baumann and Unzicker 1981:27; Ruiter and Lavigne

1985:42; Herrmann et al. 1986:435.

Lepidostoma querla Denning 1949:119-120, f. 11-11A (male) NEW SYNONYM;

Fischer 19/70:63.

Lepidostoma moneka Denning 1968:67, f. 4-4A (male) NEW SYNONYM; Canton and

Ward «1981:45%. Baumann and Unzicker 1981:27; Herrmann et al.

1986435."

This species is unique, having two short lateral bifarious points with adjacent

bases (ventral point sometimes absent). | have examined types of L. ormeum,

L. monekum, and L. querlum as well as additional specimens collected at the type

locality of L. querlum which exhibit natural variation in segment X similar to

several forms described under these three available species names.

Male. Head and its appendages as in L. (Neo.) pluviale. Wings brown,

venation and form like those of L. (Neo.) pluviale; anterior wing approximately

Weaver: North American Lepidostomatidae 81

7.3 mm long. Genitalia (f. 159) having segment X with lateral processes each

reduced to pair of short appressed bifarious points, ventral point sometimes |

reduced. Phallus like that of L. (Neo.) pluviale. Inferior appendage ventromesal

ridge with blunt apex.

Female. Head like that of L. (Neo.) pluviale. Wings brown, venation like

that of L. (Neo.) pluviale. Genitalia (f. 166) having spermathecal sclerite with

length and width subequal in ventral view and having deep dorsal emargination

in lateral view.

immatures, case and habitat. Unknown.

Life history. Recorded flight dates: 11 June - 26 August.

Distribution. Arizona, Colorado, New Mexico, Utah, and Wyoming.

Type data. Lepidostoma ormea Ross (1946): ‘Holotype. - male, Heber City,

Utah, July 23, 1939, at light, Knowlton & Harmston." Holotype at the INHS (Webb

1980). See type material examined below.

Lepidostoma querla Denning (1949): "Holotype. - male, Oak Creek Canyon,

Arizona, June 1942 (C. P. Alexander)." See type material examined below.

Lepidostoma moneka Denning (1968): ‘Holotype. - male, one mile south of

Portal, Arizona, 8 August 1965, Mont A. Cazier." See type material examined

below.

Material examined. Holotype male of L. ormeum, (INHS); holotypes male of

L. querlum and L. monekum, (DGD). ARIZONA: Oak Creek Canyon, 4 males,

39 females, 18 VI 1968, OSF & Menke, (USNM); Apache Co: Little Colorado River,

Hwy 373, Greer, 2 males, 4 females, 17 VII 1969, (USNM); Gila Co: Christopher

Cros Hwy: 260,°3 males, 13: Vil 1974: WPS, > CUSNM). COLORADO: Grand'Cot

Fraser River, Granby, male, 2 females, 4 VIII 1973, RWB & WPS, (BYUC); Eagle

Co: Eagle River, Gypsum, 2 males, female, 10 VII 1973, RWB & WPS, (BYUC).

NEW MEXICO: CatromCo* Whitewater Cr, Catwalk Picnic Area, 2 males, 2

females, 11 VI 1974, WPS, (USNM). UTAH: Duchesne Co: Uinta River, canyon

summer homes, male, female, 7 Vill 1975, RWB, (BYUC).

Lepidostoma (Neodinarthrum) pluviale (Milne)

(hi vol a2; 10: 16tetep

Olemira pluvialis Milne 1936:117, 123, 124 (male).

Lepidostoma rhino Ross 1946:268, 276, f. 4A-C (male, female) NEW SYNONYM;

Fischer 1970:637: Flint and Bueno~-1977:375, 377: Nimmo and: Scudder

1978:120; Webb 1980:98; Nimmo and Scudder 1983:80.

Lepidostoma veleda Denning 1948b:20-21, pl. 4 f. 5-5C (male, female) NEW SYN-

ONYM; Fischer 1970:68; Anderson 1976:124: LaFontaine 1981:249, 325;

Roemhild 1982:11; Baumann and Unzicker 1981:27; Ruiter and Lavigne

1985:44; Herrmann et al. 1986:435-436; Ward 1986:191.

Lepidostoma pluviale: Fischer 1970:61; Anderson 1976:120; Solomon and Leiser

197%: 472; 178)" Nimme, and Scudder: 1978120) Sehmid 19602150. 4..°542

(male); LaFontaine 1981:191, 249, 250, 251, 325; Baumann and Unzicker

1981:27; Roemhild 1982:11; McElhone and Davies 1983:2303; Nimmo and

scudder: 1963: 74, °80;}: Ruiter and Lavigne: 1985742): Hermmann et: al.

1986:435; MecElhone et al.’ 1987:281, 282.

This species is distinguished from other species of the L. (Neo.) pluviale

group by having segment X with rounded apex extending beyond the small pair

of lateral bifarious points in lateral view. The types of L. rhino and L. veledum

were examined and compared with the holotype of O. pluvialis (the genitalia of

the latter were cleared and illustrated for the first time). Significant variation

was observed in segment X of males of L. (Neo.) pluviale from a single collection

series (Fo 161) Hence, considerable differences in male genitalia within a

population of L. (Neo.) pluviale must be expected. The new synonyms listed

above were based on this interspecific variation of male abdominal segment X.

82 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

Male. Head (f. 151) with scape 0.8 mm long, maxillary palps 0.8 mm long.

Wings (f. 152) with anterior wing approximately 6.5 mm long. Genitalia (f. 160,

161). Segment X rounded apex extended beyond small lateral bifarious points;

lateral processes each reduced to pair of small lateral bifarious points, one pointed

dorsad and one ventrad. Phallus like that of other species in this group. Inferior

appendage with blunt ventromesal ridge.

Female. Head typical of female Lepidostomatidae. Wings: F.0152)° with

anterior wing approximately 7.0 mm long, light brown. Genitalia (f. 167) with

spermathecal sclerite somewhat pentagonal in ventral view, broad and dorsal

margin with shallow emargination in lateral view.

Immatures, case and habitat. All larval instars have a smooth, slightly

curved case of sand grains (f. 1). Larvae were reported by Ross (1946) to

inhabit mountain streams.

Life history. Pupae collected 14 June to 31 July. Recorded flight dates:

30 May - 9 November.

Distribution. Alberta, Baja California Norte, British Columbia, California,

Colorado, Idaho, Montana, New Mexico, Oregon, South Dakota, Utah, and

Wyoming.

Type data. Olemira pluvialis Milne (1936): "Holotype. - male, 3 male par-

atypes ‘Creede (8800'), Colo., 9-VII, F. M. Carpenter,’ in the M.C.Z.; allotype

female, 1 male, 1 female paratypes, ‘Cultus L., B.C., 4-Vll-1932, W. E. Ricker,

2 males, 2 females, same data but 7-VI-1934, 23-VI-1932, 24-VI-1932, 9-XI-1933,

respectively, in the author's collection; 11 other males from B.C. and Wyo., 29-V

to 22-Vil, 5 in the Can. Nat. Collection, 6 with the author.” At the MCZC | have

seen the male holotype, "M.C.Z. Type 22275"; and 5 paratypes (3 males and 2

females), "M.C.Z. Paratypes 22275." See type material examined below.

Lepidostoma rhino Ross (1946): "Holotype. - male, R. Santo Domingo,

Rancho San Antonio, 1000' elevation, Baja California, Mexico, May 24, 1937, H.

J. Rayner. Allotype. - female, same data as holotype. Paratypes. - 4 males, 4

females, same data as holotype." Holotype at the INHS (Webb 1980). See type

material examined below.

Lepidostoma veleda Denning (1948b): "Holotype male, Albany County,

Wyoming, Snowy Range Mountains, near Centennial, Wyoming, August 3, 1947 (D.

G. Denning). Allotype. - female, same data as for holotype. Paratypes. - 9

males, 2 females, Woods Landing, Wyoming, August 10, 1947 (D. G. Denning); 3

males, Poudre River, 15 miles west of Teds Place, Colorado, August 17, 1947 (D.

G. Denning); 3 males, 4 females, Poudre River, approximately 15 miles east of

Cameron Pass, Colorado, August 19, 1947 (D. G. Denning); 10 males, Rocky

Mountain. National Park) Colorado, August, 9,;°°1947. (D-°G. Denning). incall

instances the species was collected only from clear, swift flowing, mountain

streams." Male and female paratypes in the INHS (pers. comm. J. D. Unzicker).

See type material examined below.

Material examined. Holotype male, paratypes 2 males of O. pluvialis,

(MCZC); holotype male, paratype female of L. rhino, (INHS); paratypes 2 males

of Li: veledum; (DGD). ALBERTA: Highwood River, Longview, 5 males, 4

females; 4 Vith 1968, APN, (UASM). “BRITISH COLUMBIA: Penticton, Ellis Cr,

at Hwy 97, male pupa, 4 pupae, 22 VI 1956, R Coleman, (USNM). COLORADO:

Boulder: Co:°° S$ St Vrain’ Cr, along Rt 7, W of Lyons, male pupa; 3 pupae, 31

Vil 1966, (ROME); Middle Boulder Cr, Hwy 119, Burker Res., male, 6 VIII 1973,

RWB & WPS, (USNM); Jefferson Co: Elk Cr, Hwy 285, 2 males, 2 females, 8 VIII

1973, RWB & WPS, (USNM). IDAHO: Teton River, Teton Basin, 4 males, 2

females, 11 VII 1966, R Nave, (USNM); Clearwater River, Wild Goose Cpg,

immature 7 VII 1974 (male emerged 20 VII), R Rood, (OSUO); Blaine Co: Sawtooth

Natl Forest, Warm Spring Cr, 16 males, 12 females, 30 VII 1978, RFS, (JSW);

Ketchum, male, 27 VIII 1978, RFS, (JSW); Lemhi Co: 4.5 mi E of Carmen, Carmen

Cr loemetes’ 26: females, -20: Vit 1963, SDS:o (GSW); Shoshone Co: 10: mi:E of

Avery, 24 males, 4 females, 23 VII 1966, DS Horning, (JSW); Valley Co: 1 mi

S' of Yellow. Pine; 7 males, 24 females,’ 3 Vilh: 1964)" SDS, (JSW). MONTANA:

Weaver: North American Lepidostomatidae 83

Yellowstone Natl Pk, near Gardiner, 2 males, 2 females, 16 VIII 1962, PJ Spangler,

(USNM). NEW MEXICO: Colfax Co: 2 mi SE of Eagle Nest, 6 males, 15 females,

15216 VET * Ov 4e Ele Todds (USNM} 51 ini Wo Utei-?k, 2 males. (13-16 Vit 4974, EL

Todd, (USNM); Lincoln Co: Rio Bonito, 1.5 mi W of Angus, male, 2 females, 16-17

Vi 1980; “MWS, CJSW) 6 “OREGON? Baker’ Co: Spring Cr, 2 males-12 Vi 1869:

male, female, 9 VII 1969; 2 males, 4 females, 12 VI! 1970, JHB, (USNM); Baker,

Pine Crj:4500', 4 males; 14 WH 1967) KG, €OSU0); °N -Powder River, S mr W of

North Powder, 2) males, female, 13 Vil 1964, oNHA?*(OSU0O} “Union Cor 427m

SW of La Grande, Hwy 244, Meadow Cr, male pupa, 14 VI 1974, NHA, (OSUO).

SOUTH DAKOTA: Hardy, 2 males, 7 VII 1965, RW Hodges, (USNM). UTAH:

Cache Co: Logan, 4 males, 2 females, 24 VI 1974; 7 males, 6 females, 1 VII 1974;

7 males, 21 females, 8 VII 1973, GF Knowlton, (USNM); Logan Canyon, male,

female, 4 VII 1937, Smith & Harmston, (USNM); Summit Co: Yellow Pine Cr, 7

mi E of Kamas, 4 males, female, 28 VII 1973, WPS, (USNM); Logan River, larvae,

| 1932; .male pupa, 8:Y1l 1992, 0° King, (USNM) : > Utah Cor America Pk Caw, 2

males, 5 females, 19 VI 1976, RFS, (JSW); Wasatch Co: Daniels Cr, Whiskey

Springs Picnic Area, male, 3 VIII 1973, RWB & WPS, (USNM); Lemmons Grove,

Provo River, many adults, July, light trap, (BYUC). WYOMING: Yellowstone

Natl Pk, Noris Cpg, 2 males, 22 VIII 1955, KM Fender, (USNM); Teton Co: 11

mi NE of Jackson, Gros Ventre, ¢ males, 31 females, 29 VII 1978, (USNM).

Lepidostoma (Neodinarthrum) rayneri Ross

Cf. 9 162, ° 168)

Lepidostoma .rayneri Ross °1941:117-118, pl. 13 #. .96-96A (male); Fischer

1970:63; Anderson 1976:121-122; Nimmo and Scudder 1978:120; Webb

1980:98; LaFontaine 1981:251: Roemhild 1982:11. Nimmo and Scudder

1983 :80;

Lepidostoma crypta Denning 1964:243-244, f. 4-4B (male) NEW SYNONYM.

Lepidostoma fischeri Denning 1968:67-68, f. 5-5B (male) NEW SYNONYM; Anderson

1976:119; McElravy and Resh 1987:88.

Similar to L. (Neo.) errigenum and apornum, this species differs by having

segment X with a very reduced apicoventral hook and inferior appendage with a

very broad ventromesal ridge with sharp apical angle extending to base of second

article.

Male. Head and its appendages as in L. (Neo.) pluviale. Wings brown,

venation and form like that of L. (Neo.) pluviale. Anterior wing 6.8 mm long.

Genitalia (f. 162) having segment X in lateral view with lateral processes, each

reduced to lateral pair of bifarious points, with ventral point smaller than dorsal

point, additional minute points sometimes present on posterior face between

bifarious points. Phallus like that of L. (Neo.) pluviale. Inferior appendage

with ventromesal ridge broad and with sharp apical angle extending posteriorly

to base of second article.

Female. Head like that of L. (Neo.) pluviale. Wings brown, venation like

that of L. (Neo.) pluviale. Genitalia (f. 168) similar to those of L. (Neo.)

errigenum and ormeum. Spermathecal sclerite very broad in ventral view, dorsal

margin with deep, curved emargination in lateral view.

Immatures, case and habitat. Pupal cases similar to those of L. (Neo.)

pluviale. \mmatures inhabiting mountain streams.

Life history. Pupae collected 15 June to July. Recorded flight dates: 2

June - 1 August.

Distribution. British Columbia, California, Oregon, and Washington.

| Type data. Lepidostoma rayneri Ross (1941): "Holotype. - male, Convict

Creek, Mono County, California, August 1, 1938 (H. J. Rayner). Allotype. -

female, same data as for holotype [INHS]. Paratypes. - 1 male, 1 female, same

data as for holotype; 1 male, Molalla, Molalla River, Clackmas County, Oregon,

84 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

July 1, 1935 (S. G. Jewett, Jr.). Paratypes are in the collection of Illinois

Natural History Survey and The Academy of Natural Sciences of Philadelphia.”

Holotype at the INHS (Webb 1980). See type material examined below.

Lepidostoma crypta Denning (1964): "Holotype. - male, Scott Mountain, 19

air miles north of Trinity Center, Trinity County, California, 5300', 5 Juby 1963;

C. D. MacNeill and V. B. Whitehead. The type is to be deposited in the collection

of the California Academy of Sciences, Golden Gate Park, San Francisco, Cali-

fornia." See type material examined below.

Lepidostoma fischeri Denning (1968): “Holotype. - male, Crater Lake

National Park, Oregon, 31 July 1964, Joe Schul and J. D. Vertrees, collected in

sphagnum bog." See type material examined below.

Material examined. Holotype male, allotype female of L. rayneri, (INHS);

holotypes males of L. cryptum and L. fischeri, (DGD). BRITISH COLUMBIA:

N of Vancover, Lynn Cr, male & female pupae, 18 VII 1946, HHR, (INHS).

CALIFORNIA: Yosemite Natl Pk, Tuolumne River, male pupa, pupae, 16 VI 1961,

GBW, (ROME); Stanislaus Natl Forest, Tuolumne River, pupae 18 VI 1961 (some

reared to adult), GBW, (ROME); Madera Co: Central Camp, 5500', 9 males, 13

femaies, 30 Vil 1983,.J Larson, (DGD); Mono Co:'.Conviet:Cr,::7200';- 20-21 July

1963, 5 males, 8 females, HDK, (USNM). OREGON: Ochoco Forest Cpg, near

Pineville, 2 males, 12 females, 26 VII 1963, JHB, (USNM); Crook Co: Marks Cr,

Hwy 26, 1 mi W of Ochoco Summit, pupae 15 VI 1974 (male & 3 females emerged

30.Vi)J, NHA, (Q@SU0); Lane Co: 40 mi E.of Eugene, 41 males, 14 females, 7 VI

1964, KG, (OSUO); Josephine,Co:,. 4 mi-N of*Grants Pass, adults, 2 VI 1965,

KG, (OS5U0);, Polk Ce: . Mill Cr, near Buell) male, 9 VII 1977, NHA, (OSUO).

WASHINGTON: Okanogan Co: Winthrop, 2 males, 42 females, 26 VII 1962, JFG

Clarke, (USNM).

Lepidostoma Subgenus Lepidostoma Rambur

This subgenus includes five North American species, L. (L.) carolina, lati-

penne, stigma, tibiale, and togatum, and the generotype, hirtum of Europe. The

other species of Lepidostoma from the Old World are tentatively included in this

subgenus, also.

Male. Head (f. 169) with scape usually shorter than that of female. Max-

illary palps two-segmented with segmentation obscured, apical segment often

flexible, extensible, and bearing scalelike setae. Wing venation of three types:

Chicunique for ty.Cle), stigma (f: 172); (2).similar: for) lL. (L299 fatipenné. and

carolina Af... 113); end. (2) shared by Lb. {L.) “togatum and»tibiale: (£2174).

Anterior wing with thyridial cell reduced or open, costal cell reduced to small

setose basal pocket, modification of veins converging at arculus, Cul unbranched

in L. (L.) stigma, tibiale, and togatum, small setose fold sometimes present in

anal region, anal loops absent, in L. (L.) latipenne and carolina broad with M

and posterior veins appressed together. Hindwing with Fl only, R5 fused with

R4, Cul unbranched; 1+2A forming petiolate fork in L. (L.) stigma, tibiale and

togatum, but fused completely in L. (L.) flatipenne and carolina. Genitalia (f.

175-179) with segment IX short. Segment X with lateral processes (possibly

intermediate appendages) usually horn-shaped and asymmetrical or reduced,

dorsomesal process reduced to pair of small lobes. Phallus with phallic apodeme

and phallobase short and collar-shaped, phallic shield with lateral straps artic-

ulating with basal plate of inferior appendages and lateral ventroposterior corners

of segment X, endothecal membrane short, parameres absent, phalicata curved

ventrad at base. Inferior appendages with main article long and narrow, apices

acuminate or fingerlike; second article usually in subapical mesal position, but

long and directed dorsoposteriorly in L. (L.) latipenne and carolina; vertical

basodorsal process capitate; basomesal process absent in L. (L.) tibiale.

Female. Head (f. 170) with scape longer than that of male. Wings: (fs 473,

174) typical of Lepidostomatidae. Genitalia (f. 180-184) having spermatheca with

Weaver: North American Lepidostomatidae 85

microtrichia forming conspicuous reticulate patterns, often being unique for spe-

cies. Spermathecal sclerite usually broader than long in ventral view, sometimes :

with lateral processes in ventral view.

Immatures, case and habitat. Cases of immatures are typical panel type,

usually tapered posteriorly and square in cross-section. In North America most

lepidostomatids are endemic to mountain streams and springs, and certainly are

most diversified in mountainous regions. However, North American species of this

subgenus seem to be eurythermic, inhabiting environments such as third to fifth

order streams or the littoral regions of lakes. This is suggested by the fact that

in the southeastern Coastal Plains region (where few other lepidostomatids occur)

L. (L.) carolina and latipenne are very common.

Distribution. Widespread throughout the Nearctic region, exclusive of the

Southwest; Alabama and Georgia to Newfoundland to Oregon to Yukon.

Key to Adults of Nearctic Lepidostoma (L.) Species

(males and females)

la. Male inferior appendage without basomesal process (f. 178); female sper-

matheca lightly reticulated anteriorly (f, 183); Georgia to Virginia |. >. 44"

ee ee a a re oe tr eye tibiale (Carpenter), p. 89.

1b. Male inferior appendage with basomesal process (f. 175-177, 179); female

spermatheca heavily reticulated anteriorly (f. 1O0 12). VOAO ar ,.. Z

2a. Male scape with bizarre mesal process, bearing large saber-shaped seta and

pair of capitate setae (f. 171); female spermatheca with anterior half strongly

reticulated and constricted, fingerlike (f. 182); British Columbia to Montana

tO Oregon to Colorado (ee ee es stigma Banks, p. 88.

2b. Male scape without mesal process or with small mesa! lobe (f. 170); female

spermatheca oval, approximately a third or less of anterior portion reticulated

ROO VBL ABE ie Ae Yan Te ai ian Mie OS 3

3a. Male lateral process of segment X horn-shaped (f. 179); female less than a

third of anterior portion of spermatheca reticulate (4.184): “Alabama and

Georgia to Newfoundland’ to Northwest Territories .2...0...0-)......00.

OMT nae eae ah gg NORE ges as Ogee eV ae enlace togatum (Hagen), p. 90.

3b. Male lateral horn-shaped processes of segment X absent or reduced (f. 175,

176); female with approximately a third of anterior portion of spermatheca

reticulate (f. 180, 181) Sian eV Nk ea ea Cay 4 A eet rc Oe od Lee a 4

4a. Large anterior wing, length greater than 7 mm; male inferior appendage with

slender middorsal process extending obliquely above main process in lateral

view (f. 176); female spermathecal sclerite wider than long in ventral view

(fs 181); Georgia t6°Nova Stolle (4. edn eo. latipenne (Banks), p. 86.

4b. Small anterior wing, length less than 7 mm; male inferior appendage with

broad middorsal process extended horizontally beyond main process in lateral

view (f. 175); female spermathecal sclerite about as wide as long in ventral

view (f. 180); North and South Carclina’ ....... carolina (Banks), p. 85.

Lepidostoma (Lepidostoma) carolina (Banks)

CEO FES A 80)

Notiopsyche carolina Banks 1911:356-357, pl. 13 f. 28 (male).

Lepidostoma carolina: Wray 1967:56; Fischer 1970:39-40; Morse et al. 1980:91;

Unzicker et al. 1982:9.106, 9.107.

86 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

This species is closely related to L. (L.) /latipenne, but it is much smaller

in size with anterior wing approximately 6 mm long. The male differs by having

the inferior appendage second article with an oblique broad apex, extending

posteriorly parallel to, and beyond the main article. The female spermatheca is

also similar to that of L. (L.) latipenne, but its spermathecal sclerite is narrower

in width about as wide as long in ventral view.

Male. Scape not highly modified, 0.9 mm long. Maxillary palps fingerlike,

0.5 mm long. Wings with venation similar to that of L. (L.) latipenne. Anterior

wing brown, approximately 6.0 mm long, very broad, venation similar to that of

L. (L.) latipenne. Hindwing about 5.0 mm long. Genitalia (f. 175) having seg-

ment X with lateral processes reduced to two small, slightly asymmetrical points

at apices of mesal processes in dorsal view. Phallus without parameres, slightly

curved near its base. Inferior appendage first article with fingerlike apex;

second article parallel to first, and broadening towards its apex being truncated

and flattened with two lateral points in dorsal view, ventromesal lobe extending

below first article in lateral view.

Female. Scape 1.0 mm long. Wings brown, venation like that of L. (L.)

latipenne. Anterior wing 6.0 mm long. Genitalia (f. 180) with anterior third of

spermatheca reticulated. Spermathecal sclerite about as wide as long in ventral

view.

Immatures, case and habitat. Unknown.

Life history. Recorded flight dates: 29 March - 19 November.

Distribution. This species is endemic to the Coastal Plains of North and

South Carolina.

Type data. Banks (1911): ‘From Southern Pines, North Carolina, 23 April

(Manee)." Ross (1938b): “Lectotype. - male, Southern Pines, North Carolina,

April 28, A. H. Manee, No. 11704." | have seen the lectotype and one other

male syntype at the MCZC.

Material examined. SOUTH CAROLINA: Aiken Co: Beulah Pond, near

Windsor 3. amales, wlOh We MG2i WH. Gross (INAS) )o 4: males: 1b) IX! -1974;2 PHC,

(CUCC); Savannah River Plant, Upper Three Runs Creek, males, 19-20 XI 19/6;

76. males, 3.females, 29 II! 1977; 14 males, female, 22. VIII 1977; 61 males, 15-16

MS SIs sDDH: BSPo CCUCE)s sbid,.. hiaker .Crom6omaless QOchM 10773 male; 4:1X

TIS AsDDHi Ge, SP) eeUCe) .

Lepidostoma (Lepidostoma) latipenne (Banks)

Gheat6S.) et Foy aye: Gs oo 18nd

Notiopsyche latipennis Banks 1905:216 (male).

Lepidostoma latipennis: Flint and Wiggins 1961:293, 295, f. 11A-B, (female); Wray

1967:56; Etnier and Schuster 1979:19; Neves 1979:171; Parker and Voshell

1981:6; Lake 1984:221.

Lepidostoma sp.: binmieker 496684. 14 JOe 830) SO) SRY 6Oo5 Fs 23,0. b female)

[probably /atipenne].

Lepidostoma latipenne: Fischer 1970:58-59; Roy and Harper 1975:1084; Roy and

Harper 19/9: 162; Morse et al. 1980:91; Unzicker etal, 1982:9.106, 9.107.

This species is closely related to L. (L.) carolina, but somewhat larger, with

anterior wing 8-9 mm in length. The male is distinguished by having inferior

appendage with a long second article pointing dorsoposteriorly. The female's

spermatheca is similar to that of L. (L.) carolina, but its spermathecal sclerite

is much wider than long in ventral view.

Male. Head (f. 169) having scape 1.0 mm long with small rounded basomesal

process and many stout setae. Maxillary palps highly modified with short cylin-

drical basal segment and flexible expansible apical segment bearing different types

of modified setae, ventral setae usually more stout and dark brown, dorsal setae

more scalelike and lighter in color. Mesonotum (f. 169) having scutellum without

Weaver: North American Lepidostomatidae 87

typical pair of mesal warts. Wings (f. 173) with anterior wing broad, approxi-

mately 8.5 mm long; veins of anterior region between C and M widely separated,

but those of anal region closely appressed and converging at distal arculus, small

basal fold along Cu2. Hindwing also broad with 1A and 2A fused. Genitalia (f.

176) having segment X lobelike without lateral processes. Phallus semicircular.

Inferior appendage first article long and fingerlike with several large apical setae

about as long as basodorsal process; second article sinuate, nearly as long as

basodorsal process, pointing dorsoposteriorly and originating from midlength mesal

position of first article; basomesal process slender.

Female. Head (f. 170) with scape 1.3 mm long. Mesonotum (f. 170) having

scutellum with typical pair of mesal warts. Wings (f. 173) with anterior wing

brown, 8.5 mm long. Genitalia (f. 181) with spermatheca similar to that of L.

(L.) carolina, anterior third with reticulate pattern, but spermathecal sclerite

slightly broader than long in ventral view.

Immatures, case and habitat. A few series of larvae have been associated

with adults, but further study is necessary to determine their distinguishing

characteristics. larvae construct a panel type case which is square in cross

section and slightly tapered posteriorly. Immatures are apparently endemic to

second and third order streams of the Atlantic Coastal Plains and Piedmont

regions.

Life history. Recorded flight dates: 15 April - 19 November.

Distribution. Alabama, Delaware, Georgia, New Hampshire, North Carolina,

Nova Scotia, Pennsylvania, South Carolina, Tennessee, and Virginia.

Type data. Banks (1905): "One male from Black Mountains, North Carolina,

June (Beutenmiller)." Al/lotype. - female (Carpenter 1933); Smoky Mountains,

North Carolina, Bryson City, Deep Creek, 2000’, 27 VIII 1930, P. J. Darlington.

Notiopsyche latipennis Banks, New Lectotype. - male, Black Mountains,

North Carolina, Vip MCZC "Type 117037”

Material examined. ALABAMA: Talledega Natl Forest, Shoal Cr at Shoal

Cr Cpg, mate, 6° 1K" 1Gl, “SCH ve: Handley). (SCH; Bipb Co: bitte Sehuitz: Cr

at spring, male, 6 X 1982, SCH, (SCH). GEORGIA: Clarke Co: Beech Creek,

13° males, ‘female, 5-6V°1955* 12. «mates. (female) T2-13°V 1958) (DCs Seatt, ft CUCC):

Union Co: Vogel St Pk, Wolf Creek, 4 males, 30-31 V 1980; male, female, 22 VII

1972, JCM, (CUCC). NORTH CAROLINA: Swain Co: Great Smoky Mts Natl Pk,

Mingus Cr, 7 males, female, 9-10 IX 1983, JSW, (JSW); Haywood Co: Great Great

Smoky Mts Natl Pk, Chestnut Branch, at Big Cr Cpg, male, 21-22 VIII 1982, JSW

& PJS, (JSW); Yancey Co: Black Mountains, South Toe River, 13 males, female,

2 1X 1978, JSW;° (ASW). “PENNSYLVANIA: ‘Forest: Co: Kellettville, Ross Run,

2 males, 27-28 VIII 1976, JSW, (JSW); Pigeon, Marienville Rod & Gun Club,

Hatchery Spring, 3 males, 27-28 VII 1976, JSW, (JSW). SOUTH CAROLINA:

Aiken Co: Savannah River Plant, Tinker Cr, 7 males, 4 females, 1 VI 1977; male,

4 1X 1976; male, 19-20: Xt). DDH €rSP,) 0CUCC):” tbids :\Woper Three’ Runs’ ‘Creek:

2: female pupae, 28 June 1977, DDH & SP, (CUCC): Anderson Co: Broadway

Watershed, Rock Cr, 3 males, 6 females, 3-4 VI 1980; 4 larvae, 31 II! 1981, JSW

& EM Taylor, (JSW); Pendleton, Aldwood, male, 1 V 1976, JCM, (CUCC); Oconee

Co: Crane: Cr, S mi N° of Tamassea, 4 mates; 26-27 V 1960, \JSW 6) TRw, tISw):

10 males, 7 females, 2-3 VI 1980, JSW & EMM, (JSW); E fork Chattooga River at

US Fish Hatchery, N of Walhalla, 2 males, female, 18 VIII 1969; male, 13 VIII 1969,

JCM, (CUCC); Pickens Co:. Wildcat Cr, 9 km NW of Clemson, males ¢ females,

1B IM EK 1968, JOM, PHC, (CRC C hs 28 larmvat 230 ere Pee (eC) 4

larvae, 117M 1978, JSWie FRW, €JSWhe Table Rock St: PkiiCartieok Cry. male; 30

VII 1969)) A ‘Deuglas; ({CUCCI 2 males, OB) X 969 GOM,) (CU Table Rock St

Pk, S Saluda River, male, 29: V 1969: 1 male, 13 X 1969, JCM, €CUCC):; Cleveland,

S. Saluda River, 2 males, 27 VIII 1969, JCM, (CUCC); Rocky Bottom, Eastatoe

Cry male; 19 ¢VP : 1969," male PS WT 1960. mate, 2) Waele) 4868) GOM) COUCC}.,

TENNESSEE: Blount Co: Great Smoky Mts Natl Pk, Rabbit Cr, male, 22-23 VIII

1982, JSW & PJS, (JSW); Sevier Co: Great Smoky Mts Natl Pk, at Chimney Top,

88 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

W Prong Little Pigeon River, male, 2 females, 15-16 VIII 1982, JSW & PJS, (JSW).

VIRGINIA: Shenandoah Natl Pk, Thornton River, male, 2 IX 1961, OSF, (USNM).

Lepidostoma (Lepidostoma) stigma Banks

(fas Te bpclil2 badd £182)

Lepidostoma, stigma Banks.190/:125, pl; -8;-4.. 10. (female): Fischer 1970:64-65;

Nimmo and Scudder 1983:74, 80; Nimmo and Wickstrom 1984:4.

Lepidostoma knowltoni Ross 1938a:1/75, f. 117 (male) NEW SYNONYM: Newell and

potter. 19/3:20, . Fischer, 19/0:98: . Anderson. «1976:41:19; > Webb. 1980:97;

LaFontaine 1981:250; Roemhild 1982:11; Baumann and Unzicker 1981:27:

Ruiter and Lavigne 1985:41; Herrmann et al. 1986:434.

The associations of female with males for each of the five Known North

American species of the subgenus Lepidostoma and the examination of the female

type of L. stigma Banks, revealed that L. knowltoni Ross is a junior synonym

of by (b.) stigma:...1 he male: genitalia. are. very similar: to -those of Lk. (62)

togatum, but L. (L.) stigma is distinguished by having a unique basomesal

process on its antennal scape (f. 171). The anterior half of the female

spermatheca is reticulated (f. 182); those of other species are no more than a

third reticulated.

Male. Scape (f. 171) 1.0 mm long, with oblique mesal process bearing very

large saber-shaped seta and two other stout capitate setae. Maxillary palp

apparently similar to that of L. (L.) togatum with flexible setous apical segment.

Wings brown. Anterior wing (f. 172) venation highly modified, fork of M long

and narrow, Cul unbranched, Cu2 and 1A atrophied, 2A elongated and adjacent

to Cul, both Cul and 2A terminating distally in hind margin separately, 3A short

and only vein terminating basally in hind margin, typical configuration of veins

at arculus absent. Genitalia (f. 177) having segment X with long pair of asym-

metrical, horn-shaped lateral processes, left one longer than right one in dorsal

view: Phalhus..similar to that of L. (L.) togatum, parameres absent, phalicata

curved near its base. Inferior appendage with first article acuminate in lateral

view; second article subapical mesal process of first article, shorter than

basodorsal process; basodorsal process long, clavate; basomesal process

acuminate, about twice as long as its basal width.

Female. Antennal scape 1.2 mm long, cylindrical. Wing venation similar to

that of L. (L.) togatum. Anterior wing about 9.0 mm long. Genitalia (f. 182)

with anterior half of spermatheca cylindrical and conspicuously reticulate;

spermathecal sclerite in ventral view with lateral processes truncated.

Immatures, case and habitat. Examination of one male pupa indicates that

final instars of L. (L.) stigma construct typical panel cases, square sagittally

with individual pieces square. Herrmann et al. (1986) reported that it inhabits

rivers.

Life history. Pupae collected 12 July. Recorded flight dates: 8 July - 30

August.

Distribution. Northern Pacific Coastal region, central and northern Rocky

Mountain regions including British Columbia, Colorado, Montana, Oregon, Utah,

Yukon, and Wyoming.

Type data. Lepidostoma stigma Banks (1907): "One female from Colorado

(Oslar), and two from Boulder, Colo., Aug. 9 (Cockerell), at light." Ross

(1938b): “Lectotype. - female, Boulder, Colorado, August 9, at light, T. D.

A. Cockerell, No. 11692." | have seen the lectotype and two other syntypes at

the MCZC. )

Lepidostoma knowltoni Ross (1938a): "Holotype. - male, Clinton, Utah, June

21, 1936, G. F. Knowlton. Paratype. - 1 male, Utah, same data as for holotype.”

Holotype at the INHS (Webb 1980).

Weaver: North American Lepidostomatidae 89

Material examined. Lectotype female of L. stigma, (MCZC). BRITISH

COLUMBIA: 2.7 km W of Topley, Hwy 16, 2 males, 18 VI! 1979, APN, (CNCI);

1 km. E of ‘Redstone, Chilcotin River, Hwy 20, 13 males, 7 females; 16 VII-1979,

APN, (CNCI). OREGON: Grant Co: Seneca, Silves River, male, 2 females, 1

VII 1976, SDS, (JSW); 1 mi NE of Tabernash, Meadow Cr, male, 7 VIII 1973, DLP,

JP &@ SDS, (SDS); Harney Co: Blitzen River, 3 mi E of Frenchglen, 4400', male

pupa, 3 pupae, collected 12 VII 1964 (2 males emerged 13 VII!), NHA, (OSUO ¢&

ROME). YUKON: Seaforth Cr, Alaska Hwy, 72.6 km W of Teslin Bridge, 9 males,

4 females, 24 VII 1979, APN, (UASM).

Lepidostoma (Lepidostoma) tibiale (Carpenter)

bie. lay tes)

Neuropsyche tibialis Carpenter 1923:39;. pl... 142 3; pli-2 #215) 16: (male; female).

Lepidostoma rileyi Denning 1948a:257, f. 9-9B, 10-10B (male, female); Wray

1967:56; Fischer 1970:66.

Lepidostoma tibialis: Wray 1967:56; Etnier and Schuster 1979:20; Parker and

Voshell 1981:6.

Lepidostoma tibiale: Fischer 1970:65-66; Unzicker et al. 1982:9.106, 9.107.

The male of this species is similar to that of L. (L.) togatum, but is dis-

tinguished by having its inferior appendage without basomesal process. The

female is unique, having spermatheca with small clusters of microtrichia in a

reticulate pattern anteriorly.

Male. Antennal scape 0.5 mm long, cylindrical. Maxillary palp similar to

thatoeok 0 yid Li) togatum: Wings brown, venation similar to that of L. (L.)

togatum. Anterior wing about 5.0 mm long. Genitalia (f. 178). Segment X with

small dorsomesal lobes and prominent horn-shaped lateral processes, left process

slightly longer than right one in dorsal view. Phallus without parameres, phali-

cata not strongly curved at its base. Inferior appendage first article with

rounded apex bearing several stout setae; second article subapical mesal lobe,

much shorter than basodorsal process; basodorsal process clavate and long;

basomesal process absent.

Female. Antennal scape 1.0 mm long. Wings with venation typical for female

lepidostomatids. Anterior wing 6.0 mm _ long. Genitalia (f. 183) having

spermatheca anterior third with many small clusters of microtrichia in reticulate

pattern. Spermathecal sclerite with two small lateral lobes in ventral view.

Immatures, case and habitat. Unknown.

Life history. Recorded flight dates: 3 March - 27 August.

Distribution. Endemic to the southern Piedmont and Coastal Plains regions

of Alabama, Georgia, North Carolina, South Carolina, Tennessee, and Virginia.

Type data. Neuropsyche tibialis Carpenter (1933): "Holotype (male) and

allotype: Bryson City, Deep. Creek, North Carolina, Avo. 27): 1980%-P. J.

Darlington, collector. Paratypes: 1 male, Bryson City, North Carolina, Aug 23;

7 males, Bryson..City, Aug. 26; 3 males, Bryson’City;: Aug! 27>: 24males, Bryson

City, Aug. 23; 3 females, Bryson City, Aug. 26; 1 female, Nantahala Gorge,

North Carolina, Aug. 25, 1930. All types in the Museum of Comparative Zoology."

Male syntype in the INHS (J. D. Unzicker). At the MCZC | have seen the male

type, 9M. C. Ze Type,19442" 2 \.and 14 paratypes:.(9 males, .d: femates), Mic: Z.

Paratype 19442."

Lepidostoma rileyi Denning (1948a): "Holotype. - male, Ela, North Carolina,

Mays 30.0.. W941 ee: Sue Sau? Bagten: Allotype. - female, same data as_ holotype.

Paratypes. - 2 females, same data as _ holotype. Types deposited in the

entomological collection of. the University of Minnesota.

Material examined. ALABAMA: Cheaha Cr, upstream of L. Chinnabee, Mt.

Cheaha St Pk, male, 27 V 1981, SCH € O'Neil, (SCH). GEORGIA: Union Co:

Baxter, Toccoa River, Hwy 60, male, 7 females, 25 IX 1972, (INHS). SOUTH

90 Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

CAROLINA: Oconee Co: Chattooga River, Hwy 28, 2 males, 22 females, 16 VII

1969, JCM, (CUCC). VIRGINIA: Giles Co: New River, McCoy, male, IX 1973,

B Strickler, (USNM); New River, 5 mi N of Radford, male, 2 VI 1971, AB Gurney,

(USNM).

Lepidostoma (Lepidostoma) togatum (Hagen)

(fo hiAg aio 4645

Mormomyia togata Hagen 1861:273.

Silo pallidus Banks 1897:29.

Pristosilo canadensis Banks 1899:212 (male).

Lepidostoma togatum: Ellis 1962:5, 14; Etnier 1965:151; Blickle and Morse

1966:11; Wray 1967:56; Unzicker 1968:4, 14, 19, 50, 51, f. 24 (female);

Fischer 1970:66-67; Unzicker et al. 1970:174; Hilsenhoff et al. 1972:22,

28; Longridge and Hilsenhoff 1973:180; Roy and Harper 1975:1085; Resh

19752153: Misenhotf.. 19%0:25%, : Roy sand. (Harper 1979:163; “Williams: -and

Williams 1979:2407, 2410; Etnier and Schuster 1979:20; Masteller and Flint

1979:169: Neves 1979:171; Mingo and Gibbs 1980:46; Schmid 1980:150, f.

528-529, 531-539 (male, female); Parker and Voshell 1981:6; LaFontaine

1987:249...250...251> Hileenhof? 1981:3165 Unzieker et al.- 198259: 106, ° 9.107;

Marshall and Larson 1982:78, f. 154; Huryn and Foote 1983:/792; Lake

1984:221; Nimmo 1986b:96.

Goerodes togatum: Corbet et al. 1966:1293; Nimmo 1966:692.

This species is very similar to the generotype, L. (L.) hirtum, of Europe.

It differs only in respect to minor details of the male anterior wing and genitalia.

In comparison to the North American species, the male of L. (L.) togatum is most

similar to that of L. (L.) stigma, from which it differs by not having a highly

modified antennal scape; L. (L.) togatum and stigma are apparently allopatric

species.

Male. Scape about 0.7 mm long and with short, rounded mesal bump bearing

several long stout setae. Maxillary palp two-segmented, basal segment

sclerotized, apical segment flexible and with many scalelike setae. Wings (f. 174).

Anterior wing 8.0 mm long, with small basal costal cell bearing many stout setae,

F1 and F3 present, Cul unbranched, and anal region with basal setose pouch.

Genitalia (f. 179) having segment X with small pair of dorsomesal lobes and large,

horn-shaped, asymmetrical lateral processes, right process longer than left one

in dorsal view. Phallus without parameres, phalicata curved near its base.

Inferior appendage first article fingerlike with several stout apical setae; second

article short, subapical mesal lobe; basodorsal process clavate and long; basomesal

process acuminate, about four times as long as its basal width.

Female. Antennal scape 1.0 mm long. Wings (f. 174). Genitalia (f. 184,

199) having spermatheca with less than a fourth of its anterior portion reticulate.

Spermathecal sclerite in ventral view with pair of lateral processes truncated

obliquely. Unzicker (1968: f. 24) provided illustrations of female genitalia which

are apparently of this species; his figure shows an extra spermathecal pouch

which is not shown in f. 199.

Immatures, case and habitat. Immatures inhabit lentic and lotic habitats

indicating that the ecological range of L. (L.) togatum is very wide.

Life history. Mature pupae collected 10 April to 16 July. Recorded flight

dates: 9 April - 30 September.

Distribution. This species is the most widespread lepidostomatid in North

America. It occurs throughout most of the USA and Canada east of the Rocky

Mountains; Alabama, Alberta, Arkansas, Connecticut, Delaware, District of

Columbia, Georgia, Kentucky, Maine, Maryland, Michigan, Minnesota, Missouri,

Newfoundland, New Hampshire, New York, North Carolina, North Dakota,

Weaver: North American Lepidostomatidae 91

Northwest Territories, Ohio, Ontario, Pennsylvania, Quebec, Rhode Island,

Saskatchewan, South Carolina, Tennessee, West Virginia, and Wisconsin.

Type data. Mormomyia togata Hagen (1961): "Hab. Washington; St.

Lawrence River, Canada (Osten Sacken)."” Ross (1938b): "Lectotype. - female,

St. Lawrence River, Canada, 1859, Osten Sacken, No. 10942." | have seen the

lectotype and one other syntype at the MCZC.

Silo pallidus Banks (1897): "Several specimens from Ithaca, N.Y." New

Lectotype. - male, Ithaca, New York, 2 July, without a red type label, selected

by Ross, 1973, but unpublished. One syntype female, same data, but with red

MCZC label, "Type 11690."

Pristosilo canadensis Banks (1899): "Sherbrooke, Canada, June (Bégin)."

New Lectotype. - male, same data as above, MCZC "Type 11689."

Material examined. ALABAMA: Cheaha Cr upstream of L. Chinnabee, Mt.

R., Hwy 31, male, 4 |X 1982,,-SCH, (SCH). ALBERTA: Edmonton, male, female,

17 VII 1955, GE Ball, (UASM); Muskeg River, N of Ft McMurry, male pupa, 21

Vl 1978, APN, (UASM); Sturgeon River, Hwy 43, 2 mi W of Onoway, male, 9

females, 18 V! 1966, .APN, CUASM); Sundance, male; 33 females; 623: VWht1955, B

Hockin, (UASM); LaBiche River, Hwy 63,--male, 29 VI 1978, APN, (UASM);

Wandering River, Hwy 63, male, 29 VI 1978, APN, (UASM); Blindman River, Hwy

12, Rimbey, 10 males, 7 females, 25 VI 1967, APN, (UASM); Lesser Slave Lake,

E of Wagner, male, female, 19 VII 1965, APN, (UASM). MAINE: Aroostook Co:

Big Black River (T15 R13), male, female, 25 VII 1961, A Brower, (USNM); Oxbow,

male, 5 females, 9-10 VII 1961, A Brower, (USNM). MARYLAND: Plummer's Isl.,

male, EA Schwartz, (USNM). MISSOURI: Carter Co: Big Spring St Pk, male,

19. V. 1972, RWB, C(USNM);.lron: Cow Annapolis, “Champion Spring; male’ pupa, 'S

larvae, 10 IV 1950, GA Ulrich, (INHS); Maries Co: Paydown Spring, Hwy 42, 7

mi W of Belle, 10 males, 3 females, 27 V 1972, BA Sussman, (USNM); Oregon

Co:, Graveyard Spring, male, 12 IV 1972, RWB, (USNM); Shannon Co: Alley

Spring, Ozark Natl Riverway, 60 males, 15 females, 19 V 1972, RWB, (USNM);

Ibid, 19’ males, 20 femates, 26 V 1973, JCM, (CUCC): Round Spring; Hwy 19,

Colesville, Clove Brook, 10 males, 2 females, 11 VI 1977, DE Ashton & JSW,

(JSW). NEW YORK: Lewis Co: Whetstone, Gulf St Pk, male, 3 females, 21 VI

1973, JCM, (CUCC). NORTHWEST TERRITORIES: Great Slave Lake, Simpson

lel, 2). males, TA WT: 1964s) male ZEON TT (Abe) OD Larson, | CUASME. NOVA

SCOTIA: Queens Co: Lake Kejimkujik, male, female, 15 VI 1958, male, 7 females,

2 VII 1957; 6 females, 1961, DC Ferguson, (USNM). PENNSYLVANIA: Pike

Co: Metamoras, Delaware River, 6 males, 12 females, 11 VI 1977, DE Ashton §&

JSW, (JSW). QUEBEC: Norway Bay, 9 males, 21 females, 4-10 VI! 1974, OSF,

(USNM). RHODE ISLAND: Washington Co: Richmond, 2 females, 20 VI 1971,

AG Lavallee, (CUCC). SASKATCHEWAN: Makwar, Hwy 55, Rapid View male, 4

females, 20 VI 1979, APN, (UASM); Sandy Lake, Prince Albert Natl Pk, larva,

male pupa, 16 VII 1970, ROME staff, (ROME). SOUTH CAROLINA: Oconee Co:

Chattooga River, Hwy 28).2 males, TO 1k 1969) JCM, (CUCC)..: VIRGINIA:

Pulaski Co: New River, 5 mi N of Radford, male, 2 VI 1971, AB Gurney, (USNM).

WEST VIRGINIA: Pendleton Co: Smoke Hole St Pk, Briggs Run, male, female,

9 IV 1977, D & M Davis, (USNM). WISCONSIN: Douglas Co: Gordon, male, 2

females, 20 VII 1967, A Lemke, (USNM).

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Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

LIST OF FIGURES

‘agi 1-6. Larval cases:

Ot) eS =

Lepidostoma pluviale, stone case.

Lepidostoma libum, stone case.

Lepidostoma acarolum, stone case.

Lepidostoma ontario, panel case.

Lepidostoma knulli, turret case.

Lepidostoma frosti, turret case.

Figures 7-9. Larval heads, ventral:

Theliopsyche melas.

Lepidostoma ontario.

Lepidostoma frosti.

Figures 10-13. SEM micrographs:

10.

13%

Theliopsyche corona, modified seta of male forewing, 2,000 X.

Lepidostoma griseum, male: A, mesonotum, 120 X; B, tibial spurs, 110 X.

Lepidostoma americanum, setae of the costal cell, male forewing: A, 500 X;

Be 1 gO0O eGo t De O00) OS:

Lepidostoma cascadense, setae in male forewing anal fold: A, 200 X; B, 500

Figures 14-18. Secondary sexual characteristics of Theliopsyche:

14,

Theliopsyche corona, male head and thorax, left lateral.

Theliopsyche corona, male: A, head, pronotum and mesonotum, dorsal; B,

right tibiae and tarsi, mesal.

Theliopsyche corona, wings: A, male; B, female.

Theliopsyche parva: <A, male wings; B, female hindwing.

Theliopsyche melas, male hindwing.

Figures 19-23. Male genitalia of Theliopsyche, left lateral (A); terga IX and X,

dorsal (B); left inferior appendage, dorsal (C); phallus, dorsal (D):

Theliopsyche corona, North Carolina, Wild Cherry Branch.

Theliopsyche epilonis, holotype.

Theliopsyche grisea, Pennsylvania, Whisky Spring.

Theliopsyche melas, paratype.

Theliopsyche parva, lectotype: E, segment VII, ventral.

Figures 24-26. Female genitalia, left lateral (A), and ventral (B):

24.

pase

Theliopsyche corona.

Theliopsyche epilonis.

Theliopsyche parva, lectoallotype.

Figures 27-56. Secondary sexual characteristics of Lepidostoma (Nosopus):

24s

26%

29,

Lepidostoma recinum, male, head, left lateral.

Lepidostoma licolum, male: A, head, pronotum and mesonotum, dorsal; B,

head, left laperall,

Lepidostoma podagrum, head, prothorax and foreleg, left lateral: A, male;

B, female.

Lepidostoma quercinum, male: A, head, pronotum and mesonotum, dorsal;

B, head, left lateral.

Lepidostoma americanum, male head, left lateral.

Lepidostoma modestum, male: A, head, pronotum and mesonotum, dorsal:

B, head, left lateral.

Lepidostoma cinereum, male, left scape, dorsal.

Figures 5/-96.

Weaver: North American Lepidostomatidae 103

Lepidostoma unicolor, male, head, pronotum and mesonotum, dorsal.

Lepidostoma mexicanum, male, head, left lateral.

Lepidostoma cascadense, male: A, left scape, dorsal; B, maxillary palp,

extended, left lateral; C, mesoscutellum, dorsal.

Lepidostoma hoodi, male, head, left lateral.

Lepidostoma spicatum, male: A, head, dorsal; B, left scape,

expanded, dorsal; C, head, left lateral.

mesal portion

Lepidostoma

verodum, male head, left lateral.

reosum, male wings.

asteneum, male wings.

jewetti, male wings.

licolum, male wings.

americanum, male wings.

podagrum, wings: A, male;

modestum, wings: A, male;

unicolor, wings: A, male;

roafi, male wings.

delongi, male wings.

B, female.

Lepidostoma acarolum, male wings.

Lepidostoma frontale, male wings.

Lepidostoma heveli, male wings.

Lepidostoma lacinatum, male wings.

Lepidostoma hoodi, wings: A, male; B, female.

Lepidostoma cascadense, wings: A, male; B, female.

Lepidostoma verodum, wings: A, male; B, female.

Male genitalia of Lepidostoma (Nosopus), left lateral (A); terga

IX and X (B); left inferior appendage, dorsal (C); phallus, left lateral (D);

phallus, dorsal (6):

Lepidostoma

reosum, holotype.

asteneum, holotype.

recinum, holotype.

jewetti, holotype.

licolum, paratype.

Lepidostoma americanum, Pennsylvania, Maul Spring.

Lepidostoma bryanti, Pennsylvania, Ross Run.

Lepidostoma costale, Pennsylvania, Pigeon.

Lepidostoma prominens, Quebec, Grand Lac Jaques Cratier.

Lepidostoma podagrum, \|daho, Teton River.

Lepidostoma quercinum, Oregon, Camp Adair.

Lepidostoma roafi, Washington, Cle Elum River.

Lepidostoma compressum, paratype.

Lepidostoma lydia, Pennsylvania, Maul Spring (phallus expanded).

Lepidostoma modestum, North Carolina, Whitewater Falls: (phalius expanded)

F, phallus, ventral.

Lepidostoma ontario, New Hampshire, Mt. Pleasant Brook.

Lepidostoma pictile, New York, Allegany State Park (phallus expanded).

Lepidostoma bakeri, paratype.

Lepidostoma cinereum, Washington, Yakima River:

paremeres, dorsal.

F, phalicate and

Lepidostoma

frosti, Pennsylvania, Maul Spring.

knulli, holotype.

unicolor,, |daho, Wind Lake,

acarolum, New Mexico, Glenwood.

aztecum, holotype.

delongi, Sonora, Esperanza.

denningi, holotype.

frontale, holotype.

Contrib. Amer. Ent. Inst., Volume 24, Number 2, 1988

84. Lepidostoma heveli, holotype.

85. Lepidostoma lacinatum, paratype.

86. Lepidostoma mexicanum, Arizona, Portal:

8/. Lepidostoma quila, paratype.

88. Lepidostoma rectangulare, holotype.

89. Lepidostoma reimoseri, paratype.

90. Lepidostoma stienhauseri, holotype.

91. Lepidostoma talamancense, Costa Rica, Ojo de Aqua.

92. Lepidostoma ermanae, holotype.

93. Lepidostoma hoodi, Washington, Minotaur Creek.

94. Lepidostoma spicatum, paratype.

95. Lepidostoma cascadense, holotype.

96. Lepidostoma verodum, California, Sagehen Creek.

F, phalicata and parameres, dorsal.

Figures 97-121. Female genitalia of Lepidostoma (Nosopus), left lateral (A) and

spermathecal sclerite, ventral (B):

97. Lepidostoma recinum.

98. Lepidostoma jewetti.

99. Lepidostoma americanum.

100. Lepidostoma bryanti.

101. Lepidostoma costale.

102. Lepidostoma prominens.

103. Lepidostoma podagrum.

104. Lepidostoma quercinum.

105. Lepidostoma roafi.

106. Lepidostoma lydia.

107. Lepidostoma modestum.

108. Lepidostoma ontario.

109. Lepidostoma pictile.

110. Lepidostoma bakeri.

111. Lepidostoma cinereum.

112. Lepidostoma frosti.

113. Lepidostoma knulli.

114. Lepidostoma unicolor.

115. Lepidostoma acarolum.

116. Lepidostoma mexicanum.

117. Lepidostoma cascadense.

118. Lepidostoma ermanae.

119. Lepidostoma hoodi.

120. Lepidostoma verodum.

121. Lepidostoma deceptivum, lectotype: C, spermatheca, left lateral.

Figures 122-126. Secondary sexual characteristics of Lepidostoma (Mormomyia):

122. Lepidostoma griseum, male: A, head, left lateral; B, mesonotum, dorsal.

123. Lepidostoma morsei, male mesonotum, dorsal.

124. Lepidostoma vernale, male: A, head, pronotum and mesonotum, dorsal; B,

head, left lateral.

125. Lepidostoma griseum, wings: A, male; B, female.

126. Lepidostoma vernale, wings: <A, male; B, female.

Figures 127-144. Male genitalia of Lepidostoma (Mormomyia), left lateral (A);

terga |X and X (B); inferior appendages, dorsal (C); phallus, left lateral

(D2. phallus, “dorsal CE):

127. Lepidostoma griseum, Pennsylvania, Maul Spring.

128. Lepidostoma morsei, holotype.

129. Lepidostoma carrolli, paratype: F, phallus expanded, left lateral.

130. Lepidostoma serratum, holotype.

131. Lepidostoma etnieri, holotype.

Lepidostoma

Figures 142-150.

and spermathecal sclerite, ventral (B):

Lepidostoma

Figures 151-153.

Ve ks

152:

1335

Weaver: North American Lepidostomatidae 105

excavatum, holotype.

libum, holotype.

sommermanae, Pennsylvania, Maul Spring.

flinti, paratype.

glenni, holotype.

sackeni, Pennsylvania, Maul Spring.

vernale, Pennsylvania, Chestnut Bridge Hollow.

lobatum, holotype.

mitchelli, holotype.

styliferum, holotype.

Female genitalia of Lepidostoma (Mormomyia), left lateral (A)

griseum.

serratum.

excavatum.

libum.

sommermanae.

flinti.

sackeni.

vernale.

mitchelli.

Secondary sexual characteristics of Lepidostoma

(Neodinarthrum):

Lepidostoma pluviale, male: A, head, pronotum and mesonotum, dorsal;

B, head, left lateral.

Lepidostoma pluviale, wings: A, male; B, female.

Lepidostoma canthum, forewings: A, male; B, female.

Figures 154-162.

terga IX and X, dorsal (B); left inferior appendage, ventral (C); left

inferior appendage, dorsal (D); phallus, left lateral (E):

Lepidostoma

Male genitalia of Lepidostoma (Neodinarthrum), left lateral (A);

canthum, California, Potreros.

baxea, holotype.

apornum, holotype.

errigenum, paratype.

lotor, holotype.

ormeum, holotype.

pluviale, holotype: F, segment X, caudal.

Lepidostoma pluviale, segment X, left lateral of 12 males from the same light

trap collection, Utah, Provo River.

Lepidostoma

Figures 163-168.

(A) and spermathecal sclerite, ventral (B):

Lepidostoma

rayneri, holotype.

Female genitalia of Lepidostoma (Neodinarthrum), left lateral

baxea.

canthum.

errigenum.

ormeum.

pluviale: C, spermathecal sclerite, vagina and segment |X,

ventral; D, segments VII, VIII| and IX, ventral; E, spermathecal sclerite

and vagina, lateral.

Lepidostoma

rayneri.

106

Contrib. Amer.

Figures 169-174.

(Lepidostoma):

169.

170.

TET.

Ti2..

174.

Lepidostoma

Ent. last.;: Volume 24, Number 2, 1988

Secondary sexual characteristics of Lepidostoma

latipenne, male: A, male head, pronotum and mesonotum,

dorsal; B, male head, maxillary palp expanded, left lateral; C, male

maxillary

Lepidostoma

Figures 1/5-179.

terga |X and X, dorsal (B); left inferior appendage, dorsal (C); phallus,

left lateral (D):

Lepidostoma

lL epidostoma

Lepidostoma

Figures 180-184.

and spermathecal sclerite, ventral (B); spermatheca, left lateral (C):

Lepidostoma

palp contracted, left lateral.

latipenne, female: A, head, dorsal; B, mesoscutellum, dorsal.

stigma, male, left scape, dorsal.

stigma, male forewing.

latipenne, wings: A, male;

togatum, wings: A, male;

B, female.

Male genitalia of Lepidostoma (Lepidostoma), left lateral (A);

carolina, South Carolina, Upper Three Runs Creek.

latipenne, South Carolina, Crane Creek.

stigma, Oregon, Senica.

tibiale, South Carolina, Chattooga River.

togatum, Pennsylvania, Delaware River.

Female genitalia of Lepidostoma (Lepidostoma), left lateral (A)

carolina.

latipenne.

stigma.

tibiale.

togatum.

ADA

4 | sees SB)

= ae

wen a

SS . SS

. ~ : S — = ee a oe ee ee a eS Re ee

: : = S < SSS ee es oe Zs es oe - : : y

a SSS Ss eS ars

his Sg a ae pats, eA me Z

ps See oe

Zs P Z,

yi) ' ih Ni

W il)

80A

79B

78C

79C

80C

78D

79D

81D

ES sy’,

87C

88C

89B 89C

90C 90D

20°

100800;

oP S05 0S 0%

157C 157D

160D 160F

Zs sali eile

elt ak b v7

138 Contrib. Amer. nt.

Inst., Volume 24, Number 2, 1988

INDEX

Valid names are in roman type, with principal references in bold.

Synonyms,

preoccupied names, and epithets of inappropriate gender are in italics.

acarola, Lepidostoma 4/7

acarolum, Lepidostoma (Nosopus) 8, 17,

23 86; At

Alepomyia 19

Alepomyiodes 19

alexanderi, Lepidostoma 51

americana, Lepidostoma 28

americana, Olemira 19, 28, 29

americanum, Lepidostoma (Nosopus) 1,

Pia sO, ot oe 40. 42543

americanum group, Lepidostoma

tNosopus). 18,19, 21, 26

Annulipalpia 9

Aopsyche 10

aporna, Lepidostoma 7/9

apornum, Lepidostoma (Neodinarthrum)

POA BS

Arcadopsyche 19

astanea, Lepidostoma 25

astaneum, Lepidostoma (Nosopus) 1, 21,

29; 26

astaneum group, Lepidostoma (Nosopus)

7 19,2 20,28

Atomyia 19

Atomyiodes 3, 15, 19

aztecum, Lepidostoma (Nosopus) 24, 46,

ea iis

bakerit, Lepidostoma (Nosopus) 17, 22,

40, 41, 44, 45;.-47,. 60

basalis, Lasiocephala 6

baxea, Lepidostoma (Neodinarthrum) 76,

Pa pee fs

bispinosa, Atomyiodes 19, 51

bispinosa, Lepidostoma 51

bispinosus, Atomyiodes 19, 51

bryanti, Alepomyia 19, 29

bryanti, Lepidostoma (Nosopus) 21, 28,

20, ot ee OU

calense, Lepidostoma 1

calensis, Lepidostoma 26

Calocidae 4

canadensis, Pristosilo 16, 90, 91

cantha, Lepidostoma //

canthum, Lepidostoma (Neodinarthrum)

Tey tt

canthum group, Lepidostoma

(Neodinarthrum) 8, 75, 76

carolina, Lepidostoma (L.) 84, 85, 86,

carolina, Notiopsyche 16, 85

carrolli, Lepidostoma (Mormomyia) 62,

655° 67

cascadense, Lepidostoma (Nosopus) 1,

BONG 20,000, 06.87

cascadensis, Jenortha 19, 5/7, 58

cascadensis, Lepidostoma 5/

cinereum, Lepidostoma (Nosopus) 1, 16,

167 19,227.40, 41, 44

cinereus, Silo 41, 42

compressa, Lepidostoma 36

compressum, Lepidostoma (Nosopus) 22,

30) Set, oe

corona, Theliopsyche 8, 10, 11, 12, 13,

costale, Lepidostoma (Nosopus) 21, 28,

Oi," 4

costalis, Lepidostoma 30

costalis, Olemira 30

crassicornis, Neoseverinia 6

crypta, Lepidostoma 83, 84

cryptum, Lepidostoma 1, 84

dafila, Lepidostoma (Nosopus) 23, 46,

deceptiva, Thremma 60

deceptivum, Lepidostoma (Nosopus) 1,

16, 60

delongi, Lepidostoma (Nosopus) 23, 46,

48, 52

denningi, Lepidostoma (Nosopus) 1, 24,

A6;49 32°93

Dinarthrella 54

Dinarthrodes 6

Dinrarthrum 6°75

epilone, Theliopsyche 12

epilonis, Theliopsyche 11, 12, 14

Eremopsyche 3, 19

ermanae, Lepidostoma (Nosopus) 1, 6,

PE wp, Oa we

errigena, Lepidostoma /9

errigenum, Lepidostoma (Neodinarthrum)

16, 79,03

etnieri, Lepidostoma (Mormomyia) 1, 62,

65, 67 ,

excavatum, Lepidostoma (Mormomyia) 16,

Gey, Or, Oo, v4

fischeri, Lepidostoma 1, 83, 84

Weaver:

flinti, Lepidostoma (Mormomyia) 62, 70,

al, ¢2,5°95,) 08

Flura 16

frontale, Lepidostoma (Nosopus) 23, 46,

49.03)

frontalis, Eremopsyche 19, 49

frontalis, Lepidostoma 49

frosti, Atomyia 43

frosti, Lepidostoma (Nosopus) 17, 22,

A, 41, Ag, 4

glenni, Lepidostoma (Mormomyia) 62, 70,

goedeni, Lepidostoma 1, 25

Goerodes 3

gracilicornis, Mormonea 15

grisea, Lepidostoma 63

grisea, Phanopsyche 60, 63

grisea, Theliopsyche 9, 11, 13, 14, 15

griseum, Lepidostoma (Mormomyia) 6, 29,

60, 61, 6323/6457 65,665. 65, 770,71,

griseum group, Lepidostoma (Mormomyia)

60, 61, 63

griseus; Silo 10, \13

heveli, Lepidostoma (Nosopus) 23, 48,

49, 53

hirta, Phryganea 15

hirtum, Lepidostoma (L.) 4, 5, 84, 90

hoodi, Lepidostoma (Nosopus) 6, 17, 20,

Oo, . co, DeaneT,

hoodi group, Lepidostoma (Nosopus) 5,

175020, -54

iranicum, Dinarthrum 6

irrorata, Crunoecia 5

Jenortha 19

jewetti, Lepidostoma (Nosopus) 4, 19,

24, 265 ae

jewetti group, Lepidostoma (Nosopus) 18,

9, 24, 26,28

knowltoni, Lepidostoma 1, 88

knulli, Lepidostoma (Nosopus) 17, 22,

40, 41. 44, 45, 66

lacinatum, Lepidostoma (Nosopus) 23, 46,

473249,).60) See 530 54

latipenne, Lepidostoma (L.) 6, 84, 85,

latipennis, Lepidostoma 86

latipennis, Notiopsyche 16, 86, 87

leechi, Lepidostoma 44

leonilae, Lepidostoma (Nosopus) 24, 46,

North American Lepidostomatidae 139

Leamidostottear 1502; )3904) 1b ,0F, Op O5-410,

19, 20 36) 48, BO). 13484

Lepidostoma (subgenus) 1, 2, 8, 9, 18,°

84, 85

Lepidostomatidae 1, 2, 3, 10

lepidostomatinae 3

liba, Lepidostoma 68

libum, Lepidostoma (Mormomyia) 16, 63,

67, 68,69

libum group, Lepidostoma (Mormomyia)

5, 16,6), OZ, Bie

licola, Lepidostoma 2/

licolum, Lepidostoma (Nosopus) 21, 26,

lobatum, Lepidostoma (Mormomyia) 62,

rk a

lotor, Lepidostoma (Neodinarthrum) 76,

lydia, Lepidostoma (Nosopus) 22, 35,

86. S76

melas, Theliopsyche 10, 11, 14

mexicana, Lepidostoma 51

mexicana, Olemira 19, 51

mexicanum, Lepidostoma (Nosopus) 17,

205 (AGA 5 AO 030 61, 3

mexicanum group, Lepidostoma (Nosopus)

ele poles eae 323 7°46

Micrasema 5

mira, Lepidostoma 57, 58

mirum, Lepidostoma 1

mitchelli, Lepidostoma (Mormomyia) 62,

67 OO Ia. 74

mitchelli group, Lepidostoma (Mormomyia)

eZ. 73

modesta, Atomyia 19, 37

modestum, Lepidostoma (Nosopus) 22,

30, 36,) BT 2G yraee 4447

modestum group, Lepidostoma (Nosopus)

6759) 1B ABs S20 SS 4G

moneka, Lepidostoma 80, 81

monekum, Lepidostoma 1, 80, 81

Mormomyia (subgenus) 1, 2, 5, 8, 9, 16,

to, VS, 60, 61,085

Mormonia 15

morsei, Lepidostoma (Mormomyia) 1, 61,

63, 64

Neodinarthrum (subgenus) 1, 2, 5, 8,

02 16,..48) Feo9s

Neuropsyche 16

Nosopus (subcenvs)}els2y4.55,. Gps,

9 BB i 205 OF 2075

Notiopsyche 16

oaxacense, Lepidostoma (Nosopus) 23,

46, 52

oaxacensis, Lepidostoma 52

140 ContribsiAmen: Ent:

Oeconesidae 4

Olemira 19

Oligopsyche 16

ontario, Lepidostoma (Nosopus) 22, 35,

3B yi on

ormea, Lepidostoma 80

ormeum, Lepidostoma (Neodinarthrum)

iL, LOy 80S), 88

pallidus, Silo 90, 9h

Paradinarthrodes 3

parva, iheliopsycher HO) ol2.c13, 14

Phanopsyche 60

pictile, Lepidostoma (Nosopus) 1, 22, 35,

36:38) 189

pictilis, . Mormonia::28,;.38, 39

pilosa, Goera 5

pleca, Lepidostoma 5/, 58

pluviale, Lepidostoma (Neodinarthrum)

Poe foie, Fi, Ono, ~o0 9681, 83

pluviale group, Lepidostoma

(Neodinarthrum) 75, 76, 78

piuviclis, Olemira (9) 18 e482

podager, Lepidostoma 31

podager, Nosopus 19, 31

podagrum, Lepidostoma (Nosopus) 4, 6,

Ay Gus 2OU NV 8t, 38

podagrum group, Lepidostoma (Nosopus)

FRAT NR 3s PG 1)

Pristosilo 16

prominens, Arcadopsyche 19, 31

prominens, Lepidostoma (Nosopus) 21,

285; 80 31

guercina, Lepidostoma 33

quercinum, Lepidostoma (Nosopus) 4, 21,

Suit 32 83

guerla, Lepidostoma 80, 81

guerlum, Lepidostoma 1, 80, 81

qguila, Lepidostoma (Nosopus) 24, 46, 52

Quisilo 10

rayneri, Lepidostoma (Neodinarthrum)

Lis FOO es

recina, Lepidostoma 26

recinum, Lepidostoma (Nosopus) 1, 21,

ZO 20

rectangulare, Lepidostoma (Nosopus) 24,

AG), 49,108

reimoseri, Lepidostoma (Nosopus) 24, 46,

| Oo 3 1S

reosa, Lepidostoma 24

reosum, Lepidostoma (Nosopus) 16, 19,

20, 24

reosum group, Lepidostoma (Nosopus)

160d 9a 2A

rhino, Lepidostoma 1, 81, 82

Inst., Volume 24, Number 2, 1988

rileyi, Lepidostoma 89

roafi, Atomyia 34

roafi, Lepidostoma (Nosopus) 23, 33,

347446 ,190) 59

roafi group, Lepidostoma (Nosopus) 18,

220.33

roemhildi, Lepidostoma 1, 56, 5/

sackeni, Lepidostoma (Mormomyia) 61,

6B FO, 7

sackeni, Mormomyia 7/1, 7/2

Sericostomatidae 1, 3

serratum, Lepidostoma (Mormomyia) 62,

65, 66

serratum group, Lepidostoma

(Mormomyia) 65

sommermanae, Lepidostoma (Mormomyia)

16; 6 6341°67>. (GB4069

spicata, Lepidostoma 56, 5/

spicatum, Lepidostoma (Nosopus) 1, 6,

720 B40 3h 56

squamulosum, Lepidostoma 15

steinhauseri, Lepidostoma (Nosopus) 23,

46. 49. 53

stigma, Lepidostoma (L.) 1, 84, 85, 88,

89, 90

strophe, Lepidostoma 1, 41

strophis, Lepidostoma 41, 42

stylifer, Lepidostoma /4

styliferum, Lepidostoma (Mormomyia) 62,

73, 14

swannanoa, Lepidostoma 1, 38, 39

talamancense, Lepidostoma (Nosopus) 1/7,

2A ay 2. OF

tallapoosa, Theliopsyche 9, 11, 13, 15

Tasimiidae 3

Thelteopsy chew) 33 4.3.5) 0. 8,028, *10;

da 24

tibiale, Lepidostoma (L.) 84, 85, 89

tibialis, Lepidostoma 89

tibialis, Neuropsyche 16, 89

togata, Mormomyia 15, 90, 91

togatum, Goerodes 90

togatum, Lepidostoma (L.) 4, 84, 85, 88,

89, 90

Trichovespula 3

unicolor, Lepidostoma (Nosopus) 8, 18,

We Oe Ban BA AO AS Ci dl AG

unicolor, Mormomyia 44, 45

unicolor group, Lepidostoma (Nosopus)

Gy, 22. M2 eh (4Qs 4G

veleda, Lepidostoma 81, 82

veledum, Lepidostoma 1, 81, 82

Weaver: North American Lepidostomatidae 14]

vernale, Lepidostoma (Mormomyia) 8, 62, verodum, Lepidostoma (Nosopus) 5, 6,

63) 6S; 66,67) G6. SOR FO Ea. 17) 203 34, 59

t3 44

vernale group, Lepidostoma (Mormomyia) weaveri, Lepidostoma (Nosopus) 22, 35,

6b, G2) 108.73 36, 39

vernalis, Lepidostoma /2 wisconsinense, Lepidostoma 29

vernalis, Mormonia 60, 7/2 wisconsinensis, Lepidostoma 19, 29

vernalis group 15, 60

veroda, Lepidostoma 59 Yamatopsyche 3

=e. F

Wats

¥ 1

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PASSE nt

hariay ee

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Contributions

of the

American Entomological Institute

Volume 24, Number 3, 1988

Pison in the New World: a revision

(Hymenoptera: Sphecidae: Trypoxylini)

Arnold S. Menke

uh,

Big Sat

[itst)

rly

' bs

W ral i

Pison in the New World: a revision

(Hymenoptera: Sphecidae: Trypoxylini)

Arnold S. Menke

Systematic Entomology laboratory

Agricultural Research Service, USDA

c/o U. S. National Museum, Washington DC 20560

ABSTRACT

The genus Pison Jurine in the Western Hemisphere is revised. Identification keys

to the Neotropical Region. All species are described, diagnosed, and their geographic

ranges outlined. Illustrations accompany the key and descriptions. Twenty nine new

species are described: abathes (Ecuador, Bolivia, Guyana), abothrum (Colombia,

Brasil), arachniraptor (Panama to Bolivia, Brasil), aranevorax (Colombia, Ecuador,

Peru, e Brasil), brasilium (Brasil), chrysops (Costa Rica to Argentina), cooperi (Costa

Rica to n Bolivia, c Brasil, Dominica), delicatum (South America), dementia (se

Brasil), doqqonum (Mexico), eu (Mexico to Peru & Suriname), euryops (Colombia,

Brasil, Argentina), erebus (Colombia), eyvae (Colombia, Ecuador, Bolivia), fritzi

(Ecuador, Brasil, Argentina), gnythos (Colombia, Ecuador, Guyana, Trinidad), larsoni

(Ecuador, Peru, Bolivia), lillo (Argentina), longicorne (Mexico to Argentina), martini

(Ecuador), nosferatu (Venezuela), oaxaca (Mexico), pentafasciatum (s Brasil),

phthinylla (Ecuador), sphaerophallus (Colombia, Ecuador, Peru, Guyana, Suriname, n

Brasil), styphopteron (Colombia, Peru), sylphe (Peru), vincenti (Ecuador, Guyana), and

wasbaueri (Argentina). One species is synonymized: flavolimbatum Turner, 1917 =

cressoni Rohwer, 1911. Pison laeve Smith, known only from its type specimen and

sometimes treated as a North American species, is interpreted as an Australasian

taxon and a diagnosis based on the type is presented.

Twelve species groups are established for the New World fauna, but the use of

subgenera is abandoned. Three generic names that have been treated as subgenera in

the past, Pisonoides Smith, Krombeiniellum Richards, and Entomopison Menke are

placed in synonymy with Pison. Some characters that are important from a

phylogenetic standpoint in Pison and related genera are analyzed to determine

polarity. Incorporation of the Crabroninae with the Larrinae is discussed. The status

of Pisonopsis as a genus is reviewed, a new generic character described, and a key to

its five species is presented. Two new species of Pison are described from New

Guinea that are important for an understanding of generic limits: woji and pistillum.

CONTENTS

TPO OGUOERORY ao cs eo a ha ene a ye eos pala ay ie cee a ]

Beknowledqements:, «k's aa yw whe OE ay ee eo ee a a ee, ee l

Mathods ard Technicides 6c ccaie aia he! oak Hk a hea 8 ee A nae es 2

Distinctions between Pisom:and Pigonapsis . . 4. .) « s. <4) aWPRNA or eae ses 3

Gienue Pisom JUGS: . i oxo GE dn we iparlen BL Ag aU Bab ey. ie lea ee ty 5

SUPGENOPS | 6 cya k. 6 le ee ee ee ek ow ee) GN nts 4 «4 5

Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

SPEGIES, GPUS |i NE re WOME ti Se EG ee ee Oe eg 7

che ey 0 gh cage oe “RRR USAR GTO) Rare UMS Re a track Vey ea Meg Aire 8

(ae a oP 1 OL) Re a IP CEG SO 13

I og ieee ak) aces Ape ars | he Bane nT Reg edman cry ig ULE St ey 18

mey TO species OF NeW WORE Piste eee Be ee Vee See ee 8 19

HST ERE OR eG.0 5 aR UG SCT SSE SICA BNR C8 AS 8 EU Sen ea 24

Oe he RCT Ceo erate vain ial ag anes aie ME ar tra ian Cai Agel cucay NTN Maat ae a 2>

ORO Fie ats pa ae alee ere er Wa COU RMN RMR, Ug: Ge. 26

OR Ie Rt cele GOs ee ST OS a ale Ca ue 27

DEST Rt a ti aa RE RS RN SOK RR Gg igs Gove a Za

Peiog OrOue 20k | ach aio aS NOL ig Ve Ue rs eon MMe" aAntar naib, NY bt 30

POE Pe Ree Un he are RLT UN RMN TS eg Meee | 3]

ba Pera Gere oi ais hoa heap tet al tS Malia a es lL ge 32

I re Eee hehe Mt te Me, Sie NUR ORIEN ESR, GRU Mats Se ay pret oe Sg 33

BROT HPOM Tere le a fe OO ha aaa ea Saye ety 33

OG CG etre ee a I OM URI, Gog ThA ALES g LAP og ty ao

BU a td PRES 0h hig Ae a SRG) AMG A CNS eg hay 35

Fae elo gs) 8 a No) Ne a eT na OE Vai INS ok Mark ae nen Pasa SD Ong en ne ee ae 37

BRT Cs Ta ae Eyer a Grr We Bate eg impengTM SIE Nag gs ees FE A i eg 38

PRUNE CPCI ee ree rg Eg ke LE ARATE | PRIDE TS Love tg ig MPa ta HOB OY Sac 39

CUT Ts UR cla) co kN er ee A ce a sel ae ae 40

SER Power ar ey te OMe Lr oc et AEN Sg SRI Ig TRG OC eS ea g 41

eae eB og UM Rie Ta Sa TNE SIREN 013s US Aa ORE Re 42

ees ee rn OEE CEES oe. Games Obl Ay MNS oy RSI 43

PPO COM eye eRe ee kk BN tse es all Goth CNIS PS Tg PRIN mig eg lager gl Gang 44

EOF a rN NN ee hE by PM A Sn SAE MS ge ey 45

EOE es 8 CSE GE CAR A LOMO eS SOR ise a a ee a Se ie a 46

PRO EN ae Carrey Mal aay rata ee ap Nema AT RA tage MANS CASS Sead nee. See gig SS See 46

aly a mug NUNN agi Cigc MAC GlegteT Rh TURN gly eae Mec age Loge ge cS tig Thay Nak SN al Stan 47

ey aia TN de ah eta mya tie Aa) SMI ge gue gt ARAN hs ge he yes ge 48

DNAS Sat us Nah GAN UNAM War Oman SLAM APM N ig SRT oN le ON 8 aC a 49

BO Oy VO leh ieee ak aya wl et ak ae NB, Gel ea We Sia!) hog liege cle 50

eR NS i ae a SUN CL AL SRM Eat ee oes NR NSW Mee K ak Tips og gue ah lh 51

FOU) ANN iy Se at EP UREN iN SLU) SQA Sg gs ae ag sy thal Sg! at gu hla Tonge GUN ts el 53

AN a RVR GE ihe en har MeiaUg Nigh gl Side Naa Sal ae aes ke gage Oke tas a dels ae Ob 55

Ly ta ee ate MCN oN Wag Vel Pini weal ig Hgiiiigiscgldk gs gROMpetumen AG INR | OTA Ok Wg 57

aah he a) OPS re nae eee eee a 58

Mee MERE gy Ce RM ENERO UI TMU CEREAL IAC ie Rav OR OR 58

PER OU OST oes TERI Di SRR SAA se SAE Uva NS Pay ee il a a De

ee EERE cael) Le. 4 ag MUN TERNS RI ASR en eGR Tbe ROO Con a ME RL Ge Ra a 61

i Net Rik Bil has Nigh call (SnMel i ik ak WE el opty: ge few twee 62

ee CUNT ni chs ag, WN ICON CR Mhas Wag GCN MoO MM CMR INH i QC Wis Neg. gia ean BMG Nal tag 64

Pera Toe yy GL Le CL ARREST ER A CE TU Aa Ce ea eA EAS RUD ES a 66

ROCHE dn eMC Cu A Gnu MAI MSM aR SUN OM UN ge og ti 67

COTO ONES ILS) 2 eS RAR ae HO TSR ncn NTU NARESH PARSON TEU Rey RUNS Tv Dae URN ME a 68

ORS Or aR Na ae OAS Cn hia Sit CORI as my CA XN URE ia VAN St lie ROC UR aS AGL Aik)

Menke: Pison in the New World iii

TU IN aa) ssh Oi eg a i iy Rid SCO Ld ne ml in ede pa

GHerisn WRINOlaA hil iiyt thes. olan tke | oie pitas ner RU ATE lye Baan 71

CAR Mpg ch Ce RRP Cee ea aN LE ec psec anne 2 gam cE aM see 0 Iain yh geese) i We Hb

CS GAVAXIF HORS PCIe zie a as Fie hE ag i Pi ay a meno hia Eg eae i a gi 74

Convexifrone Fascnenberg eo ijeed. £0i.oes ape Sur aaa apace a i ail 75

WASOSUGE) NETIC fr Gia a eh Dee ane a pe Ne ae ee a ne el 76

COGr ar: Terie isk: eg a Ee ise ae Mi ie vane to aryl MN cetip tia 77

lonaicorne Menine > un gria! | sce: 2 eR aes eid pall oem) aun ein ed ee 78

P UST POI ie INS RE UR Ty Bh Ertan aa an ea aie ee 80

rlosm: Sieh Lite BUR eh Pe ate eon alta ea Nae ARO seasinin 81

gureolaciaie Stream ery iy I PANT AEN eI 84

Vinceniki Menken Sure So re a ees ie tami i ed B4

Oaxaca Mienlee i: We POU BU RS TH IS SE A Tatty UF he th le 85

Gv Enos: Miniter Whee “IC i a i a ae ce a ps ir a 86

SOMAGTOOMa US PORK: aie ip eo a ah tg ALE Ue ein alg aA aia ED 88

PRE be RORLEY BOD ey LINED Ue GI AO ALN) RN RR AMHR ccc laae 90

is tae@ve: Sih: (ence ae Gi A aire er rial cae a A Ste ae ec an 90

New species of Pison from New Guinea ...........4.. fae Or aa aoe 90

st tL eirra: DW ie rye ay okie eg aT Rt Rt ladies ried aid 91

TUS Cig a cic): Maan eee ie gaat PTO ELUNE) Mae Usa lia hI Ati VneA MAMMA LoCrer wo HRM amy’ 92

Key bo spaciesual Pisomensis (ia) eine ie ln ag aa cle lel a OU hh at 93

Later tere ibe ils ai athe tba ee eer a i ER ar nN ans lal I aa 94

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INTRODUCTION

The spider predator genus Pison Jurine is cosmopolitan, but in the Western

Hemisphere it is essentially a neotropical taxon. All but one of the 44 species

treated here are restricted to that region. The only nearctic species, agile (Smith), is

not native: it was introduced into northeastern North America from the Orient during

World War II. Another species attributed to the North American fauna, laeve Smith,

appears to belong in the Indoaustralian area based on my examination of its type (see

Appendix, p. 90).

This study has been based on 1580 specimens of Pison from the Western

Hemisphere, of which 484 were chilense Spinola, a commonly collected endemic in

Chile and Argentina. When I began my review of the genus, only 15 neotropical

species were known, but I now recognize 43 species in the region, 29 of them new.

This swells the world total for Pison to somewhere around 200 described species

making it one of the larger genera of Sphecidae. More neotropical species await

detection, perhaps as many as 20, and of course the large Australian fauna has yet to

be subjected to a modern review. Worldwide, the genus may eventually contain

nearly 300 species.

This revision has to be considered as only a beginning framework for several

reasons. Some species are known only from one sex or even one specimen. I have

studied a number of specimens that represent additional undescribed species, but |

have not attempted to describe them because they are singletons and are in a species

group (the cressoni group) whose species are often difficult to distinguish even when a

lot of material is available. Furthermore, several species treated here may

ultimately prove to be complexes. Finally, in a few instances I have been able to

separate males, but their corresponding females seem identical. Clearly much more

collecting is needed before a definitive revision can be achieved. Such a revision will

also require rearing of nest material to positively associate sexes.

I have not attempted a comprehensive phylogenetic analysis of the New World

fauna of Pison. Such an analysis would have to be considered preliminary because the

very large Australian fauna (about 50 named species of Pison), which may have some

Gondwanian links with the South American fauna, is poorly known, and the rest of the

world species of the genus are not well documented morphologically. Lack of a

comprehensive world collection of Pison, as well as time constraints imposed on my

study, have precluded an in-depth world survey of the genus. I have analyzed a

number of characters, however, that are phylogenetically and morphologically

important in the genus, in the Trypoxylini, as well as in the Larrinae. I have

concluded that the Crabroninae should be incorporated with the Larrinae.

In the Appendix at the end of this paper I describe two new species of Pison from

New Guinea that are important from the standpoint of understanding the

morphological diversity in the genus. A key to the five species of the New World

genus Pisonopsis Fox, a taxon separated with difficulty from Pison, is also found in

the Appendix. :

ACKNOWLEDGMENTS

During the writing phase of this project I sought and received advice, opinions

and counsel from a number of people: Wojciech Pulawski, James Carpenter, E. Eric

Grissell, Maria Alma Solis, David Wahl, Richard Bohart and Karl Krombein. Their

thoughts and help are appreciated. Drafts of the manuscript were critically reviewed

by Wojciech Pulawski, James Carpenter, and Ray Gagne and their comments enabled

me to considerably improve the ms. Duane Hope demonstrated how to mount

genitalia on gold wires with polyvinyl acetate (PVA) for SEM photomicrography.

Brian Kahn, Susann Braden, and Walter Brown came to my assistance whenever I ran

into problems on the scanning electron microscope. Leslie Brothers provided a

provincial map of Ecuador. Bonnie Farmer corrected bird names cited in biology

sections. Vera Lee assisted in preparing camera ready copy. To all these people I

extend my sincere thanks.

Z Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

Colin Vardy of the British Museum (Natural History), Max Fischer of the

Naturhistorisches Museum, Vienna, the late Eberhard K6nigsmann of the Humboldt

Museum, Berlin, and J. O. Hising of the Martin Luther Universitat, Halle, generously

lent type material. Other research material was borrowed from the following

collections. Abbreviations used in the species treatments are listed first.

AEI - American Entomological Institute, Gainesville (Henry Townes).

AMNH - American Museum of Natural History, New York (Marjorie Favreau).

BERLIN - Zoologischen Museum der Humboldt Universitat, Berlin (Eberhard

KOnigsmann, F. Koch).

BMNH - British Museum (Natural History), London (Colin Vardy, Michael Day).

BPBM - Bernice P. Bishop Museum, Honolulu (Gordon Nishida).

CAS - California Academy of Sciences, San Francisco (Wojciech Pulawski).

CMP - Carnegie Museum, Pittsburg (George Wallace).

CNC - Canadian National Collection, Ottawa (Lubomir Masner).

COOPER - Martin Cooper Collection, Lyme Regis, Dorset, England

CSDA - California State Department of Agriculture, Sacramento (Marius Wasbauer).

CU - Cornell University, Ithaca (George C. Eickwort).

DIAS - Braulio F. de Souza Dias Collection, Fundacao IBGE, Brasilia.

FRITZ - Manfredo Fritz Collection, Salta, Argentina.

FSDA - Florida State Department of Agriculture, Gainesville (Lionel Stange).

GEMBLOUX - Faculte des Sciences agronomiques de l'Etat, Gembloux (Jean

Leclercq).

GENEVA - Muséum d'Histoire Naturelle, Geneva (Claude Besuchet).

HALLE - Martin Luther Universitat, Halle, East Germany (J. O. Husing).

HEH - Henry Hespenheide Collection, Los Angeles.

IBGE - Fundacao IBGE, Brasilia (Braulio de Souza Dias).

KU - University of Kansas, Lawrence (George Byers, Robert Brooks, John Wenzel).

LECLERCQ - Jean Leclercq Collection, Liege, Belgium (Jean Leclerca).

LEIDEN - Rijksmuseum van Natuurlijke Historie, Leiden (Kees van Achterberg).

LILLO - Instituto Miguel Lillo, Tucuman (Abraham Willink, Lionel Stange).

MCZ - Museum of Comparative Zoology, Cambridge, Mass. (James Carpenter).

MLSU - Zoological Museum of the Moscow Lomonosov State University, Moscow

(Alexander V. Antropov).

OSU - Oregon State University, Corvallis (George Ferguson).

PARIS - Muséum National d'Histoire Naturelle, Paris (Simone Kelner-Pillault).

PMA - Provincial Museum of Alberta, Edmonton (Albert Finnamore).

RAW - Anthony Raw Collection, Brasilia.

UCD - Richard M. Bohart Museum, University of California, Davis (Robert Schuster).

UCM - Universidad Central de Venezuela, Maracay (F. Fernandez-Yépez).

USNM - National Museum of Natural History, Washington DC. (Arnold Menke).

USU - Bee Biology and Systematics Lab., USDA, Utah State University, Logan (Terry

Griswold).

VIENNA - Naturhistorisches Museum, Vienna. (Max Fischer)

ZMC - Universitetets Zoologiske Museum, Copenhagen (Ole Lomholdt).

METHODS AND TECHNIQUES

I have not provided complete label data for all species. For common, described

taxa I have given only the country, province, and locality followed by the

abbreviation of the owner of the material in parentheses. The number and sex of

specimens from any one locality is usually given only for new species; males are

noted however, for species in which that sex is poorly represented. Collectors names

are cited only for material of new species. Some of my new species may prove to

represent complexes of two or more species. In these cases I have not always

included all material studied in the paratype series. Instead atypical specimens are

listed as metatypes, i.e., non-type material.

Menke: Pison in the New World 3

Extensive use of scanning electron photomicrographs has been made to illustrate

various body parts. Genitalia and a few other structures were coated with gold

palladium, but most material was photographed uncoated, particularly clypei,

mandibles, and propodei. A Cambridge Stereoscan 100 set at 10 KV was used for

coated material. Uncoated material was scanned at 2 KV on the same machine using

a low KV anode, or occasionally on a Hitachi S-570. Uncoated specimens were held

in a special stub similar to that described by Stork and Claugher (1987). This stub

(fig. 1) permits leaving specimens on their pins or points during SEM work, and allows

photographs of holotypes and unique material to be made without damage. Uncoated

material was often soaked in an undiluted solution of the commercial cleaning agent,

409, and then run through water, alcohol, and chloroform. The last causes setae to

remain erect after drying. Sometimes cleaning in chloroform was enough. Debris

was removed from the body by using a small piece of Scotch Magic Transparent Tape

stuck to the end of an insect pin. Dust and other small particles adhere to the tape.

Male genitalia and sternum VIII were glued to the end of fine gold wire with

polyvinyl acetate (PVA) dissolved in a mixture of acetone and 99% isopropyl! alcohol.

The wires were then glued to standard SEM stubs with Elmer's Glue-All (fig. 2).

These structures were not cleared, but were often cleaned in an undiluted solution of

409 cleaning agent. The PVA was usually nearly dry on the wire by the time it was

applied to a specimen, so to facilitate adhesion of these small objects they were

soaked in 99% isopropyl alcohol immediately before touching the wire to them.

A Wild M-400 photographic microscope was used to take pictures of wings and

some propodei.

Figures 1-2. SEM mounting techniques. 1, stub drilled to accept insect pin to

permit photography of uncoated type specimens and other unique material. 2, male

genitalia and sternum VIII mounted on gold wires that are glued to standard stub.

DISTINCTIONS BETWEEN PISON AND PISONOPSIS

These two genera are separated by rather tenuous differences (see Bohart and

Menke, 1976:330). Pisonopsis has a notched mandible, but so do some Pison.

However, Wojciech Pulawski (in litt.) has discovered another mandibular feature that

ultimately may be the only character that will permit maintaining Pisonopsis as a

discrete genus. In Pison the flattened lateral face of the mandibular base is margined

posteriorly (or below depending on your viewpoint) by a rather sharp edge (condylar

ridge of Michener and Fraser, 1978) that extends from the lower condyle to the notch

(figs. 4-6), or to the midpoint of the much more common notchless mandible (fig. 3).

This condition occurs in all other trypoxylines and Larrinae that I have examined

4 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

except Pisonopsis. In Pisonopsis the lateral face of the mandible has a rounded ridge

that parallels the condylar ridge of the mandible delimiting a groove (figs. 7-8). This

secondary structure is best developed in clypeata Ashmead, areolata (Spinola), and

australis Fritz, and it appears to be an autapomorphy for Pisonopsis. Clearly the

mandible in the Sphecidae needs further study since it may offer defining features

not heretofore appreciated.

The graduli found on abdominal sterna III-IV of Pisonopsis do not occur in Pison,

but a practical problem of using them as a generic character is that they are often

barely visible due to telescoping of the abdominal segments, and they are especially

Figures 3-8. Left mandible in lateral view. 3-6, Pison species (arrow indicates

condylar ridge); 3 is arachniraptor, 4 is species from New Guinea, 5 is convexifrons, 6

is qnythos. 7-8, Pisonopsis species (arrow indicates secondary ridge); 7 is birkmanni,

8 is areolata.

Menke: Pison in the New World 5

weak in areolata from South America. The marginal cell of Pisonopsis is rounded

apically, rather than acuminate as in most Pison. But the cell is rounded also in

some Pison (eremnon for example).

The basal flagellomeres of the male antenna in Pisonopsis are asymmetrically

swollen in all species. The asymmetry of the first or first and second flagellomeres is

rotated 90 to 180° in relation to flagellomeres II or III-V or VI. Some species of Pison

have asymmetrical flagellomeres also, and it is not clear if the male antennal

structure in Pisonopsis can be used as a generic character. This needs more study.

Since no key is available for the five species of Pisonopsis, I have included one in

the Appendix (p. 93). This key was partly developed by R. M. Bohart but has been

revised by me and expanded to include the South America species.

GENUS PISON JURINE

Pison Jurine (in Spinola), 1808:255. Type species: Pison jurini Spinola, 1808, monotypic.

Pisonoides F. Smith, 1858:104. Type species: Pison obliteratum F. Smith, 1858,

monotypic. NEW SYNONYMY.

Krombeiniellum Richards, 1962:118. New name for Paraceramius Radoszkowski, 1887:

432. Type species: Paraceramius koreensis Radoszkowski, 1887, automatic.

NEW SYNONYMY.

Entomopison Menke, 1968a:1. Type species: Pison pilosum Smith, 1873, original

designation. NEW SYNONYMY.

The generic description in Bohart and Menke (1976:332) adequately characterizes

Pison, but one character was overlooked. A nearly universal feature of the genus is a

pit or transversely elongate depression anteromedially on the pronotum (figs. 16, 95).

This is usually obscured by the back of the head, but it is an important apomorphic

trend in Pison. I examined about 50 Old World species of Pison and only four were

found to lack a pronotal depression. These were obliteratum Smith, anthopus

(Brullé), multistrigatum Turner, and kohlii Bingham. In the New World a pit is absent

in the euryops and krombeini groups and a few species in the cressoni group.

Trypoxylon and Pisoxylon apparently always lack these depressions. Species of

Pisonopsis and Aulacophilus have either a pit or transverse depression. I have not

seen material of Aulacophilinus.

The discovery of two unusual species in New Guinea (see Appendix, p. 90)

complicates the traditional generic concept. These two taxa have an unusually

elongate claviform first abdominal segment (figs. 347-350), and at first glance look

like members of Trypoxylon. But both have three submarginal cells and two

recurrent veins in the forewing (figs. 345-346), and an anterodorsal pronotal pit.

Unlike nearly all Pison species, these two wasps have enlarged eye facets beneath the

eye notch (figs. 341-344), a universal feature of Pisoxylon and most members of

Trypoxylon s. s. Interestingly the facets are fairly uniform in size in most (all?)

species of the subgenus Trypargilum of Trypoxylon. The antennal sockets are

contiguous with the clypeus in one of the species of Pison from New Guinea, and

slightly above the frontoclypeal suture in the other. The sockets are far above the

clypeus in Pisoxylon and Trypoxylon except for some groups within the subgenus

Trypargilum. Pison lobiferum Arnold from Madagascar (Arnold, 1945) has three

submarginal cells and a clavate first abdominal segment but its wing venation is quite

different, and it apparently lacks a dorsolateral propodeal ridge. Thus a close

relationship to the two New Guinea wasps seems unlikely.

SUBGENERA

Four subgenera were recognized by Bohart and Menke (1976): Pison s.s.,

Pisonoides, Krombeiniellum and Entomopison. I now believe that subgenera are

unwarranted as discussed below.

6 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

Pisonoides was proposed (Smith, 1858) for Pison obliteratum, an Indonesian

species with two submarginal cells and a clavate first abdominal segment, but Turner

(1916) used it for all species of Pison with two submarginal cells although he was

aware that it was not a monophyletic subgenus. Two-celled wings have arisen

independently a number of times in unrelated species or groups of Pison, and I

(Menke, 1968a) restricted Pisonoides to three species with a clavate abdomen and two

submarginal cells (obliteratum, difficile Turner and icarioides Turner). lLomholdt

(1985:15) treated Pisonoides as a genus and further refined its definition by including

only species (unspecified) with a "constriction" between abdominal segments I and II.

Lomholdt (1985:4) listed Pisonoides exclusum Turner as the species he studied but

Turner (1916) described its first abdominal segment as not elongate. I have studied

obliteratum and difficile, and based on their overall morphology they have

convergently developed two-celled wings and an elongate gaster and are not closely

related. Thus there is no reason to maintain Pisonoides and I am synonymizing it

under Pison. Pison difficile is of interest in that it is one of the very few species of

the genus with a noncarinate hindcoxa (see p. 17).

Krombeiniellum has been used for species in which the eyes are densely

microsetose (fig. 9), and was originally used for taxa with only two submarginal cells.

I (Menke, 1968b) expanded the subgeneric concept to include species with setose eyes

and three submarginal cells. My review of the neotropical fauna of Pison has shown

that the three-celled species are not closely allied to the two-celled forms and that

there are two disparate groups of species with two-celled forewings (agile and

stangei groups). Furthermore, Pison delicatum, a new species with asetose eyes and

three submarginal cells, shares several features with the agile, stangei and krombeini

groups: a short, clavate antenna; nonspiny legs (fig. 72); a velvety covering of short

body setae; and thick tarsal claws (fig. 70). If Krombeiniellum is to be recognized as

a subgenus, it must be redefined on the basis of these latter features, some of which

are apomorphies. I am not certain, however, that this would result in a monophyletic

taxon and am simply synonymizing the subgenus with Pison.

Two neotropical species groups of Pison, the convexifrons and pilosum groups,

are unusual in that the mandible has a notch on the posterior surface (figs. 5-6). I

(Menke, 1968a) established the subgenus Entomopison for these species. This taxon

has two apomorphies: the notched mandible and the polished, asetose “bearing”

surface on the anterior margin of the pronotum (figs. 288, 295). The mandibular

notch was thought to be unique to these New World species until Wojciech Pulawski

returned from New Guinea in 1987 with a Pison (CAS), probably undescribed, that has

a small notch. Actually the notch in this wasp is more like a “step” (fig. 4) that is

delimited basad by a tooth-like angle. A similar but smaller step is present on the

mandible of the Australasian and western Pacific island inhabitant, Pison ignavum

Turner. Neither of these species have a broad pronotal bearing surface. In the two

New World species groups this pronotal structure takes on two very different forms: a

mediodorsal expansion of the polished rim that is not free (fig. 288, convexifrons

group), and a broad lamella that is free from the rest of the notum (fig. 295, pilosum

group). The two types represent convergent specializations in my opinion. Similar

lamelliform structures are found in two other neotropical groups of Pison, the fritzi

and krombeini groups, but in these the surface is setose, at least in part, and not

always highly polished (fig. 45). Based on the foregoing, the convexifrons and pilosum

groups probably represent two unrelated monophyletic lineages and I have decided to

synonymize Entomopison under Pison.

Finally, Antropov and Pulawski (in prep.) are describing a fossil species of Pison

from Baltic amber that has a notched mandible similar to that found in the

convexifrons and pilosum groups. I have examined the unique specimen, and it

appears to have an elongate anterodorsal pronotal pit that is bordered anterad by an

overhanging lamelliform carina. It is impossible to determine if there is a polished

bearing surface as in the convexifrons group, but there is no free lamella like that

found in the pilosum group. The labrum appears quadrangular and is not emarginate,

and the occipital carina is incomplete ventrad, ending well before the hypostomal

carina. In these respects the fossil is like the convexifrons group. The pitted

Menke: Pison in the New World 7

episternal sulcus is straight, ending ventrad before curving forward. The metapleural

flange looks like it is somewhat lamelliform, and almost as broad as a hindocellus.

Hindtarsomeres II-IV each have a plantula. The propodeal dorsum has ridges that

extend from the base in an oblique fashion, and a median, longitudinal carina. The

fossil differs from present day species of the convexifrons and pilosum groups in at

least two ways: the propodeal side is not delimited above by a crenulate ridge or line

of foveae, and the second recurrent vein terminates on the third submarginal cell.

Clearly the fossil represents a third lineage with a notched mandible.

SPECIES GROUPS

The 44 New World species segregate into 12 groups, all but one of which (the

conforme group) are defined by clear apomorphic features. Most of these groups

appear to be endemic to the New World, but the agile group is an Old World

assemblage with an introduced species in North America. The chilense group may

have relatives in Australia.

Some of the characters useful in defining groups include: form of the mandible

(position of teeth, presence of notch); shape and details of clypeal lobe; shape of

labrum; form of the occipital carina; whether or not the frons is strongly swollen;

proportions of head (eye length vs. distance between eye notches); presence or

absence of dense setae on eye; presence or absence of an anterodorsal lamella or boss

on the pronotum; presence or absence of a crenulate ridge at the top of the propodeal

side; sculpture of propleuron; form of the episternal sulcus; wing venation; and form

of male sternum VIII and genitalia.

The presence on the eye of short, dense setae is an apomorphic trait common in

several groups (fig. 9), but some species of Pison have a few short setae widely

scattered over the eye (cressoni group for example). In general a naked eye is the

common condition.

Figure 9. Left eye of Pison plaumanni showing eye setation.

8 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

The form of male sternum VIII is fairly uniform within a species group. It may

be simply truncate or rounded apically (fig. 17), the plesiomorphic state, or the apex

may be emarginate (fig. 238). The emargination is often bounded laterally by a blunt

prong, or pseudosting (fig. 254).

The makeup of the male genitalia varies among the 12 species groups, some being

robust, some laterally compressed, and some more or less dorsoventrally compressed.

The gonostyle is a simple structure in the conforme, fritzi, eremnon, delicatum,

agile, and euryops groups. In the pilosum group the gonostyle is often a simplified,

large plate-like affair. The gonostyle has a ventral lobe in the krombeini, cressoni,

and convexifrons groups, and there are two ventral lobes in the chilense group. I

regard the presence of one or two gonostylar lobes as an apomorphy. The volsellar

lobes are very large in the conforme, fritzi, eremnon, krombeini, and cressoni groups,

but are quite small in the agile, stangei, convexifrons and pilosum groups. In the

eremnon group, the volsellar lobes are represented by two long, narrow structures.

Descriptions and discussions of apomorphies are provided for most of the species

groups. Descriptions have not been given for the two monotypic groups (eremnon and

delicatum), but their species descriptions and discussions are adequate.

SPECIES CHARACTERS

Described here are features used in species descriptions and group diagnoses that

need clarification or merit discussion. Morphological terms are from Bohart and

Menke (1976) except that for surface sculpture I have used the terminology developed

by Harris (1979). I have used the terms "“carina" and “ridge” more or less

interchangeably. A lamelliform carina is one that is elevated into a thin blade-like

structure.

Structures that have not been studied carefully for use in species discrimination

are mouthparts, proportions and details of leg segments especially the tarsi, and

details of wing venation such as proportions of cells and cell veinlets and the form of

individual veins and cells. Apparent differences in the form of the propodeum

(length, angle between dorsum and hindface, etc.) have been noted in some species

but these are difficult to quantify and have usually been ignored.

Labrum: Sometimes the labrum is short, broad and is described as “transverse”.

Most often the labrum is "quadrangular", i.e., more or less truncate apically with

perpendicular sides. It is deeply emarginate in the euryops and pilosum groups with

the result that usually only two finger-like lobes are visible beneath the clypeal

margin.

Clypeus: The clypeus is one of the more important structures for species

discrimination. The free edge often has an impunctate, asetose rim or "lip", and

usually projects as a median lobe whose shape varies from triangular to truncate or

rounded. The lobe may bear several "teeth". When viewed from below, the free edge

of the clypeus is often clearly "thickened" or "double-edged". In the latter instance

the thickening is margined behind by a carina. Often the thickest part is the section

between the median lobe and the lateral end point of the clypeal margin. The clypeal

surface is often covered by metallic silver setae that are appressed and obscure the

surface sculpture. Mixed with these short setae are longer setae that are directed

downward, and in some cases they are quite bristly and form a “brush” that surpasses

the edge of the clypeus (fig. 6).

Frons: The common appearance is a gently convex surface, but in the

convexifrons group the frons is strongly swollen toward the antennal sockets. In the

pilosum group some species have a small, impunctate polished depression or “dimple”

on the midline above the sockets. This feature is large in gnythos, new species.

Antenna: The antenna varies from moniliform with elongate flagellomeres (two

or more times as long as wide) to clavate where the flagellomeres are progressively

Menke: Pison in the New World 2

shorter and broader toward the apex. Measurements of length and width of

flagellomeres were made with an ocular micrometer at a magnification of 50x.

Width measurements were always taken at the apex of an article, and the microsetae

on the surface were not included. The male antenna sometimes has tyli or other

modifications on some of the flagellomeres. When present, tyli occur on the surface

that is on the inside of the antennal curl, and great care must be used in examining

the flagellum for the presence of these and other specialized structures. Good light

and high magnification (50X) are essential, and often the antenna has to be relaxed

and uncurled in order to see the surface bearing these structures. A “simple” antenna

is one in which the flagellum has no visible modifications.

Head measurements: The "upper interocular distance", or UID, is the shortest

distance between the eyes at the vertex - it is always measured behind the ocellar

triangle. The “lower interocular distance", or LID, is the shortest distance between

the eyes adjacent to the clypeus (or near level of antennal sockets in some

extralimital species). Eye length is measured vertically and is compared with the

distance between the eye notches on the inner orbits (fig. 10). The “ocellocular

distance", or OOD, is the measurement made between the inner orbit of the eye and

the hindocellus. This is compared to the “hindocellus diameter", or HOD. All of

these measurements were made with an ocular micrometer at a magnification of 50X

to maximize accuracy.

Figure 10. Face of Pison eu, female, showing how to measure eye length and

distance between eye notches.

10 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

Occipital carina: This is usually “incomplete”, i.e, it does not form a complete

circle. The lower ends of the carina either terminate just before the midline of the

head, or just before meeting the hypostomal carina, or sometimes actually meet the

hypostomal carina.

Pronotal pit and lamella: Because the anterior portion of the pronotum is usually

hidden under the rear of the head, its structure has not been studied, at least in the

Sphecidae. But in Pison I discovered two types of features that merit the attention

of anyone working on the family. One character is the development in some taxa of

the narrow polished anterior rim into either a broad boss or a broad, free lamella.

The second is the presence of a round pit or transversely lengthened depression

behind the anterior margin of the pronotum. Both features can only be viewed

adequately when the head is tilted forward.

The normally narrow polished rim of the pronotum is occasionally broadened

dorsally into a polished boss (convexifrons group, fig. 288) that is suggestive of a

bearing surface for the back of the head (this may not be its true function). The

more usual modification however, is the extension of the polished rim upward and

backward as a broad, free lamella. In the pilosum group it extends laterad to about

the level of the pronotal lobe (figs. 294-295), and the area directly beneath it is

smooth and polished. Sometimes this lamella is present laterally but narrows toward

midline where it is paralleled posteriorly by a lamelliform, setose carina that borders

the anterodorsal pronotal depression (fig. 45, fritzi n. sp.). The lamella may be

asetose and polished (pilosum and fritzi groups) or minutely setose (krombeini group).

The “anterodorsal pit of the pronotum” varies from a small round pit whose

diameter is less than that of a hindocellus (fig. 95), to a “transversely elongate

depression" (fig. 67) that may or may not be bordered anteriorly by a sharp edge or

carina, the latter sometimes lamelliform, dull, and setose (delicatum, fritzi, n. spp.).

When lamelliform this carina can be confused with the anterodorsal lamella described

above, but the latter is usually polished and often asetose. When the pit is

transformed into an elongate depression its surface is impunctate and shiny. When I

refer to its "length" in descriptions and discussions, I mean its transverse length, i.e.,

perpendicular to the longitudinal axis of the body.

Pronotal collar: This is a fairly simple structure in most species, but it may be

thick or thin and comparison to the width of the metanotum is usually made.

Occasionally the “humerus”, or lateral corner of the collar, is angular, the angle being

part of a transversely oriented ridge (fig. 99). In some species the collar has a

median prominence (fig. 288), or the anterior face may be flattened at the middle.

Scutal flange: The lateral edge of the scutum adjacent to the tegula is reflexed

upward forming a flange of variable width.

Lateral ridge of propodeum: Some groups of Pison have a ridge that extends ina

gradual arc from the spiracle to the propodeal socket (fig. 27). Since it is situated on

the interface between the propodeal dorsum and side, it delimits the top of the

latter. Usually associated with the dorsal side of this ridge are cross—carinae of

variable length that are perpendicular to it. This usually makes the ridge appear

crenulate. Sometimes the cross-carinae are reduced to very short teeth along the

dorsal side of the ridge. In some species, the ridge is replaced by a row of large

foveae as in the pilosum group. In a few species the ridge (larsoni n. sp.) or row of

foveae (pilosum Smith) vary in development from present to non-existant.

Forewing: The marginal cell is typically acuminate distally. This is enhanced by

the slight extension of vein R; beyond the end of the cell. In eremnon Menke the RS

vein curves forward at the cell apex where it meets R) perpendicularly (fig. 53). A

somewhat intermediate condition occurs in chilense (fig. 253). Although three

submarginal cells is the norm in Pison, with the second petiolate, two-celled wings

are characteristic of the agile and stangei groups (fig. 12), and are also found in a few

species of other groups (fig. 11). This reduction in submarginal cells occurs two

Menke: Pison in the New World ll

ways. Loss of the outer veinlet (Ir-m) of the second submarginal cell results in the

two-celled condition seen in some species of the cressoni and euryops groups. In

these the remaining veinlet is usually angulate (fig. 11). Diminution (figs. 87-88) and

eventual loss of the second submarginal cell is another way of achieving a two-celled

wing (agile and stangei groups). Here the resulting veinlet between the remaining

submarginal cells is more or less straight (fig. 12). The shape of submarginal cell III

varies at the group level. The inner and outer veinlets (Ir-m, 2r-m) may meet on the

marginal cell or be well separated there. In rare instances the two veinlets join

before reaching the marginal cell resulting in a petiolate third submarginal cell. The

termination point of the recurrent veins (Im-cu and 2m-cu) varies considerably in

Neotropical Pison, sometimes even within a species. The point of divergence of the

forewing media in relation to crossvein cu-a is useful in some species. Typically the

media diverges after cu-a, but occasionally it is before cu-a as in abothrum and

brasilium.

Figures 11-12. Right forewing. ll, Pison styphopteron n. sp. (holotype). 12,

Pison plaumanni Menke.

12 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

Legs: The hindcoxa has one or two longitudinal carinae dorsally. The inner one

is always present and sometimes it is partially lamelliform. Apparent differences

between species in the degree and shape of the lamelliform part of this carina seem

unreliable, however. An outer carina is present in the convexifrons and pilosum

groups, but in most Pison it is usually present only on the apical one half, or less. It

is absent in a few species. The male foretrochanter is usually simple, but it has a

sharp spine in some species of the pilosum group (figs. 292-293). A plantula occurs at

the ventral apex of tarsomeres II-IV in some species (figs. 13-14, fritzi and pilosum

groups, chilense, eremnon), but usually only tarsomere IV has one, and they appear to

be absent in the krombeini and stangei groups and in a few other species. Usually the

legs have some conspicuous spines at the apex of the tarsomeres, but in the

krombeini, agile, stangei and delicatum groups the legs are essentially unarmed (fig.

72). The claw is unusual in these latter forms also. Instead of gradually tapering to

the apex, the claw is of fairly uniform width until just before the apex (fig. 70). The

claws in males of the pilosum group are asymmetrical on the foreleg, and sometimes

on the middle leg. In this group the anterior claw is deformed and lamelliform (fig.

14).

Figures 13-14. Details of male foretarsus of Pison pilosum. 13, ventral view of

tarsomeres III-IV showing plantulae. 14, ventral view of tarsomeres III-V showing

deformed claw.

Menke: Pison in the New World }3

Gaster: This is the definitive abdomen and the segments are numbered I-VI or

VII depending on sex. Segment I is somewhat elongate in a few species and I have

expressed this by measuring the length of tergum I along its midlength in dorsal view,

and comparing that to the apical width. This is somewhat imprecise because the

midlength can vary depending on how the segment is oriented. The first two or three

terga are sometimes rather strongly convex with result that in lateral profile there is

a strong constriction between terga I and II (pilosum group, fig. 301). In a few species

of the cressoni group, especially in the female, tergum II has a median hump (figs.

162, 251). More commonly tergum II may have a pair of low swellings, one on each

side of the midline, as in the pilosum group, eremnon, and others. The apical margin

of tergum I is sometimes double-edged (euryops and pilosum groups, fig. 335), or it

may be depressed subapically forming an apical band (eremnon), or the edge may be

reflexed as in the krombeini group. Some species have tergal fasciae of setae

(chilense group for example) or yellow bands (cressoni group). Male tergum VII is

generally simple, but it is laterally compressed in some species of the convexifrons

group, truncate apically in the chilense group, and ends in two rounded, reflexed lobes

in eremnon. Tergum VI in eremnon is deeply V-notched apically (fig. 56). Although

the sterna are sometimes modified in the females (slight median hump on sternum II

in conforme, for example), specializations on the venter occur mainly in males and

are presumed to have a role in copulation. The male of eremnon, which has the most

sternal embellishments among New World Pison, has transverse flanges on sterna

II-III, setal depressions on IV-VI, and sterna VI-VII are emarginate apically (figs.

58-59). The male of delicatum has decumbent mats of setae on sterna III-V (fig. 68).

Sternum VII sometimes has depressions of various shapes as in the pilosum group (figs.

304-306). Sternum VIII may be truncate apically (fig. 108), rounded apically (fig. 18),

emarginate apically (fig. 69), or end in two strong prongs (fig. 254).

Male genitalia: These structures usually offer good specific differences in New

World Pison and I have used scanning electron photomicrographs to illustrate them.

The ventral side of the genitalia offers the most diagnostic features, particularly the

structure of the gonostyle. The volsella in New World Pison is usually represented by

a pair of lobes associated with the inner margin of the gonostyle. They are often

quite large as in the cressoni group (fig. 145), but in the pilosum group the volsellar

lobes are much reduced (fig. 308) and sometimes scarcely defined. See species group

discussions for further details.

CHARACTER ANALYSIS

Discussions of characters and their polarity in the subfamily Larrinae were given

by Bohart and Menke (1976) and Lomholdt (1985). Lomholdt presented a cladistic

analysis and a reclassification of the Larrinae. The following review of characters is

largely restricted to those features that are of significance in Pison and not polarized

by Bohart and Menke (1976:224). In my attempts at polarizing characters I have

studied examples of most genera in all subfamilies of the Sphecidae, but I have

examined as many species of each genus in the Trypoxylini as possible. I have also

reviewed Brothers’ (1975) important paper on aculeate phylogeny, as well as

Carpenter (1981) on the Vespidae. Determining polarity has sometimes been

problematical and developing a hypothesis of mandibular notch polarity has been

especially unsatisfying. What is really needed is a thorough morphological study of

the mandible throughout the Sphecidae similar to the one done on bees by Michener

and Fraser (1978) in which all grooves, ridges, setal rows, etc. are analyzed and

homologies developed.

The plesiomorphic state for each character is listed first.

Mandibular notch: |. absent. 2. present.

There can be scarcely any doubt that the notched mandible is an apomorphy

within the Sphecidae as a whole, but how to interpret it within the subfamily

Larrinae, where it is a common feature, is a problem. Lomholdt (1985:19, 21)

14 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

regarded a notchless mandible as the plesiomorphic condition in the Larrinae sensu

lato, within which he incorporated the Crabroninae (Oxybelini and Crabronini). He

stated that notchless mandibles occur in “a very few larrine genera”, but

unfortunately he did not list them. Bohart and Menke (1976:225) declined to polarize

the presence of a mandibular notch, but suggested that the notch could be the

plesiomorphic state. The notch is universal, or nearly so, in the majority of the 39

genera in the Larrinae sensu stricto (Crabroninae excluded), and is absent primarily in

genera that are ffairly specialized (Nitela, Auchenophorus, Aulacophilus,

Aulacophilinus, Pisoxylon, Trypoxylon, Sanaviron, Bohartella, Dalara and Paraliris).

Lomholdt (1985:19, 45) considered the simple mandible in Nitela and Auchenophorus

to be a reversal from the apomorphic notched state, and presumably that was his

interpretation of the presence of simple mandibles in the 30 or so genera of the

Crabronini.

Regardless of the polarization adopted, homoplasy is involved, and support for

either hypothesis can only be obtained by a thorough phylogenetic analysis of all

genera in the Larrinae and Crabroninae. In lieu of such an analysis I am adopting the

most parsimonious hypothesis of mandibular notch polarity, one that minimizes

reversals. Thus I regard a notchless mandible as the plesiomorphic state in Larrinae.

I also hypothesize that highly evolved genera like Nitela, Auchenophorus, Trypoxylon,

and Bohartella have retained a primitive mandible (instead of a reversal as espoused

by Lomholdt). Reversals may have occurred from the apomophic notched condition in

genera like Liris, Holotachysphex and Tachytes (Larrini) where a few species have

simple mandibles. Notchless mandibles also occur sporadically in a few species of

some other genera in the Larrinae s. s., and most species of Solierella (Miscophini)

have simple mandibles. Bohart and Menke (1976:292) and Lomholdt (1985:165)

believed the notchless mandible in Solierella to be a reversal, but it seems more

parsimonious to regard the genus as having retained the plesiomophic notchless

mandible in the majority of its species, especially since it is regarded as one of the

least specialized genera in the Miscophini by these three authors.

A mandibular notch is not unique to the Larrinae s.s. The condition occurs in

eight of the approximately 45 genera of the Crabroninae and also in three small

unrelated genera: Dinetus (Astatinae), Laphyragogus (Laphyragoginae), and

Xenosphex (Xenosphecinae). Several important apomorphies are shared by the

Larrinae and Crabroninae (single midtibial spur, simple volsella, and larvae with

preapical anus”), but Bohart and Menke (1976:30, 222) did not combine them into one

subfamily. Lomholdt (1985), however, has taken this step and I am in accord. The

mandibular notch is an additional derived feature of the Larrinae and Crabroninae

which, although not present in all genera, suggests that it is logical to place the

crabronine tribes Oxybelini and Crabronini within a restructured Larrinae. Four of

the five genera of Oxybelini have notched mandibles although the state is not

universal in all of them. Only four genera of Crabronini have notched mandibles and

they were regarded by Bohart and Menke (1976) as among the most primitive in the

tribe, a judgement that is probably erroneous due to the presence of various

apomorphic states, one, of course, being the mandibular notch. I hypothesize that

most genera in the Crabronini have retained the plesiomorphic notchless mandible

although a few reversals may have occurred.

The function of the notch is unknown although Lomholdt (1985:19) declared that

it and “the row of stiff bristles presumably serve to clean and protect the opposite

* Lomholdt (1985:21) regards the absence of larval papillae as an additional apomorphy, but

Evans (1964) treated this state as the plesiomorphic condition in Sphecidae. Lomholdt's idea

only requires reduction of the papilla along several different phyletic lines, but Evans‘

polarity requires reversals as discussed by him (Evans, 1964:232-233): reduction of the

antennal papilla and then redevelopment of long papillae "by at least two stocks of

Sphecidae". In bees Michener (1953) regarded the absence of papillae as an apomorphy. More

study of this character seems appropriate. Lomholdt also introduced a new apomorphy for the

Larrinae, the presence of a "genal tentorial arm", but he added that "the significance of

this character can be evaluated better after studying more sphecid genera". He also used a

carinate hindcoxa as a synapomorphy for the Larrinae but see my discussion on page 17.

Menke: Pison in the New World 15

mandible". How a mandibular notch can clean any part of the opposite mandible

escapes me, and it is more likely that the “bristles” are used in digging, i.e., part of

the psammophore. It is conceivable that the notch might be used in cleaning an

antenna or part of a leg, but no one has described such activity to my knowledge.

The notch is common in fossorial genera and it may have some function in nest

building, although the fact that some fossorial genera in the Crabronini have

notchless mandibles confounds this idea. The absence of a mandibular notch

sometimes coincides with a xylicolous or mud nest building lifestyle as in

Trypoxylon. Some Pison are ground nesters (chilense, several Australian species - see

Evans, 1981) and they have notchless mandibles. Members of the convexifrons and

pilosum groups of Pison are nonfossorial so far as known, and all of them have

notched mandibles. The small genus Holotachysphex is also perplexing; five of its six

species have notchless mandibles, but the one with notched mandibles and the only

species whose biology is known, turneri (Arnold), is a twig nester (Gess, 1978). In the

final analysis, there is no strong evidence for associating the mandibular notch with

fossorial or nonfossorial habits.

Labrum: |. quadrangular, free margin entire. 2. bilobed.

The labrum is a fairly simple structure in Pisonopsis, Aulacophilus and many

species of Pison, but the apex is variably emarginate in some Pison and Trypoxylon.

The extreme elaboration of the emargination results in a labrum that has two long,

finger-like lobes (Pisoxylon, some Pison, some Trypoxylon), an obvious apomorphy.

Anterior polished rim of pronotum: 1. narrow. 2. broadened.

Generally the anterior rim of the pronotum in the Sphecidae is narrowly polished,

presumably as a bearing surface for the back of the head. This is the plesiomorphic

condition. This rim is sometimes elaborated into a mediodorsal triangle as in Prionyx

(Sphecinae), or a broad dorsal boss as in some Pison. Sometimes the rim is broadened

over much of its length as in Ampulex (Ampulicinae), Polemistus and Arpactophilus

(Pemphredoninae), Psammaletes and Ammatomus (Gorytini), Paranysson and

especially Mesopalarus (Miscophini), or it may be broadened laterally as in Vechtia

(Crabronini). These various elaborations are obvious apomorphies.

Anterodorsal lamella of pronotum: |. absent. 2. present.

In some Pison the narrow polished rim is elaborated into a broad, free lamella, an

obvious apomorphy. Within the Trypoxylini this lamella is not unique to Pison. It

occurs in Aulacophilus and some Trypoxylon (rugifrons for example). In a cursory

survey of the other sphecid groups I have found a similar structure in the Sphecinae

(Trigonopsis, Podium), the Pemphredonini (Pemphredon, Stiqmus), the Scapheutini

(Scapheutes, Bohartella), the Crabronini (Entomocrabro, Quexua, Chimila) and the

Gorytini (Sagenista, Pseudoplisus oraclensis). A similar but possibly not homologous

lamelliform structure occurs in some Pison and elsewhere in the Sphecidae

(Neodasyproctus, Dasyproctus and Ectemnius of the Crabronini for example). The

anterodorsal pronotal pit or transverse depression is often delimited cephalad by a

carina which, in some taxa, is elaborated into a lamella that resembles the polished

anterodorsal lamella, but it is dull, setose and has at its base the normal, narrow

polished, pronotal rim. The presence of the latter suggests that this type of lamella

is not homologous with the polished lamelliform extension of the pronotal rim. In any

event, both forms of lamellae are obvious apomorphies.

Anterodorsal pit of pronotum: |. absent 2. present.

The anterior part of the pronotum of many sphecids has a pit, a pair of pits, a

series of pits, or a transverse depression that may be sulciform. These are believed

to be homologous but some of the linear depressions have a more anterior location

and may represent independent developments. Pronotal depressions are absent in

most Ampulicinae, Laphyragoginae, Nyssoninae, Philanthinae, Entomosericinae and

Xenosphecinae, and this is the common state in Pompilidae and many Vespidae s.l.

based on my cursory survey of those two groups. On this evidence a plain pronotum is

the plesiomorphic condition in Sphecidae (Carpenter and Cumming, 1985, regarded

16 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

similar pronotal pits or “foveae” as apomorphic in Eumeninae). I regard the single,

round, median pit seen in some Pison as the simplest apomorphic state. Often the pit

is transversely elongate and one or both margins may be cariniform, and this

represents further specialization. The channel-like or sulciform depression seen in

some sphecids (most Sphecinae, Astatinae, some Pemphredonini, Scapheutini, some

Crabronini, some Trypoxylini) may be a further extension of this trend, or else a

completely independent specialization. In genera having species with a pronotal

lamella, a polished linear depression is sometimes closely associated and it is likewise

possibly a completely independent development instead of a highly derived pit. Its

location is more forward than the normal location of the pit.

I have made a cursory examination of the pronotum throughout the Sphecidae

and present my findings below. In the Trypoxylini I have examined many species of

each genus, but only one or two species of each genus in other sphecid groups. My

survey of the genera in the Pemphredoninae and Nyssoninae has been even less

thorough. Thus the following discussion is preliminary, but it does give some

indication of the variation of a new character complex.

In the Trypoxylini only Pisonopsis, Aulacophilus, and Pison have depressions (I

have not seen material of Aulacophilinus Lomholdt, 1980). I examined a cross section

of species in Trypoxylon (including Trypargilum) and the pronotum always lacks a pit

or transverse depression. The same is true of Pisoxylon. Although most species of

Pison have a pronotal pit or transverse depression, the plesiomorphic pitless condition

is occasionally seen (obliteratum, xanthopus, multistrigatum and kohlii in the Old

World; species of the euryops, krombeini and pilosum groups in the New World).

In the Miscophini the pronotum is pitless in Nitela, Auchenophorus, Mesopalarus,

some Solierella, some Plenoculus, some Miscophus, some Saliostethus and in

Nanomiscophus. Lyroda is unique in having a linear series of large, deep pits that

closely parallel the polished rim of the pronotum - an autapomorphy for the genus.

The forward location of these pits may mean that they are not a homologue of the

pronotal pit. I suspect they are not. Other genera in the Miscophini have a

transversely elongate pronotal pit (Plenoculus, Sphodrotes, Larrisson, Paranysson,

Aha and some Solierella and Miscophus). Some Saliostethus have two pits. I have not

examined Sericophorus or some of the endemic African genera. In the Oxybelini and

Crabronini the pronotum may be pitless, or more commonly have a transverse

depression that is sometimes vaguely defined. In some genera there are two discrete

pits (Enoplolindenius, Foxita, Arnoldita, Chimiloides, Lestica, for example).

Anacrabro and Encopognathus have a deep, transverse groove. In the Bothynostethini

the pronotum is plain in all species of Bothynostethus examined, and Willinikiella

argentina (Schrottky) only has a vague pronotal depression. I have not examined

Sanaviron (Vardy, 1987). Most genera in the Larrini seem to have a pair of elongate

pits or a single elongate pit that may be weakly impressed. Some species of

Tachysphex have a small circular pit. In Palarus (Palarini) the pronotum has a pair of

transversely elongate grooves that are narrowly separated at the midline by a bridge.

In some species there is a transverse groove behind these two depressions that is

similar to the condition found in some nyssonine wasps (see below).

In the Pemphredoninae the pronotum displays a variety of states. The Psenini

that were examined have a transverse impression at the base of the collar, but it is

often vague, and it is not clear if this is homologous with the pronotal pit. In the

Pemphredonini the pronotum may be pitless (Diodontus, Polemistus), have a

transverse channel that may be deep (Pemphredon, Stigmus) or shallow

(C arinostigmus), or a pair of shallow pits (Spilomena, Ammoplanops, Pulverro). The

front margin of the pronotum is deeply emarginate at the middle in the last two

genera, a feature that may have taxonomic importance.

Although most nyssonine taxa seem to have a plain pronotum, Heliocausus

(including Tiguipa and Acanthocausus - Heliocausini) has a transverse depression at

the base of the collar. Similar depressions occur in the gorytin genera Sphecius,

Handlirschia and Kohlia; the depression is broadly oval and deep in the last genus.

Stizus and Stizoides (Stizini) and some genera in the Bembecini (Hemidula, Stictia,

Rubrica, Zyzzyx, and Trichostictia) have a deep pit also, and a thin, sulciform

depression often extends laterad from the area. The nyssonine "pit", by virtue of its

Menke: Pison in the New World 17

location at the base of the collar, may not be homologous with the pronotal

depressions found in other sphecid subfamilies.

Although a pitless pronotum is regarded as plesiomorphic here, it seems likely to

me that reversals have occurred in some sphecid genera.

Lateral ridge of propodeum: 1. absent. 2. present.

This ridge is a common feature in many genera of the Oxybelini, Crabronini, and

Trypoxylini but is infrequent elsewhere in the Larrinae, and the rest of the Sphecidae

for that matter. Within the Trypoxylini it is prevalent in Pison and Trypoxylon but

not universal in either. Because the Oxybelini and Crabronini are generally regarded

as among the most highly evolved larrines, and because the ridge is restricted

primarily to the Larrinae, I am considering the presence of this ridge to be an

apomorphy. I suspect that reversals may have occurred in Trypoxylon, Pison, and

perhaps in other genera, but further study is needed to be sure.

Hindcoxal carinae: 1. absent. 2. inner carina present. 3. inner and outer carinae

present.

Brothers (1975:524, 544) considers a noncarinate hindcoxal dorsum to be

plesiomorphic in the Aculeata, and in the Sphecidae. Within the Sphecidae a

noncarinate coxa is typical of some species of Ampulex (Ampulicinae), all genera in

the Sphecinae, a few genera in the Pemphredoninae, all genera in the Astatinae, the

Laphyragoginae, all Nyssoninae except Nyssonini, some Alyssonini and some Gorytini,

and all Philanthinae except Pseudoscolia and some Cerceris.

According to Lomholdt (1985:20) the hindcoxa is carinate "in most larrine genera”

and “a very similar structure is present in Dinetus" (Astatinae). But in fact the

hindcoxa is noncarinate in all genera in the Larrini (rare exceptions), some species of

Palarus (Palarini), some Trypoxylini (Pisoxylon, some species of the subgenus

Trypargilum, and rarely in Pison), the genus Belomicroides (Oxybelini), approximately

half of the genera in Crabronini, and some genera of Miscophini (some Lyroda,

apparently all Plenoculus, a few Miscophus, apparently all Sphodretes, many

Paranysson, and all Aha). Finally, three species of Dinetus available to me, pictus

(Fabricius), cereolus Morice, and psammophilus Kazenas, lack hindcoxal carinae.

When a carina is present it is typically found along the inner edge of the dorsum.

Sometimes it is lamelliform and may be extruded into a tooth near the coxal base. In

some sphecids, there are two carinae, an inner one and an outer one (Entomosericus,

Bohartella, Enchemicrum, Oxybelus, Encopognathus, Entomognathus, Lindenius, some

Palarus, some Larrisson, some Auchenophorus, some Pison, most Pisonopsis). The

outer one is often restricted to the apical one-half of the coxa. In rare cases, the

outer carina is the only one present (Belomicrus, Chimila, some Lestica for

example). The hindcoxal carinae vary in form and length in different sphecid groups,

and I suspect that these carinae have arisen independently a number of times; the

possibility of a few reversals also needs to be examined. Carpenter (1981) regarded

the hindcoxal carina as homoplasious in the Vespidae.

Lomholdt (1985:21) cites the carinate hindcoxa as one of five synapomorphies for

the subfamily Larrinae, adding that "no such carina occurs elsewhere in the sphecid

wasps”. But this in incorrect. My cursory survey of the Sphecidae indicates coxal

carinae occur in several other subfamilies. In the Ampulicinae apparently all

members of the Dolichurini have a coxal carina and so do some species of Ampulex

(Ampulicini). I have not surveyed the Pemphredoninae thoroughly but a carina is

present in Ammopsen, Stigmus, Ammoplanops, Pulverro, and some Pemphredon.

Apparently all genera in the Nyssonini and the gorytin genera Clitemnestra,

Ochleroptera, Olgia and Argogorytes (some) have coxal carinae. A short carina is

found in some species in the Alyssonini, and some species of the philanthine genera

Cerceris and Pseudoscolia.

In summary, a hindcoxal carina is a common feature in the Larrinae, but it is not

universally present nor is it unique to the group by any means. Thus, it cannot be a

synapomorphy for the Larrinae.

18 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

Tarsal claws: 1. symmetrical, gradually attentuate to apex. 2. asymmetrical or

suddenly attentuate near apex.

In Pison the claws of each leg are nearly always identical and typically they

narrow gradually toward the apex. In the pilosum group, however, the anterior claw

of the foreleg and sometimes the midleg is deformed in the male, an apomorphy.

Another apparent apomorphy is the peculiar thick claw found in some species where

attenuation to the apex occurs suddenly (delicatum, agile, stangei and krombeini

groups).

Male sternum VIII: 1. apex not emarginate. 2. apex with semicircular or V-shaped

emargination. 3. emargination bounded by a pair of blunt projections.

The emargination of sternum VIII results in the presence of a pseudosting that

may be elaborated upon by the development of small projecting lobes. I consider

these to be derived traits.

Gonostyle of male genitalia: 1. narrowly elongate distally. 2. with short distal lobe.

An elongate, setose gonostyle is the common condition in Larrinae, and most

Pison, but in the pilosum group the gonostyle is often represented by a large plate

that has at most a short apical setose lobe.

Gonostylar appendages: 1. none present. 2. one accessory lobe present ventrally.

3. two accessory lobes present ventrally.

A fairly simple gonostyle was considered by Bohart and Menke (1976) as the

plesiomorphic state. A few New World species of Pison have elaborate gonostyli

including one or two lobes ventrally.

Volsellar lobes: 1. large. 2. small

Reduction of the volsella in the Sphecidae is generally accepted as the

apomorphic condition (Bohart and Menke, 1976; Lomholdt, 1985). Within the Larrinae

the volsella is sometimes absent (Paranysson, Plenoculus, Solierella, Nitela - all in

Miscophini) or reduced to a small setose area or lobe on the inner basoventral area of

the gonostyle (Aha, Miscophus, Saliostethus, Saliostethoides, Miscophoides,

Namiscophus - all Miscophini). The majority of larrine genera have a well defined

volsella, however, and this is true for most species of New World Pison. But the

volsellar lobes are much reduced in the pilosum group. The elongate volsellar lobes

of the cressoni and eremnon groups may be elaborations, so that reversals can not be

ruled out.

BIOLOGY

Data are available for only 7 of the 44 species treated here and much of it is

fragmentary so that no generalizations can be made. Nothing is known for 7 of the

12 species groups recognized. Pison chilense excavates burrows in the ground and

cells are separated by mud partitions. The other species, agile, cressoni, aranevorax,

longicorne, pilosum and sphaerophallus, make nests in sheltered situations such as

bird nests, wasp nests, tents of lasiocampid moths, and the underside of leaves.

These non-groundnesters make mud cells that are sometimes arranged in rows,

sometimes randomly grouped. Spiders are provisioned. Chalcidoid parasitoids have

been reared from the nests of some species. See individual accounts of the above

species for further details.

10.

ll.

Menke: Pison in the New World 19

KEY TO SPECIES OF NEW WORLD PISON™

(unknown: males of abathes, duckei, erebus, eyvae,

martini, nosferatu, neotropicum, pentafasciatum,

and phthinylla, and females of lillo and oaxaca)

Mandible with: posterobasal, notch: (£iGS...3- +6) iccidiucsisseiensavsevaddugiawserniorsecwedseeeeces 2

Mandible entire, without posterobasal NOtCh (fig. 3) .......cccccsscccecsccccecccccecccccocs 19

Females. (six Gastral SQGMENES): ssdawacecshesascunscccssvicnunediacesoeonabes De alata apralal aubiiaae ati >

Males (seven gastral: segments): cutee utilise se iki lis imi ekiaehidevandadeasd oni beswenevens 10

Anterior margin of pronotum with broad, non-lamelliform, polished area that is

restricted:to middorsuni (figs 208) Skreet 4

Anterior margin of pronotum with broad, polished, free lamella that extends

laterad to level of pronotal lobe (Figs: 294-295) scicssiidicdvcccdecéscucesacesewebssndends 7

Flagellomeres VII-IX nearly three times as long as wide; clypeal lobe as in

fig. 285; propodeal hindface unridged (or largely so) .... longicorne Menke, p. 78

Flagellomeres VII-IX about as long as wide; clypeal lobe as in fig. 270; propodeal

hindlace with transverse TidOe 0k, i aealie hes aa inde Aaa aSon oe 5

Propodeal dorsum finely, transversely carinate or striatopunctate (fig. 268)

sh vin win Wn wd ed pow Salis a lad ale balck wear ald ila ya ra convexifrons Taschenberg, p. 75

Propodeal dorsum smooth except for median Carina (fig. 269) ........seccscssscscesceess 6

Mesopleural punctation sparser toward mesopleural sulcus (fig. 280); Amazon

basin and nerewerd. oe ee a re cooperi Menke, p. 77

Mesopleural punctation uniformly dense to mesopleural sulcus (fig. 279);

POTATO RESET A LGCTIE IVE: sivisicinvauionerninnventnsienann divas Uvoncat bernie ed wasbaueri Menke, p. 76

Gaster with obvious constriction between terga I-II in lateral profile (fig. 301),

tergum I strongly HUMPET .........csseeeeesees COL OSUH) CONMMDION Vissi escdsinanevsdivnescons 8

Gaster without constriction, terqum') Not humped o.ielbu WL ioe eka sd eedlgsadee 9

Antenna completely black .........ceeeeeeee pilosum Smith & vincenti Menke, p. 81, 84

Flagellomeres I-II pale beneath; se Brasil, Paraguay .... aureofaciale Strand, p. 84

Propodeal dorsum with median carina and lateral line of foveae; clypeal lobe

arcuate (fig. 324); frons with ocellar-sized polished depression (fig. 323)

PPLE ANTE RE Te Cre eT LeU eRe EDT REPO E nT Uy Fey et mr Tuts rete qnythos Menke, p. 86

Propodeal dorsum without median carina or lateral line of foveae (fig. 337);

clypeal lobe triangular (fig. 333); frons without ocellar-sized polished

Gepression above SOCKELS .......cscccecescccccesccceccscceeees sphaerophallus Menke, p. 88

Foretrochanter with sharp spine on underside (figs. 292-293) ......ssccesccssccceeccees 1]

F oretrachanter WIthOUt SRINe ON: UNGSTBIGE au iidnindssvenses cnwadh Hebaddeens Gemrwecddadeescces 14

Foretrochanter spine located near middle of segment (fig. 293); propodeal dorsum

without median longitudinal carina; propodeum without crenulate groove

laterally celine Gece ea hi GH bie sphaerophallus Menke, p. 88

Foretrochanter spine located at apical third of segment (fig. 292); propodeal

dorsum with median, longitudinal carina, at least basally; propodeum with

crenulate groove laterally ...........sccseesees (pilosa 'GOMiDlaK): sispissnsnndveneeice 12

* Head measurements should be made at a magnification of at least 50X.

2h.

a2,

Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

Sternum VII flat, without circular or oval depression at middle (fig. 303); anterior

claw of. midieq normal, not distorted ih o..ccsecccesceesccossescoseds pilosum Smith, p. 81

Sternum VII with circular or oval depression at middle (may be concealed under

S VI) (figs. 304-306); anterior claw of midleg distorted ............cccccccscececcveees 15

Sternum VIII narrow and depression on VII circular (fig. 304); s. Brasil, Paraguay

sn sac eating acai reach abc s naesarearite'g aiden ane Sten cEMB rae ee, Scare aureofaciale Strand, p. 84

Sternum VIII broad and depression on VII elongate (fig. 305); n. South America

as dislacc a aise Baa Ui cae teu Gia RV aaa SMS ECL RARE LN Tae Ui wanouL cues eetanwataules vincenti Menke, p. 84

... SLernuoi VIE reunded. apically: (Tigs..306, 570)... ccserate tare eu le RA US ss edie de 15

Sternum. Vill emargmate apically (Tig. 27 Sic, stisaensiesvaveusncessyhscaswdsesadeveedes souewence 16

Tergum VII compressed laterally, forming a median ridge; frons without

ocellus-sized polished spot between sockets and midocellus; UID more than

GOcialy LD. is. siccesd immed eer earthe enewteel nas Me Sea ee, Sets oaxaca Menke, p. 85

Tergum VII without median ridge, apical margin reflexed; frons with polished

ocellus-sized median spot between sockets and midocellus; UID less than one

Re EP 2s So oe i ee eee Eee) Lee Sas fe oe edek bees dada qnythos Menke, p. 86

Flagellomeres VIII-X almost twice as long as broad; propodeal hindface smooth or

PRSAR IN OO wavicruisena biiaih wre ataittabeain au euaimetiahe aug eeu naldosin'eau tun gw oeiies longicorne Menke, p. 78

Flagellomeres VIII-X about as long as broad, shorter and thicker than I-III;

propodeal hindface with strong transverse ridging (figs. 268-269) ..........sse00. 17

Propodeal dorsum cross-carinate (fig. 268) ......... convexifrons Taschenberg, p. 75

Propadeal Gersiim. largely smoot Cig? 269)? oii Ae aes ah oe ai easie sees 18

Gonostyle of genitalia with long, curving spine ventrally (fig. 282); Bolivia, Brasil

POPOL 8 Gee ICA AD ONAN IC A rico s.sinis inessicascnnienwna rider mtaiinnes cooperi Menke, p. 77

Gonostyle with short ventral spine (figs. 276-277); northern Argentina

wvinth dhicde lias SARS ANE RU AL. Ualbig gt Bia W Nisin gse eek wseeawa iE cod Soles Seda wasbaueri Menke, p. 76

Eye covered densely with short setae (fig. 9) .......csccccccsscccccccccccccccccccccsssscscoens 20

Eye bare or with only few widely scattered Setae ..........ccccssccccccccccccccsscsscceeees 26

Propodeal side without dorsal carina or ridge; forewing with two or three

CSTR a to ere se busy isin pels Waa ediaaig ea vane ea de nau Deri g basen acatagbas iaweosoceniebeeebcuhavaess 21

Propodeal side delimited dorsad by carina or crenulate ridge that extends from

petiole socket area to spiracle (carina may be obscured by setae); forewing

with two submarginal Cells ...........ccececeeescees (GEENGO) QrOUP) waderdivecdvoesiiceene 25

Forewing with two submarginal cells, the second not petiolate; tergum I simple

apically, not reflexed nor with subapical transverse depression; northeastern

Narth: AMerica. iimnieavec dics (ACME POPOUD): iuswnsrsdsserscandewsodes agile (Smith), p. 38

Forewing with three submarginal cells, the second petiolate; tergum I with

reflexed apical rim (with subapical transverse depression); Neotropical

stbbepcd a orceceaudlo tra manta cadences spate CMe aes Crome ih GFOUP) scicstacdeuneuenes dwveaceiaiadesevseesaseceZ

Propodeal dorsum and hindface densely covered with appressed silver or gold

setae that obscure shiny surface; sternum I with appressed silver setae; disk of

scutum shiny, parapsidal and admedian lines sulciform

A ARS EE oh RUG eG eas cA BA Nahe whe ns CON krombeini Menke, p. 45

Propodeal dorsum with dense appressed setae at base, but apex and hindface

sparsely setose, not obscuring dull surface; sternum I with erect pale setae

(not silver); scutum dull, parapsidal and admedian lines not sulciform

Spleen a eth seme Was lite tet ante ah anne Wn Sceieh tae ea a ile aI oa ae neotropicum Menke, p. 46

Z's

26%

27.

Bye

33.

Menke: Pison in the New World Zi

Ocellocular distance 1.38 to 1.65X hindocellus diameter; propodeal dorsum

smooth, punctate; eastern and southeastern Brasil ...... plaumanni Menke, p. 40

Ocellocular distance 1.08 to 1.31X hindocellus diameter; propodeal dorsum

VERIBUIG: sas Fetradekvcbl pier aecaniauanlarnt cates TORR athe oie rettad Meade. 24

Upper interocular distance .76 to .80X the distance between eye notches, the

latier shallow (iGe./ 8). ows bac cenatisen ean edends uibadeeniees Cease abathes Menke, p. 43.

Upper interocular distance .59 to .66X the distance between eye

nobehes: eye NOLChes Geem (1G. 9). was decpavcuadheenoucn saceiild vous Ri Dectesited faa anctmtereak as 25

Pronotum rounded laterally, humerus at most with faint ridge (fig. 94); propodeal

dorsum mostly ridged or striatopunctate; occipital carina complete ventrad;

BOlIViay ATO UiNig. cistc esr ccse tua leetiocet cay cecmaaa kl tute ncenen tates stangei Menke, p. 41

Pronotum angulate laterally, humerus with obvious ridge (fig. 98); propodeal

dorsum mostly punctate (ridging restricted to base and short cross-carinulae

associated with median carina); occipital carina interrupted ventrad; Panama

EO YCOMUPAIMEG SED co clacscngs cams tanncae danse naweunsccaneanasnaes CeCe duckei Menke, p. 42

Propodeal side delimited dorsad by carina or crenulate ridge that extends from

petiole socket area toward spiracle (figS. 27, 46) .......cccccccsccccccecccccccecccecccecs 27

Propadeal side without Corsal:@arinia Gr PAGE ie, vissesss ccavansassuneecebassdseneteensanadees 39

Eye length 10-20% less than distance between eye notches (fig. 10); propodeal

dorsum without median longitudinal carina, surface smooth, punctate; labrum

ending in two fingerlike lobes ..............ceeee. COVEN OS OU cdt ca sicae engine's co « 28

Eye length equal to or greater than distance between eye notches (fig. 15) (if 10%

less then propodeal dorsum with median carina); propodeal dorsum with or

without median longitudinal carina; labrum arcuate, or truncate or shallowly

SMISMIM ALE BOLO AL Wi is) vag uk is eign umeteilieste sautatecan dee daguaretien ects named ncldes hakow eee eke: 34

Femaies, Gastar With G visible tera: vcsisveiactanee lias acy eterna eunccsceasreabiee 29

Males; caster with 7 Wigiile tera aii. i. alee cell cuit ate buna veageleues ss bine da hee se censamakae dase 31

Forewing with two submarginal cells, outer veinlet of submarginal cell I angled

(fig. 11); clypeal lobe quadrangular, lower lip roundly triangular (figs. 124-125)

diet nighie nani ac sraea ant aie ned SAA AMT REL Tae ERR oan ee styphopteron Menke, p. 50

Forewing with three submarginal cells; ClyPEUS VALiOUS .........cceccceescccesccccscceees 30

Propodeal hindface with coarse, transverse ridges from top to bottom; upper

interocular distance essentially equal to lower interocular distance

PEEL ERE PEEL MEM Ee Py NIM Bc OM MEAT My MMR DN Lay gM Asif hy euryops Menke, p. 48

Propodeal hindface smooth above, ridges present only next to petiole socket;

upper interocular distance at most .8X as long as lower interocular distance

4.049 GMAW MOMS Com Ris RAD SAME Le ran gine s <kARA Te CAPO C ks Kien hunted Taree eink Ebvo een ds eu Menke, p. 47

Forewing with two submarginal cells, outer veinlet of submarginal cell I angled

(fig. 11); flagellum without linear tyli or asetose, shiny areas

an d\0 94h e paied bn ind We $MM GaN CARE cae Uae Bacaimiatals cadets Maes aunt oo styphopteron Menke, p. 50

Forewing with three submarginal cells (if only two then outer veinlet of I is

straight); flagellomeres with asetose polished areas or linear tyli present on

Dre Rt Hci se sinus as vat ccaepeah nevenmnh Sens puck cata hen asa mamas tip tes ates tak vyiints Ms E IN ete tl un ges eOliig 32

Propodeal hindface smooth above, with transverse ridges only near petiole socket

_tinig cd oacp bh gdiheg sate cx teased amas cae hd aa da To Pataca 0 st ade a ice nace eu Menke, p. 47

Propodeal hindface with coarse transverse ridges from top to bottom ............- 33

F lagellomeres V-VI asymmetrically swollen in profile (fig. 126)

soeERWAULevaaheds sUaeeaseaeadeeaad DUAEAWEN Ue OER AALT CLs Ed Radda ered ea ke EEG et ak euryops Menke, p. 48

Flagellomeres V=Vi HOt SWOLIGM sccecccvdasccdscocsaccaserctvsedbuscsedesens lillo Menke, p. 49

34.

35.

36.

42.

Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

Occipital carina a complete circle, separated from hypostomal carina by at least

an ocellus diameter; pronotum with broad, transverse lamella anterodorsally

(Hg AS eae PRG 2. CGI) echo aiveceees uc haushederts iesasdsadebust, eosee oes 35

Occipital carina incomplete ventrally, or contiguous with hypostomal carina;

pronotum with or without anterodorsal lamella ................sceccescceccesccecceeccees 36

Carinae of propodeal dorsum mostly transverse, giving way to punctures laterally

(fig. 47); anterior margin of pronotum with broad lamella laterally at level of

pronotal lobe Ghigi AS) sisiieresar ieee eiveaides cei dees ees Gavel leee fritzi Menke, p. 31

Carinae of propodeal dorsum oriented posterolaterally, extending nearly to

lateral ridge (fig. 49); anterior margin of pronotum without lamella at level of

BEONOUGP TORE ell uh US A a paseo iain nosferatu Menke, p. 32

Tarsomeres without conspicuous stout setae distally (fig. 71); episternal sulcus

ending ventrad without curving forward to anterior margin of mesopleuron;

propodeal dorsum with median longitudinal carina but smooth overall,

SDDEALIIG HRDUNCEALE Wliscicsdvteuscoveswoeberevenvuaesctedvedereees delicatum Menke, p. 35

Tarsomeres armed distally with conspicuous stout setae; episternal sulcus curving

forward ventrally, reaching anterior margin of mesopleuron; propodeal dorsum

punctate and/or covered by Carinae ...........eeee. (conforime Group) ........2....... ay

Disk of propodeal dorsum smooth, punctate, shiny, with median longitudinal

depression that contains carina (fig. 25) ........csecescesecesees conforme Smith, p. 25

Disk of propodeal dorsum obliquely carinate or striatopunctate, with or without

median longitudinal depression and carina (figs. 28-29) ..........ccecccccccseceeceeees 38

Propodeal side almost entirely smooth, punctate; Mexico?

SEEPS AVE a is iG SERIE URES RSIS GTA. OE RAL Or nn PE Une mALSoor OHNE eam uA doggonum Menke, p. 26

Propodeal side striatopunctate on dorsal half (fig. 26); Ecuador to Bolivia

ES OU Se RR Ui a SOS ON ee Pe CON De Salada A iii some larsoni Menke, p. 29

Tergum I sharply depressed subapically, forming a broad band; propodeal side

with a few large punctures scattered among dense, fine punctation (fig. 57);

HOdy Ta FP PONG Or MIGKE I leasiiecsin ceaeanelees eremnon Menke, p. 33

Tergum I without sharp subapical depression; punctures of propodeal side of

LIPLFOPI SL ee Oy ol 2 TTRETT OF TOSS Veiiiee va eeliea luis eliiceeea siecle secsels eleswessVelstdvicinecsee 40

Propodeal dorsum with many carinae, at least basally (figs. 29, 31, 263), or

OEP PACOOUNICURUG cee iiiorceccsder cbse ssseuensieeWausbagussUCeNbeeGucUDbnsceseUNescnreedsdsee¥ebsnyere 4]

Propodeal dorsum at most with median, longitudinal carina, surface smooth

eDN TRL aL 8 0 SOROS UNRE Se MSIE sya eae (GRO OGAL OWE VARs fecaasedgdavaesizeiceess. 44

Pronotal collar thin, knife-edged, closely appressed to and lower than scutum;

appressed silver setae confined to lower frons ........ (conforme group) ......... 42

Pronotal collar as thick as metanotum, rounded, and nearly as high as scutum;

head and thorax with extensive appressed silver setae .... (chilense group) .... 43

Ridges of propodeal dorsum obliquely oriented (fig. 29)

SLM Ot eerie Teas U Tate uM Uae buiw ec eacksh peas ee kaweaa eae some larsoni Menke, p. 29

Ridges of propodeal dorsum longitudinally oriented when present (fig. 31)

SRG Ni NMR NR aN ALA TL On RUAN Ne acl he lkardts eyvae Menke, p. 27

Most species in this group are separated reliably only by rather subtle differences in the

female clypeus and male genitalia. I have attempted to use other, more obvious differ-

ences in the key (color for example), realizing that they may not work for all material.

Comparison with descriptions and figures should be made after arriving at a name in the

key.

43.

44,

45.

50.

33;

Menke: Pison in the New World 23

Wing veins amber; erect setae of head, thorax and legs black; terga I-III with

bright silver fasciae; propodeal dorsum obliquely carinate/striatopunctate

cha des dig MeWiela Liiva g oR lr ee EE eek ee chilense Spinola, p. 71

Wing veins black; erect setae white; tergal fasciae dull; propodeal dorsum

longitudinally carinate, interspaces impunctate ................. sylphe Menke, p. 73

Forewing with two -submaernginal cells (fig..132) cal. ieiealab, ween, we .. 45

Forewing with three. submarginal celle (Tics PBV ii ci ccsncsescesinnwssnedameda chia tlie seers 47

Legs yellowish brown except coxae; first segment of gaster almost entirely

yellowish brown; female clypeus as in fig. 249; female tergum II strongly

humped (hig 2 Si) whiner ed al RS phthinylla Menke, p. 70

Legs largely or completely black; gastral segment I black except pale apical band

sometimes present on tergum; female clypeus as in figs. 160, 239, 241; female

ber Quin: TE ee Mme 325 HA A Is aa ld oh ae eed 46

Ocellocular distance .25 to .40X hindocellus diameter; female clypeus as in

FiGSe ZOO 2 a eal se Ete ee he aranevorax Menke, p. 68

Ocellocular distance .71X hindocellus diameter; female clypeus as in fig. 160

POE R TR MEER ON MMMM SCS APNE FAV MRE ENE cit OER a ys) 0 Ek LD Ae ABR WE erebus Menke, p. 57

Legs entirely yellowish brown (coxae and hindfemur above sometimes black) ... 48

Legs black, or, at most, femora and tibiae a mixture of black and pale brown .. 50

Forewing media diverging from M+Cu before crossvein cu-a; female clypeus

as in fig. 230; male clypeus as in fig. 231 ...........ccccceeees abothrum Menke, p. 67

Forewing media diverging from M+Cu after Cuma ............ccsccccccccsccccscescccccsceecs 49

Gastral segment I all black, no yellow apical band on tergum; tergum II strongly

convex in lateral profile (fig. 162); tergum I longer than wide

AE ee RIL AMO Ke RADE a TID Te BAM SUL AMRIT Na ig toe a PAWS a alld BIE CNH ORR martini Menke, p. 58

Gastral segment I completely or extensively yellowish brown, tergum with yellow

apical band; tergum II not unusually swollen; tergum I as long as wide

i isils aletiitasalaietird vie an vibe satay dusts daa Haw areuaeteed uae Kaiie MU e rar una ea ales maculipenne Smith, p. 59

Scutellum more sparsely punctate posteriorly where very fine punctures are

mixed with large punctures (fig. 142); female clypeal lobe reflexed (fig. 136)

PPE eo ROE MEMEO RE IIe NEA) MARR ORAM RIL EM DRE) CRIT NOMI 2omN cressoni Rohwer, p. 53

Scutellum wniformly punctate, punctures of One Size iii ee 51

Upper interocular distance .86-.91X lower interocular distance in female,

Fae OUR haa LBB AT NA adn ERS be Fae LU MEAL eaRERE ETON MAE LE TUR Tn ACME CE OE 52

Upper interocular distance .57-.79X lower interocular distance in female,

oF BK Ry aerate ala Weald aaa al Dept an a als ul a Ree ab dabe i ai lis, 53

Female clypeus as in fig. 177; male flagellum with linear tyli at least on articles

IV-VI; forewing media diverging from M+Cu before crossvein cu-a or

COT ee ie Cee ee i brasilium Menke, p. 61

Female clypeus as in fig. 221; male flagellum without tyli; forewing media

diverging after cu-a or interstitial with it ................e00. dementia Menke, p. 66

Propodeum largely covered by dense gold setae that obscure surface and

ocellocular distance in female usually .82-1.16X hindocellus diameter, and in

male .93-1.18X HOD; female clypeus as in figs. 149-151

si nnin ald ohhh ea HMA RG NOMEN ET IU OT NM ua eal aii aes chrysops Menke, p. 55

Propodeal surface largely or entirely visible, but if dense setae present on

dorsum then ocellocular distance in female no more than .69X hindocellus

diameter and in male no more than .85X HOD; female clypeus different ...... 54

24 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

54. Ocellocular distance 1.23xX HOD; terga I-V with broad, yellow apical bands

BAS eer cree y Sr Ane Sine Ree RL MO Sa Ig Sg OM pp a cele a pentafasciatum Menke, p. 58

- Ocellocular distance .33 to .85X HOD; yellow bands confined to terga I-III ...... 55

55. Female clypeus as in figs. 204, 206, 209; male flagellum with linear tyli on

Sruicies they OF VICE. 259) oo... seven catanescotenveiwaes es caainvees cameronii Kohl, p. 64

- Female clypeus as in figs. 166, 168-169, 186, 189, 192, 194; male flagellum

without tyli

56. Female clypeus with short median lobe whose margin is straight or slightly

sinuate (figs. 166, 168-169), clypeal rim not elevated above adjacent

punctation; male clypeus as in fig. 170; wing membrane sometimes yellowish

dues «alesibilaw ogc We lewis ce emialsio on ddaloa Go modo ea a ckeelceGiat. Cabdbce teat maculipenne Smith, p. 59

- Female clypeal margin above labrum with concavity bearing four angles or

teeth (figs. 186, 189, 194), clypeal rim elevated above adjacent punctation

(figs. 188, 196); male clypeus as in fig. 197; wing membrane never yellowish

Bs Bowe upbeat vou vases «rude | See JOE esl bau GEs Re Aub bate caOae Salo wt arachniraptor Menke, p. 62

Conforme Group

(Figs. 15-40)

Description:

Frons moderately swollen; labrum quadrangular; female mandible with weak

mesal tooth on inner margin; male antenna simple, flagellomeres elongate; clypeal

disk swollen, punctate; eye length slightly greater than or equal to distance between

eye notches (fig. 15); occipital carina ending just before joining apex of hypostomal

Carina; pronotum without anterodorsal lamella, but with anterodorsal pit that is

sometimes transversely elongate (fig. 16), this depression margined behind by carina,

surface behind carina shiny and usually with several ridges; scutal flange of variable

width; tegula- punctate, setose on basal half, impunctate, smooth beyond; propodeal

side sometimes delimited dorsad by crenulate ridge that extends from petiole socket

to spiracle (absent in eyvae, sometimes absent in larsoni); propleuron sparsely

punctate on disk; lower end of episternal sulcus curving forward to anterior margin of

mesopleuron (evanscent ventrad in eyvae and females of larsoni); male foretrochanter

without spine; outer carina of hindcoxa weak, reduced to distal ridge; hindtarsomere

IV with small plantula; male tarsal claws symmetrical; forewing media diverging

after crossvein cu-a; forewing marginal cell acuminate apically; three submarginal

cells, inner and outer veinlets of III usually broadly separated on marginal cell; hamuli

divided into two groups; gaster without yellow bands; tergum I single-edged apically;

male sternum VIII rounded or truncate apically, not emarginate; genitalia robust,

gonostyle slender and densely fringed with long setae apicoventrally, volsellar lobe

large, feebly setose, aedeagus unusually broad (dorsal view, figs. 36, 40), penis valves

fused over most of their dorsal length, each with apical horn (figs. 34, 38).

Included species:

Pison conforme, doggonum, eyvae, and larsoni.

Discussion:

The main features of this small group are the incomplete occipital carina, the

projecting, rounded female clypeal lobe, the coarse cross-ridging of the propodeal

hindface, and the non-emarginate male sternum VIII. The dorsolateral propodeal

ridge is not universal and can only be termed an apomorphic trend. The globular form

of the male genitalia is distinctive but males are unknown in two species. The

conforme group does not appear to have a single universal apomorphic character

state, and in fact it may be a paraphyletic group. When males of all species are

known, perhaps an apomorphy will be found in the genitalia.

Pison larsoni and eyvae are sister species, sharing the thin, closely appressed

pronotal collar, the absence (except in some larsoni) of a crenulate propodeal ridge,

Menke: Pison in the New World 25

and the evanescing episternal sulcus ventrad in the female. Pison conforme and

doggonum have a thicker collar and a well formed crenulate ridge on the propodeum.

Pison conforme, by virtue of its transverse pronotal pit and smooth propodeal dorsum,

has a somewhat isolated position. Pison doqgonum is fairly similar to larsoni in

sculpture and general habitus.

Pison conforme Smith

(Figs. 15-17, 19-20, 25, 33-36)

Pison conforme Smith, 1869:297. Holotype male: "Mexico" (BMNH).

Description, female (81 specimens):

Black; wings clear or slightly infumate. Frons with appressed silver setae on

either side of antennal socket, sometimes dense enough to obscure sculpture, clypeus

sometimes with similar setae laterally; erect setae on body pale; tergum I with

broadly interrupted apical silver fascia.

Free margin of labrum with weak indentation. Clypeal lobe as in fig. 19, free

margin not thickened or double-edged, surface smooth, polished. Frons dull,

shallowly punctate, punctures one to two diameters apart; eye length slightly greater

than distance between eye notches (fig. 15); UID 0.47-0.53X LID; OOD 0.33X HOD or

less, usually about 0.20X HOD; flagellomere I length slightly less than to slightly

more than 3X apical width, II about the same, III-X longer than broad.

Anterodorsal pit of pronotum transversely elongate, length equal to UID or

greater, carina behind pit sometimes lamelliform (fig. 16); collar thin, laterally with

linear impression that makes humeral angle prominent, ridgelike (fig. 16); scutum dull

or weakly shiny, punctate, punctures usually slightly larger than those of frons and

deeper, more sharply defined, separated by less than puncture diameter to two or

more diameters (punctures finer than on frons and several diameters apart in

specimens from Ecuador), interspaces imbricate (Harris, 1979), hind margin of scutum

with short ridges; scutellum shiny or somewhat dull, punctures somewhat smaller than

those of scutum; metanotum with micropunctures; disk of propodeal dorsum smooth,

shiny, with sparse, fine, setigerous punctures similar in size to those of metanotum,

usually with median longitudinal carina (sometimes reduced or absent in specimens

from Ecuador) that extends from base to about middle of dorsum (fig. 25), this carina

usually in depression (depression absent in specimens from Ecuador), base of dorsum

with short perpendicular carinae; propodeal hindface with coarse, well spaced,

cross-ridges that are continued onto posterolateral surface of dorsum, interspaces

shiny, smooth; propodeal side smooth, shiny, finely punctate, punctures slightly

larger, denser than those on propodeal dorsum; mesopleuron shiny, punctate,

punctures same size as those on scutum and less than diameter to several diameters

apart; metapleuron shiny, with pinprick punctures.

First recurrent vein usually ending near middle of second submarginal cell, rarely

interstitial between I-II, second recurrent vein ending on submarginal cell III or

interstitial between II-III.

Gaster shiny, terga I-II finely punctate, punctures several diameters apart,

sternum II swollen in profile, often with vague gibbosity on midline at apical third.

Length 6.5-8.5 mm.

Male (33 specimens):

Same as female except: clypeus usually covered by dense, appressed silver setae;

Clypeal lobe as in fig. 20; UID 0.55-0.62X LID; OOD usually 0.25X HOD; scutum

usually shiny; first recurrent vein usually ending near base of second submarginal cell,

and second recurrent interstitial between II-III, but variety of other arrangements

observed; apical margin of tergum VII arcuate; sternum II sometimes swollen,

sometimes with vague gibbosity; sterna II-VI each with one or more long, erect setae

posterolaterally; sternum VIII apex roundly truncate, slightly indented (fig. 17);

genitalia as in figs. 33-36, aedeagus slightly curved in lateral profile, apical horns not

sharply pointed (fig. 34); length 5.5-7.5 mm.

26 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

Discussion:

Pison conforme is commonly collected in Central America and northwestern

South America. Within the conforme group the smooth, punctate propodeal dorsum

with a median longitudinal carina usually set in a depression is distinctive, although

the carina is absent in some Ecuadorian material. The transversely elongate pronotal

pit, not easily seen unless the head is tipped forward, distinguishes conforme from

larsoni and doqgonum. Tergum VII is rounded apically in males of conforme, but in

larsoni it is truncate. The aedeagus also differs between these two species (compare

figs. 33-36 & 37-40).

I have studied Smith's type.

Range:

West central Mexico to Peru.

Material examined:

MEXICO, Jalisco: 9 mi. s Guadalajara (MCZ); Morelos: Cuernavaca (CU, USNM);

Vera Cruz: Jalapa (UCD, USNM); Rio Blanco (UCD); Rinconada (UCD); Fortin de las

Flores (FSDA); Orizaba (UCD); Cordoba (GENEVA); Chiapas: San Cristobal de las

Casas (CU, CNC); Bochil (UCD); Nachic (CU); L. Montebello N. P. (CNC).

GUATEMALA: Sta. Emilia, Pochuta (USNM); San Jeronimo B. V. P. (AMNH);

Helvetia, San Sebastian (MCZ); Las Sabanetas, Barbarena (MCZ); Moca, Guatalon

(MCZ). EL SALVADOR: 3 mi. w Quezaltepeque (UCD); Mt. San Salvador (UCD).

COSTA RICA: Turrialba (BERLIN); Santa Ana (MCZ); San José, San Antonio de

Escazu (COOPER); San Pedro de Montes de Oca (USNM); San Jose (USNM, BMNH);

Alajuela, Chomogo area (USNM); Monteverde (USU); Escazu (USU). COLOMBIA,

Valle: Pefias Blancas, 10 km w Cali (CSDA, FSDA); Candelaria, Finca San Luis

(CSDA); Pance CVC, 15 km w Cali (CSDA); Cali (BMNH); Boyaca: Muzo (MCZ);

Cauca: Chisquio El Tambo (OSU); San Andres de Pisimbala, 60 km e Popayan (BMNH);

Maaqdalena: N. Sierra Nevada de S., Marta, Rio Buritaca (BMNH); Vista Nieve, San

Lorenzo Mt. (USNM); Putumayo: Mocoa (BMNH); Cundinamarca: 3 km n Alban

(AMNH). ECUADOR, Pinchincha: Tinalandia near Sto. Domingo (PMA), 47 km s

Santo Domingo, Rio Palenque Sta. (CNC); Prov.?: Naranjapata (BPBM); Huigra (CU).

PERU, Madre de Dios: Avispas (MCZ).

Pison dogqgonum Menke, n. sp.

(Figs. 24, 27-28)

Description, holotype female:

Black; wings clear. Eye notches and lower frons with appressed silver setae that

obscure sculpture, clypeus more sparsely covered by appressed silver setae; rest of

head and thoracic vestiture pale or silvery; tergum I with silver fascia laterally,

adjacent area of tergum II with some appressed silver setae.

Free margin of labrum straight. Clypeal lobe as in fig. 24, free margin not

thickened or double-edged, surface smooth, not highly polished. Frons dull, shallowly

punctate, punctures one to two diameters apart; eye length slightly greater (1.03X)

than distance between eye notches; UID 0.47X LID; OOD 0.06 HOD; flagellomere I

length 3X apical width, II almost as long, IHI-X longer than broad.

Anterodorsal pit of pronotum oval, greatest width about equal to hindocellus

diameter; collar thin, humeral angle not ridgelike; scutum dull, densely punctate,

punctures same size as on frons but mostly less than diameter apart, hindmargin of

scutum with short ridges; scutellum weakly shining, punctation similar to scutum;

metanotum more finely punctate than scutellum; disk of propodeal dorsum

impunctate, weakly shiny, with median longitudinal carina and many fine, somewhat

irregular, oblique carinae that fade posterolaterally where surface becomes punctate

(figs. 27-28); propodeal hindface with several widely spaced transverse ridges that

become weaker dorsad; propodeal side shiny, smooth (except for a few ridges

adjacent to spiracle), with shallow, setigerous punctures that are one to two

Menke: Pison in the New World 27

diameters apart; mesopleuron smooth, shiny, punctate, punctures same size as those

on scutum but one to three diameters apart; metapleuron microridged above, smooth,

shiny, impunctate below.

First recurrent vein ending on submarginal cell I on left forewing, interstitial on

right forewing, second recurrent vein interstitial on left wing, ending on submarginal

cell III on right wing.

Gaster shiny, terga with very fine pinprick punctures; sternum II normal, not

swollen.

Length 7 mm.

Male:

Unknown.

Discussion:

Pison doggonum is rather enigmatic since only one specimen is known, but

several features separate it from larsoni, its most similar relative: free margin of

labrum straight (usually shallowly emarginate in larsoni), hindocellus nearly touching

eye (OOD 0.11-0.30X HOD in females of larsoni), pronotal pit as broad as hindocellus

(smaller than ocellus in larsoni), sculpture of propodeal dorsum fine (coarser in

larsoni, compare figs. 27-28, 29-30), and propodeal side nearly all punctate

(diagonally striatopunctate on dorsal half in larsoni). Pison doqggonum is a more finely

punctate wasp than larsoni and the latter species has a thinner pronotal collar.

Range:

Presumably Mexico. The type specimen is labelled simply, "Mejico, Mus. Drews."

Type:

Holotype female: "Mejico" (ZMC).

Etymology:

The name doggonum is a Latinization of the American slang expression, “dog

gone it", in this instance a reflection of my frustration at having only one specimen

with poor collecting data.

Pison eyvae Menke, n. sp.

(Figs. 23, 31-32)

Description, holotype female:

Black, wings clear. Body with pale erect vestiture (darkest on scutum) that is

longest on propodeal hindface, mesopleural venter and underside of coxae and

trochanters; lower frons with dense, appressed, tarnished silver setae that obscure

sculpture; no tergal fasciae.

Labrum quadrangular, margin entire. Clypeus with prominent truncate median

lobe with rounded corners, free edge not thickened (fig. 23). Frons dull, granular,

shallowly punctate, punctures one to two diameters apart. Eye length 1.04X distance

between eye notches; UID 0.48X LID; OOD 0.20X HOD. Flagellomeres I-III each 3X

as long as apical width, flagellomere X length 2X apical width.

Pronotum with transversely elongate anterodorsal pit that is margined behind by

sharp lamelliform carina whose length is less than UID; collar thin, closely appressed

to and lower than scutum. Scutal flange broad posterad, strongly reflexed. Scutum

and scutellum dull, densely punctate, most punctures less than diameter apart.

Metanotum weakly shiny, densely micropunctate. Propodeum shiny; dorsum densely,

finely punctate but with about dozen, short, coarse ridges at base, the two center

ridges extending posterad toward apex delimiting linear depression (fig. 32), dorsum

laterally with gently arcuate, linear depression that extends from base to near apex;

hindface impunctate, with about seven coarse cross-ridges that curve onto propodeal

side where they change to fine striatopunctation; propodeal side densely punctate on

28 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

lower half, finely obliquely striatopunctate above but without delimiting ridge.

Mesopleuron shiny, densely punctate, punctures one to two diameters apart.

Episternal sulcus evanescent as it curves forward ventrad, not clearly reaching

pleural margin. Metapleuron shiny, densely micropunctate.

First recurrent vein ending at middle of second submarginal cell, second

recurrent vein ending on third submarginal cell.

Gaster shiny, finely punctate, punctures several diameters apart on terga I-II.

Length 9 mm.

Variation in females (8 specimens):

Wings sometimes weakly infumate (Colombia). Appressed setae of frons

sometimes silver (Colombia). Margin of clypeal lobe sometimes arcuate. UID

0.453-0.49X LID (Colombia, Ecuador), or 0.52 LID (Bolivia); OOD 0.17-0.21X HOD

(Colombia, Ecuador), or 0.25-0.26X HOD (Bolivia). Carina behind anterodorsal pit of

pronotum very high, longer than UID (Bolivia). Propodeal dorsum varies: often only

one median, longitudinal carina present (fig. 31), sometimes all basal ridges may be

short, i.e, no long median longitudinal carina or carinae, and disk only punctate

(Bolivia), or many basal ridges may extend posterad changing to fine, arcuate

striatopunctation that covers most of disk with middle ridge strongest and straight

(Colombia, fig. 31), lateral arcuate linear depression sometimes absent; propodeal

side nearly entirely punctate or irregularly covered with oblique carinulae that are

continuations of coarse ridges of hindface (this latter condition occurs in specimens

with striatopunctate dorsum). First recurrent vein occasionally interstitial between

submarginal cells I-II or ending near base of second submarginal; second recurrent

vein sometimes interstitial between submarginals II-III. Length as short as 7.0 mm.

Male:

Unknown.

Discussion:

Pison eyvae differs from other members of the conforme group, except larsoni,

in having an episternal sulcus that becomes evanscent ventrally as it curves forward

toward the pleural margin. Pison eyvae is similar to larsoni in the narrow, closely

appressed pronotal collar, female clypeal shape, head measurements, and wing

venation, but the sculpture of the propodeal dorsum, though variable, is distinctive in

eyvae. The basal ridges are more or less longitudinally oriented, and when long, they

curve in a gentle arc toward the apex (figs. 31-32). The uppermost of the lateral,

anterad extensions of the coarse ridges on the hindface are similarly curved when

viewed from above, thus accentuating the affect. Pison eyvae usually lacks a

crenulate ridge at the top of the propodeal side (weakly formed in some specimens),

but the ridge is sometimes present in larsoni.

Range:

Known only from Colombia, Ecuador and Bolivia.

Types:

Holotype female: ECUADOR, Morona-Santiago: Macas, 1100 m, II-2-1982, M.

Cooper (COOPER).

Paratypes (8 females): COLOMBIA, Valle: Lago Calima, 3 mi. behind dam,

tropical wet forest, VII-16-75, R. C. Wilkerson (CSDA, FSDA). ECUADOR: same

data as type; Pichincha: Nambillo Valley nr. Mindo, 1450 m, VI-26-87, VII-2-87, M.

Cooper (COOPER). BOLIVIA, La Paz: Chulumani, 1700 m, IV-3-79, M. Cooper

(BMNH), Coroico, 1700 m, V-22-79, M. Cooper (BMNH).

Etymology:

The name eyvae is an arbitrary, euphonious combination of letters.

Menke: Pison in the New World 29

Pison larsoni Menke, n. sp.

(Figs. 18, 21-22, 26, 29-30, 37-40)

Description, holotype male:

Black; wings slightly infumate. Frons with appressed tarnished silver setae

between and lateral to antennal sockets: rest of head and body vestiture pale but

generally brownish; no tergal fasciae.

Free margin of labrum semicircularly emarginate. Clypeal lobe polished (fig.

22). Frons dull, punctate, punctures separated by one to two diameters, interspaces

appearing granulate; eye length equal to distance between eye notches; UID 0.59xX

LID; OOD 0.36X HOD; flagellomere I length about 2.5x apical width, remaining

flagellomeres progressively shorter, VIII-X about as long as broad, flagellomeres II-IV

each with a tiny, pale, oval sensory area on ventral side toward base, that on III the

largest.

Anterodorsal pit of pronotum transversely oval, greatest width shorter than

hindocellus diameter; collar thin, closely appressed to and lower than scutum,

humeral area not ridgelike or otherwise prominent. Scutum dull, punctures same size

as those on frons, but somewhat deeper and mostly one diameter apart, interspaces

granular; hindmargin of scutum with short ridges. Scutellum weakly shiny, more

sparsely punctate than scutum. Metanotum densely, finely punctate. Propodeal

dorsum shiny, with oblique ridges basally and cross-ridges along center line, these

changing to striatopunctation laterally; propodeal hindface with irregular, coarse

cross-ridges that become weaker dorsad; propodeal side shiny, obliquely

striatopunctate on dorsal half and near hindcoxa, punctate elsewhere (similar to fig.

26), side with irregular crenulate ridge dorsally. Mesopleuron shiny, smooth,

punctate, punctures larger than on scutum and about one diameter apart.

Metapleuron striatopunctate above, finely punctate below.

First recurrent vein of forewing ending on submarginal cell II, second recurrent

vein interstitial between submarginal cells II-III.

Gaster shiny, terga I-II finely punctate, punctures one to three diameters apart;

tergum VII somewhat drawn out apically, truncate; sterna II-III with one long, erect

seta posterolaterally, IV-VI each with two long, erect setae posterolaterally; sternum

VIII broad, angularly rounded apically (fig. 18); genitalia as in figs. 37-40, aedeagus

strongly arcuate in lateral profile (fig. 37), apical horns acuminate (fig. 38).

Length 7.5 mm.

Female (7 specimens):

Same as male except: free margin of labrum shallowly emarginate or truncate;

clypeusy.asy in, fig, 214 «ID. 0y50+0654* LID: OOD:-0:11-.30« HOD; flagellomere |

length 3X apical width, VII-IX longer than wide: scutal punctures mostly half

diameter apart; scutellar punctures slightly smaller than those of scutum, less than

diameter apart; propodeal dorsum usually with median, longitudinal carina that may

be short (figs. 29-30), oblique ridging usually longer, coarser (fig. 29); propodeal side

not clearly delimited by crenulate ridge in 5 Ecuador specimens from Nambillo,

striatopunctation sometimes covers all of side; episternal sulcus evanscent ventrally

as it curves forward; second recurrent vein usually interstitial between submarginal

cells II-III, but ending on II in one specimen and on III in another; sternum II with one

long, erect seta posterolaterally as in male, but succeeding sterna with more

numerous long setae that are more mesal; length 6.5-7.5 mm.

Discussion:

Pison larsoni has a thin pronotal collar that is lower than and closely appressed

to the scutum, a character state shared only with eyvae. The sculpture of the

propodeal dorsum, though variable in both species, separates the two: ridges, when

present, are primarily longitudinal in eyvae (figs. 31-32), but oblique or transverse

and always present in larsoni (at least in females, figs. 29-30). In five of the eight

specimens of larsoni the crenulate ridge delimiting the top of the propodeal side is

evanescent or absent, a constant characteristic of eyvae. Like eyvae, the female

episternal sulcus is evanscent ventrally in larsoni. The striatopunctate upper part of

30 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

the propodeal side of larsoni separates it from doqgonum. The ocellocular distance is

wider in larsoni than doggonum but too few specimens are available to determine the

reliability of this difference. The genitalia of larsoni differs from that of conforme

especially in the strongly arched aedeagus and sharp apical horns (compare figs.

37-38 & 33-34).

_ Range:

Known only from Ecuador, Peru and Bolivia.

Types:

Holotype male: BOLIVIA, La Paz: San Jacinto, 1700 m, XII-5/8-55, L. Pefia (KU).

Paratypes (7 females): ECUADOR, Morona-Santiago: Cord. de Cutucu, 6 km e

Macas, 1100 m, V-22-87, M. Cooper (COOPER); Pichincha: Nambillo Valley nr. Mindo,

1450 m, VI-27/28-87, VIII-15-87, M. Cooper (COOPER). PERU, Cuzco: Valle del Rio

Coshipata, Hacienda Santa Isabel, 1700 m, I-3-52, F. Woytkowski (LLILLO).

Etymology:

Pison larsoni is dedicated to Gary Larson whose "The Far Side" cartoons have

brought much laughter to me and many other entomologists.

Fritzi Group

(Figs. 41-52)

Description:

Frons swollen; eye asetose; male antenna with vague sensory areas on some

flagellomeres; female clypeal lobe trapezoidal or roundly truncate, male clypeal lobe

triangular; labrum quadrangular; female mandible with weak mesal tooth on inner

margin; occipital carina a complete circle, separated from hypostomal carina by

hindocellus diameter or slightly more; pronotum with transversely elongate, polished

depression that is margined anteriorly by broad, thin, setose, overhanging lamella;

pronotal collar thick but front face flattened broadly at middle; thorax elongate,

length in lateral profile about 2X height; scutal flange narrow, upturned; tegula

densely punctate on inner half, impunctate beyond; propodeal side delimited dorsad

by crenulate ridge that extends from petiole socket to spiracle; propodeal dorsum

finely cross-carinate and/or striatopunctate; propleuron punctate; lower end of

episternal sulcus curving forward to edge of mesopleuron; outer carina of hindcoxa

reduced to short, evanescent, distal ridge; male foretrochanter without spine;

plantulae present on tarsomeres II-IV or IIJ-IV; male tarsal claws symmetrical;

forewing media diverging after crossvein cu-a; forewing with three submarginal cells;

endpoint of recurrent veins highly variable: first to submarginal cell I or II or

interstitial, second to submarginal cell II or III or interstitial; gaster without yellow

bands; tergum I single-edged apically; male sternum VIII with semicircular

emargination apically; genitalia dorsoventrally flattened, penis valve without apical

horn, gonostyle densely covered by long, coarse setae ventrally, volsellar lobes feebly

setose.

Included species:

Pison fritzi and nosferatu.

Discussion:

This small group of rather dull, densely punctate wasps is characterized by the

complete occipital carina that is separated from the hypostomal carina, the dorsal

lamella associated with the elongate depression of the pronotum, the thick pronotal

collar, the lateral ridge of the propodeum, the apically emarginate male sternum VIII,

and the elongate body. The fritzi group is otherwise similar to the conforme group

but the male genitalia are robust in the latter and dorsoventrally flattened in the

former. Apomorphies are the pronotal lamella, transverse pronotal pit, the propodeal

ridge, and emarginate male sternum VIII. The greatly elongate, polished, flat,

Menke: Pison in the New World 31

pronotal depression of this group is clearly homologous with the anterodorsal pit of

smaller dimensions common in most species of Pison. One species, fritzi, has two

lamellae on the pronotum: the one associated with the anterodorsal pit, and one that

arises from the front margin of the notum. The last is lamelliform only laterally,

narrowing toward the midline of the pronotum.

Pison fritzi Menke, n. sp.

(Figs. 41, 43, 45-47, 50-52)

Description, holotype female:

Black; wings clear. Clypeus laterally and lower half of frons except for narrow

zone above each antennal socket, densely covered with appressed tarnished silver

setae that obscure sculpture, disk of clypeus sparsely covered with somewhat darker

setae; terga I-II with brownish fasciae apically; vestiture of thorax (except on collar

and propodeum posteriorly) and gaster brownish.

Free margin of labrum slightly arcuate; clypeal lobe roundly trapezoidal (similar

to fig. 41), its polished surface curving down to free margin, clypeal disk variably

densely punctate. Frons dull, densely but shallowly punctate, punctures separated by

about half a diameter; eye length greater than distance between eye notches (latter =

O.91X eye length); UID 0.67X LID; OOD 0.42X HOD; flagellomere I length 2.5xX

apical width, remaining flagellomeres progressively shorter.

Anterior margin of pronotum with broad, partly setose, polished lamella at level

of pronotal lobe that narrows dorsad where it is paralleled behind by setose lamella

that overhangs greatly elongate depression (similar to figs. 43, 45), length of

depression about three-fourths width of collar, depression delimited by carina

posteriorly; collar thicker than metanotum; scutum dull, with same dense punctation

as frons; scutellum weakly shiny, somewhat less densely punctate than scutum;

metanotum weakly shiny, more finely punctate than scutellum; propodeal dorsum

with many oblique carinae basally that blend with many parallel, arcuate,

cross-carinae along midline that change to dense, oblique striatopunctation laterally

(similar to fig. 46), no median longitudinal carina; propodeal hindface with several

coarse cross-ridges at petiole socket but striatopunctate above; propodeal side

densely punctate, almost striatopunctate, punctures separated by half diameter or

less, interspaces shiny; mesopleuron densely punctate, interspaces weakly shiny,

punctures separated by one diameter or less and larger than elsewhere on body;

metapleuron densely punctate, punctures smaller, shallower ventrad; plantulae

present on tarsomeres II-IV.

First recurrent vein interstitial between submarginal cells I-II, second recurrent

ending on submarginal cell III.

Tergum I more coarsely punctate than succeeding terga, punctures about same

size as those on scutellum and separated by less than puncture diameter; posterior

margin of tergum I broadly depressed (equal to width of fascia or breadth of

metanotum at midline).

Length 9 mm.

Variation in females (9 specimens):

Labrum sometimes truncate; UID 0.63-0.71X LID; OOD 0.44X HOD in Ecuador

specimen; propodeal dorsum with well formed median longitudinal carina in one

specimen from Nova Teutonia, Brasil; cross-ridges of propodeal hindface present

nearly to top in some specimens; apical depression of tergum I especially deep in

Ecuador specimen; end point of recurrent veins highly variable with many

combinations observed even in different wings of one specimen; length 7-9.5 mm.

Male (1 specimen):

As in female except: clypeal lobe triangular; distance between eye notches

O.94X eye length; UID 0.86 LID; OOD 0.62X HOD; flagellomere I length slightly

more than 2X apical width, III-V with vague sensory areas ventrally and slightly

52. Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

asymmetrically swollen; scutal punctures larger, deeper than those of frons; tergum

VII narrowly truncate; genitalia as in figs. 50-52; apicoventral surface of gonostyle

densely clothed with long, coarse, curved setae; penis valve slender, shorter than

gonostyle.

Discussion:

The most diagnostic feature of fritzi is the anterolateral lamella on the

pronotum (fig. 45). Pison nosferatu has only an anterodorsal setose lamella. The

orientation of the carinae on the propodeal dorsum also differs between the two

species. They are arcuately transverse in fritzi, but in nosferatu all the carinae are

oriented obliquely at about a 459 angle (compare figs. 46-47 & 48-49).

Range:

Pison fritzi is known from Ecuador, southern Brasil and northwestern Argentina.

Types:

Holotype female: BRASIL, Santa Catarina: Nova Teutonia, XII-1-1955, F.

Plaumann (USNM).

Paratypes (10 specimens): ECUADOR, Tungurahua: Pastaza Valley, 1600 m,

VIII-25-87, 9, M. Cooper (COOPER). BRASIL, Santa Catarina: Nova Teutonia,

various dates, 6 9%, F. Plaumann (MCZ, UCD); Blumenau, I-1885, ?%, Hetschko

(VIENNA); Guanabara: Represa do Rio Grande, VI-66, 9, M. Alvarenga (AE/J).

ARGENTINA, Salta: Rosario Lerma, IJ-85, ¢6, M. Fritz (FRITZ).

Etymology:

It is a pleasure to name this wasp after one of South America’s best wasp

collectors, Manfredo Fritz.

Pison nosferatu Menke, n. sp.

(Figs. 42, 44, 48-49)

Description, holotype female:

Black; wings slightly infumate. Frons with appressed tarnished silver setae

between eye and antennal socket and between the sockets; terga I-III with brownish

fasciae apically, IIJ-IV with weak fasciae; vestiture of body generally pale or

brownish.

Free margin of labrum weakly concave; clypeal lobe double-edged (similar to fig.

42); eye length only slightly greater than distance between eye notches (latter =

0.96X eye length); UID 0.63X LID; OOD 0.30K HOD; flagellomere I length almost 3X

apical width; frons dull, punctation as described for fritzi but somewhat coarser.

Anterior margin of pronotum without lamella laterally, but with setose lamella

that overhangs anterodorsal depression, the latter not delimited posteriorly by carina

(similar to fig. 44); collar about as thick as metanotum; scutum dull, punctation same

as fritzi but somewhat coarser; scutellum weakly shiny, punctation as in fritzi,

metanotum as described for fritzi but punctures somewhat larger; propodeal dorsum

covered by obliquely oriented carinae, those that cross midline forming rounded V's,

carinae changing to striatiopunctation posterolaterally, dorsum without clearly

defined median longitudinal carina (fig. 49); propodeal hindface with three widely

spaced, coarse cross-ridges, interspaces shiny, impunctate, upper third of hindface

punctate; propodeal side densely punctate changing to vertical striatopunctation

dorsad; mesopleuron shiny, densely punctate, punctures largest on body, separated by

one or two diameters and changing to weak horizontal striatopunctation beneath

scrobe; metapleuron striatopunctate above, obscurely punctate below; plantulae

present on tarsomeres III-IV.

Recurrent veins interstitial in left forewing, first ending on second submarginal

cell and second ending on third submarginal cell in right wing.

Gaster as described for fritzi except that depressed margin of tergum I narrower

Menke: Pison in the New World 55

than breadth of metanotum and less sharply defined.

Length 9.5 mm.

Variation in females (1 specimen):

OOD 0.37X HOD; propodeal hindface with cross-carinae to top, upper carinae

somewhat less coarse (fig. 48); length 9 mm.

Male:

Unknown.

Discussion:

Pison nosferatu is similar to fritzi, but the absence of a polished lamella

laterally on front edge of the pronotum is distinctive. The female clypeal lobe

differs between the two species also (compare figs. 41-42). The punctation of

nosferatu is somewhat coarser than that of fritzi, a difference difficult to appreciate

without material of both species. The carinae of the propodeal dorsum are more

oblique in nosferatu, being oriented at an angle of about 459 to the linear axis. In

fritzi the carinae are more transverse (compare figs. 46-47 & 48-49).

Range:

Known only from Venezuela.

Types:

Holotype female: VENEZUELA, Araqua: Rancho Grande, 1100 m, [-22/23-1978,

blacklight in cloud forest, J. B. Heppner (USNM). One paratype female with same

data (USNM).

Etymology:

The name nosferatu, a noun in apposition, is from the 1922 German movie

Nosferatu, based on the story of Dracula by Bram Stoker. In the movie, Nosferatu

was a pseudonym for the vampire Dracula.

Eremnon Group

Pison eremnon Menke

(Figs. 53-63)

Pison eremnon Menke, 1968a:5. Holotype female: Santarém, Brasil (CMP).

Description:

Body completely black, leading edge of fore and hindwings darkly infumate.

Head and thorax (including the coxae, trochanters and femora) with much long, erect

black setae. Tergum I and sterna with similar erect vestiture, but sparser on latter.

Pale pubescence present only posterolaterally on propodeum and short.

Head, thorax and terga I-II mostly bipunctate: large punctures scattered among

many finer punctures (fig. 57). Nearly all punctures of disk of scutum and scutellum

large (larger than any others on body), separated by one to six diameters, interspaces

smooth, shiny. Propodeal dorsum covered with small punctures (no large punctures)

that change to striatopunctation laterally. Terga III-V or VI densely, finely punctate,

dull or weakly shiny. Tergum III with few slightly larger punctures scattered over

surface. Sterna II-V shiny, irregularly punctate; punctures small, densest peripherally

but mixture of large and small sparse punctures present on disk of female. Male

sterna III-VI largely impunctate except laterally. Hindtibia with two to four short,

very stout setae on outer surface that are borne on slight elevations, as well as

several similar but more slender setae scattered over the surface in pits. Midtibia

sometimes with single, stout seta.

34 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

Head differs strongly between sexes: male with broad face and short clypeus in

comparsion to female (see figs. 54-55). Median lobe of clypeus with broad, smooth

shiny lip in both sexes that is not thickened. Female clypeus without brush. Eye

asetose. Male UID 0.72X LID, female UID 0.64X LID. OOD 3.2X HOD in male, 1.38X

HOD in female. Occipital carina broadly incomplete ventrally. Female mandible

with weak median tooth on the inner margin, but that of male edentate. Mandible

socket nearly closed off by extension of hypostoma. Anterodorsal pronotal pit

transversely elongate, deep, margined posterad by sharp lamelliform carina; width of

pit about 0.66X UID. Collar as high as scutum, thicker than metanotum, in female

with median elevation suggestive of Lyroda. Tegula impunctate on outer half.

Propodeum with median longitudinal carina dorsally but no ridge present between

spiracle and petiole socket. Propodeum posterolaterally with 5 or 6 Coarse,

transverse ridges above petiole socket that become progressively finer dorsad.

Propleuron mostly punctate, but with small, shiny impunctate area present laterally

on disk. Episternal sulcus curving forward ventrad attaining pleural margin.

Hindcoxa with fine outer carina that evanesces basad. Tarsomeres II-IV with

plantulae. Forewing marginal cell rounded distally (fig. 53). Submarginal cell I

longer, narrower than typical for New World members of genus, resulting in petiole of

submarginal cell II ending at midpoint of marginal cell (fig. 53). Recurrent veins

terminate on submarginal cells II and III, respectively. Forewing media diverges from

M+Cu basad of crossvein cu-a. WHamuli of hindwing not divided into two groups.

Tergum I elongate, apical width slightly less than three-fourths its length, and

hindmargin broadly depressed forming bandlike margin that is as broad as

metanotum. Male abdomen unusual: tergum VI deeply emarginate, VII ending with

pair of rounded lobes (fig. 56); sternum II strongly swollen in profile, it and

subsequent sterna bearing subapical transverse setose flanges that diminish

progressivly posterad, being represented on sterna IV-VI by setose depressions (figs.

58-59); sternum VIII ending in pair of blunt lobes (fig. 60). Genitalia laterally

compressed; gonobase elongate, with longitudinal crest dorsally (figs. 61-62);

gonostyle long, narrow, setose; and volsellar lobes elongate, setose apically, narrow in

ventral view (fig. 63); aedeagus broad in dorsal view (fig. 61) but otherwise a fairly

simple structure without defined head or ventral spine.

Length 12.5-16 mm.

Discussion:

Pison eremnon is the largest New World species of the genus, and it has many

unique attributes. The shiny, irregularly bipunctate, black body, the dark erect setae

of the head, thorax and legs, the wing venation and infumate leading edge of the

forewing, the nearly closed mandible socket, the stout setae of the hindfemur, the

elongate tergum I with its depressed bandlike hindmargin, and the male abdominal

features readily identify eremnon. The male is identified here for the first time and

is represented by a single specimen from Pachitea, Peru (BERLIN). The long volsellar

lobes are distinctive (fig. 63).

The distally rounded forewing marginal cell and elongate submarginal cell I (fig.

53), the abdominal structure especially in the male (figs. 58-59), the nearly closed

mandible socket, and head dimorphism (figs. 54-55) are unique attributes of eremnon

that isolate it from all other the neotropical groups in the genus. I regard the nearly

closed mandibular socket, the head dimorphism, and abdominal structures as

apomorphies of the eremnon group.

The nearly closed mandible socket is an interesting feature since it approaches

the condition typical of the Philanthinae, where a closed socket is one of the prime

characteristics that separate that subfamily from the Larrinae. The head of the male

of eremnon is reminiscent of some species of Philanthus because of the notched eyes,

the broad face and narrow clypeus.

Strong head dimorphism occurs in the Old World species regale Smith and strandi

Yasumatsu and their males have abdominal peculiarities suggestive of eremnon. But

in regale and strandi the propodeum has a crenulate ridge between the spiracle and

petiole socket, the mandible socket is broadly open, the mandible has a subbasal tooth

on the cutting edge, the wing venation is different, and the male genitalia are

Menke: Pison in the New World be.

dissimilar. A relationship between eremnon and these two Old World species, if any,

does not appear close.

Range:

Colombia to Peru, Guyana and northern Brasil.

Material examined (19 females, 1 male):

COLOMBIA, Putumayo: Mocoa, 600 m, various dates, M. Cooper (BMNH); Villa

Garzon, 8 km s Mocoa, various dates, M. Cooper (BMNH, COOPER); Amazonas: La

Chorerra, VIII-14/23-76, M. Cooper (BMNH); Leticia, VIII-21/29-74, M. Cooper

(BMNH); Meta: La Macarena, X-7/XI-29-76, M. Cooper (BMNH); Caqueta: Yuruyaco,

73 km sw Florencia, II-5-79, M. Cooper (BMNH). ECUADOR, Morona-Santiago:

Taisha, 500 m, I-21-82, M. Cooper (COOPER). PERU, Pasco: Puerto Bermudez, 500

m, XI-8-84, M. Cooper (COOPER); Huanuco: Rio Pachitea, Tessman, (BERI_IN).

GUYANA: Wanaina, N.W. District, III-31, J. Meyers (BMNH).

Delicatum Group

Pison delicatum Menke, n. sp.

(Figs. 64-76)

Description, holotype male:

Black, mandible yellowish brown, inner surface of foretibia brown; wings weakly

stained but forewing with clear area near third submarginal cell. Body without long

erect setae, and mostly covered by fairly dense, pale, short setae that impart a

velvety appearance; clypeus and lower half of frons with dense, appressed silver setae

that obscure sculpture; terga I-III with broadly interrupted silver fasciae.

Labrum short, broad, free margin entire, slightly arcuate; clypeal free edge

thickened, without prominent median lobe, but with very short, broad lobe defined by

polished part of thickened edge, this lobe with weak, rounded, median tooth (similar

to fig. 66); frons moderately swollen, dull, contiguously micropunctate, appearing

granular; eye asetose, eye length 0.95X distance between eye notches; LID 0.83X

UID; OOD 1.33X HOD; antenna clavate, flagellomere I length slightly more than 2X

apical width (10:4), V-X broader than long; occipital carina incomplete ventrally,

ending just before hypostomal carina.

Pronotum with transversely elongate, polished anterodorsal depression whose

length is equal to about half width of collar, this depression delimited anteriorly by

high lamella (similar to fig. 67); collar slightly thicker than metanotum, noncarinate,

but with shallow, anteromedian depression that narrows top of collar at midline;

thorax shiny; scutal flange narrow, upturned; scutum densely micropunctate,

punctures separated by about puncture diameter; tegula completely densely,

micropunctate; scutellum margined anteriorly by narrow sulcus that is vaguely

foveolate, punctures of scutellum sparser than on scutum; metanotum impunctate;

propodeal dorsum with median longitudinal carina in shallow depression, surrounding

surface smooth, sparsely covered with setose micropunctures; propodeal hindface

with coarse cross-ridges that become weaker toward top; propodeal side with

crenulate ridge between spiracle and petiole socket, side smooth, densely but very

shallowly punctate, punctures about two diameters apart, lower end of propodeal

spiracle surrounded by chain of small foveolae; propleuron finely punctate; episternal

sulcus ending ventrally, not curving forward to anterior margin of mesopleuron;

punctation of mesopleuron similar to scutum; mesopleural sulcus paralleled anteriorly

by row of foveolae; metapleuron more sparsely, finely punctate than mesopleuron;

metapleural flange not lamelliform.

Forewing media diverging after crossvein cu-a; forewing with three submarginal

cells, inner and outer veinlets of III narrowly separated on marginal cell, recurrent

vein I ending on submarginal I, recurrent vein II ending on submarginal cell II.

Legs smooth, devoid of conspicuous spiniform setae at apices of tarsomeres:

36 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

tarsomere IV with tiny plantula; tarsal claw thick to just before apex where it

terminates in a sharp, somewhat hooked point; midcoxa margined anterad by ridge

associated with precoxal sulcus; dorsum of hindcoxa with strong inner carina, but

outer carina reduced to short evanescent distal ridge.

Distal edge of tergum I simple, surface of tergum I weakly shining, coriarius (see

Harris, 1979) and obscurely punctate, punctures about a diameter apart; tergum II

shiny, smooth, punctate; distal margin of tergum VII with broad, obtuse emargination;

sterna III-V with discal mats of decumbent, long, white setae (as in figs. 68-69);

sternum VIII with broad, apical V-notch (as in fig. 69); genitalia as in figures 73-76,

aedeagus broad in dorsal view (fig. 76) and with unusual head (figs. 73-75) that has a

nearly circumferential lamella ventrally that is joined at midventer by another

lamella; gonostyle with usual long setae but subapically with area of dense, very short

setae (fig. 74); volsellar lobes elongate, feebly setose.

Length 5 mm.

Variation in males (20 specimens):

Mandible varies from yellow to nearly black. Mid and hindtibiae sometimes

brownish within, sometimes legs completely black. Eye length sometimes as short as

0.90X distance between eye notches. LID sometimes only 0.76X UID. OOD

1.28-1.46X HOD. Ridges of propodeal hindface sometimes evanescent laterally and

dorsad, occasionally hindface has only two or three ridges just above petiole socket

with remainder smooth, punctate. Forewing media sometimes interstitial with cu-a.

First recurrent vein sometimes interstitial between submarginal cells I-II in one or

both wings, second recurrent rarely ending on third submarginal. Tergum I sometimes

smooth, shiny, and clearly punctate. Length to 7 mm.

Female (22 specimens):

As in male except: edge of clypeus sometimes yellowish brown; vestiture

sometimes dull silver; mandible with median cleft on inner margin, but no tooth;

labrum arcuate; clypeus without prominent median lobe, free edge thickened at

middle and produced into arcuate lower lip (figs. 64-65); eye length 0.92-0.97X

distance between eye notches; LID 0.74-0.84X UID; OOD 1.24-1.53X HOD;

flagellomere I length 2.5-3X apical width; punctation of scutellum about as dense as

on scutum, metanotum with similar but sparser punctures; cross-ridges of propodeal

hindface as variable as in male; veinlets of submarginal cell III sometimes meeting on

marginal cell; first recurrent vein ending on first submarginal cell in 50% of the

specimens, interstitial in about 50%, rarely both ending on second submarginal cell;

length 6-8 mm.

Discussion:

Pison delicatum, a commonly collected species, has a distinctive velvety

appearance due to a general covering of short setae on the body and an absence of

erect setae. This look is enhanced by the dense micropunctures distributed generally

over the body. Distinctive features of the species are the straight episternal sulcus,

the ridge at the top of the propodeal side, the very elongate depression of the

pronotum (fig. 67), and the hair mats on male sterna III-V (fig. 68-69). The elongate

pronotal depression is similar to the condition in the fritzi group, and the lamella

bordering it is probably homologous with the structure found in that group and the

krombeini group. The lamellae of the aedeagal head are distinctive and make the

genitalia unlike any other New World species (figs. 73-74).

Apomorphies of the delicatum group are the broad face (fig. 64), the clavate

antenna with distal flagellomeres wider than long, the non-spiny legs with peculiar

broad tarsal claws (figs. 70-72), the straight (incomplete) episternal sulcus, the

propodeal ridge, the long pronotal depression (fig. 67), and the hair mats of the male

sterna (figs. 68-69). The antenna, absence of erect body setae, tarsal claws and

completely punctate tegula are shared with the krombeini and stangei groups, but

they have densely setose eyes (an apomorphy) and an episternal sulcus that curves

forward ventrally to the anterior margin of the mesopleuron. The krombeini group

differs from the delicatum group in possessing a broad lamella on the anterior

Menke: Pison in the New World 37

margin of the pronotum (an apomorphy), in lacking a lateral ridge on the propodeum,

and in having a complete occipital carina. The male abdominal sterna have

specialized areas in delicatum and krombeini groups, but they are dissimilar and

presumably this is convergence.

The delicatum and stangei groups share the propodeal ridge that extends between

the propodeal spiracle and petiole socket, but in the stangei group there are only two

submarginal cells and the eye is setose (both apomorphies). The pronotum has a

simple, median pronotal pit anteriorly and the collar lacks an anteromedian

depression in the stangei group.

Pison agile and delicatum share a straight episternal sulcus, a completely

punctate tegula, and have similar legs and antenna, but the setose eyes, lamelliform

metapleural flange, and two submarginal cells are apomorphies of the agile group

that separate it from the delicatum group. The occipital carina is a complete circle

in the agile group.

Range:

Pison delicatum occurs from Colombia and Venezuela to northern Argentina and

southern Brasil.

Types:

Holotype male: VENEZUELA, Zulia: Los Angeles del Tucuco, IV-15/16-1981, A.

S. Menke and L. Hollenberg (USNM).

Paratypes (22 females, 20 males): COLOMBIA, Vaupes: Mitu, V-16-74, M.

Cooper (BMNH); Meta: Cord. Macarena, II-15/28-76, M. Cooper (BMNH); Valle:

Caula, VII-10-75, R. Wilkerson, malaise trap (CSDA), finca San Luis near Candelaria,

IX-7/9-75, J. Lattke (CAS). VENEZUELA, Borburata, Car., III-15-40, P. Anduze

(CU). ECUADOR, Napo: Coca on Rio Napo, V-65, L. Pefia (AMNH), Tena,

XII-9/14-71, M. Cooper (BMNH), Limoncocha, VI-15/28-76, S. & J. Peck (CNC);

Zamora-Chinchipe: Timbara, IV-4-65, L. Peha (AMNH). PERU, Cuzco: Quillabamba,

XII-23/27-83, L. Huggert (PMA); Huanuco: Tingo Maria, VI-21-82, malaise trap,

Wasbauer and Slansky (CSDA); Amazonas: Rodriquez de Mendoza, 1500 m, V-16-82,

M. Cooper (COOPER); Junin: Valle Chanchamayo, 800 m., 1939, VIII-12-51, W.

Weyrauch (LILLO), San Ramon, Valle Chanchamayo, 800 m., II-12-40, W. Weyrauch

(AMNH). BOLIVIA, Beni: Rio Itenez about 4 km. above Costa Marques (Brasil),

IX-12/18-64, Bouseman and Lussenhop (AMNH); La Paz: Chulumani, 1700 m.,

IV-2-79, M. Cooper (BMNH); Coroico (BERLIN). PARAGUAY: San Pedro Cororo-Rio

Ypane, XII-1/4-83, malaise trap, M. Wasbauer (CSDA); Rio Ypane, Cororo, XI-79, M.

Fritz (FRITZ). ARGENTINA, Misiones: El Dorado, XII-70, Foerster (FRITZ);

Cataratas del Iguazu, XI-5/9-70, Porter and Stange, (LILLO); Salta: Oran, Abra

Grande, X-18/25-68, C. Porter (MCZ). BRASIL, Bahia: Itabuna, XI-15-82, II-21-83,

F. P. Benton (BMNH); Sao Paulo: Ribeirao Pruto, X-10-65, R. Neilsen (UCD);

Campinas, 1903, Hempel (VIENNA); Espirito Santo: Colatina, X-69, F. M. Oliveira

(AE]I).

Non-type material (one headless specimen): BRASIL, Para: Almeirim, XII-16-02,

A. Ducke (VIENNA).

Etymology:

The name delicatum is the Latin word meanings delicate, soft, or tender.

Agile Group

(Figs. 77, 81-85)

Description:

Frons strongly swollen; face not broad, eye length slightly greater than distance

between eye notches (fig. 77); labrum quadrangular, free margin entire; free margin

of female clypeus with rounded median lobe whose edge is slightly thickened (fig. 77);

female mandible with one inner tooth that is located slightly beyond midpoint; eye

38 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

densely covered with short setae; antenna clavate, outer flagellomeres broader than

long; occipital carina a complete circle, narrowly separated from apex of hypostomal

carina; pronotum with small, round anterodorsal pit but no lamella; scutal flange

moderately broad; tegula completely punctate; scutellum bordered anterad by row of

foveolae; propodeum punctate and with median longitudinal carina on dorsum but

without lateral ridge at top of side; propleuron slightly gibbous posterolaterally, disk

polished, impunctate; episternal sulcus straight, ending ventrad without curving

forward; mesopleuron with subomaulus that often almost joins lower end of episternal

sulcus; mesopleural sulcus paralleled anteriorly by row of foveolae; metapleural

flange broadly lamelliform posteriorly (fig. 81); forewing media diverging after cu-a;

forewing with two submarginal cells, second not petiolate, recurrent veins ending on

submarginal cell I and II, respectively, or second interstitial between I-II; midcoxa

bordered anterad by high ridge that is associated with precoxal sulcus; legs smooth,

devoid of conspicuous spiniform setae at apices of tarsomeres; female foretrochanter

without carina; hindcoxa with low inner carina dorsally, but without outer carina;

hindtarsomere IV with very tiny plantula; tarsal claw thick to just before apex;

tergum I simple apically; male sternum VIII narrow, entire or weakly indented at

apex; genitalia laterally compressed, gonostyle simple but fringed ventrally with

coarse, arcuate setae, volsellar lobes small, weakly setose.

Included species:

Pison agile in the North America, others (including agile) in the Oriental and

Palearctic Regions: browni (Ashmead), differens Turner, hissaricum Gussakovskij, and

possibly erythropus Kohl and rothneyi Cameron.

Discussion:

The following characterize the agile group, an Old World assemblage with one

species adventive in North America: eyes setose, occipital carina complete, pronotum

with only a small median pit anteriorly, subomaulus present, episternal sulcus

straight, two submarginal cells, hindcoxal dorsum without an outer carina, and

propodeum without ridge laterally. I have studied examples of browni and differens

as well as agile. Although sternum VIII of the male is bluntly rounded with a slight

indentation in agile (fig. 82), it can be narrower and more pointed as in browni from

the Philippine Islands (see fig. 239 in Tsuneki, 1983).

Apomorphies of the agile group are the setose eyes, the clavate antenna, the two

submarginal cells, the presence of a subomaulus, the straight episternal sulcus, the

broad metapleural flange (fig. 81), and the form of the tarsal claws. The subomaulus

is merely a short spur in browni.

Earlier I (Menke, 1968b) called this the koreense group but Krombein (1979)

indicated that agile was a senior synonym of Radoszkowski's name.

Pison agile (Smith)

(Figs. 77, 81-85)

Parapison agilis Smith, 1869:300. Holotype female: Ceylon (BMNH).

Paraceramius koreensis Radoszkowski, 1887:433. Holotype female: Korée (Mus.

Krakow?)

Pison koreense, Krombein, 1958a:166. Adventive in North America (Illinois, Virginia).

Pison koreense, Krombein, 1958b:189. Catalog.

Paraceramius koreensis, Richards, 1962:118. Paraceramius preoccupied,

Krombeinielleum proposed as replacement name.

Pison koreensis, Iwata, 1964:1. Biology.

Pison koreense, Krombein, 1967:394. Records from Maryland and Michigan.

Pison koreense, Menke, 1968a:3, 7. Taxonomy, listed.

Pison koreense, Menke, 1968b:1100, 1102. Taxonomy, keyed.

Pison koreense, Sheldon, 1968:107. Biology.

Pison koreense, Bohart and Menke, 1976:333, 337. Listed.

Menke: Pison in the New World 59

Pison agile, Krombein, 1979:1641. Synonymized koreense with agile, record from

Kansas.

Discussion:

The following combination of characters separate agile from all other New World

Pison: eye densely setose, metapleural flange broadly lamelliform (fig. 81), forewing

with only two submarginal cells, propodeum without a dorsolateral ridge, and surface

of tergum I dull in contrast to the shiny surface of II. Sternum VIII and the male

genitalia are shown in figures 82-85.

This is the only Pison known in North America, and it is adventive from the

Oriental Region (Krombein, 1958a). I have not studied the types of koreense or agile.

Biology:

Iwata (1964) reported on agile nests in Japan, and Sheldon (1968) observed the

species in Illinois and described the biology in detail, including illustrations of the

larva. Both authors used the name koreense. Iwata illustrated the cells of agile, and

described them as “very thin and fragile” and said that “clearly [they] may be the .....

most fragile mud cell{s] made by Japanese Aculeata." He said that a nest (16 studied)

consisted of one to 21 separate but contiguous cells arranged irregularly (average

number of cells/nest = 7.6). They were glued to vertical protected surfaces, and the

exterior surface had a "ripple relief." Iwata found 6 immature spiders of the genus

Araneus in one cell.

Sheldon reported that agile made its nests in sheltered situations also, and even

used empty cells of Trypoxylon politum (Say) nests. One to 12 fragile cells, arranged

randomly, made up a nest. Adult spiders of the genus Dictyna representing two

species, were the prey found in four cells, but immatures of the same genus were also

provisioned. The average number of spiders per cell was 25.2. Sheldon stated that

the female carried the paralyzed spider in her mandibles. The wasp egg was laid on

the last spider provisioned. The eulophid parasitoid Melittobia chalybii Ashmead was

reared from one cocoon of agile that Sheldon found in an unsealed cell of Trypoxylon

politum (Iwata found no parasitoids in his observations).

Range:

Apparently widespread in the Oriental Region (Sri Lanka, India) and eastern Asia

(China, Japan, Korea). In North America agile is known from Kansas, Illinois,

Michigan, New York, Maryland, and Virginia.

Material examined (12 specimens):

ILLINOIS: Palisades Park (USNM); Kickapoo State Park (USNM). MICHIGAN:

Gun Lake Mal. Tr. (BMNH). NEW YORK: Ithaca (CU); Lewisboro (AMNH).

VIRGINIA: McLean (USNM, BMNH).

Stangei Group

(Figs. 9, 12, 78, 86-108)

Description:

Scape, mandible, mouthparts, and most of fore and mid legs yellowish brown.

Body devoid of erect setae; frons not swollen; face not broad, eye length greater than

distance between eye notches; free margin of labrum arcuate or quadrangular; female

clypeus not double-edged but margin sometimes thickened, a median lobe weakly or

not differentiated; male clypeus tridentate, thickened; female mandible with one

inner subapical or median tooth or simple; eye densely covered with short setae;

antenna clavate, outer flagellomeres broader than long; occipital carina usually

incomplete below, ending just before reaching midventral line, occasionally a

complete circle narrowly separated from apex of hypostomal carina; pronotum with

small round or transversely elongate pit anterodorsally; pronotal collar without

anteromedian depression but sometimes with obliquely transverse ridge; scutal flange

40 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

narrow, upturned; parapsidal lines sulciform in female; tegula completely

micropunctate; scutellum margined anteriorly by row of contiguous foveolae;

propodeal side delimited dorsad by crenulate ridge that extends from petiole socket

area to spiracle; propleuron punctate, with gibbosity posterolaterally; episternal

sulcus curving forward ventrally, attaining anterior margin of mesopleuron but often

weakly impressed there; mesopleural sulcus paralleled anteriorly by row of foveolae;

metapleural flange narrow; forewing media diverging after cu-a; forewing with two

submarginal cells (three in rare exceptions, figs. 87-88), II not petiolate in two-celled

wing (figs. 12, 86); both recurrent veins ending on submarginal cell I, or second

interstitial between I-II, or ending on II; midcoxal cavity bordered anterad by high

ridge associated with precoxal sulcus; legs smooth, devoid of conspicuous spiniform

setae at apices of tarsomeres; tarsomeres apparently without plantulae; tarsal claw

thick to just before apex; female foretrochanter not carinate ventrally; dorsum of

hindcoxa with low or very weak inner carina and no outer carina or only an apical

vestige; apical rim of tergum I simple; male sternum VIII narrow, apex truncate or

with shallow emargination; gonostyle of male genitalia long, slender, simple, setose

ventrally, volsellar lobes small, aedeagus without ventral spine.

Included species:

Pison abathes, duckei, plaumanni and stangei.

Discussion:

The following characterize the stangei group: eye densely setose (fig. 9),

pronotum with small median pit anteriorly (fig. 95), episternal sulcus complete,

forewing usually with only two submarginal cells (but see below), and propodeum with

dorsolateral ridge. In this assemblage the true second submarginal cell has been

reduced to the point of disappearing, but in one male a tiny second submarginal is

present in both wings (figs. 87-88). The eye notch is unusually shallow in one species,

abathes. Members of the stangei group are among the smallest species of Pison

known.

Apomorphies of the group include the setose eyes, the clavate antenna, the form

of the tarsal claws, the two submarginal cells, and lateral propodeal ridge.

In terms of species discrimination, the stangei group is the one of the most

vexing assemblages in the New World fauna. More material has been available than

when I first treated the group (Menke, 1968b), but differences between stangei and

duckei are still not as precise as one would like, and the male of the latter is still

unknown. Pison plaumanni remains the most distinctive species and its male is now

known. One new species has been discovered. Finally a male from Ecuador (AMNH)

has three submarginal cells, the second represented in each forewing by a much

reduced cell (figs. 87-88). The wing condition of this specimen may be a freak

occurrence, but in any case it is probably an undescribed species. The male genitalia

are fairly similar in plaumanni, stangei and the specimen with three submarginal cells.

Pison plaumanni Menke

(Figs. 9, 12, 89-90, 95, 101-104)

Pison plaumanni Menke, 1968b:1105. Holotype female: Nova Teutonia, Santa Catarina,

Brasil (UCD).

Description:

Pedicel, basal part of flagellum, and clypeal margin often yellowish brown.

Female clypeus with weakly defined lobe: it is thickened, truncate, and its edge has a

transverse dimple (fig. 89; the clypeal outline in Menke, 1968b, fig. 7, is inaccurate).

Male clypeus as in fig. 90. UID 0.68-0.69X distance between the eye notches in

females, 0.72X in males. LID 0.69-0.71X UID in females, 0.63-0.66X in males.

Ocellocular distance greater than other species of stangei group (OOD 1.38-1.65X

HOD). Occipital carina broadly interrupted ventrad in comparison with other species,

gap equal to length of foretarsomere I or only slightly shorter (17:20). Anterodorsal

Menke: Pison in the New World 4l

pronotal pit circular to transverse, usually margined posterad by carina. Pronotal

collar rounded laterally, no trace of transverse ridge or humeral angle (fig. 95). Disk

of propodeal dorsum smooth, punctate, with median longitudinal carina that is often

in depression; carina usually reaches propodeal apex or nearly so, and frequently

many short cross-carinae associated with it. Propodeal hindface varies from

punctate to transversely ridged and punctate. Propodeal side smooth, punctate,

without ridges except sometimes just beneath spiracle. Second recurrent vein

interstitial between submarginal cells I-II, or ending just inside I. Male sternum VIII

slightly emarginate at apex, lateral margins parallel or slightly diverging basad (fig.

102).

Length 5-6.5 mm.

Discussion:

This is the largest species in the stangei group. The broad ocellocular distance,

the rounded humerus of the pronotal collar (and absence of an oblique ridge), and

smooth propodeal dorsum and side are the most diagnostic features. The eighth

sternum of the male is not constricted before the apex in contrast to that of stangei.

I have examined two females from Mocoa, Putumayo, Colombia (BMNH) that

resemble plaumanni in the clypeus, the broadly interrupted occipital carina, the

rounded humerus of the pronotal collar, the propodeal sculpture, and body size.

However, the ocellocular distance is only slightly greater than an ocellus diameter

(OOD 1.1l-1.14X HOD), the LID is 0.76-0.78X the UID, and the UID is only 0.61-0.62X

the distance between the eye notches. Thus these specimens may represent an

undescribed species.

Range:

Pison plaumanni is known only from eastern and southeastern Brasil.

Material examined (26 females, 12 males):

BRASIL, Bahia: Itabuna, VIII-7-83, III-18-84, F. Benton (BMNH); Aguas

Vermelhas, XII-83, M. Alvarenga (AEI). Sao Paulo: Sao Paulo, various dates, V. Alin

(MLSU). Santa Catarina: Nova Teutonia, various dates, F. Plaumann (UCD, MCZ,

USNM, BMNH). "Brazilia", 1886 (USNM).

Pison stangei Menke

(Figs. 86, 91-92, 94, 96, 105-108)

Pison stangei Menke, 1968b:1102. Holotype male: Amaicha, Tucuman,

Argentina (LILLO).

Description:

Antenna and fore and midlegs occasionally entirely or largely black. Female

clypeus with prominent median lobe the edge of which is variably thickened (fig. 91).

Male clypeus as in fig. 92. UID 0.59-0.62X the distance between eye notches in

females, 0.62-0.64 in males. LID 0.74-0.81X UID in females, 0.76-0.79X in males.

OOD 1.08-1.27X HOD. Occipital carina often a complete circle narrowly separated

from apex of hypostomal carina, but sometimes weakly complete or narrowly

incomplete at midventral line. Anterodorsal pronotal pit circular, not margined

posterad by carina. Pronotal collar rounded laterally although weak transverse ridge

present (figs. 94, 96). Propodeal dorsum with median carina that fades posterad,

usually ending well before apex; disk of dorsum variably sculptured: coarsely to finely

obliquely ridged or striatopunctate, or ridging becoming transverse on apical half to

two-thirds, or surface obscurely striatopunctate, or even mostly just punctate.

Propodeal hindface transversely ridged, ridges sometimes evanescent laterad.

Propodeal side obliquely or horizontally ridged above, punctate below. Second

recurrent vein ending on submarginal cell I or sometimes interstitial between I-II.

Male sternum VIII slightly concave at apex, lateral margins slightly constricted

42 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

subapically (fig. 108). Male genitalia as in figs. 105-107, aedeagus with midventral

ridge (figs. 105-106).

Length 4.5-6 mm.

Discussion:

The rounded humerus of the pronotal collar and the fine ridges on the upper half

of the propodeal side are the most obvious diagnostic features of stangei. The weak

transverse ridge on the collar (fig. 96) is a condition intermediate between plaumanni,

which has none (fig. 95), and duckei and abathes, both of which have a well developed

oblique ridge (figs. 97, 99). Except for the difference in the collar, stangei is similar

to duckei, and their head measurements are essentially identical. The occipital

carina is often a complete circle in stangei, but it is always interrupted in duckei.

When I described stangei (Menke, 1968b), I used the ridging of the propodeal dorsum as

a diagnostic feature for separation from duckei, the latter having a mostly punctate

dorsum. But the sculpture of the dorsum varies considerably in stangei, some

specimens being mostly punctate and therefore similar to duckei. Thus propodeal

sculpture is not reliable for separating these two species.

Range:

Pison stangei is known only from northwestern Argentina and Bolivia.

Material examined (14 females, 4 males):

BOLIVIA, Tarija?: Las Carreras, I-4-56 (KU). ARGENTINA, Salta: Rosario de

Lerma, XI-10/18-83, M. Wasbauer (CSDA), Pocitos, I-1971, M. Fritz, Martinez and

Fritz (USNM). Tucuman: Horco Molle, Parque Sierra San Javier, 700 m, various

dates, L. A. Stange (I_LILLO), Jardin del Lillo, IX-19-67, C. Porter (MCZ), Tucuman,

XI-24-65, L. A. Stange (LILLO); Santiago del Estero: Suncho Corral, XII-28-75, L.

Stange (LILLO).

Pison duckei Menke

(Figs. 97-98)

Pison duckei Menke, 1968b:1103. Holotype female: Para, Brasil (VIENNA).

Description (female only):

Pedicel and flagellomere I usually yellowish, clypeal margin yellowish in type,

hindleg largely yellowish brown in material from Trinidad, midleg black above in

specimen from Brasilia. Clypeus with prominent median lobe whose edge is thickened

(similar to stangei). UID 0.60-0.66X distance between eye notches. LID 0.70-0.79X

UID. OOD 1.1-1.31X HOD. Occipital carina interrupted ventrad. Anterodorsal

- pronotal pit circular, margined posterad by short carina. Pronotal collar with

oblique, transverse ridge that makes humerus angulate when viewed from above (figs.

97-98). Propodeal dorsum punctate, with median longitudinal carina that ends at

middle or slightly beyond, many short cross-carinulae associated with median carina,

base of dorsum with up to 10 oblique carinae that only rarely extend onto disk (one

Trinidad specimen); hindface with fine cross-carinulae that become evanescent

dorsad; propodeal side variable: sometimes with several horizontal ridges only

beneath spiracle, sometimes upper area more extensively ridged. Both recurrent

veins ending on first submarginal cell.

Length 4-4.5 mm.

Discussion:

Pison duckei is similar to stangei. The essential features of duckei for

separation from stangei are: pronotal collar with obvious oblique transverse ridge

that makes humerus clearly angulate (fig. 97), occipital carina interrupted, and

propodeal dorsum mostly punctate. The degree of the pronotal angulation varies

however, and the punctate propodeal dorsum is approached by some specimens of

Menke: Pison in the New World 43

stangei. The sculpture of the propodeal side is often similar between the two species.

The male of duckei is unknown. The male with three submarginal cells discussed

under the stangei species group is not the unknown male of duckei, although its head

measurements fall within the range of the species. The sculpture of its propodeal

dorsum is more like stangei, the humerus of the pronotal collar is not angulate, and of

course the wing has three submarginal cells.

Range:

Pison duckei is known from scattered localities between Panama and central

Brasil.

Material examined (11 females):

PANAMA: Las Cumbres, XII-8/28-82, H. Wolda (PMA). COLOMBIA, Tolima:

Armero, I-30/II-5-77, E. L. Peyton (USNM). VENEZUELA, Guarico: Hato

Masaguaral, 44 km s Calabozo, 5-3/10-85, Menke & Carpenter (USNM). TRINIDAD:

St. Augustine, various dates, E. Callan, J. Noyes (BMNH). BRASIL, Para: Para

(Belém), I-18-01, A Ducke (holotype) (VIENNA); Federal District: Brasilia, Lago Sul,

NI-7-77, A. Raw (BMN#).

Pison abathes Menke, new species

(Figs. 78, 93, 99-100)

Description, holotype female:

Black, following yellow brown: palpi, mandible, labrum, clypeal margin, scape

and pedicel beneath, margin of pronotal lobe, fore and midlegs except coxae, and

hindtrochanter.

Labrum trapezoidal in shape, shallowly concave (fig. 93); clypeus with arcuate

median lobe whose margin is slightly thickened; eye notches shallow (fig. 78), UID

0.8X distance between eye notches; LID 0.68X UID; OOD 1.27X HOD; occipital

carina complete but weak opposite apex of hypostomal carina and separated from

latter; anterodorsal pit of pronotum circular, smaller than hindocellus; collar with

oblique, transverse ridge that makes humerus angulate in dorsal view (figs. 99-100);

scutum, scutellum and metanotum weakly shining, finely, nearly contiguously

punctate, punctures largest on scutum; propodeal dorsum shiny, with about 8 short,

coarse carinae at base, middle one extending only about one-third distance to apex,

disk of dorsum irregularly transversely carinulate changing to punctation laterally,

apex of dorsum delimited from hindface by short carina; hindface shiny, with coarse

cross-ridges; propodeal side longitudinally carinulate on upper one-fifth, remainder

punctate; both recurrent veins ending on submarginal cell I.

Length 4.5 mm.

Variation in females (3 specimens):

Clypeus completely black (Bolivia), hindfemur yellow brown (Bolivia, Guyana).

Edge of clypeal lobe not thickened (Bolivia, Guyana). UID 0.76-0.78X distance

between eye notches; LID 0.67-0.70X UID. OOD 1.13-1.27X HOD. Propodeal dorsum

largely punctate in Bolivian specimen, median carina extending half distance to apex,

the latter without transverse carina; dorsum in Guyana specimen more like holotype

but median carina almost reaching apex, the latter with only suggestion of a

transverse carina. Propodeal side with only two carinulae just beneath spiracle

(Bolivia, Guyana). Second recurrent vein interstitial in one wing of Guyana paratype.

Length 4-4.5 mm.

Male:

Unknown.

44 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

Discussion:

The shallow eye notches, angulate pronotal humerus, and trapezoidal labrum are

the salient features of abathes. The variation seen in the sculpture of the propodeal

dorsum is disconcerting and it may be that the Bolivia and Guyana material is not

conspecific with the Ecuadorian females. However, there is also considerable

variation in the propodeal sculpture of stangei. It is possible also that the shallow

eye notches of abathes simply represent extreme variation of duckei, the only other

species with angular pronotal humeri. But lacking intergrades I have decided to treat

these specimens as a new species.

Range:

Ecuador, Bolivia and Guyana.

Types:

Holotype female: ECUADOR, Pichincha: Rio Palanque Res. Sta., II]-1983, malaise

trap, Sharkey and Masner (PMA).

Paratypes (three females): ECUADOR, same data as type (PMA). BOLIVIA,

Santa Cruz: Rio Mamore, 2 km n mouth of Rio Chapare, VII-31-65, J. Bouseman

(AMNH). GUYANA: Mazaruni, clearing, in colony house, VIII-26-37, O. Richards

(BMNH).

Etymology:

The name abathes is a Greek word that means shallow, a reference to the

distinctive eye notches of this species.

Krombeini Group

(Figs. 79-80, 109-116)

Description:

Frons not strongly swollen; face broad, eye length about 10% shorter than

distance between eye notches (figs. 79-80); labrum transverse, hidden, free margin

arcuate or obtusely triangular, entire; free margin of female clypeus not thickened or

double-edged, outline a low obtuse triangle or with median lobe; female mandible

with subapical inner tooth; occipital carina a complete circle, broadly contiguous

with apex of hypostomal carina; eye densely covered with short setae; antenna

clavate, outer flagellomeres broader than long; anterior edge of pronotum with broad,

thin, setose lamella that extends laterad to level of pronotal lobe; pronotum without

anterodorsal pit; scutal flange broad, upturned; tegula evenly punctate; scutellum

with unpitted narrow sulcus anteriorly; propodeum punctate, with median longitudinal

carina on dorsum but without lateral ridge, hindface without cross-ridges; propleuron

with short, high lamelliform projection posterolaterally; episternal sulcus curving

forward ventrally, attaining anterior margin of mesopleuron; mesopleural sulcus

paralleled by series of foveolae; forewing media diverging after cu-a; forewing with

three submarginal cells, II receiving both recurrent veins; midcoxal cavity delimited

anterad by high ridge associated with precoxal sulcus; legs smooth, devoid of

conspicuous spiniform setae at apices of tarsomeres; female foretrochanter with

carina ventrally; outer dorsal carina of hindcoxa represented by high apical lamella;

tarsi without plantulae; tarsal claw thick to just before apex; tergum I with reflexed

apical rim that is lamelliform; male sternum VIII semicircularly emarginate apically,

flanked by two prongs (fig. 110); male genitalia as in figs. 113-116, gonostyle setose,

with narrow, ventral, subapical lobe (fig. 115); volsellar lobes large, setose apically;

aedeagal head with ventral process (see fig. 13 in Menke, 1968b).

Included species:

Pison krombeini and neotropicum.

Menke: Pison in the New World 45

Discussion:

The following define the krombeini group: face broad, eye setose, occipital

carina complete, pronotum with setose lamella along anterior margin, propodeum

without ridge laterally, forewing with three submarginal cells, tegula entirely

punctate, tergum I apex reflexed, and male sternum VIII emarginate, bounded by

prongs. The tergal character needs some elaboration. The lamella-like apical margin

of tergum I is not the real rim. The true edge of the tergum can be seen beneath this

broad, upturned flange - thus, tergum I is really double-edged. Earlier I (Menke,

1968b) described the female mandible as having two subapical teeth, but there is only

one tooth in krombeini, and only a suggestion of a second subapical tooth in

neotropicum.

Apomorphies of the group are the broad face, the setose eye, the pronotal

lamella, the carinate female foretrochanter, the reflexed and double-edged margin of

tergum I, and the two prongs at the apex of male sternum VIII. A similar but asetose

pronotal lamella also occurs in the pilosum group. The male of krombeini has

transverse depressions on sterna III-IV, but the male of neotropicum is unknown; thus,

at present, it is impossible to list this specialization as an additional apomorphy for

the group.

Because I described both species earlier (Menke, 1968b), I am presenting only

brief diagnoses and distribution records here.

Pison krombeini Menke

(Figs. 79, 109-110, 112-116)

Pison krombeini Menke, 1968b:1105. Holotype male: British Honduras (= Belize)

(USNM).

Diagnosis:

The dense metallic setae of the propodeal dorsum and hindface is the most

obvious recognition feature. The female clypeus (fig. 109) differs from that of

neotropicum (fig. lll). The thorax is shiny, the pronotal collar has a prominent ridge

anterolaterally, and the parapsidal and admedian lines of the scutum are deeply

impressed. The metapleural flange is narrow. Terga I-II have broad silver or golden

apical fasciae, and sternum I has appressed silver setae that are directed basad. The

forewing has a cloud at the apex of the first submarginal cell and the leading edge of

the wing is faintly infumate. The inner and outer veinlets of submarginal cell III are

separated on the marginal cell by an ocellus diameter or less, sometimes joining

there, or even forming a petiole. The depressions on male sterna III-IV are

presumably diagnostic also (fig. 112). The aedeagus has a large ventral prong (not

visible in figs.), the apices of the two penis valves curl away from each other (figs.

114-115), and the gonostyle has a curled fingerlike process on its inner, ventral

margin near the apex (figs. 114-115). Length 6-7.5 mm.

Range:

Southern Mexico to Colombia.

Material examined (13 females, 3 males):

MEXICO, Vera Cruz: Cordoba (GENEVA); Tabasco: Teapa, March, H. H. Smith

(BMNH); Yucatan: Chichen Itza, VII-20-52, J. & D. Pallister (AMNH). BELIZE:

“British Honduras", I-O8 (USNM). GUATEMALA: El Salto, Escuintla, VI-28-34, F. X.

Williams (BPBM), Helvetia, San Sebastian, IV-21/22-31, J. Bequaert ((MCZ), Sta.

Emilia, Pochuta, I/II-31, J. Bequaert (MCZ). COSTA RICA, Santa Rosa Park,

II-13-78, D. Janzen (AEI). PANAMA: Barro Colorado I., VIII-15/IX-26-28, P. Rau

(note 7756) (USNM). COLOMBIA, Arauca: Tame, VII-8/17-76, M. Cooper (BMNH);

Magdalena: Rio Frio, VII-13-27, G. Salt (BMNH); Valle: Cali and vic., 3-4000 feet,

I-8/20-72, H. Evans (MCZ).

46 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

Pison neotropicum Menke

(Figs. 80, 111)

Pison neotropicum Menke, 1968b:1106. Holotype female: Rio de Janeiro, Brasil

(USNM).

Diagnosis (male unknown):

Distinctive features of neotropicum are the shape of the female clypeus (fig. Ill),

the dull thoracic dorsum, absence of appressed metallic setae on the propodeal

hindface and apex of the dorsum, absence of silver setae on sternum I and absence of

metallic fasciae on terga I-II. The collar has a transverse ridge anteriorly, but it is

not prominent laterally. The admedian and parapsidal lines of the scutum are not

depressed. The metapleural flange is about as broad as an ocellus diameter. The

leading edge of the forewing is moderately infumate, and the inner and outer veinlets

of submarginal cell III are widely separated on the marginal cell. Length 8.5 mm.

No additional material has been found since the species was described.

Range:

Known only from southeastern Brasil.

Material examined (6 females):

BRASIL, Guanabara: Rio de Janeiro, X-38/I-39, R. C. Shannon (USNM, UCD).

Euryops Group

(Figs. ll, 117-130)

Description:

Frons not swollen; face broad, eye length 10-20% shorter than distance between

eye notches (fig. 117, 119); female clypeus with roundly truncate median lobe, its

apex with lower lip of variable shape; male clypeus with sharp, triangular median

lobe; female mandible with cleft near middle of inner margin, but no tooth; labrum

with deep, U-shaped emargination resulting in two apical, setose, fingerlike lobes;

occipital carina joining apex of hypostomal carina; pronotum without anterodorsal

lamella or pit; lateral margin of scutum a broad upturned flange; tegula uniformly

punctate; scutal flange broad; propodeal dorsum without median longitudinal carina;

propodeal side with crenulate ridge that extends from petiole socket nearly to

spiracle; disk of propleuron largely impunctate, shiny; lower end of episternal sulcus

curving forward ventrally, usually attaining anterior margin of mesopleuron; dorsum

of hindcoxa with strong inner carina, but outer carina reduced to short evanescent

distal ridge; forewing media diverging from M+Cu after crossvein cu-a although

sometimes nearly interstitial; forewing usually with three submarginal cells, rarely

two (fig. ll); tergum I narrowly double-edged apically; male sternum VIII with broad,

shallow emargination apically; male genitalia laterally compressed, aedeagus with

broad, apically rounded, laterally compressed head whose ventral margin has a

hooklike angle (not visible in SEM photo, figs. 128-130), volsellar lobes slender,

acuminate, apical half of gonostyle narrow, setose ventrally (figs. 128-129).

Included species:

Pison eu, euryops, lillo, and styphopteron.

Discussion:

The following define the euryops group, one of the most distinctive in the New

World fauna: face broad, labrum with U-shaped emargination, pronotum without

anterodorsal pit, propleuron largely impunctate, tegula completely punctate, and

tergum I double-edged. The broad face, labral form and double-edged tergum are

apomorphies. The labral and tergal character states also occur in the pilosum group

but neither is universal there.

Members of the euryops group have very similar sculpture and vestiture. Species

Menke: Pison in the New World 47

differences are found mainly in clypeal shape, facial proportions, flagellar features,

and sculpture of the propodeal hindface. The loss of the second submarginal cell in

one species, styphopteron, appears to be a consistant feature, but only three

specimens are known. The genitalia display relatively minor differences between the

four species, but insufficient male material of lillo, styphopteron and eu has

hampered assessment of their usefulness in species. discrimination. The

asymmetrically swollen flagellomeres in males of euryops are distinctive, but the few

males available of the other species does not permit positive conclusions on the

reliability of the apparent antennal differences observed.

Pison eu Menke, n. sp.

(Figs. 117-118, 121, 128-130)

Description, holotype female:

Black; wings clear except for cloud over submarginal cell II. Clypeus and lower

half of frons with dense, appressed, silver setae that obscure sculpture; terga I-II

with silver apical fasciae that are interrupted at middle; erect setae of body pale,

longest on metanotum and propodeum.

Clypeal lobe arcuate, its lower lip with broad, shallow emargination (similar to

fig. 118). Frons shiny, finely, densely punctate (punctures | diameter apart or less);

eye length 0.88X distance between eye notches; UID 0.77X LID. OOD 0.85X HOD.

Flagellomere I length 3.5X apical width, remaining flagellomeres progressively

shorter.

Thorax shiny, scutum and scutellum very finely punctate, punctures sparser than

those of frons (1-4 diameters apart); metanotum impunctate; propodeal dorsum

largely smooth but disk with sparse, pinprick punctures that become denser

peripherally and along midline; propodeal hindface with several transverse ridges just

above petiole socket, upper part of hindface smooth, with scattered pinprick

punctures; propodeal side smooth, uniformly covered with pinprick punctures (several

diameters apart). Mesopleuron smooth, densely punctate, punctures 1-2 diameters

apart anteriorly and about same size as those on scutum, becoming sparser, finer

toward mesopleural sulcus; row of foveolae along lower half of mesopleural sulcus.

Metapleuron same as propodeal side.

Forewing with three submarginal cells, recurrent vein I ending on submarginal

cell II, recurrent vein II ending on submarginal cell III (nearly interstitial in right

wing).

Tergum I with fine pinprick punctures similar to those on propodeal dorsum; II

more densely punctate.

Length 7.5 mm.

Variation in females (15 specimens):

Lower clypeal lip not indented in single specimen from Oaxaca. UID 0.76-0.80X

LID. Punctation varies in coarseness from specimen to specimen. Tergum III

sometimes has apical fascia. Recurrent vein II is interstitial between submarginal

cells II and III] in eleven specimens. Occasionally forewing uniformly but weakly

infumate. Length up to 9.5 mm.

Male (2 specimens):

As in female except: mandible as in fig. 121; clypeal lobe as in fig. 121. UID

0.93-0.95X LID; OOD = HOD in Peruvian specimen. Flagellomeres VI-VII each with

weak linear tylus (Venezuelan specimen) or narrowly linear shiny area (Peruvian

specimen). Genitalia as in figs. 128-130. Length 6.5 mm.

Discussion:

The female clypeal lobe and the shape of its lower lip are immediately diagnostic

in that sex (fig. 118). The non-ridged upper part of the propodeal hindface is also

distinctive in both sexes, although styphopteron is similar. The latter species,

48 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

however, has only two submarginal cells and other differences. The upper interocular

distance is 0.73-0.80X the lower interocular distance in females of styphopteron and

eu. In euryops the UID and LID are essentially equal. Flagellomere I is more than 3X

as long as wide in females of eu and usually slightly less than 3X as long as wide in

euryops. This flagellomere is about 4X as long as wide in females of styphopteron.

The difference in the flagellum of the two males is disconcerting, but no other

differences between them are apparent, and I am assuming that they are conspecific.

Range:

Pison eu occurs from southern Mexico to southern Peru and eastward to

Suriname, but I have no records from the Amazon basin proper.

Types:

Holotype female: COLOMBIA, Narino: Barbacoas, IV-2-1974, M. Cooper (BMNH).

Paratypes (17 specimens): MEXICO, Oaxaca, 44 mi. w Tehuantepec, VII-21-52, 9,

Gilbert and MacNeil (UCD). HONDURAS: La Ceiba, 2, F. J. Dyer (USNM). COSTA

RICA, Guanacaste: Santa Rosa Park, VII-15-77, 2, D. H. Janzen (AEI); La Selva Res.

Sta., VI-ll/17-86, ?%, Hanson and Bohart (USU). COLOMBIA, Meta: Restrepo,

VI-18-74, 2, L. A. Stange (LILLO); Rio Duda, III-8/12-76, 2, M. Cooper (BMNH);

Arauca: Tame, VII-1/7-76, ?, M. Cooper (BMNH); Putumayo: Villa Garzon, 400-550

m, XII-25-87, ?, M. Cooper (COOPER). PERU, Huanuco: Tingo Maria, VI-23-82, ¢,

malaise trap, Wasbauer and Slansky (CSDA); Cuzco: Quincemil, 750 m, X-16/31-62,

?, L. Pena (MCZ). VENEZUELA, Araqua: El Limon, 480 m, IV-24/29-V-1-73, 6, 22,

malaise trap, C. Rosales (UCM). GUYANA: Essequibo River, Moraballi Creek,

X-8-29, 22 (BMNH). SURINAME: Paramaribo, VII-20-60, XI-22-57, 29, P. v.

Doesburg (LEIDEN).

Etymology:

The name eu, treated as a noun in apposition, is based on the Greek prefix

meaning true or good, a reference to the fact that the species is valid.

Pison euryops Menke, n. sp.

(Figs. 119-120, 122, 126-127)

Description, holotype female:

Same as described for eu except: forewing lightly infumate except for clear spot

at submarginal cell III; terga I-II with broadly interrupted fascia; clypeal lobe roundly

trapeziform, its lower lip arcuate (similar to fig. 120); eye length 0.80X distance

between eye notches; UID essentially equal to LID; OOD 1.31X HOD; flagellomere I

length less than 3X apical width (19.5:7.5); punctures of scutum and scutellum 1 to 2

diameters apart, metanotum with sparse pinprick punctures; propodeal dorsum

punctures slightly larger than those on scutum, | to 2 diameters apart; propodeal

hindface completely covered by coarse, transverse ridges; second recurrent vein

interstitial between submarginal cells II and III; tergum I with punctation similar to

scutum.

Length 8 mm.

Variation in females (15 specimens):

Eye length up to 0.87X distance between eye notches. OOD usually 1.25X HOD,

but OOD = HOD in one specimen from Guanabara. Thoracic punctation varies in

degree of coarseness from specimen to specimen. Recurrent vein I occasionally

interstitial between submarginal cells I-II. Length 6.5-9 mm.

Male (9 specimens):

As in female except: inner margin of mandible sinuate (fig. 122); clypeal lobe a

simple triangle, sides straight (fig. 122); LID 0.83-0.95X UID; OOD 1.33X HOD or

slightly more; flagellomeres III-VI with linear, asetose, shiny areas ventrally (fig.

Menke: Pison in the New World 49

127), IV-VII or VIII swollen toward apex in profile, V-VI most strongly so (fig. 126);

second submarginal cell greatly reduced in two specimens from Argentina (smaller

than ocellus, absent in one wing of one specimen); first recurrent vein ending most

commonly on submarginal cell II, but sometimes interstitial or ending on I, second

recurrent vein usually interstitial but sometimes ending on submarginal II or III;

genitalia similar to eu. Length 6-7.5 mm.

Discussion:

The arcuate shape of the lower lip of the female clypeal lobe (fig. 120), and the

asymmetrically swollen flagellomeres of the male antenna (fig. 126) are diagnostic

for euryops. The completely ridged propodeal hindface is an additional character

that is shared only with lillo. Females of euryops differ from eu and styphopteron in

the essentially equal lower and upper interocular distances.

The reduction of the second submarginal cell in the males from Argentina does

not occur in the single female known from that country.

Range:

Pison euryops is known from Colombia, central and southern Brasil and northern

Argentina.

Types:

Holotype female: BRASIL, Santa Catarina: Nova Teutonia, XI-13-1961, 92, F.

Plaumann (MCZ).

Paratypes (24 specimens): COLOMBIA, Meta: Cord. Macarena, II-15/29-76, ?, M.

Cooper (BMNH). BRASIL, Distrito Federal: Res. Ecol. IBGE, various dates 1981-82, 4

g, 3 ?, (IBGE); Guanabara: Represa Rio Grande, VI-67, 9, M. Alvarenga (AEI); Rio de

Janeiro, IX-38, ?, R. C. Shannon, (USNM); Santa Catarina; Nova Teutonia, various

dates, 7 2, 36, F. Plaumann (MCZ, UCD, LACM, KU, OSU); Corupa, XII-40, 9, A.

Maller (AMNH). ARGENTINA, Entre Rios: Salto Grande, Concordia, IX-64, 9?

(GEMBLOUX); Misiones: Puerto Rico, XI-5/13-70, 6, Porter & Stange (LILLO); Dos

de Mayo, XII-73, 6, M. Fritz (FRITZ).

Etymology:

The name euryops is based on the Greek words eurys (= broad) and ops (= face),

and is a reference to the broad head.

Pison lillo Menke, n. sp.

Description, holotype male:

Black; mandible, tegula and tarsi brown; wings clear; clypeus and lower half of

frons densely covered by pale, gold, appressed setae that obscure sculpture; setae of

thorax and gaster pale; terga I-II with broadly interrupted fasciae.

Mandible and clypeal lobe same as euryops; frons weakly shiny, finely, densely

punctate, punctures about one diameter apart; eye length 0.82X distance between eye

notches; LID 0.90X UID; OOD 1.55X HOD; flagellomere I length slightly less than 3X

apical width, III-VI with narrow, asetose shiny areas ventrally, that on III elevated as

a tylus, V-VI not asymmetrically swollen (flagellomeres X-XI missing on left antenna,

III-XI missing on right antenna).

Thorax shiny, punctation similar to euryops and eu, propodeal hindface

completely covered by coarse, transverse ridges.

Forewing with three submarginal cells, recurrent vein I ending on I, recurrent

vein II ending on III, but both nearly interstitial.

Genitalia similar to eu.

Length 7 mm.

Female:

Unknown.

50 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

Discussion:

Pison lillo is known from a single male that is similar to euryops, differing

mainly in having non-swollen flagellomeres. Both species have similar facial

measurements and have ridges to the top of the propodeal hindface.

Range:

Known only from northern Argentina.

Type:

Holotype male: ARGENTINA, Tucuman: Trancas, Tacanas, III-3-1948, R.

Golbach (LILLO).

Etymology:

The name lillo, a noun in apposition, is a dedication to the Instituto Miguel Lillo.

Pison styphopteron Menke, n. sp.

(Figs. 11, 123-125)

Description, holotype female:

Black; wings clear; clypeus and lower half of frons with dense, appressed silver

setae that obscure sculpture; terga I-II with broadly interrupted tarnished silver

fasciae; erect setae of body pale, longest on metanotum and propodeum.

Clypeal lobe reflexed, quadrangular, corner rounded, lower lip triangular, tip

rounded (similar to fig. 124). Frons shiny, punctate, punctures one to two diameters

apart; eye length 0.89X distance between eye notches; UID 0.73X LID; OOD 0.60X

HOD. Flagellomere I length slightly more than 4X apical width, remaining articles

progressively shorter.

Thorax shiny, scutal punctures similar to frons, one to three diameters apart,

scutellum strongly gibbous and more sparsely punctate than scutum, metanotum very

finely punctate; propodeal dorsum and upper half of hindface smooth, very finely,

sparsely punctate, lower part of hindface with several transverse ridges; propodeal

side smooth, sparsely, finely punctate. Mesopleuron smooth, punctures similar to

those on scutum but sparser, finer, toward mesopleural sulcus; row of foveolae along

lower half of mesopleural sulcus. Metapleuron with fine, pinprick punctation.

Forewing with two submarginal cells via fusion of II and III, outer veinlet of

submarginal cell I angled, appendiculate (fig. 11); first recurrent vein interstitial in

right wing, ending on submarginal II in left wing.

Tergum I more sparsely punctate than II.

Length 7 mm.

_ Variation in females (1 specimen):

Eye length 0.85X distance between eye notches; OOD slightly more than 0.75X

HOD; flagellomere I length almost 4X apical width. Scutellum with median

longitudinal impression. Propodeal hindface with traces of cross-ridging on upper

half. Outer veinlet of submarginal cell I not appendiculate, recurrent vein I ending on

submarginal cell II. Length 8.5 mm.

Male (1 specimen):

As in female except: inner margin of mandible a simple arc (fig. 123); clypeal

lobe as in fig. 123. Eye length 0.91X distance between eye notches; UID 0.93X LID;

OOD 0.90X HOD; flagellomere I length slightly less than 3X apical width;

flagellomeres simple. Thorax as described for holotype. Genitalia similar to eu but

gonostyle apex comparatively narrower dorsoventrally and apical margin of aedeagus

angled. Length 5.5 mm.

Discussion:

The presence of only two submarginal cells and’ the angled outer veinlet of

Menke: Pison in the New World | 51

submarginal cell I are unique features of styphopteron. The shape of the female

Clypeal lobe and lip are distinctive (figs. 124-125), as is the arcuate cutting edge of

the male mandible and simple male flagellum. All species of the euryops group have

a gibbous scutellum but it is most strongly swollen in females of styphopteron. This

is a difficult character to use because it is a matter of degree and there is variation

in the two females available.

Range:

Known only from Colombia and Peru.

Types:

Holotype female: PERU, Madre de Dios: Avispas, 400 m, X-1/15-1962, L. Pefa

(MC Z).

Paratypes (1 male, 1 female): COLOMBIA, Putumayo: Puerto Asis, VII-4-74, M.

Cooper, ¢ (BMNH). PERU, Cuzco: Quincemil, 750 m, X-16/31-62, ?, L. Pefia (USNM).

Etymology:

The name styphopteron is a combination of the Greek words stypho (= contract)

and pteron (= wing), a reference to the reduction (or contraction) of the submarginal

cells in the forewing of this species.

Cressoni Group

(Figs. 3, 131-252)

The cressoni group, a neotropical endemic, is the largest and most difficult

assemblage of New World Pison; its species are fairly monotonous morphologically,

diagnostic characters are few, and some are variable. Adding to this malaise is the

fact that the male is unknown in four species (erebus, martini, pentafasciatum,

phthinylla), and not positively associated in a fifth (maculipenne). Finally, I have

studied twelve specimens, mostly females, that undoubtedly represent a number of

undescribed species. I am reluctant to attempt description of them because each is

represented by one sex and only one to three specimens. In the cressoni group, a

single female is usually not sufficient to determine if it represents a distinct taxon,

or is simply a variant of a known species. Several species recognized here,

maculipenne, chrysops, and arachniraptor, may contain hidden species. Obviously my

treatment of this section of the genus has to be considered a beginning framework.

Species characters are confined mainly to features of the head (shape of clypeal

margin, ocellocular distance, details of male flagellum, etc.), occasional wing

venational differences, and male genitalia. The descriptions that follow will,

therefore, be rather brief. The following description outlines the morphology

common to nearly all species in the cressoni group.

Description:

Body black but terga I-III often rimmed with creamy yellow bands, legs and/or

body sometimes extensively yellow brown; forewing usually infumate along leading

edge (medial through marginal cells), rest of wing clear to yellowish. Vestiture pale

except sometimes brownish on gaster, mostly appressed, erect setae short and

confined to propodeum and mesopleural venter when present.

Frons, most of thorax, and gaster densely, finely punctate, punctures no more

than diameter apart on thorax, and often nearly contiguous.

Female mandible with inner subapical tooth (fig. 133), male mandible simple

apically. Labrum quadrangular or transverse, free margin straight or arcuate, usually

with median notch. Clypeal free margin with liplike rim that is impunctate, asetose,

and usually shiny; this rim broadest at middle and often projecting as median lobe

that may bear three or four toothlike lobes in female, one or three in male; clypeal

margin thickened or double-edged especially laterally. Antenna not elongate, distal

flagellomeres about as long as broad. Male antenna sometimes with tyli, flagel-

52 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

lomeres sometimes asymmetrically swollen. Frons moderately swollen, dull,

punctures often shallow. Occipital carina ending just short of hypostomal carina, or

narrowly interrupted at midventral line where it is narrowly separated from

hypostomal carina. Pronotum without anterodorsal lamelliform flange, but usually

with oval to broadly transverse anterodorsal pit that is obscured by decumbent setae,

pit rarely absent (abothrum, aranevorax); collar noncarinate, as thick as metanotum

or slightly thinner. Scutal flange nearly as broad as hindocellus, not strongly

upturned; scutum dull or weakly shiny. Scutellar punctation usually of one size,

uniform, punctures mostly diameter apart or less. Tegula entirely punctate or

impunctate on outer half. Propodeum smooth, shiny, finely punctate, without

dorsolateral crenulate ridge between spiracle and petiole socket, sometimes with

weak median longitudinal ridge on dorsum, often with weak cross-ridges on hindface

just above petiole socket. Disk of propleuron impunctate, usually polished.

Episternal sulcus incomplete ventrad, disappearing as it turns toward anteroventral

margin of pleuron. Metapleural flange not lamellate. Forewing media usually

diverging from M+Cu after crossvein cu-a (before cu-a in abothrum, brasilium);

forewing usually with three submarginal cells, the second usually receiving both

recurrent veins (fig. 131); only two submarginal cells in three species (aranevorax,

erebus, phthinylla) due to loss of distal veinlet of the true second submarginal cell

(fig. 132); submarginal cell III often very long, its length often approximately half

that of I and its anterior veinlet often as long as or longer than posterior veinlet of

submarginal cell II (fig. 131). Hamuli of hindwing divided into two groups. Hindcoxa

only with apical remnant of outer dorsal carina, inner carina usually complete;

tarsomeres weakly spinose apicoventrally, plantula present on IV or apparently absent

(abothrum, aranevorax, dementia, phthinylla), claws normal. Gaster dull or shiny,

tergal margins single-edged; male tergum VII blunt apically or conical, male sternum

VIII semicircularly notched apically; genitalia slender, volsellar lobes usually elongate

and fringed apically with setae, inner margin of gonostyle with midventral lobe that

varies from foliaceous to digitiform, apical half of gonostyle usually densely clothed

with long setae ventrally, and with short setae on inner surface, aedeagus usually

slender, without defined head or ventral spine.

Included species:

Pison abothrum, arachniraptor, aranevorax, brasilium, cameronii, chrysops,

cressoni, dementia, erebus, maculipenne, martini, pentafasciatum, phthinylla.

Discussion: )

The most distinctive features of the cressoni group are: the subapically dentate

female mandible, the polished impunctate propleuron, the incomplete episternal

sulcus, the elongate third submarginal cell, the infumiate anterior margin of the

forewing, and the smooth propodeum (no crenulate ridge at top of side). In the New

World fauna the tendency for the terga to be margined by yellow is unique to the

cressoni group. Another attribute of the group is the tendency for the legs and body

to be partially yellowish or reddish brown. Some species appear to lack tarsal

Plantulae: abothrum, aranevorax, dementia and phthinylla. Subapically dentate

mandibles also occur in the krombeini and stangei groups.

Apomorphies of the group include the subapically dentate female mandible, the

incomplete episternal sulcus, the polished impunctate propleuron, the elongate third

submarginal cell, the notched male sternum VIII, and the ventral lobe of the

gonostyle. The yellow tergal bands found in many species, and the loss of the second

submarginal cell in three species (aranevorax, erebus, phthinylla) are apomorphic

trends of the cressoni group, the last character state found elsewhere in Pison.

The male genitalia offer reliable species differences. Those of cressoni and

chrysops are unusual in that the gonostyle is comparatively stouter than in other

species, is sparsely setose ventrally and lacks setae on the inner surface. The

volsellar lobes are unusually stout and sparsely setose in these two species, and they

are probably closely related. The aedeagus most often is acuminate apically in the

cressoni group, but in aranevorax and abothrum it ends in an expanded “head”.

Menke: Pison in the New World 53

The midventral lobe of the gonostyle is least developed in these two species also, but

it is a long, narrow blade-like or finger-like structure in maculipenne, brasilium,

arachniraptor, cressoni and chrysops. Pison maculipenne, brasilium and arachniraptor

seem Closely related based on their genitalic structure.

Pison cressoni Rohwer

(Figs. 131, 133-148)

Pison cressoni Rohwer, 1911:570. Holotype female: Nicaragua, San Antonio USNM).

Pison flavolimbatum Turner, 1917:112. Lectotype female: Guyana, Issororo (BMNH),

present designation. NEW SYNONYMY.

Pison sp., Oniki, 1970a:726. Nest.

Pison sp., Oniki, 1970b:354. Nest, parasite.

Description:

Infumation of forewing brown, uniform through medial, submarginal I and

marginal cells. Head black, except scape usually yellowish brown at least on lower

surface, and frequently scape, pedicel, and flagellomere I (or II in some males)

entirely yellowish brown. Thorax black except foretibia rarely brownish in male.

Gaster entirely black but usually terga I-II or III with apical creamy yellow bands of

variable width. Metallic setae of face, gena, pronotal collar, and propodeal dorsum

usually golden (lower facial setae sometimes silver, propodeal setae rarely brownish).

Female clypeal margin with short median lobe whose free edge is sinuate and

reflexed upward (surface of median lobe with transverse crease, fig. 136), sinuation

delimiting three or four weak angles or more protuberant teeth (depending on

whether or not middle angle or tooth is emarginate resulting in two points)(figs.

137-138), lower margin of lateral tooth continued laterad as carina delimiting broadly

thickened lateral part of clypeus (ventral view, fig. 135); male clypeus with three

prominent teeth at middle that are not reflexed, middle tooth slender (Central

America and Santa Catarina, Brasil, fig. 139) or broad basally and blunt apically

(Colombia, Trinidad to Peru and Sao Paulo, Brasil, fig. 140), margin of outer tooth

extended laterad as carina just as in female but delimiting broader area (fig. 141).

Male flagellomeres without tyli, not asymmetrically swollen. UID in female

0.67-0.75X LID; UID in male 0.77-0.84X LID. OOD in female usually 0.72-0.92X

HOD, but 1.0-1.15X HOD in largest specimens; OOD in male 1.0-1.12X HOD.

Occipital carina ending just as it reaches apex of hypostomal carina. Anterodorsal

pit of pronotum deep, usually smaller than hindocellus but sometimes transversely

elongate. Tegula punctate only on inner half. Punctation of scutellum usually

conspicuously sparser on posterior one-half to one-third, interspaces shiny, some

micropunctures irregularly mixed with macropunctures in this area (fig. 142), these

micropunctures (actually pores - see fig. 143) present even when macropunctation is

fairly uniform over entire scutellum (in some males from Central America punctation

is fairly uniform and micropunctures very scarce). Male sternum VIII constricted

before apex, apex rounded lateral to emargination (fig. 148). Genitalia as in figs.

144-147, midventral lobe of gonostyle long, slender, curving inward in ventral view

(fig. 146).

Length 8-12.5 mm (female), 8-10 mm (male).

Discussion:

Pison cressoni, a commonly collected species, is recognized by the reflexed lobe

of the female clypeus (fig. 136), the broad ocellocular distance (0.72-1.15X HOD), the

sparsely bipunctate posterior part of the scutellum (fig. 142), the shape of male

sternum VIII (fig. 148) and the genitalia (figs. 144-147). Within the cressoni group,

the bipunctate scutellum is unique to this species. The “micropunctures" are actually

pores (fig. 143) but at 50X magnification they look like punctures. These pores are

often clustered in groups of 2 to 6. The genitalia of cressoni only resemble those of

chrysops, and the two species are probably more closely related to each other than to

54 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

others in the group. Male sternum VIII is constricted before the apex in both species,

unlike others in the cressoni group. The midventral lobe of the gonostyle is elongate

in both species, but the shape of sternum VIII and the volsellar lobes differ between

them (compare figs. 144-148 & 155-159).

The variation in cressoni is vexing. Males from Central America and also the

southernmost part of the species' range (Santa Catarina, Brasil) have narrow,

fingerlike clypeal “teeth", while elsewhere the teeth are shorter, and the middle one

is broad at the base (compare figs. 139-140). The male genitalia are uniform over the

range of the species however. Some variation in the outline of the female clypeal

lobe is also evident (figs. 133-134), but the reflexed lip is a fairly constant feature.

In specimens with a weakly reflexed lip, the OOD:HOD and/or scutellar punctation

generally provide positive identification. The punctation of the scutellum varies

from sparse on the posterior half to nearly uniform over the entire surface. But in

every case a few micropunctures are visible (50X magnification) on the posterior half.

Pison cressoni resembles maculipenne, another commonly collected species, and

the two are largely sympatric. Pison maculipenne differs in having a non-reflexed

(flat) clypeal lobe surface in the female, a narrower ocellocular distance (OOD in

female 0.43-0.65X hindocellus diameter except up to 0O.75X HOD in largest

specimens), a narrower UID (UID in female 0.57-0.66X LID), a uniformly

monopunctate scutellum, a differently shaped male sternum VIII, and the genitalia.

Furthermore, the wings of maculipenne are often amber colored, and the body

sometimes has yellowish brown areas that are often extensive. In particular,

laterotergite I of maculipenne is usually yellowish brown, while in cressoni it is

black. The occipital carina of maculipenne is a complete circle that is tangential

with the hypostomal carina or narrowly separated from it. In cressoni the occipital

carina ends just before reaching the hypostomal carina.

I have examined the type of cressoni. Rohwer indicated that his specimens were

males but in fact the holotype and paratype are females. I have seen two of Turner's

three syntypes of flavolimbatum, and selected one as lectotype.

Range:

Southern Mexico to Trinidad, Peru and Santa Catarina, Brasil. Apparently

widespread in South America.

Material examined (96 females, 35 males):

MEXICO, Veracruz: UNAM reserve, 32 km n Catemaco, I-10-82, J. W. Wenzel

(USNM). BELIZE: Toledo, Punto Gorda (BMNH). GUATEMALA: E1 Salto, Escuintla

(BPBM); Sa. Emilia, Pochuta (MCZ). HONDURAS: nr. Tegucigalpa, emerged from

nest of Eutachyptera psidii (Lasiocampidae)((USNM). NICARAGUA: San Antonio

(USNM). COSTA RICA: Golfito (MCZ). PANAMA: Barro Colorado I., Phil Rau note

#7812 (USNM, AMNH). COLOMBIA, Arauca: Tame (BMNH); Valle: Cali (MCZ);

Vaupes: Mitu, M. Cooper note 85 (BMNH); Magdalena: N. Sierra Nevada de S. Marta,

Rio Buritaca (BMNH); Becerril, Quebrada Socorpas (USNM); Caqueta: Yuruyaco, 73

km sw Florencia (BMNH); Putumayo: Mocoa (BMNH). ECUADOR, Napo: Limoncocha

(USNM); Pichincha: Nanegal (COOPER); Morona-Santiago: Cord. de Cutucu, 6 km e

Macas (COOPER). PERU, Junin: Valle Chanchamayo (LILLO); Huanuco: Cueva de

Las Pavas, nr. Tingo Maria (LILLO); Monson Valley, Tingo Maria (CAS); 67 km e Tingo

Maria (CAS); Tingo Maria (CSDA, MCZ, USNM, AMNH); Loreto: Pucallpa (USNM).

TRINIDAD: Maracas Valley, from mud cells (USNM, BMNH). GUYANA: Bartica

(CU); Waratuk (AMNH); Demerara River (CU); Issororo (BMNH); North West District

(BMNH). SURINAME: Raleigh-Vallen (USNM); Paramaribo (LEIDEN); Brownsberg

(PMA). BRASIL, Para: Belém (UCD, VIENNA); nr. Belém, ex cells in bird nests

(Phaethornis ruber and Tolmomyias flaviventris) (USNM); Federal District: Brasilia

(RAW, BMNH, DIAS; Minas Gerais: Juiz de Fora (RAW); Lassance (CU); env.

Passa-Quatro (PARIS); Lagoa Santa (HALLE). Sao Paulo: no location (USNM); no

location, ex nest of Phaethornis squalidus (USNM); Santa Catarina: Blumenau

(VIENNA).

Menke: Pison in the New World 55

Biology:

Oniki (1970a, b) discovered the cells of a Pison sp. inside the lower part of nests

of the hummingbird, Phaethornis ruber. Voucher material in the USNM has enabled

me to identify the wasp as cressoni. Oniki examined the nests of three other

hummingbird species in her Belém study area, but none contained cells of Pison,

suggesting that cressoni prefers P. ruber nests, at least at that location. Eight out of

13 bird nests examined by Oniki contained cressoni cells. Such nests had a small

entrance near the bottom and contained up to 13 mud cells of the wasp. The opening

of a cell faced downward and three to five spiders were provisioned. Besides rearing

examples of cressoni from one such nest, Oniki obtained a female of the genus

Brachymeria, possibly a parasitoid of the wasp although this chalcidid genus does not

commonly attack sphecids. She suggested that possibly more than one Pison nest

occurred in some of the bird nests.

Oniki subsequently obtained specimens of Pison cressoni from mud cells in the

nest of the flycatcher Tolmomyias flaviventris (Tyrannidae) near Belém (voucher

material in USNM), but she has not published details as yet. A chrysidid of the genus

Trichrysis determined as "species 6" (USNM) by R. M. Bohart emerged from some of

the cells. The USNM collection also has an old Brasilian specimen of cressoni with

the notation "Nester im Neste von Phaethornis squalidus”.

Pison cressoni makes nests in other sheltered situations. The material from

Veracruz, Mexico, was reared from mud cells that were built inside an abandoned

nest of the vespid genus Polybia (Wenzel, in litt.). The single specimen listed above

from Honduras emerged from the larval tent of the lasiocampid moth Eutachyptera

psidii. Notes by Martin Cooper on a specimen of cressoni from Mitu, Colombia (85)

say "seen entering a rolled-up mosquito net suspended from ceiling of a room. When

net opened remains of very thin and cylindrical mud cell found attached to netting."

Mud cells of cressoni from Barro Colorado I., Panama, collected by Zetek (AMNH)

have small, wool-like fibers imbedded in them suggesting that they were in some kind

of nest. I have also studied material of cressoni from Trinidad that Callan reared

from mud cells without annotation as to substrate.

Pison chrysops Menke, n. sp.

(Figs. 149-159)

Description, holotype female:

Infumation of forewing leading edge uniformly brown from apical half of medial

cell through apex of marginal cell, wing membrane practically clear. Body black

except as follows: scape obscurely brownish; palpi yellowish brown; apical half of

mandible reddish brown except apex dark; pronotal lobe, tegula, scutal flange, femora

beneath, and laterotergites I-II reddish brown. Clypeus, lower frons and gena with

dense appressed silver setae; frontal setae changing to gold at level of eye notch and

continuing narrowly dorsad along orbit to level of ocelli; pronotal collar, hind margin

of scutum, metanotum, and propodeum (except middle of hindface and side adjacent

to metapleuron) densely covered with appressed gold setae; meso- and metapleuron

sparsely covered with appressed silver setae. Terga I-III with broad, creamy yellow

bands apically, sternum II with narrow apical creamy yellow band.

Clypeal margin without median lobe but smooth polished rim broadened at

middle, upper edge of rim not elevated above adjacent clypeal punctation; clypeal

edge with four weak angles, the middle pair of which surround cuplike pit in ventral

view (similar to figs. 151-152), the margin between these two middle angles slightly

reflexed upward; in ventral view margin thin at middle (except for pit between

middle angles), but moderately thickened laterad of angles (similar to fig. 152). UID

0./2X LID. OOD 0.96X HOD. Occipital carina incomplete ventrally, ending near

apex of hypostomal carina. Pronotum with transversely elongate anterodorsal pit.

Tegula impunctate on outer half. Scutellar punctures separated by less than diameter

to two diameters. Media of forewing diverging from M+Cu just after crossvein cu-a.

Hindtarsomere IV with plantula.

Length 7.5 mm.

56 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

Variation in female (32 specimens):

Scape occasionally all black; legs rarely all black (Guyana); scape, pedicel, and

flagellomere I yellow brown (Peru; Salta, Argentina); clypeal margin reddish brown

(Colombia; Salta, Argentina). Anterior half of propodeal side or even entire side

sometimes without dense, appressed gold setae, and such setae rarely restricted to

disk of dorsum and interface between hindface and side (Tena, Ecuador; Guyana;

Belém, Brasil). Tergum III sometimes without yellow band (Santa Catarina, Brasil), or

band very thin (Santa Catarina, Brasil; n Argentina); sternum II usually without yellow

band. Clypeal margin variable (figs. 149-151), lateral angles sometimes weak or

absent, middle pair sometimes strongly reflexed. UID 0.65-0.79X LID. OOD

0.82-1.16X% HOD, except in two extreme specimens: 0.55X (Ecuador, 60 km s

Guayaquil) and 0.60X (Colombia, Barbacoas). There is a strong tendency for the UID

and OOD to become proportionally greater toward the southern part of the range (s

Brasil, n Argentina). Media of forewing usually diverging well beyond crossvein cu-a,

the distance equal to about one-third of cu—a length. Length 7-8 mm.

Male (7 specimens):

Differs from female as follows: scape pedicel, flagellomeres I-IV, labrum, and

mouthparts sometimes yellow brown (Bolivia); foreleg reddish brown (Bolivia).

Appressed setae on gena gold and on propodeal side silver (Bolivia), appressed

mesopleural setae silver (El Limon, Venezuela). Sternum II without yellow band.

Clypeal margin with three teeth, edge thickened in ventral view (figs. 153-154).

Flagellum simple. UID 0.78-0.84X LID. OOD 0.93-1.18% HOD. Media of forewing

sometimes interstitial with crossvein cu-a (Palmar, Costa Rica). Sternum VIII

constricted before apex (fig. 159). Genitalia as in figs. 155-158; midventral lobe of

gonostyle long, slender, the tip usually incurved in ventral view (fig. 157). Length 7-8

mm.

Discussion:

Pison chrysops is one of the more commonly collected and wide ranging

neotropical species. The dense covering of gold setae on the propodeum is the most

conspicuous recognition feature, although the propodeum of some cameronii and

arachniraptor is partially covered by gold setae. The broader ocellocular distance of

chrysops usually separates it from these two species however, and the female clypei

differ (compare figs. 149-151, 186-188, & 204-210). The female clypeus is variable

but its thin margin and lack of a median lobe are fairly diagnostic for separation from

simple flagellum, clypeal teeth, sternum VIII, and genitalia are diagnostic in

combination. The genitalia and sternum VIII are most similar to cressoni. The

uniformly punctate scutellum of chrysops distinguishes it from cressoni.

Two female specimens are rather disconcerting. Both have an unusually narrow

ocellocular distance: OOD is 0.60% HOD in a specimen from Barbacoas, Colombia

(COOPER) and 0.55X HOD in a specimen from 60 km s Guayaquil, Ecuador (MCZ). I

have not included these in the paratype series although I am fairly certain that they

are conspecific with chrysops.

Range:

Pison chrysops is known from Costa Rica to northern Argentina and southern

Brasil.

Types:

Holotype female: COSTA RICA, Punta Arenas: Palmar, II-1950, D.O. Allen

(USNM).

Paratypes (30 females, 7 males): COSTA RICA, Guanacaste: I-22/23-77, D. H.

Janzen (AEI); Punta Arenas: Golfito, XII-1948, P. & D. Allen (MCZ), Esquinas nr.

Golfito, 1948, P. & D. Allen (MCZ), Palmar, V/VI-1950, D. O. Allen (USNM).

PANAMA, Canal Zone: Barro Colorado I., VI-14-39, Zetek (USNM). VENEZUELA,

Araqua: Rancho Grande, V-12-46, W. Beebe (AMNH), El Limon, 480 m, V-21/22-73,

C. J. Rosales (UCM); Distrito Federal: La Guaira, VII-4-1900, Lyon and Robinson

Menke: Pison in th New World 57

(USNM). COLOMBIA, Magdalena: Rio Frio, X-15-27, G. Salt (BMNH); Meta:

Restrepo, 500 m, VIII-1936, J. Bequaert (MCZ). PERU: Diversos, P. Weiss (USNM).

ECUADOR, Napo: Tena, 500 m, IV-11/28-76, M. Cooper (BMNH). BOLIVIA, Beni: Rio

Itenez at mouth of Rio Baures, IX-30-64, J. Bouseman (AMNH). GUYANA: Rupununi

Savannah, II-7-34, J. Meyers (USNM). SURINAME: Paramaribo, I-7-58, P. van

Doesburg (LEIDEN). BRASIL, Para: Belém, VI-1903, A. Ducke (VIENNA); Sao Paulo:

Campinas, II-1924, F. X. Williams (BPBM); Santa Catarina: Nova Teutonia, various

dates, Fritz Plaumann (UCD, MCZ, OSU, BMNH). ARGENTINA, Salta: Rio Pescado

nr. Oran, IX-24-71, C. Porter (LILLO), Pocitos, I-1971, M. Fritz (FRITZ); Entre Rios:

Pronunciamiento, I-1/5-65 (LECLERCQ).

Metatypes (2 females): COLOMBIA, Narifo: Barbacoas, 50 m, VIII-16-84, M.

Cooper (COOPER). ECUADOR: Balao Chico, 60 km s Guayaquil, IV-23/29-63, L.

Pena (MCZ).

Etymology:

The name chrysops is a Greek word meaning gold-colored, in this case a

reference to the dense covering of gold setae on the propodeum.

Pison erebus Menke, n. sp.

(Fig. 160)

Description, holotype female:

Infumation of forewing brown and uniform through medial, submarginal I and

marginal cells. Body black except scape, pedicel and flagellomere I reddish brown.

Clypeus, lower frons, gena, and dorsum of collar with dense appressed gold setae.

Clypeal margin with short median lobe whose free edge is broadly concave,

surface of lobe with semicircular impression (fig. 160), edge thickened laterally as in

cressoni (ventral view) and with median dimple. UID 0.72X LID. OOD 0.71X HOD.

Occipital carina narrowly interrupted ventrad, its ends tangential with hypostomal

carina apex. Anterodorsal pit of pronotum transversely oval, length equal to

hindocellus diameter. Tegula uniformly punctate. Scutellum uniformly punctate,

punctures of one size. Forewing with two submarginal cells through loss of outer

veinlet of true second submarginal cell.

Length 8 mm.

Male:

Unknown.

Discussion:

Pison erebus is one of three species in the cressoni group that have only two

submarginal cells. It differs from phthinylla and aranevorax in having a broader

OOD, and a distinctive clypeus (fig. 160). The reddish brown antennal base and

golden haired clypeus and lower frons contrast distinctively with the black body, but

they may prove to be an unreliable features when more material is available. Pison

erebus resembles cressoni but the uniform scutellar punctation, wing venation and

absence of yellow tergal bands separate it from the latter species.

Range:

Known only from the holotype collected in Colombia.

Type:

Holotype female: COLOMBIA, Caqueta: Yuruyaco, 73 km sw Florencia,

II-8-1979, M. Cooper (BMNH).

Etymology:

Erebus, a noun in apposition, is the name of the Roman god of darkness, a

reference to the predominant black color of the species.

58 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

Pison pentafasciatum Menke, n. sp.

(Fig. 161)

Description, holotype female:

Infumation of forewing yellowish brown, darkest on submarginal cell I and

marginal cell, rest of wing clear with yellowish brown veins. Body black except for

broad yellow apical bands on terga I-V (as wide as metanotum), those on III-IV

bisinuate on basal side; sterna II-IV with narrower apical yellow bands; laterotergites

I-II brownish translucent. Head without obvious metallic setae, surface sculpture of

face not obscured.

Clypeal margin with short median lobe (fig. 161), its free edge with blunt median

tooth whose apex is dimple-like in ventral view, edge of lobe not thickened in ventral

view, but clypeal margin lateral to it thickened. UID 0.77X LID. OOD 1.23X HOD.

Occipital carina incomplete ventrally, ending well before midline of head.

Anterodorsal pit of pronotum transversely elongate, length greater than hindocellus

diameter. Tegula impunctate on outer half. Scutellum broadly convex.

Length 9 mm.

Male:

Unknown.

Discussion:

Pison pentafasciatum is easily recognized by the broad yellow bands on the first

five terga, the very broad ocellocular distance, and the absence of appressed metallic

facial setae. The clypeus is also fairly distinctive (fig. 161).

Range:

Known only by the holotype from southeastern Brasil.

Type:

Holotype female: BRASIL, Santa Catarina: Cauna, XII-1945, A. Maller (UCD).

Etymology:

The name pentafasciatum is a combination of the Greek word penta (= five) and

the Latin fascia (= band) and refers to the yellow abdominal bands.

Pison martini Menke, n. sp.

(Figs. 162-165)

Description, holotype female:

Infumation of forewing leading edge brown, darkest on marginal cell, wing

membrane pale brownish behind. Following body parts yellowish brown: scape,

pedicel, flagellomere I, labrum, palpi, pronotal lobe, and legs except coxae and

dorsum of hindfemur; following are reddish brown: clypeal lobe, mandible (dark at

tip), rim of petiole socket, and laterotergite I; gaster without creamy yellow tergal

bands. Following with dense, appressed gold setae: clypeus, lower frons and narrowly

along inner orbit to level of ocelli, gena, and top of pronotal collar; setation of rest

of thorax including legs paler, longest on propodeum; gaster with decumbent brownish

setae.

Clypeal margin with short median lobe whose edge is slightly trisinuate in frontal

view, surface of lobe smooth, flat (similar to fig. 163); edge of clypeus slightly

thickened as seen in ventral view, hind edge of lobe sinuate (similar to fig. 165). UID

0.60X LID. OOD 0.64X HOD. Occipital carina incomplete ventrad, ending near apex

of hypostomal carina. Anterodorsal pit of pronotum deep, transversely elongate,

slightly longer than hindocellus diameter. Tegula with punctures near outer edge as

well as on inner half, small discal area impunctate. Propodeal dorsum with fine

longitudinal carina at base. Hindtarsomere IV with tiny plantula. Thorax and gaster

Menke: Pison in the New World D9

elongate, tergum I length (measured in dorsal view) 1.22X apical width. Tergum II

swollen in lateral profile in contrast to adjoining terga (fig. 162).

Length ll mm.

Female variation (3 specimens):

Flagellomere I sometimes black; hindfemur sometimes entirely yellowish; UID

0.57-0.62X LID; OOD 0.60-0.64X HOD; clypeal lobe truncate in two specimens (fig.

164); fine carina of propodeal dorsum extends to the latters apex in one specimen;

length to 12 mm.

Male:

Unknown.

Discussion:

Pison martini, the largest species of the cressoni group, has a distinctive,

essentially truncate, clypeal lobe (figs. 163-164) although it is similar to that of

maculipenne. The body is noticibly elongate in comparison to stocky species such as

cressoni and maculipenne, but this is rather difficult to quantify. I have measured

the length of tergum I as seen in dorsal view and contrasted that with its dorsal

width. The tergum of martini is clearly longer than wide, while in cressoni and

maculipenne tergum I is about as long as wide or slightly shorter than wide. The

humped tergum II of martini is distinctive (fig. 162) but may prove to be variable

when more material is available. Pison phthinylla, a species with only two

submarginal cells (martini has 3), has a similarly humped tergum II. The color pattern

of martini (yellowish legs, essentially all black gaster, golden appressed setae on

head, and somewhat paler setae on rest of body) is distinctive too, but that may also

prove to vary.

Range:

Known only from one locality on the amazonian side of Ecuador.

Types: |

Holotype female: ECUADOR, Morona-Santiago: Macas, 1,100 m, II-1-1982, M.

Cooper (BMNH).

Paratypes (3 females): two with same data as type except II-2-82 and II-9-84

(COOPER, USNM); one from Cord. Cutucu, about 6 km e Macas, about 1000 m,

X-25-78 (BMNH#H).

Etymology:

The name martini is a dedication to Martin Cooper who has probably collected

more neotropical material and species of Pison than any other person. Without his

material this revision would have been far less complete.

Pison maculipenne Smith

(Figs. 166-176)

Pison maculipenne Smith, 1860:80. Holotype female: Brasil, "Ega" (= Tefé) (BMNH).

Description:

Infumation of forewing variable: dark or light brown and uniform through medial,

submarginal and marginal cells, or absent from submarginal cell or restricted to

marginal cell; wing membrane often yellowish. Head usually entirely black except

scape often brownish beneath; clypeus, mandible and mouthparts largely yellowish

brown in occasional specimens (Ecuador and along Amazon River); scape, pedicel and

flagellomere I, or I-II or III yellowish in same material. Thorax usually black except

pronotal lobe, tegula, dorsum of petiole socket and closing face of femora sometimes

paler; upper surface of tibiae occasionally also pale; tarsomeres I in some specimens

(including both males) largely white or yellowish; thorax, including legs, extensively

60 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

yellowish brown in some Ecuadorian material (Tena) with black restricted to scutum

(sometimes partly pale), scutellum, metanotum, propodeal dorsum and _ side,

mesopleural venter (sometimes partly pale), and base of mid and hindcoxae; thorax

even more yellowish brown in some Brasilian specimens (holotype, Tefé) and (Para):

black restricted to metanotum and propodeal side. Gaster usually black except

laterotergite of tergum I yellowish or reddish, apex of tergum I and sometimes II

translucent or with creamy yellow band; reddish tint of gaster sometimes more

extensive: segment I largely pale (some Colombian and Ecuadorian material), segment

I and tergum II occasionally extensively or completely pale (Trinidad, some

Colombian and Ecuadorian material, holotype from Tefé), abdomen extensively pale

in one specimen from Tena (terga II-VI and sterna II-V black only basally) and one

from Para (only small dark area on tergum III). Metallic setae of clypeus, gena

(typically sparse) and elsewhere usually silver, golden only on specimens that have

extensively yellowish bodies, and then entire head often densely gilded.

Female clypeal margin with short median lobe whose free edge is typically

weakly sinuate (fig. 166) or even just straight, sinuations sometimes forming three or

four “teeth” (especially strong in s Brasil, figs. 168-169), lobe not reflexed (polished

surface flat and usually raised above adjacent punctation), free margin of clypeus

lateral to lobe narrowly thickened (ventral view, fig. 167); male clypeus with three

prominent teeth (fig. 170), margin of outer tooth sloping laterad beneath and behind

Clypeal margin forming broad, thick edge in ventral view (fig. 171). Male flagellum

simple. UID 0.57-0.66X LID in female; 0.69-0.71X LID in male. OOD in female

usually 0.43-0.65X HOD, but up to 0.75X HOD in largest specimens; OOD 0.72X HOD

in male. Occipital carina complete, tangential with or slightly separated from apex

of hypostomal carina. Anterodorsal pit of pronotum transversely elongate, as long or

longer than hindocellus diameter. Tegula sometimes completely punctate although

sparsely so on outer half. Propodeal dorsum sometimes with fine, longitudinal carina

at base. Male sternum VIII as in fig. 176. Genitalia as in figs. 172-175, midventral

lobe of gonostyle elongate, its apical half dorsoventrally compressed (fig. 172).

Length 8.5-11.5 mm in female, 8.5 mm in male.

Discussion:

Pison maculipenne, a commonly collected wasp, is a puzzling species for two

reasons. Over its range it displays considerable color variation, some of which does

not seem to follow any particular pattern. The other problem concerns the

identification of the male; I have three possible candidates and I am not positive that

I have selected the correct one.

The wing membrane color alone (whether it is clear or yellowish) suggests at

first that two species are involved. For example, both types occur on Barro Colorado

I. in Panama. However, I have been unable to find any other basis for separating the

two color forms as separate species. Perhaps this wing color variation is

microenvironmentally induced. MacLean, Chandler and MacLean (1978) demonstrated

that color variation in Polistes was highly correlated with nest microclimate (relative

humidity-temperature). Another color variable is the striking xanthochroism of a

few specimens in Ecuador and Brasil, including the holotype. Intermediate stages

between the common, essentially black type to the extreme yellowish forms, suggests

that a broad spectrum of color morphs probably exist. Much more collecting will be

needed before the extent and significance of this xanthochroism can be analyzed.

Selecting the male of maculipenne from three possible candidates has been

frustrating. Head measurements, configuration of the occipital carina, and clypeal

details do not offer a means of correlating one of the three with the female, and I

have resorted to wing membrane color alone. Collecting locality information is of no

use since the three male species are sympatric with the female distribution, some

having been taken at the same locales as some of the females. The males differ in

antennal structure. The flagellum is simple in one male species (Canal Zone, Panama

- PMA, Venezuela - USNM); flagellomeres II-VI have raised linear tyli in another

(Colombia - BMNH); and in the third flagellomeres III-VII are asymmetrically swollen

and associated with them are narrow to welt-like tyli (Peru - MCZ; Colombia -

Menke: Pison in the New World 61

Cooper; Brasil - AMNH). The genitalia of the three male species are fairly similar.

In each the tip of the midventral lobe of the gonostyle is paper-thin in side view. In

ventral view these paired appendages diverge in two of the male species, in the third

(the male with antennal tyli) they curve outward resembling the horns of a cow.

Minor differences in the volsellar lobes have been noted. There are slight differences

in the clypeus among the three male species. The two males from Panama and

Venezuela have a yellowish wing membrane, a characteristic of many females of

maculipenne, and a feature unknown in other species of the cressoni group. On that

basis I have treated these two specimens as the male of the species.

The male enigma poses one obvious question: does my treatment of maculipenne

possibly represent a complex of two or three extremely similar species, the males of

which are represented by the three types discussed above? Only rearing of nestmates

can resolve this conundrum.

I have examined two possible “types” of maculipenne from the BMNH, one from

"Ega" and one from “Para", but only the former agrees perfectly with Smith's

description. The color pattern of the Para specimen does not agree with the original

description, and this locality was not listed by him. Smith does not suggest that he

had more than one specimen, and I consider the wasp from Ega as the holotype.

Pison maculipenne is similar to cressoni and differences are discussed under that

species. Pison martini is perhaps the closest relative of maculipenne. The female

clypeus is similar in both (compare figs. 166-169 & 163-165). Pison maculipenne has

a normal tergum II, while in martini it is humped (fig. 162). Tergum I is about as long

as wide in maculipenne, but longer than wide in martini. Unfortunately the male of

martini is unknown. On the basis of the form of the midventral lobe of the gonostyle,

and other aspects of the male genitalia, maculipenne, brasilium, arachniraptor, and

possibly cameronii, are closely related.

Range:

Costa Rica to Bolivia, eastward to Trinidad and southward to southern Brasil.

Material examined (60 females, 2 males):

COSTA RICA: Golfito (UCD). PANAMA, Canal Zone: Barro Colorado I. (AMNH,

USNM, PMA). COLOMBIA, Narino: Barbacoas (BMNH, COOPER); Caqueta: Yuruyaco,

73 km sw Florencia (BMNH); Meta: La Macarena (BMNH), Cordillera Macarena

(BMNH); Putumayo: Mocoa, 550 m (BMNH, COOPER), Villa Garzon, 400 m (BMNH,

COOPER), Puerto Leguizamo, 180 m (BMNH); Cauca: Napoles, 350 m, 5 km e Puerto

Limon (COOPER). ECUADOR, Napo: Muyuna, 500 m, 5 km w Tena, (biol. note 33,

"on leaf with prey, a spider, in primary forest") (BMNH, COOPER), Tena (BPBM).

PERU, Loreto: Pucallpa (BMNH); province?: El Campamiento, Col. Perone (CU).

BOLIVIA, Beni: Rio Mamore, 10 km e San Antonio (AMNH). VENEZUELA, Bolivar: E]

Bochinche Res. Forestal Imataca, 200m (UCM); Zulia: Los Angeles del Tucuco

(USNM). TRINIDAD: Arima Valley, 800-1200 feet (AMNH). GUYANA: Rupununi

Savannah (BMNH). SURINAME: (VIENNA). BRASIL, Para: no locality (VIENNA,

BMNH); Amazonas: Tefé (BMNH); Minas Gerais: Pedra Azul (AEI); Guanabara: Rio de

Janeiro (USNM).

Pison brasilium Menke, n. sp.

(Figs. 177-185)

Description, holotype male:

Forewing without discrete dark infumation along leading edge, membrane weakly

brownish. Body black except as follows: mandible center, tegula and laterotergite I

reddish brown. Lower frons, clypeus, gena, top of pronotal collar, and metanotum

laterally with dense, appressed silver setae; propodeal dorsum and interface between

side and hindface with dense silver setae that do not obscure sculpture. Terga I-III

with broad creamy yellow apical bands.

Clypeal margin with three similar teeth, margin sinuate laterad (similar to fig.

179), edge thickened in ventral view (similar to fig. 180); polished rim not elevated

62 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

above clypeal punctation. Flagellomeres III-VII with polished, linear tyli beneath,

those on III and VII short. UID 0.98X LID. OOD 1.71X HOD. Occipital carina

incomplete ventrally, ending short of hypostomal carina apex. Pronotum with

transversely elongate anterodorsal pit. Tegula impunctate on outer half. Propodeal

hindface transversely rugosopunctate just above socket, changing to coarse, dense

transverse striatopunctation toward dorsum, punctures coarse, essentially contiguous;

punctation of propodeal side dense, punctures less than a diameter apart. Media of

forewing diverging from M+Cu before crossvein cu-a. Hindtarsomere IV with tiny

plantula. Sternum VIII similar to maculipenne. Genitalia as in figs. 181-185;

midventral lobe of gonostyle long, narrow, laterally compressed (fig. 184), attenuate

apically.

Length 6.25 mm.

Variation in males (3 specimens):

Forewing sometimes strongly infumate along leading edge; laterotergite I black;

tyli present on flagellomeres III-VIII (short, weak on III and VIII), or IV-VI, or IV-VII;

UID 0.92-0.94X LID; OOD 1.60-1.65X HOD; forewing media interstitial with crossvein

Cu-a in one specimen; length to 7 mm.

Female (3 specimens):

Body black except mandible and tegula obscurely reddish brown; clypeal margin

without median lobe but with bidentate, slightly reflexed snout (fig. 177), sometimes

small angulation present laterad, polished rim arcuately broadened above, clypeal

edge thin in ventral view (fig. 178); UID 0.87-0.90X LID; OOD 1.35-1.57X HOD;

forewing media occasionally interstitial with crossvein cu-a; length 7-8 mm.

Discussion:

The very broad upper interocular distance (UID nearly as broad as LID) and

ocellocular distance (OOD more than 1.33X HOD) are distinctive features of

brasilium, although dementia and pentafasciatum have similar proportions. The

divergence of the forewing media before crossvein cu-a in most specimens of

brasilium is also unusual in the cressoni group (abothrum is the only other species

with this feature). The presence of linear tyli on the male flagellum, and the dense,

coarse punctation of the propodeal side and hindface, with cross-ridging on the last,

are also distinctive. The genitalia are similar to maculipenne but the midventral lobe

of the gonostyle is laterally compressed (dorsoventrally compressed in maculipenne).

Range:

Known only from the Federal District of Brasil.

Types:

Holotype male: BRASIL, Federal District: Brasilia, Lago Sul, IV-7-1977, A. Raw

(BMNH).

Paratypes (3 males, 3 females): Federal District: Reserva Ecologica do IBGE, 20

km s Brasilia, various dates, malaise trap (IBGE).

Etymology:

The name brasilium is based on the type locality, Brasilia.

Pison arachniraptor Menke, n. sp.

(Figs. 3, 186-203)

Description, holotype female:

Infumation of forewing leading edge light brown, wing membrane clear. Body

black except as follows: pronotal lobe, tegula, and laterotergite I-II reddish brown.

Lower frons with dense, appressed silver setae that are continued dorsad along inner

orbit to level of ocelli, setae with slight golden tinge dorsad; clypeus not densely

Menke: Pison in the New World 63

silvered except at extreme lateral corner; gena with dense, appressed slightly golden

setae; dorsum of collar with appressed slightly golden setae, rest of thoracic setae

similarly colored or paler, but not dense except at base of propodeal dorsum and

above hindcoxa. Terga I-III with creamy yellow bands, that on III narrow.

Clypeal margin without median lobe, but with concavity as wide as labrum that

bears four angles (similar to fig. 186); margin of concavity thickened in ventral view,

middle pair of angles delimiting pit (similar to fig. 187); clypeal rim around concavity

sharply elevated above adjacent clypeal punctation (similar to fig. 186). UID 0.63xX

LID. OOD 0.36-.38X HOD (right and left OOD different). Occipital carina narrowly

incomplete ventrad, the gap not tangential with apex of hypostomal carina.

Pronotum with transversely elongate anterodorsal pit. Tegula narrowly impunctate

laterad. Media of forewing diverging from M+Cu just after crossvein cu-a.

Hindtarsomere IV with plantula.

Length 6.5 mm.

Variation in female (29 specimens):

Infumation of forewing sometimes dark brown, and membrane sometimes lightly

brownish; scape often yellowish brown, rarely scape, pedicel and flagellomere I

(sometimes base of II) rarely yellowish or reddish brown (Ecuador; Mato Grosso,

Brasil); legs except coxae, trochanters, sometimes partly brownish. Appressed facial

setae sometimes entirely golden (Ecuador). Terga I-III sometimes translucent

apically rather than yellow banded, yellow bands sometimes only on I-II. Shape of

clypeal concavity variable (figs. 189, 191-194, 196), edge often thinner in larger

specimens (figs. 190, 195). UID 0.62-0.78X LID (UID proportially greater in larger

specimens). OOD 0.38-0.66X HOD except 0.70-0.82X HOD in largest specimens.

Ends of occipital carina sometimes tangential with apex of hypostomal carina.

Tegula sometimes punctate along outer margin but with small impunctate discal

zone. Second recurrent vein occasionally interstitial between submarginal cells

II-III. Length 6.5-8 mm.

Male (2 specimens):

As in female except: appressed facial and thoracic setae silver in one specimen;

hindtarsomere I whitish in one specimen; terga I-II with broad creamy yellow bands,

III with weak band; clypeal margin with three blunt teeth, middle one broadest (fig.

197), edge thickened in ventral view (fig. 198), polished rim sharply elevated above

Cclypeal punctation (does not show in SEM photo, fig. 197); flagellum simple; UID

0.78-0.79X LID; OOD 0.70-0.77X HOD; second recurrent vein interstitial between

submarginal cells II-III in one specimen; sternum VIII similar to maculipenne;

genitalia as in figs. 199-203, apex of midventral lobe of gonostyle spatulate, twisted

(figs. 202-203); length 7 mm.

Discussion:

The polished clypeal rim is elevated above the punctate part of the clypeus (figs.

186, 188, 196) in both sexes, a unique attribute of arachniraptor. The female clypeus

typically has a broad shallow concavity containing four “teeth” (fig. 188) that is

distinctive, but some metatypes vary widely and may not be conspecific (figs. 191,

193, 196). Several metatypes (indicated with an asterisk) may prove to be chrysops, a

similar species. Pison arachniraptor has gold setae on the propodeum and

occasionally it is dense as in chrysops. The ocellocular distance is narrower in

arachniraptor than in chrysops, and the female clypei differ (compare figs. 186, 192,

194 & 149-151). The twisted, spatulate midventral lobe of the gonostyle is

distinctive in the male (fig. 202).

The clypeal margin is less thickened in some of the largest females (figs. 190,

195). These and a few other atypical females have not been included in the paratype

series.

Range:

Panama to Bolivia and Goias, Brasil.

64 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

Types:

Holotype female: ECUADOR, Zamora-Chinchipe: Zumbi, VI-10-1976, A. Langley

(USNM).

Paratypes (20 females, 2 males); PANAMA, Canal Zone: Barro Colorado I. V-12,

G. Fairchild (MCZ). COLOMBIA, Putumayo: Villa Garzon, 8 mi. s Mocoa, VII-24-78,

M. Cooper (BMNH); Villa Garzon, 400 m, X-13-84, M. Cooper (COOPER); Mocoa,

various dates, M. Cooper (BMNH, COOPER); Meta: La Macarena, XI-20/29-76, M.

Cooper (BMNH); Amazonas: Leticia, VIII-2-76, M. Cooper (BMNH). ECUADOR,

Napo: Muyuna, 500 m, 5 km w Tena, IV-22-81, M. Cooper (COOPER); Pichincha: Rio

Palanque Res. Sta., V-5/VII-25-85, S. & J. Peck (PMA); Morona-Santiago: Cord. de

Cutucu, 6 km e Macas, 1100 m, VII-3/10-81, M. Cooper (COOPER). PERU, Huanuco:

Monson Valley, Tingo Maria, XI-2-54, Schlinger & Ross (CAS); Cuzco: Quincemil, 750

m, XI-16/31-62, L. Pena (MCZ). BOLIVIA, Santa Cruz: Puerto Grether, 230 m,

IX-22-81, M. Cooper (COOPER). VENEZUELA, Bolivar: El Hormiguero Meseta de

Nuria, XII-13/17-74 (UCM). GUYANA: Mazaruni, VIII-19-1937, Richards & Smart

(BMNH). BRASIL, Para: Belém, IX-10-01, A. Ducke (VIENNA); Tucurui, I-1979, M.

Alvarenga (AEI); Rondani: Vilhena, XI-73, M. Alvarenga (AEI); Mato Grosso: Sinop,

X-75, M. Alvarenga (AEI).

Metatypes (9 females): COLOMBIA, Putumayo: Villa Garzon”, 400 m, X-10-84,

M. Cooper (COOPER), Puerto Leguizamo, 180 m, X-27/29-71, M. Cooper (BMNH), La

Hormiga near Mocoa™, IX-5-78, M. Cooper (BMNH); Cauca: Napoles”, 350 m, 5 kme

Puerto Limon, Rio Caqueta, IX-15-84, M. Cooper (COOPER). ECUADOR:

Cumbaratza, III/IV-65, L. Pena (MCZ). PERU, Madre de Dios: Quincemil, on branch

R. Manu, XI-5/16-62, L. Pena (AMNH). BRASIL, Bahia: Cepec Itabuna, III-4 &

VII-3-83, F. Benton (BMNH); Goias: Jatai, XI-1972, M. Alvarenga (FRITZ).

Etymology:

The name arachniraptor is a combination of the Greek word arachne (spider) and

the Latin word raptor (robber), and is a reference to the exclusive use of spiders as a

larval food source by these wasps.

Pison cameronii Kohl

(Figs. 204-220)

Pison fasciatum Kohl, 1884:339. Holotype female: "Mexico oder Peru” (VIENNA).

Preoccupied by Pison fasciatum Radoszkowski, 1876.

Pison cameronii Kohl, 1893:546, footnote 1. New name for fasciatum Kohl, 1884.

Description:

Infumation of forewing light or dark brown, often darkest over marginal cell.

Head black except scape often brownish beneath and palpi yellowish, scape, pedicel

and flagellomeres I-V yellowish in one male (Bahia, Brasil). Thorax black except:

edge of pronotal lobe usually paler, petiole socket area often brownish, closing face

of femora often brownish, midtarsomere I often brownish or whitish, and

hindtarsomere I often white except for apex (tarsomeres I-II whitish on all legs in

male from Mato Grosso, Brasil). Terga I-III either with translucent apical band or I

or I-II or I-III with white or creamy yellow apical bands; sternum I, tergum I

dorsobasally and its laterotergite often reddish brown. Metallic setae of face

(appressed setae sparse on female clypeus except laterally), gena and propodeum

silvery (sometimes golden on gena and propodeum).

Female clypeus with snout-like projection beneath free margin and weak tooth

lateral to it, apex of snout often narrowly notched (figs. 204-207) but sometimes

snout broad with wide, shallow emargination (figs. 209-210); margin of female

clypeus lateral to tooth thick in ventral view but attenuate laterad (fig. 208); male

clypeus with three similar stout teeth, middle one longest (fig. 211), clypeal margin in

ventral view thicker laterally than in female (fig. 212). Flagellomeres III-V or VI

with linear tyli in male (figs. 213-214). Female UID usually 0.60-0.69X LID, but

0.70-0.7/4X LID in exceptional forms (n Argentina, s Brasil); male UID 0.75-0.86X

Menke: Pison in the New World 65

LID. Female OOD usually 0.40-0.57X HOD, but in most specimens from southern

part of range (s Brasil) OOD 0.60-0.69X HOD, and in material from northern fringe of

range (Mexico) OOD 0.60X HOD; OOD rarely only 0.33-0.38X HOD (Trinidad, Peru);

male OOD usually 0.75-0.85X HOD, but 0.69X HOD in specimen from Caceres,

Brasil. Occipital carina ending on hypostomal carina just laterad of its apex.

Anterodorsal pit of pronotum transversely elongate, deep, as long as or longer than

hindocellus diameter. Tegula completely punctate. Male sternum VIII as in fig. 220.

Male genitalia as in figs. 215-219, midventral lobe of gonostyle bent laterad in

ventral view (figs. 216-218), tip of aedeagus expanded (figs. 217-218).

Length 8-10 mm (female), 7-8 mm (male).

Discussion:

Pison cameronii is a wide ranging, commonly collected species. The snout-like

projection beneath the clypeal rim is the most diagnostic feature in the female (figs.

204, 206), but it is subject to some variation. In occasional specimens from Colombia

and Brasil the snout is wide and shallowly emarginate (fig. 209). The incomplete

occipital carina, i.e., ending on the hypostomal carina, is an additional feature of the

species although not unique to it. Males can be recognized by the clypeal teeth and

the presence of linear tyli on flagellomeres III-V or VI (fig. 213). The genitalia are

also diagnostic: the tip of the aedeagus is expanded, and the midventral lobe of the

gonostyle is directed laterad (figs. 216-218). On the basis of its genitalia, cameronii

may be most closely related to arachniraptor, brasilium and maculipenne.

sometimes the propodeal dorsum and interface between the side and hindface is

densely covered with gold setae, and such specimens resemble chrysops. The female

Clypei (compare figs. 204, 206, 209 & 149-151) differ however, and most specimens of

chrysops have a broader ocellocular distance. The antennal tyli of male cameronii

separates that sex from males of chrysops.

The upper interocular distance varies in relation to the lower interocular

distance as indicated in the description, and is not a usable species character. The

UID is broadest in females from the southern end of the distribution of cameronii:

northern Argentina and southern Brasil, especially Nova Teutonia. The variation in

the length of the ocellocular distance lessens its value as a diagnostic tool. Its

variation is more random than the UID, but there is a definite tendency for the OOD

in females to be broad in material from the extreme ends of the range (Mexico and

southern Brasil).

I have examined the type. It is labeled simply "Am. A." In his original

description Kohl stated that the specimen came from Mexico or Peru but his evidence

was not given. |

Range:

southern Mexico to Colombia and Trinidad, widespread in South America where

Cameronii occurs as far south as Uruguay and northern Argentina.

Material examined (141 females of which 88 are from Nova Teutonia, Brasil, 8 males):

MEXICO, Chiapas: Palenque, 100 m (PMA); Muste, 440 m, nr. Huixtla (CNC). EL

SALVADOR: Los Chorros (UCD). COSTA RICA: Golfito (USNM); Palmar (USNM);

San José, San Antonio de Escazu (COOPER). PANAMA, Canal Zone: Barro Colorado

I, (USNM, AMNH). COLOMBIA, Caqueta: Yuruyaco, 73 km sw Florencia (BMNH);

Narino: Barbacoas, 50 m (COOPER); Meta: La Macarena (BMNH); Putumayo: Mocoa,

600 m (BMNH), Villa Garzon, 8 mi. s Mocoa (BMNH); Boyaca: Muzo, 900 m (AMNH),

Valle: Cali, 900 m (BMNH), Vaupes: Miraflores, 400 m (BMNH). ECUADOR, Napo:

Muyuna, 5 km w Tena (BMNH); Morona-Santiago: Cord. de Cutucu, 6 km e Macas,

1100 m (COOPER), Taisha, 500 m (COOPER); Pichincha: Rio Palanque Res. Sta., 250

m (PMA). PERU, Loreto: Pucallpa (BMNH); Madre de Dios: Avispas, 400 m (MCZ).

BOLIVIA, Beni: Rio Itenez at mouth of Rio Baures (AMNH), Rio Mamore at mouth of

Rio Ibare (AMNH); Santa Cruz: Est. Experimental Gral. Saavedra (LILLO).

PARAGUAY: Sommerfeld (LILLO). ARGENTINA, Salta: Pocitos (FRITZ), Corrientes:

Las Marias, Ca. Virasoro (LILLO), Ytuzaingo (FRITZ). TRINIDAD: Maracas Valley

(BMNH). GUYANA: Wismar (AMNH). BRASIL, Para: Belém (VIENNA); Bahia: Cepec,

66 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

Itabuna (BMNH); Mato Grosso: Caceres (AEI), Itaum (AEI); Sao Paulo: Caraguatatuba

(FRITZ); Guanabara: Rio de Janeiro (USNM), Represa do Rio Grande (AEI); Rio de

Janeiro: Conceigao de Macabu (AEI); Santa Catarina: Nova Teutonia (AEI, USNM,

LECLERCQ, UCD, MCZ, CU, OSU, KU). URUGUAY, Tacuarembo: 40 km nw

Tacuarembo (AMNH).

Pison dementia Menke, n. sp.

(Figs. 221-229)

Description, holotype female:

Infumation of forewing restricted to marginal cell, membrane lightly brownish.

Body black except: mandible reddish brown except basally; pronotal lobe, tegula, apex

of coxae, and lower surface of trochanters, femora and tibiae yellowish brown;

laterotergites I-II yellow brown. Clypeus laterally, lower frons, gena, top of pronotal

collar, scutal flange, and metanotum laterally with dense, appressed silver setae;

propodeum with suberect pale gold setae that are densest on dorsum and along

interface between side and hindface. Terga I-III with with broad, apical creamy

yellow bands, IV-V translucent apically.

Clypeal margin without median lobe, margin with round notch and two weak

teeth lateral to it (similar to fig. 221), polished rim broadened behind notch and

teeth, clypeal margin narrowly thickened in ventral view (similar to fig. 222). UID

0.91X LID. OOD 1.11X HOD. Occipital carina incomplete ventrad, ending short of

hypostomal carina apex. Pronotum with transversely elongate anterodorsal pit, its

length greater than hindocellus diameter. Tegula completely punctate. Tarsi

apparently without plantulae.

Length 8.5 mm.

Variation in females (7 specimens):

Infumation of forewing leading edge sometimes extending basad from marginal

cell; legs sometimes mostly black with only lower surface of mid and hindfemora

yellowish brown; appressed facial and genal setae sometimes golden; pair of teeth

lateral to clypeal notch sometimes poorly or not differentiated; UID 0.86-0.91X LID;

OOD 1.02-1.29X HOD; forewing media often interstitial with crossvein cu-a or nearly

so; length 7-8.5 mm.

Male (2 specimens):

As in female except: marginal cell only faintly infumate; antenna sometimes

yellowish brown beneath to tip and palpi and mid and hindtarsomere I sometimes

yellowish (Bahia); clypeal margin with short, median tooth and lateral sinuation (fig.

223); flagellum simple; UID 0.98-1.0X LID; OOD 1.17-1.51X HOD; sternum VIII

similar to maculipenne; genitalia as in figs. 225-229, midventral lobe of gonostyle

broad, its apex curled (figs. 225, 227); length 5.5-7 mm.

Discussion:

The clypeus is distinctive in both sexes of dementia (figs. 221, 223). The very

broad upper interocular distance (UID usually almost as broad as LID) and ocellocular

distance (OOD about equal to HOD or up to 1.5X HOD) are additional recognition

features of the species. The male genitalia are diagnostic (figs. 226-229).

Range:

Known only from eastern and southeastern Brasil.

Types:

Holotype female: BRASIL, Santa Catarina: Nova Teutonia, I]-24-1964, Fritz

Plaumann (UCD).

Paratypes (8 females, 2 males): BRASIL, Bahia: Itapetinca, 300 m, XI-69, F. M.

Oliveira (AEI); Guanabara: Rio de Janeiro, I-39, R. C. Shannon (USNM); Sao Paulo:

Menke: Pison in the New World 67

Galia Est., VII-1984, F. M. Oliveira (AEI), Sao Paulo, VII-4-77, V. Alin (MLSU); Santa

Catarina: Nova Teutonia, various dates, F. Plaumann (UCD, MCZ).

Etymology:

The name dementia, noun in apposition, is the Latin word for insanity. The

combination Pison dementia names a condition that will afflict anyone who, in the

future, decides to do further work on the Neotropical species of the genus,

particularly in the cressoni group.

Pison abothrum Menke, n. sp.

(Figs. 230-238)

Description, holotype female:

Infumation of forewing leading edge uniformly brown, wing membrane pale

brownish behind. Following parts yellowish brown: scape, pedicel, flagellomere I,

clypeus except near antennal socket, mandible (apex darker), labrum, palpi, pronotal

lobe, tegula, propodeal side above mid and hindcoxae, rim of petiole socket, precoxal

lobe, metasternum, legs except coxae dark above, gastral seqment I except distal half

of tergum black, and laterotergite II. Gaster without creamy yellow tergal bands.

The following with dense, appressed gold setae: clypeus and lower half of frons and

narrowly along inner orbit to level of ocelli, gena, and top of pronotal collar; setation

of rest of thorax including legs similar or paler, longest on propodeum. Tergum I

setae brownish gold, but setation of rest of gaster brownish.

Clypeal margin with median lobe that is depressed below surface of clypeus, lobe

almost as wide as clypeus, its margin obtusely triangular (fig. 230), its surface

smooth, asetose, edge of lobe thin in ventral view. UID 0.67X LID. OOD 0.56X

HOD. Occipital carina incomplete ventrad, ending near apex of hypostomal carina.

Pronotum without anterodorsal pit. Media of forewing diverging from M+Cu before

crossvein cu-a. Tegula sparsely punctate on outer half. Tergum II more convexly

swollen in lateral profile than tergum III. Tarsi apparently without plantulae.

Length 10 mm.

Variation in female (one specimen):

Body more extensively yellowish brown: flagellomere II (III reddish brown),

clypeus entirely, ventral margin of pronotum, scutal flange, metanotum laterally,

mesopleuron except venter, metapleuron, propodeal side, propodeal hindface

extensively, mid and hindcoxae entirely, tergum I entirely except for small brown

spot near hindmargin, base and margins of tergum II, lateral margin of tergum III, and

sternum II except for transverse brown spot near hindmargin. Setae of gaster paler.

OOD 0.60 HOD. Length 9 mm.

Male (one specimen):

As in female except: following yellowish or yellow brown: scape, pedicel,

flagellomeres I-III, lower half of clypeus (lobe dark), mandible, labrum, palpi,

pronotal lobe, tegula, scutal flange, hypoepimeral area of mesopleuron, metapleuron,

propodeal side, legs except forecoxa, and gastral segment I (except tergum with tiny

brown spot near hindmargin. Appressed facial setae a mixture of gold and silver.

Clypeus with small median lobe with rounded tooth at middle (fig. 231); clypeal

margin thickened (ventral view, fig. 232). Flagellum simple. UID 0.82 LID. OOD

0.73X HOD. Media of forewing almost interstitial with crossvein cu-a. Tergum II

not appreciably more convex than III. Sternum VIII narrow (fig. 238). Genitalia as in

figs. 233-237, aedeagus with apicoventral lobe (fig. 233), midventral lobe of gonostyle

broad, but narrowing to small rounded apex (fig. 235). Length 7 mm.

Discussion:

Pison abothrum has a number of distinctive features. The absence of a pronotal

pit separates the species from all others in the cressoni group except aranevorax

68 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

which has only two submarginal cells in the forewing. The divergence of the forewing

media before crossvein cu-a is shared with only one other species in the group,

brasilium, but the latter has broader ocellocular and upper interocular distances.

Tarsal plantulae appear to be absent in abothrum. Other species of the cressoni

group (except aranevorax, dementia and phthinylla) have these at least on

hindtarsomere IV. The female clypeal lobe of abothrum is unusual and diagnostic (fig.

230). In the male the broad midventral lobe of the gonostyle is unusual among

cressoni group species, as is the shape of the aedeagus in lateral profile (figs. 233,

235).

Pison abothrum is similar to martini in color pattern (legs almost totally

yellowish, gaster without yellow bands, etc.), and the female of abothrum has a

somewhat humped tergum II like the latter species. However martini has a deep

pronotal pit and the media diverges after crossvein Cu-a.

The male that I associate with the two Colombian females of abothrum is from

Mato Grosso, Brasil. It does not have a pronotal pit, the forewing media diverges

before crossvein cu-a (just barely), tarsal plantulae are absent, and it has the same

color pattern as the females. Thus I am confident that the male and two females are

conspecific.

Range:

Known only from Colombia and Mato Grosso, Brasil.

Types:

Holotype female: COLOMBIA, Putumayo: Villa Garzon, 400 m, X-17-1984, M.

Cooper (BMNH#H).

Paratypes (one male, one female): COLOMBIA, Meta: La Macarena, XI-1976, M.

Cooper (BMNH). BRASIL, Mato Grosso: Sinop, 129 31’ S, 550 37' W, II-1976, M.

Alvarenga (AEI).

Etymology:

The Greek word bothros means pit. The name abothrum, based on that word,

means without a pit, a reference to the absence of a pronotal pit in this species.

Pison aranevorax Menke, n. sp.

(Figs. 239-248)

Description, holotype female:

Infumation of forewing leading edge darkest on marginal cell, membrane clear.

Body black except: scape beneath, palpi, hindmargin of pronotal lobe, tegula and

foretibia yellowish brown; tergal apices and lateral margins pale, translucent (no

yellow bands). Clypeus laterally with appressed silver setae; lower frons and gena

with appressed pale golden setae.

Clypeal margin with short median lobe (fig. 239) that has median projection,

edge only slightly thickened in ventral view (fig. 240). Mandible slender in anterior

view (fig. 239). UID 0.65X LID. OOD 0.37X HOD. Occipital carina incomplete

ventrally, ending just short of hypostomal carina apex. Pronotum_ without

anterodorsal pit. Tegula sparsely punctate on outer half. Forewing with only two

submarginal cells, the reduction obtained by the loss of the outer veinlet of true II;

recurrent vein I interstitial between submarginals I-II; tarsi apparently without

plantulae.

Length 7 mm.

Variation in females (8 specimens):

Forewing membrane sometimes lightly infumate behind foremargin; scape

entirely yellowish brown, or entirely black; legs entirely black or more extensively

yellowish brown: tarsomeres I-II of mid and hindleg and apical fourth of mid and

hindfemur ventrally; clypeal lobe sometimes strongly tridentate, middle tooth cuplike

beneath (figs. 241-242), clypeal margin often more strongly thickened than in type

Menke: Pison in the New World 69

(ventral view); UID 0.54-0.64X LID; OOD 0.25-0.38X HOD; pronotum sometimes with

weak, transversely elongate, anterodorsal pit; recurrent vein I sometimes ending near

apex of submarginal cell I, sometimes ending on submarginal cell II; length 6.5-8 mm.

Male (two specimens):

As in holotype except: tarsomere I on mid and hindleg pale, basal fourth of mid

and hindtibiae yellowish brown; clypeus with short, three-toothed lobe (fig. 243);

flagellum simple; UID 0.65-0.67X LID; OOD 0.40-0.44X HOD; sternum VIII similar to

maculipenne (fig. 176); genitalia as in figs. 244-248, midventral lobe of gonostyle

broad, its ventral edge curled (figs. 244-246), aedeagus laterally compressed at apex,

its tip broad, angular (figs. 244, 247); length 6 mm.

Discussion:

The presence of only two submarginal cells in the forewing separates aranevorax

from other species of the cressoni group except erebus and phthinylla. The female

clypei differ however (compare figs. 239, 241 & 160, 249), and erebus has a broader

ocellocular distance. P. aranevorax has a normal tergum II and largely black legs,

while phthinylla has a strongly humped tergum II, largely yellowish brown legs, a

largely yellowish brown tergum I, and a deep anterodorsal pronotal pit. The

propodeum of aranevorax is short with a fairly steep hindface, but in phthinylla it is

elongate with a gradually sloping hindface (fig. 252). The compressed, angular

aedeagus of aranevorax is distinctive (figs. 244, 247).

The holotype, one male and one female paratype were reared from the same nest

by Martin Cooper, and are assuredly conspecific. However, six of the nine females

display puzzling variation in the clypeus (one of two specimens from Leticia,

Colombia, and material from Mitu, Colombia, Ecuador, Peru and Brasil). The clypeal

lobe is more strongly dentate in these (fig. 241), the polished lip is bordered above by

a fine, sharp impression, and these females also have a weakly developed anterodorsal

pit on the pronotum. The female from Ecuador (Santiago) was captured at a nest

from which a male emerged. The genitalia of this male are identical with those of

the male from Barbacoas, Colombia. Their clypei are also identical. Thus I assume

that only one species is represented by the material before me.

Range:

Known only from Colombia, Ecuador, Peru, and eastern Brasil.

Types:

Holotype female: COLOMBIA, Narino: Barbacoas, 80 m, I-2-1972, ex nest #3, M.

Cooper (BMNH).

Paratypes (8 females, 2 males): COLOMBIA, Narino: same data as type;

Amazonas: Leticia, VIII-19/23-74, M. Cooper (BMNH); Vaupes: Mitu, V-ll-74, M.

Cooper (BMNH) ECUADOR, Napo: Archidona, 500 m, IV-9-81, M. Cooper

(COOPER); Morona-Santiago: Santiago, 300 m, VII-27-87, biology note 30, M. Cooper

(COOPER). PERU, Huanuco: Monson Valley, Tingo Maria, IX-23-54, Schlinger and

Ross (CAS). BRASIL, Bahia: Itabuna, II]-17-85, F. P. Benton (BMNH).

Etymoloaqy:

The name aranevorax is based on the Latin words aranea (= spider) and vorax (=

Gluttonous). It refers to the exclusive use of spiders as a larval food source.

Biology:

Martin Cooper reared the Barbacoas material (1 male, 2 females) from a single

mud nest that was built on the under side of an oblique substrate. Cooper's brief

notes (BMNH) include crude sketches of the nest. The nest apparently consisted of 8

vertical mud tubes that were surrounded by mud “flanges”. Four of the tubes were

sealed. Two of the open tubes contained a sealed cell, one of which contained a

cocoon with larva inside, a possible parasitic larva on the outside of the cocoon, and

the remains of spiders.

Cooper also found a nest in Santiago, Ecuador (biology note 30). This nest was

70 Contrib. Amer.iEnt. Inst.; vol, 24, no. 3, 1968

attached to the underside of the leaf of an epiphytic plant. It consisted of two rows

of cells (3 and 5 cells respectively). The more or less vertical cells projected from

basal layers of mud. A male was reared from the nest and Cooper captured a female

at the nest. A chalcidoid wasp was also obtained from the nest.

Pison phthinylla Menke, n. sp.

(Figs. 132, 249-252)

Description, holotype female:

Infumation of forewing leading edge brown, restricted to marginal cell and outer

third of submarginal I, wing membrane pale brownish elsewhere. Following body

parts yellowish brown: scape, pedicel beneath, labrum, palpi, mandible except apex

dark, pronotal lobe, tegula, fore and midlegs except coxa, hindleg except coxa dark

above, gastral segment I except tergum with small dark brown spot just behind

spiracle, sternum Il; remaining terga and sterna brown but pale margined. Lower

frons, clypeus laterally, and procoxa with dense appressed gold setae; setation of rest

of body mostly sparser, yellowish brown or brownish, longest on propodeum.

Clypeal margin with median lobe that has semicircular emargination, surface of

lobe smooth and descending laterad below clypeal margin (similar to fig. 249); edge of

lobe thickened in ventral view (similar to fig. 250). UID 0.57X LID. OOD 0.38X

HOD. Occipital carina incomplete ventrally, ending just short of hypostomal apex.

Anterodorsal pit of pronotum transversely elongate, sharply delimited anteriorly but

not posteriorly. Tegula completely punctate although sparsely so on outer half.

Propodeal dorsum with fine median longitudinal carina; propodeal hindface meeting

dorsum at a broad obtuse angle (side view, similar to fig. 252). Forewing with only

two submarginal cells due to loss of outer veinlet of second submarginal cell (similar

to fig. 132). Tarsi apparently without plantulae. Thorax and gaster elongate, tergum

I length (measured in dorsal view) 1.25X apical width; tergum II swollen in lateral

profile, in contrast to tergum III (similar to fig. 251).

Length 10.5 mm.

Female variation (1 specimen):

UID 0.56X LID. OOD 0.32X HOD. Length 9.5 mm.

Male:

Unknown.

Discussion:

Pison phthinylla is similar to two other species with pale legs, martini and

abothrum, but the presence of only two submarginal cells and the shape of the female

clypeus are immediately diagnostic. Tergum II is more strongly humped in phthinylla

than in martini. The propodeum of phthinylla is elongate in side view, the angle

formed by the hindface and dorsum being a broad obtuse angle (fig. 252). The angle is

less in martini.

Pison aranevorax and erebus also have only two submarginal cells but they differ

from phthinylla in a number of ways (see aranevorax and erebus).

Range:

Known only from Ecuador.

Types:

Holotype female: ECUADOR, Morona-Santiago: Cord. de Cutucu, 6 km e Macas,

1,100 m, V-21-1981, M. Cooper (COOPER).

Paratype (one female): same data as type except 1000 m, X-27-78 (BMNH).

Etymology:

The Greek word phthinylla, a noun in apposition, means a thin or delicate woman,

a reference to the elongate form of the species.

Menke: Pison in the New World Be

Chilense Group

(Figs. 253-267)

Description:

Head, thorax and coxae, trochanters and femora beneath with long, erect setae.

Labrum quadrangular. Female mandible with weak tooth at middle of cutting edge,

male mandible without tooth. Clypeus with prominent median lobe whose edge is not

thickened. Occipital carina incomplete ventrally, ending just before hypostomal

carina. Pronotum with anterodorsal pit, but no lamella. Pronotal collar about as

thick as metanotum, about as high as scutum. Lateral flange of scutum narrow.

Tegula impunctate on outer half. Propodeum entirely striatopunctate or ridged, but

without crenulate ridge at top of side. Propleuron punctate. Episternal sulcus

curving forward ventrad, attaining anterior margin of pleuron. Hindcoxa with inner

dorsal carina, outer carina complete or only with apical remnant. Tarsomeres spinose

apically, claws normal; hindtibia with few short, stout setae scattered over outer

surface. Forewing media diverging before or after crossvein cu-a; marginal cell

acuminate distally although somewhat bluntly so in chilense; three submarginal cells

present, III not strongly narrowed on marginal cell; termination of recurrent veins

variable. Hamuli of hindwing more or less divided into two groups. Margin of tergum

I single-edged; tergum VII of male truncate apically. Male sternum VIII ending in two

prongs. Male genitalia complex: gonostyle with long setae on outer (lateral) surface,

and two ventral lobes (figs. 258, 264); volsellar lobes feebly setose; aedeagus with

well defined head, but no ventral tooth or spine.

Included species:

Pison chilense and sylphe.

Discussion:

The main features of the chilense group are: the long, erect body setae, the

many propodeal carinae, the absence of a crenulate ridge at the top of the propodeal

side, the emarginate and pronged male sternum VIII, and the complex gonostyle of the

male genitalia (with accessory lobes). The last two features are apomorphies. The

long, erect body setae and tergal fasciae may be additional apomorphies. Pison

chilense has tarsal plantulae, but these are apparently absent in sylphe. The closest

allies of the chilense group, which appears restricted to the Andes, may be found in

Australia based on material that I have seen from that continent.

Pison chilense Spinola

(Figs. 253-254. 256-260)

Pison chilense Spinola, 1851:326. Syntypes, male and female: "Coquimbo" and

"Santiago", Chile (TURIN).

Pison chilense, Janvier, 1928:87. Biology.

Pison chilensis, Casolari and Moreno, 1980:114. Lists three syntypes from "Chili".

[Notes taken by R. M. Bohart in 1960 during a visit to the museum in Turin

indicate that there are 2 males and | female].

Description:

Margin of labrum slightly arcuate or straight in female, slightly concave in male;

edge of female clypeal lobe arcuate, surface smooth, shiny; male clypeal lobe

triangular, surface smooth, shiny; flagellomeres II-IV of male antenna swollen

beneath, flagellomeres I-IX each with depressed linear placoid beneath that become

progressively smaller toward article IX which only has tiny circular placoid; pronotum

with anterodorsal, transverse pit whose length is about three-fourths distance across

hindocelli, pit overlain by lamella anterad; propodeum entirely striatopunctate,

obliquely so dorsally, transversely so on hindface, dorsum without median longitudinal

q2 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

carina; outer carina of hindcoxa restricted to apical remnant; forewing media

interstitial with cu-a or diverging from M+Cu before it; marginal cell apex bluntly

acuminate (fig. 253); submarginal cell II occasionally incomplete through loss of

either inner or outer veinlet; first recurrent vein of forewing ending on submarginal

cell I or II, or interstitial between I and II, second recurrent vein ending on III; tergum

VII of male truncate or even concave; sternum VIII as in fig. 254; gonostyle with

setose midventral lobe whose upper end projects laterad as a horn (figs. 256, 258), and

a more basal lobe that is scimitar-like in ventral view (fig. 259), and has patch of

stout setae basally (fig. 257), outer surface of gonostyle setose, dorsal edge with few

widely spaced long, sinuate, stouter setae (fig. 260); volsellar lobes angular in ventral

view (fig. 259).

Discussion:

This commonly collected Chilean and Argentinean endemic is easily recognized

by its large size (8-11.5 mm long), the amber tegulae and wing veins (at least

basally), the long dark erect setae of the head and thorax (including the coxae,

trochanters and femora), and the broad silver fasciae of terga I-III. The tergal

fasciae give chilense the appearance of the genus Tachytes. The face and top of the

pronotal collar are covered by dense appressed silver setae and this type of setation

also occurs on the mesopleuron just beneath the scrobe, on the gena, and on the outer

surface of the femora.

I have not studied the Spinola’s syntypes but have read R. M. Bohart's notes on

them taken during his visit to the museum in Turin in 1960. His notes and Spinola's

description are sufficient to positively identify chilense.

Range:

Chile between 309-379 latitude (Coquimbo Prov. to Concepcion Prov.) and the

provinces of Tucuman and Neuquen in Argentina. Records from Argentina are few

but they suggest that chilensis probably occurs in the Andes from Tucuman to

Neuquen.

Material examined (482 specimens):

CHILE, Coquimbo: 8 km n San Pedro DeQuile (MCZ); El Panque (MCZ); Tongoy

(MCZ); Ovalle (MCZ); Hda. Illapel, 900-1200 m (OSU). Aconcaqua: Cta. El Melon

(MCZ). Valparaiso: El Salto (BMNH, MCZ); Galalea (ZMC); Valparaiso (MCZ); La

Campana nr Limache (MCZ). Santiago: Queb. de La Plata, La Rinconada, Maipu

(LILLO, CAS); Pefalolén (BMNH); Limache (BMNH); 20 km e Santiago (ZMC);

Apoquindo (LILLO, MCZ); Olivicura (MLSU); Quebrada El Peumo (MLSU, KU, MCZ,

OSU); Quebrada S. Ramon (MLSU); 5 km e Cuesta de Ibache (CAS); Camelo (LILLO,

KU); El Canelo (OSU); Curicavi (OSU); Nido de Aguila (OSU). O'Higgins: La Leonera

(MCZ). Colchaqua: 3 km n Callejones (CAS). Maule: Forel Carrizalillo (USNM);

Cauquenes (BERLIN). Nuble: 17.5 km s Curanipe (USNM). Concepcion: Concepcion

(BMNH). Province unknown: Nido de Aguila (MCZ); Olmué (BMNH); Tonlemo (KU);

Estero del Tempo (KU); Margamarga (MCZ); Illapel (MCZ). ARGENTINA, Tucuman:

Tafi del Valle, La Quebradita (LILLO). Neuquen: Collon Cura (LILLO).

Biology:

Janvier (1928) described and illustrated the nest of chilense. The species is a

ground nester, excavating burrows in stream banks. The soil is moistened with water

from the adjacent streams to facilitate excavation. Cells are arranged in linear

fashion and each is separated by a wall of mud.

Pison chilense has long, erect setae on the gena, propleuron, and coninal surface

of the forecoxa, trochanter and femur. The mid and hindlegs of chilense also have

long setae on the trochanter and femur. The lower and upper surfaces of the

mandible have a linear series of long setae set in grooves. This vestiture (except on

the mandible) is scattered and may not be identical to the psammophore described by

Evans (1981) on some Australian ground nesting species of the genus.

Menke: Pison in the New World 73

Pison sylphe Menke, n. sp.

(Figs. 255, 261-267)

Description, holotype female:

Black, wings slightly infumate. Erect setae of body pale; lower frons and clypeus

with appressed silver setae; terga I-IV with broad silver fasciae.

Margin of labrum entire. Clypeus with triangular median lobe (fig. 261). Frons

weakly swollen, dull, no obvious punctation. Eye length 0.94X distance between eye

notches; UID 0.59X LID; OOD 0.40X HOD. Flagellomere I length 3X apical width,

II-III slightly less than 3X, IX slightly longer than wide.

Pronotum with very small, transversely elongate anterodorsal pit (half

hindocellus diameter). Scutum dull, imbricate, shallowly punctate, punctures one to

two diameters apart, closer than that anteriorly. Scutellum flattened, weakly shiny,

imbricate, punctate, punctures more well defined than those of scutum. Propodeum

entirely covered by closely spaced, fairly regular ridges (fig. 263), interspaces

smooth, impunctate, shiny, ridges on dorsum longitudinal and continuing onto

hindface where they converge toward midline, ridges on propodeal side oblique and

curving onto hindface. Mesopleuron weakly shiny, more coarsely, sharply punctate

than scutum, punctures one-half to two diameters apart.

Forewing media diverging after cu-a. Both recurrent veins ending on

submarginal cell II.

Hindcoxa with complete but low outer carina on dorsal surface. Terminal

tarsomeres and claws missing on all legs, tarsal plantulae apparently absent.

Gaster weakly shiny, closely, finely punctate.

Length 9 mm.

Male:

As in female except: fasciae on terga II-IV weaker than that on I. Labrum with

slight indentation. Clypeal lobe quadrangular but with small, median tooth (fig. 262).

Frons with shallow but well defined punctation, punctures mostly one diameter

apart. UID 0.72 LID; OOD 0.82 HOD. Flagellomere I length 3X apical width, II

2.5X apical width, III 2X apical width, IX slightly longer than wide. Ridges of

propodeal dorsum slightly divergent toward apex, middle ridge straight, stronger than

others; ridges of dorsum not continuing onto hindface in convergent manner, hindface

with many cross-ridges, the uppermost of which are continuous with those on

propodeal side, ridges of propodeal side changing to striatopunctation ventrad. First

recurrent vein interstitial between submarginals I-II on right wing. Sternum VIII as in

fig. 255. Genitalia as in figs. 264-267. Length 7.5 mm.

Discussion:

The many closely spaced carinae with impunctate intervals on the propodeal

dorsum, and the black body with pale erect hair immediately identify sylphe. The

shape of the clypeal lobe in both sexes, the absence of tarsal plantulae, the simple

male flagellum and complex genitalia are also diagnostic. The volsellar lobes are

rounded in ventral view (fig. 266) and the basalmost lobe of the gonostyle extends

laterad as a sharp point in ventral view (fig. 266).

Range:

Known only from northern Peru at elevations above 4800 feet (1500 m).

Types:

Holotype female: PERU, Cajamarca: Cajamarca, 2700 m, III-1942, W. Weyrauch

(ttt):

Paratype male: PERU, La Libertad: Samne, ca. 40 km ne Trujillo, 1500 m,

VII-12/17-75, Porter and Stange (LILLO).

714 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

Etymology:

The name sylphe, a noun in apposition, is based on the French word sylphe which

means "a fairylike spirit of the air".

Convexifrons Group

(Figs. 5, 268-291)

Description:

Frons swollen (inflated); eye asetose; male antenna simple, flagellum without tyli

or other modifications; mandible with posterior notch (fig. 5), female mandible with

mesal tooth or cleft on cutting edge; lower half of female clypeus with long,

somewhat decumbent setae that project over clypeal margin forming a "brush";

female clypeal lobe triangular or quadrangular; male clypeus with polished,

triangular, median lobe (fig. 271); labrum quadrangular, apex entire or with obtuse

emargination; occipital carina incomplete ventrad, ending on or near hypostomal

carina; anterior edge of pronotum with polished plate-like area at middorsum that is

not free (fig. 288), with deep, transversely elongate depression behind it that has

sharp hindmargin (fig. 288); scutal flange broad, somewhat reflexed; tegula

impunctate on outer half, punctate adjacent to scutum; propodeal side delimited

dorsad by crenulate ridge (figs. 268-269); propleuron punctate, disk not largely

polished; metapleural flange narrower than hindocellus diameter; tergum-I apex

single-edged; forewing media diverging from M+Cu after crossvein cu-a; marginal

cell acuminate distally; forewing with three submarginal cells, III strongly narrowed

toward marginal cell, its veinlets often joining there or even forming petiole; first

recurrent vein ending on submarginal cell I or II, second ending on submarginal II or

interstitial between II-III]; hamuli of hindwing divided into two groups; male

foretrochanter without spine; anterior claw of male foreleg normal, not distorted;

dorsum of hindcoxa usually delimited by complete outer and inner carinae, inner

carina usually lamelliform in part; tarsomeres III-IV, or IV with plantulae; gaster

without yellow tergal bands; male tergum VII simple, not laterally compressed; male

sternum VII simple, VIII broadly concave apically (fig. 278); genitalia somewhat

laterally compressed, gonostyle elongate, setose, usually with midventral spine or

finger-like lobe (figs. 273, 282), volsellar lobes small (figs. 274, 290).

Included species:

Pison convexifrons, cooperi, longicorne, wasbaueri.

Discussion:

The main features of this group, all of which are apomorphies, are the

mandibular notch; the swollen frons; the non-lamelliform polished area at the top of

the pronotum that does not extend very far laterally; the deep, transverse depression

behind it; the propodeal ridge; and the apically emarginate male sternum VIII.

Three species, convexifrons, cooperi and wasbaueri, are very similar structurally

and form a monophyletic unit within the convexifrons group. The propodeal hindface

of these species is cross-carinate (figs. 268-269), the antennal flagellomeres are

about 2X as long as wide and the outer ones are much shorter, the female clypeal

lobe is triangular, the female labrum is emarginate, and the gonostyle of the male

genitalia has a spinose process ventrally (figs. 273, 277, 282). Although the three are

not strikingly different externally, the male genitalia are diagnostic.

The fourth species, longicorne, is unique in a number of ways. The propodeal

hindface is largely smooth, the flagellomeres are 2.5 to 3.75X as long as wide and the

articles remain long to the tip, the hindocellus is much farther from the eye than in

convexifrons and its siblings, the pronotal collar has a prominent median elevation

(fig. 288), the female clypeal lobe is roundly quadrangular, and the female labrum is

not notched. The male genitalia lack a ventral spine on the gonostyle and the

volsellar lobes are larger than in the convexifrons complex (fig. 290).

Menke: Pison in the New World te

Wing venation, particularly the end point of the recurrent veins, varies within

each species as indicated in the descriptions. The size and shape of submarginal cells

II-III also varies, and in exceptional specimens the second is open through loss of a

veinlet. In two species (some aureofaciale and some longicorne) the veinlets of

submarginal cell III join at the marginal cell, and sometimes they join before meeting

it, forming a petiole.

Pison convexifrons Taschenberg

(Figs. 5, 268, 270-275)

Pison convexifrons Taschenberg, 1870:18. Lectotype female: Nov. Frib. (Nova

Friburgo, Rio de Janeiro, Brasil), present designation (HALLE).

Description, male (2 specimens):

Body black, hindtarsus brownish. Appressed facial setae silver. Abdomen

without silver fasciae.

Labrum quadrangular, obtusely indented. Clypeus as in fig. 271, frons weakly

shiny, densely punctate. UID 0.73X LID. OOD 0.57X HOD. Flagellomeres I-IV 2X as

long as apical width, V 1.33X as long as apical width, VI-X progressively shorter.

Scutum and scutellum dull, densely punctate, punctures less than puncture

diameter apart. Propodeum shiny, dorsum with median, longitudinal carina that fades

posterad, surface with many carinae that are oblique basally, but transverse on apical

half, dorsum punctate laterally; hindface impunctate, but with coarse cross-carinae

similar to those on dorsum; propodeal side with oblique or horizontal carinae dorsad,

these changing to weak striatopunctation or simple punctation ventrad. Mesopleuron

dull, densely punctate (mostly less than 1 diameter apart), punctures below scrobe

coarser than those on scutum. Metapleuron finely, horizontally carinate above, finely

punctate, shiny below. First recurrent vein interstitial or ending near base of second

submarginal cell; second recurrent vein ending near apex of second submarginal cell

or interstitial.

Sternum VIII apex broadly, shallowly concave (similar to fig. 278). Genitalia as

in figs. 272-275; outer surface of gonostyle apex covered by long setae (fig. 272),

ventral edge of gonostyle with curved fingerlike process at about midpoint (figs.

273-274).

Length 5 mm.

Female (18 specimens):

Similar to male except: tarsi black, terga I-II occasionally with silver fasciae

laterally. Clypeus as in fig. 270. UID 0.50-0.60X LID. OOD O.10-0.25X HOD.

Flagellomere I length slightly more than 2X apical width. Transverse carinae of

propodeal dorsum usually finer than in male (fig. 268), sometimes changing to

striatopunctation laterally, or evanscent, median carina sometimes in broad trough.

First recurrent vein ending near apex of Ist submarginal cell in half of the material,

interstitial in the others; second recurrent vein usually ending near apex of

submarginal cell II but occasionally interstitial between II and III (right and left wings

of specimens sometimes different for both recurrent veins). Length 5.5-7.5 mm.

Discussion:

The cross-carinate propodeal dorsum (fig. 268) and male genitalia are the best

diagnostic features of convexifrons. The species is similar to wasbaueri but the setae

of the gonostyle are confined to an apical group in convexifrons (fig. 272). There are

two groups of setae in wasbaueri (fig. 276), and the propodeal dorsum is only

punctate. The two species are allopatric.

The cross-carinate propodeal dorsum of convexifrons separates it from cooperi.

The latter species has a largely smooth dorsum (fig. 269), and the genitalia are

different (compare figs. 272-275 & 281-284).

In several females the cross-carinae of the propodeal dorsum are evanscent

76 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

(Blumenau and Nova Teutonia, Brasil; Pirapo, Paraguay). I have seen normally

carinate females from the same Brasilian localities, however. Nevertheless, these

weakly carinate specimens are difficult to positively identify without associated

males. Forewing venation is variable in convexifrons. The size and shape of

submarginal cells II and III varies, and the end point of the recurrent veins may be

different on the two wings of one individual as indicated in the description. The

second submarginal cell is incomplete in one wing of one female.

Taschenberg (1870) described convexifrons from three female syntypes all of

which I have examined. A lectotype label has been placed on one of the specimens

from Nova Friburgo.

Range:

This wasp is known only from southeastern Brasil, eastern Paraguay and

northeastern Argentina.

Material examined (18 females, 2 males):

BRASIL, Rio de Janeiro: Nova Friburgo (HALLE); Guanabara: Rio de Janeiro

(HALLE); Santa Catarina: Nova Teutonia (UCD, LECLERCQ, MCZ), Corupa (AMNH),

Blumenau (VIENNA). PARAGUAY: Pirapo (FRITZ). ARGENTINA, Entre Rios:

Federacion (FRITZ).

Pison wasbaueri Menke, new species

(Figs. 276-277, 279)

Description, holotype male:

Body black, tarsi brownish. Appressed facial setae silver. Abdomen without

tergal fasciae.

Head same as convexifrons except UID 0./6X LID; OOD 0.71X HOD.

Thorax same as convexifrons except scutum weakly shining; propodeal dorsum

with median longitudinal carina and short basal carinae, but rest of surface only

punctate; first recurrent vein interstitial between submarginal cells I-II, second

recurrent ending near apex of II.

Sternum VIII same as convexifrons. Apex of gonostyle with setae along outer

ventral margin and cluster of setae distad of ventral spine (figs. 276-277).

Length 5.5 mm

Variation in males (2 specimens):

UID 0.72-0.77X LID; OOD 0.57X HOD in one specimen. First recurrent vein

ending near apex of submarginal cell I in three wings, interstitial in the fourth,

second recurrent interstitial or nearly so. Outer veinlet of third submarginal cell

incomplete in one wing. Length 4.5-6 mm.

Female (1 specimen):

Same as female convexifrons except: UID 0.61X LID; flagellomere I length

about 2.5X apical width; propodeal dorsum without transverse carinae or

striatopunctation; first recurrent vein ending on submarginal cell I; length 5 mm.

Discussion:

The absence of cross—-carinae on the propodeal dorsum of wasbaueri is the only

external difference from convexifrons. The male genitalia of wasbaueri are

distinctive: the setae of the gonostyle are divided into two groups, an apical one and

another just distal to the ventral spine (figs. 276-277). The genitalia of wasbaueri

offer the only reliable means of separating the species from males of cooperi

(compare figs. 276-277 & 281-184). The single female of wasbaueri differs from

most females of cooperi in having a uniform, dense punctation on the mesopleuron

(fig. 279). In most females of cooperi the punctures are sparser beneath the scrobe

and toward the mesopleural suture (fig. 280).

Menke: Pison in the New World 77

Range:

Known only from northwestern Argentina. The species has been collected at low

elevation Andean localities in cloud forest/deciduous subtropical thornforest

transition zones.

Types:

Holotype male: ARGENTINA, Salta: Rosario de Lerma, XII-24/27-1983, malaise

trap, M. Wasbauer (CSDA).

Paratypes (2 males, 1 female): ARGENTINA, Salta: Yuto, I-11-66, H. & M.

Townes (AEI); Tucuman: Horco Molle, Parque Sierra San Javier, 700 m, I-15-76, L. A.

Stange (LILLO).

Etymology:

This species is dedicated to Marius Wasbauer, collector of the holotype.

Pison cooperi Menke, new species

(Figs. 269, 278, 280-284)

Description, holotype male:

Body black. Appressed facial setae silver. Abdomen without tergal fasciae.

Head same as convexifrons except UID 0.67X LID; OOD 0.30X HOD; and

flagellomere I length nearly 2.5X apical width.

Thorax same as convexifrons except surface of propodeal dorsum only punctate

(similar to fig. 269), only with vestiges of basal and transverse ridging; propodeal side

with few horizontal ridges beneath spiracle, but surface mostly punctate;

metapleuron mostly punctate above; first recurrent vein interstitial in left forewing,

ending near apex of first submarginal in right wing.

Sternum VIII as in figure 278. Gonostyle with setae confined to apex (like fig.

281), ventral spine unusually long, curved (like figs. 281-282).

Length: 5.25 mm.

Variation in males (3 specimens):

UID 0.63-0.69X LID; OOD 0.25-0.40X HOD. Flagellomere I length slightly more

than 2.5X apical width. First recurrent vein received by second submarginal cell in

specimen from Ega. Length 5-5.5 mm.

Female (58 specimens):

Similar to male except: terga I-II usually silvery fasciate at least laterally, III

and IV occasionally weakly fasciate laterally. Clypeus same as convexifrons (fig.

270). UID 0.46-0.57X LID. OOD 0.20X HOD or less, occasionally nearly contiguous.

Flagellomere I length 2.33-2.75X apical width. Punctation of mesopleuron usually

much sparser along mesopleural suture (2-5 diameters apart, or even impunctate) and

sometimes beneath scrobe as well (fig. 280). Propodeal side punctation sometimes

evanscent. Most often first recurrent vein goes to submarginal cell I, and second

goes to submarginal cell II (right and left wings sometimes different), but sometimes

one or both recurrents interstitial. Length 4.5-6.5 mm.

Discussion:

The absence of cross—carinae on the propodeal dorsum of cooperi distinguishes it

from the very similar convexifrons (compare figs. 268-269), although occasional

specimens of cooperi have short cross-carinae along the median longitudinal carina.

Male genitalia have to be examined to separate cooperi from wasbaueri (compare

figs. 281-284 & 276-277). The long, curved spinose process of the gonostyle (fig.

281-282) is unique to cooperi. The process is much shorter in convexifrons and

wasbaueri. An additional feature of cooperi is the generally sparser mesopleural

punctation toward the mesopleural suture. This varies, however, and some specimens

are fairly uniformly densely punctate as in convexifrons and wasbaueri.

78 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

Evans’ (1972) record of a Pison (Entomopison) sp. near convexifrons from

Dominica refers to cooperi.

Range:

Pison cooperi, a commonly collected wasp, is known from Costa Rica, Panama,

and the West Indian island of Dominica. It occurs widely, across northern South

America and reaches northern Bolivia in the west and central Brasil in the east.

Types:

Holotype male: COLOMBIA, Amazonas: La Chorrera, VIII-14/23-1976, M. Cooper

(BMNH).

Paratypes (3 males, 57 females): COSTA RICA, Heredia: La Selva, 4 km se

Puerto Viejo, VII-21-80, ex trap nest M620-C1 (female, emerged VIII-19-80) and C-2

(male, emerged VIII-18-80), R. E. Coville (USNM); Puntarenas: Pta. Morales,

IX-10-87, W. E. Eberhard (COOPER). PANAMA: Cabima (now under the canal),

V-28-11, A. Busck (USNM). DOMINICA: Clarke Hall Est., V-30-66, G. Steyskal

(USNM); S. Chiltern, 1600', II-19-65, H. E. Evans (USNM). COLOMBIA, Amazonas: La

Chorrera, VIII-14/23-76, M. Cooper (BMNH); Leticia, VIII-21/23-74, VIII-29/1X-2-74,

M. Cooper (BMNH); El Encanto, IX-15/19-76, M. Cooper (BMNH); Boyaca: Muzo, 900

m, VII-36, J. Bequaert (MCZ); Magdalena: N. Sierra Nevada de S. Marta, Rio

Buritaca, 100 m, II-23/24-74 M. Cooper (BMNH); N. Sierra Nevada de S. Marta, Rio

Don Diego, XI-22-74, XII-3-74, M. Cooper (BMNH); Meta: Cord. Macarena,

II-15/28-76, M. Cooper (BMNH); Villavicencio, V-7-74, M. Cooper (BMNH); Narino:

Barbacoas, III-29-74, V-2/6-1976, VII-19/21-74, VIII-3-84, M. Cooper (BMNH,

COOPER); Putumayo: Villa Garzon, 8 mi. s Mocoa, VII-22/23-78, IX-8-87, M. Cooper

(BMNH, COOPER); La Hormiga near Mocoa, IX-6-78; Vaupes: Mitu, V-14-74, M.

Cooper (BMNH). ECUADOR, Napo: Muyuna, 5 km w Tena, 500 m, IV-10-81,

VIII-26-79, XI-24-78, M. Cooper (BMNH); Limoncocha, VI-15/28-76, S. & J. Peck

(CNC), Limoncocha, VI-3-77, D. Vincent (USNM); Tena, 500m, IV-11/28-76,

XII-9/14-71, M. Cooper (BMNH); Puerto Misahualli, 30 km e, 350 m, II-83, L. Masner,

M. Sharkey (PMA); Misahualli, 1 hour down Rio Napo, II-19-83, L. Huggert (PMA);

Laguna Jatuncocha, 20 km s Nuevo Rocafuerte on Rio Yasuni, II-5/7-86, Wojcicki pan

trap, Finnamore, Thormin, Blades (PMA); Morona-Santiago: Rio Upano nr. Patuca,

570 m, VII-14-87, M. Cooper (COOPER). PERU, Huanuco: Monson Valley, Tingo

Maria, X-21-54, Schlinger and Ross (CAS). BOLIVIA, Beni: Rio Mamore, about 5 km

nw mouth of Rio Grande, VIII-2-65, J. K. Bouseman (AMNH); Riberalta, XI-56, M.

Fritz (FRITZ). TRINIDAD: St. Augustine, IX-4-49, E. Callan (BMNH). GUYANA:

Mazaruni-Potaro Dist., Takutu Mountains, malaise trap, XII-3/10-83, Steiner and

Spangler (USNM); Mazaruni, VII-19-37, VII-28-37, IX-7-37, IX-21-37, Richards and

Smart (BMNH); Ryk-Over-Al, IX-17-37, Richards and Smart (BMNH); Blairmont,

VIII-23, F. X. Williams (BPBM); Bartica, V-17-01 (CU). SURINAME: Coppename

River, env. Voltz Mountain, VII-14-63, J. van der Vecht (LEIDEN). BRASIL,

Amazonas: Ega (= Tefé) (BMNH). Goias: Vila Vera, X-73, M. Alvarenga (AEI). Para:

Belém, VI-24, F. X. Williams (BPBM); Para (= Belém), X-26-01, A. Ducke (VIENNA).

Rio Branco: Vista Alegre, IX-6-24 (CU).

Metatype (one headless female): SURINAME: Paramaribo, I-1961, P. van

Doesburg (LEIDEN).

Etymology:

This wasp is dedicated to Martin Cooper, Pison collector extraordinaire.

Pison longicorne Menke, n. sp.

(Figs. 285-291)

Description, holotype male:

Body black, mandible tip and tarsi brownish. Appressed facial setae silver.

Terga I-III with silver apical fasciae.

Menke: Pison in the New World 79

labrum quadrangular, apex with shallow obtuse V notch. Clypeus as in fig. 286;

frons dull, densely punctate, with tiny polished area between midocellus and antennal

sockets; UID = 0.80X LID; OOD 0.83X HOD. Antenna elongate, length of articles

I-VI more than twice apical width (I 2.60X width), VII 2X apical width, VIII-X slightly

less than 2X apical width.

Pronotal collar with angular median prominence (similar to fig. 288). Scutum

and scutellum weakly shining, densely, finely punctate (punctures a diameter apart or

less). Propodeum shiny, dorsum with sparse pin prick punctures (3 or more diameters

apart) and extremely fine median longitudinal carina that is evanscent posterad;

hindface smooth, impunctate except dorsad; propodeal side with dense, fine

punctation similar to that on scutellum. Mesopleuron behind episternal sulcus shiny,

moderately punctate (punctures |l-2 diameters apart). Metapleuron smooth, very

finely punctate. First recurrent vein ending near apex of submarginal cell I, second

recurrent vein ending at apical third of submarginal cell II.

Sternum VIII apex with deep, arcuate emargination. Genitalia as in figs.

289-291; gonostyle broad, flat, outer surface setose (fig. 289); volsellar lobes

consisting of a pair of short, rounded structures fringed with setae (fig. 290).

Length 6.5 mm.

Variation in males (8 specimens):

Tarsi often black, appressed setae sometimes weakly golden. UID 0.72-0.85X

LID. OOD varies from 0.66X to slightly more than HOD, but usually OOD 0.75X HOD

or slightly more; single specimen in which OOD greater than HOD from Brasil, and is

largest male. Flagellomere I length 2.5-3.5X apical width. Second recurrent vein

nearly interstitial in most specimens, and is in one. Submarginal II incomplete in one

wing of specimen from Guyana. Specimens from Argentina and Santa Catarina,

Brasil have few feeble cross-ridges on propodeal hindface, but surface basically

smooth, shiny. Median longitudinal propodeal carina extends to apex in two

specimens. Length 6-8 mm.

Female (53 specimens):

Similar to male except: Legs completely black, flagellum sometimes brownish

beneath; appressed setae varying from silver to brassy. Clypeal lobe truncate with

corners rounded, its margin thickened (fig. 285). Labrum not notched. UID

0.56-0.72X LID. OOD 0.45-0.66X HOD. Flagellomeres longer, I 3.0-3.75X apical

width, VII nearly 3X apical width. Punctation of propodeal dorsum varies from sparse

to moderately dense (1-2 diameters apart) and coarser; median longitudinal carina

often strong, extending to apex; hindface usually punctate peripherally, often with

weak transverse ridges interrupted at midline. Submarginal cell II sometimes very

small, about equal to one hindocellus diameter. First recurrent vein rarely

interstitial (s Brasil), second recurrent rarely interstitial (s Brasil, Panama), both

recurrents end on submarginal II in one specimen (s Brasil). Terga I-III or IV with

fasciae. Length 8-10 mm.

Discussion:

Pison longicorne is most easily recognized by the elongate antennal

flagellomeres. All but the last are 3 or more times as long as wide in the female, and

2 or more times in the male. In other species of the convexifrons group the

flagellomeres shorten progressively toward the antennal tip, and the outer articles

are at most 2 times as long as wide in the female, less than that in the male. The

largely noncarinate propodeal hindface, the angular median prominence of the

pronotal collar (fig. 288), and the shape of the female clypeal lobe of longicorne (fig.

285) are distinctive within the convexifrons group. The male genitalia (figs. 289-291)

are also diagnostic. In particular the gonostyle is broad and flat distally, the outer

surface setose, and unlike other species in the group there is no spinose process

ventrally (fig. 289). The volsellar lobes, though small, are larger than in other species

of the group (fig. 290).

80 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

Range:

This commonly collected species occurs from northwestern Mexico to northern

Argentina.

Types:

Holotype male: PARAGUAY: San Bernardino, October, K. Fiebrig (VIENNA).

Paratypes (9 males, 52 females): MEXICO, Jalisco: Plan de Barrancas, VIII-9-65,

H. E. and M. A. Evans (MCZ); Chiapas: Palenque Ruins, VI-22-69, B. V. Peterson

(CNC). COSTA RICA: Zent, III-1924, H. W. Atkinson (USNM); Turrialba, XII-?, Heyne

(BERLIN); Golfito, XII-1948, P. and D. Allen (MCZ). PANAMA: Barro Colorado I.,

VIT-14-39, V-1939, Zetek (USNM); Barro Colorado I., III-26-24, J. C. Bradley (CU).

COLOMBIA, Vaupes: Mitu, V-17-74, M. Cooper (BMNH); Putumayo, Mocoa,

V-31/VI-7-76, I-l0O-77, M. Cooper (BMNH); Villa Garzon, IX-1-84, M. Cooper

(COOPER); Amazonas: La Chorrera, VIII-14/23-76, M. Cooper (BMNH). GUYANA:

Kartabo, VII-26-24, W. Myers (CMP); 50 mi. point, Bartica—Potaro road, VIII-30-37,

nest 160 & 160A, Richards and Smart (BMNH). SURINAME: Sipaliwini, VI-14-63, P.

van Doesburg (LEIDEN). PARAGUAY: San Bernardino, K. Fiebrig (BERLIN); sw

Saltos del Guaira, XII-8-71, L. Peha (AMNH). BRASIL, Guanabara: Reprusa Rio

Grande, X-1969, M. Alvarenga (AEI); Rio de Janiero: Itatiaia, XI-l6-39, Zikan

(LILLO); Santa Catarina: Nova Teutonia, II, XI, XII, various years, F. Plaumann (MCZ,

AMNH, CU, UCD, USNM); Cauna, XII-1945, A. Maller (AMNH); Pinhal

XI-XII-1947-1948, A. Maller (AMNH); Rio Grande do Sul: no locality, Stieglmayr

(VIENNA). ARGENTINA, Tucuman: Rio Los Sosa, XI-25-65, W. Weyrauch (LILLO);

San Pedro de Colalao, Foerster (FRITZ).

Etymology:

The name longicorne is based on the the Latin words longus and cornu and it

refers to the distinctive long antenna of this wasp.

Biology:

The Guyana specimens listed above emerged from "brittle clay cells in old

Metapolybia nest under envelope" according to field notes 160 and 160A by O. W.

Richards (in BMNH).

Pilosum Group

(Figs. 6, 13-14, 292-340)

Description:

Frons not swollen; eye asetose; male antenna simple, flagellum without tyli or

other modifications; lower half of female clypeus with long, somewhat decumbent

setae that project over clypeal margin forming a "brush" (fig. 6); female clypeal lobe

projecting, apex rounded to acutely triangular, often double-edged at apex (figs. 298,

323, 333); male clypeus with truncate or angular median lobe that has rounded tooth

(figs. 300, 334); labrum usually with V or U-shaped emargination; mandible with

posterior notch (fig. 6), female mandible with mesal tooth or cleft on cutting edge;

occipital carina incomplete ventrad, ending on or near hypostomal carina; anterior

edge of pronotum with polished, free lamella that extends laterad to level of pronotal

lobe (figs. 294-295), no pit behind lamella; scutal flange broad, somewhat reflexed;

tegula impunctate on outer half, punctate adjacent to scutum; propodeum usually

_with foveolate ridge or linear series of foveae laterally that defines upper limit of

side (except in sphaerophallus and some pilosum); propleuron punctate, disk not

largely polished; episternal sulcus sometimes evanescent ventrad as it curves toward

anterior border of pleuron; metapleural flange broad, as wide as hindocellus or

broader (fig. 302); forewing media diverging from M+Cu after crossvein cu-a;

marginal cell acuminate distally; three submarginal cells present, III strongly

narrowed toward marginal cell, its veinlets often joining there or even forming

petiole; first recurrent vein ending on submarginal cell I or II, second ending

Menke: Pison in the New World 81

submarginal II; hamuli of hindwing divided into two groups; dorsum of hindcoxa

usually delimited by inner and outer carinae (outer carina rarely absent), inner carina

usually lamelliform in part; tarsomeres with plantulae; male foretrochanter with

(figs. 292-293) or without spine; anterior claw of male foreleg (and sometimes

midleg) distorted, usually broadly lamelliform apically (fig. 296-297); gaster without

yellow tergal bands; tergum I sometimes double-edged (qnythos, sphaerophallus);

male tergum VII often laterally compressed at apex forming rounded longitudinal

ridge; male sternum VII sometimes with oval or round asetose depression, VIII rounded

or angulate apically (figs. 303-306); genitalia robust, gonostyle represented mostly by

large, broad plate, usually with short, apical setose extension (figs. 307, 311, 315,

319, 329), volsellar lobes small or scarcely defined (figs. 308, 312, 317, 330), penis

valves separating distad (dorsal view) forming a broad aedeagus.

Included species:

Pison aureofaciale, gnythos, oaxaca, pilosum, sphaerophallus and vincenti.

Discussion:

The most obvious synapomorphies defining the pilosum group are: the polished,

free lamella that extends around the anterior edge of the pronotum; the broad

metapleural flange; and the distortion of the anterior claw of the male foreleg. The

last feature is an autapomorphy, at least among New World Pison. The apical setose

lobe seen on the gonostyle in other groups of New World Pison is greatly reduced in

most species of the pilosum group and the gonostyle is represented mostly by a large,

broad plate. I regard these and the asetose depression on male sternum VII found in

three species as specializations. Other apomorphic trends in the pilosum group are:

labrum usually with U-shaped median notch, the tendency for the episternal sulcus to

evanesce as it curves forward ventrally, the presence of a spine on the male

foretrochanter of some species, the double-edged tergum I in two species, and the

laterally compressed male tergum VII of some species.

The six species of the group segregate into several lineages. The largest of these

is the pilosum species complex (aureofaciale, pilosum, vincenti) in which females are

nearly impossible to distinguish with certainty without associated males. These three

species have one synapomorphy: male foretrochanter with spine distally. Pison

Oaxaca is linked to the pilosum complex via the following synapomorphies: tergum I

strongly humped and the joint between it and tergum II strongly constricted when

viewed in lateral profile (fig. 301), male tergum VII laterally compressed, and male

sternum VII with depression.

Pison gnythos and sphaerophallus share a double-edged tergum I, a

synapomorphy, but the latter species has a spine on the male foretrochanter at the

middle, an autapomorphy. The gonostyli of sphaerophallus are completely fused,

forming a globose genital capsule, another obvious autapomorphy. In sphaerophallus

the volsellar lobes are long and finger-like and the gonostyle has a long, slender,

setose, arcuate appendage apically. If the absence of a median longitudinal carina on

the propodeal dorsum and absence of a line of foveae at the top of the propodeal side

are loss features (i.e., reversals), then these states are additional apomorphies of

sphaerophallus. The large, shiny facial "dimple" of gnythos (fig. 323) which probably

has a gladular function, is an autapomorphy that isolates it from sphaerophallus. The

genitalia of gqnythos are more like those of the pilosum complex.

Pison pilosum Smith

(Figs. 13-14, 292, 294-296, 298-303, 307-310)

Pison pilosus Smith, 1873:295. Holotype female: Ega (= Tefé, Brasil)(BMNH).

Description, male (14 specimens):

Body black, tarsomeres II-IV brown (Mexico) or I-IV light brown or yellowish

82 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

(Panama, Colombia, Trinidad, Guyana, Suriname). Following often brownish or

yellow: apex of hindtibia beneath, base of tergum VII and sterna VI-VII entirely.

Flagellomeres V-XI with brown spots beneath (Panama). Appressed facial setae pale

golden, dense, obscuring sculpture except on disk of frons; gena with similar

vestiture. Terga I-II and sometimes III with apical silver fasciae that are broadly

interrupted at midline.

Clypeal lobe with median tooth (fig. 300); labrum with U-shaped notch. Frons

dull, shallowly, densely punctate, punctures smaller than those on scutum, interspaces

granular; middle of frons with narrow, polished spot on midline. UID 0.57-0.62X

LID. OOD 0.41-0.52X HOD. Flagellomere I length about 3X apical width, II slightly

less than 3X apical width; III-X progressively shorter, outer articles only slightly

longer than wide.

Thorax shiny. Scutum densely punctate, punctures averaging less than a

diameter apart; scutellum similarly punctured, though punctures slightly smaller,

sparser; metanotum impunctate. Propodeal dorsum with median, longitudinal carina

that fades posterad, carina in depression, surface of dorsum smooth, punctate,

punctures separated by one to three diameters; punctation of dorsum extending onto

hindface which has a shiny, vertical, median impression and several cross-ridges just

above petiole socket; propodeal side smooth, punctate, punctures sparser ventrad,

side delimited dorsally by foveolate ridge. Mesopleuron smooth, densely punctate,

punctures beneath scrobe somewhat larger than those of scutum, punctures separated

by less than diameter above scrobe and in front of episternal sulcus, elsewhere up to

2 or 3 diameters apart; metapleuron with sparse to dense, fine, pinprick punctation.

Precoxal lobes delimited at base by deep foveolate sulcus. Forewing venation

variable, first recurrent vein ending on submarginal cell I or II or interstitial, these

three patterns of about equal occurrence; veinlets of submarginal cell III joining at

marginal cell or nearly so. Foretrochanter with sharp, posterior spine at distal third

(fig. 292). Anterior claw of foreleg distorted, with broad lamella apically (fig. 296),

same claw of midleg simple.

Tergum I in lateral profile strongly humped posterad, joint between I & II

constricted (similar to fig. 301); terga II-III each with pair of slightly arcuate,

transverse swellings just before apical margin; tergum VII laterally compressed at

apex forming rounded longitudinal ridge. Sternum VII flat, without basomedian

depression (fig. 303), sternum VIII as in fig. 303. Genitalia as in figs. 307-310;

gonostyle in lateral profile with short, setose apical lobe (fig. 307), which, in ventral

view (figs. 308-309), is merely the lateral edge of setose concavity, free (distal) edge

of concavity with small, median process (fig. 309); volsellar lobe small, delimiting

inner border of gonostylar concavity in ventral view (fig. 308).

Length: 6-7 mm.

Female (65 specimens):

Similar to male except: legs, antenna, and terminal gastral segments black

(apical margins of terga sometimes pale); appressed facial and thoracic setae

sometimes silvery, terga III-IV often with complete apical fasciae. Clypeus with

obtusely triangular or rounded median lobe that is double-edged at apex and often

somewhat translucent (figs. 298-299). UID 0.46-0.55X LID. OOD 0.29-0.48X HOD.

Flagellomere I length about 4X apical width or slightly less, II about 3X apical width

or slightly less, outer articles progressively shorter, X about 1.5X apical width.

Thoracic punctation sometimes more uniformly dense than in male, sometimes most

punctures nearly contiguous. Propodeal hindface often transversely punctatorugose

nearly to top; foveolate ridge of propodeal side often reduced to linear series of large

pits (fig. 302), or even absent. Both recurrent veins usually ending on submarginal

cell II, occasionally recurrent I interstitial between submarginals I and II, or rarely I

ends on submarginal cell I. Swellings of terga II-III sometimes weak, especially on

III. Length: 6-8.5 mm.

Menke: Pison in the New World 83

Discussion:

Pison pilosum is the most commonly collected species of the pilosum group.

Females of pilosum, aureofaciale and vincenti are extremely similar and I have been

unable to find morphological differences between them. Males of these three can be

separated by differences in sterna VII and VIII, and genitalia.

The only definite male/female associations of pilosum before me were reared

from nests taken at St. Augustine, Trinidad (BMNH), the Canal Zone (USNM), and

Paramaribo, Suriname (LEIDEN). I have been unable to find any differences between

these females and the single female of vincenti, so it is quite possible that some of

my females of “pilosum" are vincenti. Sternum VII is flat basally in males of pilosum

(fig. 303), sternum VIII is moderately narrow toward the apex (fig. 303), and the

genitalia (figs. 307-310) are distinctive. Females that I have associated with these

males have completely black antennae, as does the single female of vincenti known to

me (pinned with its presumed male on one pin). Females of aureofaciale have pale

areas ventrally on flagellomeres I and II and sometimes III, and the appressed facial

setae are more golden than in pilosum and vincenti.

Males of pilosum also differ from those of aureofaciale and vincenti in having an

unmodified anterior claw on the midleg. This claw is distorted and apically laminate

in the latter two species. !

Unfortunately Smith's type of pilosum is a female, and, although I have examined

it, I cannot be certain that it is conspecific with what I am interpreting as that

species.

Range:

I have seen males of pilosum from southern Mexico (Chiapas), Costa Rica,

Panama, Colombia, Trinidad, Guyana and Suriname. Females that I have interpreted

as this species have been taken within that area, but also as far north as Jalisco,

Mexico, and as far south as Bolivia and Santa Catarina in Brasil.

Material examined (79 specimens):

MEXICO, Nuevo Leon: Sierra de Picachos, 7 mi. s Cerralvo, 2, (USNM); Jalisco:

Chamela, ? (USU); Chiapas: Escuintla, 6 (USNM). COSTA RICA, Guanacaste: Rio

Corbici nr. Canas, ? (UCD); Bebedero, ? (MCZ); Santa Rosa N. Park, ¢, 2? (BMNH);

Heredia: F. La Selva, 3 km s Pto. Viejo, dé (USNM). PANAMA, Canal Zone: Barro

Colorado I., ? (UCD); ex Canal Zone, intercepted at Brownsville, Texas in Cattleya

gigas, 6, 2? (USNM). COLOMBIA, Antioquia: Fredonia, ¢ (USNM), Valle Medellin, ¢

(USNM); Meta: Cord. Macarena, ? (BMNH); Valle: Mun. Candelaria, Finca San Luis, ?

(CSDA, FSDA); Magdalena: N. Sierra Nevada de S., Marta, Rio Buritaca, ? (BMNH);

Amazonas: Leticia, ? (BMNH, CNC); Vaupes: Mitu, ? (BMNH); Caqueta: Yuruyaco, 73

km sw Florencia, ? (BMNH); Putumayo: Mocoa, ? (BMNH). VENEZUELA, Monagas:

Rio Marichal Largo (puente), @ (UCM). TRINIDAD: St. Augustine, 6, ? (BMNH);

Curepe, 2? (PMA, CNC). GUYANA: Blairmont, 6, 2? (BMNH); Georgetown, ? (BMNH),

Kaieteur, High Forest, ? (BMNH). SURINAME: Paramaribo, 6, ? (LEIDEN, BERLIN).

BRASIL, Para: Belém, ? (CSDA, BPBM, VIENNA, USNM); Santarem, ? (CMP);

Tabatinga, @? (VIENNA); Guanabara: Rio de Janeiro, 2? (USNM); Sao Paulo: Teodoro

Sampaio, ? (FRITZ); Santa Catarina: Nova Teutonia, ? (UCD). ECUADOR, Napo:

Limoncocha, ? (CNC, FSDA, USNM); Tena, 2? (BMNH, BPBM, PMA). PERU, Loreto:

Pucallpa, ? (BMNH); Huanuco: Tingo Maria, ? (CAS); Pasco: Puerto Bermudez, ?

(CU). BOLIVIA, Beni: Rio Mamore 10 km e San Antonio, ? (AMNH); Santa Cruz:

Puerto Grether, ? (COOPER).

Biology:

Some specimens bear labels indicating that they were reared from mud cells. F.

X. Williams noted that one female that he reared in Guyana “hatched ... from

Pseudagenia-like cell mass [on] underside of leaf".

84 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

Pison aureofaciale Strand

(Figs. 304, 311-314)

Pison aureofaciale Strand, 1910:174. Lectotype male: Asuncion, Paraguay, present

designation. (BERLIN).

Description, male (6 specimens):

As described for pilosum except: pale markings variably developed on antenna:

scape and pedicel varying from light brown to black, flagellomeres I-IV, or V, or VII

light brown or yellowish; apical margins of terga and sterna II-VII pale. Appressed

facial setae pale to bright golden, dense, obscuring surface of frons; tergal fasciae

pale golden, present on I-VI, that of I interrupted at middle. UID 0.58-0.65X LID.

OOD 0.47-0.63X HOD. Flagellomere I length slightly more than 3X apical width, II

3X apical width. First recurrent vein ending on submarginal cell II or interstitial

between I and II; third submarginal cell sometimes narrowly petiolate at marginal

cell. Anterior claw of midleg distorted, apically laminate. Sternum VII with circular

depression at base (fig. 304), VIII narrowing to fairly sharp apex (fig. 304). Genitalia

as in figs. 311-314; gonostyle with short, setose apical lobe whose apex is bent

ventrad hooklike (ventral view, figs. 312-313); volsellar lobes discrete.

Length 5.5-7 mm.

Female (7 specimens):

Similar to male except: Scape’ and pedicel black, flagellomeres I-II, and

sometimes III, pale, at least beneath at apex; legs black; clypeal lobe obtusely

triangular, narrowly double-edged at apex, often pale, translucent; UID 0.52-0.55xX

LID; OOD 0.38-0.51X HOD; flagellomere I length 3.5-4X apical width, II 3X apical

width or slightly less; first recurrent vein occasionally ending on submarginal cell ];

tergal swellings sometimes weak, especially on III; length 7-7.5 mm.

Discussion:

This species is so similar to pilosum that I first thought aureofaciale was nothing

more than a geographic color form of it. But the male genitalia (figs. 311-314) and

sterna VII-VIII (fig. 304) are quite different. The pale areas on the antenna aid in the

identification of aureofaciale, but the variation observed in the few specimens

available suggest that this feature is not likely to hold up over the entire range of the

species, especially in the female. I cannot find any morphological differences

between females of aureofaciale and pilosum.

I have examined Strand's 3 male and 3 female syntypes of aureofaciale and

labeled one male as lectotype.

Range:

Paraguay and Brasil. Brasilian material is represented by a male and female

labeled simply "Bahia". There are several sites with this name in Brasil, but I am

assuming that the one in the province of Santa Catarina is the most likely for these

specimens.

Material examined (13 specimens):

PARAGUAY: Asuncion, 6, ? (BERLIN); San Bernardino, ¢, ? (BERLIN); San Pedro

Cororo, Rio Ypane, ¢ (CSDA); Col. Piarerata, @? (FRITZ). BRASIL: Bahia, 6, ?

(PARIS).

Pison vincenti Menke, n. sp.

(Figs. 305, 315-318)

Description, holotype male:

Same as pilosum except: Body entirely black except tergal margins II-IV pale;

appressed facial setae silver, dense except on disk of frons; UID 0.54X LID; OOD

Menke: Pison in the New World 85

0.52X HOD; flagellomere I length slightly more than 3X apical width, II 3X apical

width; punctures of propodeal dorsum separated by puncture diameter of less on

average; mesopleural punctures beneath scrobe about same size as those on scutum

and separated by puncture diameter or less; forewing membrane weakly stained in

cellular area, first recurrent vein ending on first submarginal cell; veinlets of

submarginal cell III meeting on marginal cell; anterior claw of midleg distorted,

laminate apically; terga II-III without arcuate swellings near apical margin; sternum

VII with deep, elongate depression (fig. 305), VIII broadly rounded apically (fig. 305);

male genitalia as in figs. 315-318; gonostyle with short, setose apical lobe whose apex

is truncate in ventral view (fig. 317), and bent ventrad as digiform process (figs.

315-316); volsellar lobes distinct.

Length 7 mm.

Male variation (1 specimen):

Foretarsomeres III-IV brownish; facial setae and tergal fasciae weakly golden;

UID 0.58X LID; forewing membrane unstained in cellular area; terga II-III each with

pair of arcuate, transverse swellings just before apical margin.

Female (1 specimen):

Like holotype except: UID 0.5X LID; OOD 0.43X HOD; flagellomere I length 4X

apical width, II slightly less than 4X apical width; clypeus with obtusely triangular

median lobe that is double-edged at apex and somewhat translucent; forewing

membrane not stained in cellular area, first recurrent vein ending on second

submarginal cell; terga II-III each with pair of arcuate, transverse swellings just

before apical margin; length 8.5 mm.

Discussion:

The male of vincenti is easily recognized by the distinctive elongate depression

of sternum VII (fig. 305) and the male genitalia (figs. 315-318). The female, however,

is inseparable from pilosum. The single female known is pinned together with the

paratype male and presumably they were caught in copula. The male genitalia of

vincenti are similar to aureofaciale but the apex of the gonostyle is truncate in

ventral view (fig. 317). The gonostyle is more rounded apically in aureofaciale (fig.

313), and the form of the depression on sternum VII and shape of sternum VIII are

different in the two species (compare figs. 304-305).

Range:

Known only from Ecuador and Guyana.

Types:

Holotype male: ECUADOR, Napo: Tena, V-24-1977, D. Vincent (USNM).

Paratypes (1 male, 1 female): GUYANA: Kartabo, Bartica District, IX-6-20

(AMNH).

Etymology:

I take pleasure in naming this wasp after David Vincent, collector of the

holotype.

Pison oaxaca Menke, n. sp.

(Figs. 306, 319-322)

Description, holotype male:

Black; tarsomeres II-IV pale; face with appressed silver setae that obscure

sculpture except on disk of frons; terga without obvious fasciae.

Labrum quadrangular but with deep U-shaped emargination; distal half of stipes

(maxilla) without depression; clypeus with obtusely angular median lobe; frons dull,

86 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

closely punctate, interspaces granulate, punctures smaller than those on scutum; UID

0.58X LID; 00D 0.50 HOD; flagellomere I length 3X apical width, II nearly 3X apical

width; distal flagellomeres slightly longer than wide.

Thorax shiny; scutum densely punctate, punctures separated by less than

puncture diameter; scutellum gibbous, punctured similar to scutum but punctures

slightly smaller; metanotum with fine setigerous punctures; propodeal dorsum with

median longitudinal carina that fades posterad, rest of surface punctate, punctures

similar to those of scutum but sparser, one to two diameters apart, propodeal

hindface with similar punctation and few transverse ridges near petiole socket,

propodeal side more sparsely punctate than dorsum, side delimited dorsad by

foveolate ridge. Mesopleural punctation as dense as on scutum, but slightly coarser;

metapleuron densely punctate, but punctures much finer than those of mesopleuron;

precoxal lobes delimited at base by deep, foveolate sulcus. First recurrent vein

interstitial between submarginal cells I-II; veinlets of submarginal cell III almost

meeting on marginal cell. Foretrochanter without spine. Anterior claw of foreleg

distorted, laminate apically, same claw of midleg unmodified. Hindcoxa with

lamelliform outer dorsal carina.

Tergum I in lateral profile strongly humped, joint between I-II constricted; terga

II-III with weak paired swellings near apical margins. Tergum VII compressed

laterally at apex forming rounded longitudinal ridge. Sternum VII with circular

depression (fig. 306), VIII broadly triangular at apex (fig. 506). Genitalia as in figs.

319-322, gonostyle with short, setose apical lobe whose inner margin bears a short

spine-like process (figs. 320-321).

Length 6.5 mm.

Female:

Unknown.

Discussion:

Pison oaxaca is only known from the holotype. The species shares most of the

characters of the pilosum species complex, but the foretrochanter of oaxaca lacks a

spine. This, combined with the circular depression of sternum VII, the shape of

sternum VIII and the appearance of the genitalia, makes oaxaca easily recognizable.

The obtusely angular form of the median lobe of the clypeus also seems diagnostic.

In the pilosum species complex the median lobe of the male is basically quadrate with

a prominent point in the middle (fig. 300).

Range:

Known only from southern Mexico.

Type:

Holotype male: MEXICO, Oaxaca: Temascal, X-17-1963, K. H. Janzen (USNM).

Etymology:

The name oaxaca, a noun in apposition, is the Mexican state in which the the

species was collected.

Pison qnythos Menke, n. sp.

(Figs. 6, 323-332)

Description, holotype male:

Black; face with appressed silver setae that obscure sculpture except on disk of

frons; terga without obvious fasciae.

Labrum quadrangular but with deep U-shaped emargination; distal half of stipes

(maxilla) with large, deep, depression; clypeus similar to that of pilosum (fig. 300);

frons dull, granulate, with polished dimple on midline above antennal sockets that is

Menke: Pison in the New World 87

almost as large as midocellus; UID 0.39X LID; OOD 0.20X HOD; flagellomere I length

4X apical width, II 3X apical width; (left antenna missing, right one missing

flagellomeres III-XI).

Thorax shiny; scutum densely punctate, punctures separated by less than

puncture diameter; scutellum gibbous, punctation similar to scutum but somewhat

sparser and finer; metanotum without obvious punctation. Propodeal dorsum with

median, longitudinal carina in depression, rest of dorsum with fine, setigerous

punctation that becomes denser, coarser laterad; propodeal hindface punctate above,

with several cross-ridges below; propodeal side punctate, punctures becoming

progressively finer and denser ventrad, side delimited dorsally by row of foveae.

Mesopleural punctation above scrobe and before episternal sulcus dense, but finer

than that of scutum, rest of pleuron with larger, sparser punctures (1-3 diameters

apart); metapleuron with dense, pinprick punctures above, impunctate and smooth

below. Precoxal lobes delimited by foveolate sulcus. First recurrent vein ending on

submarginal cell I; veinlets of submarginal cell II] separated on marginal cell.

Foretrochanter without spine. Anterior claw of foreleg distorted, lamelliform.

Hindcoxa without outer dorsal carina.

No strong constriction between terga I-II of gaster; tergum I with subapical

beadlike rim. Tergum II with with pair of arcuately transverse swellings near apical

margin, III with similar but weak swellings. Tergum VII not laterally compressed.

Sternum VII flat, VIII bluntly rounded apically (fig. 328). Genitalia as in figs.

329-332; gonostyle with short, setose apical lobe whose inner margin bears a hooklike

lobe (figs. 330-331); volsellar lobes weakly defined (fig. 330).

Length 7 mm. »

Female (15 specimens):

Similar to male except: clypeus with dense, appressed silver hair only laterally;

tergum I with silver fascia laterally, II-IV with tarnished silver fasciae laterally;

forewing membrane sometimes lightly stained; labrum with shallow median

emargination, or edge entire; clypeus with rounded median lobe (fig. 324); frons with

median polished dimple that usually contains small pit (fig. 323), dimple variable in

size, somewhat smaller than to larger than midocellus, and delimited on lower edge

by tubercular swelling that is usually surmounted by short carina (fig. 325); UID

0.35-0.40X LID; OOD 0.20 HOD or less; flagellomere I length 4X apical width or

more, II 3X apical width or more, flagellomeres VII-IX about 2X apical width; thorax

somewhat more densely punctate than male, especially on propodeum and

mesopleuron; carina of propodeal dorsum often with short cross-carinae; first

recurrent vein ending on submarginal cell II or interstitial between I-II; veinlets of

submarginal cell III joining at marginal cell, sometimes cell narrowly petiolate there;

hindcoxa with low, outer carina on dorsum; length 6.5-8 mm.

Discussion:

Females of gnythos are usually easily identified by the large polished dimple-like

facial depression (fig. 323), and the form of the clypeal lobe (fig. 324). The facial

dimple is present in males but it is comparatively smaller. The male has a simple

foretrochanter and the genitalia are diagnostic (figs. 329-332). The upper interocular

distance is narrower than in other species of the pilosum group (UID 40% of LID or

less in gnythos), and the hindocellus is very narrowly separated from the eye (OOD =

0.20X HOD or less). In both sexes tergum I has a bead-like rim just before the free

margin (figs. 326-327). This character state is nearly identical to the double-edged

tergum I in sphaerophallus. The apical depression on the stipes, the weakly

emarginate female labrum, and absence of an outer carina on the hindcoxa of the

male are other distinctive characters of qnythos.

The median carina and the lateral foveolate ridge of the propodeum suggest a

close relationship with the pilosum complex, but the absence of a strong constriction

between terga I-II, the non-compressed tergum VII of the male, and simple male

foretrochanter isolate gnythos.

88 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

Range:

Known from Colombia, Ecuador, Trinidad and Guyana.

Types:

Holotype male (pinned with female): GUYANA, Kartabo, IX-4-1922 (AMNH).

Paratypes (15 females); COLOMBIA, Putumayo: Mocoa, III-26/IV-6-76,

I-1/10-77, M. Cooper (BMNH); Villa Garzon, 8 mi. s Mocoa, VII-22-78, VIII-31, Ix-29,

X-7/13-84, M. Cooper (BMNH, COOPER); Caqueta: Yuruyaco, 73 km sw Florencia,

II-12-79, M. Cooper (BMNH). ECUADOR, Morona-Santiago: Cord. de Cutucu, 6 kme

Macas, 1100 m, V-14-81, M. Cooper (COOPER); Napo: Muyuna, 500 m, 5 km w Tena,

VIII-23-81, M. Cooper (COOPER). TRINIDAD: IV-30/V-13-33, D. Vesey-Fitzgerald

(BMNH). GUYANA: Kartabo, IX-4-22 (AMNH).

Etymology:

The Greek noun gnythos means pit or hollow, a reference to the distinctive large

dimple on the female frons.

Pison sphaerophallus Menke, n. sp.

(Figs. 293, 297, 333-340)

Description, holotype male:

Black, hindtarsomeres III-IV brownish beneath. Appressed facial setae silver,

dense only laterally on clypeus and lower frons. Terga II-III with tarnished silver

fasciae laterally.

Labrum quadrangular but with deep, median V-shaped notch. Distal half of

stipes flat. Clypeus as in fig. 334; frons dull, surface minutely granulate, impunctate,

with small polished area on midline above antennal sockets; UID 0.58X LID; vertex

with transverse impression just behind ocelli; OOD 0.50X HOD; flagellomere I length

nearly 4X apical width, II nearly 3X apical width, VI-IX about as long as wide.

Thorax shiny; scutum densely punctate, punctures separated by less than

puncture diameter, nearly contiguous; scutellum gibbous, more finely, sparsely

punctate than scutum, punctures one to four diameters apart, especially posterad;

metanotum with scattered pinprick punctures. Propodeal dorsum with sparse

setigerous micropunctures separated by 3 or 4 diameters, surface smooth, without

median carina; rest of propodeum similarly punctate but punctures larger on hindface

and upper part of side; propodeum without foveolate ridge at top of side (similar to

fig. 337). Mesopleural punctation above scrobe and before episternal sulcus dense,

similar to that of scutum, but punctures sparser and larger over rest of pleuron,

separated by up to 3 diameters, interspaces smooth; metapleuron finely, sparsely

punctate; precoxal lobes delimited at base by transverse carina and foveolate sulcus.

First recurrent vein of forewing ending near apex of first submarginal cell; veinlets

of third submarginal cell joining at marginal cell. Foretrochanter with sharp spine

beneath near middle (fig. 293). Anterior claw of foreleg distorted but apical lamella

narrow, claw with angular basal tooth (fig. 297), same claw of midleg unmodified.

No strong constriction between terga I-II of gaster, tergum I narrowly

double-edged apically, terga II-III each with pair of arcuately transverse swellings

near apical margin; tergum VII not laterally compressed; sternum VII with vague,

basomedian depression, VIII broadly, bluntly rounded at apex. Genitalia as in figs.

338-340, gonostyli fused forming a spherical box upon which the volsellar lobes (fig.

339) and elongate gonostyli (fig. 338) appear as slender appendages.

Length: 6.5 mm.

Variation in males (7 specimens):

Tarsomeres II-IV sometimes almost entirely pale; UID 0.58-0.63X LID; OOD

0.44-0.71X HOD; flagellomere I length 3-3.5X apical width, II sometimes only

Menke: Pison in the New World 89

slightly more than 2X apical width; propodeal side sometimes largely impunctate

below, surface smooth, glossy; mesopleuron punctation beneath scrobe sometimes

denser, punctures 1-2 diameters apart; metapleuron sometimes impunctate below;

veinlets of third submarginal cell widely separated on marginal cell in one specimen.

Female (13 specimens):

Similar to male except: body completely black, forewing membrane sometimes

lightly stained; clypeus with acute triangular median lobe (fig. 333); UID 0.52-0.57X

LID; flagellomere I length 4X apical width or slightly less, II 3X apical width or

‘slightly less. Length 6.5-7.5 mm.

Discussion:

Pison sphaerophallus is the most easily identified species in the pilosum group.

The absence of a median carina on the smooth propodeal dorsum (fig. 337), and

absence of a lateral foveolate propodeal ridge are unique features. The double-edged

free margin of tergum I (figs. 335-336), the median position of the male

foretrochanter spine (fig. 293) and acutely triangular median lobe of the female

clypeus (fig. 333) are additional characters of sphaerophallus. The spheroid male

genital capsule is particularly unusual and immediately diagnostic (figs. 338-340).

The distortion of the anterior male foreleg claw is less developed in sphaerophallus

than other species of the pilosum group, but the presence of an angular tooth-like |

prominence at its base is distinctive (fig. 297).

Range:

Known from Colombia, Ecuador, Peru, Guyana, Suriname and the Amazon basin

of Brasil.

Types:

Holotype male: COLOMBIA, Meta: La Macarena, XI-20/29-1976, M. Cooper

(BMNH).

Paratypes (7 males, 13 females): COLOMBIA, Meta: La Macarena, XI-20/29-76,

M. Cooper (BMNH); Putumayo: Mocoa, X-30-74 (biol. note #259), M. Cooper

(BMNH), Villa Garzon, 400 m, IX-6, X-6/13-84, M. Cooper (COOPER). ECUADOR,

Napo: Muyuna, 5 km w Tena, XI-30-78, M. Cooper (BMNH); Pastaza: Rio Pastaza

road between Puyo and Macas, 900 m, V-27-87, M. Cooper (COOPER). PERU,

Huanuco: Tingo Maria, X-5-46, Weyrauch (USNM), Tingo Maria, X-9-54, Schlinger &

Ross (CAS). GUYANA: Mazaruni, nest 53A, VIII-19-37, Richards & Smart (BMNH).

SURINAME: 1963, D. Geyskes (LEIDEN). BRASIL, Para: Belém, III-10-07, A. Ducke

(VIENNA); prov. unknown: Rio Jaravi, Estirao do Equador, X-1979, Alvarenga (FRITZ).

Etymology:

The name sphaerophallus, a noun in apposition, is formed from the Greek

sphaero- meaning ball, and the Greek word phallos meaning penis. The name refers

to the distinctive male genitalia.

Biology:

Martin Cooper found a mud nest of sphaerophallus on the under side of an

epiphyte leaf in primary forest (biological note #259, BMNH). The nest was oval and

flattened and contained about 10 cells arranged in two parallel rows. Four adults

were reared from the nest. The cocoon walls were made up of microsized sand grains

presumably held together with silk.

The Guyana material listed above came from a nest, but I have not had access to

the field notes (nest #53A, BMNH).

90 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

APPENDIX

Pison laeve Smith

Pison laevis Smith, 1856:317. Holotype male: "Georgia" (BMNH).

Pison laeve, Krombein, 1979:1641. Doubts North American origin, suggests New

Georgia in Solomon Islands.

Discussion:

I have examined Smith's type of laeve, and it is unlike any New World species

known to me. The provenance of the type certainly was not the state of Georgia, and

Krombein (1979) suggested that it may have originated on one of the islands of

Melanesia or Micronesia. The type of laeve does resemble some of the smooth, shiny,

sparsely punctate species from the western Pacific Ocean that lack a dorsolateral

propodeal ridge such as nigellum Krombein and insulare Smith, but the very broad

scutal flange of laeve differentiates it from them. Colin Vardy (pers. comm.) says

that he has compared the type of laeve with "all our named species” and it is not

conspecific with any of them.

Some of the characteristics of laeve are: black, wings moderately infumate, body

with scattered erect pale setae, silver setae restricted to lower gena and lower face;

occipital carina incomplete, ending ventrad; labrum quadrangular but with mesal

emargination; clypeal lobe triangular (similar to male of insulare but not as

prominent as in that species); lower frons and clypeus with appressed silver setae;

frons dull, granular in appearance, impunctate; UID 0.64% LID; OOD = 0.22 HOD;

flagellomeres elongate, flagellomere I length 3X apical width; thorax smooth, shiny,

sparsely punctate, punctures coarsest on mesopleuron; pronotum with simple median

pit just behind anterior margin; upturned flange at lateral margin of scutum very

broad, equal to diameter of hindocellus; propodeum smooth, shiny, sparsely punctate,

dorsum with median longitudinal impression but no carina, dorsum without ridge

laterally; dorsum of hindcoxa with low inner carina but no outer carina; terga shiny,

sparsely, shallowly punctate, without fasciae; tergum VII truncate; sternum VIII

emarginate, similar to insulare.

New species of Pison from New Guinea

Two new species are described here because they share two apomorphic features

that occur rarely in the genus. The eye facets are considerably enlarged toward the

inner orbits (figs. 341-344), and gastral segment I is claviform (figs. 347-350).

Enlarged eye facets occur in the Australasian species ignavum and ruficorne but the

enlargement in these species is not nearly as striking as in the two described below.

A clavate first abdominal segment is known elsewhere in the genus (difficile,

icarioides, obliteratum, lobiferum, for example) but the degree of petiolation is most

highly developed in the two new species.

In one of the two new species, the antennal sockets are separated from the

frontoclypeal suture, a condition typical of the genus Trypoxylon and rare or unknown

in other species of Pison.

Although the two new species share several apomorphies (the two above plus a

transversely elongate anterodorsal pronotal pit, a laterodorsal propodeal carina, and

an emarginate labrum), it is not clear if they are sister species. The wing venation

differs considerably between them (compare figures 345-346), as does the makeup of

the clypeus and form of the eye notch (figs. 341, 343). On the other hand, both

species have a large fovea laterobasally on sternum II, a feature that may be unique

to them, and possibly an apomorphy.

Both species were collected in a malaise trap at the same location in

northwestern New Guinea.

Menke: Pison in the New World 91

Pison pistillum Menke, n. sp.

(Figs. 341-342, 345, 347, 349)

Description, holotype female:

Black except following yellow brown: scape, lower side of pedicel, lower

two-thirds of clypeus, basal half of mandible, palpi, tegula, hindmargin of pronotal

lobe, legs except hindtarsomeres III-V and all arolia which are black. Wings clear,

veins black. Clypeus with dense, appressed silver setae, this vestiture continued

dorsad along inner orbit to eye notch and narrowly beyond to level of midocellus;

gena with dense, appressed silver setae; thoracic vestiture short, silver, sparse on

scutum, scutellum and propodeal dorsum. Gaster largely bare but with sparse,

appressed silver setae on tergum II laterobasally and apicolaterally on tergum III.

Disk of frons weakly shiny, alutaceous, vaguely, shallowly punctate. Eye notches

acute (fig. 341), distance between them 0.84X eye length; eye facets increasingly

larger toward inner orbit, those at level of antennal sockets largest, 2X as large as

smallest facets (fig. 342). Antennal socket separated from frontoclypeal suture by

0.40X midocellus diameter; antennal flagellum elongate, flagellomere I length 4.5xX

apical width, VII length 2.3X apical width. UID 0.96 LID (just below antennal

sockets); OOD 0.18X HOD. Clypeal disk angularly swollen in lateral profile, lower

two-thirds of swelling flat; clypeal lobe present, margin thin, trilobate (fig. 342).

Labrum semicircularly emarginate apically. Mandible with subbasal tooth on cutting

edge (fig. 342), entire posterobasally. Occipital carina incomplete ventrally.

Pronotum with long, deep, transverse depression anterodorsally (length nearly

equal to UID), depression margined posteriorly by sharp carina and anteriorly by

overhanging lamelliform carina; collar non-carinate, rounded laterally. Scutum

weakly shiny, densely micropunctate, punctures about a diameter apart. Tegula

impunctate on outer third. Scutellum shiny, densely micropunctate, the punctures

smaller than those of scutum and 1-4 diameters apart. Propodeal dorsum smooth,

shiny, micropunctate similar to scutellum, with median, longitudinal, linear

impression that contains weak carina at base; propodeal hindface more coarsely,

densely but shallowly punctate than dorsum, with deep, longitudinal depression;

propodeal side delimited dorsad by sharp, arcuate carina that extends from petiole

socket nearly to spiracle, side shiny, impunctate near metapleural sulcus, but

extremely finely micropunctate elsewhere. Propleuron shiny, largely impunctate.

Episternal sulcus disappearing ventrad as it begins to curve anterad. Mesopleuron

shiny, densely micropunctate.

Forewing with three submarginal cells, third conspicuously narrowed toward

marginal cell; media diverging from M+Cu well before crossvein cu-a (fig. 345).

Hindcoxa with inner dorsal carina that ends before apex, outer dorsal carina

absent; tarsomere IV with plantula; tarsal claw tapering gradually to apex.

Gaster shiny, micropunctate, punctures separated by one to several diameters;

segment I clavate (figs. 347, 349), tergum I depressed bandlike along free edge.

Length 10.5 mm.

Male:

Unknown.

Discussion:

Pison pistillum differs from woji in wing venation (fig. 345, media diverging

before cu-a, shape of submarginal cell III); in the more elongate antennal flagellum;

in the projecting, trilobate clypeal lobe (fig. 342) and flattened clypeal disk (not

apparent in SEM photos); in the sharp eye notch; and the color and form of the first

gastral segment (compare figs. 347-350). Pison pistillum is also a larger wasp than

woji.

Distribution:

Known only from a single location in northwestern New Guinea.

92 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

Type:

Holotype female: PAPUA NEW GUINEA, Enga_ Prov.?: Baiyer River,

II-6/25-1979, 1100 m, J. Sedlacek (AEIJ).

Etymology:

The name pistillum, a noun in apposition, is the Latin word for the club shaped

pounder used in a mortar. Here it refers to the claviform first abdominal segment of

this wasp.

Pison woji Menke, n. sp.

(Figs. 343-344, 346, 348, 350)

Description, female:

Black except following yellow brown: scape, pedicel and flagellomeres I-II

beneath: mandible except tip; palpi; tegula; hindmargin of pronotal lobe; legs except

coxae, midtarsomeres II-IV, hindtarsomeres II-V, and all arolia; gastral segment I.

Wings clear, veins dark bronw. Clypeus with dense, appressed silver setae, this

vestiture continued dorsad along inner orbit to eye notch and narrowly beyond to

level of midocellus; gena with dense appressed silver setae; thoracic vestiture short,

silver, sparse on scutum, scutellum and propodeal dorsum. Abdominal vestiture short,

sparse, brownish.

Disk of frons weakly shiny, alutaceous, vaguely, shallowly punctate. Eye notches

rounded (fig. 343), distance between them 0.97X eye length; eye facets increasingly

larger toward inner orbit below notch, those at level of antennal sockets largest,

about 3X as large as smallest facets (fig. 344). Antennal socket essentially

contiguous with frontoclypeal suture; antennal flagellum moderately elongate,

flagellum I length 2.75X apical width, VII length 1.5X apical width. UID essentially =

LID (at level of sockets); OOD 0.22X HOD. Clypeal disk evenly convex; clypeal free

margin broadly arcuate, without median lobe (fig. 344), edge of margin narrowly

folded under at middle, edge otherwise not thickened. Labrum semicircularly

emarginate apically. Mandible with weak tooth at middle of cutting edge (fig. 344),

entire posterobasally. Occipital carina incomplete ventrally.

Pronotum with long, narrow, transverse depression anterodorsally (length about

two-thirds of UID), depression margined posteriorly by sharp carina and anteriorly by

sharp edge; collar non-carinate, rounded laterally. Scutum weakly shiny, densely,

shallowly punctate, punctures less than diameter apart. Tegula impunctate on outer

third. Scutellum weakly shiny, densely punctate, punctures smaller than those of

scutum, about one diameter apart. Propodeal dorsum smooth, shiny, punctate,

punctures finer than those of scutullum, one-half to one diameter apart; dorsum with

median, longitudinal carina that ends about two-thirds distance to apex; propodeal

hindface punctate similar to dorsum and with deep, longitudinal depression; propodeal

side delimited dorsad by sharp, arcuate carina that extends from petiole socket

nearly to spiracle (fig. 348), side shiny, finely micropunctate. Propleuron weakly

shiny, obscurely micropunctate. Episternal sulcus disappearing ventrad as it begins to

curve anterad. Mesopleuron shiny, obscurely micropunctate.

Forewing with three submarginal cells, third not much narrowed toward marginal

cell; media diverging from M+Cu after crossvein cu-a (fig. 346).

Hindcoxa with inner dorsal carina that ends before apex, outer dorsal carina

absent; tarsomere IV with plantula; tarsal claw tapering gradually to apex.

Gaster weakly shiny, micropunctate, punctures separated by 2 to 4 diameters;

segment I clavate (figs. 348, 350), tergum I entirely dorsal from base to spiracles (fig.

348).

Length 6.5-8 mm.

Male:

Unknown.

Menke: Pison in the New World 93

Variation:

Nothing of significance.

Discussion:

Pison woji differs from pistillum in forewing venation (fig. 346, media diverging

after cu-a, shape of submarginal cell III); the less elongate antennal flagellum; the

arcuate clypeal margin (fig. 344) and evenly convex clypeal disk (fig. 343); the

rounded eye notch; and the yellowish brown color and form of the first abdominal

segment (fig. 348). Pison woji is a smaller species than pistillum.

Types:

Holotype female: PAPUA NEW GUINEA, Enga_Prov.?: Baiyer River,

I-25/II-6-1979, L100 m, J. Sedlacek (AEI).

Paratypes (5 females): Same data as holotype except collected XII-26-1978 to

ITI-9-1979 (AEI).

Etymology:

This wasp is affectionately dedicated to Wojciech "Woj" Pulawski, sphecid

expert, masterful sphecid collector, and valued friend.

Key to species of Pisonopsis

2. Scutum and scutellum weakly shiny, uniformly densely punctate; hindwing jugal

lobe small, its length about one-third length of submedial cell: female

clypeus without nasiform process above free margin; female without pygidial

plate: Chile and Argentind: ..v.ceuceinctosgepeeeottiden teen eee areolata (Spinola)

- Disk of scutum and all of scutellum polished, both coarsely, sparsely punctate,

the last almost impunctate; hindwing jugal lobe length about half that of

submedial cell; female clypeus with nasiform process above free margin;

female with pygidial plate margined by carinae; Argentina ...... australis Fritz

3. Female with pygidial plate margined by carinae; male clypeal lobe quadrangular

but with rounded median tooth; interspaces between scutal and scutellar

PUMCTUP ES POM GMC: ciciecaessavedtesecameracnamaiune a ldielomr ius Ube clypeata Ashmead

- Female without pygidial plate; male clypeal lobe triangular; interspaces between

scutal and scutellar punetures-dulito shimyow, ids cca ave connate nier oetlbtes coun. 4

4, Pronotal lobe pale or whitish posteriorly; apex of male sternum VIII emarginate

PESWILIMG GIT OT POMS ba saniciceditewco lou tele ty sak beer oe cae birkmanni Rohwer

- Pronotal lobe all black; male sternum VIII truncate apically

wasain:asala A ain Sosa'wsalolcldieiasag'aey ah PRI Rima tncicld cia Nias Mth ENO ACN os a) aa). en eal triangularis Ashmead

The subspecies of Pisonopsis trianqularis and clypeata (i.e., californica and

occidentalis, respectively) established by Williams (1954) are not recognized by Bohart

(in litt.) who feels that they are simply variants within each species.

94 Contrib. Amer. Ent. Inst., vol. 24, no. 3, 1988

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