Bonn zoological Bulletin 67 (2): 71-77

2018 - Daza J.D. et al.

https://doi.org/10.20363/BZB-2018.67.2.071

ISSN 2190-7307

http://www.zoologicalbulletin.de

Scientific note

urn:|sid:zoobank.org:pub:AA5B7777-C36B-4E9C-B77E-272FBCBA3D7A

Things are not always as they seem:

High-resolution X-ray CT scanning reveals

the first resin-embedded miniature gecko of the genus Ebenavia

Juan D. Daza™', Jordan P. Hunziker', Aaron M. Bauer’, Philipp Wagner”*, Wolfgang Bohme*

' Department of Biological Sciences, Sam Houston State University, 2000 Avenue I, Life Sciences Building, Room 105, Huntsville,

TX 77341, USA

? Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, PA 19085, USA

3 Allwetterzoo Miinster, Sentruper Str. 315, D-48161 Miinster, Germany

* Zoologisches Forschungsmuseum A. Koenig, Adenauerallee 160, D-53113 Bonn, Germany

* Corresponding author: E-mail: juand.daza@gmail.com

'urn:lsid:zoobank.org:author:7F796C4B-BOD5-4669-8D6F-96CA494B62F5

?urn:Isid:zoobank.org:author:A54A 16B8-BEE]1-4996-9170-6CADBS539B5D1

3urn:Isid:zoobank.org:author:0575CE45-A B5C-4493-A 8A D-DD399E18284A

4urn:Isid:zoobank.org:author: FFAC2972-9F52-404B-BA9C-489C7793FF8D

*urn:Isid:zoobank.org:author: 78 BBOEBO-8A48-4A 1 D-8D53-287AD1 EF5FFA

Abstract. We identify a presumed specimen of Sphaerodactylus in amber from the Zoological Research Museum Alexan-

der Koenig as being embedded in copal, rather than amber. Further, the specimen matches the morphology not of a Hispa-

niolan gecko, but of the extant Madagascan species Ebenavia boettgeri, which occurs in a known area of copal deposits.

Key words. Sphaerodactylus, Ebenavia, CT scan, Madagascar, Osteology.

Fossil lizards embedded in amber are frequently spec-

tacular since they preserve, in high definition, the three

dimensionality of ancient organisms. To date, fossil re-

mains of squamates have been found in six amber de-

posits around the world (Daza et al., 2016). Examples

include the oldest reptile in amber (Baabdasaurus xe-

nurus) from the Early Cretaceous of Lebanon (Arnold et

al., 2002); some scales attributable to a squamate from

the Albian of France (Perrichot and Néraudeau, 2005);

14 fossil squamates from the mid-Cretaceous of Myan-

mar, including members with affinities to Iguania, Gek-

kota, Scincoidea, Anguimorpha, and Ophidia (Arnold &

Poinar, 2008; Daza et al., 2016; Fontanarrosa et al., 2018;

Daza et al., in press; Xing et al., 2018); a gekkotan and

numerous lacertids in Baltic amber (Succinilacerta suc-

cinea, BOhme & Weitschat, 1998; Borsuk-Biatynicka et

al., 1999; Yantarogekko balticus, Bauer et al., 2005; see

also Cernatisky & Augé, 2013); and many lizards from

the Miocene deposits of Mexico and Hispaniola classified

in the genus Anolis (A. electrum, Lazell, 1965; Rieppel,

1980; de Queiroz et al., 1998; Polcyn et al., 2002; Castafie-

da et al., 2014; Sherratt et al., 2015) and Sphaerodactylus

(S. dommeli, Bohme, 1984; S. ciguapa, Daza & Bauer,

2012). Copal specimens have received relatively less

attention and were reviewed in Broschinski & Kohring

Received: 04.07.2018

Accepted: 13.09.2018

(1998). The genera Phelsuma, Lygodactylus, and Gecko-

lepis have been preserved in Madagascan copal.

The study of lizards in amber has been facilitated

by the use of X-rays and High-Resolution X-ray com-

puted tomography (HRCT; Polcyn et al., 2002; Daza et

al., 2013; Castafieda et al., 2014; Sherratt et al., 2015;

Daza et al., 2016), allowing the rendering of the skele-

ton without distortion, in addition to providing incredi-

ble integumentary detail. As part of an ongoing research

project, we examined all available gecko specimens in

Miocene amber from Hispaniola preserved in amber us-

ing HRCT. One specimen was revealed to be neither a

Sphaerodactylus nor embedded in Miocene amber. The

Specimen was scanned at the Center for Nanoscale Sys-

tems, Harvard University using a Nikon (Metris) X-Tek

HMXST 225 scanner with a molybdenum target at 70KV,

135 pA, 1000 ms exposure, 3143 projections, 0.1° rota-

tion step, and no filter. The reconstructed voxel size for

the particular specimen was 14.251 um. The original data

set has been archived and is available to the public at Mor-

phosource (https://www.morphosource.org/Detail/ProjectDetail/

Show/project_id/545). Additional specimens for compar-

ison were scanned at UTCT | The University of Texas

High-Resolution CT Facility in a Xradia — Zeiss ma-

chine. The specimens were scanned with a 4X objective,

Corresponding editor: R. Peters

Published: 01.11.2018

72 Juan D. Daza et al.

Fig. 1. Specimen ZFMK 94000, in dorsal view. Scale bar equals 5 mm.

70kV/10W using variable parameters. These specimens

are part of a large data base of skull Micro-CT that in-

cludes nearly all gekkotan genera (Aaron M. Bauer digi-

tal collection). All post- processing of the scan data was

_ @Pemba (Tanzania) Seychelles

( Zanzibar

Coétivy Island

Agaléga Islands

~ Grand Comoro

@ Anjouan

Mohéli® © e Mayotte

Nosy Bé@_

@Cap D'Ambre

cue \Sambava area

Br %°

@

eest Marie

Mozambique i ateet me ae

Channel » Antanar Aa Patou

® Mauritius

Reunion @

@ Ebenavia maintimainty

@ Ebenavia inunguis

@ Ebenavia boetigeri

@ Ebenavia tuelinae

@ Ebenavia safari

@ Ebenavia robusta

0 200 km

(et hati

Fig. 2. Distributional map of Ebenavia species. Colors follow

Hawlitschek et al. (2018). Additional localities for £. mainti-

mainty taken from Nussbaum and Raxworthy (1998).

Bonn zoological Bulletin 67 (2): 71-77

performed using Avizo Lite 9.5.0 (OFEI SAS, Thermo

Fisher Scientific, 2018).

The specimen in question, from the Alexander Koe-

nig Research Museum (ZFMK 94000, Fig. 1), had been

obtained by the museum sponsoring society (Alexan-

der-Koenig-Gesellschaft) in November 2012 under the

assumption that 1t was a piece of Dominican amber with

a Sphaerodactylus (Gekkota: Sphaerodactylidae) inclu-

sion. Although the size and overall appearance is con-

sistent with that of these miniaturized geckos (Daza et

al. 2008), a more thorough analysis of this material and

comparison with the Micro-CT data base of gekkotans

indicated that this specimen is a Madagascan clawless

gecko of the genus Ebenavia in the family Gekkonidae.

Morphological data indicates that the specimen is a sub-

fossil, and that the resin is copal, not amber.

Copal from Madagascar is botanically assigned to

the fabacean species Hymenaea verrucosa (Penney et

al., 2005). Copal can be differentiated from mature res-

ins, such as amber, with Raman spectroscopic analyses,

showing more intense bands at around 1640 cm"! due to

more stretching vibrations of the v(C=C) attributed to the

olefinic group (C=CH2; Winkler et al., 2001). Thermal

analyses have also been used to characterize resins; co-

pal from Madagascar may be differentiated from amber

and other copal resins in reaching a peak in differential

thermogravimetric analysis at 384 °C, while Colombian

copal and amber from other localities peaks at 400 °C or

more (Ragazzi et al., 2003). The age of copal resins may

be only a few hundred to up to four million years old.

Some resins from Madagascar have been dated using car-

bon dating analyses to be as young as just a few decades

(Poinar, 1999; Bosselaers et al., 2010). Other estimates

©ZFMK

Miniature gecko in resin Ge)

postorbitofrontal

epipterygoid scleralossicle AV

ribs scapulocoracoid Squamosal, quadrate frontal

Q Pe = ee a

eS SSS rr Sg INES >» — prefrontal

BOLI, Fue LA LS Ce —>_ maxilla

: Qn oe Se Rie 7 > RR 3

sulin == iy 2 ~-— = premaxilla

|. radius 74 aac dentary

le r.fadius \. r5 64 -coronoid

Aly sac r. humerus r. carpus

4 2 |. humerus

Is postorbitofrontal B

frontal

t14 ectopterygoid

tis jugal prefrontal

r. humerus

r. radius

maxilla

nasals

premaxilla

r. ulna r. Carpus

r.ulna _ otooccipital Cc

r. radius basioccipital

[> . .

J =~ parabasisphenoid

i. pterygoid

ae palatine

Faxes = vomer

ee — dentary

ee maxilla

compound coronoid

bone surangular

quadrate

‘7 postorbitofrontal _

scleral ossicle epipterygoid D

prootic

prefrontal DTS ER squamosal_ ;

nasals P= eS. “ao aed . y

: Ee ee ; ; LOR Aa TS) = et he

premaxilla —?- SRS ee ae, Sc! Ze —— al

a Vi eae

naxilla ee —.

dentary -

coronoid 0 5mm

Fig. 3. Gecko in copal (Ebenavia boettgeri, ZFMK 94000) A. right lateral view, B. dorsal view; C. ventral view; D. left lateral view

indicating the major bones. Abbreviations c#, cervical vertebrae #, l#, left toe #, r#, right toe #, t#, thoracic vertebrae #.

for the age of Madagascan copal include a range from

Holocene to Recent (10,000—100 y; Schltiter & Gnielins-

ki, 1987; Lourengo, 1996; Winkler et al., 2001). In this

study, we confirmed that the specimen is embedded in

copal based on the morphological similarities with mod-

ern species, and some simple tests on the resin: 1) A hot

needle was pushed into the piece, causing rapid melting

at the point of insertion (rather than slow melting expect-

ed in amber); the melting resin released a mild fragrance

Bonn zoological Bulletin 67 (2): 71-77

(amber yields a sooty odor). 2) Under a UV lamp the

piece did not show any color change (rather than emitting

a bluish glow, as does amber).

Morphological comparisons considering members of

nearly all described gekkotan genera indicate that Ebe-

navia shares most morphological characters with ZFMK

94000. The genus occurs on Madagascar and satellite

islands, Pemba Island, Grand Comoro, Mohéli, An-

jouan, Mayote, Nosy Bé, Nosy Komba, Ile Sainte-Ma-

©ZFMK

74 Juan D. Daza et al.

px

Se .

- _

‘ 4 —

omy a : oat i a ‘¢ =~ -

~ yr

ae

— = 4 ; : ee

ene AAAS A$

nil

Gr

Leo

De iT ee

Z

-

*

\ .

« t

5mm

*

x . , — ep

WW & . om “

/ hon, eS Se PF) en

ve EW AA

. ty AIR ' Ny

Yee PEAT ETEAAD IVY

4 f rer

Fig. 4. HRCT of the skulls of three geckos. Sphaerodactylidae: A. Sphaerodactylus semasiops (MCZ R-55766); Gekkonidae:

B. Ebenavia boettgeri (ZFMK 94000), and C. Ebenavia boettgeri (CAS 66195).

tie (Nosy Boraha), Nosy Mangabe, and Ile aux Prunes

(Nosy Alafiafia), as well as Mauritius (Ramanamanyato

et al., 2002; Hawlitschek et al., 2017, 2018; Uetz et al.,

2018). Until recently Ebenavia included only two species

(E. maintimainty and E. inunguis). Ebenavia maintimain-

ty has a restricted range, being found in Toliara Province

in southwestern Madagascar (Nussbaum & Raxworthy,

1998), while the more widespread EF. inunguis was re-

cently split into four new species (viz., E. boettgeri,

Bonn zoological Bulletin 67 (2): 71-77

E. robusta, E. safari, E. tuelinae; Hawlitschek et al.,

2018, Fig. 2). Copal deposits in Madagascar are concen-

trated in the northern part of the island (1.e., Cap D’Am-

bre and the Sava Region, Geirnaert 2002), which is com-

patible with the distribution of E. safari and E. boettgeri

(Hawlitschek et al., 2018).

The specimen is embedded in a cone-shaped piece of

orange resin (Fig. 1). The preservation is exceptional,

conserving the complete anterior half of the body. It ap-

©ZFMK

Miniature gecko in resin ES

Fig. 5. Lateral view of the snout of ZFMK 94000, inset shows the separation between the rostral scale and the nostril.

pears desiccated, but the skeleton is in perfect condition,

including the skull, vertebral column (all six cervical and

14 thoracic vertebrae), ribs, pectoral girdle and forelimbs

(Figs 1, 3). The skull is intact, with the exception of the

anterior portion of the left maxillary bone, which appears

damaged, and both sclerotic rings, which are collapsed.

Even fine details of the skeleton are visible (e.g., small

sesamoids in the elbow; Fig. 3). Although having an in-

tact skeleton is not necessarily an indication of its young

geological age (for example, a Mesozoic gecko in amber

exhibits a near pristine skeleton, Daza et al., 2016), the

skeleton typically exhibits multiple fractures in the ma-

jority of Miocene Sphaerodactylus from Hispaniola. Us-

ing two X-rays from three ethanol preserved specimens

of Ebenavia boettgeri (CAS 66195 [male, based on the

presence of cloacal bones], CAS 66196 [gravid female

with 2 eggs], 16° 54' 37.08" S, 49° 54' 40.716" E, St. Ma-

rie, and USNM 495825 25° O1' 12.0" S, 46° 58' 48.0" E

[gravid female with 1 egg]) we were able to determine

that the missing portion of the precloacal region (SVL)

is between 18 and 25% of the SVL. Using these values,

Bonn zoological Bulletin 67 (2): 71-77

the estimated SVL of the copal specimen is 34.8-37.8

mm. The estimated size matches several species of Ebe-

navia, although it greatly exceeds the adult size range of

E. maintimainty (21-24 mm; Nussbaum & Raxworthy,

1998).

The specimen in copal was compared to similar-

ly sized, formalin-fixed, ethanol preserved specimens

of E. boettgeri from St. Marie (Fig. 4) and E. robusta

(ZSM296/2010; Hawlitschek et al., 2018). Shared traits

with Ebenavia include a small premaxilla with a short

ascending nasal process (long in Sphaerodactylus), fused

nasals (unfused in Sphaerodactylus), frontal broad with

flat dorsal surface (narrow and convex in Sphaerodacty-

Jus); quadrate more or less straight with a slightly convex

conch (curved and extremely convex conch in Sphaero-

dactylus), high number of foramina in the maxillary fa-

cial process (fewer foramina in Sphaerodactylus), high,

discrete splenial (fused to coronoid in Sphaerodactylus);

stapedial foramen absent (present in Sphaerodacty-

/us);dentary ending at the level of the coronoid eminence

(extending beyond the coronoid in Sphaerodactylus); and

©ZFMK

76 Juan D. Daza et al.

retroarticular process narrow (broad in Sphaerodactylus).

Morphology of the manus is very similar between the

two genera, both Ebenavia and Sphaerodactylus having

the same (plesiomorphic) phalangeal formula (2-3-4-5-3)

and similar relative length of toes, from largest to small-

est IN>IV>H>V>l1.

The copal gecko is clearly differentiated from £. main-

timainty. Head length in the E. inunguis group is 9.2—

9.5 mm (~9.5 in the copal gecko and between 5.4—5.9

in E. maintimainty), dorsal scales are partially keeled

in the E. inunguis group and the copal gecko (vs. fully

keeled in E. maintimainty), and the rostral scale is broad

in the E. inunguis group and the copal gecko (vs. narrow

in E. maintimainty). Using the key from Hawlitschek et

al. (2018), we were able to confirm that ZFMK 94000

has the rostral scale separated from the nostril (Fig. 5),

which is a character that defines Ebenavia boettgeri. This

identification 1s compatible with the fact that Madagas-

can copal mines are only found in the distribution range

of E. safari and E. boettgeri.

Even if the gecko in copal is potentially less than sev-

eral hundred years old, the material provides an historical

record of a living species of Ebenavia in the northeast of

Madagascar based on the known location of copal de-

posits (Geirnaert, 2002). The North of Madagascar is a

critical area for understanding the current distribution of

Ebenavia in Madagascar, as ancestral area reconstruc-

tions may indicate a colonization of northern Madagascar

from the Comoros Islands (Hawlitschek et al., 2017).

Acknowledgements. We thank Morris Flecks for taking the

photograph of the specimen in copal. We also thank Emma

Sherratt (The University of Adelaide), Jessie Maisano and

Matt Colbert (UT|CT, The University of Texas at Austin) for

assistance with the HRCT. We thank Rebecca Laver and two

anonymous reviewers for comments on the manuscript. The

Department of Biological Sciences sponsored Jordan P. Hun-

ziker to present partial results of this project at The Joint Meet-

ing of Ichthyologists and Herpetologists (JMIH) in Austin in

2017. Juan D. Daza received funding from NSF DEB 1657648

and the Biological Sciences Program at Sam Houston State

University. Aaron M. Bauer received funding from NSF DEB

1555968 and 1556585, and the Gerald M. Lemole Endowed

Chair Funds. PW is grateful to Jurgen Velten for an ongoing

cooperation in amber research.

REFERENCES

Arnold EN, Azar D, Ineich I, Nel A (2002) The oldest reptile in

amber: a 120-million-year-old lizard from Lebanon. Journal

of Zoology 258: 7-10

Arnold EN, Poinar G. (2008) A 100 million year old gecko with

sophisticated adhesive toe pads, preserved in amber from

Myanmar. Zootaxa 1847: 62-68

Bauer AM, Bohme W, Weitschat W (2005) An early Eocene

gecko from Baltic amber and its implications for the evo-

lution of gecko adhesion. Journal of Zoology 265: 327-332

Bohme W (1984) Erstfund eines fossilien Kugelfingergeckos

(Sauria: Gekkonidae: Sphaerodactylinae) aus Dominikani-

Bonn zoological Bulletin 67 (2): 71-77

schem Bernstein (Oligozén von Hispaniola, Antillen). Sala-

mandra 20: 212—220

Bohme W, Weitschat W (1998) Redescription of the Eocene

lacertid lizard Nucras succinea Boulenger, 1917 from Baltic

amber and its allocation to Succinilacerta n. gen. Mitteilun-

gen aus dem Geologisch-Palaontologischen Institut der Uni-

versitat Hamburg 81: 203—222

Borsuk-Biatynicka M, Lubka M, Bohme W (1999) A lizard

from Baltic amber (Eocene) and the ancestry of the crown

group lacertids. Acta Paleontologica Polonica 44: 349-382

Bosselaers J, Dierick M, Cnudde V, Masschaele B, Van Hoore-

beke L, Jacobs P (2010) High-resolution X-ray computed to-

mography of an extant new Donuea (Araneae: Liocranidae)

species in Madagascan copal. Zootaxa 2427: 25-35

Castafieda MdR, Sherratt E, Losos JB (2014) The Mexican

amber anole, Anolis electrum, within a phylogenetic context:

implications for the origins of Caribbean anoles. Zoological

Journal of the Linnean Society 172: 133-144

Cernaiisky A, Augé ML (2013) New species of the genus Ple-

siolacerta (Squamata: Lacertidae) from the Upper Oligocene

(MP28) of Southern Germany and a revision of the type spe-

cies Plesiolacerta lydekkeri. Palaeontologia 56: 79-94

Daza JD, Abdala V, Thomas R, Bauer AM (2008) Skull anat-

omy of the miniaturized gecko Sphaerodactylus roosevelti

(Squamata: Gekkota). Journal of Morphology 269: 1340-

1364

Daza JD, Bauer AM (2012) A new amber-embedded sphaero-

dactyl gecko from Hispaniola, with comments on the mor-

phological synapomorphies of the Sphaerodactylidae. Brevi-

ora 529: 1-28

Daza JD, Bauer AM, Stanley EL, Bolet A, Dickson B, Losos

JB (In press) An enigmatic miniaturized and attenuate whole

lizard from the Mid-Cretaceous amber of Myanmar. Breviora

Daza JD, Bauer AM, Wagner P, Bohme W (2013) A reconsider-

ation of Sphaerodactylus dommeli Bohme, 1984 (Squamata:

Gekkota: Sphaerodactylidae), a Miocene lizard in amber.

Journal of Zoological Systematics and Evolutionary Re-

search 51: 55—63

Daza JD, Stanley EL, Wagner P, Bauer AM, Grimaldi DA

(2016) Mid-Cretaceous amber fossils illuminate the past di-

versity of tropical lizards. Science Advances 2: e1501080

de Queiroz K, Chu L-R, Losos JB (1998) A second Anolis liz-

ard in Dominican amber and the systematics and ecological

morphology of Dominican amber anoles. American Museum

Novitates 3249: 1-23

Fontanarrosa G, Daza JD, Abdala V (2018) Cretaceous fossil

gecko hand reveals a strikingly modern scansorial morpholo-

gy: Qualitative and biometric analysis of an amber-preserved

lizard hand. Cretaceous Research 84: 120-133

Geirnaert E (2002) L’ambre — miel de fortune et mémoire de

vie. Les Editions du Piat, Saint-Julien-du-Pinet

Glaw F, Vences M (2006) Field guide to the amphibians and

reptiles of Madagascar. 3" edition. Vences & Glaw Verlag

GbR, Munich

Hawlitschek O, Toussaint EFA, Gehring PS, Ratsoavina FM,

Cole N, Crottini A, Nopper J, Lam AW, Vences M, Glaw F

(2017) Gecko phylogeography in the Western Indian Ocean

region: the oldest clade of Ebenavia inunguis lives on the

youngest island. Journal of Biogeography 44: 409-420

Hawlitschek O, Scherz MD, Ruthensteiner B, Crottini A, Glaw

F. 2018. Computational molecular species delimitation and

taxonomic revision of the gecko genus Ebenavia Boettger,

1878. The Science of Nature 105: 1-21

Lazell Jr J (1965) An Anolis (Sauria, Iguanidae) in amber. Jour-

nal of Paleontology 39: 379-382

©ZFMK

Miniature gecko in resin 77

Loureng¢o WR (1996) Premier cas connu d’un sub-fossile de

scorpion dans le copal de Madagascar. Comptes Rendus de

l’ Académie des Sciences, Paris Series Ia 323: 889-891

Nussbaum RA, Raxworthy CJ (1998) Revision of the genus

Ebenavia Boettger (Reptilia: Squamata: Gekkonidae). Her-

petologica 54: 18-34

Penney D., Ono H, Selden PA (2005) A new synonymy for

the Madagascan copal spider fauna (Araneae, Selenopidae).

Journal of Afrotropical Zoology 2: 41-44

Perrichot V, Néraudeau D (2005) Reptile skin remains in the

Cretaceous amber of France. Comptes Rendus Palevol 4:

47-51

Poinar GOJ (1999) Cenozoic fauna and flora in amber. Estudios

del Museo Ciencias Naturales de Alava 14: 151-154

Polcyn M, Rogers IJ, Kobayashi Y, Jacobs L (2002) Computed

tomography of an Anolis lizard in Dominican amber: syste-

matic, taphonomic, biogeographic, and evolutionary implica-

tions. Palaeontologia Electronica 5: 1-13

Ragazzi, E. Roghi G., Giaretta A., Gianolla P (2003) Classifi-

cation of amber based on thermal analysis. Thermochimica

Acta 404: 43-54

Ramanamanjato J-B, Mcintyre PB, Nussbaum RA (2002) Rep-

tile, amphibian, and lemur diversity of the Malahelo Forest, a

Bonn zoological Bulletin 67 (2): 71-77

biogeographical transition zone in southeastern Madagascar.

Biodiversity and Conservation 11: 1791-1807

Rieppel O (1980) Green anole in Dominican amber. Nature

286: 486-487

Schltiter T, Gnielinski Fv (1987) The East African copal. Its

geological, stratigraphic, palaeontologic significance and

comparison with fossil resins of similar age. National Muse-

um of Tanzania Occasional Paper, Dar es Salaam, Tanzania

8: 1-32

Sherratt E, Castafieda MdR, Garwood RJ, Mahler DL, Sanger

TJ, Herrel A, Losos JB (2015) Amber fossils demonstrate

deep-time stability of Caribbean lizard communities. Pro-

ceedings of the National Academy of Sciences of the United

States 112: 9961-9966

Uetz P, Freed P, Database JHTR (2018) The reptile database.

Online at http://www.reptile-database.org/ (last access: July

2, 2018)

Winkler W, Kirchner ECh, Asenbaum A, Musso M (2001) A

Raman spectroscopic approach to the maturation process of

fossil resins. Journal of Raman Spectroscopy 32: 59-63

Xing LD, Caldwell MW, Chen R, Nydam RL, Palci A, Simdées

TR, McKellar RC, Lee MSY, Liu Y, Shi HL, Wang K, Bai

M (2018) A Mid-Cretaceous embryonic-to-neonate snake in

amber from myanmar. Science Advances 4: eaat5042

©ZFMK

BHL

i

Blank Page Digitally Inserted

Bonn zoological Bulletin 67 (2): 79-99

2018 - Bartsch I.

https://doi.org/10.20363/BZB-2018.67.2.079

ISSN 2190-7307

http://www.zoologicalbulletin.de

Research article

urn:|sid:zoobank.org:pub: 142EQODF-4EFF-4D7 1-84EC-C18D985FDC3B

Freshwater halacarid mites (Acari: Halacaridae) from Madagascar —

new records, keys and notes on distribution and biology

Ilse Bartsch

Forschungsinstitut Senckenberg, DZMB, c/o DESY, Notkestr. 85, D-22607 Hamburg, Germany

E-mail: ibartsch@senckenberg.de

urn:lsid:zoobank.org:author:3DA 19B06-A7CC-4353-BF9F-EE1CC44FB145

Abstract. Four freshwater halacarid species were previously recorded from Madagascar. They had been extracted from

sandy deposits at the banks of streams and creeks. Recently another two species were found, namely Porohalacarus alpi-

nus and Soldanellonyx monardi. On the basis of individuals of these two species, as well as of additional material of the

four formerly recorded species (Limnohalacarus cultellatus, Limnohalacarus novus, Lobohalacarus weberi, and Ropo-

halacarus pallidus), morphological characters are added to previous descriptions and the male of P. alpinus is described.

A key is given to adults of Madagascar freshwater halacarid genera. Morphological differences between the presently

known African freshwater halacarids are outlined. Geographical and biological data of the six species from Madagascar

are presented. All genera and most of the species have world-wide distributions.

Key words. Halacaroidea, riverine sediments, taxonomy, biology, geography.

INTRODUCTION

The database ‘Freshwater Animal Diversity Assessment’

(FADA) presents a list of 67 freshwater halacarid species

and subspecies in 17 genera (Bartsch 2013d). Some of

the species are related to marine genera but the majority

belong to genera which are restricted to fresh or slight-

ly saline brackish water. Eight truly freshwater halacarid

Species are recorded from continental Africa, 1.e., Limno-

halacarus africanus Walter, 1935; L. fontinalis Walter &

Bader, 1952; L. major Bader, 1968; L. marlieri (Bader,

1968); L. portmanni Bader, 1967; Lobohalacarus weberi

(Romiyn & Viets, 1924); Porohalacarus alpinus (Thor,

1910); Ropohalacarus uniscutatus (Bartsch, 1982); and

Soldanellonyx monardi Walter, 1919 (Walter 1935; Wal-

ter & Bader 1952; Bader 1967, 1968; Green 1984; Green

et al. 1974; Bartsch 2008a, 201 3a, b).

Collections of the freshwater mite fauna in sandy de-

posits at the banks of Madagascar streams and creeks,

carried out by R. Gerecke and T. Goldschmidt, also in-

cluded halacarid mites. Recently, records of four species

were published (Bartsch 2013b). Sorting of additional

samples brought to light another two species, namely

Porohalacarus alpinus and Soldanellonyx monardi, and

also several more individuals of the formerly mentioned

Limnohalacarus cultellatus Viets, 1940; Limnohalacarus

novus Bartsch, 2013; Lobohalacarus weberi;, and Ropo-

halacarus pallidus Bartsch, 2013. Morphological charac-

Received: 17.07.2018

Accepted: 10.10.2018

ters of the six species from Madagascar are outlined, the

male of Porohalacarus alpinus is described, biological

details are added and discussed. The geographical distri-

bution of the species is summarized in maps.

MATERIAL AND METHODS

The halacarid mites were extracted from banks of streams

and creeks by digging pits into the sandy deposits and

filtering the seeped water (Karaman-Chappuis Method).

Collectors are Drs R. Gerecke and T. Goldschmidt. The

MD numbers refer to those in the collectors’ collection

diary (unpublished). The halacarid mites were partly

studied in a drop of glycerine, partly cleared in lactic

acid, rinsed in glycerine and mounted in glycerine jelly.

Voucher specimens are deposited in the Zoological Mu-

seum Hamburg (ZMH), Centrum fiir Naturkunde (CE-

NAK), University of Hamburg; additional material in the

author’s collection.

The presentation of the species found on Madagascar

starts with a bibliographical list which includes the first

description of the species, papers which added morpho-

logical details, recent records or keys covering large-

scale geographical areas (British Islands, Central Europe,

North America) and synonyms. The geographical regions

correspond to those outlined in Balian et al. (2008) and

Bartsch (2009). These are the Afrotropical, Palaearctic,

Corresponding editor: B. Huber

Published: 01.11.2018

80 Ilse Bartsch

Oriental, Nearctic, Neotropical, and Australian Regions,

the Pacific Islands and Antarctica. The latter region in-

cludes islands of the southern Atlantic (South Georgia)

and southern Indian Ocean (Prince Edward Islands,

Crozet Island, St Paul, and Kerguelen).

The illustrations and morphological and biological data

are of individuals from Madagascar. Rarely occurring

numbers of setae are in parentheses. A figure in square

brackets indicates the number of cases involved. Decimal

indices are added to notify the position of a structure with

reference to anterior — posterior end of the idiosoma.

Abbreviations used in the descriptions, keys, and dis-

cussion: AD, anterior dorsal plate; AE, anterior epimeral

plate; AP, anal plate; ds-1 to ds-6, first to sixth pair of

dorsal idiosomatic setae, numbered from anterior to pos-

terior; GA, genitoanal plate; gac, genital acetabula; glp,

gland pore(s), numbered glp-1 to glp-5 from anterior to

posterior; GO, genital opening; GP, genital plate; L:H,

ratio length to height; LxW, data of length and width;

L:W, ratio length to width; mxs, maxillary setae, mxs-1,

mxs-2, basal and apical pair of maxillary setae, respec-

tively; OC, ocular plate(s); P-1 to P-4, first to fourth pal-

pal segment; pas, parambulacral seta(e); PD, posterior

dorsal plate; PE, posterior epimeral plate(s); pgs, peri-

genital setae; sgs, subgenital setae. The legs, their seg-

ments and claws are numbered from I to IV from anterior

to posterior. The leg segments are trochanter, basifemur,

telofemur, genu, tibia, and tarsus. The number of setae

on the tarsi includes the solenidion but excludes the

parambulacral setae. In the illustrations, marginal setae

are shown either in dorsal or in ventral aspect but not in

both. Unless indicated otherwise, the given length of a

segment is that along its dorsal margin.

SYSTEMATICS

Annotated key to adult Afrotropical freshwater

halacarid genera and Madagascar species

la. Genu and telofemur of leg I almost equal in length.

Tarsus I with ventromedial spur and pair of short

WOMLLAUS CLAM xara. ath ar satss les, Lobohalacarus

Remarks. In the Afrotropical region, the genus Lobo-

halacarus is represented by a single species, L. we-

beri. The majority of the individuals have a frontal

spine whereas the anterior margin of the other fresh-

water species is arched or truncate.

1b. Genu of leg I shorter than telofemur I. Tarsus I with

Oe leVienbraleSClae. 0.8 t lesbo eee ore ge 2

2a. Gland pores inconspicuous or absent. .................... 3

2b. Dorsum with four to five pairs of distinct gland

pores (glp), pair of glp-1 on AD, glp-2 marginally

in striated integument, glp-4 and glp-5 in striated

intesumentlateral TOP, wisi ede ell ieee gaan 4

Bonn zoological Bulletin 67 (2): 79-99

3a. With four dorsal plates (AD, pair of OC and PD) and

four ventral plates (AE, pair of PE and GP). OC with

comea and large spot of black eye pigment (> 10 um

[fs VAINIG (Gs) fry er 8, ae he ea ee Porohalacrus

Remarks. The only Porohalacarus species known

from Madagascar and Africa, P. alpinus, is charac-

terized by its spots of dark eye pigment, a small one

on the AD and a larger one on each OC. The other

Madagascan freshwater halacarids have either faint-

ly brown or no eye pigment.

3b. With dorsal shield (AD and PD fused, OC reduced or

fused with PE) and ventral shield (AE, PE and GP

fused). Neither cornea nor eye pigment present. ......

Bhi Nid tc ld ai sania Biches 2 haiti De cd Ropohalacarus

Remarks. A single species is known from Mada-

gascar, namely Ropohalacarus pallidus. A record

of another species, R. wniscutatus, is from northern

Africa (Bartsch 2013). Distinguishing characters are

outlined below.

4a. Genital acetabula in posterior part of GP, none situated

anterior to the level of GO. Length of anal sclerites

about half that of female genital sclerites. ...............

PNR aM El 2x A PEE OR SO te Soldanellonyx

Remarks. At present, Soldanellonyx monardi is the

only Soldanellonyx species with records from the

Afrotropical region. Worldwide nine species have

been described.

Ab. Genital acetabula arranged in line along lateral

margin of GP (or area representing this plate), most

anterior acetabula situated well anterior to GO. Anal

sclerites less than 1/3 of the size of genital sclerites.

5a. OC not including platelet with gland pore. Ventral

plates AE, PE and GA separated by striated

integument. Claws I apically with few, delicate tines,

basally with lamellar process with about four tines.

pS Lae, Oa te Monee ee Se Limnohaltacarus cultellatus

5b. OC including platelet with gland pore. All ventral

plates fused. Claws I with J-shaped arranged pectines

bearing at least 1S Lome MNCS assis ccc ssjercennn. 225eesns

Ut pd td eee. ire ALR arte Limnohalacarus novus

Remarks. These two Limnohalacarus species are

recorded from Madagascar but further species are

mentioned from Africa. All African species are out-

lined below, after the presentation of L. novus.

Madagascan and related halacarid species,

notes on morphology, taxonomy, biology, and distri-

bution

Limnohalacarus cultellatus Viets, 1940

L. cultellatus Viets, 1940: 194—200, figs 5-12.

L. cultellatus, Bartsch 2011: 491-493, fig. 2A—C, 2013a:

204—206, figs 1b—h, 2a—h; Pepato & Dos Santos Costa

2015: 5-7, fig. 4A—I.

©ZFMK

Freshwater halacarid mites 81

Fig. 1A—E. Limnohalacarus cultellatus Viets, 1940, female. A. idiosoma, dorsal; B. idiosoma, ventral; C. gnathosoma, ventral;

D. leg I, medial; E. tarsus I, medial (lateral setae in broken line, lateral claw omitted). (Scale = 50 um) (AD, anterior dorsal plate;

AE, anterior epimeral plate; ascl, anal sclerite; GA, genitoanal plate; gac, genital acetabula; glp-1 to glp-5, first to fifth gland pore;

gscl, genital sclerite; OC, ocular plate; PD, posterior dorsal plate; PE, posterior epimeral plate; ph, pharyngeal plate; ti, tines; I-5,

tibia of leg I).

L. kakinadus Chatterjee & Chang, 2005: 23-27, figs 1A—

H, 2A-D.

Collecting data. South-eastern Madagascar, Fianarant-

soa, Ionilahy, stream draining area Marosaro (S from

River Ionilahy), 220 m, 21°C, 0.072 mS/cm, 12 Aug.

2001, interstitial (MD 023). — South-eastern Madagas-

car, Fianarantsoa, Ionilahy, River Ionilahy, 200 m, 23°C,

0.059(11. Aug.)-0.088(13. Aug.) mS/cm, 11/13 Aug.

2001, interstitial (MD 026). — Northern Madagascar,

Antsiranana, Antalaha, Marofinaritra, River Andranom-

enaheli, upstream confluence with River Ankavia (right

affluent below MD 135), 70 m, 22.3°C, 0.009 mS/cm,

riffle, 04 Nov. 2001 (MD136a).

Short description (Fig. 1A—E). Female: Length of

idiosoma 255-290 um [3]. OC and platelet with gland

Bonn zoological Bulletin 67 (2): 79-99

pore separated by striated integument (Fig. 1A), OC

about 1.1 times longer than wide. Corneae and spots with

eye pigment lacking. AE, PE and GA separated by stri-

ated integument (Fig. 1B). Pair of ds-6 present on anal

cone (obscured by genital sclerites). GP with four to five

pairs of gac, three to four pairs of pgs, genital sclerites

with two pairs of sgs. Gnathosoma 1.5—1.7 times longer

than wide; pharyngeal plate distinctly removed from ba-

sal margin of gnathosoma. Both pairs of maxillary setae

slender (Fig. 1C). Legs shorter than idiosoma, length ra-

tio leg I:idiosoma 0.7:1. Length of telofemur I 1.8 times

the height (Fig. 1D). Basifemora I to IV with 4, 3, 2, 1

setae, tibiae I to [V with 7, 6, 7, 6 setae. Tibiae I and II

each with one bipectinate seta, tibiae III with two bipect-

inate ventromedial setae, tibia IV with slender, smooth

ventral setae. Claws on leg I slender, apically with few

©ZFMK

82 Ilse Bartsch

e Limnohalacarus cultellatus

* Limnohalacarus novus

Fig. 2. Records of Limnohalacarus cultellatus Viets, 1940 (black spot) and Limnohalacarus novus Bartsch, 2013 (asterisk).

(8—10) minute tines (Fig. 1E), basal lamellar process with

four tines.

Male: Not present.

Juveniles: Length of deutonymph, protonymph and

larva 240 um [1], 200 um [1] and 155 um [1), respective-

ly. Dorsal aspect similar to that of females. AE of larva

with pair of epimeral pores. In nymphs GP and anal plate

separated by striated integument, in larvae GP lacking.

Remarks. Limnohalacarus cultellatus can be distin-

guished from the other African Limnohalacarus species

by combination of: dorsal and ventral plates separated

and pectines on claws I much more delicate than on

claws II to [V. Notes on African Limnohalacarus species

are given below.

Biology. At present there is no record of a male, neither

from Africa nor from other parts of the world.

One of the females included an ovoid excretory body,

125 um long, 60 um wide, the centre (10 um in diameter)

is dark, the margin hyaline and delicately stratified. One

of the females held an egg, 50 um long, 55 um wide.

Bonn zoological Bulletin 67 (2): 79-99

Geographical distribution (Fig. 2) (cf. Bartsch 2011,

2013a; Pepato & Dos Santos Costa 2015; Ojeda et al.

2016):

Afrotropical Region. — Madagascar;

Palaearctic Region. Europe: — Hungary;

Oriental Region. — India (Andhra Pradesh);

Nearctic Region. — United States (Georgia, Wisconsin ?);

Neotropical Region. — Brazil (Mato Grosso, Rio Grande

do Sul) — Cayman Islands (Grand Cayman) — El Salva-

dor — Mexico (Quintana Roo) — The Netherlands Antil-

les (Bonaire, Curacao) — Venezuela (Margarita Island).

Limnohalacarus novus Bartsch, 2013

L. novus Bartsch, 2013a: 206—210, figs 3a—h, 4a—g.

L. billabongis, Bartsch 2008b: 127, 128.

Collecting data. Central Madagascar, Antananarivo, An-

jazorobe, River Ranonisoanavola (larger stream E from

main mountain chain), 1200 m asl, 13.2°C, 0.058 mS/cm,

interstitial, 23 Jul. 2001 (MD 012). — South-eastern Mad-

agascar, Fianarantsoa, Ionilahy, stream draining area Ma-

rosaro (S from River Ionilahy), 220 m, 21°C, 0.072 mS/

cm, interstitial, 12 Aug. 2001 (MD 023). — South-east-

ern Madagascar, Fianarantsoa, Ionilahy, River Ionilahy,

©ZFMK

Freshwater halacarid mites 83

Fig. 3A—G. Limnohalacarus novus Bartsch, 2013, female. A. idiosoma, dorsal; B. idiosoma, ventral; C. excretory body, ventral; D.

genital area with ovipositor; E. gnathosoma, ventral; F. leg I, medial; G. claw I, ventral. (A—F, scale = 50 um; G, scale = 10 um)

(dgr, dark granules; hy, hyaline part; mxs-1 and mxs-2, basal and apical maxillary seta, respectively; ov, ovipositor; ph, pharyngeal

plate; P-3, third palpal segment; sgs, subgenital seta).

200 m, 23°C, 0.059(11. Aug.)-0.088(13. Aug.) mS/cm,

interstitial, 11/13 Aug. 2001 (MD 026). — South-east-

ern Madagascar, Fianarantsoa, Ionilahy, small stream

crossing the railroad east from village, 200 m, 19.9°C,

0.083 mS/cm, 15 Aug. 2001 (MD027). — Southern Mad-

agascar, Tulear, Tsimelahy, River Antarantsa, ca 1 km

upstream from village, 300 m, 20.4°C, 0.171 mS/cm,

interstitial, 04 Sep. 2001 (MD 058). — Central Mada-

gascar, Antanarivo, Ankaratra, Reserve Manjakatompo,

left affluent of River Mahiavona, EM Mantsina, 1750 m,

14.1°C, 0.003 mS/cm, interstitial, 08 Oct. 2001 (MD107).

— North-western Madagascar, Majunga, Adjamangirana,

stream in dry forest, upstream, rice field area (road to the

village of Tsaratanana), 220 m, 30.8°C, 0.008 mS/cm,

interstitial, 19 Oct. 2001 (MD115). — Northern Mada-

gascar, Antsiranana, Antalaha, Marofinaritra, River Ant-

sohibe, upstream confluence with River Ankavia (5 km

NE Amparihimena), 70 m, 25.2°C, 0.008 mS/cm, riffle,

03 Nov. 2001 (MD135). — Northern Madagascar, Antsir-

anana, Antalaha, Marofinaritra, River Andranomenaheli,

upstream confluence with River Ankavia (right affluent

below MD 135), 70 m, 22.3°C, 0.009 mS/cm, riffle, 04

Nov. 2001 (MD136a). — Northern Madagascar, Antsir-

Bonn zoological Bulletin 67 (2): 79-99

anana, Maroambihy (Sambava), left affluent of River Lo-

koho upstream from the village, 90 m, 26.0°C, 0.010 mS/

cm, interstitial, 12 Nov. 2001 (MD149). — Northern Mad-

agascar, Antsiranana, Joffreville (M. d’Ambre), River

Antomboka, downstream large cascade, 850 m, 20 Nov.

2001 (MD163). — Northern Madagascar, Antsiranana,

Joffreville (Montagne d’Ambre), River Manques in Re-

serve Fontenay, 580 m, interstitial, 21 Nov. 2001 (MD

165).

Short description (Fig. 3A—G). Female: Length 206—

289 um [20]. OC including platelet with glp-3 (Fig. 3A),

L:W of OC 1.4—1.6:1. Length ratio of PD:AD 2.6—2.7:1.

Pair of ds-2 absent. Ventral plates AE, PE and GP fused

(Fig. 3B). Area corresponding to GP with three pairs of

pgs, five to six pairs of gac and on each genital sclerite

two (three) sgs. Ovipositor short and narrow (LxW 74 x

10 um), very faintly sclerotized, genital spines very del-

icate and short, 2 um in length, and arising from minute

papillae (Fig. 3D). Anal slit less than 10 um long and

guarded by pair of short anal sclerites. Gnathosoma about

1.1 times longer than wide. Pair of mxs-2 shorter but

wider than mxs-1 (Fig. 3E). Legs shorter than idiosoma,

©ZFMK

84 Ilse Bartsch

length ratio leg I:idiosoma 0.7:1. Length of telofemur I

2.1—2.3 times the height (Fig. 3F). Tibiae I to IV with 8,

7, 7, 6 setae [14]. Ventromedial seta on tibia II and both

ventromedial setae of tibia III bipectinate; setae on legs

I and IV smooth. Claws I to IV with J-shaped arranged

pectines (Fig. 3G), extending from apical lateral flank

and along medial flank to basis of claw. On claws II to [V

basal tines partly fused to a lamellar process.

Male: Not present.

Juveniles: Length of deuto-, protonymphs and larvae

231-275 um [13], 180-226 um [7] and 142-170 [5]

um, respectively. In contrast to adults, OC and posteri-

or gland-bearing platelet separated by transverse striae.

Ventral plates AE, PE and GP of nymphs separated from

each other by striated integument, in larvae GP absent

and AE with pair of epimeral pores.

Biology. Eleven of 15 females studied held an egg. The

length x width of the eggs ranged from 35 x 35 to 100 x

50 um. The eggs are not deposited in the substratum but

attached to tibiae IV. One-third of the females had a sin-

gle cocoon fixed with a slender stem, the latter 2-3 um

wide. The length of the cocoons was about 103-113 um,

the diameter 60-64 um. Each one enclosed a single em-

bryo.

Limnohalacarus species have the excretory material

accumulated within an ovoid, compressed body (cf. Wal-

ter 1914; Petrova 1966; Ramazotti & Nocentini 1960;

Pepato & Dos Santos 2015). In the material studied,

11 (73 %) of 15 females, 9 (22%) of 27 deutonymphs

and one (22%) of six protonymphs held such a body,

but none of four larvae. The body was dorsal to the gut,

in some individuals it had a dark centre, surrounded by

concentrically arranged more or less transparent layers

(Fig. 3C). In one female the excretory body was almost

uniformly dark, in another one hyaline, the body recog-

nizable mainly because refraction of light. In one female

(length 255 um) this body was 108 um long, 71 um wide

and 41 um thick, the general size range in length was

95-158 um, in width 50-92 um. The length of the ex-

cretory body equalled 0.4—0.5 times the length of the fe-

male idiosoma. In deutonymphs the stratified body was

60-85 um long, 45-95 wide, in a protonymph 50 um

long, 36 um wide, 1.e., a length of 0.3—0.4 and 0.2 of that

of the nymphal idiosoma, respectively. Such a body was

present/absent independent of the age of the instar, e.g. it

was absent both in assumedly recently hatched females

and in ovigerous ones and also in those carrying a cocoon

(or their remnants) fixed to the tibiae.

Remarks. Adults of Limnohalacarus cultellatus and

L. novus, the two species known from Madagascar, can

easily be separated by (1) the shape of the ocular plates,

namely OC hardly longer than wide (L:W 1.1:1) and not

including the platelet with a gland pore versus OC 1.3

times longer than wide and including the platelet with

Bonn zoological Bulletin 67 (2): 79-99

gland pore, (2) the ventral plates, separated versus fused

to a shield, (3) the length of the gnathosoma, at least 1.5

times longer than wide versus 1.1 times or less, and (4)

the claws on tarsus I, with few delicate versus numer-

ous long tines. Juveniles of both species have the ventral

plates and the OC and gland-pore bearing platelet sepa-

rated but can be distinguished by the shapes of gnatho-

soma and claw I.

Compared with African species, L. novus turns out to

be most similar to LZ. portmanni. Both are of about the

same size, their length 206-289 and 217-289 um, re-

spectively, but the PD of Z. novus is not as slender as in

L. portmanni and the length ratio PD:AD is 2.6—2.7:1 in

L. novus but 3.1—3.3:1 in L. portmanni (Bartsch 2013a).

Geographical distribution (Fig. 2) (cf. Bartsch

2013a):

Afrotropical Region. — Madagascar;

Australian Region. — Australia (Queensland — Moreton

Bay, North Stradbroke Island, not Moreton Island).

Limnohalacarus species from the Afrotropical Region

The Limnohalacarus species recorded from the Afrotrop-

ical region are: L. africanus, L. cultellatus, L. fontinalis,

L. major, L. marlieri, L. novus, and L. portmanni. Rely-

ing on often vague descriptions, the most specific mor-

phological characters of the six African species are:

Limnohalacarus africanus. Characters (according to

Walter 1935, Green 1984 and Bartsch 2013a): Length

330 um, OC and gland pore-bearing sclerite separated,

ventral plates separated, GA of female with 8—9 (up to 12

according to Green 1984) pairs of gac and three pairs of

pgs, its genital sclerites with two pairs of sgs; L:W of fe-

male gnathosoma 1.6:1 (Walter 1935: p. 74), but distinct-

ly less in the deutonymph (Walter 1935: fig. 4), none of

setae on tibiae I bipectinate, pectines on claws I J-shaped

and bearing numerous long tines. Distribution: Burkina

Faso (Upper Volta) (well at Banfora) (Walter 1935), West

Cameroon (Debundska Lake) (Green et al. 1974), South

Sudan (Lake No) (Green 1984).

Limnohalacarus fontinalis. Characters (according

to Walter & Bader 1952 and Bartsch 2013a): Length

310-370 um, gland pore included in elongate OC; ven-

tral plates fused, L:W of gnathosoma 1.1:1; tibiae I to IV

with 8, 7, 7, 6 setae, respectively, claws I with numerous

long tines. Distribution: Kenya (Mombasa, from well

with slightly saline water) (Walter & Bader 1952), South

Sudan (Lake No) (Green 1984).

Limnohalacarus major. Characters (according to Ba-

der 1968): Length 465 um, distinctly larger than the other

species (their length 224-370 um), gland pore included

in elongate OC; ventral plates fused, gnathosoma slen-

der, its L:W 1.6:1. Distribution: DR Congo (Zaire) (Lake

Tanganyika) (Bader 1968).

©ZFMK

Freshwater halacarid mites 85

Limnohalacarus marlieri. Characters (according to

Bader 1968 and André & N’Dri 2012): Length of fe-

male 305 um, of male 350 um. OC elongate, 1.4 times

longer than wide. Length ratio PD:AD 3.4:1. Ventral

plates fused. Female with eight pairs of gac posterior

and posterolateral to GO. Number of gac along margins

of area representing GA not known. Gnathosoma short,

L:W 1.1-1.2:1. Limnohalacarus marlieri is very similar

to both L. fontinalis and L. portmanni, it 1s classified as

a Junior synonym and omitted in the following key. Dis-

tribution: DR Congo (Zaire) (Lake Tanganyika) (Bader

1968).

Limnohalacarus portmanni. Characters (according

to Bader 1967 and Bartsch 2013a): Length of females

224-250 um, of males 217—289 um, gland pore included

in elongate OC; length ratio PD:AD 3.1—3.3:1; ventral

plates fused, L:W of gnathosoma 1.2:1. Female with up

to seven pairs of gac and three pairs of pgs, each genital

sclerite with (two to) four sgs. All claws with J-shaped

arranged pectines. Distribution: DR Congo (Zaire) (Lake

Tanganyika, from shallow water sediment) (Bader 1968).

Characters of Limnohalacarus cultullatus and L. novus

are outlined above.

Key to adult Afrotropical Limnohalacarus species

ta. All ventral. plates/separated bo. s:2 ei. alas ieee. 2

1b. All ventral plates fused to a single shield. ............... 3

2a. Claws on tarsus I with numerous distinct tines. ........

BRE a Re cn FERATNT 2 MaghSOTSRAOG BE (conn OO africanus

2b. Claws I apically and in middle with a few delicate

tines, basally with narrow lamella bearing tines. .....

a irR baad ene Nee clin, eR Anas ettass ua e cultellatus

3a. Length of idiosoma exceeding 450 um. L:W of

SMAthMOSOMId se Oa le. ok, Au we am iat aN, major

3b. Length of idiosoma between 200-400 um. L:W of

enMAOSOmMasd Talal | haat, Oe ean las toad darths. eee de tds 4

4a. Length of idiosoma about 300-400 um. ... fontinalis

4b. Length of idiosoma between 200 and almost

SOLON ty bane We ieeens Mier cis ee ANNERENE | ttc in: 2p Itai Ee rk Uninet 5

5a. L:W of PD 2.1:1. Length ratio of PD:AD 3.1-3.3:1.

sesh sapien Pa De los el eae eh ea bol a mae: Dae portmanni

5b. L:W of PD 1.9:1. Length ratio of PD:AD 2.6—2.7:1.

Lobohalacarus weberi (Romijn & Viets, 1924)

Walterella weberi Romijn & Viets, 1924: 217-220,

figs 3-6.

Lobohalacarus weberi, Schwoerbel 1955: 147, fig. 1;

Green and MacQuitty 1987: 162, fig. 68A—D; Bartsch

2006: 128-130, fig. 5-8a—f, 2007: 74-80, fig. 2, 2011:

493-494, fig. 3A-C.

Walterella weberi quadripora Walter, 1947: 236-237,

fig. 35.

Bonn zoological Bulletin 67 (2): 79-99

Lobohalacarus weberi quadriporus, Viets 1959: pl. 4,

fig. 42, pl. 5, fig. 47.

Lobohalacarus weberi gotoensis Imamura, 1970: 455—

457, figs 1 and 2.

Lobohalacarus weberi tristanensis Bartsch, 1995: 171—

175, figs 1-13.

Species with vague descriptions but expected to belong

to L. weberi:

Halacarus processifer Walter, 1919a: 21-23, fig. 1-3

(only protonymph known);

Lobohalacarus hummelincki Viets, 1940: 191-194, fig,

1,21 and II, 310 and IV, 4;

Lobohalacarus bucharensis Jankovskaja, 1967: 109-

114, fig. 111-7), 21-7), 3(1-6) (only deuto- and proto-

nymph known);

Lobohalacarus bunurong Harvey,

figs 1-6.

1988: 363-365,

Collecting data. Central Madagascar, Antananarivo, An-

jazorobe, River Ranonisoanavola (larger stream E from

main mountain chain), 1200 m asl, 13.2°C, 0.058 mS/

cm, interstitial, 23 Jul. 2001 (MD 012). — South-east-

ern Madagascar, Fianarantsoa, Ionilahy, stream draining

area Marosaro (S from River Ionilahy), 220 m, 21°C,

0.072 mS/cm, interstitial, 12 Aug. 2001 (MD 023). —

South-eastern Madagascar, Fianarantsoa, Ionilahy, small

stream crossing the railroad east of village, 200 m, 15

Aug. 2001 (MD027). — South-eastern Madagascar, Fi-

anarantsoa, Andrambovato, stream 3 km E from the vil-

lage, upstream from the cascade, 900 m, 20 Aug. 2001

(MD 038). — Southern Madagascar, Tulear, Tsimelahy,

River Antarantsa, ca 1 km upstream from village, 300 m,

20.4°C, 0.171 mS/cm, interstitial, 04 Sep. 2001 (MD

058). — Central Madagascar, Antanarivo, Ankaratra, Re-

serve Manjakatompo, left affluent of River Mahiavona,

EM Mantsina, 1750 m, 14.1°C, 0.003 ms/cm, intersti-

tial, 08 Oct. 2001 (MD107). — Northern Madagascar,

Antsiranana, Andapa, right affluent of River Amben-

drana downstream, large cascade, 600 m, 11 Nov. 2001

(MD147).

Short description (Fig. 4A—E). Female: Length (with

frontal spine included) 249-304 [25] um, 228-273 (if

frontal spine excluded). All Madagascar specimens with

frontal spine, spine generally slender (Fig. 4C) and about

21-31 um long, one female with 19 um-long spine. Dor-

sal plates uniformly foveate, foveae 3 um wide. AD and

anterodorsal part of AE fused (Fig. 4A). Opposing edges

of AD and PD truncate. OC oblong, without cornea or

eye pigment. Dorsal setae minute, seven pairs present,

most posterior pair on PD. Ventral plates AE, PE and GA

fused to a shield (Fig. 4B). A pair of epimeral pores about

levelling with aperture of legs II. GO extending anteriad

to the level of apertures of legs IV. Area corresponding

to PE with one rather short dorsal and two long ventral

©ZFMK

86 Ilse Bartsch

Fig. 4A—-E. Lobohalacarus weberi (Romijn and Viets, 1924), female. A. idiosoma, dorsal; B. idiosoma, ventral; C. frontal spine;

D. gnathosoma, ventral; E. leg I, medial. (A, B, D, E, scale = 50 um; C, scale = 10 um) (ascl, anal sclerite; ep, epimeral pore; eu, eu-

pathid setae; fsp, frontal spine; pas, parambulacral setae; I-1 to I-6, trochanter, basifemur, telofemur, genu, tibia, and tarsus of leg I).

setae, genital area with four to five pairs of setae. Each

genital sclerite with two to three gac. Gnathosoma slen-

der, 1.8 times longer than wide. Rostrum almost extend-

ing to end of P-2 (Fig. 4D). P-2 with single dorsal seta,

P-3 with medial spur. Length ratio leg I:idiosoma 0.8:1,

following legs somewhat shorter. Telofemur and genu I

equal in length (Fig. 4E). Telofemora HI and IV with 2/0

dorsal/ventral setae. Genu I with two ventral setae, gen-

erally one seta spur-, the other seta-like, rarely both setae

spur-like. Tibia I ventrally with two pairs of setae, one

pair spiniform, one bristle-like; ventral flank of tibiae II

to IV with (zero to) one smooth and (one to) two pecti-

nate setae. Tarsus I with ventromedial spur, apically with

pair of eupathidia (short sensory setae) and pair of dou-

bled pas (Fig. 4E); tarsus III with 4/1 dorsal/ventral setae,

and tarsus IV with 3/1 setae. Lateral fossa membrane of

tarsus I enlarged, its length 7 um, height 6 um, on follow-

ing tarsi lateral and medial fossa membrane small both in

length and height. Claws on tarsus I smaller than those of

following tarsi.

To get an idea of variants in respect to external char-

acters, as commonly found in Lobohalacarus weberi

(Bartsch 1995, 2007, 2011), details of 25 uncleared fe-

males were examined in a drop of glycerine, but some

Bonn zoological Bulletin 67 (2): 79-99

females were damaged or the characters in question

obscured. The character studied were (1) frontal spine:

present [25]; (2) number of gac per genital sclerite: 2

[12], 3 [38]; (3) number of pgs in each half of genital

plate: 3 [1], 4 [22], 5 [26]; (4) combination of spines (sp)

and bristles (br) on genu I: sp/br [49], sp/sp [1]; (5) num-

ber of dorsal/ventral setae on telofemur III: 2/0 [47], 2/1

[1]; (6) number of dorsal/ventral setae on telofemur IV:

2/0 [49], 2/1 [0]; (7) number of pectinate (p) + smooth

(s) setae on tibia II: 2p+1s [46], 1p+ls [4]; (8) number

of pectinate (p) + smooth (s) setae on tibia III: 2pt+1s

[31], lpt1s [14]; (9) number of pectinate (p) + smooth

(s) setae on tibia IV: 2p+1s [48], 1p+1s [0]; (10) number

of dorsal/ventral setae on tarsus III: 4/1 [43], 4/0 [1], 3/0

[0]; (11) number of dorsal/ventral setae on tarsus IV: 4/1

[1], 3/1 [45], 3/0[2].

Male: Not present.

Juveniles: Length (frontal spine excluded) of deu-

tonymphs 230-247 [3] um, protonymphs 210 [2] um and

larvae 130-162 [4] um. All instars with slender frontal

spine, in one larva that spine very delicate. Ventral plates

AE, PE and GA separated. Tarsus I with three dorsal se-

tae, dorsolateral solenidion, enlarged lateral fossa mem-

brane, ventromedial spur, and two ventral eupathidia.

©ZFMK

Freshwater halacarid mites 87

e /obohalacarus weberi complex

Fig. 5. Records of Lobohalacarus weberi (Romijn & Viets, 1924) and species of the L. weberi complex.

Biology. Eighteen of 26 females enclosed an egg, one

female two eggs. The smallest egg was globular and had

a size of 40 x 40 um, the largest one reached a size of

110 x 56 um.

No male was found in the material from Madagascar.

Though L. weberi is one of the very wide-spread and

commonly recorded species reliable records of males are

lacking.

Remarks. Two easily recognized characters used for

identification of Lobohalacarus weberi are the frontal

spine and the ventral shield. These characters are known

to vary, though rarely. In the samples from Madagas-

car, all individuals have a pointed frontal spine and a

ventral shield. In almost all Madagascan specimens the

frontal spine is very slender, similar to that illustrated by

Schwoerbel (1955: fig. 1(4)). In general, the spine is

somewhat shorter but wider (cf. Romijn & Viets 1924:

figs 3 and 4; Bartsch 2006: fig. 5-8a and b). Among ma-

terial from Inaccessible Island, Tristan da Cunha Islands,

three out of six females had no frontal spine. In two of

these three specimens the anterior margin of the idioso-

ma was evenly arched, in one the spine was reduced to

a hood-like process (Bartsch 1995: figs 12 and 13). Out

of 92 individuals from New Zealand, one female had no

spine but an evenly rounded anterior margin, 91 females

Bonn zoological Bulletin 67 (2): 79-99

had a frontal spine (Bartsch 2007: fig. 2A—C). In the

same material one of the females had no ventral shield,

instead AE, PE and GA were separated (Bartsch 2007:

fig. 2C). Lobohalacarus weberi is expected to be highly

variable in its morphology rather than to be represent-

ed by several cryptic species. Studies on the influence of

habitat parameters on character expression do not exist.

Geographical distribution of Lobohalacarus weberi

and the Lobohalacarus weberi complex (Fig. 5) (cf.

Bartsch 2008a, b, 2011, 2014a; PeSi¢ et al. 2010; Fritz &

Feminella 2011; Stolbov et al. 2018):

Afrotropical Region. — Kenya — Madagascar — Tristan da

Cunha Islands;

Palaearctic Region. Europe: — Austria — Belgium — Unit-

ed Kingdom (England, Northern Ireland, Scotland) —

Bulgaria — Denmark — Faeroerne — Finland — France

— Germany — Greece — Hungary — Iceland — Italy —

Macedonia — Monte Negro — Poland — Portugal — Ro-

mania — Switzerland — Spain — Sweden — The Nether-

lands. Northern Africa: — Tunisia. Asia: — Iran — Japan

(L. weberi gotoensis—Imamura 1970) — Russia (Tyu-

men region) — Uzbekistan (Lobohalacarus bucharen-

sis—Jankovskaja 1967);

©ZFMK

88 Ilse Bartsch

Nearctic Region. — Canada (British Columbia, New

Brunswick, Newfoundland, Ontario, Quebec). — Unit-

ed States (Alabama, Arizona, California, Colorado,

Georgia, Illinois, New Hampshire, New Mexico, New

York, North Carolina, Rhode Island, Tennessee, Vir-

ginia);

Neotropical Region. — Chile (Magallanes) — Peru (Lake

Levandera, Lobohalacarus processifer) — Venezuela

(near Higuerote, Lobohalacarus hummelincki),

Australian Region. — Australia (New South Wales (Lobo-

halacarus sp.), Queensland, Victoria (Lobohalacarus

bunurong and Lobohalacarus sp.), the author expects

these specimens to belong to the L. weberi complex). —

New Zealand (North and South Island);

Pacific Islands. — Hawaiian Islands.

Porohalacarus alpinus (Thor, 1910)

Halacarus alpinus Thor, 1910. 348-351, figs 1 and 2.

Porohalacarus alpinus, Thor 1922: 110, 111; Bartsch

1973: 117-119, figs 1-21, 2007: 80-82, fig. 3, 2011:

497-498, fig. 6A—D; Green & MacQuitty 1987: 164,

fig. 69A—D.

Porohalacarus alpinus alpinus Viets 1927: 465-469,

figs 3-8.

Porohalacarus alpinus alpinus, Bartsch 1987: 85, 2006:

130-132, figs 5-9a-f.

Fig. 6A—G. Porohalacarus alpinus (Thor, 1910). A. idiosoma, dorsal, female; B. idiosoma, ventral, female; C. gnathosoma, ven-

tral, female; D. gnathosoma, dorsal (P-3 and P-4 omitted), female; E. leg I, medial, female; F. idiosoma, ventral, male; G. papilla

with genital acetabula, male. (A—F, scale = 50 um, G, scale = 10 um) (ch-cl, cheliceral claw; ch-f, dorsal flap-like process of

chelicera; co, cornea; eyp, eye pigment; papt+gac, papilla with genital acetabula; P-1 to P-4, first to fourth palpal segment; ret-a,

reticulate area; vs, ventral seta).

Bonn zoological Bulletin 67 (2): 79-99

©ZFMK

Freshwater halacarid mites 89

Porohalacarus alpinus brachypeltatus Viets, 1927: 469,

figs 9, 10.

Porohalacarus alpinus brachypeltatus, Bartsch 1987:

85—86, figs 2 and 3, 2006:132, 5-10a, b.

Collecting data. Southern Madagascar, Tulear, Tsimela-

hy, River Antarantsa, ca 1 km upstream from village,

300 m, 20.4°C, 0.171 mS/cm, interstitial, 04 Sep. 2001

(MD 058). — Northern Madagascar, Antsiranana, Samba-

va, Maroambihy, left affluent of River Lokoho upstream

from the village, 90 m, 26.0°C, 0.010 mS/cm, interstitial,

12 Nov. 2001 (MD149).

Short description (Fig. 6A—G). Female: Length 280-—

318 um [10]. Dorsal plates reticulate, in anterior part of

PD reticulation honey comb-like, 1n posterior part pre-

senting a longitudinal pattern. OC with cornea. AD and

OC with spots of black eye pigment (Fig. 6A), that on

AD narrow, 13 um wide, spot on OC 15-18 um in diam-

eter. Dorsal setae very small, their position as illustrated

in Fig. 6A, ds-6 on PD removed from posterior margin.

Surface of ventral plates finely porose. Genital plate with

(four to) five pairs of pgs, each genital sclerite with (four

to) five external acetabula. Interval between anterior

margin of GP and that of GO somewhat longer than the

latter’s length (Fig. 6B). Gnathosoma hardly longer than

wide (Fig. 6C). Rostrum shorter than gnathosomal base

and hardly extending to middle of P-2. Palps four-seg-

mented, lateral to rostrum, i.e., distance between pair

of P-1 more than their width. P-2 with two dorsal setae

(Fig. 4D), P-3 with medial spur. Chelicera with claw and

wide, flap-like dorsal process. Legs slender, shorter than

idiosoma, length of legs I and IV 0.7 times that of idioso-

ma. Genu I much shorter than telofemur I (Fig. 6E). Tibi-

ae I to IV with 4, 3, 2, 2 ventral setae, in addition tibiae III

and IV with short, faintly pectinate mid-segmental medi-

al seta. Tarsus I with single slender ventral seta (Fig. 6E),

following tarsi without ventral setae. Paired claws with

delicate tines.

Male: Length of idiosoma 286 um, width 185 um [1].

Dorsal aspect similar to that of females, plates similar

in shape, length:width ratio and ornamentation. Length

of AD 65 um, width 93 um, length of OC 40 um, width

20 um, length of PD 188 um, width 130 um. GP and

AP fused to GA, length of that plate 148 um, width

121 um, GA slightly wider than female GP. Length of

male GO 50 um, width 33 um, distance between GO and

anterior margin of GA 58 um, i.e., 1.1 times longer than

GO. GA with 19 pgs on one side, 20 on the other side;

one of genital sclerites with one, the other with two sgs.

Lateral to posterior part of GO a circular area present,

10-12 um in diameter, with internal bars forming re-

ticulate ornamentation (Fig. 6F, ret-a). Posterior to GO

a pair of dome-like areas, 12-14 um in diameter, each

with five gac (Fig. 6G). Each acetabulum, about 7 um

in diameter, with an inner central papilla, 3 um in diam-

eter, and surrounded by inward crooked teeth. Length of

Bonn zoological Bulletin 67 (2): 79-99

spermatopositor slightly more than that of GO. Shape of

gnathosoma and legs similar to those of females.

Juveniles: With two nymphal and a larval instar. Length

of deutonymphs 240-263 um [6], of protonymphs 186—

210 um [6] and of larvae 140-185 [6] um. All instars

with dark spots of eye pigment.

Biology. Fifteen out of 20 females were ovigerous, the

other five held no eggs. The size (length x diameter) of

the eggs was between 55 x 63 um and 88 x 75 um.

Remarks. The females in the Madagascan material

have five (rarely four) external acetabula on each genital

sclerite. In populations from other parts of the world the

number of acetabula is known to vary between four and

nine (Bartsch 2011). In Europe two varieties are known,

P. a. alpinus and P. a. brachypeltatus. Discriminating

characters in females are the length of the idiosoma,

i.e., 297-371 versus 248-270 um, and the position of

the GO in relation to length of the GP, namely distance

from anterior margin of GO to that of GP about the same

versus about half the length of GO, respectively (Viets

1927: fig. 6 versus fig 9; Bartsch 2006: fig. 5-9b versus

fig. 5-10a). Similar, nymphs of P. a. alpinus have, in con-

trast to P. a. brachypeltatus, a \arger distance between the

area with genital acetabula and the anterior margin of GP.

A rather inconspicuous difference is that the PD of P. a.

alpinus is slightly longer than that of P. a. brachypelta-

tus. In the first mentioned the ds-3 insert immediately

adjacent or in the margin of the PD, in the latter within

the narrow band of striated integument. The Madagascan

individuals belong to P. a. alpinus.

In general, all adults in populations of P. alpinus are

females. The above mentioned male is one of the very

rare exceptions. A former record of a male is one of P. a.

brachypeltatus. It was found among more than 2500

slide-mounted adults, all extracted from a population liv-

ing in Northern Germany which inhabited a small former

peat ditch, now filled with rapidly growing Sphagnum sp.

(Sphagnales) (Bartsch 1987). The water in the peat ditch

was acidic, had a low ionic concentration and a sparse

fauna (some few cladocerans, rotifers and testaceans).

Differences between the males, from Madagascar (P. a.

alpinus) and northern Germany (P. a. brachypeltatus),

are the larger size, 286 um versus 254 um, and a higher

number of pgs, namely 39 versus 20 pgs. The morpho-

logical differences between the two forms P. a. alpinus

and P. a. brachypeltatus raises the question whether these

are two separate species, subspecies or ecotypes? In

northern Europe, Porohalacrus a. alpinus inhabits a wide

range of substrata, slightly acidic to alkaline and even

oligohaline brackish water, oligo- and mesotrophic lakes,

ponds and banks of slow flowing rivers, substrata rich

in microcaverns, e.g. dense colonies of small mussels

(Dreissena sp.), colonies of sponges, bryozoans, mosses,

and biofilms on vascular plants, all these habitats have a

©ZFMK

90 Ilse Bartsch

e Porohalacarus alpinus

Fig. 7. Records of Porohalacarus alpinus (Thor, 1910) (Porohalacarus alpinus brachypeltatus included).

rich meio- and microfauna and -flora. In contrast, P. a.

brachypeltatus has been taken only in strongly acidic

Sphagnum peats with sparse associated meiofauna. Judg-

ing by the gut content of P. a. alpinus, namely brownish,

with darker and lighter particles, these mites are carniv-

orous. In contrast, the gut content of P. a. brachypeltatus

is greenish, suggesting that the mites are phytophagous.

Is P. alpinus brachypeltatus a degenerate (impoverished)

form, because of its life in an environment with low con-

centration of cations and meagre diet? Both molecular

and experimental studies, namely rearing of the mites

over several generations under different environmental

conditions, may give an answer. Phenotypic plasticity in

freshwater crustacea has been documented more than a

century ago (Woltereck 1909), and plasticity is not re-

stricted to arthropods, Cattau et al. (2018) recently de-

scribed a rapid morphological change in a bird of prey.

Geographical distribution (Fig. 7) of both P. a. alpi-

nus and P. a. brachypeltatus (Viets 1956; Green & Mac-

Quitty 1987; Bartsch 2007, 2009, 2011; Tolstikov et al.

2005; Semenchenko et al. 2010; Stolbov et al. 2018;

Pepato & da Silva Conceicao 2019 in press):

Afrotropical Region. — Madagascar (new record);

Bonn zoological Bulletin 67 (2): 79-99

Palaearctic Region. Europe: — Austria — Belgium — Great

Britain (England, Scotland, Wales) — Denmark — Fin-

land — France — Germany — Hungary — Iceland — Italy —

Norway — Poland — Russia (Saratov—Volga Biological

Station) — Sweden — Switzerland — The Netherlands —

Ukraine. Northern Africa: — Algeria. Asia: — Turkey —

Russia (Kamchatka, Sakha Republic, Tyumen region);

Australian Region. — Australia (Western Australia—Es-

perance). — New Zealand (North Island).

Nearctic Region. — United States (New Hampshire,

Rhode Island). — Canada (Ontario, Quebec);

Neotropical Region. — Brazil (Rio Grande do Sul—Poro-

halacarus cf. alpinus).

Ropohalacarus pallidus Bartsch, 2013

Ropohalacarus pallidus Bartsch, 2013b: 80-84, figs la—

h, 2a-g.

Collecting data. Central Madagascar, Antananarivo, An-

jazorobe, River Ranonisoanavola (larger stream E from

main mountain chain), 1200 m asl, 13.2°C, 0.058 mS/

cm, interstitial, 23 Jul. 2001 (MD 012). South-eastern

Madagascar, Fianarantsoa, Ionilahy, stream draining

area Marosaro (S from River Ionilahy), 220 m, 21°C,

©ZFMK

Freshwater halacarid mites 91

Fig. 8A—G. Ropohalacarus pallidus Bartsch, 2013. A. idiosoma, dorsal, female; B. idiosoma, ventral, female; C. gnathosoma, ven-

tral, female; D. leg I, medial, female; E. idiosoma and gnathosoma, ventral, protonymph; F. idiosoma, ventral, deutonymph, G. leg

IH, lateral, larva. (Scale = 50 um) (pgs, perigenital seta(e); P-1, first palpal segment; III-1, II-2+3, IN-4, II-5, HI-6, trochanter,

femur, genu, tibia and tarsus of leg III).

0.072 mS/cm, 12 Aug. 2001, interstitial (MD 023). —

South-eastern Madagascar, Fianarantsoa, Ionilahy, small

stream crossing the railroad east from village, 200 m, 15

Aug. 2001 (MD 027). — South-eastern Madagascar, Fi-

anarantsoa, Andrambovato, stream 3 km E from the vil-

lage, upstream from the cascade, 900 m, 20 Aug. 2001

(MD 038). — Southern Madagascar, Tulear, Tsimelahy,

River Antarantsa, ca 1 km upstream from village, 300 m,

20.4°C, 0.171 mS/cm, 04 Sep. 2001, interstitial (MD

058). — Tulear, Andohalela, Isaka, spring area S pass RIP

118 (km36), 700 m, 16.0—-18.4°C, 0.055—0.060 mS/cm,

10 Sep. 2001, interstitial (MD 071). — Northern Mada-

gascar, Antsiranana, Andapa, right affluent River Am-

Bonn zoological Bulletin 67 (2): 79-99

bendrana downstream, large cascade, 600 m, 11 Nov.

2001 (MD147).

Short description (Fig. 8A—G). Female: Length

203-245 [30] um, idiosoma pale, about 1.8 times longer

than wide, its anterior and posterior part narrowed. Eye

pigment lacking. AD and PD fused to an elongate dorsal

shield (Fig. 8A). Ventral shield extending dorsad and in-

cluding AE, PE, GA (Fig. 8B) and parts corresponding

to OC. Integument of dorsal shield with delicate porosi-

ty and faint foveate ornamentation, ventral plates almost

smooth. Dorsal setae very small, ds-1, ds-3 and ds-4 in

dorsal shield, ds-2 in dorsal part of ventral shield. Adanal

setae lacking. Gland pores lacking. Area of ventral shield

©ZFMK

92 Ilse Bartsch

e Ropohalacarus pallidus

* Ropohalacarus uniscutatus

Fig. 9. Records of Ropohalacarus pallidus Bartsch, 2013 (black spot) and R. uniscutatus Bartsch, 1982 (asterisk).

representing AE with two pairs of marginal and one pair

of ventral setae, areas of PE each with one marginal and

one ventral seta, and that of GP with two perigenital se-

tae. Marginal setae of idiosoma longer than ventral setae.

Three pairs of genital acetabula arranged in posterior part

of genital sclerites. Gnathosoma slightly wider than long

(Fig. 8C). Palps four-segmented, extending beyond short,

conical rostrum. Short palps and rostrum visible in dor-

sal aspect. Legs distinctly shorter than idiosoma (length

ratio leg I:idiosoma equalling 0.5:1). Distance between

apertures of pairs of legs II and HI almost equal to half

the length of idiosoma (in the other genera length of id-

iosoma more than twice the distance between these ap-

ertures). Trochanter I in one of the females studied with

delicate medial seta, in general no seta present. Ventral

setae on genu I and three ventral setae on tibia I bipecti-

nate (Fig. 6D).

Male: Not present.

Juveniles. Length of deutonymph 173—226 um [6], of

protonymph 122-173 um [3], of larva 113-128 um [2].

Nymphs with AD and PD separated by striated integu-

ment, remnants of ocular plates and marginal part of epi-

meral plates fused. Ventral plates AE, PE and GP fused to

a single ventral shield, no fissure between area represent-

Bonn zoological Bulletin 67 (2): 79-99

ing AE and GP. Ventral plates delicate; ornamented with

foveae. In both nymphal instars area representing AE

with three pairs of setae (Fig. 8E and F). In deutonymph

area representing PE with one pair of marginal and one

of ventral setae (Fig. 8F) and area of GP with one pair of

pgs. Protonymphal PE solely with pair of marginal setae.

Larval tarsus III, just as tarsi III of following instars, with

three dorsal setae, namely two fossary setae and one so-

lenidion (Fig. 8G).

Biology. Of 32 females studied 13 (41%) were oviger-

ous. In general a single egg was present, its size (length x

diameter) between 45 x 43 um and 70 x 43 um.

Remarks. As illustrated in Bartsch (2013b: fig. 2e),

AD and PD of the deutonymphs, as also those of the pro-

tonymphs, are separated from each other by a narrow,

transverse area of striated integument, but both deuto-

and protonymphs have a ventral shield, including the an-

terior and posterior epimeral and the genital plate (AE,

PE and GP). In contrast to Bartsch (2013b: fig. 2f, 2g),

no transverse rupture was recognized in the new materi-

al studied (three deutonymphs, one protonymph). In the

majority of halacarids the adults have solid exoskeletal

©ZFMK

Freshwater halacarid mites 93

plates. In several species these are fused to a dorsal or

ventral shield. Sometimes, though rarely, both the dor-

sum and venter are covered by a solid shield whereas the

juveniles have the plates separated by tensible striated

integument. Consequently juveniles can grow due to ex-

pansion of the striated integument between the dorsal and

the ventral plates (as illustrated in Halacarellus subter-

raneus — Bartsch 1972 fig. 28A, B versus C, D and E,

F versus G, H) whereas the adults can hardly increase in

size. An exception is Ropohalacarus pallidus. \n the ju-

veniles of this species, the foveate textured integument of

the ventral plates is expected to allow a slight dilatation

(cf. Bartsch 2013b). Data on the thickness of the pro- and

epicuticula are lacking.

No males have ever been found, neither of Ropohal-

acarus pallidus nor of the second species, namely

R. uniscutatus, but the number of specimens studied to

date 1s small (< 50 individuals).

Compared with other Madagascar halacarids, R. palli-

dus is in its general shape most similar to Porohalacarus

alpinus. Distinguishing characters are: the pale idioso-

ma (versus prominent spots of eye pigment), the dorsal

and ventral shield (versus dorsum with AD, OC and PD,

venter with AE, PE and GP), the ornamentation of dor-

sal shield or plates, delicately foveate versus reticulated

with longitudinal polygons, the position of apertures of

legs III, in posterior third of idiosoma (0.7) versus in its

middle (0.5).

Fig. 10A—G. Soldanellonyx monardi Walter, 1919. A. idiosoma, dorsal, female; B. idiosoma, ventral, female; C. part of gnatho-

somal base with P-1, P-2 and P-3, female; D. gnathosoma, ventral, female; E. gnathosoma, lateral, female; F. leg I, ventromedial,

female; G. idiosoma, dorsal, protonymph. (Scale = 50 um) (aps, apical seta on second palpal segment; ascl, anal sclerite; bs, basal

seta on second palpal segment; ep, epimeral pore; exb, excretory bar; eyp, eye pigment; mxs-1, basal maxillary seta; P-1, P-2, P-3,

first, second and third palpal segment).

Bonn zoological Bulletin 67 (2): 79-99

©ZFMK

94 Ilse Bartsch

e Soldanellonyx monardi

Fig. 11. Records of So/danellonyx monardi Walter, 1919.

Geographical Distribution (Fig. 9):

Afrotropical Region. — Madagascar (Bartsch 2013b and

present record).

Soldanellonyx monardi Walter, 1919

S. monardi Walter, 1919b: 238-241, fig. 4-7.

S. monardi, Green & MacQuitty 1987: 156, fig. 6SA—C;

Bartsch 2006: 142, 143, fig. 5-19a—f, 2007: 82, 83,

fig. 3, 2011: 502, fig. 11A—C, 2014a: 167-169, fig. 2A—

K.

S. monardi sarangaensis Viets, 1929: 32-34, fig. 5—7(a,

b).

Subspecies of uncertain position but expected to belong

to S. monardi are:

S. monardi hyogoensis Imamura, 1981: 292 (described in

Imamura 1959: 56, 57, fig. 2a—e).

S. monardi japonicus Imamura, 1971: 334-336, figs 2a—

c, 3a-d.

Collecting data. Central Madagascar, Antananarivo, An-

jazorobe, River Ranonisoanavola (larger stream E from

main mountain chain), 1200 m asl, 13.2°C, 0.058 mS/

Bonn zoological Bulletin 67 (2): 79-99

? Soldanellonyx sp.

cm, interstitial, 23 Jul. 2001 (MD 012). — South-east-

ern Madagascar, Fianarantsoa, Andrambovato, stream

3 km E from the village, upstream from the cascade,

900 m, 20 Aug. 2001 (MD 038). — Southern Madagascar,

Tulear, Tsimelahy River Antarantsa, ca 1 km upstream

from village, 300 m, 20.4°C, 0.171mS/cm, interstitial,

04 Sep. 2001 (MD 058). — Central Madagascar, Anta-

nanarivo, Ankaratra, Reserve Manjakatompo, left afflu-

ent of River Mahiavona, EM Mantsina, 1750 m, 14.1°C,

0.003 ms/cm, interstitial, 08 Oct. 2001 (MD 107).

Short description (Fig. 10A—G). Female: Length

270-335 um [8]. With faint spots of brown eye pigment

in anteromedian part of AD and anterior part of OC

(Fig. 1OA). AD, OC and PD with reticulate ornamenta-

tion, reticulation most distinct within pair of faint costae

of PD. Pairs of glp-1 to glp-5 and ds-1 to ds-4 as illustrat-

ed, ds-5 lacking, ds-6 on anal cone. Delicate punctation

of ventral plates presenting reticulate pattern. AE, PE and

GP separated. AE with pair of epimeral pores (Fig. 10B).

Each PE with one ventral and lateral but no dorsal seta.

Gnathosoma somewhat wider than long (Fig. 10D). Pair

of palps attached dorsally, distance between P-1 less than

their width (Fig. 10C). P-2 somewhat flattened, length

1.3 times the height (Fig. 1OE); basal seta on P-2 short;

spiniform, distal seta long and slender. P-3 with large me-

©ZFMK

Freshwater halacarid mites 95

dial spine, its length 0.9 of that of P-3. P-4 ending with a

similar wide spine. Legs shorter than idiosoma, length of

legs I and IV about 0.6 and 0.7 times that of idiosoma, re-

spectively. Length of telofemur I 1.5 times the height. In

addition to long, slender setae, telofemur, genu and tibia

of leg I with 2, 2 and 1 short and spiniform dorsal setae

and 0, 2, 2 long ventral spines, respectively. Pair of ven-

tral spines on genu I distinctly, on tibia I faintly bipect-

inate. Claws I with solid, mushroom-like arranged tines

(Fig. 10F), tines on claws on following tarsi in J-shaped

arrangement.

Male: Not present.

Juveniles: Length of deuto-, protonymphs and larvae

205-285 [8], 180-255 [7] and 143-148 um [2], respec-

tively.

Biology. Of eight females studied six held eggs. Size

of eggs 10 x 10 to 40 x 45 um (length x diameter). If

present, the excretory material 1s in form of small, brown

globuli. These are concentrated within a rod-like struc-

ture (Fig. 10G).

Remarks. Sol/danellonyx monardi is presently the only

Soldanellonyx species recorded from the Afrotropics.

Soldanellonyx chappuisi Walter, 1917 and S. visurgis

Viets, 1959, as S. monardi reported from several con-

tinents, have as yet not been taken but are expected to

be found in future studies in Africa or the Afrotropical

region. Soldanellonyx chappuisi and S. visurgis have, in

contrast to S. monardi, more slender telofemora I (length

more than 1.5 times the height), four ventral bristles or

Spines on tibia I (versus two ), more slender and longer

P-2 (length more than twice the height) and the two dor-

sal setae on P-2 are similar-sized (versus basal seta less

than half the length of distal seta). In addition, no epime-

ral pores are seen in S. chappuisi and S. visurgis but these

are present in S. monardi.

Another six So/danellonyx species are described, one

species has been collected in both southern Japan and

Kamchatka, each one of the others from a single ge-

ographical region, four from Japan, one from the Lake

Baikal.

In Soldanellonyx monardi, as also in S. chappuisi and

S. visurgis, males are extremely rare or absent. Those re-

corded by Sokolov (1952), Efford (1959) and Imamura

(1981) need confirmation. The author expects the males

of S. monardi to differ from females by a high number

(>20) of pgs arranged around the GO and, of course, the

presence of the spermatopositor. Slight differences in

the outline of the GA, as described by Sokolov (1952)

and Imamura (1981), are commonly found in females.

Sokolov (1952: fig. 91,4) presented an illustration of a

male GA with a slightly larger number of genital acetabu-

la (11-12 pairs) and pgs (5-6 pairs) but else similar to that

of females, the typical spermatopositor is not illustrated.

Imamura (1981) described the ‘penis skeleton’ as having

Bonn zoological Bulletin 67 (2): 79-99

four hook-shaped claws; the number of the genital ace-

tabula is similar to that of females. Hook-shaped claws on

the spermatopositor are else unknown in halacarid males

and those mentioned by Imamura (1981) may represent

the genital spines of an ovipositor. Efford (1959) did not

present any morphological details.

Geographical distribution (Fig. 11) (of S. monardi

and its subspecies) (Fig. 8) (Bartsch 2008a, 2011, 2014a;

Tolstikov et al. 2005; PeSi¢ et al. 2010; Stolbov et al.

2018; Pepato & da Silva Concei¢ao, in press):

Afrotropical Region. — Kenya — Madagascar;

Palaearctic Region. Europe: — Austria — Belgium — Unit-

ed Kingdom (England, Northern Ireland, Scotland,

Wales) — Bulgaria — Crimea — Croatia — Denmark —

Faeroerne — Finland — France — Germany — Hungary

— Iceland — Italy — Luxemburg — Macedonia — Monte

Negro — Portugal — Romania — Russia (Karelia, Kola

Peninsula, Lake Onega) — Switzerland — Spain — Swe-

den — The Netherlands. Northern Africa: — Tunisia.

Asia: — Japan — Russia (Kamchatka, Tyumen region);

Oriental Region. — Indonesia (Java) — Vietnam;

Australian Region. — Australia (New South Wales,

Queensland). — New Zealand (North Island);

Pacific Islands. — Hawaiian Islands;

Nearctic Region. — Canada (British Columbia, Manitoba,

New Brunswick, Newfoundland, Ontario, Quebec). —

United States (Alabama, Arizona, California, Georgia,

Indiana, Missouri, New Hampshire, New York, North

Carolina, Missouri, Oregon, Pennsylvania, Rhode Is-

land, Texas, Tennessee, Virginia). The Soldanellonyx

species, by Vinke (2013) mentioned from the North-

west Territories (Canada), may belong to S. monardi (in

Fig. 11 marked by a question mark);

Neotropical Region. — Brazil (Sao Paulo) — Chile (Ma-

gallanes) — Falkland Islands (Malvinas).

DISCUSSION

Biogeography

The halacarid species extracted from shallow water sandy

deposits at the banks of streams and creeks on Madagas-

car belong to six species in five genera. Considering that

the knowledge regarding the freshwater halacarid fauna

of the world is very sparse, it is striking that none of the

genera is restricted to Madagascar or to the Afrotropical

Region, all are cosmopolitans. At species level, a single

one (Ropohalacarus pallidus) is at present only known

from Madagascar, but this does not mean that the spe-

cies is endemic since no similar habitats have been stud-

ied in adjacent parts of Africa. The three species Lobo-

halacarus weberi, Porohalacarus alpinus and Soldanell-

onyx monardi are the ones generally found in the course

of similar studies in northern Europe, North America and

©ZFMK

96 Ilse Bartsch

New Zealand (Husmann & Teschner 1970, Bartsch 2007,

2011). The three species are spread on all continents ex-

cept for Antarctica from where no adequate substrata

have been studied in respect to their freshwater halacarid

fauna. The two Limnohalacarus species taken on Mada-

gascar are wide-spread, too. Since most Limnohalacarus

species have been found in warm-temperate and tropi-

cal regions, these may avoid cold-temperate waters. One

exception, Limnohalacarus wackeri (Walter, 1914) is re-

corded from northern European and Asian waters, from

southern Finland and Kamchatka, respectively (Paasivir-

ta 1975; Tolstikov et al. 2005).

Madagascar, as well as New Zealand, is known to have

a unique water mite fauna (Gerecke 2004; Sirvid et al.

2011). Gerecke (2004) presented a list of Hydrachnidia,

collected on Madagascar, which included 63 species. Of

these, 35 (or 56%) had only been taken on Madagascar,

and 24 (or 38%) on both Madagascar and Africa; only

four species (or 6%) proved to be wide-spread (Gerecke

2004). Since then more Madagascan species have been

described (Goldschmidt 2008; PeSi¢ et al. 2013). Simi-

lar, the New Zealand mite fauna is known to include a

very high number of endemisms; in many well studied

mite groups, terrestrial as well as aquatic mites, 80 %

of the species and 35 of the 59 (59 %) of the freshwater

genera are expected to be restricted to New Zealand (Sir-

vid et al. 2011). Because of the low reproduction rate of

halacarids, absence of dispersal instars but unusual wide

geographical distribution of not just one but several more

or less syntopic living species, Bartsch (1996, 2007)

expects these species to have colonized and spread on

Pangea since the Mesozoic or even Pre-Mesozoic. Mi-

nor morphological differences between populations are

assumedly due to phenotypic plasticity and no evidence

of cryptic speciation.

Limnohalacarus: Characters not or rarely observed

in other halacarid genera

Species of the genus Limnohalacarus demonstrate two

characters which are either restricted to this genus or

only rarely found in other halacarid genera. Unique

is that the eggs are fixed to the basal part of the tibiae

IV (Viets 1940; Ramazotti & Nocentini 1960; Petrova

1966; Bartsch 1999) but the oviposition has not been ob-

served. Up to 9 eggs, or their remnants, can be found

per leg (Bartsch 1999). The embryos in the eggs are of-

ten in different states of development. In a population

of L. wackeri from northern Italy, the development of

the eggs took about one month (Ramazotti & Nocentini

1960). In general, halacarids deposit their eggs via the

Ovipositor into a substratum (Teschner 1963; Kirchner

1969; Bartsch 1972), either singly or in clusters. When

at rest, the ovipositor is retracted and in most halacarid

females it extends internally slightly or distinctly beyond

the anterior margin of the genital foramen. The outline

Bonn zoological Bulletin 67 (2): 79-99

of the ovipositor and its genital spines are at least partly

visible through the genital plate. The extruded ovipositor

ends with genital spines. In some genera these spines are

more or less spiniform, in others palmate, either faintly

or distinctly sclerotized (cf. Bartsch 2015). In contrast

to the majority of halacarid species, the ovipositor of

Limnohalacarus novus is short and narrow (LxW 74 x

10 um), the 1-2 um-long genital spines at the end of the

Ovipositor are very faintly sclerotized, they arise from

minute papillae.

Another detail often found in Limnohalacarus species

is the combination of an ovoid, somewhat flattened body

with accumulated excretory material, and small anal

sclerites. Such an excretory body is already present in

larvae but is most conspicuous in adults (Walter 1914;

Petrova 1966; Ramazotti & Nocentini 1960; Pepato &

Dos Santos 2015). In most of the other halacarid spe-

cies, in larvae, nymphs and adults, the waste products

are concentrated within a dorsomedian bar (cf. So/danel-

lonyx monardi—F 1g. 10G; Isobatrus uniscutatus (Viets,

1939)—Bartsch 2014b: fig. 1). The length of the bar,

if present, can correspond to about half or almost two-

thirds of the idiosomal length, it is filled with white or

light brown granulated material. The ‘light mesial line’,

mentioned by Johnston (1836) in the description of

Thalassarachna basteri (Johnston, 1836), and the ‘dark

line’ in Lohmannella falcata (Hodge, 1863), described by

Hodge (1863), may represent such bars. Halacarids have

the excretory organ dorsal to the gut (Thomae 1926), def-

ecation is through the anus which is guarded by the anal

sclerites. Anatomical details in Thalassarachna basteri

have been studied by Thomae (1926).

The presence of ovoid excretory bodies, in Limno-

halacarus novus distinctly stratified, is not restricted to

Limnohalacarus species but documented also from oth-

er halacarid genera. Examples are Halacarus excellens

Lohmann, 1907 and Rhombognathus amplus Bartsch,

2013 (Bartsch 2010, 2013c) as well as other prostig-

matid mite families, e.g. the Cunaxidae (Kielczewsk1

& Wisniewski 1977). In halacarid species with such an

excretory body the anal sclerites are unusual small, just

as in L. novus. More species with small anal sclerites

are Bathyhalacarus anomalous Bartsch, 2005, Rhombo-

gnathus bulbosus Bartsch, 2005, R. cyrtonotus Bartsch,

2000, R. delicatulus Bartsch, 2000 (Bartsch 2000c,

2005a, b) but in individuals of these species no ovoid

bodies with excretory material were found. However, the

number of mites studied is very small. At present it seems

that halacarid species with a large ovoid excretory body

(20-50 % of idiosomal length) have small anal sclerites,

though not all mites with small anal sclerites have such

an ovoid body.

Little is known about feeding, almost nothing about

digestion and defecation in halacarid mites. Most

halacarids, marine as well as freshwater species, are

expected to be carnivorous. Limnohalacarus wackeri,

©ZFMK

Freshwater halacarid mites 97

for example, could be reared with a diet of small cili-

ates, Lobohalacarus weberi with pieces of oligochaetes

(Teschner 1963; Ramazotti & Nocentini 1960). Larvae

are known to show an excretory bar or ovoid body soon

after commencement of feeding (Kirchner 1969; Rama-

zotti & Nocentini 1960). Rate, periods and frequency of

defecation are not known. A general idea may give the

studies by Bowman (2017a, b) who examined the feed-

ing of the mesostigmatid Pergamasus longicornis (Ber-

lese, 1906), a carnivorous species as Limnohalacarus,

but terrestrial and with a length of almost 1300 um dis-

tinctly larger than Limnohalacarus. In P. longicornis up

to three weeks may be needed to clear the idiosoma of

excretory material (Bowman 2017a, b).

Among halacarids most species have excretory bars,

only a small number has ovoid bodies with concentri-

cally arranged layers (e.g. Rhombognathus amplus).

Among the few specimens studied of the latter species,

collected in Singapore mangroves, each following instar

had an excretory body almost similar or larger in size

than that of the preceding stage. The length of the idioso-

ma (in parentheses) and LxW data of the excretory body

are in protonymphs (209-235 um) about 37 x 25 um,

in deutonymphs (242-284 um) 37-75 x 25-30 um, in

tritonymphs (245-354 um) 127 x 30 um, and in adults

(364-415 um) 110-182 x 62-77 um. For critical ex-

amination of taxonomic details, halacarid mites have to

be cleared, 1.e., the body content has to be removed. In

R. amplus the stiff excretory body could only with dif-

ficulty be pressed through the camerostome, though the

size of the latter (LxW about 70 x 100 um) is much larger

than that of the anal slit (length about 20 um). How can

such large excreta pass through the small anal opening?

Do the mites accumulate excretory material during their

lives, from the larval to the adult stage? This seems to be

unlikely; the amount of excreta produced in the course

of the mite’s life 1s expected to be much larger than that

enclosed in the ovoid body. Most halacarids studied have

a life span of about 15 months, Limnohalacarus up to

24 months (Ramazotti & Nocentini 1960; Bartsch 1972,

1987), the major part takes the adult stage. About one

quarter of the Limnohalacarus novus females studied had

no excretory bodies. This quarter included stages from

recently hatched to cocoon-carrying females, and accord-

ingly we can expect that the small anal sclerites do not

prevent defecation. Since quiescent or moulting stages

were not represented in the samples, we do not know if at

least partly the mites can defecate during or immediately

after moulting, as long as the integument is not hardened

yet.

The two just mentioned species differ in their life style

which in turn will influence digestion and defecation.

Rhombognathus amplus is phytophagous and restricted

to the upper tidal zone, several hours a day emerged.

The delicate algal film inhabited quickly desiccates.

The mites studied were collected during low tide, con-

Bonn zoological Bulletin 67 (2): 79-99

sequently they were dehydrated and that certainly had

an influence on the compactness of the waste products

and the defecation. In contrast, Limnohalacarus species

are carnivorous and live in an at least constantly water-

logged habitat. Feeding and metabolic activities are not

hampered by desiccation and hence the wastes may be

elastic enough to be pressed through the small and nar-

row anal slit.

Acknowledgements. The halcarids have been collected by Rein-

hard Gerecke (Tubingen) and Tom Goldschmidt (Mtinchen)

during a visit to Madagascar, meant to study the aquatic water

mite fauna in river sediments. This field trip was financed by

Dave Cook (Paradise Valley, USA). Reinhard Gerecke forward-

ed the mites to me and Almir Rogério Pepato (Belo Horizonte,

Brasil) informed me about unpublished records of halacarids

from Brazil. To all my sincerest thanks.

REFERENCES

André HM, N’Dri JK (2012) Bréviaire de taxonomie des

acariens. Abc Taxa, la série de manuels dédiés aux renforce-

ments des capacités en taxonomie et en gestion des collec-

tions 13: 1-184

Bader C (1968) Wassermilben aus Zentralafrika. Annales du

Musée Royal de |’Afrique Central, Série A, Zoologie 163:

1-50

Balian EV, Segers H, Lévéque C, Martens K (2008) An in-

troduction to the freshwater animal diversity assessment

(FADA) project. Hydrobiologia 595: 3-8

Bartsch I (1972) Ein Beitrag zur Systematik, Biologie und

Okologie der Halacaridae (Acari) aus dem Litoral der Nord-

und Ostsee. I. Systematik und Biologie. Abhandlungen und

Verhandlungen des Naturwissenschaftlichen Vereins zu

Hamburg, Neue Folge 16: 155-230

Bartsch I (1973) Porohalacarus alpinus (Thor) (Halacari-

dae, Acari), ein morphologischer Vergleich mit marinen

Halacariden nebst Bemerkungen zur Biologie dieser Art. En-

tomologisk Tidskrift 94: 116-123

Bartsch I (1982) Halacariden (Acari) im StBwasser von Rhode

Island, USA, mit einer Diskussion tiber Verbreitung und Ab-

stammung der Halacaridae. Gewasser und Abwasser 68/69:

41-58

Bartsch I (1987) Zur Biologie, Okologie und Verbreitung der

siBwasserbewohnenden Halacaride Porohalacarus alpinus

(Acari). Archiv fir Hydrobiologie 111: 83-93

Bartsch I (1995) A new subspecies of the freshwater halacarid

mite Lobohalacarus weberi (Romijn and Viets) (Halacari-

dae, Acari) from a Southern Atlantic Ocean Island. Annals of

the Cape Provincial Museums, Natural History 19: 171-180

Bartsch I (1996) Halacarids (Halacaroidea, Acari) in freshwa-

ter. Multiple invasions from the Paleozic onwards? Journal of

Natural History 30: 67-99

Bartsch I (1999) Two new freshwater mites of the genus Lim-

nohalacarus (Halacaridae: Acari) from Australia. Records of

the Western Australian Museum 19: 443-450

Bartsch I (2000) Rhombognathinae (Acari: Halacaridae) from

the Great Barrier Reef, Australia. Memoirs of the Queensland

Museum 45: 165-203

Bartsch I (2005a) Halacaridae from the depths of Western Ant-

arctica (Arachnida: Acari). Senckenbergiana Biologica 85:

31-41

©ZFMK

98 Ilse Bartsch

Bartsch I (2005b) Upper littoral rhombognathines (Acari:

Halacaridae) of Singapore: description of three new species.

Cahiers de Biologie Marine 46: 273-287

Bartsch I (2006) 5. Acari: Halacaroidea. Pp. 113-157 in Ge-

recke R (ed.) StBwasserfauna von Mitteleuropa Vol. 7,2-1,

Elsevier, Spektrum Akademischer Verlag, Miinchen

Bartsch I (2007) Freshwater Halacaridae (Acari) from New

Zealand rivers and lakes, with notes on character variability.

Mitteilungen aus dem Hamburgischen Zoologischen Muse-

um und Institut 104: 73-87

Bartsch I (2008a) Freshwater halacarid mites (Halacaridae:

Prostigmata: Acari) from Tunisia, three new records and

notes on geographical distribution of these species. Entomo-

logische Mitteilungen aus dem Zoologischen Museum Ham-

burg 15: 15-27

Bartsch I (2008b) Halacarid mites (Acari) in a freshwater in-

fluenced beach of North Stradbroke Island, Moreton Bay,

Queensland. Pp 117-130 in Davie PJF, Phillips JA (eds)

Proceedings of the Thirteens International Marine Biologi-

cal Workshop. The Marine Fauna and Flora of Moreton Bay,

Queensland. Memoirs of the Queensland Museum 54 (1)

Bartsch I (2009) Checklist of marine and freshwater halacarid

mite genera and species (Halacaridae: Acari) with notes on

synonyms, habitats, distribution and descriptions of the taxa.

Zootaxa 1998: 1-170

Bartsch I (2010) Halacarus excellens Lohmann, 1907 (Acari:

Halacaridae), a new record a century later, re-description and

notes on this and other Antarctic halacarid species. Entomo-

logische Mitteilungen aus dem Zoologischen Museum Ham-

burg 15: 157-170

Bartsch I (2011) North American freshwater Halacaridae

(Acari): literature survey and new records. International

Journal of Acarology 37: 490-510

Bartsch I (2013a) The freshwater halacarid genus Limnohal-

acarus (Halacaridae, Acari), species diagnoses, distribution

and relationship. Verhandlungen des naturwissenschaftlichen

Vereins in Hamburg 47: 185-220

Bartsch I (2013b) Freshwater halacarid mites (Acari: Halacar-

idae) from Madagascar. New records and description of a

new species. Acarologia 53: 77-87. DOI: 10.1051/acarolo-

gia/20132080

Bartsch I (2013c) Upper littoral rhombognathines (Acari:

Halacaridae) from Singapore: description of a new species.

Acarologia 53: 305-313 DOI: 10.1051/acarologia/20132097

Bartsch I (2013d) Arachnida-Halacaridae checklist. World

checklist of freshwater Halacaridae species. http://fada.bio-

diversity.be/group/show/32

Bartsch I (2014a) New records of freshwater Halacaridae

(Acari) from the Oriental Region and a supplementary de-

scription of Soldanellonyx visurgis. International Journal of

Acarology 40(2): 165-173

Bartsch I (2014b). Die Meeresmilbe Jsobactrus uniscuta-

tus (Acari: Halacaridae), ein ’Européer” in australischen

Gewéassern. Verhandlungen des Naturwissenschaftlichen

Vereins zu Hamburg, Neue Folge 48: 177-188

Bartsch I (2015) The genital area of Halacaridae (Acari), life

stages and development of morphological characters and im-

plication on the classification. Zootaxa 3919: 201-259

Berlese A (1906) Monographia del genere Gamasus Latr. Redia

3: 65-304

Bowman CE (2017a) Modelling Malpighian tubule crys-

tals within the predatory soil mite Pergamasus longicornis

(Mesostigmata: Parasitidae). Experimental and Applied Ac-

arology 72: 35-59

Bonn zoological Bulletin 67 (2): 79-99

Bowman CE (2017b) Gut contents, digestive half-lives and

feeding state prediction in the soil predatory mite Pergama-

sus longicornis (Mesostigmata: Parasitidae). Experimental

and Applied Acarology 73: 11-60

Cattau CE, Fletcher RJ Jr, Kimball RT, Miller CW, Kitchens

WM (2018) Rapid morphological change of a top predator

with the invasion of a novel prey. Nature Ecology & Evolu-

tion 2 (1):108—-115. DOI: 10.1038/s41559-017-0378-1

Chatterjee T, Chang CY (2005) A new species of Limnohal-

acarus (Acari: Halacaridae) from India. Bulletin. Institut

Royal des Sciences Naturelles de Belgique, Entomologie 75:

23-27

Efford I (1959) The distribution of three British Porohalacari-

dae (Acari). Entomologist’s monthly Magazine 95: 84

Fritz KM, Feminella JW (2011) Invertebrate colonization of

leaves and roots within sediments of intermittent Coastal

Plain streams across hydrologic phases. Aquatic Sciences 73

(4): 459-469

Gerecke R (2004) The water mites of Madagascar (Acari,

Hydrachnidia): a revised list completed by original material

conserved at the Muséum national d’ Histoire naturelle, Paris.

Zoosystema 26 (3): 393-418

Goldschmidt T (2008) Taxonomical, ecological and zoo-

geographical studies on anisitsiellid water mites (Acari:

Hydrachnidia: Anisitsiellidae Koenicke, 1910) from Mada-

gascar. Zootaxa 1954: 1-120

Green J (1984) The occurence of Limnohalacarus (Acari:

Halacaridae) in Lake No, White Nile. Hydrobiologia 110:

135-136

Green J, Corbet SA, Betney E (1974) Ecological studies on

crater lakes in West Cameroon. Debundsha Lake. Journal of

Zoology, London 173: 199-223

Green J, MacQuitty M (1987) Halacarid Mites. Synopses of the

British Fauna (New Series) 36: 1-178

Harvey M (1988) A new species of Lobohalacarus from Aus-

tralia (Chelicerata: Acarina: Halacaridae). Memoirs of the

Museum of Victoria 49: 363-365

Husmann S, Teschner D (1970) Okologie, Morphologie und

Verbreitungsgeschichte subterraner Wassermilben (Limno-

halacaridae) aus Schweden. Archiv ftir Hydrobiologie 67:

242-267

Hodge G (1863) Contributions to the marine zoology of Sea-

ham Harbour. On some undescribed marine Acari. Transac-

tions of the Tyneside Naturalists’ Field Club 5(4): 298-303

Imamura T (1959) Water-mites (Hydrachnellae and Porohal-

acaridae) from the subterranean waters of Akiyoshidae Karst,

Japan. Japanese Journal of Zoology 12: 251-255

Imamura T (1970) The fauna of the insular lava caves in West

Japan. II. Porohalacaridae (Acari). Bulletin of the National

Science Museum, Tokyo 13: 455-458

Imamura T (1971) The fauna of the lava lakes around Mt. Fu-

ji-san. V. Limnohalacarinae (Acari). Bulletin of the National

Science Museum, Tokyo 14: 333-336

Imamura T (1981) Fresh-water halacarid mites from Oahu Is-

land, Hawaii. Annotationes Zoologicae Japonensis 54: 287—

292

Jankovskaja AI (1967) [On the fauna of ground-waters of mid-

dle Asia. I. Hydracarina of the fam. Porohalacaridae (Acari)

in the ground-waters of the wells in Kizylkum desert.] Trudy

Zoologi¢eskogo Instituta, Akademija Nauk SSSR, Leningrad

43: 109-117 (In Russian)

Johnston G (1836) Illustrations in British zoology. Magazine of

Natural History 9 (63): 353-357

©ZFMK

Freshwater halacarid mites 99

Kielczewski B, Wisniewski J (1977) Irregularities in the gua-

nine expel in the mite Cunaxa setirostris (Herm) (Acarina,

Cunaxidae). Acarologia 19: 619-621

Kirchner W-P (1969) Zur Biologie und Okologie von Halacarus

basteri basteri Johnson 1836 (Acari, Trombidiformes).

Oecologia (Berlin) 3: 56-69

Lohmann H (1907) Uber einige faunistische Ergebnisse der

Deutschen Stidpolar-Expedition unter besonderer Bertick-

sichtigung der Meeresmilben. Schriften des Naturwissen-

schaftlichen Vereins fiir Schleswig-Holstein 14: 1-14

Ojeda M, Rivas G, Alvarez F (2016) First record of the genus

Limnohalacarus (Acari: Halacaridae) from Mexico (Primer

registro del género Limnohalacarus (Acari: Halacaridae) de

Mexico). Revista Mexicana de Biodiversidad 87: 1131-1137

Paasivirta L (1975) Distribution and abundance of Halacaridae

(Acari, Trombidiformes) in the oligotrophic lake Paajarva,

Southern Finland. Annales Zoologici Fennici 12: 119-125

Pepato AR, Dos Santos Costa SG (2015) New records of the

genus Limnohalacarus (Halacaridae, Trombidiformes) from

southern Brazil. Check List The Journal of Biodiversity Data

11 (1): Article 1500. www. biotaxa.org/cl

Pepato AR, da Silva Concei¢aéo PH (2019) Chapter 17 Halacar-

idae. (in press) in Thorp JH (ed.) Thorp and Covich’s Fresh-

water Invertebrates, Fourth Edition, Volume HI, Keys to

Neotropical and Antarctic Fauna. Academic Press, 672 pp.

PeSi¢ V, Smit H, Datry T (2010) New records of water mites

(Acari: Hydrachnidia, Halacaroidea) from Patagonia (Chile).

Systematic & Applied Acarology 15: 151-160

PeSi¢ V, Cook D, Gerecke R, Smit H (2013) The water mite

family Mideopsidae (Acari: Hydrachnidia): a contribution to

the diversity in the Afrotropical region and taxonomic chang-

es above species level. Zootaxa 3720: 1-75

Petrova A (1966) Deux nouveaux halacariens d’Israél — Lim-

nohalacarus capernaumi n. sp. et Lohmannella heptapegoni

n. sp. International Journal of Speleology 2: 355-362

Ramazotti G, Nocentini AM (1960) Porohalacaridae (Hydra-

carina) del Lago di Mergozzo. Memorie dell’Istituto Italiano

di Idrobiologia Dott. Marcode Marchi 12: 185-200

Romijn G, Viets K (1924) Neue Milben. Archiv fur Naturge-

schichte 90: 215-225

Schwoerbel J (1955) Uber einige Porohalacariden (Acari) aus

dem siidlichen Schwarzwald. Zoologischer Anzeiger 155:

146-150

Semenchenko KA, Abé H, Boeskorov GG (2010) New data on

the water mite fauna (Acari, Hydrachnidia, Halacaroidea) of

the Sakha Republic (Yakutia). Entomological Review 90 (2):

218-229

Sirvid PJ, Zhang Z-Q, Harvey MS, Rhode BE, Cook DR,

Bartsch I, Staples DA (2010) Phylum Arthropoda Chelicerata

horseshoe crabs, arachnids, sea spiders. Chapter 6. Pp. 50-89

in Gordon DP (ed.) New Zealand Inventory of Biodiversity

2 (6)

Sokolov II (1952). Vodjanye kle8¢i. II. Halacarae. Fauna SSSR

5: 1-201

Stolbov VA, Popova VV, Sheikin SD, Tupitsyn SS (2018) Wa-

ter mites (Acariformes: Hydrachnidia, Halacaroidea) of bogs

of Western Siberia (Russia). Ecologica Montenegrina 18:

102-109

Teschner D (1963) Die Biologie, Verbreitung und Okologie der

Grundwassermilbe Lobohalacarus weberi quadriporus (Wal-

ter, 1947), Limnohalacaridae Acari. Archiv fir Hydrobiolo-

gie 59: 71-102

Bonn zoological Bulletin 67 (2): 79-99

Thomae H (1926) Beitrage zur Anatomie der Halacariden. Zoo-

logische Jahrbiicher, Abteilung 2, 47: 155-190

Thor S (1910) Die erste norwegische Sufwasserform der

Halacariden. Zoologischer Anzeiger 36: 348-351

Thor S (1922) Neue Acarina-Formen aus meinen alteren Samm-

lungen, nebst Bemerkungen tiber Arten, Gattungen und Fa-

milien. Nyt Magasin for Naturvidenskaberne 61: 91-118

Tolstikov AV, Vvedenskaya TL, Stolbov VA (2005) Preliminary

data on the fauna of fresh-water mites (non-Parasitengona) of

the Kamchatka Peninsula. Vladimir Ya. Levanidov‘s Bienni-

al Memorial Meetings 3: 309-311 (In Russian, with English

Summary)

Viets K (1927) Mitteilung ber das Vorkommen von Halacariden

in der Kiemenhohle des FluBkrebses. Verhandlungen. Inter-

nationale Vereinigung fiir theoretische und angewandte Lim-

nologie 3: 460-473

Viets K (1929) Uber StiRwasser-Halacaridae. (Vierte Mittei-

lung tber Hydracarina von den Sunda-Inseln.) Zoologischer

Anzeiger 86: 27-34

Viets K (1939) Meeresmilben aus der Adria (Halacaridae und

Hydrachnellae, Acari). Archiv fiir Naturgeschichte (Neue

Folge) 8: 518-550

Viets K (1940) Zwei neue Porohalacaridae (Acari) aus Sid-

amerika. Zoologischer Anzeiger 130: 191-201

Viets K (1956) Die Milben des StiBwassers und des Meeres.

Katalog der Halacaridae. Meeresmilben, II. Abschnitt.

Pp. 641-870 in Viets K (ed.) Die Milben des St®Bwassers

und des Meeres. Hydrachnellae et Halacaridae, Acari. VEB

Fischer Verlag, Jena

Viets K (1959) Die aus dem Einzugsgebiet der Weser bekann-

ten oberirdisch und unterirdisch lebenden Wassermilben.

Veroffentlichungen des Instituts fiir Meeresforschung in

Bremerhaven 6: 303-513

Vinke K (2013) An investigation of sampling protocols and

community involvement in stream biomonitoring in the

Sahtu Settlement Area, Northwest Territories. Master of Sci-

ence Thesis, MS00449, University of Prince Edward Island

(Canada). 293 pp. ISBN: 9780499004499

Walter C (1914) Notizen tber die StBwasserformen der

Halacariden nebst Beschreibung einer neuen Art. Archiv fiir

Hydrobiologie 9: 279-285

Walter C (1917) Schweizerische StiBwasserformen der

Halacariden. Revue Suisse de Zoologie 25: 411-423

Walter C (1919a) Hydracarinen aus den peruanischen Anden

und aus Brasilien. Revue Suisse de Zoologie 27: 19-59

Walter C (1919b) Schweizerische StiBwasserformen der

Halacariden. Revue Suisse de Zoologie 27: 235-242

Walter C (1935) Hydracarina. In Voyage de Ch. Alluaud et P.A.

Chappuis en Afrique Occidentale frangaise (Déc. 1930—Mars

1931). Archiv fir Hydrobiologie 28: 69-136

Walter C (1947) Neue Acari (Hydrachnellae, Porohalacari-

dae, Trombiidae) aus subterranen Gewassern der Schweiz

und Rumaniens. Verhandlungen der Naturforschenden Ge-

sellschaft in Basel 58: 146-238

Walter C, Bader C (1952) Mission scientifique de l’Omo. — Hy-

dracarina. Mémoires du Muséum Nationale d’ Histoire Natu-

relle, Série A, Zoologie 4: 87-236

Woltereck R (1909) Weitere experimentelle Untersuchungen

uber Artveranderung, speziell tber das Wesen quantita-

tiver Artunterschiede bei Daphniden. Verhandlungen der

deutschen zoologischen Gesellschaft 19: 110-172

©ZFMK

BHL

i

Blank Page Digitally Inserted

FORSCHUNGS

Bonn zoological Bulletin 67 (2): 101-107

2018 - Weber M. et al.

https://doi.org/10.20363/BZB-2018.67.2.101

Research article

urn: |sid:zoobank.org:pub:24D17A3F-20E2-465E-AB71-2C2AA2ADSC5D

Reliability, completeness and improvement

of our knowledge on Germany’s parasitoid wasp fauna —

a case study in Chalcidoidea (Hymenoptera)

Maximilian Weber', Ernst Klimsa’*, Gerd Reder’, Ralph S. Peters**

"4 Zoologisches Forschungsmuseum Alexander Koenig, Arthropoda Department, Adenauerallee 160, D-53113 Bonn, Germany

? Zeisigstrape 2, D-92421 Schwandorf, Germany

°Am Pfortengarten 37, D-67592 Flérsheim-Dalsheim, Germany

“Corresponding author: E-mail: rpeters@leibniz-zfmk.de

'urn:lsid:zoobank.org:author:9413F7B7-B1C4-492D-B293-D89055C45A71

?urn:Isid:zoobank.org:author:2BB938E2-57ED-4611-A43F-D8F481C506D3

3urn:Isid:zoobank.org:author:3 A8EFCFD-DB6D-47C2-A F44-C4B24297C446

4urn:Isid:zoobank.org:author:5C16658B-28A2-4D32-8B5D-1371553DDA18

Abstract. Parasitoid wasps account for a significant proportion of Germany’s insect fauna. Detailed and accurate

knowledge on this fauna is crucial in order to conduct thorough studies in, for example, ecology and conservation, and to

adequately meet the challenges related to the recently reported dramatic biodiversity loss. However, our knowledge on

many species-rich insect groups is error-prone and fragmentary. In this study, we evaluate our knowledge on the German

fauna of Chalcidoidea, one of the most species-rich parasitoid wasps groups, for reliability and completeness. We show

that more than one third of the 1,939 recorded chalcidoid species are known only from a single citation/record, and usually

records are neither detailed nor vouchered. More than two thirds of the species have never been taxonomically revised. We

also found that there are 344 species recorded from neighboring countries of Germany which might also occur in Germa-

ny. In this study, we report the first records from Germany of Calosota aestivalis Curtis, 1836 (Chalcidoidea: Eupelmidae:

Calosotinae) and Zorymus cupreus (Spinola, 1808) (Chalcidoidea: Torymidae: Toryminae). Both records are the result of

close collaboration between citizen scientists and professional entomologists, and both records include detailed collecting

information, life images and voucher specimens. We note that such collaboration is very valuable to increase our knowled-

ge on previously widely neglected taxa. The necessary strategic goal of a well-known German insect fauna, however, will

only be achieved by strongly intensifying the research on biodiversity and taxonomy of all insect taxa.

Keywords. Biodiversity; Chalcidoidea; Germany; parasitoid wasps.

ISSN 2190-7307

http://www.zoologicalbulletin.de

INTRODUCTION

Approximately 33,000 insect species have been listed to

occur in Germany (Volkl et al. 2004), and this number

is frequently used in scientific or popular publications,

comments or surveys. A significant proportion of them

belong to so-called understudied taxa. Taxa in need of

more study are, among others, many groups of Diptera

and Hymenoptera; two insect orders referred to as the

“big four” (together with Lepidoptera and Coleoptera)

because of their exceptional species diversity, even in

comparatively species-poor regions such as the western

Palaearctic (Schumann et al. 1999; Dathe et al. 2001).

There is a long tradition of citizen entomologists study-

ing certain groups of Lepidoptera and Coleoptera often

by geographic region and in astonishing detail, though

this is very unusual in many other insect groups such as

Hymenoptera (excluding Aculeata).

Received: 02.02.2018

Accepted: 16.10.2018

A reliable assessment of insect faunas is becoming in-

creasingly more important in the light of recent findings

that report dramatic losses in insect biomass and poten-

tially also species richness (Hallmann et al. 2017). Yet,

existing species lists for non-aculeate Hymenoptera, for

example, are putatively highly incomplete, error-prone or

outdated (Dathe et al. 2001; Mitroiu et al. 2015). How-

ever, an assessment of Germany’s biodiversity and its

differences and changes over time and space cannot or

should not be done without including these very spe-

cles-diverse groups.

When discussing these understudied taxa and their de-

cided importance for Germany’s biodiversity, two main

issues need to be considered: 1) How severe is the lack of

knowledge, 1.e., how reliable and complete are the pub-

lished species lists? and 2) How can our knowledge on

these groups be improved to meet the strategic goal of a

well-known German biodiversity?

Corresponding editor: X. Mengual

Published: 01.11.2018

102 Maximilian Weber et al.

In this study, we exemplarily screened and evaluated

the currently listed records of chalcidoid wasps (Chal-

cidoidea), one of the very species-rich and notoriously

understudied groups of parasitoid Hymenoptera. To date,

there are 1963 (Schmidt 2015) or 1964 (Noyes 2018)

species of Chalcidoidea checklisted for Germany (i.e.,

approximately 6 % of the German insect fauna). To eval-

uate the reliability and completeness of these records, we

checked the available information in detail (taken from

the well-curated Universal Chalcidoidea Database (Noy-

es 2018; http://www.nhm.ac.uk/our-science/data/chalci-

doids/)).

First, we checked the number of referenced records

from Germany for all species and the respective year of

publication. Few and/or old records might indicate lower

reliability of a given recorded species to actually occur

in Germany. Second, we checked how many recorded

species belong to groups (i.e., genera, subgenera, species

groups etc.) that have been taxonomically revised in the

past, especially in the last 50 years. Taxonomic revision in

groups as delicate as parasitoid wasps (delicate meaning

small-sized, species-rich, subtle species differences) can

result in significant changes of the number of recognized

species (e.g., Hansson & Shevtsova 2012; Khatib et al.

2014). Species records in unrevised taxonomic groups

need to be handled with caution per se. Third, to get an

idea of the number of species that are most likely present,

but have never been formally recorded, we searched for

species that occur in at least two neighboring countries of

Germany. These will most likely also occur in Germany.

In a second part, we show and discuss how knowledge

on Germany’s biodiversity can be improved via close

collaboration between citizen scientists and profession-

als.

Citizen scientists in entomology have excellent know-

ledge about regionally important habitats, they collect,

sort, and mount a lot of specimens, they have profound

knowledge in special groups, but also in general ento-

mology, they may be organized in groups where they reg-

ularly exchange specimens and expertise, and perform

examinations of species biology. On the other hand, pro-

fessional entomologists at museums or other research in-

stitutions have state-of-the-art infrastructure (collections,

molecular laboratories, etc.), taxonomic expertise in

groups that are not too attractive for amateurs, and they

often have the expertise to put faunistic records in a sci-

entific context or to develop new research questions from

observations, often within an international network of re-

searchers. These areas of expertise perfectly complement

each other when it comes to improving knowledge on

biodiversity, in a way that serves both public and science.

The connection and cooperation between citizen scien-

tists and professionals works well in some animal taxa,

and is also inherent part of the German Barcode of Life

initiative (GBOL) (Geiger et al. 2016). However, even

large-scale initiatives such as GBOL cannot cover all of

Bonn zoological Bulletin 67 (2): 101-107

Germany’s biodiversity. For example, most parasitoid

Hymenoptera groups have been largely excluded from

the first two phases of GBOL.

In this context, we add some records of chalcidoid

wasps new to Germany that were found through col-

laboration between citizen scientists and professional

hymenopterists. With our new records we demonstrate

that, if collection by citizen scientists and collaboration

between citizen scientists and professional entomologists

is improved, new chalcidoid species can be easily found

to complement our knowledge on local biodiversity. Ide-

ally, this includes also live pictures, biological data, fine-

scale distribution data and deposition of vouchers in sci-

entific collections, i.e., species records that are far more

valuable than “naked” lists, both in terms of scope and

scientific validity.

Both aspects of this study in concert, however, demon-

strate that collecting, collaboration and taxonomic exper-

tise also have to be significantly expanded and improved

to cover all of Germany’s species diversity in a reason-

able time.

MATERIAL & METHODS

Evaluating the Chalcidoidea species list for Germany

The superfamily Chalcidoidea comprises the following

23 extant families (Heraty et al. 2013; JanSta et al. 2017):

Agaonidae (not occurring in Germany), Aphelinidae,

Azotidae, Chalcididae, Cynipencyrtidae (not occurring

in Germany), Encyrtidae, Eriaporidae (not occurring in

Germany), Eucharitidae, Eulophidae, Eupelmidae, Eu-

rytomidae, Leucospidae, Megastigmidae, Mymaridae,

Ormyridae, Perilampidae, Pteromalidae, Rotoitidae (not

occurring in Germany), Signiphoridae, Tanaostigmati-

dae, Tetracampidae, Torymidae, and Trichogrammatidae.

To evaluate the reliability and completeness of species

records, we examined the information at Universal Chal-

cidoidea Database (UCD, Noyes 2018; http://www.nhm.

ac.uk/our-science/data/chalcidoids/). In this database, all

publications on Chalcidoidea and the data therein are im-

plemented in a timely and highly complete manner by

the enormous effort of John Noyes from Natural History

Museum, London, a Chalcidoidea expert. John Noyes

kindly provided the raw data underlying the published

content at UCD, exported as a .csv file (status as of June

2017). The dataset contained all the published records

of chalcidoid wasps for Germany and its neighboring

countries, 1.e., the Netherlands, Belgium, Luxembourg,

France, Switzerland, Austria, Czech Republic, Poland,

and Denmark. Each record includes the valid species

name, valid genus name, author and country, and the full

reference of the record, including the year of publication.

Initially, all German records were sorted according to

their reference year.

©ZFMK

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna 103

Fig 1. Calosota aestivalis (Eupelmidae: Calosotinae), a species newly recorded from Germany. A. female; B. male; C. female

during oviposition. Live pictures are not from voucher specimen.

Next, we searched for species recorded from Germany

with only one record or reference. Again, these single re-

cords were sorted according to their reference year.

Then, to locate species that have been taxonomical-

ly revised, we searched the references of all recorded

species for the key terms “revision”, “reclassification”,

“synonym”, “new combination”, “review”, “descrip-

tion”, “taxonomy” or any parts of the respective words.

These terms should be included in the titles of at least

the vast majority of taxonomic revision publications and

also cover most of the respective terms in French and

German. “New species” was not considered as species

descriptions not necessarily include a taxonomic revi-

sion. For publications whose titles did not contain exact

information about the revised taxa, the respective ab-

stract was consulted. For all positive matches the cor-

responding taxa (1.e., species, genus, tribe, family) were

marked as “revised” within the main excel sheet and the

date of publication was noted. When a taxonomic unit

was revised (e.g., genus), all subunits (e.g., species) were

listed as revised. Based on the date of publication it was

possible to determine how many of the revised species

were taxonomically revised before and during the last 50

years.

Bonn zoological Bulletin 67 (2): 101-107

Finally, we searched for species that occur in at least

two neighboring countries of Germany that have no com-

mon borderline between them. These will most likely

also occur in Germany. By specifying that the records are

from two or more non-contiguous neighboring countries

of Germany, we intended to exclude species that occur

in geographic regions or habitats that are not necessarily

present in Germany, e.g., eastern Palaearctic species or

alpine species.

New Chalcidoidea records from Germany

Authors have routinely collected species of Hymeno-

ptera in various habitats using hand nets. Live photos

were taken with Canon EOS 5 Mark II, 100mm Macro

(Torymus cupreus (Spinola, 1808)) and Nikon D7200,

60mm Macro (Calosota aestivalis Curtis, 1836). Spec-

imens were killed with ethyl acetate, mounted, labelled,

and identified. Specimen vouchers of the new records are

deposited at Zoologisches Forschungsmuseum Alexan-

der Koenig (ZFMK; Bonn, Germany).

©ZFMK

104 Maximilian Weber et al.

Fig 2. Jorymus cupreus (Torymidae: Toryminae), a species newly recorded from Germany.

Bonn zoological Bulletin 67 (2): 101-107 ©ZFMK

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna 105

Table 1. Records of Chalcidoidea from Germany and their distribution in terms of publication date. Records from neighboring

countries not included.

Year

1700-1799

1800-1899

1900-1950

1951-2000

2001-2017

RESULTS

Evaluating the Chalcidoidea species list for Germany

According to the Universal Chalcidoidea Database

(UCD) there are 1,964 species recorded from Germa-

ny. From this preliminary list, we excluded 29 species

names or entries that were either erroneously listed twice,

very recently synonymized, nomina nuda, or fossil taxa.

There are four additional species recorded from Germany

in the raw data provided by John Noyes, which are not

listed in the “regional list” for Germany tn the UCD web-

site. Consequently, our final list included a total number

of 1,939 chalcidoid species for Germany. For all these

species a total of 5,183 records were listed. Virtually all

records dated from 1951-2017 (98.13 %), and only a few

(1.87 %) were from earlier publications (for details see

Table 1).

Out of the 1,939 species listed from Germany, 764

(39.40 %) were documented with only one reference or

record. The majority of single records (726) dated from

1951 to 2017. Regarding taxonomic revision, 551 species

(28.42 %) belong to taxa that have been taxonomically

revised within the last 50 years. Only 98 species (5.05

%) were revised before 1968, resulting in 649 species

(33.47 %) that have ever been taxonomically revised.

Regarding potential new species records for Germany,

344 species were listed in at least two neighboring coun-

tries that do not have a borderline between them. These

species represent 11.3 % of the total of 3,043 species re-

corded from the neighboring countries. For these species

an occurrence in Germany can be assumed to be proba-

ble. In summary, a total of 2,283 chalcidoid species have

been recorded from Germany or can be expected to occur

in Germany.

A full list of the species recorded from Germany plus

those species recorded from neighboring countries which

do not have a borderline between them, along with spe-

cies-specific information as included in this study, 1s giv-

en in Appendix I.

Bonn zoological Bulletin 67 (2): 101-107

Proportion of records

(Total = 100 %)

Number of records

(Total = 5,183)

51.13 % 2,650

47.00 % 2,436

New Chalcidoidea records from Germany

We report for the first time for Germany the occurrence

of Calosota aestivalis Curtis, 1836 (Chalcidoidea: Eu-

pelmidae: Calosotinae) (Fig. 1) and Jorymus cupreus

(Spinola, 1808) (Chalcidoidea: Torymidae: Toryminae)

(Fig.2):

Calosota aestivalis Curtis, 1836

Calosota aestivalis Curtis, 1836: 596.

For synonyms, distribution and associates see Noyes

(2018).

Material examined. Germany: Bavaria, Landkreis

Schwandorf, Schwandorf (GMS: N 49° 18' 15.347"

W 12° 7' 53.025", DG: N 49.304263 W 12.131396),

29.iv.2015, south-facing woodpile next to small wood-

land, leg. E. Klimsa; det. E. Klimsa, G. Gibson (1 female,

deposited at ZFMK, ZFMK-HYM-00012133).

Remarks. Specimens of this species were also observed

on several occasions in the years 2016 and 2017 at wood-

piles in Schwandorf and Kallmtinz (Bavaria).

Torymus cupreus (Spinola, 1808)

Diplolepis cuprea Spinola, 1808: 212-213.

For synonyms, distribution and associates see Noyes

(2018).

Material examined. Germany: Rheinland-Pfalz, Don-

nersbergkreis, Zell (GMS: N 49° 38' 58.87" O 8°

8'5.031", DG: 49.649686 8.134731), tvy-covered hedge-

row at the village outskirts, 01.x.2015, leg. G. Reder; det.

G. Reder, vid. R. Peters (1 female, deposited at ZFMK,

ZFMK-HYM-00020525).

DISCUSSION

Results show that the majority of published records of

chalcidoid species from Germany date from the period

after the Second World War, which may indicate an in-

©ZFMK

106 Maximilian Weber et al.

crement of collecting, identification and publication ac-

tivities during that time. For approximately one third of

the recorded species only a single record is listed. Any

single event is sought to be verified or falsified to be sci-

entifically sound. We have no information on whether

these single records are correct or not; in fact, they could

well be correct, and on the other hand, also multiple re-

cords can be wrong. Ideally, we would have checked

if records are vouchered in a scientific collection and

could be verified accordingly. Our expectation was that

the number of voucher-referenced records was very low,

rendering most records not necessarily wrong but scien-

tifically flawed. A thorough check for vouchers was not

done, due to time constraints. However, when checking

this for 20 randomly chosen species, we found almost no

detailed collecting information or references to deposited

vouchers. Almost half of the records (47 %, see Table 1)

date from rather recently, which indicates that the study

of Germany’s chalcidoid fauna has accelerated, and that

the quality of the records should increase with more and

more recent records added.

The fact that only 33.5 % of the recorded species from

Germany have been taxonomically revised (based on

our search for key terms in the publication titles) might

severely flaw the reliability of the known records. Tax-

onomic revisions in chalcidoid wasps usually result in

significant additions or subtractions of recognized val-

id species. Recent examples from European chalcidoid

taxa include the Eupelmus urozonus group (Eupelmidae)

(now 21 valid species, but only nine species before revi-

sion; Khatib et al. 2014), Dibrachys cavus group (Ptero-

malidae) (now three valid species, but five species be-

fore Peters & Baur (2011) and 12 species before Gahan

(1938)), the genus Omphale (Eulophidae) (now 37 val-

id species, but 31 species before Hansson & Shevtsova

2012), and the genus Copidosoma (Encyrtidae) (now 58

valid species, but 84 species before Guerrier1 & Noyes

2005). Furthermore, Pteromalus Swederus, 1795, one of

the most species-rich genera of Chalcidoidea with 271

species listed for Germany, has never been revised, but

currently ongoing studies suggest a significant impact of

revision on the number of valid species for this genus

(unpublished). We need to stress that taxonomic work, in

terms of revisionary work for taxonomic clarity, is cru-

cial for any accurate species list and the basis for any

further application of these lists in ecology, conservation,

or biogeography. The low percentage of taxonomically

revised taxa points towards two major problems in our

knowledge of German Chalcidoidea: 1) the lists could

well be grossly wrong in terms of true species, and 2) the

number of scientists working on the taxonomy of chal-

cidoid (or any parasitoid) wasps is too low. There are

very few specialists working on these groups. Actually,

chalcidoid wasps range among the most studied and sur-

veyed parasitoid wasp taxa in Germany, with a handful of

specialists working on them; so, the situation is worse or

Bonn zoological Bulletin 67 (2): 101-107

much worse in other diverse Hymenoptera groups. The

result that only 33.5 % of the taxa have been taxonomi-

cally revised means that we have no clue about the taxo-

nomic status of two-thirds of the chalcidoid wasp species

in our own local fauna. Any study using the current list

will be substantially flawed.

The number of species from neighboring countries 1s

lower than we originally anticipated, but still, another

344 chalcidoid species are likely to be present in Germa-

ny (an increment of almost 18 % from the total). Most

of these species are recorded from the Czech Republic,

France and the Netherlands. This east-west axis includes

the major part of the neighboring species that probably

also occur in Germany. The examples of France and the

Czech Republic show the positive impact on the species

inventory by the work of taxonomic experts such as J.-Y.

Rasplus and G. Delvare in France, and Z. Bouéek in the

Czech Republic.

If one looks into the chalcidoid fauna in more detail,

new records for Germany can be found rather easily.

We complemented our manuscript with two new spe-

cies records from Germany for species already recorded

from neighboring countries, collected by citizen scien-

tists Gerd Reder and Ernst Klimsa. Both are active hy-

menopterists that frequently collect and photograph hy-

menopterans mainly in their local vicinity. Both authors

approached the senior author (RSP) of this study with

some photos and some questions on the identity of the

depicted species and sent the specimens to the ZFMK for

further examination. In collaboration with Gary Gibson,

a well-renowned expert on Chalcidoidea at the Canadian

National Collection (CNC, Ottawa, Canada), the speci-

mens were identified. Both new records included exact

collecting data as well as live pictures. This additional

information is very rare and highly welcome. For many

parasitoid species only very sparse additional informa-

tion is recorded and often none at all. Of course, infor-

mation on host association is most valuable. This infor-

mation is also missing or fragmentary for many species.

As an example, we checked the records of 200 randomly

chosen species from Germany and found no or only one

reference on host association for 17 % of the species.

The two new German records presented here illustrate

how rapid, easy and helpful joint efforts of citizen and

professional entomologists can be, even in groups such

as parasitoid Hymenoptera. Of course, many parasitoid

groups are even more problematic than Chalcidoidea and

harbor problems such as the need of taxonomic work,

minute size and no available experts that will make 1m-

provement of knowledge of the local fauna exceedingly

more difficult.

To provide reliable and complete species lists is only

part of solving the problem of parasitoid wasps being

widely neglected in science, politics and public. The

other part, although tightly linked to the production of

reliable species lists, is more taxonomic work and revi-

©ZFMK

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna 107

sions, the increased use of modern molecular and mor-

phological tools in taxonomy and species identification,

the publication of easy-to-use identification keys, and

the education and funding of taxonomic experts. We call

for future intensive collections of Germany’s chalcidoid

(and parasitoid) fauna, and for intensive collaboration

of citizen and professional entomologists as well as for

large-scale taxonomy and monitoring projects to over-

come the unbearable situation that we have no or only

meagre knowledge on a significant portion of Germany’s

biodiversity. The latest results showing that insect popu-

lations decrease at a rapid speed (Hallmann et al. 2017)

urge for fast and sweeping assessments of biodiversity in

order to drastically improve our understanding of local

fauna and to guard against potential loss of Germany’s

biological heritage.

Acknowledgements. We thank John Noyes (London) for giv-

ing access to raw data of the Universal Chalcidoidea Database

and for kind assistance with evaluating the data. Trevor Burt

(Bonn) is acknowledged for linguistic corrections and valuable

suggestions that helped to improve the manuscript.

REFERENCES

Al Khatib F, Fusu L, Cruaud A, Gibson G, Borowiec N, Rasplus

J-Y, Ris N, Delvare G (2014) An integrative approach to

Species discrimination in the Eupelmus urozonus complex

(Hymenoptera, Eupelmidae), with the description of 11 new

species from the Western Palaearctic. Systematic Entomolo-

gy 39: 806-862

Dathe HH, Taeger A, Blank SM (eds) (2001) Verzeichnis der

Hautfligler Deutschlands (Entomofauna Germanica 4).

Entomologische Nachrichten und Berichte (Dresden), Bei-

heft 7: 1-175

Gahan AB (1938) Notes on some genera and species of Chal-

cidoidea (Hymenoptera). Proceedings of the Entomological

Society of Washington 40: 209-227

Geiger MF, Astrin JJ, Borsch T, Burkhardt U, Grobe P, Hand R,

Hausmann A, Hohberg K, Krogmann L, Lutz M, Monje C,

Misof B, Moriniere J, Miller K, Pietsch S, Quandt D, Rulik

B, Scholler M, Traunspurger W, Haszprunar G, Wagele W

(2016) How to tackle the molecular species inventory for an

industrialized nation-lessons from the first phase of the Ger-

man Barcode of Life initiative GBOL (2012-2015). Genome

59: 661-670

Guerrieri E, Noyes J (2005) Revision of the European species

of Copidosoma Ratzeburg (Hymenoptera: Encyrtidae), para-

sitoids of caterpillars (Lepidoptera). Systematic Entomology

30: 97-174

Bonn zoological Bulletin 67 (2): 101-107

Hallmann CA, Sorg M, Jongejans E, Siepel H, Hofland N,

Schwan H, Stenmans W, Miller A, Sumser H, Horren T,

Goulson D, de Kroon H (2017) More than 75 percent decline

over 27 years in total flying insect biomass in protected areas.

PLoS ONE 12(10): e0185809

Hansson C, Shevtsova E (2012) Revision of the European spe-

cies of Omphale Haliday (Hymenoptera, Chalcidoidea, Eulo-

phidae). ZooKeys 232: 1-157

Heraty JM, Burks RA, Cruaud A, Gibson GAP, Liljeblad J,

Munro J, Rasplus JY, Delvare G, JanSta P, Gumovsky A,

Huber JT, Woolley JB, Krogmann L, Heydon SL, Polaszek

A, Schmidt S, Darling DC, Gates MW, Mottern J, Murray

E, Molin AD, Triapitsyn S, Baur H, Pinto JD, van Noort S,

George J, Yoder MJ (2013) A phylogenetic analysis of the

megadiverse Chalcidoidea (Hymenoptera). Cladistics 29:

466-542

Jansta P, Cruaud A, Delvare G, Genson G, Heraty J, Krizkova

B, Rasplus JY (2017) Torymidae (Hymenoptera, Chalci-

doidea) revised: molecular phylogeny, circumscription and

reclassification of the family with discussion of its biogeog-

raphy and evolution of life-history traits. Cladistics: 1-25

Mitroiu M, Noyes J, Cetkovic A, Nonveiller G, Radchenko A,

Polaszek A, Ronquist F, Forshage M, Pagliano G, Gusenleit-

ner J, Bartalucci M, Olmi M, Fusu L, Madl M, Johnson N,

Jansta P, Wahis R, Soon V, Rosa P, Osten T, Barbier Y, de

Jong Y (2015). Fauna Europaea: Hymenoptera — Apocrita

(excl. Ichneumonoidea). Biodiversity Data Journal 3: e4186

Noyes JS (2018) Universal Chalcidoidea Database. World-

WideWeb electronic publication. http://www.nhm.ac.uk/en-

tomology/chalcidoids/index.html

Peters RS, Baur H (2011) A revision of the Dibrachys cavus

species complex (Hymenoptera: Chalcidoidea: Pteromali-

dae). Zootaxa 2937: 1-30

Schmidt S (2015) Checklist of German Chalcidoidea (Hyme-

noptera). WorldWideWeb electronic publication. http://www.

zsm.mwn.de/docs_zsm/htdocs/hym/chal/e/overview.html

Schumann H, Bahrmann R, Stark A. (eds) (1999) Checkliste

der Dipteren Deutschlands. Studia dipterologica Supplement

2: 1-354

Volkl W, Blick T, Kornacker PM, Martens H (2004) Quantitati-

ver Uberblick tiber die rezente Fauna von Deutschland. Natur

und Landschaft 79: 293-295

APPENDIX I

(electronic supplement, available at www.bonnzoologicalbulletin.de)

List of Chalcidoidea species recorded from Germany, based on

the Universal Chalcidoidea Database, with some corrections of

the original data and with species-specific data on the record

and the species’ taxonomy. In red species recorded from Ger-

many; in green species recorded from neighboring countries of

Germany that do not have a border between them.

©ZFMK

BHL

i

Blank Page Digitally Inserted

APPENDIX 1

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna

(APPENDIX 1)

Supplementary material. List of Chalcidoidea species recorded from Germany, based on the Universal Chalcidoidea Database,

with some corrections of the original data and with species-specific data on the record and the species’ taxonomy. In red species

recorded from Germany; in green species recorded from neighboring countries of Germany that do not have a border between them.

family

Aphelinidae

species

Aphelinus abdominalis

Aphelinus annulipes

Aphelinus argiope

Aphelinus asychis

Aphelinus chaonia

Aphelinus circumscriptus

Aphelinus flavus

Aphelinus fusciscapus

Aphelinus longipennis

Aphelinus mali

Aphelinus notatus

Aphelinus semiflavus

Aphelinus varipes

Aphytis chilensis

Aphytis chrysomphali

Aphytis luteus

Aphytis mytilaspidis

Aphytis proclia

Centrodora acridiphagus

Centrodora amoena

Centrodora speciosissima

Centrodora tibialis

Coccobius annulicornis

Coccophagus gossypariae

Coccophagus insidiator

Coccophagus lycimnia

Coccophagus palaeolecanii

Coccophagus pulchellus

Coccophagus scutatus

Coccophagus scutellaris

Coccophagus semicircularis

Diaspiniphagus moeris

Encarsia aleurochitonis

Encarsia aurantii

Encarsia berlesei

Bonn zoological Bulletin 67 (2): 101-107

author

(Dalman)

(Walker)

Walker

(Ratzeburg)

(Nees)

(Forster)

(Haldeman)

(Ratzeburg)

Howard

(Forster)

Howard

(Mercet)

(Ratzeburg)

(Le Baron)

(Walker)

(Otten)

Forster

(Girault)

(Nees)

Ratzeburg

Gahan

(Dalman)

(Walker)

Yasnosh

Westwood

Howard

(Dalman)

(Forster)

(Walker)

(Mercet)

(Howard)

distribution

Germany

revision

(year)

2014

Oe

2014

2010

single record

(year)

2001

2003

1997

2001

©ZFMK

family

Aphelinidae

Azotidae

Chalcididae

Encyrtidae

species

Encarsia citrina

Encarsia fasciata

Encarsia formosa

Encarsia inaron

Encarsia intermedia

Encarsia leucaspidis

Encarsia margaritiventris

Encarsia perniciosi

Encarsia tricolor

Eretmocerus californicus

Eretmocerus mundus

Marietta picta

Pteroptrix bicolor

Pteroptrix longiclava

Ablerus atomon

Ablerus pinifoliae

Belaspidia obscura

Brachymeria femorata

Brachymeria minuta

Brachymeria moerens

Brachymeria obtusata

Brachymeria parvula

Brachymeria podagrica

Brachymeria rugulosa

Brachymeria secundaria

Brachymeria tibialis

Brachymeria vitripennis

Chalcis biguttata

Chalcis femorata

Chalcis myrifex

Chalcis ramicornis

Chalcis sispes

Conura xanthostigma

Euchalcis magna

Haltichella rufipes

Hybothorax graffii

Neohybothorax hetera

Psilochalcis benoisti

Psilochalcis subarmata

Acerophagus austriacus

Acerophagus malinus

Adelencyrtus aulacaspidis

Bonn zoological Bulletin 67 (2): 101-107

Maximilian Weber et al.

author

(Craw)

(Malenotti)

Gahan

(Walker)

(Ferriere)

(Mercet)

(Tower)

Forster

Howard

Mercet

(André)

(Howard)

(Girault)

(Walker)

(Mercet)

Masi

(Panzer)

(Linnaeus)

(Ruschka)

(Forster)

(Walker)

(Fabricius)

(Forster)

(Ruschka)

(Walker)

(Forster)

Spinola

Nees

(Sulzer)

Gravenhorst

(Linnaeus)

(Dalman)

Boucéek

(Olivier)

Ratzeburg

(Walker)

(Steffan)

(Forster)

(Mercet)

(Gahan)

(Brethes)

distribution

Germany

revision

(year)

1999

1952

L9SZ

lars

Keaz,

1952

NOSZ

ey,

line oy

1952

2017

1952

ey,

1952

LOSZ

1952

single record

(year)

2001

1994

2001

©ZFMK

family

Encyrtidae

species

Adelencyrtus intersectus

Ageniaspis atricollis

Ageniaspis fuscicollis

Ageniaspis testaceipes

Aglyptus rufus

Anagyrietta pantherina

Anagyrus belibus

Anagyrus matritensis

Anagyrus schmuttereri

Anagyrus schoenherri

Anagyrus securicornis

Anomalicornia tenuicornis

Anthemus pini

Anusia nasicornis

Aphycoides clavellatus

Aphycoides tenuis

Aphycus apicalis

Aphycus hederaceus

Aphycus sumavicus

Baeocharis pascuorum

Blastothrix brittanica

Blastothrix erythrostetha

Blastothrix hungarica

Blastothrix ilicicola

Blastothrix longipennis

Blastothrix sericea

Blastothrix truncatipennis

Bothriothorax altensteinii

Bothriothorax clavicornis

Bothriothorax intermedius

Bothriothorax paradoxus

Bothriothorax wichmani

Boucekiella depressa

Cerapterocerus celadus

Cerapterocerus mirabilis

Cerchysius subplanus

Cercobelus jugaeus

Charitopus fulviventris

Cheiloneurus claviger

Cheiloneurus elegans

Cheiloneurus glaphyra

Bonn zoological Bulletin 67 (2): 101—107

author

(Fonscolombe)

(Dalman)

(Ratzeburg)

(Dalman)

Ferriere

(Walker)

(Mercet)

Ferriere

(Westwood)

Domenichini

Mercet

Ferriere

Forster

(Dalman)

(Ratzeburg)

(Dalman)

(Westwood)

Hoffer

Mayr

Girault

(Walker)

Erdés

Mercet

Howard

(Dalman)

(Ferriére)

Ratzeburg

(Dalman)

Claridge

(Dalman)

Ferriere

Hoffer

(Walker)

Westwood

(Dalman)

(Walker)

Forster

Thomson

(Dalman)

(Walker)

distribution

Germany

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna

(APPENDIX 1)

revision

(year)

1916

1964

single record

(year)

2003

1999

2003

1999

2001

©ZFMK

4 Maximilian Weber et al.

revision single record

family species author distribution (year) (year)

Encyrtidae Cheiloneurus paralia (Walker) Germany

Encyrtidae Choreia inepta (Dalman) Germany

Encyrtidae Coccidencyrtus phenacocci Ferriére Germany

Encyrtidae Coelopencyrtus arenarius (Erdos) Germany

Encyrtidae Copidosoma agrotis (Fonscolombe) Germany 2013

Encyrtidae Copidosoma aithyia (Walker) Germany 2013

Encyrtidae Copidosoma albipes (Westwood) Germany 2013

Encyrtidae Copidosoma aretas (Walker) Germany 2013

Encyrtidae Copidosoma boucheanum Ratzeburg Germany 2013

Encyrtidae Copidosoma cervius (Walker) Germany 2013 2005

Encyrtidae Copidosoma chalconotum (Dalman) Germany 2013

Encyrtidae Copidosoma cuproviride Springate & Noyes Germany 2013 2005

Encyrtidae Copidosoma dius (Walker) Germany DONS 2001

Encyrtidae Copidosoma filicorne (Dalman) Germany 2013

Encyrtidae Copidosoma flagellare (Dalman) Germany 2013

Encyrtidae Copidosoma floridanum (Ashmead) Germany 2013

Encyrtidae Copidosoma fuscisquama (Thomson) Germany 2013 2005

Encyrtidae Copidosoma genale (Thomson) Germany 2013

Encyrtidae Copidosoma iracundum Erdos Germany 2013 2005

Encyrtidae Copidosoma peticus (Walker) Germany 2013

Encyrtidae Copidosoma serricorne (Dalman) Germany 2013

Encyrtidae Copidosoma sosares (Walker) Germany 2013

Encyrtidae Copidosoma terebrator Mayr Germany 2013

Encyrtidae Copidosoma thebe (Walker) Germany ZOTS

Encyrtidae Copidosoma truncatellum (Dalman) Germany 2013

Encyrtidae Copidosoma varicorne (Nees) Germany 2013

Encyrtidae Dinocarsis hemiptera (Dalman) Germany 1966 2001

Encyrtidae Dinocarsis hofferi Graham Germany

Encyrtidae Discodes aeneus (Dalman) Germany

Encyrtidae Discodes coccophagus (Ratzeburg) Germany

Encyrtidae Discodes differens Yasnosh Germany

Encyrtidae Discodes encopiformis (Walker) Germany

Encyrtidae Echthroplexis puncticollis (Thomson) Germany

Encyrtidae Ectroma fulvescens Westwood Germany 2007

Encyrtidae Ectroma reinhardi (Mayr) Germany 2007

Encyrtidae Encyrtus albitarsis Zetterstedt Germany

Encyrtidae Encyrtus aurantii (Geoffroy) Germany 1997

Encyrtidae Encyrtus flavipes Nees Germany 2001

Encyrtidae Encyrtus foersteri Mayr Germany 2001

Encyrtidae Encyrtus infelix (Embleton) Germany 1989

Encyrtidae Encyrtus infidus (Rossi) Germany

Bonn zoological Bulletin 67 (2): 101-107 ©ZFMK

family

Encyrtidae

species

Encyrtus mucronatus

Encyrtus swederi

Ericydnus apterogenes

Ericydnus baleus

Ericydnus longicornis

Ericydnus robustior

Ericydnus sipylus

Ericydnus strigosus

Ericydnus theron

Ericydnus ventralis

Eucoccidophagus semiluniger

Eugahania fumipennis

Eupoecilopoda perpunctata

Eusemion cornigerum

Eusemion longipennis

Habrolepis dalmanni

Helegonatopus dimorphus

Heterococcidoxenus schlech-

tendali

Homalotylus ephippium

Homalotylus eytelweinii

Homalotylus flaminius

Homalotylus hemipterinus

Homalotylus platynaspidis

Isodromus vinulus

Ixodiphagus hookeri

Lamennaisia ambigua

Lamennaisia nobilis

Leptomastidea bifasciata

Leptomastix epona

Leptomastix histrio

Leptomastix mayri

Mahencyrtus comara

Mayrencyrtus imandes

Mayridia merceti

Mayridia myrlea

Mayridia procera

Metaphycus asterolecanii

Metaphycus chermis

Metaphycus delos

Metaphycus insidiosus

Metaphycus maculipennis

Bonn zoological Bulletin 67 (2): 101—107

author

Ratzeburg

Dalman

Mayr

(Walker)

(Dalman)

Mercet

(Walker)

(Nees)

Trjapitzin

(Dalman)

(Hoffer)

(Ratzeburg)

(Masi)

(Walker)

(Ashmead)

(Westwood)

(Hoffer)

(Mayr)

(Ruschka)

(Ratzeburg)

(Dalman)

(De Stefani)

Hoffer

(Dalman)

(Howard)

(Nees)

(Mayr)

(Walker)

(Forster)

Ozdikmen

(Walker)

Trjapitzin

(Walker)

(Mercet)

(Fonscolombe)

Guerrieri & Noyes

(Mercet)

(Timberlake)

distribution

Germany

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna

(APPENDIX 1)

revision

(year)

199]

Lot

1991

LOD k

1991

Meee

1991

LOOT

1993

1969

2000

single record

(year)

2001

2003

2001

2010

2006

2001

1972

2001

2003

©ZFMK

6 Maximilian Weber et al.

revision single record

family species author distribution (year) (year)

Encyrtidae Metaphycus melanostomatus (Timberlake) Germany 2000

Encyrtidae Metaphycus nitens (Kurdjumov) Germany 2000

Encyrtidae Metaphycus punctipes (Dalman) Germany 2000

Encyrtidae Metaphycus silvestrii Sugonjaev Germany 2000

Encyrtidae Metaphycus stagnarum Hoffer Germany 2000 2003

Encyrtidae Metaphycus unicolor Hoffer Germany 2000 2003

Encyrtidae Metaphycus zebratus (Mercet) Germany 2000

Encyrtidae Microterys aeneiventris (Walker) Germany 2011

Encyrtidae Microterys apicipennis Bakkendorf Germany 2011 2001

Encyrtidae Microterys chalcostomus (Dalman) Germany 2011

Encyrtidae Microterys cyanocephalus (Dalman) Germany 2011

Encyrtidae Microterys duplicatus (Nees) Germany 2011

Encyrtidae Microterys ferrugineus (Nees) Germany 2011

Encyrtidae Microterys fuscipennis (Dalman) Germany 2011

Encyrtidae Microterys hortulanus Erdos Germany 2011

Encyrtidae Microterys lunatus (Dalman) Germany 2011

Encyrtidae Microterys masii Silvestri Germany 2011

Encyrtidae Microterys nietneri (Motschulsky) Germany 2011

Encyrtidae Microterys sceptriger (Forster) Germany 2011

Encyrtidae Microterys sylvius (Dalman) Germany 2011

Encyrtidae Microterys tessellatus (Dalman) Germany 2011

Encyrtidae Microterys trjapitzini Yasnosh Germany 2011

Encyrtidae Microterys zarina (Walker) Germany 2011

Encyrtidae Mira mucora Schellenberg Germany 1977

Encyrtidae Oobius zahaikevitshi Trjapitzin Germany

Encyrtidae Ooencyrtus fulvipes Hoffer Germany

Encyrtidae Ooencyrtus gravis (Nees) Germany

Encyrtidae Ooencyrtus kuvanae (Howard) Germany

Encyrtidae Ooencyrtus tardus (Ratzeburg) Germany

Encyrtidae Ooencyrtus telenomicida (Vassiliev) Germany

Encyrtidae Ooencyrtus vinulae (Masi) Germany

Encyrtidae Parablastothrix metatibialis Erdos Germany

Encyrtidae Parablastothrix montana Erdos Germany

Encyrtidae Parablastothrix plugarui Trjapitzin Germany

Encyrtidae Parablatticida brevicornis (Dalman) Germany

Encyrtidae Platencyrtus parkeri Ferriére Germany 2003

Encyrtidae Prionomastix morio (Dalman) Germany

Encyrtidae Prionomitus mitratus (Dalman) Germany 1981

Encyrtidae Prionomitus tiliaris (Dalman) Germany

Encyrtidae Pseudencyrtus eupelmoides (Ratzeburg) Germany

Encyrtidae Pseudencyrtus idmon (Walker) Germany 2007

Bonn zoological Bulletin 67 (2): 101-107 ©ZFMK

family

Encyrtidae

Eucharitidae

species

Pseudencyrtus misellus

Pseudencyrtus salicisstrobili

Pseudleptomastix brevipennis

Pseudorhopus testaceus

Psilophrys tenuicornis

Rhopus brachypterus

Rhopus parvulus

Rhopus piso

Rhopus semiapterus

Rhopus sulphureus

Sectiliclava cleone

Subprionomitus festucae

Syrphophagus aeruginosus

Syrphophagus aphidivorus

Syrphophagus herbidus

Syrphophagus hyalipennis

Syrphophagus mamitus

Syrphophagus pertiades

Syrphophagus taeniatus

Tachinaephagus zealandicus

Tetracnemoidea piceae

Tetracnemoidea spilococci

Tetracnemus diversicornis

Tetracnemus heydeni

Thomsonisca amathus

Trechnites fuscitarsis

Trechnites insidiosus

Trichomasthus albimanus

Trichomasthus cyaneus

Trichomasthus cyanifrons

Trichomasthus dignus

Trichomasthus frontalis

Trichomasthus marsus

Tyndarichus melanacis

Tyndarichus navae

Tyndarichus scaurus

Zaomma eriococci

Zaomma hirsuta

Zaomma lambinus

Eucharis adscendens

Stilbula cyniformis

Bonn zoological Bulletin 67 (2): 101-107

(APPENDIX 1)

author

(Dalman)

(Linnaeus)

(Ferriere)

(Ratzeburg)

Graham

(Mercet)

(Walker)

(Mercet)

(Westwood)

(Walker)

(Mayr)

(Dalman)

(Mayr)

(Dalman)

(Mayr)

(Walker)

(Forster)

Ashmead

(Erdos)

Ferriere

Westwood

(Mayr)

(Walker)

(Thomson)

(Crawford)

Thomson

(Dalman)

Khlopunov

Alam

(Walker)

(Dalman)

Howard

(Walker)

(Ferriere)

(Ratzeburg)

(Walker)

(Fabricius)

(Rossi)

distribution

Germany

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna

revision

(year)

1970

2012

2009

2002

single record

(year)

2001

2006

2014

2001

2009

2003

1999

2001

©ZFMK

family

Eulophidae

species

Aceratoneuromyia granularis

Achrysocharoides acerianus

Achrysocharoides atys

Achrysocharoides carpini

Achrysocharoides cilla

Achrysocharoides cruentus

Achrysocharoides insignitellae

Achrysocharoides latreillii

Achrysocharoides nigricoxae

Achrysocharoides niveipes

Achrysocharoides robiniae

Achrysocharoides splendens

Achrysocharoides suprafolius

Achrysocharoides usticrus

Achrysocharoides zwoelferi

Allocerastichus doderi

Anaprostocetus acuminatus

Aprostocetus aethiops

Aprostocetus annulatus

Aprostocetus apama

Aprostocetus aquaticus

Aprostocetus arenarius

Aprostocetus aristaeus

Aprostocetus artemisicola

Aprostocetus boreus

Aprostocetus brachycerus

Aprostocetus bruzzonis

Aprostocetus calamarius

Aprostocetus caudatus

Aprostocetus cecidomyiarum

Aprostocetus ciliatus

Aprostocetus citrinus

Aprostocetus citripes

Aprostocetus clavicornis

Aprostocetus collega

Aprostocetus crino

Aprostocetus diversus

Aprostocetus elongatus

Aprostocetus emesa

Aprostocetus epicharmus

Aprostocetus eriophyes

Bonn zoological Bulletin 67 (2): 101-107

Maximilian Weber et al.

author

Domenichini

(Askew)

(Walker)

Bryan

(Walker)

Hansson

(Erdos)

(Curtis)

(Delucchi)

(Thomson)

Hansson & Shevtsova

(Delucchi)

(Askew)

(Erdos)

(Delucchi)

Masi

(Ratzeburg)

(Zetterstedt)

(Forster)

(Walker)

(Erdos)

(Walker)

Graham

(Delucchi)

(Thomson)

(Masi)

Graham

Westwood

(Bouché)

(Nees)

(Forster)

(Thomson)

(Zetterstedt)

(Ratzeburg)

(Walker)

(Forster)

(Walker)

(Taylor)

distribution

Germany

single record

(year)

2001

©ZFMK

family

Eulophidae

species

Aprostocetus escherichi

Aprostocetus eupatorii

Aprostocetus flavovarius

Aprostocetus fulvipes

Aprostocetus gaus

Aprostocetus gratus

Aprostocetus hagenowii

Aprostocetus leptoneuros

Aprostocetus leucone

Aprostocetus longicauda

Aprostocetus longiscapus

Aprostocetus luteus

Aprostocetus lycidas

Aprostocetus lysippe

Aprostocetus mandanis

Aprostocetus metra

Aprostocetus microscopicus

Aprostocetus minimus

Aprostocetus mycerinus

Aprostocetus neglectus

Aprostocetus novatus

Aprostocetus orithyia

Aprostocetus ovivorax

Aprostocetus pachyneuros

Aprostocetus pallipes

Aprostocetus pausiris

Aprostocetus percaudatus

Aprostocetus phineus

Aprostocetus phragmiticola

Aprostocetus pseudopodiellus

Aprostocetus ptarmicae

Aprostocetus pygmaeus

Aprostocetus roesellae

Aprostocetus rubi

Aprostocetus rubicola

Aprostocetus rumicis

Aprostocetus salictorum

Aprostocetus serratularum

Aprostocetus strobilanae

Aprostocetus subplanus

Aprostocetus tanaceticola

Bonn zoological Bulletin 67 (2): 101-107

author

(Szelény1)

Kurdjumov

(Nees)

(Forster)

(Walker)

(Giraud)

(Ratzeburg)

(Walker)

(Thomson)

(Ratzeburg)

(Walker)

(Rondani)

(Ratzeburg)

(Walker)

(Domenichini)

(Walker)

(Silvestri)

(Ratzeburg)

(Dalman)

(Walker)

(Silvestri)

(Walker)

Graham

(Bakkendorf)

Graham

(Zetterstedt)

(Nees)

Graham

(Ratzeburg)

Graham

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna

(APPENDIX 1)

distribution

Germany

single record

(year)

2001

©ZFMK

10

family

Eulophidae

species

Aprostocetus terebrans

Aprostocetus trjapitzini

Aprostocetus vassolensis

Aprostocetus venustus

Aprostocetus veronicae

Aprostocetus xanthopus

Aprostocetus zosimus

Asecodes congruens

Asecodes erxias

Asecodes lagus

Asecodes lucens

Asecodes turcicum

Astichus arithmeticus

Astichus maculatus

Astichus solutus

Aulogymnus aceris

Aulogymnus arsames

Aulogymnus euedoreschus

Aulogymnus fumatus

Aulogymnus gallarum

Aulogymnus obscuripes

Aulogymnus skianeuros

Baryscapus adalia

Baryscapus agrilorum

Baryscapus anasillus

Baryscapus bonessi

Baryscapus bruchophagi

Baryscapus conwentziae

Baryscapus crassicornis

Baryscapus daira

Baryscapus diaphantus

Baryscapus endemus

Baryscapus evonymellae

Baryscapus galactopus

Baryscapus garganus

Baryscapus gradwelli

Baryscapus impeditus

Baryscapus lotellae

Baryscapus nigroviolaceus

Baryscapus oophagus

Baryscapus pallidae

Bonn zoological Bulletin 67 (2): 101-107

Maximilian Weber et al.

author

Erdos

(Kostjukov)

Graham

(Gahan)

Graham

(Nees)

(Walker)

(Nees)

(Walker)

(Nees)

(Forster)

Hedqvist

Forster

(Walker)

(Ratzeburg)

(Linnaeus)

(Mayr)

(Ratzeburg)

(Walker)

(Ratzeburg)

Graham

Askew

(Gahan)

(Ferriere)

(Erdos)

(Walker)

(Bouché)

(Ratzeburg)

(Domenichini)

Graham

(Nees)

(Delucchi)

(Nees)

(Otten)

Graham

distribution

Germany

single record

(year)

2001

2010

2001

2013

2001

2013

©ZFMK

family

Eulophidae

species

Baryscapus servadeii

Baryscapus spartifoliellae

Baryscapus sugonjaevi

Baryscapus turionum

Ceranisus menes

Ceranisus pacuvius

Chaenotetrastichus grangeri

Chaenotetrastichus semiflavus

Chrysocharis acoris

Chrysocharis acutigaster

Chrysocharis amasis

Chrysocharis amyite

Chrysocharis antoni

Chrysocharis assis

Chrysocharis avia

Chrysocharis budensis

Chrysocharis chlorus

Chrysocharis clarkae

Chrysocharis crassiscapus

Chrysocharis elongata

Chrysocharis entedonoides

Chrysocharis equiseti

Chrysocharis eurynota

Chrysocharis foliincolarum

Chrysocharis gemma

Chrysocharis idyia

Chrysocharis illustris

Chrysocharis laomedon

Chrysocharis laricinellae

Chrysocharis liriomyzae

Chrysocharis mediana

Chrysocharis nautius

Chrysocharis nephereus

Chrysocharis nigricrus

Chrysocharis nitetis

Chrysocharis nitidifrons

Chrysocharis orbicularis

Chrysocharis pallipes

Chrysocharis pentheus

Chrysocharis phryne

Chrysocharis pilosa

Bonn zoological Bulletin 67 (2): 101-107

author

(Domenichini)

Graham

(Kostjukov)

(Hartig)

(Walker)

(Erdos)

(Girault)

(Walker)

Hansson

(Walker)

Hansson

(Walker)

Hansson

Erdos

Graham

Yoshimoto

(Thomson)

(Walker)

Hansson

Graham

(Christ)

(Walker)

Graham

(Walker)

(Ratzeburg)

Delucchi

Forster

(Walker)

(Thomson)

(Walker)

Graham

(Nees)

(Walker)

Delucchi

distribution

Germany

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna

(APPENDIX 1)

revision

(year)

2011

1996

1985

11

single record

(year)

1993

2001

©ZFMK

12

family

Eulophidae

species

Chrysocharis polyzo

Chrysocharis prodice

Chrysocharis pubens

Chrysocharis pubicornis

Chrysocharis purpurea

Chrysocharis submutica

Chrysocharis viridis

Chrysonotomyia germanica

Cirrospilus argei

Cirrospilus diallus

Cirrospilus elegantissimus

Cirrospilus elongatus

Cirrospilus lyncus

Cirrospilus pictus

Cirrospilus salatis

Cirrospilus singa

Cirrospilus staryi

Cirrospilus viticola

Cirrospilus vittatus

Closterocerus lanassa

Closterocerus lyonetiae

Closterocerus pannonicus

Closterocerus ruforum

Closterocerus trifasciatus

Colpoclypeus florus

Crataepus marbis

Dahlbominus fuscipennis

Dermatopelte budensis

Derostenus gemmeus

Derostenus punctiscuta

Dichatomus acerinus

Dicladocerus albitarsis

Dicladocerus euryalus

Dicladocerus westwoodii

Diglyphus chabrias

Diglyphus crassinervis

Diglyphus isaea

Diglyphus minoeus

Diglyphus pachyneurus

Diglyphus poppoea

Diglyphus pusztensis

Bonn zoological Bulletin 67 (2): 101-107

Maximilian Weber et al.

author

(Walker)

Delucchi

(Zetterstedt)

Bukovskii

Graham

(Nees)

(Erdos)

(Crawford)

Walker

Westwood

Boutek

Walker

(Nees)

Walker

Boutek

(Rondan1)

Walker

(Walker)

(Ferriere)

(Erdos)

(Krausse)

Westwood

(Walker)

(Zetterstedt)

Erdos & Novicky

Westwood

Thomson

Forster

(Ashmead)

(Haliday)

Westwood

(Walker)

Erdos

(Walker)

Graham

Walker

(Erdos & Novicky)

distribution

Germany

revision

(year)

1985

1990

1984

2003

1987

single record

(year)

2001

1959

2001

©ZFMK

family

Eulophidae

species

Diglyphus subplanus

Dimmockia brevicornis

Elachertus aequalis

Elachertus aeruginosus

Elachertus artaeus

Elachertus charondas

Elachertus coerulescens

Elachertus cyaneus

Elachertus deplanatus

Elachertus ditissimus

Elachertus facialis

Elachertus fenestratus

Elachertus gallicus

Elachertus inunctus

Elachertus isadas

Elachertus laevis

Elachertus lateralis

Elachertus lunatus

Elachertus pilosiscuta

Elachertus plagiatus

Elachertus pulcher

Elachertus reticulatus

Elachertus timidus

Elachertus walkeri

Elasmus flabellatus

Elasmus nudus

Elasmus polistis

Elasmus schmitti

Elasmus unicolor

Elasmus viridiceps

Elasmus westwoodi

Entedon abdera

Entedon aequilongus

Entedon calcicola

Entedon canaliculatus

Entedon caudatus

Entedon cavicornis

Entedon chalybaeus

Entedon cioni

Entedon cionobius

Entedon confinis

Bonn zoological Bulletin 67 (2): 101-107

(APPENDIX 1)

author

(Erdos)

Forster

(Walker)

Nees

Forster

(Ratzeburg)

Forster

Nees

Erdos

Nees

(Walker)

(Forster)

(Spinola)

Forster

Bouéek

Forster

(Erdos)

Ratzeburg

Forster

(Ratzeburg)

(Fonscolombe)

(Nees)

Burks

Ruschka

(Rondani)

Thomson

Giraud

Walker

Ratzeburg

Graham

(Forster)

Ratzeburg

Thomson

Ratzeburg

distribution

Germany

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna

revision

(year)

1964

1995

1208

1995

1:99 5:

1995

L999

LOS9

1999

13

single record

(year)

1968

2001

1972

2001

1968

2001

©ZFMK

14

family

Eulophidae

species

Entedon connexus

Entedon costalis

Entedon crassiscapus

Entedon diotimus

Entedon ergias

Entedon gracilior

Entedon heyeri

Entedon hylotomarum

Entedon inconspicuus

Entedon incultus

Entedon insignis

Entedon lixi

Entedon longiventris

Entedon longus

Entedon luteipes

Entedon mecini

Entedon methion

Entedon oxys

Entedon parvicalcar

Entedon philiscus

Entedon pinetorum

Entedon procioni

Entedon punctiscapus

Entedon rumicis

Entedon setifrons

Entedon tenuitarsis

Entedon thomsonianus

Entedon tibialis

Entedon transparens

Entedon unicostatus

Entedon vaginulae

Entedon xanthostoma

Entedon zanara

Entedonomphale bicolorata

Entedonomphale carbonaria

Euderomphale chelidonii

Euderus agrili

Euderus albitarsis

Euderus viridis

Eulophus abdominalis

Eulophus albitarsus

Bonn zoological Bulletin 67 (2): 101-107

Maximilian Weber et al.

author

Ratzeburg

Dalman

Erdos

Walker

Graham

(Ratzeburg)

Ratzeburg

Askew

Erdos

Ratzeburg

Bouéek

Ratzeburg

Askew

Walker

Askew

Thomson

Walker

Ratzeburg

Erdos

Thomson

Graham

Askew

Thomson

Erdos

(Nees)

Ratzeburg

(Ratzeburg)

Walker

(Ishii)

(Erdos)

Erdos

Bouéek

(Zetterstedt)

Thomson

Nees

Ratzeburg

distribution

Germany

revision

(year)

1999

1989

1999

jee,

1999

Ios

1999

1989

1999

foes

1999

single record

(year)

2001

2003

2001

1966

2001

©ZFMK

family

Eulophidae

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna 15

species

Eulophus bifasciatus

Eulophus binotatus

Eulophus blancardellae

Eulophus breviramulis

Eulophus cecidomyiarum

Eulophus cephalotes

Eulophus chrysomelae

Eulophus coccorum

Eulophus cyanescens

Eulophus depressus

Eulophus dubitabilis

Eulophus emicans

Eulophus foveolatus

Eulophus inconspicuus

Eulophus larvarum

Eulophus nitidulus

Eulophus pennicornis

Eulophus polycerus

Eulophus ramicornis

Eulophus rupicapra

Eulophus semicupreus

Eulophus smerinthicida

Eulophus tabidus

Eulophus thespius

Eulophus vagus

Euplectrus bicolor

Hemiptarsenus fulvicollis

Hemiptarsenus ornatus

Hemiptarsenus unguicellus

Hemiptarsenus wailesellae

Hemiptarsenus waterhousii

Holarcticesa clinius

Holcotetrastichus rhosaces

Hyssopus geniculatus

FAyssopus nigritulus

Hyssopus olivaceus

FHyssopus tephridus

FAyssopus thymus

Kocourekia debilis

Melittobia acasta

Mestocharis bimacularis

Bonn zoological Bulletin 67 (2): 101-107

(APPENDIX 1)

author

Nees

Forster

Bouché

Forster

Ratzeburg

Nees

Ratzeburg

Bouéek

Nees

Forster

Nees

(Linnaeus)

Nees

Forster

(Fabricius)

Forster

Nees

Bouéek

Nees

Walker

Nees

(Swederus)

Westwood

(Nees)

(Zetterstedt)

Nowicki

Westwood

(Walker)

(Hartig)

(Zetterstedt)

(Thomson)

Yefremova

Girault

(Ratzeburg)

(Walker)

(Dalman)

distribution

Germany

revision single record

(year) (year)

2001

1963

2003

2003 2001

2001

1988

©ZFMK

16

family

Eulophidae

species

Mestocharis maculata

Microlycus biroi

Microlycus heterocerus

Minotetrastichus frontalis

Minotetrastichus platanellus

Minotetrastichus prolongatus

Miotropis unipuncta

Necremnus aenigmaticus

Necremnus artynes

Necremnus cosconius

Necremnus folia

Necremnus leucarthros

Necremnus metalarus

Necremnus tidius

Neochrysocharis albiscapus

Neochrysocharis aratus

Neochrysocharis chlorogaster

Neochrysocharis clinias

Neochrysocharis cuprifrons

Neochrysocharis dimas

Neochrysocharis formosus

Neochrysocharis microstoma

Neochrysocharis nunbergi

Omphale acuminata

Omphale aethiops

Omphale aetius

Omphale betulicola

Omphale brevis

Omphale breviventris

Omphale chryseis

Omphale clymene

Omphale clypealis

Omphale connectens

Omphale grahami

Omphale incognita

Omphale lugens

Omphale lugubris

Omphale matrana

Omphale nitens

Omphale obscura

Omphale phruron

Bonn zoological Bulletin 67 (2): 101-107

Maximilian Weber et al.

author

(Forster)

Erdos

Thomson

(Nees)

(Mercet)

Graham

(Nees)

Gibson

(Walker)

(Nees)

(Walker)

Erdos

(Walker)

(Erdos)

(Walker)

Erdos

(Walker)

(Westwood)

(Graham)

(Szczepansk1)

Giswijt

Graham

(Walker)

Graham

(Walker)

(Thomson)

Graham

Giswijt

Hansson & Shevtsova

(Nees)

Askew

Erdos

Graham

(Forster)

(Walker)

distribution

Germany

revision

(year)

1988

2015

1990

single record

(year)

2001

2015

2001

2012

2001

2012

2001

©ZFMK

family

Eulophidae

species

Omphale rubigus

Omphale salicis

Omphale stelteri

Omphale sulciscuta

Omphale telephe

Omphale theana

Omphale varipes

Omphale versicolor

Oomyzus galerucivorus

Oomyzus gallerucae

Oomyzus incertus

Oomyzus pegomyae

Oomyzus scaposus

Oomyzus sempronius

Parasecodella obscura

Pediobius alaspharus

Pediobius alcaeus

Pediobius brachycerus

Pediobius calamagrostidis

Pediobius cassidae

Pediobius chilaspidis

Pediobius claridgei

Pediobius clita

Pediobius crassicornis

Pediobius dactylicola

Pediobius epeus

Pediobius epigonus

Pediobius eubius

Pediobius facialis

Pediobius festucae

Pediobius flaviscapus

Pediobius foliorum

Pediobius lysis

Pediobius metallicus

Pediobius nigritarsis

Pediobius phalaridis

Pediobius phragmitis

Pediobius phyllotretae

Pediobius planiventris

Pediobius polanensis

Pediobius pyrgo

Bonn zoological Bulletin 67 (2): 101-107

(APPENDIX 1)

author

(Walker)

(Haliday)

(Bouéek)

(Thomson)

(Walker)

(Thomson)

(Nees)

(Hedqvist)

(Fonscolombe)

(Ratzeburg)

Graham

(Thomson)

(Erdos)

(Thomson)

(Walker)

(Thomson)

Dawah

Erdos

Bouéek

Dawah

(Walker)

(Thomson)

Dawah

(Walker)

(Giraud)

Dawah

(Thomson)

(Geoffroy)

(Walker)

(Nees)

(Thomson)

Dawah

Bouéek

(Riley)

(Thomson)

Bouéek

(Walker)

distribution

Germany

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna

revision

(year)

2003

1995:

17

single record

(year)

2012

2001

©ZFMK

18

family

Eulophidae

species

Pediobius saulius

Pediobius termerus

Pediobius tetratomus

Platyplectrus chlorocephalus

Platyplectrus laeviscuta

Platyplectrus pannonica

Pnigalio agraules

Pnigalio cristatus

Pnigalio cruciatus

Pnigalio epilobii

Pnigalio longulus

Pnigalio monilicornis

Pnigalio nemati

Pnigalio obscurus

Pnigalio pectinicornis

Pnigalio phragmitis

Pnigalio soemius

Pnigalio tardulus

Pnigalio tricuspis

Pnigalio tridentatus

Pronotalia carlinarum

Pronotalia inflata

Quadrastichus misellus

Quadrastichus pedicellaris

Quadrastichus sajoi

Rhicnopelte crassicornis

Sigmophora brevicornis

Sphenolepis pygmaea

Stenomesius rufescens

Stepanovia aurantiaca

Stepanovia eurytomae

Sympiesis acalle

Sympiesis dolichogaster

Sympiesis euspilapterygis

Sympiesis flavopicta

Sympiesis gordius

Sympiesis grahami

Sympiesis gregori

Sympiesis kelebiana

Sympiesis notata

Sympiesis sericeicornis

Bonn zoological Bulletin 67 (2): 101-107

Maximilian Weber et al.

author

(Walker)

(Thomson)

(Nees)

(Thomson)

(Erdos)

(Walker)

(Ratzeburg)

Boucéek

(Zetterstedt)

(Westwood)

(Ratzeburg)

(Linnaeus)

(Erdos)

(Walker)

(Nees)

(Erdos)

(Thomson)

(Szelényi & Erdos)

Graham

(Delucchi)

(Thomson)

(Szelény1)

(Nees)

(Panzer)

Nees

(Retzius)

(Ratzeburg)

(Nees)

(Walker)

Ashmead

(Erdos)

Boutek

(Walker)

Erdos

Boutéek

Erdos

(Zetterstedt)

(Nees)

distribution

Germany

revision

(year)

1984

1977

single record

(year)

2001

2007

2001

©ZFMK

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna 19

(APPENDIX 1)

revision single record

family species author distribution (year) (year)

Eulophidae Sympiesis solitaria Szelényi Germany 2005

Eulophidae Sympiesis viridula (Thomson) Germany

Eulophidae Sympiesis xanthostoma (Nees) Germany

Eulophidae Tamarixia actis (Walker) Germany

Eulophidae Tamarixia monesus (Walker) Germany

Eulophidae Tamarixia pronomus (Walker) Germany

Eulophidae Tamarixia pubescens (Nees) Germany

Eulophidae Tamarixia upis (Walker) Germany

Eulophidae Tetrastichus atratulus (Nees) Germany 1953

Eulophidae Tetrastichus atrocoeruleus (Nees) Germany 1953 2001

Eulophidae Tetrastichus brachyopae Graham Germany 1953

Eulophidae Tetrastichus capitatus (Ratzeburg) Germany 1953 2001

Eulophidae Tetrastichus clito (Walker) Germany eS)

Eulophidae Tetrastichus coeruleus (Nees) Germany 1953

Eulophidae Tetrastichus halidayi (Graham) Germany 1953

Eulophidae Tetrastichus heeringi Delucchi Germany 1953

Eulophidae Tetrastichus hylotomarum (Bouché) Germany 1953

Eulophidae Tetrastichus ilithyia (Walker) Germany 1953

Eulophidae Tetrastichus inunctus (Nees) Germany 1953 2001

Eulophidae Tetrastichus julis (Walker) Germany Ls:

Eulophidae Tetrastichus legionarius Giraud Germany 1953

Eulophidae Tetrastichus lyridice (Walker) Germany 1953 2001

Eulophidae Tetrastichus miser (Nees) Germany 1953

Eulophidae Tetrastichus murcia (Walker) Germany 1953

Eulophidae Tetrastichus ooctonus (Kawall) Germany 1953

Eulophidae Tetrastichus polyporinus Askew Germany 1953 2007

Eulophidae Tetrastichus setifer Thomson Germany 1g53 2002

Eulophidae Tetrastichus sinope (Walker) Germany 1953 1963

Eulophidae Tetrastichus telon (Graham) Germany 1953

Eulophidae Thripastichus gentilei (Del Guercio) Germany 1966

Eulophidae Trjapitzinichus evanescens (Ratzeburg) Germany

Eulophidae Xanthellum szabopatayi Moczar Germany

Eupelmidae Anastatus bifasciatus (Geoffroy) Germany

Eupelmidae Anastatus catalonicus Bolivar y Pieltain Germany

Eupelmidae Anastatus giraudi (Ruschka) Germany 2001

Eupelmidae Anastatus insignis (Forster) Germany 2001

Eupelmidae Anastatus japonicus Ashmead Germany

Eupelmidae Anastatus oscari (Ruthe) Germany

Eupelmidae Brasema stenus (Boucéek) Germany 2006

Eupelmidae Calosota acron (Walker) Germany 2010 2001

Eupelmidae Calosota grylli Erdos Germany 2010 1993

Bonn zoological Bulletin 67 (2): 101-107 ©ZFMK

20

family

Eupelmidae

Eurytomidae

species

Calosota metallica

Calosota obscura

Calosota vernalis

Calymmochilus dispar

Eupelmus annulatus

Eupelmus atropurpureus

Eupelmus azureus

Eupelmus brachynterae

Eupelmus fuscipennis

Eupelmus hartigi

Eupelmus linearis

Eupelmus microzonus

Eupelmus splendens

Eupelmus stramineipes

Eupelmus urozonus

Eupelmus vesicularis

Eusandalum coronatum

Eusandalum elongatum

Eusandalum flavipenne

Eusandalum inerme

Eusandalum walkeri

Merostenus excavatus

Merostenus rostratus

Metapelma nobile

Aximopsis nodularis

Bruchophagus astragali

Bruchophagus ater

Bruchophagus gibbus

Bruchophagus phlei

Bruchophagus platypterus

Bruchophagus roddi

Eurytoma abieticola

Eurytoma abrotani

Eurytoma aciculata

Eurytoma adleriae

Eurytoma afra

Eurytoma aloineae

Eurytoma appendigaster

Eurytoma aquatica

Eurytoma arctica

Eurytoma aspila

Bonn zoological Bulletin 67 (2): 101-107

Maximilian Weber et al.

author

(Gahan)

Ruschka

Curtis

Bouéek & Andriescu

Nees

Dalman

Ratzeburg

(Schwagrichen)

Forster

Giraud

Nikol’skaya

Dalman

(Retzius)

(Thomson)

(Ruschka)

Ruschka

(Ratzeburg)

(Curtis)

(Dalman)

(Ruschka)

(Forster)

(Boheman)

Fedoseeva

(Walker)

(Boheman)

(Erdos)

(Walker)

Gussakovskiy

Ratzeburg

(Panzer)

Ratzeburg

Zerova

Boheman

(Burks)

(Swederus)

Erdos

Thomson

(Walker)

distribution

Germany

revision

(year)

2010

2016

1967

2017

1959

single record

(year)

2001

2006

2001

2013

1958

2001

©ZFMK

family

Eurytomidae

species

Eurytoma aterrima

Eurytoma baldingerae

Eurytoma brunniventris

Eurytoma castor

Eurytoma caulicola

Eurytoma centaureae

Eurytoma collaris

Eurytoma compressa

Eurytoma coxalis

Eurytoma crassinervis

Eurytoma curculionum

Eurytoma curta

Eurytoma cynipsea

Eurytoma danilovi

Eurytoma danuvica

Eurytoma dentata

Eurytoma erdoesi

Eurytoma extincta

Eurytoma flavimana

Eurytoma goidanichi

Eurytoma gracilior

Eurytoma hypochoeridis

Eurytoma jaceae

Eurytoma kangasi

Eurytoma maura

Eurytoma mayri

Eurytoma microneura

Eurytoma morio

Eurytoma neesii

Eurytoma nobbei

Eurytoma nodulosa

Eurytoma obscura

Eurytoma onobrychidis

Eurytoma oophaga

Eurytoma pediaspisi

Eurytoma petiolata

Eurytoma phalaridis

Eurytoma pinetorum

Eurytoma pistaciae

Eurytoma pollux

Eurytoma pumila

Bonn zoological Bulletin 67 (2): 101-107

(APPENDIX 1)

author

(Schrank)

Erdos

Ratzeburg

Claridge

Zerova

Claridge

Walker

(Fabricius)

Erdos

Thomson

Mayr

Walker

Boheman

Zerova

Erdos

Mayr

Szelényi

Ratzeburg

Boheman

Boutek

Dalla Torre

Claridge

Mayr

Hedqvist

Boheman

Ashmead

Ratzeburg

Boheman

Walker

Mayr

Ratzeburg

Boheman

Nikol’skaya

Silvestri

Pujade i Villar

Forster

Graham

Ratzeburg

Rondani

Claridge

Forster

distribution

Germany

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna

revision

(year)

2014

21

single record

(year)

2001

2006

2001

2006

2001

©ZFMK

22

family

Eurytomidae

species

Eurytoma punctulata

Eurytoma robusta

Eurytoma rosae

Eurytoma roseni

Eurytoma rufa

Eurytoma rufipes

Eurytoma scabra

Eurytoma serratulae

Eurytoma setigera

Eurytoma sphegum

Eurytoma strigifrons

Eurytoma striolata

Eurytoma truncatella

Eurytoma verticillata

Eurytoma wachtli

Mangoma salicis

Sycophila biguttata

Sycophila concinna

Sycophila fasciata

Sycophila flavicollis

Sycophila mellea

Sycophila submutica

Systole albipennis

Systole atratula

Tetramesa aciculata

Tetramesa affinis

Tetramesa agrostidis

Tetramesa airae

Tetramesa albomaculatum

Tetramesa brachypodii

Tetramesa brevicollis

Tetramesa brevicornis

Tetramesa brischkei

Tetramesa calamagrostidis

Tetramesa cylindrica

Tetramesa eximia

Tetramesa flavipes

Tetramesa foersteri

Tetramesa fulvicollis

Tetramesa fumipennis

Tetramesa giraudi

Bonn zoological Bulletin 67 (2): 101-107

Maximilian Weber et al.

author

Forster

Mayr

Nees

Claridge

Zerova

Walker

Forster

(Fabricius)

Mayr

(Fabricius)

Thomson

Ratzeburg

Zerova

(Fabricius)

Mayr

(Walker)

(Swederus)

(Boheman)

(Thomson)

(Walker)

(Curtis)

(Thomson)

Walker

(Dalla Torre)

(Schlechtendal)

(Hedicke)

(Howard)

(Schlechtendal)

(Ashmead)

(Schlechtendal)

(Walker)

(Schlechtendal)

(Giraud)

(Forster)

(Hedicke)

(Walker)

(Schlechtendal)

distribution

Germany

single record

(year)

2001

2006

2001

2006

2001

©ZFMK

family

Eurytomidae

Leucospidae

Mymaridae

species

Tetramesa hordei

Tetramesa hyalipennis

Tetramesa laevigata

Tetramesa linearis

Tetramesa longicornis

Tetramesa longula

Tetramesa maritima

Tetramesa novalis

Tetramesa petiolata

Tetramesa phleicola

Tetramesa phragmitis

Tetramesa poae

Tetramesa puccinellae

Tetramesa scheppigi

Tetramesa schlechtendali

Tetramesa schmidti

Leucospis dorsigera

Leucospis gigas

Leucospis intermedia

Alaptus auranti

Alaptus extremus

Alaptus fusculus

Alaptus minimus

Alaptus novickyi

Alaptus pallidornis

Alaptus schmitzi

Alaptus stammeri

Anagrus atomus

Anagrus avalae

Anagrus bakkendorfi

Anagrus ensifer

Anagrus foersteri

Anagrus incarnatosimilis

Anagrus incarnatus

Anagrus nigriceps

Anagrus subfuscus

Anaphes aries

Anaphes brachygaster

Anaphes brevitarsis

Anaphes compressus

Anaphes depressus

Bonn zoological Bulletin 67 (2): 101-107

(APPENDIX 1)

author

(Harris)

(Walker)

(Hedicke)

(Walker)

(Dalman)

(Hedicke)

Zerova

(Walker)

(Hedicke)

(Erdos)

(Schlechtendal)

Zerova

(Schlechtendal)

(Hedicke)

Fabricius

Illiger

(Mercet)

Soyka

Walker

Westwood

Soyka

Forster

Soyka

(Linnaeus)

Soyka

Debauche

(Ratzeburg)

Soyka

Haliday

(Smits van Burgst)

Forster

Debauche

(Debauche)

(Soyka)

distribution

Germany

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna

revision

(year)

1959

1959)

1959

2016

2009

pu

single record

(year)

2001

1921

2001

©ZFMK

24

family

Mymaridae

species

Anaphes diana

Anaphes discolorisimilis

Anaphes dorcas

Anaphes exiguus

Anaphes flavipes

Anaphes flavus

Anaphes fuscipennis

Anaphes gauthieri

Anaphes germaniacus

Anaphes gracillimus

Anaphes intermedius

Anaphes leptoceras

Anaphes longicornis

Anaphes luna

Anaphes malchinensis

Anaphes medius

Anaphes neospecialis

Anaphes ovipositor

Anaphes pannonicus

Anaphes parallelipennis

Anaphes parvus

Anaphes pilicornis

Anaphes quadraticornis

Anaphes rectipennis

Anaphes regulus

Anaphes serenus

Anaphes silesicus

Anaphes sulphuripes

Anaphes variatus

Anaphes wertaneki

Arescon dimidiatus

Camptoptera cardui

Camptoptera magna

Camptoptera papaveris

Camptoptera punctum

Caraphractus cinctus

Cleruchus janetscheki

Cleruchus pluteus

Cosmocomoidea atra

Cosmocomoidea oxypygus

Erythmelus agilis

Bonn zoological Bulletin 67 (2): 101-107

Maximilian Weber et al.

author

(Girault)

(Soyka)

(Debauche)

(Soyka)

(Forster)

(Soyka)

Haliday

Debauche

Ozdikmen

(Soyka)

(Debauche)

Walker

(Girault)

(Soyka)

Soyka

(Soyka)

Soyka

(Soyka)

(Forster)

(Soyka)

Walker

(Soyka)

(Curtis)

(Forster)

Soyka

Forster

(Shaw)

Walker

Novicky

Enock

(Forster)

(Enock)

distribution

Germany

revision

(year)

2003

2014

2011

2014

2012

2003

single record

(year)

2001

1999

1973

2001

2014

2011

2001

2003

©ZFMK

family

Mymaridae

species

Erythmelus flavovarius

Erythmelus panis

Eustochus atripennis

Gonatocerus fuscicornis

Gonatocerus longicornis

Gonatocerus minimus

Gonatocerus pictus

Litus cynipseus

Lymaenon litoralis

Lymaenon longior

Lymaenon novickyi

Mymar pulchellum

Ooctonus hemipterus

Ooctonus insignis

Ooctonus notatus

Ooctonus sublaevis

Ooctonus vulgatus

Polynema capillatum

Polynema crassicorne

Polynema euchariforme

Polynema flavipes

Polynema foersteri

Polynema fumipenne

Polynema fuscipes

Polynema gracile

Polynema gracilior

Polynema laetum

Polynema latipenne

Polynema malkwitzi

Polynema marginatum

Polynema neustadti

Polynema novickyi

Polynema ovatum

Polynema ovulorum

Polynema pusillum

Polynema sachtlebeni

Polynema schmitzi

Polynema spectabile

Polynema stammeri

Polynema vitripenne

Stephanodes similis

Bonn zoological Bulletin 67 (2): 101-107

(APPENDIX 1)

author

(Walker)

(Enock)

(Curtis)

(Walker)

Nees

Forster

(Haliday)

Haliday

(Haliday)

(Soyka)

Curtis

Haliday

Walker

Forster

Haliday

Soyka

Forster

Haliday

Walker

Soyka

Walker

Haliday

(Nees)

Soyka

Forster

Soyka

(Soyka)

Soyka

(Linnaeus)

Haliday

Soyka

(Soyka)

Soyka

(Forster)

distribution

Germany

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna

revision

(year)

2003

2007

2013

2004

1996

2010

2006

1997

25

single record

(year)

2001

1881

2001

1978

2001

1946

2001

©ZFMK

26

family

Mymaridae

Ormyridae

Perilampidae

species

Stethynium triclavatum

Ormyrus caeruleus

Ormyrus chalybeus

Ormyrus cingulatus

Ormyrus cosmozonus

Ormyrus diffinis

Ormyrus gratiosus

Ormyrus nitidulus

Ormyrus orientalis

Ormyrus papaveris

Ormyrus pomaceus

Ormyrus punctulatus

Ormyrus rufimanus

Ormyrus versicolor

Ormyrus violaceus

Chrysolampus aeneicornis

Chrysolampus anguliventris

Chrysolampus attenuatus

Chrysolampus brevicornis

Chrysolampus coeruleovirens

Chrysolampus dubius

Chrysolampus ellipticus

Chrysolampus excellens

Chrysolampus foersteri

Chrysolampus fuscimanus

Chrysolampus gibbosus

Chrysolampus gilvipes

Chrysolampus granulatus

Chrysolampus indubitatus

Chrysolampus interruptus

Chrysolampus laevipetiolatus

Chrysolampus pachymerus

Chrysolampus pallitarsis

Chrysolampus punctatus

Chrysolampus rufitarsis

Chrysolampus scapularis

Chrysolampus splendidulus

Chrysolampus subcarinatus

Chrysolampus subsessilis

Chrysolampus tenuiscapus

Chrysolampus thenae

Bonn zoological Bulletin 67 (2): 101-107

Maximilian Weber et al.

author

Enock

Walker

(Ratzeburg)

(Forster)

Forster

(Fonscolombe)

(Forster)

(Fabricius)

Walker

(Perris)

(Geoffroy)

(Ratzeburg)

Mayr

Forster

Ratzeburg

Nees

Forster

Della Torre

Forster

(Forster)

Ratzeburg

(Spinola)

Forster

Nees

Forster

(Walker)

distribution

Germany

revision

(year)

2014

single record

(year)

2001

1844

2001

1860

2001

2014

2001

2014

©ZFMK

family

Perilampidae

Pteromalidae

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna

species

Chrysolampus transversus

Chrysomalla roseri

Perilampus aeneus

Perilampus angustus

Perilampus auratus

Perilampus aureoviridis

Perilampus chrysonotus

Perilampus cristatus

Perilampus cuprinus

Perilampus intermedius

Perilampus laevifrons

Perilampus micans

Perilampus minutalis

Perilampus neglectus

Perilampus nitens

Perilampus ruficornis

Perilampus ruschkai

Perilampus tristis

Ablaxia robusta

Acroclisis nigricornis

Aggelma violacea

Anisopteromalus calandrae

Anogmus hohenheimensis

Anogmus hungaricus

Anogmus piceae

Anogmus strobilorum

Anogmus vala

Apsilocera bramleyi

Arthrolytus discoideus

Arthrolytus maculipennis

Arthrolytus ocellus

Asaphes suspensus

Asaphes vulgaris

Caenacis capnopterus

Caenacis flavipes

Caenacis inflexa

Caenacis lauta

Callitula bicolor

Callitula elongata

Callitula ferrierei

Callitula pyrrhogaster

Bonn zoological Bulletin 67 (2): 101-107

(APPENDIX 1)

author

Forster

(Rossius)

Nees

(Panzer)

Walker

Forster

Boutek

Dalman

Steffan

Boutek

Walker

(Fabricius)

Hellén

Mayr

Hedqvist

Forster

(Zetterstedt)

(Howard)

(Ratzeburg)

(Erdos)

(Ruschka)

(Thomson)

(Walker)

Graham

(Nees)

(Walker)

(Nees)

Walker

(Ratzeburg)

Masi

(Ratzeburg)

(Walker)

Spinola

(Thomson)

Boutek

(Walker)

distribution

Germany

revision

(year)

2014

1961

21

single record

(year)

2001

1985

2001

1985

2001

1966

©ZFMK

28

family

Pteromalidae

species

Capellia cecidomyiae

Capellia orneus

Catolaccus ater

Cea pulicaris

Cecidolampa barbotini

Cecidostiba docimus

Cecidostiba fungosa

Cecidostiba geganius

Cecidostiba semifascia

Cerocephala cornigera

Cerocephala rufa

Cheiropachus quadrum

Chlorocytus alticornis

Chlorocytus breviscapus

Chlorocytus diversus

Chlorocytus formosus

Chlorocytus harmolitae

Chlorocytus inchoatus

Chlorocytus longicauda

Chlorocytus phalaridis

Chlorocytus polichna

Chlorocytus spicatus

Chlorocytus terminalis

Chlorocytus ultonicus

Cleonymus apicalis

Cleonymus cyaneus

Cleonymus elongatus

Cleonymus eximius

Cleonymus laticornis

Cleonymus viridinitens

Coelopisthia areolata

Coelopisthia eurynota

Coelopisthia extenta

Collentis suecicus

Colotrechnus subcoeruleus

Conomorium amplum

Conomorium patulum

Coruna clavata

Cratomus megacephalus

Cryptoprymna atra

Cyclogastrella simplex

Bonn zoological Bulletin 67 (2): 101-107

Maximilian Weber et al.

author

(Ratzeburg)

(Walker)

(Ratzeburg)

Walker

Askew

(Walker)

(Geoffroy)

(Walker)

Westwood

(Walker)

(Fabricius)

Graham

(Walker)

Boutek

Graham

(Thomson)

Graham

(Walker)

Graham

Forster

Walker

Forster

Askew

(Forster)

(Walker)

(Graham)

Thomson

(Walker)

Walker

(Fabricius)

(Walker)

distribution

Germany

revision

(year)

1961

1972

19hz

1972

single record

(year)

2001

©ZFMK

family

Pteromalidae

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna

(APPENDIX 1)

species

Cyrtogaster clavicornis

Cyrtogaster vulgaris

Dibrachoides cionobius

Dibrachoides dynastes

Dibrachys affinis

Dibrachys fuscicornis

Dibrachys lignicola

Dibrachys microgastri

Dibrachys verovesparum

Diglochis crinifrons

Diglochis paludicola

Diglochis sylvicola

Dimachus cingulum

Dinarmus acutus

Dinotiscus aponius

Dinotiscus avrupanensis

Dinotiscus colon

Dinotiscus eupterus

Dinotiscus isvicrensis

Dipara petiolata

Dirhicnus clandestinus

Dirhicnus ramealis

Endomychobius endomychi

Epicopterus choreiformis

Erdoesia tessellata

Erdoesina alboannulata

Eulonchetron torymoides

Eumacepolus einersbergensis

Eumacepolus pulcher

Eumacepolus saxeseni

Euneura lachni

Euneura saetosa

Euneura sopolis

Eunotus acutus

Eunotus areolatus

Eunotus cretaceus

Eunotus nigriclavis

Eunotus obscurus

Eunotus parvulus

Eurydinota leptomera

Gastracanthus erythrogaster

Bonn zoological Bulletin 67 (2): 101-107

author

Walker

Graham

(Forster)

Masi

(Walker)

Graham

(Bouché)

Peters & Baur

(Forster)

Abraham

(Walker)

(Nees)

(Thomson)

(Walker)

Doganlar

(Linnaeus)

(Walker)

Doganlar

Walker

(Forster)

(Nees)

(Walker)

Westwood

Bouéek

(Ratzeburg)

(Thomson)

(Ratzeburg)

Graham

(Ashmead)

(Delucchi)

(Walker)

Kurdjumov

(Ratzeburg)

Walker

(Forster)

Masi

Forster

(Dalla Torre)

distribution

Germany

revision

(year)

1965

1987

2011

2007

Log

1972

19FZ

1972

29

single record

(year)

2001

2005

2011

2001

2007

2001

1999

2001

1999

2001

©ZFMK

30

family

Pteromalidae

species

Gastracanthus pulcherrimus

Gastrancistrus acontes

Gastrancistrus amabilis

Gastrancistrus aphidum

Gastrancistrus ater

Gastrancistrus autumnalis

Gastrancistrus claviger

Gastrancistrus fulvicornis

Gastrancistrus fulvicoxis

Gastrancistrus fuscicornis

Gastrancistrus glabellus

Gastrancistrus picipes

Gastrancistrus pusztensis

Gastrancistrus rosularum

Gastrancistrus salicis

Gastrancistrus torymiformis

Gastrancistrus undulatus

Gastrancistrus xylophagorum

Glyphognathus convexus

Glyphognathus flammeus

Glyphognathus laevigatus

Glyphognathus nitidus

Gyrinophagus luteipes

Habritys brevicornis

Halticoptera aenea

Halticoptera circulus

Halticoptera collaris

Halticoptera corrusca

Halticoptera crius

Halticoptera dimidiata

Halticoptera elongatula

Halticoptera flavicornis

Halticoptera izzetbaysali

Halticoptera laevigata

Halticoptera mustela

Halticoptera patellana

Halticoptera plana

Halticoptera polita

Halticoptera smaragdina

Halticoptera triannulata

Halticoptera vehbikoci

Bonn zoological Bulletin 67 (2): 101-107

Maximilian Weber et al.

author

Westwood

Walker

(Girault & Dodd)

(Ratzeburg)

(Nees)

(Walker)

Forster

(Walker)

Graham

Walker

(Nees)

(Erdos)

(Ratzeburg)

(Nees)

(Ratzeburg)

(Delucchi)

Ruschka

(Ratzeburg)

(Walker)

(Gravenhorst)

(Walker)

(Forster)

Graham

(Spinola)

Doganlar

Thomson

(Walker)

(Dalman)

(Forster)

(Walker)

(Curtis)

(Erdos)

Doganlar

distribution

Germany

revision

(year)

2006

single record

(year)

2001

1966

2001

2009

2001

2006

2001

2006

2001

2006

2003

2006

©ZFMK

family

Pteromalidae

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna

species

Hemitrichus seniculus

Heydenia pretiosa

Hobbya stenonota

Holcaeus calligetus

Holcaeus glabriculus

Holcaeus gorgasus

Holcaeus siccatorum

Holcaeus stenogaster

Holcaeus stylatus

Homoporus apharetus

Homoporus arestor

Homoporus destructor

Homoporus febriculosus

Homoporus femoralis

Homoporus fulviventris

Homoporus gibbiscuta

Homoporus luniger

Homoporus nypsius

Homoporus semiluteus

Homoporus subniger

Hyperimerus pusillus

Isocyrtus laetus

Janssoniella ambigua

Janssoniella caudata

Lampoterma bianellatum

Lampoterma viride

Lamprotatus brevicornis

Lamprotatus claviger

Lamprotatus picinervis

Lamprotatus simillimus

Lamprotatus splendens

Lamprotatus truncatus

Lariophagus distinguendus

Lariophagus puncticollis

Lariophagus rufipes

Lariophagus teutonus

Leptomeraporus nicaee

Macroglenes Bouceki

Macroglenes brevicornis

Macroglenes chalybeus

Macroglenes compressus

Bonn zoological Bulletin 67 (2): 101-107

(APPENDIX 1)

author

(Nees)

Forster

(Ratzeburg)

(Walker)

(Nees)

(Walker)

(Ratzeburg)

(Walker)

Graham

(Walker)

(Say)

(Girault)

(Forster)

(Walker)

(Thomson)

(Nees)

(Walker)

Walker

Graham

Kerrich

Graham

(Thomson)

Thomson

Delucchi

Westwood

(Fonscolombe)

(Forster)

(Miller)

Hedqvist

(Della Torre)

(Walker)

(Graham)

(Nees)

(Haliday)

(Forster)

distribution

Germany

revision

(year)

1961

2010

31

single record

(year)

2001

1969

2001

1881

2001

1999

2001

©ZFMK

32

family

Pteromalidae

species

Macroglenes conjungens

Macroglenes gramineus

Macroglenes penetrans

Macroglenes varicornis

Meraporus foveolatus

Meraporus glaber

Meraporus graminicola

Meraporus modestus

Merismus megapterus

Merismus nitidus

Merismus rufipes

Merismus splendens

Merismus viridis

Merisus flagellatus

Merisus splendidus

Mesopolobus aequus

Mesopolobus albitarsus

Mesopolobus amaenus

Mesopolobus bidentis

Mesopolobus citrinus

Mesopolobus clavatus

Mesopolobus clavicornis

Mesopolobus crassipes

Mesopolobus diffinis

Mesopolobus dilutipes

Mesopolobus dubius

Mesopolobus fagi

Mesopolobus fasciiventris

Mesopolobus fuscipes

Mesopolobus gemellus

Mesopolobus graminum

Mesopolobus incultus

Mesopolobus laticornis

Mesopolobus longicollis

Mesopolobus mediterraneus

Mesopolobus morys

Mesopolobus nobilis

Mesopolobus phragmitis

Mesopolobus rhabdophagae

Mesopolobus semiclavatus

Mesopolobus sericeus

Bonn zoological Bulletin 67 (2): 101-107

Maximilian Weber et al.

author

(Graham)

(Haliday)

(Kirby)

(Haliday)

(Forster)

(Szelény1)

Walker

(Forster)

Walker

(Walker)

Walker

Graham

(Delucchi)

Boutek

Walker

(Walker)

(Ratzeburg)

(Forster)

(Ratzeburg)

(Walker)

(Ratzeburg)

(Walker)

Askew & Lampe

Westwood

(Walker)

Baur & Muller

(Hardh)

(Walker)

Graham

(Mayr)

(Walker)

(Erdos)

(Graham)

(Ratzeburg)

(Forster)

distribution

Germany

revision

(year)

2010

2007

single record

(year)

2001

2013

2001

2007

2001

©ZFMK

family

Pteromalidae

species

Mesopolobus spermotrophus

Mesopolobus subfumatus

Mesopolobus tarsatus

Mesopolobus teliformis

Mesopolobus tibialis

Mesopolobus typographi

Mesopolobus verditer

Mesopolobus xanthocerus

Metacolus azurescens

Metacolus azureus

Metacolus unifasciatus

Metastenus concinnus

Miscogaster discedens

Miscogaster elegans

Miscogaster maculata

Miscogaster necopina

Miscogaster rufipes

Mokrzeckia pini

Muscidifurax raptor

Nasonia vitripennis

Neocatolaccus proximus

Nodisoplata diffinis

Norbanus obscurus

Norbanus scabriculus

Notanisus sexramosus

Ormocerus latus

Ormocerus vernalis

Oxysychus pilosulus

Pachycrepoideus vindemmiae

Pachyneuron aphidis

Pachyneuron coccorum

Pachyneuron flavipes

Pachyneuron formosum

Pachyneuron gibbiscuta

Pachyneuron grande

Pachyneuron groenlandicum

Pachyneuron innoxius

Pachyneuron leucopiscida

Pachyneuron muscarum

Pachyneuron piceae

Pachyneuron ratzeburgi

Bonn zoological Bulletin 67 (2): 101-107

author

Hussey

(Ratzeburg)

(Nees)

(Walker)

(Westwood)

(Ruschka)

(Norton)

(Thomson)

(Ratzeburg)

Forster

Walker

(Otten)

Walker

Delucchi

Walker

(Hartig)

Girault & Sanders

(Walker)

(Forster)

(Walker)

(Masi)

(Nees)

(Erdos)

Walker

(Thomson)

(Rondani)

(Bouché)

(Linnaeus)

(Forster)

Walker

Thomson

(Holmgren)

(Forster)

Mani

(Linnaeus)

(Ratzeburg)

Ozdikmen

distribution

Germany

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna

(APPENDIX 1)

revision

(year)

2007

1970

2010

2015

2

single record

(year)

1971

2001

1978

2001

1884

©ZFMK

34

family

Pteromalidae

species

Pachyneuron solitarium

Pachyneuron vitodurense

Pandelus flavipes

Panstenon oxylus

Pegopus inornatus

Peridesmia congrua

Peridesmia discus

Perniphora robusta

Phaenocytus glechomae

Platygerrhus affinis

Plutothrix bicolorata

Plutothrix coelius

Plutothrix trifasciata

Polycystus oscinidis

Pseudocatolaccus nitescens

Psilocera obscura

Psilocera punctifrons

Psilocera verticillata

Psilonotus achaeus

Psilonotus adamas

Psilonotus hortensia

Psychophagus omnivorus

Pteromalus aberrans

Pteromalus abieticola

Pteromalus acicularis

Pteromalus acuminatus

Pteromalus aequus

Pteromalus aerosus

Pteromalus agilis

Pteromalus albescens

Pteromalus albipennis

Pteromalus alternans

Pteromalus ambiguus

Pteromalus angustus

Pteromalus anomalipennis

Pteromalus apicalis

Pteromalus arborivagus

Pteromalus atramentarius

Pteromalus aurantiacus

Pteromalus aureolus

Pteromalus aurifacies

Bonn zoological Bulletin 67 (2): 101-107

Maximilian Weber et al.

author

(Hartig)

Delucchi

(Forster)

(Walker)

Ruschka

(Forster)

(Walker)

(Spinola)

(Walker)

(Thomson)

Kurdjumov

(Walker)

Walker

(Thomson)

(Forster)

Walker

(Walker)

Forster

Ratzeburg

Forster

Ratzeburg

Walker

Forster

Nees

Forster

Ratzeburg

(Thomson)

Forster

distribution

Germany

revision

(year)

1961

1992

eZ,

1992

single record

(year)

1966

1958

2001

©ZFMK

family

Pteromalidae

species

Pteromalus aurinitens

Pteromalus barycerus

Pteromalus bedeguaris

Pteromalus berylli

Pteromalus bifoveolatus

Pteromalus blandus

Pteromalus breviscapus

Pteromalus brunnicans

Pteromalus cardui

Pteromalus carinatus

Pteromalus caudiger

Pteromalus cerinopus

Pteromalus chalcophanes

Pteromalus chalybaeus

Pteromalus chlorospilus

Pteromalus chrysis

Pteromalus chrysos

Pteromalus cioni

Pteromalus cionobius

Pteromalus clavipes

Pteromalus coerulescens

Pteromalus coeruleus

Pteromalus colosseus

Pteromalus compactus

Pteromalus compos

Pteromalus concinnus

Pteromalus conformis

Pteromalus conoideus

Pteromalus conopidarum

Pteromalus crassus

Pteromalus cryptocephali

Pteromalus cubocephalus

Pteromalus cupreus

Pteromalus curculionoides

Pteromalus cylindraceus

Pteromalus cyniphidis

Pteromalus dahlbomi

Pteromalus dalmanni

Pteromalus decipiens

Pteromalus depressus

Pteromalus devorator

Bonn zoological Bulletin 67 (2): 101-107

author

Forster

(Thomson)

Walker

Forster

Ratzeburg

(Erdos)

Forster

(Graham)

Forster

Nees

(Walker)

Forster

Walker

(Thomson)

(Erdos)

Forster

Ratzeburg

Forster

(Graham)

Ratzeburg

(Bouéek)

Forster

Ratzeburg

Forster

Nees

(Bouché)

Forster

(Linnaeus)

Ratzeburg

Forster

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna

(APPENDIX 1)

distribution

Germany

a5

single record

(year)

2001

1966

2001

1841

1834

2001

1834

2001

©ZFMK

36

family

Pteromalidae

species

Pteromalus diadema

Pteromalus diatatus

Pteromalus difficilis

Pteromalus dimiduis

Pteromalus diminuator

Pteromalus dirutor

Pteromalus dispar

Pteromalus divitissimus

Pteromalus dolichurus

Pteromalus ecarinatus

Pteromalus egregius

Pteromalus elatus

Pteromalus elevatus

Pteromalus elongatus

Pteromalus eminens

Pteromalus esuriens

Pteromalus euurae

Pteromalus exiguus

Pteromalus exoletus

Pteromalus exsertus

Pteromalus extensus

Pteromalus facilis

Pteromalus fagi

Pteromalus fasciatus

Pteromalus faunigena

Pteromalus ferox

Pteromalus fervidus

Pteromalus festivus

Pteromalus flavipalpis

Pteromalus foersteri

Pteromalus fugax

Pteromalus furtivus

Pteromalus fuscopalpus

Pteromalus genuinus

Pteromalus gnavis

Pteromalus gracillimus

Pteromalus gratiosus

Pteromalus guttula

Pteromalus habilis

Pteromalus herbaceus

Pteromalus hercyniae

Bonn zoological Bulletin 67 (2): 101-107

Maximilian Weber et al.

author

Ratzeburg

Schmidt

Forster

Dalla Torre

Forster

(Curtis)

Dalla Torre

(Thomson)

Forster

(Walker)

Ratzeburg

Forster

Askew

Forster

Ratzeburg

(Thomson)

Forster

Ratzeburg

Dalla Torre

Forster

Dalla Torre

Forster

Ratzeburg

Forster

Ratzeburg

distribution

Germany

revision

(year)

single record

(year)

2001

1841

2001

1841

2001

1841

2001

©ZFMK

family

Pteromalidae

species

Pteromalus hieracii

Pteromalus honestus

Pteromalus hyalopterus

Pteromalus hypocyaneus

Pteromalus ignobilis

Pteromalus illustratus

Pteromalus immundus

Pteromalus impressifrons

Pteromalus inanis

Pteromalus incertus

Pteromalus inclytus

Pteromalus inconspicuus

Pteromalus inermis

Pteromalus infelix

Pteromalus infestus

Pteromalus infinitus

Pteromalus inquilinus

Pteromalus insignis

Pteromalus intermedius

Pteromalus jejunus

Pteromalus laetus

Pteromalus laevis

Pteromalus laricinellae

Pteromalus latreillei

Pteromalus lazulinus

Pteromalus lepidotus

Pteromalus leptogaster

Pteromalus leptostictus

Pteromalus limbatus

Pteromalus lineolatus

Pteromalus lugens

Pteromalus lutulentus

Pteromalus macrocerus

Pteromalus maculiscapus

Pteromalus mandibulatus

Pteromalus mariae

Pteromalus maurus

Pteromalus melancholicus

Pteromalus melanocerus

Pteromalus melanochlorus

Pteromalus microneurus

Bonn zoological Bulletin 67 (2): 101-107

author

(Thomson)

Forster

Dalla Torre

Forster

Dalla Torre

Forster

(Walker)

Forster

Ratzeburg

Forster

Ratzeburg

Forster

Dalla Torre

Forster

Dalla Torre

Ratzeburg

Dalla Torre

Forster

Ratzeburg

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna

(APPENDIX 1)

distribution

Germany

Ml

single record

(year)

2001

1841

2001

1848

2001

1841

2001

1844

©ZFMK

38

family

Pteromalidae

species

Pteromalus microps

Pteromalus micros

Pteromalus mixtus

Pteromalus mobilis

Pteromalus molestus

Pteromalus monochrous

Pteromalus musaeus

Pteromalus nanulus

Pteromalus napaeus

Pteromalus naucus

Pteromalus navis

Pteromalus nebulosus

Pteromalus neesii

Pteromalus neglectus

Pteromalus nigricans

Pteromalus nobilis

Pteromalus nodulosus

Pteromalus nuperus

Pteromalus obductus

Pteromalus obscurus

Pteromalus obvolitans

Pteromalus ochrocerus

Pteromalus opacus

Pteromalus opimus

Pteromalus ornatus

Pteromalus pachygaster

Pteromalus pachymerus

Pteromalus papaveris

Pteromalus parietinae

Pteromalus patro

Pteromalus pellucidiventris

Pteromalus pellucidus

Pteromalus picinus

Pteromalus pilosellus

Pteromalus planiusculus

Pteromalus platyphilus

Pteromalus pogonochoeri

Pteromalus polychlori

Pteromalus polycyclus

Pteromalus pomacearum

Bonn zoological Bulletin 67 (2): 101-107

Maximilian Weber et al.

author

(Graham)

Dalla Torre

Forster

Walker

Dalla Torre

Forster

Ratzeburg

Dalla Torre

Ratzeburg

Forster

Ratzeburg

Forster

Nees

Forster

(Thomson)

Forster

(Graham)

Walker

Ratzeburg

Forster

Walker

Ratzeburg

Forster

Ratzeburg

distribution

Germany

revision

(year)

single record

(year)

2001

1848

2001

1841

2001

2005

2001

©ZFMK

family

Pteromalidae

species

Pteromalus praeceps

Pteromalus praelongus

Pteromalus praepes

Pteromalus praepotens

Pteromalus princeps

Pteromalus propinquus

Pteromalus psyllus

Pteromalus pulcherrimus

Pteromalus pullus

Pteromalus punctum

Pteromalus pungens

Pteromalus puparum

Pteromalus pusillus

Pteromalus pygmaeanae

Pteromalus pygmaeus

Pteromalus questionis

Pteromalus racemosi

Pteromalus ramulorum

Pteromalus rapax

Pteromalus ratzeburgii

Pteromalus regius

Pteromalus relevatus

Pteromalus rhombicus

Pteromalus saltatorius

Pteromalus sapphireus

Pteromalus scandiae

Pteromalus semotus

Pteromalus sequester

Pteromalus similis

Pteromalus simplex

Pteromalus sincerus

Pteromalus singularis

Pteromalus smaragdinus

Pteromalus solidus

Pteromalus sonchi

Pteromalus sparsus

Pteromalus sphaerogaster

Pteromalus splendidus

Pteromalus strobilobius

Pteromalus subaequalis

Pteromalus sublaevis

Bonn zoological Bulletin 67 (2): 101-107

author

Forster

(Linnaeus)

Forster

Ratzeburg

Forster

Ratzeburg

Forster

Dalla Torre

Forster

(Graham)

(Walker)

Walker

Forster

Janzon

Forster

Ratzeburg

Forster

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna

(APPENDIX 1)

distribution

Germany

39

single record

(year)

2001

1844

2001

1852

2001

1993

2001

1852

2001

©ZFMK

40

family

Pteromalidae

species

Pteromalus subniger

Pteromalus subpunctatus

Pteromalus subterraneus

Pteromalus sulphuripes

Pteromalus sybarita

Pteromalus sylvarum

Pteromalus syntomus

Pteromalus terebrans

Pteromalus tessellatus

Pteromalus tibialis

Pteromalus timidus

Pteromalus tricollis

Pteromalus tripolii

Pteromalus troglodytes

Pteromalus unicolor

Pteromalus uyari

Pteromalus vaginatus

Pteromalus vaginulae

Pteromalus vallatus

Pteromalus vallecula

Pteromalus varians

Pteromalus variolosus

Pteromalus velox

Pteromalus veneris

Pteromalus ventricosus

Pteromalus verticalis

Pteromalus vibulenus

Pteromalus vicarius

Pteromalus vicinus

Pteromalus violarum

Pteromalus viridicans

Pteromalus vorax

Rakosina deplanata

Rhaphitelus ladenbergii

Rhaphitelus maculatus

Rhicnocoelia constans

Rhopalicus atricornis

Rhopalicus guttatus

Rhopalicus magdalis

Rhopalicus opisthotomus

Rhopalicus quadratus

Bonn zoological Bulletin 67 (2): 101-107

Maximilian Weber et al.

author

Forster

Ratzeburg

Forster

Ratzeburg

Nees

Dalla Torre

Forster

(Graham)

Dalla Torre

Forster

Ozdikmen

Forster

Ratzeburg

Forster

Ratzeburg

(Spinola)

Forster

Dalla Torre

Forster

(Walker)

Ratzeburg

Forster

Dalla Torre

Forster

Bouéek

(Ratzeburg)

Walker

(Walker)

(Forster)

(Ratzeburg)

distribution

Germany

revision

(year)

1979

1989

single record

(year)

1841

2001

1852

2001

1852

2001

1852

2001

©ZFMK

family

Pteromalidae

species

Rhopalicus tutela

Rhopalicus virescens

Roptrocerus mirus

Roptrocerus polychromus

Roptrocerus xylobius

Roptrocerus xylophagorum

Sceptrothelys deione

Sceptrothelys grandiclava

Sceptrothelys intermedia

Sceptrothelys occultus

Schimitschekia populi

Schizonotus latus

Schizonotus sieboldi

Scutellista obscura

Seladerma berani

Seladerma bicolor

Seladerma breve

Seladerma coeruleovirens

Seladerma convexum

Seladerma diffine

Seladerma diutinum

Seladerma geniculatum

Seladerma globosum

Seladerma laetum

Seladerma scaea

Seladerma simplex

Semiotellus diversus

Semiotellus mundus

Semiotellus punctifrons

Semiotellus rujanensis

Spalangia cameroni

Spalangia crassicornis

Spalangia endius

Spalangia erythromera

Spalangia fuscipes

Spalangia nigra

Spalangia nigripes

Spalangia nigroaenea

Spalangia rugulosa

Spalangia subpunctata

Spalangiopelta alata

Bonn zoological Bulletin 67 (2): 101-107

author

(Walker)

(Ratzeburg)

(Walker)

Forster

(Ratzeburg)

(Walker)

Graham

(Forster)

Boucéek

(Walker)

(Ratzeburg)

(Forster)

(Delucchi)

Walker

(Forster)

Walker

(Walker)

(Delucchi)

(Zetterstedt)

(Delucchi)

Walker

(Walker)

(Thomson)

(Walker)

(Nees)

Bouéek

Perkins

Bouéek

Walker

Forster

Nees

Latreille

Curtis

Forster

Boucek

distribution

Germany

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna

(APPENDIX 1)

revision

(year)

1958

2009

4]

single record

(year)

2001

©ZFMK

42

family

Pteromalidae

species

Spaniopus amoenus

Spaniopus dissimilis

Spaniopus peisonis

Sphaeripalpus fuscipes

Sphaeripalpus punctulatus

Sphaeripalpus sericeus

Sphaeripalpus viridis

Sphegigaster clavicornis

Sphegigaster cuscutae

Sphegigaster intersita

Sphegigaster nigricornis

Sphegigaster pallicornis

Sphegigaster pedunculiventris

Sphegigaster truncata

Spintherus dubius

Staurothyreus cruciger

Stenomalina bicolor

Stenomalina communis

Stenomalina dives

Stenomalina epistena

Stenomalina fallax

Stenomalina favorinus

Stenomalina gracilis

Stenomalina iera

Stenomalina illudens

Stenomalina laticeps

Stenomalina liparae

Stenomalina micans

Stenomalina oxygyne

Stenomalina spectabilis

Stenoselma nigrum

Stictomischus gibbus

Stictomischus groschkei

Stictomischus longiventris

Stictomischus miniatus

Stictomischus nitentis

Stictomischus obscurus

Stictomischus scaposus

Stictomischus tumidus

Stinoplus lapsanae

Syntomopus incisus

Bonn zoological Bulletin 67 (2): 101-107

Maximilian Weber et al.

author

Forster

Walker

(Erdos)

(Walker)

(Forster)

(Thomson)

Forster

(Forster)

Ferriére

Graham

(Nees)

(Spinola)

Thomson

(Nees)

Graham

(Forster)

(Nees)

(Walker)

(Forster)

(Walker)

(Giraud)

(Olivier)

(Walker)

(Forster)

Delucchi

(Walker)

Delucchi

Thomson

Delucchi

(Walker)

Thomson

(Walker)

Graham

Thomson

distribution

Germany

revision

(year)

1972

2011

single record

(year)

2001

2003

2001

1999

2001

1841

©ZFMK

family

Pteromalidae

species

Syntomopus incurvus

Syntomopus oviceps

Syntomopus thoracicus

Systasis encyrtoides

Systasis tenuicornis

Termolampa pinicola

Theocolax elegans

Theocolax formiciformis

Thinodytes cyzicus

Tomicobia pityophthori

Tomicobia seitneri

Toxeuma acilius

Toxeuma fuscicorne

Trichomalopsis acuminata

Trichomalopsis caesareus

Trichomalopsis exigua

Trichomalopsis fucicola

Trichomalopsis germanica

Trichomalopsis hemiptera

Trichomalopsis microptera

Trichomalopsis peregrina

Trichomalopsis potatoriae

Trichomalopsis punctata

Trichomalopsis tigasis

Trichomalus annulatus

Trichomalus apertus

Trichomalus bracteatus

Trichomalus campestris

Trichomalus cinctus

Trichomalus conifer

Trichomalus coryphe

Trichomalus cristatus

Trichomalus elongatus

Trichomalus exquisitus

Trichomalus flagellaris

Trichomalus frontalis

Trichomalus fulgidus

Trichomalus fulvipes

Trichomalus generalis

Trichomalus germanus

Trichomalus glabellus

Bonn zoological Bulletin 67 (2): 101—107

(APPENDIX 1)

author

Walker

Thomson

Walker

Bouéek

(Westwood)

Westwood

(Walker)

(Bouéek)

(Ruschka)

(Walker)

Walker

(Graham)

(Dalla Torre)

(Walker)

(Graham)

(Walker)

(Lindeman)

(Graham)

(Ratzeburg)

(Walker)

(Forster)

(Walker)

(Forster)

(Walker)

(Forster)

Delucchi & Graham

(Forster)

Graham

(Thomson)

(Forster)

(Walker)

(Forster)

(Della Torre)

(Forster)

distribution

Germany

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna

revision

(year)

2007

43

single record

(year)

2001

1841

2001

©ZFMK

44

family

Pteromalidae

Signiphoridae

Tanaostigmatidae

species

Trichomalus gynetelus

Trichomalus helvipes

Trichomalus inscitus

Trichomalus intestinarius

Trichomalus lepidus

Trichomalus lonchaeae

Trichomalus lucidus

Trichomalus nanus

Trichomalus notabilis

Trichomalus obsessorius

Trichomalus perfectus

Trichomalus pexatus

Trichomalus pilosus

Trichomalus posticus

Trichomalus repandus

Trichomalus robustus

Trichomalus rufinus

Trichomalus rugosus

Trichomalus statutus

Trichomalus tenellus

Tricolas xylocleptis

Trigonoderus bimaculatus

Trigonoderus cyanescens

Trigonoderus filatus

Trigonoderus immaculatus

Trigonoderus occultus

Trigonoderus princeps

Trigonoderus pulcher

Tritneptis affinis

Tritneptis diprionis

Tritneptis klugii

Urolepis maritima

Urolepis rufipes

Vrestovia fidenas

Xestomnaster chrysochlorus

Xestomnaster mazares

Xiphydriophagus meyerinckii

Yusufia acerina

Chartocerus subaeneus

Thysanus ater

Tanaostigmodes megalarus

Bonn zoological Bulletin 67 (2): 101-107

Maximilian Weber et al.

author

(Walker)

(Forster)

Boutéek

(Walker)

(Forster)

(Walker)

(Ratzeburg)

(Walker)

Delucchi & Graham

(Forster)

(Walker)

Boucéek

(Nees)

(Forster)

Walker

(Nees)

(Forster)

Westwood

Walker

(Nees)

Gahan

(Ratzeburg)

(Walker)

(Ashmead)

(Walker)

(Ratzeburg)

(Bouéek)

(Forster)

Walker

(Walker)

distribution

Germany

revision

(year)

2007

1993

single record

(year)

2001

2005

©ZFMK

family

Tetracampidae

Torymidae

species

Dipriocampe diprioni

Epiclerus nomocerus

Epiclerus panyas

Epiclerus temenus

Foersterella erdoesi

Foersterella reptans

Platynocheilus cuprifrons

Tetracampe impressa

Bootanomyia bohemianii

Bootanomyia dorsalis

Bootanomyia stigmatizans

Cryptopristus caliginosus

Eridontomerus laticornis

Eridontomerus syrphi

Glyphomerus stigma

Glyphomerus tibialis

Idiomacromerus papaveris

Idiomacromerus terebrator

Megastigmus aculeatus

Megastigmus atedius

Megastigmus bipunctatus

Megastigmus brevicaudis

Megastigmus pictus

Megastigmus pinus

Megastigmus rosae

Megastigmus spermotrophus

Megastigmus strobilobius

Megastigmus suspectus

Microdontomerus annulatus

Monodontomerus aeneus

Monodontomerus aereus

Monodontomerus dentipes

Monodontomerus laricis

Monodontomerus minor

Monodontomerus obscurus

Monodontomerus vicicellae

Podagrion pachymerum

Pseudotorymus arvernicus

Pseudotorymus leguminus

Pseudotorymus militaris

Pseudotorymus napi

Bonn zoological Bulletin 67 (2): 101-107

author

(Ferriére)

(Masi)

(Walker)

Boutek

(Nees)

Forster

(Ratzeburg)

(Fabricius)

(Walker)

(Forster)

(Fabricius)

Forster

(Forster)

(Masi)

(Swederus)

Walker

(Swederus)

Ratzeburg

(Forster)

Parfitt

Boutek

Wachtl

Ratzeburg

Borries

(Spinola)

(Fonscolombe)

Walker

(Dalman)

Mayr

(Ratzeburg)

Westwood

(Walker)

Ruschka

(Boheman)

(Amerling & Kirch-

ner)

distribution

Germany

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna

(APPENDIX 1)

revision

(year)

1958

2016

1958

2016

2003

2016

2005

45

single record

(year)

2001

1993

1998

©ZFMK

46

family

Torymidae

species

Pseudotorymus papaveris

Pseudotorymus salicis

Pseudotorymus salviae

Pseudotorymus sapphyrinus

Pseudotorymus tarsatus

Torymoides kiesenwetteri

Torymus abbreviatus

Torymus affinis

Torymus angelicae

Torymus approximatus

Torymus argei

Torymus armatus

Torymus arundinis

Torymus associatus

Torymus auratus

Torymus austriacus

Torymus azureus

Torymus basalis

Torymus baudysi

Torymus bedeguaris

Torymus caudatus

Torymus cerri

Torymus chloromerus

Torymus cingulatus

Torymus confinis

Torymus confluens

Torymus cultriventris

Torymus cyaneus

Torymus difficilis

Torymus druparum

Torymus eglanteriae

Torymus epilobii

Torymus erucarum

Torymus fagi

Torymus fagineus

Torymus fastuosus

Torymus flavipes

Torymus formosus

Torymus fuscicornis

Torymus galii

Torymus geranii

Bonn zoological Bulletin 67 (2): 101-107

Maximilian Weber et al.

author

(Thomson)

Ruschka

(Fonscolombe)

(Nees)

(Mayr)

Boheman

(Fonscolombe)

(Walker)

Forster

Boutéek

Boheman

(Walker)

Forster

(Miller)

Graham

Boheman

(Walker)

Boutek

(Linnaeus)

Boheman

(Mayr)

(Walker)

Nees

(Walker)

Ratzeburg

Walker

Nees

Boheman

Mayr

Graham & Gijswijt

(Schrank)

(Hoffmeyer)

Graham

Boheman

(Walker)

Boheman

(Walker)

distribution

Germany

revision

(year)

1998

single record

(year)

2001

1841

2001

1852

1834

2001

©ZFMK

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna

family

Torymidae

Trichogrammatidae

species

Torymus heyeri

Torymus igniceps

Torymus impar

Torymus kaltenbachi

Torymus laetus

Torymus longicalcar

Torymus microcerus

Torymus microstigma

Torymus micrurus

Torymus minutus

Torymus nitidulus

Torymus nobilis

Torymus notatus

Torymus pascuorum

Torymus pastinacae

Torymus persicariae

Torymus phillyreae

Torymus poae

Torymus quercinus

Torymus resinanae

Torymus roboris

Torymus rosariae

Torymus rubi

Torymus ruschkai

Torymus scutellaris

Torymus speciosus

Torymus spinosus

Torymus tanaceticola

Torymus tipulariarum

Torymus varians

Torymus ventralis

Aphelinoidea bischoffi

Asynacta exigua

Chaetostricha walkeri

Chaetostrichella pungens

Lathromeris danica

Lathromeris germanica

Lathromeris scutellaris

Oligosita foersteri

Oligosita subfasciata

Ophioneurus signatus

Bonn zoological Bulletin 67 (2): 101-107

(APPENDIX 1)

author

Wachtl

Mayr

Rondani

Forster

(Walker)

Graham

(Walker)

Bouéek

Forster

(Walker)

Boheman

(Walker)

Bouéek

Graham & Gijswijt

Mayr

Ruschka

(Hoffmeyer)

Boheman

Ratzeburg

(Walker)

Graham & Gijswijt

(Schrank)

(Hoffmeyer)

(Walker)

Boheman

(Kamijo)

Ruschka

Zetterstedt

(Walker)

(Fonscolombe)

(Novicky)

(Nees)

(Forster)

(Mayr)

(Kryger)

(Girault)

Forster

Girault

Westwood

Ratzeburg

distribution

Germany

revision

(year)

1998

47

single record

(year)

2001

1840

2001

1840

2001

2013

1852

1998

2010

2001

1971

2001

1914

©ZFMK

48

family

Trichogrammatidae

Aphelinidae

Azotidae

Chalcididae

species

Poropoea stollwerckii

Prestwichia aquatica

Prestwichia solitaria

Pseudoligosita nigripes

Trichogramma brassicae

Trichogramma cacaeciae

Trichogramma cephalciae

Trichogramma chilonis

Trichogramma dendrolimi

Trichogramma embryophagum

Trichogramma euproctidis

Trichogramma evanescens

Trichogramma minutum

Trichogramma semblidis

Trichogramma zeirapherae

Trichogrammatoidea stammeri

Ufens foersteri

Aphelinus daucicola

Aphelinus flaviventris

Aphelinus humilis

Aphelinus subflavescens

Aphytis diaspidis

Aphytis hispanicus

Centrodora livens

Coccophagus silvestrii

Encarsia lutea

Protaphelinus nikolskajae

Pteroptrix dimidiata

Ablerus celsus

Belaspidia nigra

Brachymeria inermis

Dirhinus hesperidum

Hockeria bifasciata

Hockeria unicolor

Lasiochalcidia dargelasii

Lasiochalcidia guineensis

Lasiochalcidia indescripta

Neochalcis fertoni

Bonn zoological Bulletin 67 (2): 101-107

Maximilian Weber et al.

author

Forster

Lubbock

Ruschka

(Giraud)

Bezdenko

Marchal

Hochmut & Martinek

Ishii

Matsumura

(Hartig)

(Girault)

Westwood

Riley

(Aurivillius)

Walter

(Novicky)

(Kryger)

Kurdjumov

Mercet

(Westwood)

(Howard)

(Mercet)

(Walker)

Compere

(Masi)

(Yasnosh)

Westwood

(Walker)

(Siebold)

(Fonscolombe)

(Rossi)

Walker

(Latreille)

(Steffan)

Boutek

(Kieffer)

revision

distribution (year)

Germany

Germany 2004

Germany 1982

Germany

Germany 1984

Germany 1997

Germany

Germany 2011

Czech Republic, France

Czech Republic, Netherlands

Czech Republic, Netherlands,

France

Austria, France, Netherlands,

Switzerland, Poland

France, Czech Republic

Denmark, Czech Republic,

Austria

Czech Republic, France

Denmark, France

France, Switzerland, Czech

Republic, Poland

Czech Republic, Poland, France

Austria, Belgium, Czech Repub-

lic, France, Switzerland

Czech Republic, France, Austria

France, Czech Republic, Austria

Austria, France, Czech Republic

Austria, Czech Republic, France,

Netherlands, Poland

France, Austria, Czech Republic

Czech Republic, France

France, Czech Republic, France

single record

(year)

2001

1914

2008

2001

2011

©ZFMK

family

Chalcididae

Encyrtidae

species

Neochalcis osmicida

Proconura nigripes

Psilochalcis subaenea

Achalcerinys lindus

Ageniaspis citricola

Anagyrus aligarhensis

Anagyrus bohemanni

Anagyrus Bouceki

Anthemus funicularis

Anthemus leucaspidis

Arrhenophagus chionaspidis

Asitus phragmitis

Bothriothorax aralius

Bothriothorax serratellus

Cerchysiella planiscutellum

Cheiloneurus boldyrevi

Cheiloneurus submuticus

Comperiella bifasciata

Copidosoma anceus

Copidosoma ancharus

Copidosoma arenarium

Copidosoma bolivari

Copidosoma charon

Copidosoma cyaneum

Copidosoma fadus

Copidosoma gloriosum

Copidosoma ratzeburgi

Copidosoma subalbicorne

Copidosoma tibiale

Dicarnosis helena

Discodes trjapitzini

Ectroma arenarium

Ginsiana carpetana

Globulencyrtus politus

Bonn zoological Bulletin 67 (2): 101-107

author

(Saunders)

(Fonscolombe)

(Masi)

(Mercet)

Logvinovskaya

Agarwal & Alam

(Westwood)

Hoffer

(Bakkendorf)

Mercet

Aurivillius

(Ferriére)

(Walker)

(Dalman)

(Mercet)

Trjapitzin & Agekyan

Thomson

Howard

(Walker)

Erd6és

Mercet

Guerrieri & Noyes

Hoffer

(Walker)

(Mercet)

Mercet

(Hoffer)

Hoffer

Herthevtzian

(Erdos)

(Mercet)

(Hoffer)

distribution

Czech Republic, France, Swit-

zerland

France, Czech Republic

Czech Republic, France

Czech Republic, France, Nether-

lands

France, Poland

Czech Republic, France

Austria, Czech Republic, France,

Netherlands, Poland

Czech Republic, Netherlands

Czech Republic, Denmark

France, Poland

Czech Republic, France, Poland,

Switzerland

Czech Republic, France

Denmark, Netherlands

Czech Republic, Denmark,

Netherlands, Switzerland

Czech Republic, France, Nether-

lands

Czech Republic, France, Nether-

lands

Czech Republic, Denmark

Czech Republic, France

Denmark, France, Netherlands,

Switzerland

Czech Republic, France, Nether-

lands

Czech Republic, France

Austria, Czech Republic, France

Czech Republic, Denmark,

France

Czech Republic, France

Austria, Czech Republic, France

Czech Republic, France

Czech Republic, Denmark,

Netherlands

Austria, Czech Republic, France,

Netherlands

Austria, Czech Republic, France

Czech Republic, France, Austria

Denmark, Austria, Czech Re-

public

Czech Republic, Denmark, Fran-

ce, Netherlands, Poland

Czech Republic, France

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna

(APPENDIX 1)

revision

(year)

49

single record

(year)

©ZFMK

50

family

Encyrtidae

Eulophidae

species

Homalotyloidea erginus

Homalotyloidea nowickyi

Hoplopsis minuta

Isodromus flaviscutum

Lyka submetallica

Metaphycus ater

Metaphycus dispar

Metaphycus flavovarius

Metaphycus flavus

Metaphycus hageni

Metaphycus hanstediensis

Metaphycus nadius

Metaphycus pappus

Metaphycus petitus

Microterys cedrenus

Microterys jalysus

Microterys madyes

Microterys matritensis

Microterys subcupratus

Microterys triozae

Mohelniella silhavyi

Moraviella inexpectata

Ooencyrtus pitvocampae

Parasauleia trjapitzini

Paratetracnemoidea malenotti

Prochiloneurus bolivari

Pseudococcobius obenbergeri

Psyllaephagus lusitanicus

Syrphophagus annulipes

Syrphophagus ariantes

Syrphophagus quercicola

Trechnites alni

Trechnites flavipes

Xanthoectroma aquilinum

Achrysocharoides butus

Apotetrastichus postmarginalis

Bonn zoological Bulletin 67 (2): 101-107

Maximilian Weber et al.

author

(Walker)

Hoffer

(Fabricius)

Hoffer & Trjapitzin

Mercet

(Mercet)

(Howard)

Daane & Caltagirone

Bakkendorf

(Walker)

(Mercet)

(Dalman)

(André)

Hoffer

(Mercet)

Hoffer

(Mercet)

Mercet

(Novickij)

(Mercet)

(Thomson)

(Walker)

(Hoffer)

Erdés

(Mercet)

Mercet

(Walker)

(Bouéek)

revision

distribution (year)

Czech Republic, France, Nether-

lands

Czech Republic, Netherlands

Czech Republic, France, Nether-

lands

Czech Republic, France

Austria, Czech Republic, Den-

mark, France

Czech Republic, France

Czech Republic, France, Nether-

lands

France, Czech Republic

Denmark, France

Czech Republic, Denmark

Poland, Czech Republic, France,

Netherlands

Czech Republic, Denmark,

Netherlands

Czech Republic, Denmark

France, Denmark

Czech Republic, Denmark

Denmark, Czech Republic

Czech Republic, France

Czech Republic, Denmark

Czech Republic, France

Czech Republic, Denmark

France, Poland

Czech Republic, France

Austria, Czech Republic, France,

Poland

Czech Republic, Denmark

Czech Republic, Netherlands

France, Czech Republic, France

Czech Republic, Netherlands

Czech Republic, Denmark,

Netherlands

Czech Republic, Denmark,

France, Poland

Czech Republic, France

Czech Republic, Netherlands,

Poland

Czech Republic, France, Austria

single record

(year)

©ZFMK

family

Eulophidae

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna

species

Aprostocetus agrus

Aprostocetus amenon

Aprostocetus apiculatus

Aprostocetus arrabonicus

Aprostocetus capitigenae

Aprostocetus celtidis

Aprostocetus cerricola

Aprostocetus cyniphidum

Aprostocetus dauci

Aprostocetus distichus

Aprostocetus domenichinii

Aprostocetus epilobii

Aprostocetus eurytus

Aprostocetus fonscolombei

Aprostocetus forsteri

Aprostocetus menius

Aprostocetus oreophilus

Aprostocetus orestes

Aprostocetus planiusculus

Aprostocetus polygoni

Aprostocetus rhipheus

Aprostocetus rufus

Aprostocetus setosulus

Aprostocetus subanellatus

Aprostocetus suevius

Aprostocetus tenuiradialis

Aprostocetus tymber

Aprostocetus viridescens

Aprostocetus viridinitens

Aprostocetus westwoodii

Aprostocetus xanthomelas

Aulogymnus testaceoviridis

Aulogymnus trilineatus

Baryscapus berhidanus

Baryscapus elasmi

Bonn zoological Bulletin 67 (2): 101-107

(APPENDIX 1)

author

(Walker)

Graham

(Erdos)

Graham

(Erdos)

(Ratzeburg)

Graham

(Erdos)

Graham

(Walker)

Graham

(Walker)

(Forster)

Graham

(Thomson)

(Erdos)

(Walker)

(Bakkendorf)

Graham

(Walker)

Graham

(Walker)

(Forster)

Graham

(Fonscolombe)

Graham

(Erdos)

(Mayr)

Erdés

(Graham)

revision

distribution (year)

Czech Republic, France, Nether-

lands

Czech Republic, Netherlands

Czech Republic, France

Czech Republic, Netherlands

Austria, Netherlands

Czech Republic, France, Nether-

lands

Austria, Belgium, Czech Repub-

lic, France

Austria, Netherlands

Czech Republic, France

Czech Republic, Netherlands

Czech Republic, France, Nether-

lands

Czech Republic, France

Austria, Czech Republic, France,

Switzerland, Austria

Czech Republic, France

Czech Republic, France, Swit-

zerland

Czech Republic, France

Czech Republic, Netherlands

Czech Republic, Denmark,

France, Netherlands

Czech Republic, France

Czech Republic, France, Nether-

lands

Czech Republic, Netherlands

Czech Republic, France, Nether-

lands

Czech Republic, Netherlands,

France, Poland

Switzerland, France

Czech Republic, France

Austria, Czech Republic, France,

Netherlands, Poland

Czech Republic, France

France, Poland

Sil

single record

(year)

©ZFMK

52

family

Eulophidae

species

Baryscapus fossarum

Baryscapus szocsi

Chrysocharis albicoxis

Chrysocharis amanus

Chrysocharis argyropezae

Chrysocharis collaris

Chrysocharis loranthellae

Chrysocharis paradoxa

Chrysocharis pilicoxa

Elachertus longipetiolus

Elasmus arcuatus

Elasmus platyedrae

Elasmus rufiventris

Entedon hercyna

Entedon nigrini

Entedon pallicrus

Entedon pseudonigritarsis

Entedon sparetus

Entedon squamosus

Hemiptarsenus autonomus

Hemiptarsenus zilahisebessi

Ionympha ochus

Microlycus virens

Neochrysocharis arvensis

Neotrichoporoides cavigena

Neotrichoporoides gordensis

Neotrichoporoides mediter-

raneus

Neotrichoporoides viridima-

culatus

Omphale admirabilis

Omphale coilus

Omphale cornula

Omphale euphorbiae

Omphale isander

Omphale parma

Oomyzus anomalus

Pediobius deplanatus

Pnigalio hirtulus

Bonn zoological Bulletin 67 (2): 101-107

Maximilian Weber et al.

author

Graham

(Erdos)

Erdés

(Walker)

Graham

Erdos

Hansson

(Thomson)

Bouéek

Ferriere

Walker

Boucek

Erdos

Walker

Thomson

(Mercet)

Erdos

(Walker)

Erdos

Graham

(Fullaway)

(Westwood)

(Walker)

Hansson & Shevtsova

(Walker)

Hansson & Shevtsova

Graham

Boucéek

(Erdos)

revision

distribution (year)

Czech Republic, France, Nether-

lands

Czech Republic, France

Czech Republic, France, Nether-

lands, Poland

Czech Republic, Netherlands

Czech Republic, France

Czech Republic, Netherlands

Czech Republic, France

Austria, France, Czech Republic

Czech Republic, France

Czech Republic, Netherlands

Czech Republic, Denmark,

France

Czech Republic, France, Nether-

lands

Czech Republic, Netherlands

Czech Republic, France

Czech Republic, Netherlands

Austria, Czech Republic, France

Czech Republic, France, Poland

Czech Republic, Netherlands

Czech Republic, France

Czech Republic, Denmark

Czech Republic, France

Austria, Czech Republic, France,

Netherlands

Czech Republic, Netherlands,

Poland

Denmark, France, Netherlands

Czech Republic, France, Nether-

lands

Austria, Czech Republic, France

Denmark, France

Czech Republic, France

Czech Republic, Netherlands

Czech Republic, France

single record

(year)

©ZFMK

family

Eulophidae

Eupelmidae

Eurytomidae

Leucospidae

Mymaridae

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna

species

Pnigalio mediterraneus

Pronotalia hungarica

Quadrastichus anysis

Quadrastichus citrinus

Quadrastichus lasiocerus

Quadrastichus pteridis

Quadrastichus thysanotus

Quadrastichus vacuna

Quadrastichus xanthosoma

Sympiesis gyorfii

Tamarixia leptothrix

Tamarixia tremblayi

Tetrastichus agrilocidus

Tetrastichus leocrates

Tetrastichus leptosoma

Tetrastichus pachycerus

Tetrastichus sodalis

Calosota aestivalis

Eupelmus falcatus

Eupelmus muellneri

Eupelmus pini

Eupelmus pullus

Reikosiella hungarica

Bruchophagus squamea

Eurytoma gyorfii

Eurytoma laricis

Eurytoma stenostigma

Eurytoma tilicola

Sycophila scorzonerae

Sycophila variegata

Systole bipunctata

Systole hofferi

Leucospis biguetina

Alaptus minimus

Anagrus brocheri

Bonn zoological Bulletin 67 (2): 101-107

(APPENDIX 1)

author

Ferriere & Delucchi

(Erdos)

(Walker)

(Thomson)

(Graham)

Graham

(Forster)

(Walker)

(Graham)

Erdos

Graham

(Domenichini)

Graham

(Walker)

Graham

Curtis

(Nikol’skaya)

Ruschka

Taylor

Ruschka

(Erdos)

(Walker)

Erdos

Yano

Thomson

Hedqvist

(Mayr)

(Curtis)

Erdos

(Kalina)

Jurine

Westwood

Schulz

revision

distribution (year)

France, Austria

Czech Republic, France

Czech Republic, France, Austria

Czech Republic, France, Nether-

lands

Austria, Netherlands

Czech Republic, Switzerland

Czech Republic, France, Nether-

lands, Poland, Switzerland,

Austria

Czech Republic, Netherlands

Austria, Czech Republic, France,

Poland

Czech Republic, Netherlands

Czech Republic, Netherlands,

Poland

Denmark, France, Netherlands

Czech Republic, France

Czech Republic, France, Nether-

lands, Poland

Czech Republic, France, Swit-

zerland

France, Czech Republic

Czech Republic, Netherlands,

Austria, France, Poland

Austria, Czech Republic,

Netherlands, Poland

Czech Republic, France

France, Czech Republic, France

Czech Republic, France

Poland, Netherlands

Czech Republic, France

Austria, Czech Republic, France

France, Czech Republic, Austria,

Netherlands, Switzerland

Czech Republic, France

Austria, Czech Republic, France,

Switzerland

Netherlands, Switzerland

Austria, Belgium, Denmark,

Netherlands, Switzerland

53

single record

(year)

©ZFMK

54

family

Mymaridae

Ormyridae

Perilampidae

Pteromalidae

species

Anaphes declinatus

Anaphes pallidus

Camptoptera franciscae

Cleruchus taktochno

Cosmocomoidea tremulae

Erythmelus gracilis

Polynema altitudine

Polynema quadruplex

Ormyrus wachtli

Perilampus eximius

Ablaxia parviclava

Ablaxia squamifera

Acrocormus semifasciatus

Anogmus laricis

Apelioma pteromalinum

Apsilocera verticillata

Arthrolytus slovacus

Bairamlia fuscipes

Caenocrepis arenicola

Chlorocytus pilosus

Chlorocytus planus

Cleonymus brevis

Cleonymus obscurus

Cyclogastrella clypealis

Dinotoides tenebricus

Ecrizotes filicornis

Erythromalus nubilipennis

Erythromalus rufiventris

Gastrancistrus acutus

Gastrancistrus amaboeus

Gastrancistrus clavatus

Gastrancistrus dispar

Gastrancistrus hamillus

Gastrancistrus puncticollis

Gastrancistrus vagans

Glyphognathus laevis

Bonn zoological Bulletin 67 (2): 101-107

Maximilian Weber et al.

author

(Soyka)

(Debauche)

Triapitsyn

(Bakkendorf)

(Howard)

(Soyka)

Mayr

Masi

(Thomson)

Thomson

Bouéek

(Thomson)

Bouéek

Graham

Waterston

(Thomson)

Graham

(Walker)

Boucek

Walker

Boucéek

(Walker)

(Thomson)

(Walker)

Walker

(Thomson)

Graham

Walker

(Thomson)

Westwood

(Delucchi)

revision

distribution (year)

Netherlands, Poland, France

Czech Republic, Netherlands

Austria, Belgium

Belgium, Denmark, Poland

Denmark, Netherlands

France, Poland

Austria, Netherlands

Netherlands, Austria

Austria, Czech Republic, France

Czech Republic, France

Czech Republic, Netherlands,

France

Belgium, Czech Republic,

Netherlands

Czech Republic, Netherlands,

Poland, Switzerland

Austria, Netherlands, Poland

Belgium, Czech Republic

Czech Republic, Netherlands

Netherlands, Switzerland, Fran-

ce, Czech Republic

Austria, Czech Republic, France

Czech Republic, Netherlands

Netherlands, Czech Republic

Czech Republic, France, Swit-

zerland

Czech Republic, France

Belgium, Czech Republic, Fran-

ce, Switzerland

Czech Republic, Belgium,

Netherlands

Czech Republic, Netherlands

Austria, Czech Republic,

Netherlands

Belgium, Czech Republic,

Netherlands

Czech Republic, Netherlands

Belgium, Czech Republic

Czech Republic, Netherlands

Belgium, Czech Republic,

France

Czech Republic, Netherlands

single record

(year)

©ZFMK

family

Pteromalidae

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna

species

Gyrinophagus aper

Halticoptera hippeus

Heteroprymna longicornis

Holcaeus compressus

Holcaeus gracilis

Homoporus pulchripes

Kaleva corynocera

Lamprotatus annularis

Lamprotatus crassipes

Lamprotatus novickyi

Lonchetron fennicum

Macroglenes paludum

Macromesus amphiretus

Melancistrus mucronatus

Mesopolobus aspilus

Mesopolobus maculicornis

Mesopolobus pinus

Mesopolobus prasinus

Mesopolobus trasullus

Mesopolobus zetterstedtii

Micradelus acutus

Micradelus rotundus

Norbanus calabrus

Norbanus cerasiops

Norbanus meridionalis

Notanisus versicolor

Novitzkyanus cryptogaster

Panstenon agylla

Pezilepsis dentifera

Platygerrhus dolosus

Platygerrhus ductilis

Platygerrhus longigena

Platygerrhus millenius

Platygerrhus unicolor

Bonn zoological Bulletin 67 (2): 101-107

(APPENDIX 1)

author

(Walker)

Erd6és

Graham

(Walker)

Thomson

(Delucchi)

Graham

(Graham)

Walker

(Thomson)

(Walker)

(Giraud)

Hussey

(Walker)

(Dalla Torre)

Graham

Walker

(Masi)

Walker

Boutek

(Walker)

(Thomson)

(Walker)

Graham

Szczepanski

Graham

revision

distribution (year)

Austria, Belgium, Czech Re-

public

Belgium, Czech Republic

Czech Republic, Netherlands

Belgium, Czech Republic,

Netherlands

Czech Republic, Belgium,

Netherlands

Czech Republic, France, Swit-

zerland

Belgium, Czech Republic

Belgium, Czech Republic,

Netherlands

Czech Republic, Netherlands,

Austria

Netherlands, Austria, Switzer-

land

Czech Republic, France, Poland

Czech Republic, Netherlands,

Switzerland

Czech Republic, Denmark,

France, Poland

Czech Republic, Netherlands

Czech Republic, Denmark

Austria, Czech Republic, France

Denmark, Netherlands, Poland

Austria, Netherlands

Belgium, France, Czech Repub-

lic, Switzerland

Czech Republic, Belgium,

Poland

Belgium, Czech Republic,

Netherlands

Czech Republic, Netherlands

Czech Republic, France

Czech Republic, Netherlands

Czech Republic, Belgium

Czech Republic, France, Nether-

lands, Poland

Czech Republic, Netherlands,

Poland

Czech Republic, France, Nether-

lands

Netherlands, Poland

Czech Republic, Netherlands

55

single record

(year)

©ZFMK

56

family

Pteromalidae

Signiphoridae

Torymidae

species

Plutothrix acuminata

Psilocera concolor

Psilocera confusa

Psilocera crassispina

Pteromalus altus

Pteromalus apum

Pteromalus crassinervis

Pteromalus eudecipiens

Rhicnocoelia grahami

Rhicnocoelia impar

Scutellista caerulea

Seladerma tarsale

Spilomalus quadrinota

Stenomalina fervida

Stichocrepis armata

Stinoplus pervasus

Synedrus transiens

Systasis annulipes

Systasis parvula

Tomicobia promulus

Toxeuma subtruncatum

Trichomalopsis submarginata

Trigonoderus sokanowskii

Trychnosoma punctipleura

Veltrusia rara

Clytina giraudi

Exopristus trigonomerus

Idiomacromerus mayri

Megastigmus milleri

Megastigmus rafni

Megastigmus specularis

Monodontomerus rugulosus

Bonn zoological Bulletin 67 (2): 101-107

Maximilian Weber et al.

author

(Thomson)

Graham

(Thomson)

(Walker)

(Retzius)

(Thomson)

Ozdikmen

Boucek

(Walker)

(Fonscolombe)

(Walker)

Graham

Forster

(Walker)

Thomson

(Walker)

Graham

(Thomson)

Novicky

(Thomson)

Boucéek

Erdos

(Masi)

(Wachtl)

Milliron

Hoffmeyer

Walley

Thomson

revision

distribution (year)

Belgium, Netherlands, Czech

Republic, France

Czech Republic, Netherlands

Czech Republic, France, Nether-

lands

Belgium, Czech Republic, Fran-

ce, Netherlands

Czech Republic, Belgium, Fran-

ce, Netherlands

Belgium, France, Netherlands,

Czech Republic, Denmark,

Switzerland

Czech Republic, Belgium,

Netherlands

Czech Republic, Netherlands,

Switzerland

Belgium, Czech Republic

Belgium, Netherlands

France, Netherlands, Czech

Republic

Belgium, Czech Republic,

Netherlands

Czech Republic, Netherlands

Belgium, Czech Republic

Austria, Czech Republic,

Netherlands, Switzerland

Czech Republic, France, Nether-

lands

Czech Republic, Netherlands

Belgium, Czech Republic,

Netherlands

Austria, Belgium, Czech Re-

public

Czech Republic, Belgium,

Netherlands

Czech Republic, Netherlands

Denmark, Switzerland

Czech Republic, France, Nether-

lands

Czech Republic, Netherlands

Czech Republic, Denmark

Czech Republic, France

Austria, Czech Republic, France

Belgium, Denmark, France,

Netherlands

Belgium, Denmark, France,

Netherlands

Denmark, France, Netherlands

France, Austria, Czech Republic,

Netherlands

single record

(year)

©ZFMK

Reliability, completeness and improvement of our knowledge on Germany’s parasitoid wasp fauna

family

Torymidae

Trichogrammatidae

species

Podagrion splendens

Pseudotorymus krygeri

Torymus aceris

Torymus acrophilae

Torymus amurensis

Torymus anthobiae

Torymus calcaratus

Torymus chlorocopes

Torymus corni

Torymus cupreus

Torymus curticauda

Torymus curtisi

Torymus euphorbiae

Torymus filipendulae

Torymus giraudianus

Torymus grahami

Torymus hederae

Torymus hylesini

Torymus juniperi

Torymus nigritarsus

Torymus orobi

Torymus pulchellus

Torymus purpurascens

Torymus scaposus

Torymus socius

Torymus stenus

Torymus thymi

Torymus veronicae

Bloodiella andalusica

Chaetostricha dimidiata

Chaetostricha doricha

Mirufens longicauda

Monorthochaeta nigra

Oligosita collina

Oligosita engelharti

Oligosita impudica

Bonn zoological Bulletin 67 (2): 101-107

(APPENDIX 1)

author

Spinola

Hoffmeyer

Boucek

Ruschka

(Walker)

Ruschka

Nees

Boheman

Mayr

(Spinola)

Graham & Gijswijt

(Walker)

Graham & Gijswijt

(Hoffmeyer)

Boucek

(Walker)

Graham

(Linnaeus)

(Walker)

Mayr

Thomson

(Fabricius)

(Thomson)

Mayr

Graham

Ruschka

Nowicki

Walker

(Walker)

(Blood)

Blood

Walker

Kryger

revision

distribution (year)

Czech Republic, France

Denmark, Netherlands

Czech Republic, France

Austria, Belgium, Czech Re-

public

Austria, Czech Republic, France,

Netherlands

Austria, Netherlands

Belgium, Czech Republic, Fran-

ce, Austria, Netherlands

Czech Republic, Netherlands

Austria, Czech Republic, France

Czech Republic, Austria, Den-

mark, Netherlands

Czech Republic, Netherlands

France, Austria

Czech Republic, Netherlands

France, Austria

Czech Republic, France

Czech Republic, France, Nether-

lands

Czech Republic, France, Nether-

lands

France, Austria, Czech Republic

Czech Republic, France, Nether-

lands, Austria

France, Austria

Czech Republic, Netherlands

France, Czech Republic

Czech Republic, Netherlands

Austria, Czech Republic, France

Czech Republic, Netherlands

Czech Republic, Denmark

Austria, Belgium

France, Poland

Czech Republic, Denmark

Czech Republic, France, Poland

Czech Republic, France

Czech Republic, Denmark,

Poland

Netherlands, Czech Republic,

Denmark

57

single record

(year)

©ZFMK

58 Maximilian Weber et al.

BHL

i

Blank Page Digitally Inserted

Bonn zoological Bulletin 67 (2): 109-116

2018 - Behr N. & Rédder D.

https://doi.org/10.20363/BZB-2018.67.2.109

ISSN 2190-7307

http://www.zoologicalbulletin.de

Research article

urn:|sid:zoobank.org:pub:939B24A9-ED39-4CF6-82D7-EB7E5933343C

Larval development stages and husbandry

of the Rice Frog Microhyla mukhlesuri Hasan et al., 2014

(Anura: Microhylidae)

Nils Behr' & Dennis Rédder**

'? Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany

“Corresponding author: E-mail: d.roedder@leibniz-zfmk.de

'urn:lsid:zoobank.org:author:5A BSOEE2-D5BD-4A B9-90CE-426B4745681E

> urn:Isid:zoobank.org: author: 1 79FOE77-9745-48EE-A 81 1-FD00627CC994

Abstract. We describe captive management and larval development of Microhyla mukhlesuri, a recently described

microhylid frog from Bangladesh, southern Yunnan, Thailand, Laos, and Vietnam, at the scientific animal keeping facility

of the Zoological Research Museum Alexander Koenig (ZFMK). Beginning at Gosner stage 25, for each larval stage

detailed characteristics are provided and additionally developmental time is compared to other members of the genus

Microhyla. Herein, we present first observations on captive reproduction of the species.

Key words. Conservation breeding, larval staging, tadpole morphology.

INTRODUCTION

The genus Microhyla Tschudi, 1838 currently comprises

41 species of small ground-dwelling frogs which are com-

monly referred to as rice frogs (Frost 2018). The group 1s

widely distributed across Asia, occurring from the Japa-

nese Ryukyu Islands and China to the north, across In-

dia, Sri Lanka and South-east Asia to the islands Suma-

tra, Borneo, Java, and Bali in the southeast (Frost 2018).

Morphological characteristics comprise a generally small

body size and a narrow mouth, a brown to reddish dorsal

coloration with variable dark markings, smooth skin on

dorsum, absence of vomerine teeth and paratoid glands,

fingers without webbing and a hidden tympanum covered

with skin (Poyarkov et al. 2014; Seshradi et al. 2016).

Microhyla mukhlesuri Hasan, Islam, Kuramoto, Ku-

rabayashi & Sumida, 2014 was recently separated from

its sister taxon M. fissipes Boulenger, 1884 and is not

yet listed by the IUCN Red List of Threatened Species

(Hasan et al. 2014; IUCN 2018). Yuan et al. (2016) sug-

gested that MZ mukhlesuri is distributed in Bangladesh,

southern Yunnan, Thailand, Laos, and Vietnam and thus

suppose that it also occurs in Myanmar and Cambodia,

resulting in a wide distribution in Southeast Asia. For

a few microhylid species of the genus Microhyla infor-

mation on breeding ecology, captive management, and

also larval staging tables are available (e.g., Shimizu &

Ota 2003; Narzary & Bordoloi 2013; Wang et al. 2017).

There 1s still a lack of information on the ecology and life

history, including larval development, of MZ mukhlesuri.

Received: 27.04.2018

Accepted: 25.10.2018

Although this species might currently not be threatened

by extinction, there are at least three species in the genus

Microhyla that are currently listed as Endangered (..e.,

M. pulchella Poyarkov, Vassilieva, Orlov, Galoyan, Tran,

Le, Kretova & Geissler, 2014, M. sholigari Dutta & Ray,

2000 and M. zeylanica Parker & Osman-Hill, 1949) and

one even as Critically Endangered (1.e., M. karunaratnei

Fernando & Siriwardhane, 1996) by the respective IUCN

Red List accounts (Biju et al. 2004; Manamendra-Arach-

chi & de Silva 2004a; Manamendra-Arachchi & de Silva

2004b; IUCN SSC Amphibian Specialist Group 2017).

Therefore, captive management and information on lar-

val development of M. mukhlesuri presented in this paper

might be used analogously for more threatened closely

related species.

Herein, we describe different tadpole stages of Micro-

hyla mukhlesuri for the first time and present our captive

management methods for this species in the scientific an-

imal keeping facility of the Zoological Research Museum

Alexander Koenig (ZFMK), Bonn, Germany. Further-

more, we documented the tadpoles’ body surface every

two to three days to examine general growth within the

larval stage.

MATERIAL AND METHODS

Species identification. Since Microhyla species are often

cryptic and hence hard to distinguish from each other,

species identification was confirmed by DNA barcoding

Corresponding editor: W. Bohme

Published: 01.11.2018

110

Nils Behr & Dennis R6dder

Fig. 1. Live specimens of Microhyla mukhlesuri at the ZFMK. A: adult frog; B: juvenile frog one week after metamorphosis.

Table 1. Developmental data of the fastest developing tadpoles. Stage: developmental stage according to Gosner (1960);

Age: number of days after hatching; Diagnostic features: characteristic features of the respective stage; L: length, D: diameter.

Stage Age [d]

25

26

27

28

29

30

31

32

33

34

35-36

37

38-39

40

41

42

43

44

45

46

1

Diagnostic features

Body nearly transparent, single pigment cells visible on whole body, wide mouth with obvious

mouthparts, well developed eyes, closed operculum covers gills, spiracle forms ventrally on

left side

Hind limb buds start to develop (L < % D)

Further development of hind limb buds (L > % D)

Further development of hind limb buds (L > D)

Further development of hind limb buds (L > 1.5 D)

Further development of hind limb buds (L > 2 D), limbs become slightly bent

Foot paddle starts to develop

First slight indentation on foot paddle visible (between toes 4 and 5)

Second indentation on foot paddle visible (between toes 3 and 4)

Third indentation on foot paddle visible (between toes 2 and 3)

Fourth indentation on foot paddle visible (between toes 1 and 2)

All toes completely separated

Metatarsal tubercle and subarticular tubercle appear

Foot tubercles, toe pads completely developed

Forelimbs visible under the transparent skin, atrophy of the mouthparts begins

Forelimbs emerge, mouth positioned anterior to nostril

Mouth angle between nostril and eye, tail begins to atrophy

Mouth angle beneath eye, tail already strongly reduced

Mouth angle posterior to eye, only a tail stub 1s left; coloration and pattern slightly developed

Tail completely resorbed, metamorphosis completed; development of coloration and pattern

completed

Larval development stages and husbandry of the Rice Frog Microhyla mukhlesuri 111

using a fragment of the mitochondrial 16S rRNA. Se-

quences were obtained as described in Koch et al. (2013).

The final sequence (GenBank Accession MH232034)

was compared with sequences of Microhyla species

available in GenBank and following the definition of

M. mukhlesuri as proposed by Yuan et al. (2016).

Captive management of adult frogs. A group of 20 adult

Microhyla mukhlesuri (Fig. 1A) originating from Viet-

nam was purchased from a commercial importer in 2017

and housed in a terrarium measuring 60 x 60 x 40 cm

(1 x w x h) in the scientific animal keeping facility of

the ZFMK. The terrarium was filled with remineralized

osmosis water up to a depth of about 5 cm. One half

of the bottom was covered with a filter pad measuring

40 x 20 x 6 cm as land part. Leaf litter (mainly Fagus syl-

vatica and Quercus robur) was scattered on the filter pad

and in the water part to provide hiding places. Addition-

ally, some live plants (1.e., Elodea sp. and Microsorum

pteropus) were placed in the water. Both air and water

temperatures ranged from 20 °C to 26 °C and humidi-

ty varied between 60% and 80%. For illumination LED

light strips (Solar Stinger 1100mm Sunstrip Dimmable

Driver) were used and the photoperiod was set between

8:00 and 20:00 h. The frogs were fed with young crickets

(Acheta domesticus and Gryllus assimilis) and flightless

fruit flies (Drosophila melanogaster and D. hydei) every

Bonn zoological Bulletin 67 (2): 109-116

a *

Fig. 2. Tadpoles of Microhyla mukhlesuri in Gosner stages 28 to 33. These stages are defined by the growth of hind limbs and

foot paddles. Stage 28: hind limbs just became longer than wide (diameter); Stage 30: hind limbs are at least two times as long as

wide (diameter); Stage 32: a first indentation appears on the foot paddles, which will later separate toes 4 and 5; Stage 33: second

indentation appears on the foot paddles between the developing toes 3 and 4.

two to three days. All prey items were dusted with differ-

ent vitamin and mineral powders (1.e., herpetal Amphib,

herpetal Mineral + Vitamin D3 and herpetal Complete

Terrarium) and furthermore crickets were gut loaded

with fresh vegetables.

Rearing setup for tadpoles. Tadpoles were transferred

into a rearing tank measuring 30 x 30 x 30 cm. This tank

was filled with remineralized osmosis water to a depth of

about 25 cm and an aquarium heater (SERA SE008710,

50W) was used to keep the water temperature at 24 °C

to 25 °C. A few aquatic plants (1.e., E/odea sp. and Mic-

rosorum pteropus) and some dried leaves of Terminalia

catappa and Fagus sylvatica were added to the water as

natural bacteriostatics and fungistatics (Chitmanat et al.

2005). Tadpoles were fed with a mixture of crushed fish

food (Sera® vipan and Tetra Tablets TabiMin) and Spiru-

lina flakes every two to three days. In addition, one stone

that was overgrown with algae was placed in the tank

and was replaced every other day when most of the al-

gae were eaten. Furthermore, Daphnia pulex and differ-

ent aquatic snails (i.e., Physella sp. and Planorbella sp.)

were added to eat possible food remains. As no filtration

was used, half of the water was changed once per week.

The following water parameters were measured in

the rearing tank: NO, < 0.05 mg/l, NO, 7.5 mg/l, NH, <

0.05 mg/l, Cu < 0.1 mg/l, KH 3°dH, GH 5°dH, pH 6.5—

s

©ZFMK

112 Nils Behr & Dennis R6dder

Table 2. Comparative larval development time of four species in the genus Microhyla: M. mukhlesuri, M. fissipes, M. ornata, M.

okinavensis. We adjusted the staging data of the other species to fit the staging system after Gosner (1960) and started counting

from stage 25 onwards. * The species identified as M. ornata by Narzary & Bordoloi (2013) might in fact be M mukhlesuri fol-

lowing Yuan et al. (2016), genetic analyses are necessary; ** Specimens from the Ryukyu Archipelago identified as MZ. ornata by

Shimizu & Ota (2003) have been assigned to M. okinavensis by Matsui et al. (2005).

Age [d]

Stage Microhyla mukhlesuri Microhyla fissipes

(Own data) (modified after

Wang et al. 2017)

25 1 l

26 10 6

27 12 9

28 14 12

29 — 15

30 16 15

31 17 17

32 19 19

33 19 19

34 | 22,

35 24 24

36 24 24

37 27 26

38 — 29

39 — 33

40 30 33

41 32 36

42 33 38

43 35 38

44 35 39

45 36 40

46 37 4]

7.5. Lighting was equivalent to the adult setup. When

the first tadpoles had developed hind legs a piece of cork

bark was added to the aquarium to provide a small land

area for metamorphosed frogs.

Rearing setup for juvenile frogs. Metamorphosed

frogs were transferred into plastic boxes measuring

33 x 21 x 28 cm with fine mesh lids on both narrow

sides. One side of this container was heightened to create

a gradient and a water part with a depth of about 2 cm

on the lower side. One layer of Hygrolon®, an artificial

and highly hygroscopic material originally developed for

cultivating orchids and ferns, was used as ground layer

to keep the air humidity on a high level of about 70% to

80%. Furthermore, a big layer of dry leaves (mainly Fa-

gus sylvatica and Quercus robur) and moss was added to

the box. Temperatures ranged between 21 °C and 26 °C

and illumination was the same as in the adult and tadpole

Bonn zoological Bulletin 67 (2): 109-116

Microhyla ornata* Microhyla okinavensis **

(modified after (modified after

Narzary & Bordoloi 2013) Shimizu & Ota 2003)

1.0 1.0

2.5 5.5

3.0 ES

LS 95

126 11.5

16.5 i Di

19.5 13.5

22.5 15.5

255 1535

28.5 [75

31.5 18.5

33.5 18.5

35.5 20.5

38.5 23.5

40.5 29.5

41.5 295

42.5 33.5

43.5 35.5

44.0 35.5

44.5 36.5

45.5 37.5

46.5 38.5

setups. In the first few weeks after metamorphosing the

juvenile frogs were fed with tropical springtails (Co/lem-

bola sp.), later on they were additionally fed with dusted

Drosophila hydei.

Data acquisition and evaluation. To document growth

and development of the tadpoles every two to four days

photos of 10 randomly chosen tadpoles were taken from

10" of June to 1% of September 2017. On the last day

of growth documentation only the five remaining larvae,

which had not completed metamorphosis at that time,

were photographed. For this the tadpoles were trans-

ferred into a Petri dish, which was lightened from below

to increase the contrast between tadpole and background

in the recorded photos. All photos were taken with a dig-

ital camera (Olympus TG-2). Additionally, morphologi-

cal data of the fastest developing tadpoles were recorded

to document the developmental stages. The photos were

©ZFMK

Larval development stages and husbandry of the Rice Frog Microhyla mukhlesuri 113

Fig. 3. Gosner stages 40 to 46 of Microhyla mukhlesuri tadpoles, representing the last steps to metamorphosis. Stage 40 (dorsal

view): development of hind limbs and toe pads is completed and foot tubercles have developed; Stage 41 (dorsal view): the fore-

limbs have become well visible under the tadpoles’ transparent skin, the mouthparts begin to atrophy; Stage 42 (ventral view):

both fully developed forelimbs have emerged, mouth angles are positioned anterior to the nostrils; Stage 43 (dorsal view): tail de-

generation begins, mouth angle is positioned between nostril and eye; Stage 44 (ventral view): the tail is already strongly reduced

and the mouth angles are now positioned beneath the eyes; Stage 45 (dorsal view): the tail is greatly reduced and the mouth angles

are positioned posterior to the eyes, additionally, the coloration has already slightly developed; Stage 46 (dorsal view): with the

completed reduction of the tail and further development of the coloration, metamorphosis is finished.

114

analyzed with the tool SAISAQ (Kurth et al. 2014) on

the open source statistics platform R (R Developmental

Core Team 2016). This software package semiautomat-

ically processes image files and computes the surface

area of a tadpole, which is highly correlated with its body

mass. This method is non-invasive and therefore suitable

for repeated measurements on (small) live animals with-

out causing much handling stress. Larval stages were

examined following the universal anuran larvae staging

table developed by Gosner (1960). We identified tadpole

stages between Gosner stage 25 and 46, starting with

the stage in which the tadpoles were found and finishing

with completion of metamorphosis. Voucher specimens

were deposited in the herpetological collection of the

Zoological Research Museum Alexander Koenig, Bonn

(ZFMK 101119-101122 [adults], ZFMK 101528 [juve-

nile], ZFMK 101529 [metamorph]), and ZFMK 101530 —

101531 [larvae]).

RESULTS

A total of 79 tadpoles were found on 10" of June 2017 in

the water part of the adult breeding group terrarium, no

unhatched or unfertilized eggs were left. When detected,

all tadpoles were free swimming without yolk sac and

showed a strong fleeing reaction when disturbed. They

had a mainly transparent body with scattered pigment

cells, a wide mouth with already completely developed

mouthparts, and the gills were covered by the operculum.

Following these features, we determined them as Gosner

stage 25 at the day of finding (Table 1). At this stage,

tadpoles had a mean body surface of about 0.057 cm?

(Fig. 4). Stages 26 to 30 ranged from day 10 to day 16

and were defined by different growth stages of the hind

limb buds (Fig. 2). We were not able to differentiate stage

29. At stage 30, the limbs became slightly bent. The fol-

lowing stages 31 to 37 were characterized by the devel-

opment and indentation of the foot paddle, finishing with

the complete separation of all toes, and lasted from day

17 to 27 (Fig. 2). Stages 38 and 39 were not document-

ed. On day 30 the first tadpoles reached stage 40, charac-

terized by developed foot tubercles and the completion

of the toe pads. Stages 41 to 46, the metamorphs stages,

were characterized by the completion of metamorphosis

and ranged from days 32 to 37 (Fig. 3). At stage 41 the

developed forelimbs were well visible under the tadpoles’

transparent skin. Additionally, the atrophy of the mouth-

parts began in this stage. Forelimbs emerged at stage 42

at day 33, and the mouth angle was positioned anterior

to the nostril. Within the next three stages the mouth an-

gles became translocated more to the distal end of the

tadpoles’ head, until they were positioned posterior to the

eyes at stage 45 at day 36. Furthermore, absorption of

the tail began in stage 43 and was finished in stage 46.

Bonn zoological Bulletin 67 (2): 109-116

Nils Behr & Dennis R6dder

Size [cm*]

00 01 02 03 04 05 06 07

Fig. 4. General growth in tadpoles of Microhyla mukhlesuri,

based on body surfaces computed with SAISAQ (Kurth et al.

2014). Blue spots represent sizes of individual tadpoles; mean

size at the respective day is plotted in the red graph.

Together with the fully developed coloration at this stage

(Fig. 1B), the metamorphosis was completed. The fastest

developing tadpole finished metamorphosis as early as

day 37 after finding, the slowest developing one at day

98, while most (32) metamorphosed between day 73 and

80. Body surface reached its highest peak at day 71 with

a mean surface of 0.490 cm? and the biggest individu-

al measuring 0.689 cm? (Fig. 3). Afterwards, when most

tadpoles reabsorbed their tail to finish metamorphosis,

general body surface decreased slightly. Directly after the

completion of metamorphosis the freshly morphed frogs

measured between 0.221 cm? and 0.333 cm?, and had a

snout-vent length of 6 mm to 7 mm.

DISCUSSION

Our results summarize the first larval staging for

free-swimming tadpole stages of Microhyla mukhlesuri.

As only already hatched tadpoles were found, we were

not able to document early embryonic stages of this spe-

cies. Future captive breeding efforts at the scientific an-

imal keeping facility of the ZFMK will be necessary to

complete the staging table.

We assume that the tadpoles had hatched only one or

two days before we found them in the breeding group

tank. Hence, based on our study tadpoles needed between

38-40 and 98-100 days to complete metamorphosis at

water temperatures of 24 °C to 25 °C. This is very similar

to the findings of Wang et al. (2017) for the closely relat-

ed sister taxon Microhyla fissipes, which needed 43 days

in total and about 41 days after hatching in late stage 23

to complete the metamorphosis at a water temperature of

22.9 °C to 25.4 °C. Furthermore, we found high similar-

ities to other species of the genus Microhyla. Measuring

from stage 25 onwards, tadpoles of M. ornata complete

metamorphosis in about 46.5 days at a water temperature

of 25 °C to 27 °C (Narzary & Bordoloi 2013), and tad-

©ZFMK

Larval development stages and husbandry of the Rice Frog Microhyla mukhlesuri 115

poles of M. okinavensis in about 38.5 days at 19 °C to

26 °C (Shimizu & Ota 2003). Detailed comparisons of

developmental time of different tadpole stages are doc-

umented in Table 2. Mohammad Ridzuan (2013) found

that tadpoles of MZ nepenthicola needed 14 days from

stage 33 to stage 46, while tadpoles of the aforemen-

tioned species needed 18 to 23 days (1.e., M. mukhlesuri:

18d, M. fissipes: 22 d, M. ornata: 21 d, M. okinavensis:

23 d; Table 2). This difference might be due to differ-

ent rearing setups, especially temperature (information

missing for M. nepenthicola) and amount of provided

food usually have high influences on the rate of larval

development (e.g., Duellman & Trueb 1986; Harkey &

Semlitsch 1988; Marian & Pandian 1985).

The number of tadpoles we found (n=79) is rather

small compared to clutch sizes reported for M. fissipes

(209 to 564 eggs [Wang et al. 2017]), M. ornata (300 to

510 eggs [Narzary & Bordoloi 2013]) and M. okinaven-

sis (220 to 910 eggs [Shimizu & Ota 2003]). Hence, we

suppose that it was either the first clutch of a recently

sexually matured and still not fully grown female, which

might produce smaller egg clutches (e.g., Gibbons &

McCarthy 1986), or that the majority of either eggs, pos-

sibly infertile eggs, or tadpoles had already been eaten by

adult frogs. Future breeding efforts might help to collect

sufficient data on clutch sizes.

The combined information about larval development,

captive breeding and management of these frogs gath-

ered in different studies might become an important fac-

tor if measures of conservation necessitate captive breed-

ing programs for the preservation of endangered species

and for restocking programs. Currently, the IUCN Red

List of Threatened Species (2018) lists four species of the

genus Microhyla as Endangered or even Critically En-

dangered (1.e., M. pulchella, M. sholigari, M. zeylanica

and M. karunaratnei). Furthermore, for seven species of

the genus there is not yet enough data available for a clas-

sification by the IUCN Red List and ten species are not

even listed at all (Frost 2018; IUCN Red List of Threat-

ened Species 2018). The endangered species might ben-

efit from the knowledge gained in our and similar studies

as husbandry analogues already today.

Acknowledgements. We want to thank Timo Hartmann and

David Hornes, who helped with the captive management of the

frogs, and Ursula Bott, helping with her excellent proof reading

skills. Morris Flecks and Claudia Etzbauer helped with the mo-

lecular analyses. We are grateful for financial support from the

Alexander-Koenig-Gesellschaft, Bonn. Furthermore, we want

to thank Anna Rauhaus and Thomas Ziegler, who provided

helpful comments to an early version of the manuscript.

REFERENCES

Biyu SD, Dutta S, Bhuddhe GD, Vasudevan K, Srinivasulu C

(2004) Microhyla sholigari. The IUCN Red List of Threat-

Bonn zoological Bulletin 67 (2): 109-116

ened Species 2004: e.T57893A11688938. _ http://dx.doi.

org/10.2305/IUCN.UK.2004.RLTS.T57893A 11688938 .en.

Downloaded on 09 April 2018

Chitmanat C, Tongdonmuan K, Khanom P, Pachontis P, Nun-

song W (2005) Antiparasitic, antibacterial, and antifungal

activities derived from a 7erminalia catappa solution against

some Tilapia (Oreochromis niloticus) pathogens. Acta Horti-

culturae 678: 179-182

Duellman WE, Trueb L (1986) Biology of amphibians. Mc-

Graw-Hill, New York

Frost DR (2018) Amphibian species of the world: an online

reference. Version 6.0 (Date of access 18. April 2018). Elec-

tronic Database accessible at http://research.amnh.org/herpe-

tology/amphibia/index.html. American Museum of Natural

History, New York, USA

Gibbons MM, McCarthy TK (2011) The reproductive output of

frogs Rana temporaria (L.) with particular reference to body

size and age. Journal of Zoology 209: 579-593

Gosner KL (1960) A simplified table for staging anuran em-

bryos and larvae with notes on identification. Herpetologica

16: 183-190

Harkey GA, Semlitsch RD (1988) Effects of temperature on

growth, development, and color polymorphism in the Ornate

Chorus Frog Pseudacris ornata. Copeia 1988: 1001—1007

Hasan M, Islam M, Kuramoto M, Kurabayashi A, Sumida M

(2014) Description of two new species of Microhyla (Anura:

Microhylidae) from Bangladesh. Zootaxa 3755: 401-418

IUCN Red List of Threatened Species. Version 2017-3. <www.

iucnredlist.org>. Downloaded on 10 April 2018

IUCN SSC Amphibian Specialist Group. (2017) Microhyla

pulchella. The IUCN Red List of Threatened Species 2017:

e.T73728100A73728104. http://dx.doi.org/10.2305/IUCN.

UK.2017-2.RLTS.T73728100A73728104.en. Downloaded

on 09 April 2018

Koch C, Venegas PJ, R6dder D, Flecks M, Bohme W (2013)

Two new endemic species of Ameiva (Squamata: Tetidae)

from the dry forest of northwestern Peru and additional infor-

mation on Ameiva concolor Ruthven, 1924. Zootaxa 3745:

263-295

Kurth M, Hornes D, Rédder D (2014) SAISAQ: A novel tool

for semiautomatic image based surface area quantification.

North-Western Journal of Zoology 10: 217—220

Manamendra-Arachchi K, de Silva A (2004a) Microhyla

karunaratnei. The IUCN Red List of Threatened Species

2004: —e.T57883A11686175. _ http://dx.do1.org/10.2305/

TUCN.UK.2004.RLTS.T57883A 11686175.en. Downloaded

on 09 April 2018

Manamendra-Arachchi K, de Silva A (2004b) Microhyla zey-

lanica. The IUCN Red List of Threatened Species 2004:

e.T57895A 11689438. http://dx.doi.org/10.2305/IUCN.

UK.2004.RLTS.T57895A 11689438.en. Downloaded on 09

April 2018

Marian MP, Pandian TJ (1985) Effect of temperature on de-

velopment, growth and bioenergetics of the bullfrog tadpole

Rana tigrina. Journal of Thermal Biology 10: 157-161

Mohammad Ridzuan ASB (2013) Staging of late larval devel-

opmental stages of the microhylid frog, Microhyla nepen-

thicola (Anura: Microhylidae). University of Malaysia Sar-

awak (Bachelor Thesis)

Narzary J, Bordoloi S (2013) Study of normal development

and external morphology of tadpoles of Microhyla ornata

and Uperodon globulosus of the family Microhylidae (Am-

phibia: Anura) from North East India. International Journal

of Advanced Biological and Biomedical Research 3: 61—73

©ZFMK

116 Nils Behr & Dennis R6dder

Poyarkov NA, Vassilieva AB, Orlov NL, Galoyan EA, Dao

Tran Thi Anh, Le DTT, Kretova VD, Geissler P (2014) Tax-

onomy and distribution of narrow-mouth frogs of the genus

Microhyla Tschudi, 1838 (Anura: Microhylidae) from Viet-

nam with descriptions of five new species. Russian Journal

of Herpetology 21: 89-148

R Development Core Team (2016): A Language and Environ-

ment for Statistical Computing. R Foundation for Statistical

Computing, Vienna, Austria

Shimizu S, Ota H (2003) Normal development of Microhyla

ornata: the first description of the complete embryonic and

larval stages for the microhylid frogs (Amphibia: Anura).

Current Herpetology 22: 73-90

Seshadri KS, Singal R, Priti H, Ravikanth G, Vidisha MK &

Saurabh S, Pratik M, Kotambylu VG (2016) Microhyla later-

Bonn zoological Bulletin 67 (2): 109-116

ite sp. nov., a new species of Microhyla Tschudi, 1838 (Am-

phibia: Anura: Microhylidae) from a laterite rock formation

in south west India. PLoS ONE 11(3): e0149727

Wang S, Zhao L, Liu L, Yang D, Khatiwada J, Wang B, Ji-

ang J (2017) A complete embryonic developmental table of

Microhyla fissipes (Amphibia, Anura, Microhylidae). Asian

Herpetological Research 8: 108-117

Yuan ZY, Suwannapoom C, Yan F, Poyarkov NA, Nguyen SN,

Chen HM, Chomdej S, Murphy RR, Che J (2016) Red River

barrier and Pleistocene climatic fluctuations shaped the ge-

netic structure of Microhyla fissipes complex (Anura: Micro-

hylidae) in southern China and Indochina. Current Zoology

62: 531-543

©ZFMK

Bonn zoological Bulletin 67 (2): 117-128

2018 - Galunder K. & Rédder D.

https://doi.org/10.20363/BZB-2018.67.2.117

ISSN 2190-7307

http://www.zoologicalbulletin.de

Research article

urn:|sid:zoobank.org: pub:3646985F-4 125-49CF-BBF2-8EDAACA3AA58

Developmental Ecology and Larval Staging

in Polypedates otilophus (Boulenger, 1893)

(Anura: Rhacophoridae)

Katharina Galunder! & Dennis Rédder?*

“Zoological Research Museum Koenig, Adenauerallee 160, D-53113 Bonn, Germany

* Corresponding author: E-mail: d.roedder@leibniz-zfmk.de

'urn:|sid:zoobank.org:author:F2603E00-55EC-435B-880A-C3E746C68939

?urn:lsid:zoobank.org:author: 1 79FOE77-9745-48EE-A 81 1-FD00627CC994

Abstract. Tadpoles of Polypedates otilophus originating from two different foam nests were raised at water temperatures

between 19°C and 27°C. A larval staging table according to Gosner is provided for the first time for the genus. The clutch

sizes of the foam nests differed (19 and 49 eggs, respectively), but only 14 and 17 tadpoles hatched at stage 25. The first

tadpole of nest one completed metamorphosis after 120 days and the first tadpole of nest two after 131 days. Before meta-

morphosis is completed, the metamorphs developed significant brown striation which is first visible on the hind legs and

subsequently also on the dorsal side of the body. Detailed characteristics of each larval stage are provided. We herein pro-

vide the first detailed report on the larval development of P. otilophus, which can be used as surrogate species for captive

management of other Polypedates taxa. Most interestingly, the temporal development of larvae in this species appears to

be extremely plastic and strongly depending on ambient temperature.

Key words. Developmental ecology, tadpole morphology, environmental plasticity.

INTRODUCTION

The genus Polypedates, which is distributed in East-

ern India, southeastern Asia, the Philippines, and Bor-

neo (Frost 2017), was first described by J. J. Tschudi in

1838. Currently 24 species of the genus are recognized

(Frost 2017) and 15 are listed at the IUCN Red List of

Threatened Species (IUCN 2017). One species 1s list-

ed as “Endangered” (Polypedates insularis Das, 1995)

and four species are listed as “Data Deficient” by IUCN

(Polypedates chlorophthalmus Das, 2005; P. hecticus Pe-

ters, 1868; P. occidentalis Das & Dutta, 2006; and P. zed

[Dubois, 1986]) and ten species are listed as Least Con-

cern (Polypedates colletti [Boulenger, 1890]; P. cruciger

Blyth, 1852; P. leucomystax |Gravenhorst, 1829]; P. mac-

rotis [Boulenger, 1891]; P. maculatus [Gray, 1830]; P.

megacephalus Hallowell, 1860; P. mutus [Smith, 1940];

P. otilophus [Boulenger, 1893]; P. pseudocruciger Das &

Ravichandran, 1998; and P. taeniatus [Boulenger, 1906];

IUCN 2017).

Polypedates otilophus was first described as Rhaco-

Phorus otilophus by Boulenger in 1893, who character-

ized Rhacophorus otilophus by its much depressed head

which is large and a little broader than long. Furthermore,

according to the original description, it possesses a point-

ed snout which is a little longer than the diameter of the

orbit and a nostril which is close to the tip of the snout.

Received: 06.04.2018

Accepted: 25.10.2018

The forehead is concave and the fingers long with rudi-

mentary webs. The tips are dilated into rather large disks

and the toes are two-thirds webbed, but the disks are

smaller than those of the fingers. The skin texture of the

dorsum is finely granular, whereas the skin of the belly

and the lower surface of the thighs are coarsely granular.

The dorsal coloration is pale olive with dark grey spots

and longitudinal streaks (Fig. 1). Further, the hind limbs

are dark cross-banded which become thinner and denser

on the concealed surfaces of the hind limbs. Males have

internal vocal sacs. A male specimen measured in Bon-

gon, North Borneo, had a snout to vent length (SVL) of

80 mm (Boulenger 1893) and Iskandar (2004) reported

that females can reach up to 100 mm SVL.

Polypedates otilophus is \isted as Least Concern by

IUCN (2017) because of its wide distribution, its pre-

sumed large population and because of the tolerance of

a degree of habitat modification. According to the most

recent assessment the species is unlikely to be declin-

ing fast enough to qualify for being listed in a higher

threat category. Matsui et al. (2014) noted that the spe-

cles occurs at many sites in Borneo and on Sumatra at

elevations below 1,000 m a.s.l. The species is arboreal

prefering lowland forests in flat and hilly terrain. Under

natural conditons breeding takes place in temporary rain

pools; specimens are also frequently found in disturbed

Corresponding editor: W. Bohme

Published: 01.11.2018

118 Katharina Galunder & Dennis Rodder

habitats, such as logged areas at the forest edge, which

apparently do not possess a threat (Inger et al. 2004).

There are few studies available reporting on the re-

production and tadpole development of species of the

genus Polypedates. Tapley and Girgin (2015) raised 14

clutches of P. otilophus and reported that tadpoles need

74 to 84 days to reach metamorphosis at 22°C to 26°C.

In this study foam nests were five times produced in the

early morning about 6 a.m., wherein the entire process of

the nest construction took about 45 minutes. Three of the

nests, which were dissected within 24 hours, contained

42 to 119 eggs. The authors reported that the tadpoles

hatched after approximately ten days and that the first

tadpole metamorphosed after 74 days. Metamorphosis

within the cohort took place within ten days (Tapley &

Girgin 2015).

Chakravarty et al. (2011) reported for Polypedates

teraiensis the entire development from ovum fertiliza-

tion up to emergence of the froglet with 58 days at 26°C

to 32°C. They examined five different foam nests of

which clutch sizes varied between 67 and 127 white-co-

loured eggs. Some eggs located on the outermost surface

of the foam nest sometimes did not develop and turned

pale yellow due to desiccation. Embryos of P. teraiensis

hatched at stage 20 and stayed within the foam nest until

stage 22 (sensu Gosner 1960). In another study, Tamuly

& Dey (2014) reported on the larval morphology and de-

velopment of Polypedates teraiensis within 42 days after

hatching at temperatures between 26°C and 33°C. Under

Bonn zoological Bulletin 67 (2): 117-128

these conditions the keratodent jaws developed at Gosner

stage 25 and were assimilated at stage 42.

Yorke (1983) presented data on the survival of embry-

os and larvae for Polypedates leucomystax. The average

embryonic mortality was 34% in field-collected egg

masses, wherein fertilization rates were approximately

100%. The pooled mortility data showed that 98% oc-

curred prior to tail-bud stage. In this study the highest

frequency and proportion of mortality occurred in the

early neural stages, whereas no embryonic mortality was

found beyond tail-bud stages. The mortality increased in

stages 31 to 35 (Yorke 1983).

Hsu et al. (2012) stated that breeding in Polypedates

braueri on the Bagua Terrace takes place from March to

August though tadpoles can be found during the entire

year, hibernating in man-made water containers in low-

land orchands. Laboratory experiments showed that the

overwintering is facultative and can be initialized by low

temperatures and limited food, wherein the role of food

availability was confirmed in the wild (Hsu et al. 2012).

Information on captive breeding of P. otilophus is

scarce. Iskandar (2004) reported that P. otilophus does

not do well in captivity. Janzen (2014) and Tapley and

Girgin (2015) reported on husbandry and breeding events

in captivity. Detailed information on larval staging and

development are currently lacking.

©ZFMK

Larval development in Polypedates otilophus 119

Fig. 2. Tanks used for captive breeding of Polypedates oti-

lophus: the terrarium of the initial breeding group (A); one of

the aquaria used for tadpole raising (B); and one of the terraria

for raising the froglets after metamorphosis (C).

MATERIAL AND METHODS

Captive Management and Breeding

In 2014, 27 tadpoles of Polypedates otilophus were do-

nated to the Zoologisches Forschungsmuseum Alexander

Koenig (ZFMK) by a private German breeder who pre-

viously reported on captive management of the species

(Janzen 2014). At the time the present study was con-

ducted our breeding group consisted of 20 specimens,

which were kept in a terrarium in the animal keeping

unit of the ZFMK measuring 60 x 80 x 140 cm Ixbxh

(Fig. 2A). They were fed ad libitum with adult crickets

(Acheta domesticus or Gryllus assimilis) two or three

times per week and irregularly with flies (Musca domes-

tica), which were fed with fresh fruit or fruit puree to

absorb more vitamins.

The terrarium was divided into two different parts: a

land part and a water part (25 x 80 x 18 cm Ixbxh) con-

taining a water pump (Eheim Universalpumpe, Typ 1260

210) connected to a sprinkler system and some Crypto-

coryne sp. The land part consisted of several layers of

filter padding reaching a total height of 20 cm and was

equipped with different plants (Monstera deliciosa and

Syngonium sp.) and some branches, partly extending

from the water part to the land part. The back side as

well as the right side of the terrarium were covered with

Hygrolon® to keep the humidity high.

Next to natural daylight, LED light strips (Solar Sting-

er 1100 mm Sunstrip Dimmable Diver) served as light

Bonn zoological Bulletin 67 (2): 117-128

source between 8 a.m. and 8 p.m. In order to stimulate

reproduction, the sprinkler system was activated daily

for about three hours, resulting in a relative humidity

between 65% and 90%. Air temperature varied between

20°C and 30°C and the water temperature varied between

20°C:and:28°G.

Under these conditions two foam nests were produced,

wherein the first one was attached on a leaf above the

water part on February 26, 2017 and the second nest was

attached to the glass wall above the water part on March

£5) 201-7.

Raising of tadpoles and froglets

To provide suitable water conditions fresh osmosis wa-

ter was remineralized in an aquarium (50 x 40 x 30 cm

Ixbxh), which was equipped with aquatic plants (Crypto-

coryne sp.), dried leaves (Fagus sylvatica) and circulated

with a water pump (Eheim Powerhead 650). After ex-

tracting water for exchanges on a weekly basis the aquar-

ium was refilled with fresh osmosis water.

The leaf which contained the first foam nest (group 1)

was cut off the plant and placed in a second terrarium (50

x 40 x 40 cm Ixbxh) to avoid disturbances of the adults.

It was similarly equipped as the terrarium of the adults

and automatically sprayed with water three times per day

for each 30 seconds. A box (25 x 18 x 7.5 cm Ixbxh)

providing remineralized osmosis water was placed under

the foam nest to allow hatching of group 1 in a moni-

tored environment. The second foam nest could not be

placed in a separate terrarium without damage as it was

attached to a glass wall. In order to collect the hatching

tadpoles of group 2 a plastic box (10 x 10 x 10 cm Ixbxh)

was placed underneath it. Foam nests were checked daily

and hatched tadpoles were moved into aquaria (see be-

low). Both foam nests were torn apart when no tadpole

hatched after at least one week and the remaining eggs

were counted and photographed. Air and water tempera-

ture ranged between 21°C and 22°C during the develop-

mental phase of both groups.

Tadpoles of both groups were separated in two similar

aquaria (30 x 30 x 30 cm Ixbxh, water level 25 cm) pro-

viding equivalent environmental conditions (Fig. 2B).

Each aquarium contained aquatic plants (Cryptocoryne

sp.), algae-covered stones, dried leaves of Fagus sylvat-

ica, which served as additional food, and snails (Phy-

sella sp. and Planorbella sp.) to remove food remains.

Tadpoles were fed three times per week ad libitum. Due

to availability the food composition changed during the

larval phase, starting with a mixture of three different

minced fish foods (Sera Vipan GroBflocke XL-Hauptfut-

ter fiir alle Zierfische, O.S.I. Spirulina Flakes and Tetra

Tablets TabiMin). Tadpoles of group 1 were fed with this

mixture until day 108, whereas the tadpoles of group 2

were fed with it until day 91. Later on the tadpoles were

©ZFMK

120 Katharina Galunder & Dennis Rodder

IMG_4327.JPG

original

1000 2000 3000

cumulative score

classification

1000 2000 3000

-150 -50 O 1000 2000 3000

Fig. 3. Example of a tadpole picture which was processed with SAISAQ to calculate the body size. The original photo is transferred

into a monochromatic Figure and a threshold is computed to select the most appropriate color intensity to delimit the largest dark

object, which in turn is measured in terms of the total number of pixels. Based on this pixel score the surface is computed using a

standard.

fed with fish food tablets (Tetra Tablets TabiMin, 1—2

tablets/aquarium).

Two-thirds of the water of each aquarium was ex-

changed weekly with remineralized osmosis water and

the algae-covered stones were exchanged. No artificial

light source was provided as the aquaria were placed in

a room providing daylight. The water temperature varied

between 19°C and 27°C (group 1) and 25°C (group 2).

During an unexpected cold period in Germany, the water

temperature of both aquariums sank to 19°C which was

counterbalanced by a heater (Sera Automatic Heater 50

W), thus reestablishing a minimum of 21°C afterwards.

A piece of floating cork was placed in each aquarium to

enable the metamorphosed froglets to leave the water.

Froglets were moved to terraria (50 x 40 x 40 cm Ix-

bxh; Fig. 2C) which were equipped similarly to those of

the adults, with a filter pad as substrate layer and a small

water part (1.5 cm depth) and Hygrolon® to keep hu-

midity high. The terraria were automatically misted with

water three times per day for 30 seconds each resulting in

a relative humidity between 70 and 90 %. The land parts

of both terraria were equipped with Monstera deliciosa,

Pilea sp., bromeliads, different mosses and ferns, and as

light source in both terraria LED light strips were used

(Solar Stinger 1100 mm Sunstrip Dimmable Diver). The

photoperiod was set to daylight hours between 8 a.m.

and 8 p.m.. Air temperature varied between 19.0°C

and 25.0°C and water temperature between 20.0°C and

255°C"

Froglets were fed with flightless fruit flies (Drosophila

melanogaster and D. hydei) and crickets (Acheta domes-

ticus) two to three times per week, which were fed with

fresh vegetables, fruit or fruit puree to absorb vitamins.

Additionally, prey items were dusted with mineral or vi-

tamin powder (herpetal Amphib, herpetal Mineral + Vi-

tamin D3 and herpetal Complete Terrarium).

Bonn zoological Bulletin 67 (2): 117-128

Data Collection and Evaluation

The development of the nests, tadpoles and froglets was

monitored between March 2™ and August 1‘, 2017. After

the tadpoles hatched, standardized photos were taken of

every tadpole every day for a week (Canon Eos 550D) as

described in Kurth et al. (2014). Afterwards the interval

was extended and photos were taken two to four times

per week until day 114 depending on the developmental

progress of the tadpoles. When the tadpoles metamor-

phosed to froglets, photos were taken of every specimen

before they were placed into the terrarium belonging to

their groups. Photos of the frogs were taken weekly.

The growth of the tadpoles was determined using the

SAISAQ pipeline (Kurth et al. 2014), programmed in

the open source statistics software R (R Developmental

Core Team 2017). SAISAQ analyses photograph files

and computes the surface of a tadpole (Fig. 3), which is

highly correlated with tadpole mass and can be used to

describe body size in tadpoles (Kurth et al. 2014).

RESULTS

Development and Staging

During the night from February 25 to February 26 and the

night from March 14 to March 15, the foam nests were

produced, while nest 2 was larger and not as compact as

nest 1. The part of the foam nest containing the eggs sank

in a fluid transition. Both foam nests were beige to yel-

low coloured becoming darker with drying. On day 9 the

foam nest of group 1 showed the first holes when the first

tadpoles hatched (Fig. 4). On day 10 and 11, the foam

nest sank further. From day 14 onwards, the foam started

to disorganize, holes became larger and remaining eggs

became visible (Fig. 4). The foam nest of group 2 started

disorganization on day 6 and most tadpoles had hatched

on day 10 (Fig. 4). The clutch of group 1 contained 49

©ZFMK

Larval development in Polypedates otilophus 121

Fig. 4. Foam nests of Polypedates otilophus. Upper row shows the development of the nest of group 1 (5, 10, 13, 16, and 19 days),

bottom row group 2 (1, 6, 10, 13, and 16 days).

eggs including 35 unhatched eggs, whereas the clutch of

group 2 contained 19 eggs including 2 unhatched.

On day 11 the first two tadpoles of group 1 hatched.

Afterwards, five tadpoles hatched on day 12 and another

five tadpoles the next day. The last two tadpoles hatched

on day 14. On day 8 the first tadpole of group 2 hatched.

On day 9, thirteen tadpoles hatched and on day 10 the

last three did.

A detailed staging table according to Gosner (1960)

is provided in Appendix 1 and corresponding photos

are provided in Figs 4-5, and 7-9. The tadpoles hatched

when they were in Gosner stage 25 (Fig. 6). Between day

21 and day 50 the hind limb buds grew (Fig. 8). On day

41 the first tadpoles of group 1 reached Gosner stage 30.

The first tadpoles of group 1 reached stage 31 on day 51.

Stages 32 to 36 were hard to identify because the muscu-

lature of the tail overlapped the foot bud partly. On day

84 only the first tadpole of group 1 reached stage 37. At

this time two tadpoles of group 1 were in stage 30, sev-

en tadpoles of group 1 were in stages 31 to 33 and the

remaining three tadpoles of group 1 were in stages 34 to

36. The first tadpole completed metamorphosis after 120

days (group 1). The other tadpoles were slower in their

development with the last specimen completing meta-

morphosis on day 169.

Some tadpoles of both groups showed a spinal curva-

ture in the area between the body and the tail fin (Fig. 9)

Bonn zoological Bulletin 67 (2): 117-128

starting on day 55 when a weak spinal curvature became

visible in some tadpoles of group 1. On day 84, most

tadpoles of group 1 showed the spinal curvature which

became more distinctive in the course of time. The spinal

curvature vanished after completing metamorphosis hav-

ing apparently no effects on the frogs.

Comparing the development in the two groups it be-

came evident that the first froglet of group 2 complet-

ed metamorphosis later than the first froglet of group 1,

although hatching earlier. In group 1 one specimen de-

veloped faster than the remaining cohort while in group

2, the tadpoles metamorphosed shortly after one another

(Gosner stages 37 to 40).

Mortality

During the study, five tadpoles died in group 1, and none

in group 2. The tadpoles died on days 18, 109, 117, 128

and 137. Noteworthy, other tadpoles fed on dead speci-

mens proving for the first time adelphophagy in P. ofi-

lophus.

Growth Rate

The tadpoles of group 2, which hatched earlier than the

tadpoles of group 1, hatched also with a smaller body

size (Fig. 6; group 1: 0.176 cm?—0.306 cm’; group2:

©ZFMK

[22 Katharina Galunder & Dennis R6dder

Fig. 5. Larval development of Polypedates otilophus. Gosner stage 25, 14 days old (A); stage 25, 39 days, ventral view (B); stage

29-30, 48 days (C); stage 31-32, 59 days (D); stage 36, 81 days (E); stage 37-38, 87 days (F); stage 39, 96 days (G); stage 39-40,

99 days (H); stage 39-40, 102 days (1); stage 39-40, 105 days (J); stage 42, 109 days (K); stage 43-44, 111 days (L); stage 46, 120

days (M).

0.184—0.247 cm?’). Larger body sizes were reached by

tadpoles of both groups which hatched last. Body sizes

were similar in both groups between day 15 and day 26

(0.7 cm’). Afterwards, the body sizes differed between

the two groups between day 27 and day 67. During this

period, the body sizes of group 1 were larger than the

body sizes of group 2. The body sizes of group 1 devel-

oped from the range of 0.687 cm?—0.868 cm” to the range

of 1.287 cm?—2.618 cm’. At their peaks, the body sizes

of the largest tadpoles of both groups had a difference

of 0.697 cm’. The body sizes of group 2 varied between

0.525 cm? and 0.823 cm? at the beginning of this period

on day 27 and between 1.187 cm? and 2.744 cm? at the

end of this period on day 67. From day 67 to day 79, the

body sizes of the two groups were similar. Between day

67 and 79, there was only one day at which a tadpole of

group | clearly had a larger body size which was than

the other tadpoles (3.246 cm? on day 75). Later on, there

was a period between day 80 and day 102 when the body

sizes of group | were again larger than the body sizes of

group 2. The body sizes of group | varied between the

range of 1.386 cm?’ and 3.589 cm? at the beginning of

this period and the range of 2.094 cm? and 4.545 cm? at

Bonn zoological Bulletin 67 (2): 117-128

the end of this period; whereas the body sizes of group 2

varied between the range of 1.147 cm? and 2.912 cm” at

the beginning and the range of 1.726 cm? and 3.877 cm?

at the end of this period.

Between day 103 and day 129 the body sizes of the

two groups of tadpoles were similar and the body sizes

of some tadpoles of group 2 were larger than those of

group 1. The body size span of group 1 varied between

2.017 cm? and 4.232 cm? at the beginning and between

4 456 cm? and 7.339 cm’ at the end of this period. On day

103 the range of the body sizes of group 2 was between

1.528 cm? and 4.271 cm’. At the end of this time peri-

od, the range of the body sizes was between 3.256 cm?

and 6.178 cm”. The maximum body size on day 115 had

a tadpole of group 2 with 6.181 cm’. After day 130 the

remaining tadpoles of group 1 had larger body sizes

than the remaining ones of group 2. At this time, some

tadpoles went ashore. The body sizes of group 2 were

only collected until day 140, but around this time the

body sizes of the tadpoles of group 1 were noticeably

larger than the body sizes of group 2, for example, on

day 135 the maximum body size of a tadpole of group

1 was 8.094 cm’, whereas the maximum body size of a

©ZFMK

Larval development in Polypedates otilophus

123

10

Group 1 Group 2

9 e tadpoles e tadpoles

% metamorphosis % metamorphosis

7

% 2

— 6 rer teats

oC © e.° 7% 3

— @ a) $334 “

7 sp piss. LSe *°

3 ort tae See oe >

e t e

Oo 4 oo Oa 8% fF2 90% 88 ¢ ¢

oo ee Evae Paes ® %

ty: $s3.185e5.9. °

3? ih ge 2 ° e —

3 Er pee has Corer

Gog heals cen’.

? 1 ee 8 ae o

2 % be ee a H

ee ieee

4 3 Tso 0°,

San e @ bd

1

0) T T T T T T T T T T T T T T T T T T T T T T T T T T T

0 10 20 30 40 50 60 70 80 90 100 110 120 130 140 150 += 160

Days after foam nest production

Fig. 6. Comparison of the growth rates of both groups of tadpoles, metamorphs and juvenile frogs.

tadpole of group 2 was 6.811 cm? on day 136. Moreover,

the maximum body size of all recorded body sizes had a

tadpole of group 1 with 8.864 cm? on day 142. After this

day, the body sizes of the tadpoles of group | decreased.

Metamorphosis

Froglets entered the landpart between day 112 and day

157. Body sizes ranged from 4.130 to 9.477 cm’. The

first tadpoles went ashore with a body size of 4.130 cm?

and 6.534 cm? on day 112 and 124 (group 1) highlighting

a huge difference of the body sizes of the first two tad-

poles (Fig. 6). On day 125, three tadpoles went ashore

(Fig. 6; two specimens belonging to group 2; 5.450 cm?

and 6.413 cm?). Tadpoles of group | left the water ear-

lier starting on day 112 than those of group 2 (day 125),

while in each cohort the first tadpoles going ashore had

smaller body sizes than those which went ashore later.

Mortality and Growth Rate of the Froglets

During the study, no metamorphosed frog died. The first

frog which went ashore on day 112 had a body size of

4.130 cm? (Fig. 6), which was reduced to 3.876 cm?

on day 120. Two frogs went ashore with body sizes of

5.450 cm? and 6.413 cm? on day 125 and later on day

129, with their body sizes decreasing to 5.307 cm? and

Bonn zoological Bulletin 67 (2): 117-128

4.723 cm’. At this time, their body size span was only at

0.584 cm’. The smallest body size had a frog of group

1 with 3.495 cm? on day 126. The three frogs of group

1 had a body size span of 1.419 cm? at the beginning.

Moreover, it became evident that all body sizes of these

three frogs developed similarly. From day 126 to day

132 the body sizes decreased, whereas the body sizes in-

creased from day 132 to day 138.

DISCUSSION

In comparison to the clutch size of 44 to 119 eggs of

P. otilophus reported of Tapley and Girgin (2015), the

clutch size of 19 and 49 eggs in our study seemed very

small. The frogs of our study were sexually mature, but

they were apparently not full-grown. Tapley and Girgin

(2015) report that three foam nests which were observed

by the authors were deposited on leaves and the other

eleven clutches were deposited on the glass walls of a

vivarium. A similar behaviour of the frogs was evident in

our breeding group.

Chakravarty et al. (2011) reported for Polypedates

teraiensis that this species begins to breed sporadically

after the first few rains of the rainy season. Reproduction

in our group of P. otilophus was also induced by an ar-

tificial rainy season. Chakravarty et al. (2011) observed

©ZFMK

124 Katharina Galunder & Dennis Rodder

pela F

°

, ‘, Dent TPT -S hae

Fig. 7. Enlarged views of the mouths of Polypedates otilophus tadpoles from three different perspectives: a tadpole of group 1 on

day 15 (A); a tadpole of group 1 on day 16 (B); a tadpole of group 2 on day 20 (C); and a tadpole of group 2 on day 22 (D).

that commonly foam nests were deposited on vegetation

above shallow temporary water, but also that some in-

dividuals of P. teraiensis deposited some foam nests on

logs or walls of human habitations far from water, which

desiccated and decayed. This behaviour is similar to the

just described behaviour of P. otilophus depositing eggs

on glass walls in captivity.

The large number of eggs found in the foam nest of

group | was conspicuous, but it is not clear, whether

the eggs were unfertilized or whether the tadpoles died

during the early development before they could hatch.

Yorke (1983) reported for Polypedates leucomystax that

the embryonic mortality was 34% and that all eggs in all

observed foam nests were fertilized. Only two eggs of

group 2 did not hatch. One of these two eggs was visi-

ble through the glass side from day 8 onwards. Chakra-

varty et al. (2011) reported for P. teraiensis that some

eggs which were found on the outermost surface of the

foam nest, which may not develop and turned pale yel-

low due to desiccation. The two eggs of group 2 which

were found in the foam nest after the tadpoles hatched

were intensively yellow. Moreover, this foam nest was

the one which was exposed to drier environmental condi-

tions resulting in a dry and hard consistency. There was a

huge difference to the consistency of the foam of group |

Bonn zoological Bulletin 67 (2): 117-128

of which the foam nest was regularly sprayed with water.

Probably the reason why the tadpoles of group 2 hatched

earlier from day 8 to day 10 than those of group 1 which

hatched from day 10 to day 14.

Tapley and Girgin (2015) reported that P. otilophus

tadpoles hatched on about day 10 at a temperature be-

tween 22°C and 26°C, but it was not clear whether the

ambient temperature was the same where the foam nests

were deposited (Tapley & Girgin, 2015). The ambient

temperature where the foam nest of group 1 was deposit-

ed was about 21.5°C and the temperature where the foam

nest of group 2 was placed was about 22°C suggesting

constancy in developmental time despite different ambi-

ent temperatures.

Gosner (1960) reported that the embryos of most spe-

cies hatch between the stages 17 to 20. In our study tad-

poles of Polypedates otilophus dropped into the water at

stage 25. Chakravarty et al. (2011) reported for Polype-

dates teraiensis that tadpoles of this species hatched in

stage 20 and they dropped into the water in stage 22

suggesting some delay between hatching and leaving the

nest. Such a delay may also be present in P. otilophus,

wherein the trigger of leaving the nest is currently un-

known but may be related to rain fall or emerging pred-

ators.

©ZFMK

Larval development in Polypedates otilophus 125

x 3 4 "3

Li ie: i ae

PR eo a ee Fal a

Fig. 8. Development of the hind extremities in Polypedates otilophus. Hind limb bud of a tadpole of group | in stage 26 on day 21

(A), in stage 28 on day 29 (B), in stage 30 on day 48 (C), in stage 31 on day 51 (D), in stage 36 on day 81 (E), in stage 37 on day

84 with five separated toes (F), in stage 39 on day 96 (G), in stage 39-40 on day 99 (H) and in stage 39-40 on day 105 (I).

In our study tadpoles of P. otilophus developed the

lower and upper jaw sheaths in stage 25. In contrast to

this, the upper and lower jaw sheaths of P. teraiensis

developed already in stage 22 and the jaw sheaths had

completely disappeared in stage 42. The reduction of

the jaw sheaths happened in both species at the same

time, but the development of the jaw sheaths is earlier

in P. teraiensis than in other species in which the forma-

tions of keratodonts and jaw sheaths are slightly delayed

(Chakravarty et al. 2011).

The first tadpoles of group 1 needed 120 days for the

entire metamorphosis, whereas the first tadpoles of group

2 completed their metamorphosis after 131 days. This

time which the tadpoles needed differs hugely from the

period of time which the P. otilophus of Tapley and Gir-

gin (2015) required, completing metamorphosis between

74 days and 84 days. They reported only ten days for the

complete metamorphosis of the first tadpole and the last

metamorphosed tadpole. In our group 1, there was a dif-

ference of 49 days between the complete metamorphosis

of the first and the last tadpole indicating huge plasticity.

One factor could be lower water temperatures, but con-

ditions were largely similar in both studies: Tapley and

Girgin (2015) raised their tadpoles at 22°C to 26°C while

Bonn zoological Bulletin 67 (2): 117-128

ours were raised between 19°C and 27°C. Moreover, Ta-

pley and Girgin (2015) reported that their tadpoles were

fed daily. In contrast, our tadpoles were fed only three

times per week but aditionally fed with dried leaves and

algae-covered stones available ad libitum.

The factors food availability and temperature are im-

portant ones and strongly affect developmental times.

Hsu et al. (2012) reported for Polypedates braueri that

these tadpoles hybernate if food ressources are limited

and, when the ambient temperatures are too low for com-

pleting metamorphosis in one year. Furthermore, the au-

thors reported on laboratory experiments in which some

tadpoles raised at 15°C never reached metamorphosis

(Hsu et al. 2012). These results highlight two points: On

the one hand, the development of the tadpoles is influ-

enced by low temperatures so that their development de-

celerates or is disrupted. On the other hand, the quantity

of food is pivotal. These factors could be the reason of

the different lengths of the larval phase in our study com-

pared to previous publications. Our two groups were ex-

posed to lower temperatures at a different point of time in

their development. That might be the reason why group 2

was behind the development of group 1 at the beginning

on day 36, before both groups got a heater. On day 36, it

©ZFMK

126 Katharina Galunder & Dennis Rodder

Fig. 9. Specimens of Polypedates otilophus with spinal curvatures: a tadpole of group | on day 55 (left) and a tadpole of group 1

on day 124 (right).

was visible that the tadpoles of group 1 had larger body

sizes than the tadpoles of group 2. At this point of time,

the water temperature of group 1 was only about 19°C.

After the heater was in use, the water temperature was

not lower than 21°C. Furthermore, group 2 comprised

more tadpoles. Maybe, the larger group size has caused a

slower development.

In contrast to this hypothesis, Chang et al. (2014) re-

ported for Rhacophorus moltrechti that the tadpoles

reared under water temperatures of 17°C and 22°C and

increased tadpole density, enhanced their larval growth,

translating into greater metamorphic mass without chang-

ing time to metamorphosis or decreasing survival rates.

This process was only reported on the tadpoles which

were raised under the just described temperatures. The

tadpoles which were raised under a water temperature of

27°C did not show this kind of development (Chang et al.

2014).

Some tadpoles died in group 1, whereas none died in

group 2. The first tadpole died on day 18. At this time,

tadpoles of group 1 were in stage 25. The next tadpole

died on day 109. There is a huge time span during which

no tadpole died. Yorke (1983) reported for Polypedates

leucomystax that the mortality increased between the

31th and 35th day. Moreover, the pooled mortality data

showed that 98% of the mortality occurred prior to the

tail-bud stage, but in contrast to this, most tadpoles of

group | died after the tail-bud stages. Some of the tad-

poles of group 1 which died had an extremely distinct

spinal curvature. Although the species appears to tolerate

a broad range of temperature regimes, more studies are

necessary to determine the optimal temperature for larval

development.

Acknowledgements. We would like to thank Morris Flecks

who helped with the camera equipment and arrangements of

plates. Furthermore, thanks to Timo Hartmann, David Hornes

Bonn zoological Bulletin 67 (2): 117-128

and Nils Behr who assisted with the animal care. Ursula Bott

helped with her excellent proofreading skills. Very helpful

comments by Thomas Ziegler significantly improved the man-

uscript.

REFERENCES

Boulenger GA (1893) 4. Descriptions of new reptiles and ba-

trachians obtained in Borneo by Mr. A. Everett and Mr. C.

Hose. Proceedings of the Zoological Society of London,

1893: 522-528

Chakravarty P, Bordoloi S, Grosjean S, Ohler A, Borkotoki

A (2011) Tadpole morphology and table of developmental

stages of Polypedates teraiensis (Dubois, 1987). Alytes 27:

85-115

Chang YM, Tseng WH, Chen CC, Huang CH, Chen YF, Hatch

KA (2014) Winter breeding and high tadpole densities may

benefit the growth and development of tadpoles in a subtrop-

ical lowland treefrog. Journal of Zoology 294: 154-160

Frost DR (2017) Amphibian species of the world: an online ref-

erence. Version 6.0. Electronic Database accessible at http://

research.amnh.org/herpetology/amphibia/index.html. Amer-

ican Museum of Natural History, New York, USA. Accessed

20 Jul 2017

Gosner KL (1960) A simplified table for staging Anuran em-

bryos and larvae with notes on identification. Herpetologica

16: 183-190

Hsu JL, Kam YC, Fellers GM (2012) overwintering tadpoles

and loss of fitness correlates in Polypedates braueri tadpoles

that use artificial pools in a lowland agroecosystem. Herpe-

tologica 68: 184-194

Inger R, Stuebing R, Iskandar D, Mumpuni (2004) Polypedates

otilophus. The IUCN Red List of Threatened Species. ac-

cessible at http://dx.doi.org/10.2305/IUCN.UK.2004.RLTS.

T58962A11862860.en. UK. Accessed at 28 August 2017

Iskandar DT (2004) The amphibians and reptiles of Malinau re-

gion, Bulungan research forest, East Kalimantan: annotated

checklist with notes on ecological preferences of the species

and local utilization. CIFOR, Jakarta

IUCN 2017. The IUCN Red List of Threatened Species. Ver-

sion 2017-3. http://www.iucnredlist.org. (download: 06 Au-

gust 2017)

©ZFMK

Larval development in Polypedates otilophus 127

Janzen P (2014) Ohrenfrosch — ein noch wenig bekannter Gast

im Terrarium. Reptilia 107: 40-43

Kurth M, Hérnes D, Réodder D (2014) SAISAQ: A novel tool

for semiautomatic image based surface area quantification.

North-Western Journal of Zoology 10: 217—220

Liem DSS (1970) The morphology, systematics, and evolution

of the Old World treefrogs (Rhacophoridae and Hyperolii-

dae). Fieldiana Zoology 57: vii + 145

Matsui M, Hamidy A, Kuraishi N (2014) A new species of

Tamuly D, Dey M (2014) Larval morphology and development

of tree frog Polypedates teraiensis (Dubois, 1987). Current

World Environment, Bhopal 9: 182—187

Tapley B, Girgin SM (2015) Captive husbandry and breeding

of file-eared tree frogs, Polypedates otilophus (Boulenger,

1893) (Amphibia: Anura: Rhacophoridae). The Herpetologi-

cal Bulletin 132: 5-8

Tschudi JJ (1858) Klassifikation der Batrachier. Mit Bertick-

sichtigung der fossilen Tiere. Reprint 2014 by Dogma Ed.,

Bremen

Yorke CD (1983) survival of embryos and larvae of the frog

Polypedates from Sumatra, Indonesia (Amphibia: Anura).

Species Diversity 19: 1-7

Polypedates leucomystax in Malaysia. Journal of Herpetol-

ogy 17: 235-241

APPENDIX I

Appendix 1. Larval development of Polypedates otilophus according to the Gosner (1960) stages. Gr. refers to group | and 2 re-

spectively, Day refers to the number of days after nest deposition.

Date [dd.mm. Pa Characteristic Traits

Gr 1103-2017 The eggs are round and brown coloured with a different intensity;

Gr 2522:03.2017 ees)

il eceoscer man The tadpoles dropped into the water; the body of the tadpole was silver coloured underneath the

22.03; 2017, eyes in the lateral view, whereas the body of the tadpole was yellow coloured in dorsal view; the

part between the eyes to the tail fin was yellow coloured; the tadpoles were flesh-coloured on the

ventral side; the tail fin was transparent with black and silver dots and with a significant visible

musculature; the black dots were mainly visible from the transition of the body to the tail fin to two-

thirds of the tail fin; variations: the black pigmentation on the whole tail fin; some show a stronger

pigmentation on the whole tail fin; mostly the lower as well as the rear tail fin area were free of dots;

the number, the order and the size of the dots were individually different; most tadpoles had also

black dots around the eyes, on eye level and on the yellow coloured area as well; these dots were also

individually variable in size, number and order; the silver dots on the body were on the same area

like the black dots, but in comparison to the black dots, the silver dots were visible only above the

USC UIALUFE on the tail fin; aa ae was more or less oval in the dorsal view; the body became

ahs:

Gr 19 122032017 rar 12032017 | 15 [om Bi i a tee eat a upper jaw sifeaiite black an rows developed (Fig. 7A/B);

G15 2s 27203.2017

gre One third of the musculature of the tail fin became thicker (Fig. 5A);

| Gr. 2: 30.03.2017 | 2530;03:2017

Gr. 1: 18.03.2017 | 21 | Tadpoles developed the hind limb buds (Fig. 8);

x 2:03.042017| 20 |

26 | Gr. 1: 22.03.2017

Gr. 2: 05.04.2017 | 22

ge eies The hind limb buds increased their growth;

| Gr. 2: 09.04.2017,

Gr, -1: 26,03 2017 | °29

Gr. 1: 01.04.2017 The hind limb buds had approximately 1.5 % size of their diameter;

Gr. 2: 17.04.2017

©ZFMK

Tadpoles increased their body weight; the body shape was oval; the body did not become wider

beyond the eyes;

The growth of the hind limb buds still went on; the colouring of the tadpoles was still the same;

Bonn zoological Bulletin 67 (2): 117-128

128 Katharina Galunder & Dennis Rodder

Appendix 1. (continued)

Date [dd.mm. a Characteristic Traits

30 Gr. 1: 07.04.2017 41 | The hind limb buds grew to the double size of their diameter; the body was silver coloured with

40. | black dots around the mouth from the ventral view (Fig. 5B);

pot At this time the first tadpoles reached this stage; the other tadpoles were still in stage 30;

| Gr. 2: 20.05.2017, 2: 20.05.2017

tesa Gr. 1: 21.04.2017 a Some tadpoles had a light spinal curvature in the transition between the body and the tail fin

(Fig. 9A);

ae

31-35 Gr. 1: 25.04.2017 | 1: 25.04.2017 The spinal curvature became more distinctive by the concerned tadpoles of group 1; the tadpoles

Gr. 2: see ee = grew;

0 06

31-35 | Gr. 1: 01.05.2017 a The spinal curvature became thicker in the concerned tadpoles of group 1;

<i All toes with the exception of the first and the second ones were separated; no pigmentation on the

Gr. 2: 04.06.2017 hind legs visible; the body shape was rounder;

Gr. 1: 20.05.2017 | 84 | All five toes were separated in one tadpole of group | (Fig. 8D); the foot was flesh-coloured with

chee lee: es gots most tadpoles of group 1 developed a spinal curvature; the spinal curvature became more

0.06 ] ] : ] ,

37-38 | Gr. 1: 23.05.2017 ara The hind legs became thicker: it seemed like the hind legs were ruffled; during this time the skin of

Gr. 2: 22.06.207 | 100 | the hind legs looked transparent so that the underlying skeleton was visible;

37-38 ee 90__} The pigmentation on the hind legs increased (Fig. 8F);

| Gr. 2: 25.06.2017,

38 | Gr. 1: 29.05.2017 ees) The inner metatarsal tubercle was formed; the skin on the hind legs still seemed ruffled; moreover,

a Beer ee | Ban the skin on the hind legs became slowly non-transparent and yellow coloured;

06.20 Q

39 Gr. 1: 01.06.2017 96 | The hind legs were non-transparent and yellow coloured;

Gr. 2: 28.06.2017

39-40 Peer eon The hind legs had a light striation with brownish stripes;

Gr. 2: 01.07.2017

39-40 | Gr. 1: 07.06.2017 | 102 | The hind legs show a significant brown striation; the body has a very light brown striation; the

oe aD are poorly visible under the skin; there was only a light thickening visible where the

39-40 | Gr. 1: 10.06.2017 | 105 | The brown striation caine more Sronouiiced on the body;

Gr. 2: 10.07.2017

Sees The forelimbs are visible under the skin; the brown striation of the body was significant and the basis

| Gr. 2: 13.07.2017 | 2713.07 20 UF of the tail fin also had the brown striation (group 2);

| Gr. 1: 14.06.2017, 1: 14.06.2017 | 109 | Brown striation of the body is significant and the basis of the tail fin shows also a brown striation

group 1); forelimbs protruded;

43-44 | Gr. 1: 16.06.2017 | 111 | The tail fin started to regress; first frogs went ashore; they often held onto the glass walls of the

Gr. 2: 17.07.2017 aquarium instead of sitting on the cork;

eee There was only a stub of the original tail fin;

| Gr. 2: 19.07.2017 | 2: 19.07.2017

Gr. 1: 25.06.2017 | 120 | The metamorphosis of the first tadpoles was completed;

Bonn zoological Bulletin 67 (2): 129-143

2018 - Sinclair B.J. & Saigusa T.

https://doi.org/10.20363/bzb-2018.67.2.129

ISSN 2190-7307

http://www.zoologicalbulletin.de

Research article

urn:|sid:zoobank.org:pub:CD343A48-8F65-40DD-81A9-9F46E 18A4331

Revision of Francis Walker’s female types of

North American Rhamphomyia Meigen (Diptera: Empididae)

Bradley J. Sinclair’ & Toyohei Saigusa?

“Canadian National Collection of Insects & Canadian Food Inspection Agency, OPL-Entomology,

K.W. Neatby Bldg., C.E.F, 960 Carling Ave., Ottawa, ON, KIA 0C6, Canada

?17-1-402 Baikoen 2-chome, Chuo-ku, Fukuoka-shi 810-0035, Japan

“Corresponding author: E-mail: bradley.sinclair@canada.ca

'urn:Isid:zoobank.org:author:45 16327F-B73E-456C-927F- 1 8EFBOB9E08B

2urn:Isid:zoobank.org:author:390F7B16-1B9D-419F-BOCD-F3279B89DF1D

Abstract. Five Nearctic species described by Walker (1849) on the basis of female specimens are revised and illustrated.

Rhamphomyia agasicles Walker is recognized as a subjective synonym of R. minytus Walker and by First Reviser action,

the latter is chosen as the senior of the two names. The following additional new synonyms are proposed: R. dana Walker,

1849 and R. valga Coquillett, 1895 = R. poplitea Wahlberg, 1844; and R. pulla Loew, 1861 = R. cophas Walker, 1849.

Rhamphomyia mallos Walker could not be associated with any recent specimens and only several old female specimens

were found to be conspecific with R. ecetra Walker.

Key words. Dance flies, Nearctic, new synonyms.

INTRODUCTION

There are some 210 species of Rhamphomyia Meigen,

1822 described from North America (Melander 1965;

Poole 1996; Bartak 2002; Saigusa 2012), which likely

only represents about 25 % of the total diversity in the

Nearctic Region. As with most Empidoidea, species con-

cepts are based primarily on differences in the male gen-

italia and species based on females alone are not easily

identified subsequently. The British entomologist, Francis

Walker described ten species of Rhamphomyia, of which

six were unfortunately based solely on female specimens

(Walker 1849, 1857). Many of the species described by

Walker were collected in subarctic and high boreal Can-

ada (Danks 1981) and deposited in the British Museum

insect collection. These species, consisting of notoriously

brief descriptions were included in Walker’s cataloguing

of the thousands of specimens in the museum’s collection

(see Evenhuis 2018). Smith (1971) revised many of the

Nearctic species of Empididae s. /at. described by Francis

Walker, but was not able to associate males with the six

female-based species.

The Canadian National Collection of Insects (CNC)

houses vast holdings of empidoids collected worldwide,

including the genus Rhamphomyia. Building on the ef-

forts of earlier curators, primarily Curran, Shewell and

Chillcott (see Cumming et al. 2011), T. Saigusa sorted

all available specimens of Rhamphomyia to subgenus,

Species group and species during a four month visit in

Received: 18.09.2018

Accepted: 06.11.2018

1985. In addition, extensive notes and pencil sketches of

the male genitalia and male and female legs were made

which have permitted identification by subsequent cura-

tors and students.

During a study of Canadian Arctic Rhamphomyia (Sin-

clair et al. in prep.), it was decided to study Walker’s fe-

male Rhamphomyia types in order to clarify their iden-

tification by associating, 1f possible, the male sex. The

association of males with the female types was accom-

plished rather rapidly for three of the five species (type

of sixth species presumed lost) and is a testament to the

quality and breadth of the CNC Rhamphomyia collection.

The results are discussed below, with each species rede-

scribed and illustrated to facilitate future species identi-

fication.

MATERIAL & METHODS

Specimens were borrowed from or housed in the follow-

ing institutions: BMNH — The Natural History Museum,

London; CNC — Canadian National Collection of Insects,

Arachnids and Nematodes, Ottawa; MZH — Finnish Mu-

seum of Natural History, Helsinki; NHRS — Naturhis-

toriska Riksmuseet, Stockholm; UGIC — University

of Guelph Insect Collection, Guelph; USNM — United

States National Museum of Natural History, Washington,

D.C. Only five of stx Walker types were examined, but all

species are listed below.

Corresponding editor: X. Mengual

Published: 15.11.2018

130 Bradley J. Sinclair & Toyohei Saigusa

Digital images of legs and wings were taken with a

Leica camera model DFC425C managed by a Leica Dig-

ital Imaging System. Terms used for adult structures fol-

low those of Cumming & Wood (2017).

Label data for primary types are cited from the top

downward, with the data from each label in quotation

marks. Labels are cited in full, with original spelling,

punctuation, and date, and label lines are delimited by a

slash (/). Additional information is included in square [ ]

brackets. The repository of each type is given in paren-

theses. Refer to Smith (1971) for explanation of collec-

tors and data labels of the Walker types.

TAXONOMY

Rhamphomyia cophas Walker, 1849

(Figs 1-5)

Rhamphomyia cophas Walker, 1849: 499. Type locality:

New York, USA.

Rhamphomyia pulla Loew, 1861: 330. Type locality:

Connecticut, USA. Syn. nov.

Note about synonymy. Although the holotype of Rham-

Phomyia cophas is in very poor condition (Figs 1, 3),

features including leg colouration, body size and mid

tibia chaetotaxy allowed for association with identified

specimens of R. pulla. Although the male holotype of

R. pulla, housed in the Museum of Comparative Zool-

ogy (see: http://140.247.96.247/mcz/Species_record.

php?id=1016) possesses yellow femora and tibiae, fe-

male legs are much darker in this species, especially the

mid and hind legs.

Type material examined. Rhamphomyia cophas. HO-

LOTYPE 9, labelled (Fig. 2): “Type [green margined

circle]”; “?? Foster/ New York”; “N.York [on reverse

side: 44/ 90”; “Walker’s/ measurements/ in error/ EAW

31.3.00”; “One of Walkers/ series so named./ EAW

[on reverse side: “Rhamphomyia/ cophas/ Walk.]”;

“not Ent. Club.”; “Cophas,”; “New York.”; “Holo-/

type [red margined circle]; “BMNH(E) #/ 246916”;

“NHMUKO10210621 [data matrix code]” (BMNH). The

holotype is in poor condition, with left midleg, both hind

legs, left fore tarsomeres 2—5 and abdomen missing; left

wing is slide mounted (see Smith 1971, pl. 2, fig. 7).

Additional material examined. CANADA. Nova Sco-

tia: Cape Breton Highlands NP, Pleasant Bay, 25-29.

vi.1984, dry and wet mixed forest, H.J. Teskey (2 3,

2 299, CNC). Ontario: Coldwater, 20, 30.v.1959, J.G.

Chillcott (8 3, 6 9, CNC); Midland, swampy woods,

2, 26.v.1959, J.G. Chillcott (3 64, 3 99, CNC); Orillia,

7.Vi.1925, 28.vi.1926, 16.vi.1927, C.H. Curran (5 33,

1 2, CNC) (Fig. 4); Osgoode, 22.v.1964, J.R. Vockeroth

Bonn zoological Bulletin 67 (2): 129-143

(1 4g, CNC); Ottawa, 30.v.1982, J.R. Vockeroth (1 3,

CNC) (Fig. 5); Simcoe, 2.vi.1939, G.E. Shewell (1 3,

CNC). Quebec: Beach Grove, 7.vi.1965, D.G.F. Cobb

(1 4, CNC); Duncan Lake, nr. Rupert, 10.vi.1971,

J.F. McAlpine (1 3, CNC); Gatineau Pk, King Mtn,

45°29'N, 75°51'W, 1.vi.2011, B.J. Sinclair (1 4, CNC);

Knowlton, 20.vi.1927, G.S. Walley (1 4, CNC); Mt. St.

Hilaire, 4.vi.1963, J.G. Chillcott (1 4, CNC); Norway

Bay, 20.vi.1939, E.G. Lester (1 @, CNC); Old Chel-

sea, 12.vi.1964, J.R. Vockeroth (1 4, CNC); Rigaud,

11.vi.1981, J.R. Vockeroth (1 ¢@, CNC). UNITED

STATES OF AMERICA. Georgia: Towns Co., Tray

Mtn, summit, 26.v.2000, G. & M. Wood (1 3’, CNC).

Michigan: Charlevoix Co., 31.v.1960, R. & K. Dreisbach

(3 33,1 2, CNC); Delta Co., 11.vi.1960, R. & K. Dreis-

bach (1 3, 1 9, CNC); Gratiot Co., 25.v.1958, R. & K.

Dreisbach (2 34, 1 9, CNC); Mackinac Co., 7.vi.1957,

7.vi.1960, R. & K. Dreisbach (1 &, 4 9, CNC); Mecosta

Co., 15.vi.1957, R. & K. Dreisbach (1 ¢, 1 9, CNC);

Missaukee Co., 31.v.1957, R. & K. Dreisbach (1 3’, 1 9,

CNC); Ontonogan Co., 18.v1.1960, R. & K. Dreisbach

(3 33, 1 2, CNC); Roscommon Co., 31.v.1957, R. &

K. Dreisbach (1 4, 1 9, CNC). New York: McLean

Reserve, nr. Dryden, 11.vi.1964, J.G. Chillcott (1 ¢,

2 92, CNC). North Carolina: Great Smoky Mtns NP,

Noland Divide Tr., 1700 m, 35°33'58"N, 83°28'37"W,

4 .vi.2001, J.M. Cumming (2 6, CNC). Ohio: Hocking

Co., Coovert Reserve, 19.v.2003, hollow, J.M. Cumming

(2 83, CNC).

Diagnosis. Males of this species are distinguished by the

dark abdomen, yellow femora and tibiae, with at least

apex of hind tibia darkened; scape and pedicel yellow-

ish; and upper digitiform process of sternite 7 projecting

horizontally, lower process hook-like, arched dorsally;

phallus with lower loop rectangularly curved and ridged

expansion only slightly broader than shaft of phallus.

Female have a darkened, shiny abdomen, mid and hind

femora dark with yellowish tip and fore femur mostly

dark with apical half yellowish.

Redescription. Wing length 6.2-7.4 mm. Male. Head

dark in ground-colour, with greyish pruinescence on

face, frons, postgena and occiput; oral margin shiny red-

dish. Holoptic, eyes with ommatidia very slightly larger

on upper half of eye. Frons divergent towards antennal

sockets, bare. Margins of face slightly divergent. Ocel-

lar triangle with pair of slender ocellar setae, longer than

postocular setae. Upper half of occiput bearing row of

stout postocular setae, stouter than ocellar setae; lower

postocular setae mostly slender and longer. Occipital

setae black and stout. Antenna with scape, pedicel and

extreme base of postpedicel yellow; postpedicel mostly

dark, nearly 4x longer than basal width; stylus length

subequal to basal width of postpedicel. Palpus yellow,

bearing long, slender setae. Clypeus bare, reddish and

©ZFMK

Revision of Francis Walker’s female types of North American Rhamphomyia Meigen [31

ie

NHMUK010210621 ?

Figs 1-5. Rhamphomyia cophas Walker. 1. Holotype, female, dorsal view; 2. Holotype labels; 3. Holotype, lateral view; 4. Recent

specimen, female, lateral view; 5. Male terminalia, lateral view. See Additional material examined section for locality details of

recent specimens.

132 Bradley J. Sinclair & Toyohei Saigusa

glossy; labrum dark and glossy, longer than eye height;

labellum dark and bearing many fine setae.

Thorax dark with dense grey pruinescence; brown to

black vittae beneath acrostichal and dorsocentral rows

extending to prescutellar depression; posterior corner of

postpronotal lobe and postalar ridge yellowish brown.

Pleura with yellowish brown highlights about suture

lines. Prosternum bare; proepisternum at fusion point

with prosternum with several dark setae; upper part of

proepisternum in front of anterior spiracle bare. Antepro-

notum with dense row of long, stout setae. Postpronotal

lobe with 1 outstanding seta and 8 or more shorter setae

of various thicknesses; acrostichal setae uniserial, two-

thirds length of dorsocentral setae; dorsocentral setae

uniserial, increasing in length posteriorly, prescutellar

seta longer than lateral scutellar seta; 1 presutural su-

pra-alar seta (= posthumeral), with several surrounding

fine setulae; 3-4 notopleural setae, with several fine set-

ulae anteriorly; 0 prealar setae; 1 postsutural supra-alar

seta; 1 postalar seta and several shorter setae; 1 long

apical pair and shorter lateral pair of marginal scutellar

setae. Laterotergite with cluster of long, dark setae. Ante-

rior and posterior spiracles blackish brown.

Legs long, stout; femora and tibiae yellow, apex of

hind tibia often brown; coxae brown becoming yellowish

apically. Coxae with lateral row of black setae. Femo-

ra with dense white ventral pile. Fore femur with row

of anteroventral and posteroventral setae increasing in

length apically. Fore tibia with long, pale ventral pubes-

cence; 2—3 anterodorsal and posterodorsal setae; apex

with several dark setae. Mid femur with anteroventral

and posteroventral rows of short, even-length stout se-

tae; 1 dark anterior preapical seta. Mid tibia with antero-

ventral and posteroventral row of short stout setae; 2—3

anterodorsal and posterodorsal setae. Hind femur longer

and stouter compared to other femora; anteroventral row

of short stout setae increasingly closer together distally;

two rows of 3-5 stout anterior setae on apical third. Hind

tibia with long, ventral pubescence; 5—6 black anterodor-

sal and posterodorsal setae, shorter than width of tibia;

apex with several dark setae; 1 long seta in posteroapical

comb. Tarsomere | of all legs slender; ventral margin of

all tarsomeres with dense ventral pile; hind tarsomeres

with several pairs of spine-like ventral setae.

Wing lightly infuscate; pterostigma elongate, distinct;

basal costal seta present. Cell dm shorter than cell bm;

CuA+CuP complete, reaching wing margin without

weakening; alular incision acute; margin of calypter with

dark setae. Halter with whitish knob and yellowish stalk.

Abdomen dark brown, basal segment and tergites 7 and

8 with whitish pruinescence, remaining segments shiny;

setae pale brown, lateromarginal setae on segments

2 and 3 long, stouter and darker. Sternites 2-6 with pair

of very long, divergent median marginal setae, nearly as

long as length of sclerite. Tergite 7 similar in width to

tergite 6. Sternite 7 tapered apically to truncate margin,

Bonn zoological Bulletin 67 (2): 129-143

with 2 rows of long stout setae, becoming longer pos-

teriorly; posterolateral margin expanded into cup-like

process, bearing pair of short digitiform processes: up-

per process short, horizontally projected; lower process

hook-like, arched dorsally. Tergite 8 more thickly scle-

rotized than tergite 7, expanded laterally at mid-length,

fused to sternite. Sternite 8 slender, compressed between

expanded posterolateral margins of sternite 7; bearing

long, stout setae. Terminalia (Fig. 5) dark brown, phallus

pale brown. Hypandrium slender, strap-like, wrapping

around base of phallus. Epandrium subrectangular, with

rounded apex; posterior half clothed in long, stout setae;

slightly longer than length of cercus. Subepandrial lobe

lacking; bacilliform sclerite with dense, erect short setae.

Cercus subrectangular with rounded apex; dorsal margin

with triangular medial lobe at mid-length. Phallus slen-

der with two deep loops, with lower loop rectangular-

ly curved; sharply bent at mid-length at point of ridged

expansion; expansion only slightly broader than shaft of

phallus. Ejaculatory apodeme narrow, subtriangular; hor-

izontal lamella on ventral margin.

Female. Similar to male except (Fig. 4): frons with

greyish pruinescence, glossy medially; margin with row

of setae, uppermost shorter. Acrostichal row initially

uniserial, biserial on posterior half. Mid and hind fem-

ora dark with yellowish tip and fore femur mostly dark

with apical half yellowish; hind femur inflated, broader

than other femora; femora without ventral pile. Abdomen

mostly shiny, except for apical segments; setae pale, ex-

cept pair of dark, divergent setae on sternites 2—5; dark,

pilose pleural membrane, indicating retracted pleural

sacs. Cercus long and slender, length 4—5 times width.

Geographic distribution. This species is widespread in

eastern North America, ranging from northern Ontario

(Canada), south to North Carolina and Georgia (USA)

and as far west as Michigan.

Remarks. Although the holotype of R. cophas is in poor

condition, very greasy with abdomen and most of the

legs missing, it can be readily assigned to the subgenus

Calorhamphomyia Saigusa, 1963 on the basis of lustrous

clypeus, leg colouration, thoracic chaetotaxy (prealar

setae absent) and general body size. Saigusa (1963) as-

signed R. pulla to this subgenus.

Rhamphomyia ecetra Walker, 1849

(Figs 6-10)

Rhamphomyia ecetra Walker, 1849: 500. Type locality:

Georgia, USA.

Type material examined. Rhamphomyia ecetra: HO-

LOTYPE 9, labelled (Fig. 8): “Type [green margined

circle]”; “One of Walkers/ series so named./ EAW

[on reverse side: “Rhamphomyia/ ecetra/ Walk.]”;

©ZFMK

Revision of Francis Walker’s female types of North American Rhamphomyia Meigen 133

BMNH(E) #

246919 *

: 8

NHMUK 010210624

Figs 6-10. Rhamphomyia ecetra Walker. 6. Holotype, female, dorsolateral view; 7. Recent specimen, female antenna, lateral view;

8. Holotype labels, left label underside of corresponding label in centre row; 9. Recent specimen, female, lateral view; 10. Recent

specimen, female, anterior view. See Additional material examined section for locality details of recent specimens.

134 Bradley J. Sinclair & Toyohei Saigusa

“Ecetra,”; “Georgia.”; “Pararhamphomyia”’; “Holo-

/ type [red margined circle]; “BMNH(E) #/ 246919”;

“NHMUKO10210624 [data matrix code]” (BMNH). The

holotype is missing both hind tibia and tarsus, right fore

tibia and tarsi and left wing is slide mounted (see Smith

iOFl spl 3: tig: 1).

Additional material examined. UNITED STATES OF

AMERICA. Georgia (3 9°, USNM). North Carolina:

Morrison (3 9°, USNM) (Figs 7, 9, 10).

Diagnosis. Females of this species are distinguished by

the dense presutural supra-alar setae; 3 pairs of scutellar

setae; dark legs with dorsal and ventral pennate setae on

hind femur and tibia, mid femur and tibia and fore tibia,

and dorsally only on fore and mid basitarsus; abdomen

shiny with reddish margins; cell dm short, distinctive

shape, shorter than cell bm; basal costal seta present.

Redescription. Wing length 4.1-4.66 mm. Female.

Head dark in ground-colour, with greyish pruinescence

on face, frons, postgena and occiput; oral margin shiny

reddish. Dichoptic, ommatidia even-sized throughout.

Frons and face broad, subequal in width; frons with row

of setulae along eye margin (Fig. 7). Ocellar triangle with

pair of long ocellar setae, longer than postocular setae.

Upper half of occiput bearing row of stout postocular

setae, stouter than ocellar setae; lower postocular setae

brown, paler than upper setae, slender and shorter. Oc-

cipital setae black and stout; postgenal setae paler than

occipital setae, long and slender. Antenna brown; scape

longer than pedicel; postpedicel nearly 3= longer than

basal width (Fig. 7); stylus length about equal to length

of scape. Palpus dark, slender, bearing numerous long,

dark setae. Clypeus bare, with greyish pruinescence; la-

brum dark and glossy, nearly 1.5 times longer than eye

height; labellum dark and bearing many pale setae, lon-

ger than palpal setae.

Thorax dark with dense grey pruinescence; grey-

ish-white vittae between acrostichal and dorsocentral

rows; dark vittae beneath acrostichal and dorsocentral

rows extending to prescutellar depression; posterior cor-

ner of postpronotal lobe and postalar ridge brownish.

Prosternum bare; proepisternum at fusion point with

prosternum with several long, slender setae; upper part

of proepisternum in front of anterior spiracle bare. An-

tepronotum with dense row of stout setae. Postpronotal

lobe with 1—2 outstanding setae, clothed in numerous

long, slender setae; acrostichal setae biserial, subequal in

length to dorsocentral setae (Fig. 10); dorsocentral setae

multiserial, increasing in length posteriorly, prescutellar

seta slightly shorter than lateral scutellar seta; presutural

supra-alar clothed with numerous slender setae, similar

to dorsocentral setae, occasionally with 1—2 outstanding

setae (posthumeral); numerous long, slender anterior

notopleural setae, similar to presutural supra-alars; 3—4

Bonn zoological Bulletin 67 (2): 129-143

stronger posterior notopleural setae; 2—3 prealar setae

and 1 postsutural supra-alar seta, with numerous shorter

setae; 1 postalar seta; 1 long apical pair and 2 shorter lat-

eral pairs of marginal scutellar setae. Laterotergite with

cluster of long, dark setae. Anterior and posterior spira-

cles brownish.

Legs short, slender, brown with grey pruinescence on

coxae. Fore coxa with row of long slender anterolateral

setae; lateral regions of mid and hind coxae with similar

setae. Fore femur with row of fine anteroventral setae,

shorter than width of femur. Fore tibia densely clothed in

setae, dorsal setae pennate, subequal in length to width of

tibia; ventral setae slightly pennate, shorter than width of

tibia; apex with several stout subapical setae. Mid femur

with dense white ventral pile; anteroventral row of short,

even-length setae; posteroventral row of pennate setae

nearly as long as width of femur; dorsal setae slightly

pennate. Mid tibia with anterodorsal and posteroventral

row of pennate setae; 1 stout anterodorsal seta at mid-

length and several subapical setae. Hind femur (Fig. 9)

with dense white ventral pile; dorsal margin with pennate

setae; posteroventral row of pennate setae longer than

dorsal row; anteroventral row of setae short and slender.

Hind tibia with dorsal and ventral pennate setae, shorter

than width of tibia; 3-4 anterodorsal and posterodorsal

setae; 1 long seta in posteroapical comb. Tarsomere | of

fore and mid legs with short dorsal pennate setae; hind

leg with 34 stout anterodorsal and posterodorsal setae,

longer than tarsomere; 3-4 stout anteroventral and pos-

teroventral setae, subequal to width of tarsomere.

Wing infuscate; pterostigma elongate, distinct; long

basal costal seta present. Cell dm shorter than length of

cell bm (Fig. 9); CuA+CuP reaching wing margin with

weakening at mid-length; alular incision acute; margin of

calypter with long brown setae. Halter brown.

Abdomen shiny with reddish lateral margins and dark

posterior margins; setae dark and numerous. Cercus long

and slender, with fine setae.

Male. Unknown.

Geographic distribution. This species is possibly re-

stricted to the southern Appalachian Mountains of Geor-

gia and North Carolina (USA).

Remarks. Rhamphomyia ecetra is assigned to the

R. (Pararhamphomyia) plumifera group sensu Saigu-

sa (unpubl. data) (or perhaps R. obscura group sensu

Bartak & Kubik 2009) and appears very similar to the

species complex of R. brevis Loew, 1861/R. corvina

Loew, 1861. Rhamphomyia ecetra is characterized by

three pairs of scutellar setae and most specimens of the

brevis/corvina complex have two pairs of scutellar setae.

Males from Georgia are required to make further conclu-

sions concerning the identification of this species.

©ZFMK

Revision of Francis Walker’s female types of North American Rhamphomyia Meigen 135

$.a- :

Pacachamphowysa

det.K.G.V.Smith,1966

®

x

;

Que of Walkers | tYbe

= Series go nained,

Malle Maer. : fais” BMNH(E) #

-Mallos 246922

ay

-

* .

NHMUK 010210625 1 3 s!

a

t

7 Rey ETDS i oy: aa

- ig

eee ad

Figs 11-15. Rhamphomyia mallos Walker. 11. Holotype, female, fore femur, head, lateral view; 12. Holotype, antenna, head, ante-

rior view; 13. Holotype labels, left two labels underside of corresponding labels in centre row; 14. Holotype, lateral view; 15. Hind

leg, posterior view.

136 Bradley J. Sinclair & Toyohei Saigusa

Rhamphomyia mallos Walkker, 1849

(Figs 11-15)

Rhamphomyia mallos Walker, 1849: 502. Type locality:

St. Martin’s Falls [Ogoki], Albany River, Ontario, Can-

ada.

Type material examined. Rhamphomyia mallos: HO-

LOTYPE 9, labelled (Fig. 13): “Type [green margined

circle]”; “44/ ?? [underside: Hudson’s/ Bay]’; “One

of Walkers/ series so named./ EAW [on reverse side:

“Rhamphomyia/ mallos/ Walk.]”; “Mallos,”; “St. Mar-

tin’s Falls,”; “s.g./ Pararhamphomyia/ det. K.G.V. Smith,

1966”; “Holo-/ type [red margined circle]; “BMNH(E)

#/ 246922”; “NHMUK0O10210625 [data matrix code]”

(BMNH). Holotype is missing the left foreleg, fore tib-

ia and tarsi and left hind tarsomere 2—5; the left wing is

slide mounted (see Smith 1971, pl. 3, fig. 2).

Diagnosis. The holotype female is characterized by pale

legs, bare prosternum, pennate setae on mid- and hind-

legs and thickened setae dorsally on fore femur; postped-

icel short, rounded basally; abdomen with pennate setae

laterally on segments 3 and 4 (on pleura?).

Redescription. Wing length 7.3 mm. Female. Head

dark in ground-colour, with greyish pruinescence on

face, frons, postgena and occiput: oral margin pruines-

cent. Dichoptic, ommatidia with uppermost smaller.

Frons and face broad, subequal in width; frons with row

of setulae along eye margin, uppermost shorter. Ocellar

triangle with pair of ocellar setae. Upper half of occiput

bearing row of stout postocular setae; lower postocular

setae slender and shorter. Occipital setae black and stout;

postgenal setae long and slender. Antenna (Fig. 12) with

scape and pedicel pale brown, postpedicel darker; scape

1.5 times longer than pedicel; postpedicel with broad

base, 4 times longer than pedicel; stylus greater than half

length of postpedicel. Palpus yellowish brown, slender,

bearing numerous long, dark setae. Clypeus not visible:

labrum dark and glossy, nearly 1.3 times longer than eye

height (Fig. 11); labellum dark and bearing many long,

dark setae.

Thorax dark with dense grey pruinescence; brownish

vittae beneath acrostichal and dorsocentral rows; later-

al scutum with apparent brownish ring encircling base

of major setae; posterior corner of postpronotal lobe and

postalar ridge brownish. Prosternum bare; proepisternum

at fusion point with prosternum with 3 long, slender setae;

upper part of proepisternum in front of anterior spiracle

bare. Antepronotum with row of stout setae. Postpronotal

lobe with 1 outstanding seta, clothed in numerous long

and short, slender setae; acrostichal setae biserial, sub-

equal in length to dorsocentral setae; dorsocentral setae

uniserial, biserial anteriorly, prescutellar seta subequal in

length to scutellar setae; dorsocentral row curved towards

Bonn zoological Bulletin 67 (2): 129-143

postpronotal lobe anteriorly; 3 presutural supra-alar setae

(posthumeral) and 2 posterior presutural supra-alar setae;

4—5 anterior notopleural setae and 3 posterior notopleural

setae in oblique row; 3-4 prealar setae; 1 postsutural su-

pra-alar setae; 1 postalar seta; 2 pairs of marginal scutel-

lar setae. Laterotergite with cluster of long, dark setae.

Anterior and posterior spiracles yellowish brown, conco-

lourous with halter.

Legs long, pale or yellowish brown with very thin grey

pruinescence on coxae. Fore coxa with row of long slen-

der anterolateral setae; lateral region of mid and hind cox-

ae with similar setae. Fore femur with row of anteroven-

tral and posteroventral setae; stronger anterodorsal setae

on basal third. Fore tibia lost. Mid femur with antero-

ventral row of short, even-length setae, longer near base;

posteroventral row of pennate setae nearly half length of

femur width; anterodorsal setae narrowly pennate, slight-

ly shorter than setae of posteroventral row. Mid tibia with

anterodorsal and posteroventral row of pennate setae;

row of anteroventral setae short. Hind femur with row of

anterodorsal and posteroventral pennate setae (Fig. 15);

setae of anteroventral row short and slender. Hind tibia

with row of anterodorsal and posteroventral pennate se-

tae, shorter than width of tibia; 5-6 posterodorsal setae;

1 long seta in posteroapical comb. Tarsomere | of all legs

without pennate setae.

Wing infuscate, especially along veins (Fig. 14);

pterostigma elongate, distinct; basal costal seta absent.

Cell dm slightly shorter than length of cell bm; CuAt+CuP

reaching wing margin with weakening at mid-length; al-

ular incision nearly right angled; calypter with dark setae.

Halter yellowish brown.

Abdomen dark brown with black setae; pennate setae

laterally on segments 3 and 4. Cercus long and slender,

with fine setae.

Male. Unknown.

Geographic distribution. The label data is likely incor-

rect (see Remarks) and this species is possibly found in

southeastern North America.

Remarks. Rhamphomyia mallos initially appeared very

similar to R. (Dasyrhamphomyia) villipes Coquillett,

1900 on the basis of pennate setae on legs and large size,

but the latter species has more setae on the fore femur,

proboscis distinctly longer, postpedicel elongate and

without abdominal pennate setae. No specimens could be

found that matched the distinctive combination of fea-

tures of this female, especially the abdominal pennate se-

tae. It is doubtful such a large sized species from northern

Ontario would be absent from the CNC, which could in-

dicate that the specimen is mislabelled and possibly orig-

inates from the southeastern United States of America.

Expanding the search, we found that R. mallos is rather

similar to R. testacea Loew, 1862, with reduced setae

on the fore femur and broad postpedicel, but the length

©ZFMK

Revision of Francis Walker’s female types of North American Rhamphomyia Meigen 17,

of the pennate setae of the latter species is longer than

the width of corresponding leg segment (see MCZ type

collection: —http://140.247.96.247/mcz/Species_record.

php?id=13637). There is also an undescribed brownish

Species in the CNC from Virginia, Tennessee, Arkansas

and possibly Mississippi and Texas, where the females

possess abdominal pennate setae, very similar shaped

postpedicel and very long pennate setae on all legs. The

latter species is currently assigned to a group of species

similar to R. pectinata Loew, 1861. Unfortunately all

these species differ from R. mailos in antennal colour,

thoracic chaetotaxy and length of the pennate setae on

the legs.

Rhamphomyia minytus Walker, 1849

(Figs 16-20)

Rhamphomyia minytus Walker, 1849: 502. Type locali-

ty: St. Martin’s Falls [Ogoki], Albany River, Ontario,

Canada.

Rhamphomyia agasicles Walker, 1849: 499. Type local-

ity: St. Martin’s Falls [Ogoki], Albany River, Ontario,

Canada. Syn. nov.

Note about synonymy. Rhamphomyia minytus and

R. agasicles were described in the same paper by Walker

(1849). We consider these names to be subjective syn-

onyms, with the former based on a male and the latter on

a female of the same species. Acting as the First Revis-

er, we select R. minytus as the senior synonym (Article

24.2.2 of the Code, ICZN 1999).

The shiny scutum, anepisternum and abdomen and the

broadened hind tarsomere 1 allowed for the association

of the sexes. The illustrations of the male terminalia and

hind tarsi by Smith (1971, figs 13, 14) facilitated identifi-

cation of additional males specimens and direct examina-

tion of the male holotype of R. minytus was un-necessary.

Type material examined. Rhamphomyia agasicles:

LECTOTYPE 9, labelled (Fig. 18): “Type [green mar-

gined circle]”; “One of Walkers/ series so named./ EAW

[on reverse side: “Rhamphomyia/ agasicles/ Walk.]”;

“Agasciles,’; “St. Martin’s Falls,”’; “LECTO-/ TYPE

[blue margined circle]; “Pararhamphomyia [written by

Smith]; “BMNH(E) #/ 246915”; “NHMUKO010210622

[data matrix code]” (BMNH). The lectotype is in good

condition, with the left wing slide mounted (see Smith

197 ply fis6).

Additional material examined. CANADA. British

Columbia: Alaska Hwy, mi 392, Summit Lake, 4500

ft, 2-4.vii.1959, E.E. MacDougall (1 9, CNC); same

locality, 5300 ft, 18.vi.1959, R.E. Leech (1 9, CNC)

(Fig. 19); same locality, 5000 ft, 23.v1.1959, R.E. Leech

Bonn zoological Bulletin 67 (2): 129-143

(1 ¢, CNC) (Figs 17, 20); same locality, 4500 ft, 23-24.

vi.1959, R.E. Leech (1 2, CNC).

Diagnosis. Females of this species are distinguished by

the shiny scutum, anepisternum and abdomen; hind fe-

mur with short posterodorsal pennate setae; hind tibia

with short anteroventral setae; hind tarsomere 1 as broad

as apex of hind tibia; wings broad, darkly infuscate with

pale base. Males are distinguished by the broad and se-

tose hind tarsomere 1; base of hind tibia with cluster of

posterior setae; cercus and epandrium narrow and elon-

gate with base of both shiny.

Redescription. Wing length 3.5-3.8 mm. Male. Head

dark in ground-colour, with greyish pruinescence on face,

frons, postgena and occiput; oral margin shiny reddish.

Holoptic, eyes with ommatidia larger on upper half of

eye, smaller on lower half. Frons divergent towards an-

tennal sockets, bare. Margins of face slightly divergent;

greyish pruinescence laterally, glossy medially. Ocellar

triangle shiny with pair of ocellar setae, shorter than post-

ocular setae. Occipital setae posterior to ocellar triangle

stouter than postocular setae. Postocular setae long and

slender; postgenal setae similar to postoculars. Anten-

na dark; scape slightly longer than pedicel; postpedicel

nearly 3x longer than basal width; stylus length about

half length of scape. Palpus dark, slender, bearing several

long, subapical setae. Clypeus bare and glossy; labrum

dark and glossy, slightly longer than eye height; labellum

dark and bearing many dark setae, subequal or longer

than palpal setae.

Thorax dark brown with dense grey pruinescence;

anepisternum shiny; dorsum of scutum shiny, with an-

terior face of postpronotal lobe and lateral margin from

notopleuron to scutellum and prescutellar depression

pruinescent. Prosternum bare; proepisternum at fusion

point with prosternum with several setae; upper part

of proepisternum in front of anterior spiracle bare. An-

tepronotum with row of stiff setae. Postpronotal lobe

with 1 outstanding seta and several shorter, finer setae;

acrostichals biserial, slightly shorter than dorsocentral

setae; dorsocentrals biserial to prescutellar depression,

increasing in length posteriorly, uniserial prescutellar se-

tae shorter than apical scutellar setae, dorsocentral row

curved towards postpronotal lobe anteriorly; 1 presutural

supra-alar seta (posthumeral) and several shorter setulae;

3 notopleural setae, with several setulae; 0 prealar setae;

1 postsutural supra-alar seta; 1 postalar seta; 1 long api-

cal pair and short lateral pair of marginal scutellar setae.

Laterotergite with cluster of long, dark setae. Anterior

and posterior spiracles concolourous with pleura.

Legs dark brown, somewhat shiny. Fore coxa with

row of 5-6 stiff anterolateral setae; lateral regions of

mid and hind coxae with similar setae. Fore femur with

row of fine posteroventral setae. Fore tibia with several

rows of dorsal setae, longer than width of tibia; posterior

©ZFMK

138 Bradley J. Sinclair & Toyohei Saigusa

19 r 20

Figs 16—20. Rhamphomyia minytus Walker. 16. Lectotype of R. agasicles Walker, female, dorsolateral view; 17. Recent specimen,

male terminalia, lateral view; 18. Lectotype of R. agasicles Walker, labels, left label underside of corresponding label in centre row;

19. Recent specimen, female, lateral view; 20. Recent specimen, male hind leg, anterior view. See Additional material examined

section for locality details of recent specimens.

Revision of Francis Walker’s female types of North American Rhamphomyia Meigen 139

face clothed with fine short setae. Mid femur with an-

teroventral and posteroventral rows of stout setae, longer

than width of femur. Mid tibia clothed with stout ventral

setae, shorter than width of tibia. Hind femur (Fig. 4E)

slightly twisted medially on apical third; basal half with

row of posterior setae, subequal to width of femur; api-

cal third with row of posterodorsal setae increasing in

length apically. Hind tibia straight, with anterodorsal se-

tae, increasing in length and stouter apically; base of hind

tibia with cluster of posterior setae; 1 long seta in pos-

teroapical comb. Hind tarsomere | greatly expanded and

flattened, broader than tibia (Fig. 20), with dense dorsal

setae, longer than width of segment; tarsomeres 2 and 3

somewhat swollen with long dorsal setae.

Wing lightly infuscate; pterostigma elongate, lightly

infuscate; without basal costal seta. Cell dm subequal in

length to cell bm; CuA+CuP weak, reaching wing margin

as crease; alular incision right angled; margin of calypter

with brown setae. Halter pale yellowish brown.

Abdomen dark brown, shiny, with longer ventral setae;

dorsolateral setae pale brown. Tergite 8 reduced to half-

length of sternite; sternite 8 with short rounded posterior

margin bearing long setae, longer than length of sclerite.

Terminalia (Fig. 17) dark brown (undissected). Epandri-

um elongate, slightly constricted beyond middle, bearing

many fine setae, longer along ventral margin and apical-

ly; middle outer face with dense setae; lower basal half

polished. Cercus elongate, 3/4 length of epandrium; ex-

panded beyond mid-length; polished on basal half; dorsal

margin at base bearing many short, fine setae; inner sur-

face on apical half with numerous slender setae. Phallus

filamentous, with slight inward curve towards middle,

forming loop extending beyond epandrium and recurved

between cerci; ejaculatory apodeme and hypandrium not

examined in undissected specimen.

Female. Similar to male except frons shiny with pale

lateral setulae, with greyish pruinescence above anten-

nae; hind femur with short posterodorsal pennate setae,

hind tibia with short anteroventral setae; hind tarsomere 1

as broad as apex of hind tibia (Fig. 16); wings broader

than male, darkly infuscate with pale base (Fig. 19).

Geographical distribution. This species is known from

northern British Columbia and northern Ontario (Cana-

da).

Remarks. Rhamphomyia minytus is assigned to the R.

(Pararhamphomyia) caudata (Zetterstedt, 1838) group,

and in North America this species group also includes

R. priapulus Loew, 1861 and R. ursinella Melander, 1928

and several undescribed species.

Bonn zoological Bulletin 67 (2): 129-143

Rhamphomyia poplitea Wahlberg, 1844

(Figs 21-28)

Rhamphomyia poplitea Wahlberg, 1844: 107. Type local-

ity: Quickjock, Sweden.

Rhamphomyia dana Walker, 1849: 502. Type locality: St.

Martin’s Falls [Ogoki], Albany River, Ontario, Cana-

da. Syn. nov.

Rhamphomyia valga Coquillett, 1895: 428. Type locali-

ty: New Hampshire, USA. Syn. nov.

Note about synonymy. Females of this species are rather

distinctive on the basis of the very long pennate setae

(Figs 23-24, 27) and R. dana matched females identified

by Chillcott (1959) as R. valga. Rhamphomyia valga is

assigned to the R. basalis Loew, 1864 group defined on

the basis of the highly modified male hind leg (Chillcott

1959). All species of the R. basalis group were examined

in detailed in both the USNM and CNC and in combina-

tion with the key to females in Chillcott (1959) only one

species (R. valga) matched the female Walker type.

Following the initial identification of R. dana with

R. valga, it was learned that the European species,

R. poplitea should also be assigned to the R. basalis

group. Only digital images of the holotype of R. popli-

tea were studied. On the basis of identical colour of the

abdomen, shape of the male hind leg and male terminalia

R. valga is considered a synonym of R. poplitea.

Type material examined. Rhamphomyia dana. HO-

LOTYPE 9, labelled (Fig. 25): “Type [green margined

circle]; “One of Walkers/ series so named./ EAW [on re-

verse side: “Rhamphomyia/ dana/ Walk. ]”; “Dana,”; “St.

Martin’s Falls,”; “Pararhamphomyia/ det. K.G.V. Smith,

1968”; “Holo-/ type [red margined circle]; “BMNH(E)

#/ 246917”; “NHMUK010210623 [data matrix code]”

(BMNH). The holotype is in good condition, with right

postpedicel missing and left wing slide mounted (see

Smith 1971, pl. 2, fig. 8).

Rhamphomyia poplitea. HOLOTYPE <, labelled

(Fig. 22): “Lp./ in. [Lapponia interior]; “Bhn [Boheman

collection]”; “Typus”; “poplitea, Wahlb. [unit tray la-

bel]” (NHRS).

Rhamphomyia_ valga. HOLOTYPE <, labelled:

“White Mts./ Morrison.”; “Collection/ C.V. Riley”:

“Type/ No. 3217/ U.S.N.M. [red label]”; “Rhamphomy-

ia/ valga/ Coq.” (USNM).

Additional material examined. CANADA. New

Brunswick: Kouchibouguac NP, 5—14.vii.1977, J.R.

Vockeroth, G.A. Calderwood (1 4, 2 29, CNC). New-

foundland & Labrador: Fogo Island, vil.1929, C.W.

Johnson (1 3, CNC); Goose Bay, 24.viii.1947, W.E.

Beckel (1 3, CNC); same locality, 7.vii.1952 (2 99,

CNC). Ontario: Iroquois Falls, 21—30.v1.1987, J.R.

Vockeroth (2 92, CNC); Thunder Bay Distr., Neys Prov.

©ZFMK

140 Bradley J. Sinclair & Toyohei Saigusa

os a 7

Saitama Mast il

‘ j So

, rl

|

Way "is os hal Mo dé Lbs.

22

Figs 21—22. Rhamphomyia poplitea Wahlberg. 21. Holotype, male, lateral view; 22. Holotype label, upper three labels attached to

specimen; lower label pinned to unit tray (photographs: Y. Brodin).

Pk, Dune Trail, 48°46'52"N, 86°36'53"W, MT, yellow

pans, 7—19.vii.2002, M. Buck (2 99, UGIC); Thunder

Bay Distr., Prairie R. at Hwy 17, 38 km E Terrace Bay,

48°48'N, 86°47'W, boggy spruce MT, 15—19.vii.2002,

M. Buck (2 64, 4 28, UGIC). Quebec: Fort Chimo,

22.vi1.1948, R.H. MacLeod (1 2, CNC); same locality,

7-9.vii.1954, J.F. McAlpine, E.E. Sterns (2 64, CNC);

Indian House Lake, 9.vii—15.vili.1954, W.R. Richards

(31 dav, 8 99, CNC) (Figs 24, 27, 28); James Bay

Rte, km 204.5, 50°58'59"N, 77°38'2"W, black spruce/

Sphagnum, MT, 7—-16.vii.2001, M. & B. Buck (1 9,

UGIC) (Fig. 26); James Bay Rte, km 578.4, 53°32'40"N,

77°40'40"W, dry lichen conifer forest, yellow pans,

9-15.vii.2001, M. & B. Buck (1 2, UGIC); Knob Lake,

54°47'N, 66°47'W, 16.vil.1948, E.G. Munroe (1 9,

CNC); La Ferme, 1.vii.1943, A. Robert (1 3, CNC);

Laniel, 19.vi.1939, F.P. Ide (1 9, CNC); same locality,

21.vi.1941, A.R. Brooks (1 <4, CNC); Parke Reserve, Ka-

mouraska, 11—17.vii.1957, G.E. Shewell (3 299°, CNC).

Yukon: Dempster Hwy, mi 87, 8—12.vii.1973, G.&D.M.

Wood (1 2, CNC); Dempster Hwy, Eagle R. crossing;

MT, 9-10.vii.1985, S.A. Marshall (1 ¢, UGIC). FIN-

LAND. Enontekis, Palmén (1 4, 1 9, MZH), Enontekis,

R. Frey (1 9, MZH); Muonio, R. Frey (1 3, MZH); Tien-

su (1 4¢, MZH). UNITED STATES OF AMERICA.

Alaska: King Salmon, Naknek River, 3—14.vit.1952,

J.B. Hartley (1 3, 1 9, CNC).

Diagnosis. Females of this species are distinguished by

long, broad and apically truncate dorsal and ventral pen-

nate setae on the mid and hind femora and mid and hind

Bonn zoological Bulletin 67 (2): 129-143

tibia, longer than width of corresponding leg segment,

and slender dorsal pennate setae on hind tarsomere 1; ab-

domen yellowish brown to pale brown, contrasting with

grey thorax. Males are distinguished by the highly mod-

ified hind legs (characteristic of basalis group) and form

of the male terminalia.

Redescription. Wing length 4.5-4.9 mm. Male. Head

dark in ground-colour, with greyish pruinescence on

face, frons, postgena and occiput; oral margin shiny red-

dish. Holoptic, eyes with ommatidia larger on upper half

of eye, smaller on lower half. Frons divergent towards

antennal sockets, bare. Margins of face slightly diver-

gent. Ocellar triangle with pair of ocellar setae, shorter

than postocular setae. Upper third of occiput bearing row

of stout postocular setae, longer than ocellar setae; lower

postocular setae slender and shorter. Occipital setae black

and stout, extending ventrally; postgenal setae similar to

occipital setae. Antenna dark; scape slightly longer than

pedicel; postpedicel nearly 3x longer than basal width;

stylus length about equal to length of scape. Palpus dark,

Slender, bearing single long, subapical seta. Clypeus

bare, with greyish pruinescence; labrum dark and glossy,

slightly longer than eye height; labellum dark and bear-

ing many dark setae, shorter than palpal seta.

Thorax dark with dense grey pruinescence; brownish

vittae beneath acrostichal and dorsocentral rows extend-

ing to prescutellar depression; posterior corner of post-

pronotal lobe and postalar ridge brownish. Prosternum

bare; proepisternum at fusion point with prosternum with

1 stiff seta and sometimes several thinner setae; upper

©ZFMK

Revision of Francis Walker’s female types of North American Rhamphomyia Meigen 141

| N

;

’

f \

4

\

S.

~

N

.

y

a

23

ea

atacham pho mya

det. K.G.V. Smith, 196

2/ 28

Figs 23—28. Rhamphomyia poplitea Wahlberg. 23. Holotype of R. dana Walker, female, lateral view; 24. Recent specimen, female

hind leg, anterior view; 25. Holotype of R. dana Walker, labels, left label underside of corresponding label in centre row; 26. Recent

specimen, male terminalia, lateral view; 27. Recent specimen, female, lateral view; 28. Male hind leg, anterior view. See Additional

material examined section for locality details of recent specimens.

©OZFMK

Bonn zoological Bulletin 67 (2): 129-143

142 Bradley J. Sinclair & Toyohei Saigusa

part of proepisternum in front of anterior spiracle bare.

Antepronotum with row of setae. Postpronotal lobe with

1 outstanding seta and several shorter, finer setae; acro-

stichals biserial, slightly shorter than dorsocentral setae;

dorsocentrals uniserial, increasing in length posteriorly,

prescutellar seta slightly shorter than scutellar setae, dor-

socentral row curved towards postpronotal lobe anterior-

ly; 1 presutural supra-alar seta (posthumeral) and 1 poste-

rior supra-alar seta (opposite anterior notopleural) longer

and stouter; 1 anterior and 3—4 posterior notopleural se-

tae, with several setulae near anterior notopleural seta;

2 prealar setae; 1 postsutural supra-alar setae; 1 postalar

seta; 1 long apical pair and short lateral pair of margin-

al scutellar setae. Laterotergite with cluster of long, dark

setae. Anterior and posterior spiracles yellowish brown.

Legs short, brown with pale “knees”. Fore coxa with

row of 5-6 stout anterolateral setae; lateral regions of

mid and hind coxae with similar setae. Fore femur with

row of fine anteroventral setae. Fore tibia with pale ven-

tral pubescence. Mid femur with anteroventral row of

short, even-length stout setae; posteroventral row of stout

setae longer, greater than half width of femur. Mid tibia

with anteroventral and posteroventral rows of very short,

stout setae, shorter than ventral setae of femur. Hind

femur greatly swollen, of uniform width on apical half

(Figs 21, 28); anteroventral row of setae weak on basal

half, stronger preapically; posteroventral row slender and

long basally, decreasing evenly to setulae at apex. Hind

tibia geniculate at base; straight, without crest of flattened

setae and not flattened; setae of anteroventral row stout;

anterodorsal setae numerous, some at least twice width of

tibia; 1 long seta in posteroapical comb. Tarsomere 1 of

all legs slender, with dense ventral setae.

Wing lightly infuscate (Fig. 21); pterostigma elongate,

normally pale; short basal costal seta present. Cell dm

subequal in length to cell bm; CuA+CuP unpigmented,

reaching wing margin as crease; alular incision obtuse;

margin of calypter with brown setae. Halter pale yellow-

ish brown.

Abdomen brown, paler than thorax; posterior margins

of tergites and sternites paler; clothed in dark setae, lon-

ger along posterior margin; setae finer and somewhat

paler on ventrolateral margin. Tergite 7 narrower than

sternite 7, with posterolateral and posterior margin more

thickly sclerotized. Tergite 8 reduced to half-length of

sternite 8, narrowed laterally; posterior margin of sternite

8 with long setae, longer than length of sclerite 8. Ter-

minalia (Fig. 26) lighter brown, with apex of epandrium

darkened. Hypandrium slender, wrapping around base of

phallus. Epandrium subtriangular, produced posterior into

slender rounded projection closed in very short, stout se-

tae; posterior half clothed in long setae, some longer than

half-length of epandrium. Subepandrial lobe digitiform;

slightly arched with rounded apex; with 6-8 long setae

on apical half. Cercus subrectangular with truncate apex,

shorter than subepandrial lobe. Phallus broad and even

Bonn zoological Bulletin 67 (2): 129-143

throughout basal half; apical half strongly tapered and

arched. Ejaculatory apodeme subtriangular, very large,

as broad as length of hypandrium; horizontal lamella on

posterior margin.

Female. Similar to male except: wing darkly infuscate

(Fig. 27), broader than in male. Legs with long, broad

and apically truncate dorsal and ventral pennate setae on

the mid and hind femora and mid and hind tibiae, longer

than width of corresponding leg segment, and slender

dorsal pennate setae on hind tarsomere | (Figs 23-24,

27). Abdomen yellowish brown to pale brown, contrast-

ing with grey thorax.

Geographic distribution. This species ranges across

Canada and North America, from New Hampshire to

Alaska (see Additional material examined). In the Pa-

laearctic Region, R. poplitea occurs in Sweden, Finland

and across Russia (Shamshev 2016).

Remarks. This species (as valga Coquillett) was includ-

ed in the R. basalis group by Chillcott (1959). This spe-

cies group 1s defined by the enlarged femur-tibia joint of

the male hind leg (Fig. 28) (Chillcott 1959) and is as-

signed to the subgenus Pararhamphomyia Frey, 1922.

Chillcott (1959) listed three European species that appear

to be assigned to this species group on the basis of male

genitalia, but apparently overlooked R. poplitea.

Rhamphompia tristis Walker, 1857

Rhamphomyia tristis Walker, 1857: 148. Type locality:

“United States”.

Type material. Not available (see Smith 1971).

Remarks. According to Smith (1971), the type specimen

has not been identified in BMNH and the Oxford Univer-

sity Museum of Natural History.

Acknowledgements. We are indebted to the following curators

and their respective institutions for the loan of specimens: Dun-

can Sivell (BMNH), Pekka Vilkamaa (MZH), Steve Marshall

(UGIC), Yngve Brodin (NRMS) and Allen Norrbom (USNM).

Torsten Dikow (USNM) kindly assisted BJS during a visit to

the Smithsonian Institution. Igor Shamshev (St. Petersburg)

and Miroslav Bartak (Prague) are thanked for comments on an

earlier draft.

REFERENCES

Bartak M (2002) Nearctic species of Rhamphomyia subgenus

Megacyttarus (Diptera: Empididae). Acta Universitatis Caro-

linae, Biologica 46: 3-215

Bartak M, Kubik S (2009) Two new east Palaearctic Rhampho-

myia (Pararhamphomyia) (Diptera: Empididae). Entomolog-

ical News 120: 76-86

©ZFMK

Revision of Francis Walker’s female types of North American Rhamphomyia Meigen 143

Chillcott JG (1959) Studies on the genus Rhamphomyia Mei-

gen: a revision of the Nearctic species of the basalis group of

the subgenus Pararhamphomyia Frey (Diptera: Empididae).

The Canadian Entomologist 91: 257—275

Coquillett, D.W. (1895) Revision of the North American Em-

pidae — A family of two-winged insects. Proceedings of the

United States National Museum 18 (1896), 387-440

Cumming JM, Sinclair BJ, Brooks SE, O’Hara JE, Skeving-

ton JH (2011) The history of dipterology at the Canadian

National Collection of Insects, with special reference to the

Manual of Nearctic Diptera. In: Festschrift commemorating

the coordinators of the Manual of Nearctic Diptera and their

contributions to building the Canadian National Collection

of Insects. Part 1. The Canadian Entomologist 143: 539-577

Cumming JM, Wood DM (2017) [Chapter] 3. Adult morphol-

ogy and terminology. Pp. 89-133 in: Kirk-Spriggs AH &

Sinclair BJ (eds) Manual of Afrotropical Diptera. Volume 1.

Introductory chapters and keys to Diptera families. Suricata

4, South African National Biodiversity Institute, Pretoria

Danks HV (1981) Arctic arthropods. A review of systematics

and ecology with particular reference to the North American

fauna. Entomological Society of Canada, Ottawa

Evenhuis NL (2018) The life and work of Francis Walker

(1809-1874). Fly Times, Supplement 2: 1-101

Loew H (1861) Diptera Americae septentrionalis indigena.

Centuria prima. Berliner Entomologische Zeitschrift 5: 307—

359

Melander AL (1965) Family Empididae (Empidae, Hybotidae).

Pp. 446-481 in: Stone A, Sabrosky CW, Wirth WW, Foote

RH & Coulson JR (eds) A Catalog of the Diptera of America

Bonn zoological Bulletin 67 (2): 129-143

north of Mexico. United States Department of Agriculture,

Agriculture Handbook No. 276

Poole RW (1996) Diptera. Pp. 15-604 in: Poole RW & Gen-

tili P (eds), Nomina Insecta Nearctica. A Check List of the

Insects of North America. Volume 3: Diptera, Lepidoptera,

Siphonaptera. Entomological Information Services, Rock-

ville, MD

Saigusa T (1963) Some new species of the genus Rhamphomy-

ia from Japan, with descriptions of two new subgenera (Di-

ptera, Empididae). Sieboldia 3 (1): 131-166

Saigusa T (2012) A new Asio-Nearctic subgenus of Rhampho-

myia (Diptera: Empididae: Empidinae). The Canadian Ento-

mologist 144: 291-322

Shamshev IV (2016) An annotated checklist of empidoid flies

(Diptera: Empidoidea, except Dolichopodidae) of Russia.

Proceedings of the Russian Entomological Society 87: 3—183

Sinclair BJ, Vajda EA, Saigusa T, Shamshev IV & Wheeler TA

(in prep.) Rhamphomyia Meigen of the Canadian Arctic Ar-

chipelago, Greenland and Iceland (Diptera: Empididae)

Smith KGV (1971) A revision of Francis Walker’s types of

North American Empididae (Diptera). Bulletin of the Brit-

ish Museum (Natural History), Entomology 26 (8): 347-370,

pls 1-3

Wahlberg E (1844) Nya Diptera fran Norrbotten och Lulea

Lappmark. Ofversigt af Kongl. Vetenskaps-Akademiens

Forhandlinger 1: 106-110

Walker F (1849). List of specimens of dipterous insects in the

collection of the British Museum. Part II, London.

Walker F (1857) Characters of undescribed Diptera in the col-

lection of W.W. Saunders, Esq., F.R.S., &c. Transactions of

the Entomological Society of London 4: 119-158

©ZFMK

BHL

i

Blank Page Digitally Inserted

Bonn zoological Bulletin 67 (2): 145-169

2018 - Jansen J.J.F.J.

https://doi.org/10.20363/bzb-2018.67.2.145

ISSN 2190-7307

http://www.zoologicalbulletin.de

Research article

urn:|sid:zoobank.org:pub:7CE40B75-3E2E-41B7-9705-E5E7BBE2AB77

Specimen labelling errors:

birds collected on the Falkland Islands prior to 1861,

now in Naturalis Biodiversity Center, Leiden, Netherlands

Justin J. F. J. Jansen

Naturalis Biodiversity Center, Postbus 9517, NL-2300 RA Leiden, Netherlands

E-mail: justin jansen@naturalis.nl

urn:lsid:zoobank.org:author:4C F8D990-C4E6-4073-9B9 1 -6424E0C 1BA54

Abstract. Historical museum specimens are often documented by labels or by handwritten notations on the pedestal

underside. However, mistakes and misinterpretations of specimen data often occur due to a complete lack of such data or

in the transcription of data from original sources (either in the form of original labels or personal communications). This

paper discusses the errors and omissions in data found on 89 specimens collected in the Falkland Islands before 1861,

present or formerly present in Naturalis Biodiversity Center (Leiden, Netherlands), resulting from label substitution and

transcription errors. Errors and omissions in data include location, collector, collecting locality and date. The corruption

of original data shows that the gravest error made at that time was the disposal of the original labels.

Key words. Bird collection, Ornithology, Falkland Islands, Charles Abbott.

INTRODUCTION

Ornithology in the early nineteenth century flourished

worldwide, but up to 1861 relatively few expeditions

with natural history objectives collected birds in the Falk-

land Islands (Jansen & van der Mije 2015). The Falklands

rose to fame when Charles Darwin published accounts

of the birds and mammals encountered (Darwin 1839,

1859) when HMS Beagle visited in 1833 and 1834. We

find published accounts of other expeditions up to 1861

(Lesson & Garnot 1826-30, Darwin 1839, Freycinet

et al. 1837, MacGillivray 1852), when Captain Charles

Compton Abbott left the Falklands (Abbott 1860, 1861).

To get a better understanding of the birds collected in

the Falkland Islands, all specimens in the Naturalis Bio-

diversity Center, Leiden, Netherlands (hereafter Natu-

ralis) or noted in the literature that were collected in the

Falkland Islands or arrived at Naturalis prior to 1 January

1862, were analysed. In total, 89 specimens, represented

by mounts or skins, were examined. A number of these

specimens have been previously cited by earlier authors

(Schlegel 1862a, b, c, d, 1863a, b, c, d, 1864, 1865, 1866,

1867, van Grouw & Steinheimer 2008).

The labelling of bird specimens in Naturalis has been

the subject of research by a few previous authors, with

a special focus on the removal of original labels (Mees

1953, Mees & Fisher 1986, Rasmussen & Prys-Jones

2003). However, their focus was on a single labelled bird

of Edgar Leopold Layard’s (Mees & Fisher 1986) and

Karl B.H. von Rosenberg’s birds from Indonesia (Mees

1953, Rasmussen & Pr¥ys-Jones 2003). No extensive re-

Received: 29.12.2017

Accepted: 28.11.2018

search has yet been carried out on a large series of birds

with regard to their label substitution and transcription

errors. The research conducted for this paper was carried

out in order to establish how much such ‘vandalism’ of

Falkland Islands birds has occurred in Naturalis.

Jansen & van der Mije (2015) noted that various sourc-

es in addition to the known expeditions collected birds

and mammals in the Falkland Islands. The original sourc-

es are therefore often hard, or in most cases even impos-

sible, to trace.

Major expeditions and collectors that collected birds in

the Falklands include:

¢ L’Uranie, 15 February—27 April 1820. ‘65+’ birds

(appendix 1) that arrived at the Museum national

d’Histoire naturelle, Paris (hereafter MNHN) have

annotations that they were collected in the Falklands

(MNHN archives).

¢ La Coquille, 18 November—18 December 1825. 42

birds (appendix 2) that arrived at the MNHN have

annotations that they were collected in the Falklands

(MNHN archives).

° HMS Beagle, 1 March—-S April 1833 and

11 March—7 April 1834. 33 birds were collected by

Charles Darwin and Syms Covington on the Falk-

lands (Steinheimer 2004). And 24 specimens from

the Falklands were collected by Captain Robert Fitz-

roy on the HMS Beagle expedition and donated in by

Sir William Burnett in 1838 (Natural History Muse-

um Tring, UK (herafter NHMUK)).

Corresponding editor: T. Topfer

Published: 05.12.2018

146 Justin J. F. J. Jansen

¢ HMS Erebus and Terror, 6 April—8 September 1842.

142 birds and eggs donated to the NHMUK by Rob-

ert McCormick in 1890 and 1891 (Sharpe 1906).

¢ HMS Rattlesnake, 5—25 July 1850. Specimens were

presented to the NHMUK by Owen Stanley in 1850,

1854 and 1855, totalling 219 specimens, and 16 birds

by William McGillivray in 1851 (Sharpe 1906).

¢ Charles Compton Abbott (1821-1887), August

1857—December 1860. The Hobart (Tasmania)

born officer entered the army at 22 July 1844, and

served in the 13", 20", 75", and 47" Regiments. He

commanded the detachment at the Falkland Islands

for four or five years (became captain in March

1858). He returned to Australia, and after serving

in the North Glouchester Militia in 1862-69, he

was transferred to the Queen’s Country Royal Ri-

fles, he retired from service in 1876. Abbott was in

command of the British forces stationed at Stanley

on East Falkland. Abbott made various excursions

into the interior, both in the north and in the south

of East Falkland and lost no opportunity to collect

eggs and skins (Darwin 1871, Jansen & van der Mije

2015). Specimens collected by Abbott ended up in

a number of collections. No less then 46 specimens

(possibly as many as 76) in Naturalis were received

directly from Abbott, as indicated by Abbott being

mentioned on the label and/or pedestal.

27 birds in Naturalis acquired from Gustav Adolph Frank

Sr., for which there are no specific collectors mentioned

on their labels, most likely originate with Abbott also,

although this remains speculative. While Abbott may

have sold his birds to Frank and John Gould, he probably

did so only to Gould, who in turn sold them to Frank,

and Frank in turn to others such as the merchant Charles

Jamrach and the Zoological Society of London (ZSL).

Of four Falkland birds examined in the NHMUK from

the ZLS, only one mentions Abbott as its source on its

label (though this is not the original label). Specimens

from Abbott arrived in the NHMUK via John Henry

Gurney (in particular raptors), an 1859 shipment from

Gould (comprising 132 birds and eggs) and from the ZLS

(Sharpe 1906). The Abbott labels are easy recognizable

as they are printed labels, with Abbott’s own handwriting

on them (Figs. 2, 3).

This paper is not intended to re-label the specimens

from archival research and no such research was execut-

ed, although known information is given.

MATERIAL AND METHODS

The 89 birds discussed in this article were found by re-

searching the relevant literature (Schlegel 1862a, b, c,

d, 1863a, b, c, d, 1864, 1865, 1866, 1867) and by visits

to Naturalis on 18 December 2012, 11 February 2013

Bonn zoological Bulletin 67 (2): 145-169

and 3 December 2015. Additional visits were made to

Museum national d’Histoire naturelle, Paris (hereafter

MNHN) on 2-12 April 2013 and to the Natural History

Museum, Tring (hereafter NHMUK) on 12-13 Novem-

ber 2013, 18 June 2015 and 23 March 2016. These visits

were made to establish if original labels on Falkland Is-

lands specimens were present. 25 Falkland specimens in

MNHN were examined and 103 Falkland specimens in

NHMUK. Literature was also consulted to find addition-

al information on dates when and where specific speci-

mens were collected, and by whom.

Aware that specimens are assigned to “Falklands” or to

“East Falkland” on the basis of label data, could well be

the location from where it is mailed/shipped to Europe.

However, besides some penguin species no odd records

were found. Research revealed that the main collector

Abbott is known to travelled extensively on East Falk-

land and collected all kinds of natural history material

(Lanjouw & Staffleu 1954), and the known expeditions

did not ship specimens from the Falklands to Europe.

The two research questions are:

¢ What were the transcription errors made on the

available sources?

¢ Did the specimens at Naturalis once had these orig-

inal labels?

RESULTS

The specimens

No synonyms are given, only the actual English and Lat-

in name. The framework as set by Steinheimer 2010 is

followed with some slight modifications.

Abbreviations

Loc = locality,

Col =collected by,

Acq = Acquisition history,

Tax = taxidermy,

Pub = Publications mention the specimen or collecting

trip.

Taxonomy and nomenclature follow del Hoyo & Collar

(2014, 2016).

ANATIDAE

Upland Goose Chloephaga picta leucoptera (J.F.

Gmelin, 1789) [RMNH.AVES.230417]. Loc: Falklands

(ca. 51° 41' 0" S, 59° 10' 0" W). Date: November—De-

cember 1822. Col: voyage La Coquille, R.P. Lesson / P.

Garnot. Age/sex: adult 4. Acq: June 1835, in exchange

with the Paris Museum. Status: extant. Accessed: 18

©ZFMK

Specimen labelling errors: birds collected on the Falkland Islands prior to 1861 147

December 2012. Tax: mounted. Pub: Lesson & Garnot

1826-1830; Schlegel 1866.

Pedestal label: [two hands] Cat n°1 / /eucoptera / Chloéphaga

reat | (Gmelin) / 4 / verzameld tijdens / de reis van

“La Coquille” / Falkland eilanden [two hands].

Pedestal underside: [one hand] Anser magellanicus ¢ / Ans-

er leucopterus / Anas leucoptera Lath / Oie a ailes blanches

Buff vol 9 / Brown III talo 40 / Cat N° / voy La Coquille /

Malouines.

Schlegel 1866 (Tome VI, 31: 99-100): Anser magellanicus, Cat

1, ¢ adulte, Malouines, voyage de /a Coquille.

Remarks: The collection site on the label (“Falkland eilan-

den’’) differs from that on the underside and Schlegel 1866

(“Malouines”). Several Latin names are cited, those on the

label and underside differ; the primary name on the latter

however agrees with Schlegel 1866. The notation of “adulte”

is present only in Schlegel 1866, and is lacking on the label

and underside. Gmelin is noted as the name source on the

label, Buffon as the source on the underside. Only Schlegel’s

handwriting appears under the base of the pedestal.

Mentioned in the June 1835 exchange list between Paris and

Leiden as ‘Oie des torres Magellanicus’. The stand is ‘new’

(e.g. it has Temminck’s handwriting) and does not contain

any original information / handwriting by Dufresne (the

mounted birds from La Coquille now present in MNHN have

Dufresne’s handwriting on the pedestal underside). A possi-

ble explanation is that it arrived in Leiden as an unmounted

skin and was subsequently mounted there.

Upland Goose Chloephaga picta leucoptera (J.F.

Gmelin, 1789) [RMNH.AVES.230418]. Loc: Falklands

(ca. 51° 41'0" S, 59° 10' 0" W). Date: —. Col: —. Age/sex:

adult 4. Acq: —. Status: extant. Accessed: 18 December

2012. Tax: mounted. Publ: Schlegel 1866.

Pedestal label: [two hands]. Cat n°2 / Jeucoptera / Chloéphaga

mageHaniea | (Gmelin) / 3 / Falkland eilanden.

Pedestal underside: [one hand] Anser magellanicus ¢ / Anser

leucopterus | Anas leucoptera Lath. / Oie a ailes blanches

Buff v9 / Brown III talo 40 / Cat N°2 / Malouines [one hand].

Schlegel 1866 (Tome VI, 31: 99-100): Anser magellanicus, Cat

2, 3 adulte, Malouines.

Remarks: The collection site on the label (“Falkland eilan-

den’’) differs from that on the underside and Schlegel 1866

(“Malouines”). Several Latin names are cited; those on the

label and underside differ; the primary name on the latter

however agrees with Schlegel 1866. The notation of “adulte”

is present only in Schlegel 1866, and is lacking on the label

and underside. Gmelin is noted as the name source on the

label, Buffon, Latham and Brown as sources on the under-

side. Only Schlegel’s handwriting appears under the base of

the pedestal.

In June 1835 two specimens where sent from the Paris Mu-

seum as part of an exchange; this could be one of those birds.

Most likely collected by Lesson / Garnot in Nov—Dec 1822.

One specimen of Chloephaga picta is missing from HMS

Beagle (Steinheimer 2004).

Upland Goose Chloephaga picta leucoptera (J.F.

Gmelin, 1789) [RMNH.AVES.230419]. Loc: Falklands

(ca. 51° 41' 0" S, 59° 10' 0" W). Date: —. Col: —. Age/sex:

adult 2. Acq: —. Status: extant. Accessed: 18 December

2012. Tax: mounted. Pub: Schlegel 1866.

Bonn zoological Bulletin 67 (2): 145-169

Pedestal label: [two hands] Cat n°3 / leucoptera / Chloéphaga

mageHaniea / (Gmelin) / 9 / Falkland eilanden.

Pedestal underside: [one hand] Anser magellanicus 9 / Anas

magellanica Lath / Oie der Terres magellaniques Buff / pl

Pub / 1006 / unreadable femelle / Cat N°3 / Malouines.

Schlegel 1866 (Tome VI, 31: 99-100): Anser magellanicus, Cat

3, Femelle adulte, Malouines.

Remark: The collection site on the label (“Falkland eilanden’’)

differs from that on the underside and Schlegel 1866 (“Ma-

louines”). Several Latin names are cited; those on the label

and underside differ; the primary name on the latter however

agrees with Schlegel 1866. The notation of “adulte” is pres-

ent only in Schlegel 1866, and is lacking on the label and

underside. Gmelin is noted as the name source on the label,

Buffon and Latham as sources on the underside. Only Schle-

gel’s handwriting appears under the base of the pedestal.

In June 1835 two specimens where sent from the Paris Mu-

seum as part of an exchange; this could be one of those birds.

Most likely collected by Lesson / Garnot in Nov—Dec 1822.

One specimen of Chloephaga picta is missing from HMS

Beagle (Steinheimer 2004).

Kelp Goose Chloephaga hybrida malvinarum J.C. Phil-

lips, 1916 [RMNH.AVES.230431]. Loc: Falklands (ca.

51° 41' 0" S, 59° 10’ 0" W). Date: —. Col: —. Age/sex:

adult 4. Acq: —. Status: extant. Accessed: 18 Decem-

ber 2012. Tax: mounted. Pub: Schlegel 1866.

Pedestal label: [two hands] Cat n°l / Chloophaga hybrida /

(Meta) / 3 malvinarum Phillips / Falkland eilanden.

Pedestal underside: [one hand] Anser antarcticus 3 ad /

Sparm. mus (ext. note la 9 pl 5) / Cat N°1 / Malouines.

Schlegel 1866 (Tome VI, 31: 98-99): Anser antarcticus, Cat

1, d adulte, au plumage d’un blanc uniforme, Malouines:

bec noir, mais entre les narines et le front rougeatre avec des

taches noires.

Remarks: The collection site on the label (“Falkland eilan-

den’’) differs from that on the underside and Schlegel 1866

(“Malouines”). Several Latin names are cited; those on the

label and underside differ; the name on the latter however

agrees with Schlegel 1866. The notations of “ad” and “adul-

te” are present on the underside and in Schlegel 1866, but

lacking on the label. Phillips is noted as the name source on

the label, Sparmann’s Museum as the source on the under-

side. Only Schlegel’s handwriting appears under the base of

the pedestal.

Kelp Goose Chloephaga hybrida malvinarum J.C. Phil-

lips, 1916 [RMNH.AVES.230432]. Loc: Falklands (ca.

51° 41' 0" S, 59° 10' 0" W). Date: November—December

1822. Col: voyage La Coquille, R.P. Lesson / P. Garnot.

Age/sex: adult 9. Acq: June 1835, in exchange with Paris

Museum. Status: extant. Accessed: 18 December 2012.

Tax: mounted. Pub: Lesson & Garnot 1826-1830; Schle-

gel 1866.

Pedestal label: [two hands] Cat. n°2 / Chloephaga hybrida /

(Metna} / 2 malvinarum Phillips / Verzameld tijdens / de

reis van / “La Coquille” / Falkland eilanden [two hands].

Pedestal underside: [one hand] Anser antarcticus 2 Vieill /

Sparm Mus Carl pl 37 / Less Coquille pl 50 / De antarctique

femelle / Cat N°2 / Coquille Malouines [one hand].

©ZFMK

148 Justin J. F. J. Jansen

Schlegel 1866 (Tome VI, 31: 98-99): Anser antarcticus, Cat 2,

Femelle adulte, Malouines, du voyage de /a Coquille, abso-

lument semblable a I’ individu figure par Lesson.

Remark: The collection site on the label (“Falkland eilanden’’)

differs from that on the underside and Schlegel 1866 (“Ma-

louines”’). Several Latin names are cited; those on the label

and underside differ; the name on the latter however agrees

with Schlegel 1866. The notation of “adulte” is present only

in Schlegel 1866, and is lacking on the label and underside.

Phillips is noted as the name source on the label, Vieillot,

Sparmann’s Museum, and Lesson as the sources on the un-

derside. Only Schlegel’s handwriting appears under the base

of the pedestal.

Mentioned in the June 1835 exchange list between Paris and

Leiden as ‘Oie des torres Magellanicus’ .

Kelp Goose Chloephaga hybrida malvinarum J.C. Phil-

lips, 1916 [RMNH.AVES.230433]. Loc: Falklands (ca.

51° 41' 0" S, 59° 10' 0" W). Date: in either 1858 or 1859.

Col: C.C. Abbott. Age/sex: young 3. Acq: via John Gould

and Gustav Adolph Frank Sr. Status: extant. Accessed: 18

December 2012. Tax: mounted. Pub: Abbott 1860, 1861;

Gould 1859; Schlegel 1866; Sclater 1860, 1861, 1862.

Pedestal label: [two hands] Cat n°3 / Chloephaga hybrida /

Melina / 3 juv. malvinarum Phillips / Abbott coll: / Frank

1860 / Falkland eilanden.

Pedestal underside: [one hand] Anser antarcticus / 3 jong /

Pooten geel, iris en bek zwart / Cat N°3 / Reis van Capitain

Abbott / Frank 1860 / Falkland.

Schlegel 1866 (Tome VI, 31: 98-99): Anser antarcticus, Cat 3,

S dans la livree de passage, Malouines, voyage du Capitaine

Abbot, acquis en 1860: pieds jaunes, vec et iris noirs (Abbot).

Remarks: The collection site on the label (“Falkland eilan-

den”) differs from that on the underside “Falkland”) and

Schlegel 1866 (“Malouines”’). Several Latin names are cited;

those on the label and underside differ; the name on the latter

however agrees with Schlegel 1866. Phillips is noted as the

name source on the label; no name source appears on the un-

derside. Only Schlegel’s handwriting appears under the base

of the pedestal.

Army Captain Charles Compton Abbott was stationed at

Stanley, East Falklands in 1859-1861 in the Falkland Islands

Detachment. He collected many birds and attached printed

labels to his specimens. These labels had his full name and

that of his regiment printed on one side, on the other side, he

noted in pen the year, collecting location and details of the

bare parts. These original labels can be found in NHMUK.

So, contra the underside and Schlegel 1866, Abbott did not

collect while on a “voyage” or “Reis”; moreover, his name is

misspelled in Schlegel 1866. This specimen was acquired via

Gustav Adolph Frank, the Amsterdam-based merchant who,

though he did not collect a single bird himself, exchanged

and purchased birds on a large scale with the likes of the

Paris Museum and John Gould, both known to possess spec-

imens from the Falklands (in particular those collected by

C.C. Abbott). However, Frank is not mentioned as the source

of this specimen in Schlegel 1866.

Ashy-headed Goose Chloephaga poliocephala P.L.

Sclater, 1857 [RMNH.AVES.230436]. Loc: Falklands

(ca. 51° 41'0" S, 59° 10' 0" W). Date: —. Col: —. Age/sex:

Bonn zoological Bulletin 67 (2): 145-169

adult <4. Acq: —. Status: extant. Accessed: 18 December

2012. Tax: mounted. Pub: Schlegel 1866.

Pedestal label: [one hand] Cat n°2 / Chloéphaga poliocephala

/ GR. Gray / 3 / Falkland eilanden.

Pedestal underside: [one hand] Anser poliocephala / Bernicla

inornatus 3 / Cat n°2 / Bernicla inornatus Gray / Gen of

Birds (unreadable) / (unreadable line) / Malouines.

Schlegel 1866 (Tome VI, 31: 101): Anser poliocephala, Cat 2,

Adulte, Malouines.

Remarks: The collection site on the label differs from that

on the underside and Schlegel 1866. Three generic and two

specific names are cited; those on the label and underside

differ; the primary binomial on the latter however agrees

with Schlegel 1866. The identification as ¢ is absent from

Schlegel 1866 and as an adult from the label and underside.

The only name source mentioned is Gray on the underside.

Only Schlegel’s handwriting appears under the base of the

pedestal.

Ruddy-headed Goose Chloephaga rubidiceps P.L.

Sclater, 1861 [RMNH.AVES.230443]. Loc: Falklands

(ca. 51° 41' 0" S, 59° 10' 0" W). Date: —. Col: —. Age/

sex: adult 4. Acq: via Gustav Adolph Frank Sr. Status:

extant. Accessed 18 December 2012. Tax: mounted. Pub:

Schlegel 1866.

Pedestal label: [one hand] Cat n°1. / Chloéphaga rubidiceps /

Sclater / 3 / Falkland eilanden.

Pedestal underside: [one hand] Bernicla / Anser rubidiceps. /

tres 1860 / 3 / Catal N° 1 / Malouines.

Schlegel 1866 (Tome VI, 31: 102): Chloephaga rubiceps, Cat

1, ¢ adulte, Malouines.

Remarks: The collection site on the labeldiffers from that on

the underside and Schlegel 1866. Two generic names are cit-

ed; those on the label and underside differ; the binomial on

the label however agrees with Schlegel 1866. The identifica-

tion as adult is present only in Schlegel 1866. The only name

source mentioned is Sclater on the label. Only Schlegel’s

handwriting appears under the base of the pedestal.

This specimen was acquired through the Amsterdam-based

dealer Gustav Adolph Frank (see Kelp Goose RMNH.

AVES.230433 above) and arrived on 30 April 1860, as indi-

cated on the underside.

Ruddy-headed Goose Chloephaga rubidiceps P.L.

Sclater, 1861 [RMNH.AVES.230445]. Loc: Falklands

(ca. 51° 41' 0" S, 59° 10' 0" W). Date: —. Col: —. Age/sex:

—. Acq: —. Status: extant. Accessed 18 December 2012.

Tax: mounted. Pub: Schlegel 1866.

Label: [one hand] Cat n°3. / Chloéphaga rubidiceps / Sclater /

Falkland eilanden.

Pedestal underside: [one hand] Chloéphaga rubidiceps / Falk-

land.

Schlegel 1866 (Tome VI, 31: 102): not mentioned.

Remark: Notably missed by Schlegel 1866, yet only Schlegel’s

handwriting appears under the base of the pedestal. The only

name source mentioned is Sclater on the label. The collection

site on the label (“Falkland eilanden’’) differs from that on the

underside (“Falkland”).

Falkland Steamer Duck Tachyeres brachypterus

Latham, 1790 [RMNH.AVES.230636]. Loc: Falklands

©ZFMK

Specimen labelling errors: birds collected on the Falkland Islands prior to 1861 149

(ca. 51° 41' 0" S, 59° 10' 0" W). Date: in either 1858 or

1859. Col: C.C. Abbott. Age/sex: adult 4. Acq: via John

Gould and Gustav Adolph Frank Sr.; though not spec-

ified, most likely arrived on 30 April 1860. Status: ex-

tant. Accessed: 18 December 2012. Tax: mounted. Pub:

Abbott 1860, 1861; Gould 1859; Schlegel 1866; Sclater

1860, 1861, 1862.

Pedestal label: absent.

Pedestal underside: [one hand] (unreadable) / Fuligula ci-

nerea | Cat N°1 / N°.3 / Loggerhead Duck / ¢ / Tachyeres

brachyptera (Lath) / Reis van Capitain Abbott / (Frank) 1860

/ Falkland.

Schlegel 1866 (Tome VI, 31: 13): Fuligula cinerea, Cat 1, ¢

adultes, Malouines, voyage de Mr. Abbot, acquis en 1860.

Remarks: The collection site on the underside differs from that

in Schlegel 1866. Two generic and two specific names are

cited, with the primary binomial on the underside agreeing

with Schlegel 1866. Both the underside and Schlegel 1866

identify the specimen as male, but only the latter identifies

it as an adult. The only name source mentioned is Latham

on the underside. Only Schlegel’s handwriting appears under

the base of the pedestal.

This specimen was acquired in 1860 from the Amster-

dam-based dealer Gustav Adolph Frank (see Kelp Goose

RMNH.AVES.230433 above).

Falkland Steamer Duck JTachyeres brachypterus

Latham, 1790 [RMNH.AVES.230637]. Loc: Falkland

(ca. 51° 41' 40" S, 57° 51' 10" W). Date: in either 1858 or

1859. Col: C.C. Abbott Age/sex: adult 3. Acq: via John

Gould and Gustav Adolph Frank Sr.; though not spec-

ified, most likely arrived on 30 April 1860. Status: ex-

tant. Accessed: 18 December 2012. Tax: mounted. Pub:

Abbott 1860, 1861; Gould 1859; Schlegel 1866; Sclater

1860, 1861, 1862.

Pedestal label: absent.

Pedestal underside: [one hand] (unreadable)/ Fuligula cine-

rea | Cat N°2 / Loggerhead Duck / 2 / Tachyeres brachyp-

tera (Lath) / Van Cap Abbott / Falkland Island Detachment /

Frank / Falkland.

Schlegel 1866 (Tome VI, 31: 13): Fuligula cinerea, Cat 2, 9

adultes, Malouines, voyage de Mr. Abbot, acquis en 1860.

Remarks: The collection site on the underside differs from that

in Schlegel 1866. Two generic and two specific names are

cited, with the primary binomial on the underside agreeing

with Schlegel 1866. Both the underside and Schlegel 1866

identify the specimen as female, but only the latter identifies

it as an adult. The only name source mentioned is Latham

on the underside. Only Schlegel’s handwriting appears un-

der the base of the pedestal. The text on the underside clear-

ly indicates the former existence of an original label on the

specimen.

The bird was acquired through the Amsterdam-based dealer

Gustav Adolph Frank and arrived on 30 April 1860 (see Kelp

Goose RMNH.AVES.230433 as for C.C. Abbott).

Falkland Steamer Duck JTachyeres brachypterus

Latham, 1790 [—]. Loc: Falkland (ca. 51° 41' 40" S, 57°

51' 10" W). Date: —. Col: —. Age/sex: —. Acq: —. Status:

Bonn zoological Bulletin 67 (2): 145-169

non-extant; not found on 18 December 2012 or 11 Febru-

ary 2013. Tax: mounted. Pub: Schlegel 1866.

Schlegel 1866 (Tome VI, 31: 13): Fuligula cinerea, Cat 3, Indi-

vidue en mue, Malouines.

Remarks: Present location of specimen and the original col-

lector unknown.

Crested Duck Lophonetta specularioides specularioides

(P.P. King, 1828) [RMNH.AVES.231375]. Loc: Falk-

lands (ca. 51° 41' 0" S, 59° 10' 0" W). Date: either 1858

or 1859. Col: C.C. Abbott. Age/sex: adult 3. Acq: via

John Gould and Gustav Adolph Frank Sr. Status: extant.

Accessed 18 December 2012. Tax: mounted. Pub: Abbott

1860, 1861; Gould 1859; Schlegel 1866; Sclater 1860,

1861, 1862.

Pedestal label: [two hands] Cat n°2 / specularioides King /

Anas eristata-Gmetin / 3 / Abbott coll / Frank 1860 / Falk-

land.

Pedestal underside: [one hand] Anas cristata Gmel / Cat N°3

/ & / eye red, feat and beak lead color / Capt Abbot / Frank

1860 / Falklands eilanden / (unreadable).

Schlegel 1866 (Tome VI, 31: 39): Anas cristata, Cat 3, ¢

adultes, iles Malouines, voyage du Capitaine Abbot, acquis

en 1860, Iris rouge, pieds et bec coleur de plomb.

Remarks: The collection site on the label, underside and in

Schlegel 1866 differ. Two specific names are cited, with the

primary binomial on the underside agreeing with Schlegel

1866. The label, underside, and Schlegel 1866 all identify

the specimen as male, but only the latter identifies it as an

adult. Two name sources are mentioned, King on the label

and Gmelin on the underside. The catalogue number on the

label doesn’t correspond with the catalogue number on the

underside and in Schlegel 1866. Only Schlegel’s handwriting

appears under the base of the pedestal.

The bird was acquired through the Amsterdam-based dealer

Gustav Adolph Frank and arrived on 30 April 1860 (see Kelp

Goose RMNH.AVES.230433 as for C.C. Abbott).

Crested Duck Lophonetta specularioides specularioides

(P.P. King, 1828) [RMNH.AVES.231376]. Loc: Falk-

lands (ca. 51° 41' 0" S, 59° 10' 0" W). Date: either 1858

or 1859. Col: C.C. Abbott. Age/sex: adult 2. Acq: via

John Gould and Gustav Adolph Frank Sr. Status: extant.

Accessed 18 December 2012. Tax: mounted. Pub: Abbott

1860, 1861; Gould 1859; Schlegel 1866; Sclater 1860,

1861, 1862.

Pedestal label: [one hand] Cat n°3 / specularioides King / Anas

cristata-Gmetn / 9 / Abbott coll / Frank 1860 / Falkland

eilanden.

Pedestal underside: [one hand] Anas cristata Gmelin / Cat N°

4/2 / Reis van / Capt Abbot / Frank 1860 / +860 / Oost

Falkland.

Schlegel 1866 (Tome VI, 31: 39): Anas cristata, Cat 4, 9

adultes, iles Malouines, voyage du Capitaine Abbot, acquis

en 1860, Iris rouge, pieds et bec coleur de plomb.

Remarks: The collection site on the label, underside and in

Schlegel 1866 differ. Two specific names are cited, with the

primary binomial on the underside agreeing with Schlegel

1866. The label, underside, and Schlegel 1866 all identify

the specimen as female, but only the latter identifies it as an

adult. Two name sources are mentioned, King on the label

©ZFMK

150 Justin J. F. J. Jansen

and Gmelin on the underside. The catalogue number on the

label doesn’t correspond with the catalogue number on the

underside and in Schlegel 1866. Only Schlegel’s handwriting

appears under the base of the pedestal.

The bird was acquired through the Amsterdam-based dealer

Gustav Adolph Frank and arrived on 30 April 1860 (see Kelp

Goose RMNH.AVES.230433 as for C.C. Abbott).

Silver Teal Spatula versicolor fretensis (P.P. King, 1831)

[RMNH.AVES.231525]. Loc: East Falkland (ca. 51° 41’

40" §, 57° 51' 10" W). Date: either 1858 or 1859. Col:

C.C. Abbott. Age/sex: adult 4. Acq: via John Gould and

Gustav Adolph Frank Sr. Status: extant. Accessed 18 De-

cember 2012. Tax: mounted. Pub: Abbott 1860, 1861;

Gould 1859; Schlegel 1866; Sclater 1860, 1861, 1862.

Pedestal label: [two hands] Anas Cat n°6 / Ouergteduta-ver-

sicolor / 3 / Abbott coll / Frank 1861 O. Falkland eilanden.

Pedestal underside: [one hand] Anas versicolor, Vieillot / Cat

n°6 (major) / 3’ / van Cap. Abott / Frank 1860 Oost Falkland.

Schlegel 1866 (Tome VI, 31: 57): Anas versicolor, Cat 6, &

adulte, ile orientales des Malouines, voyage du Capitaine Ab-

bot, acquis en 1860.

Remarks: The collection site on the label, underside and in

Schlegel 1866 differ. Two generic names (one crossed out)

are cited, with the primary binomial on the underside agree-

ing with Schlegel 1866. The label, underside, and Schlegel

1866 all identify the specimen as female, but only the latter

identifies it as an adult. Only one name source is mentioned,

Vieillot on the label. Only Schlegel’s handwriting appears

under the base of the pedestal.

The bird was acquired through the Amsterdam-based dealer

Gustav Adolph Frank and arrived on 30 April 1860 (see Kelp

Goose RMNH.AVES.230433 as for C.C. Abbott).

Silver Teal Spatula versicolor fretensis (P.P. King, 1831)

[RMNH.AVES.231526]. Loc: East Falkland (ca. 51° 41’

40" S, 57° 51' 10" W). Date: either 1858 or 1859. Col:

C.C. Abbott. Age/sex: adult 2. Acq: via John Gould and

Gustav Adolph Frank Sr. Status: extant. Accessed 18 De-

cember 2012. Tax: mounted. Pub: Abbott 1860, 1861;

Gould 1859; Schlegel 1866; Sclater 1860, 1861, 1862.

Pedestal label: [two hands] Anas Cat n°7 / Ouerquedulta versi-

color / €Vieillot} / 2 / Abbott coll: / Frank 1860 O. Falkland

eilanden.

Pedestal underside: [one hand] Anas versicolor / (major) /

Catal. N°7. / 2 / Beak base yellow, remainder (unreadable)

(Abbot) / van Cap. Abott / (Frank 1860) Oost Falkland.

Schlegel 1866 (Tome VI, 31: 57): Anas versicolor, Cat 7, 9

adulte, ile orientales des Malouines, voyage du Capitaine Ab-

bot, acquis en 1860.

Remarks: The name giving of the location of collecting is dif-

ferent in underside, label and Schlegel 1866. The sexing of

the specimen is noted in all both sources, the ageing however

is only present in Schlegel 1866. Only Schlegel’s handwrit-

ing under the base of the pedestal.

The collection site on the label, underside and in Schlegel

1866 differ. Two generic names are cited (one crossed out),

with the primary binomial on the underside agreeing with

Schlegel 1866. The label, underside, and Schlegel 1866 all

identify the specimen as female, but only the latter identifies

Bonn zoological Bulletin 67 (2): 145-169

it as an adult. No name sources are mentioned. Only Schle-

gel’s handwriting appears under the base of the pedestal.

The bird was acquired through the Amsterdam-based dealer

Gustav Adolph Frank and arrived on 30 April 1860 (see Kelp

Goose RMNH.AVES.230433 as for C.C. Abbott).

Cinnamon Teal Spatula cyanoptera cyanoptera (Vieil-

lot, 1816) [RMNH.AVES.231631]. Loc: East Falkland

(ca. 51° 41' 40" S, 57° 51' 10" W). Date: either 1858

or 1859. Col: C.C. Abbott. Age/sex: adult 3. Acq: via

John Gould and Gustav Adolph Frank Sr. Status: extant.

Accessed 18 December 2012. Tax: mounted. Pub: Abbott

1860, 1861; Gould 1859; Schlegel 1866; Sclater 1860,

1861, 1862.

Pedestal label: [two hands] Anas Cat Nr6 / Ouerqteduta cyan-

optera / ¢Vieillot} / 4. / Abbott coll. / Frank, 1861 / Falkland

eilanden.

Pedestal underside: [one hand] Anas cyanoptera / Cat N°6 /

3 / Iris geelrood, pooten geel / snavel zwart / (Abbot) / van

Capt Abbot / Frank 1861. Oost Falkland.

Schlegel 1866 (Tome VI, 31: 51-52): Anas cyanoptera, Cat 6,

S adulte, iles Falkland, voyage du Capitaine Abbot, acquis

en 1861.

Remarks: The collection site on the label, underside and

in Schlegel 1866 differ. Two generic names are cited (one

crossed out), with the primary binomial on the underside

agreeing with Schlegel 1866. The label, underside, and

Schlegel 1866 all identify the specimen as male, but only

the latter identifies it as an adult. Only one name source is

mentioned, Vieillot on the label. Only Schlegel’s handwriting

appears under the base of the pedestal.

The bird was acquired through the Amsterdam-based dealer

Gustav Adolph Frank and arrived on 30 April 1860 (see Kelp

Goose RMNH.AVES.230433 as for C.C. Abbott).

Cinnamon Teal Spatula cyanoptera cyanoptera (Vieil-

lot, 1816) [RMNH.AVES.231625]. Loc: East Falkland

(ca. 51° 41' 40" S, 57° 51' 10" W). Date: either 1858

or 1859. Coll. C.C. Abbott. Age/sex: 9. Acq: via John

Gould and Gustav Adolph Frank Sr. Status: extant. Ac-

cessed 18 December 2012. Tax: mounted. Pub: Abbott

1860, 1861; Gould 1859; Schlegel 1866; Sclater 1860,

1861, 1862.

Pedestal label: [two hands] Anas Cat N°11. / Ouerqueduta dis-

cors. (L)/ 2 / Abbott coll 1860 / Falkland eilanden.

Pedestal underside: [one hand] Anas discors / Cat N°9/ ° / Iris

zwart, pooten en bek bruinachtig / Cap Abbot / Oost-Falk-

land.

Schlegel 1866 (Tome VI, 31: 50-51): Anas discors. Cat 9, Fe-

melle, iles Falkland, voyage du Capitaine Abbot, acquis en

1860.

Remarks: The collection site on the label, underside and

in Schlegel 1866 differ. Two generic names are cited (one

crossed out), with the primary name on the underside agree-

ing with Schlegel 1866. The label, underside, and Schlegel

1866 all identify the specimen as female, but none identify

the age. No name source is mentioned. Only Schlegel’s hand-

writing appears under the base of the pedestal. The identifi-

cation is incorrect.

©ZFMK

Specimen labelling errors: birds collected on the Falkland Islands prior to 1861 151

The bird was acquired through the Amsterdam-based dealer

Gustav Adolph Frank and arrived on 30 April 1860 (see Kelp

Goose RMNH.AVES.230433 as for C.C. Abbott).

Chiloe Wigeon Mareca_ sibilatrix (Poeppig, 1829)

[RMNH.AVES.230928]. Loc: Falklands (ca. 51° 41' 0"

S, 59° 10' 0" W). Date: in either 1858 or 1859. Col: C.C.

Abbott. Age/sex: adult 4. Acq: via John Gould and Gus-

tav Adolph Frank Sr. Status: extant, accessed 18 Decem-

ber 2012. Tax: mounted. Pub: Abbott 1860, 1861; Gould

1859; Schlegel 1866; Sclater 1860, 1861, 1862.

Pedestal label: [one hand] Cat n°3 / Mareca sibilatrix / (Pop-

pig) / @ / Abbott coll. / Frank, 1860 / Falkland eilanden.

Pedestal underside: [one hand] Anas chiloensis / Cat N°5 / ¢ /

Capt Abbot / (Frank 1860) / Oost Falkland.

Schlegel 1866 (Tome VI, 31: 46): Anas chiloensis, Cat (No nu-

mber), 3 adulte, iles Falkland, voyage du Capitaine Abbot,

acquis en 1860.

Remark: The collection site on the label, underside and in

Schlegel 1866 differ. The label, underside, and Schlegel

1866 all identify the specimen as male, but only the latter

identifies it as an adult. Only one name source is mentioned,

Poeppig on the label. The bird was acquired through the Am-

sterdam-based dealer Gustav Adolph Frank and arrived on

30 April 1860 (see Kelp Goose RMNH.AVES.230433 as for

C.C. Abbott). Only Schlegel’s handwriting appears under the

base of the pedestal.

Yellow-billed Pintail Anas georgica spinicauda Vieillot,

1816 [RMNH.AVES.231462]. Loc: East Falkland (ca.

51° 41' 40" S, 57° 51’ 10" W). Date: in either 1858 or

1859. Col: C.C. Abbott. Age/sex: adult 4. Acq: via John

Gould and Gustav Adolph Frank Sr. Status: extant, ac-

cessed 18 December 2012. Tax: mounted. Pub: Abbott

1860, 1861; Gould 1859; Schlegel 1866; Sclater 1860,

1861, 1862.

Pedestal label: [two hands] Anas Cat n°3 / Dafite-spinicauda /

€Vieillot} / 3 / Abbott coll. / Frank, 1860 / Falkland eilanden.

Pedestal underside: [one hand] Anas spinicauda / Cat N°3 /

S / (unreadable) zwart, snavel geel, bovenkaak aan de / kant

van top met een zwarten band (zeldzaam) / van Capit Abbott

/ Frank 1860 / Oost Falkland.

Schlegel 1866 (Tome VI, 31: 38-39): Anas spinicauda, Cat 3,

3 adulte, iles Malouines, voyage du Captaine Abbot, acquis

en 1860.

Remark: The collection site on the label, underside and in

Schlegel 1866 differ. The label, underside, and Schlegel

1866 all identify the specimen as male, but only the latter

identifies it as an adult. Only one name source is mentioned,

Vieillot on the label. The bird was acquired through the Am-

sterdam-based dealer Gustav Adolph Frank and arrived on

30 April 1860 (see Kelp Goose RMNH.AVES.230433 as for

C.C. Abbott). Only Schlegel’s handwriting appears under the

base of the pedestal.

Yellow-billed Pintail Anas georgica spinicauda Vieillot,

1816 [RMNH.AVES.231463]. Loc: East Falkland (ca.

51° 41' 40" S, 57° 51' 10" W). Date: in either 1858 or

1859. Col: C.C. Abbott. Age/sex: adult 2. Acq: via John

Gould and Gustav Adolph Frank Sr. Status: extant, ac-

Bonn zoological Bulletin 67 (2): 145-169

cessed 18 December 2012. Tax: mounted. Pub: Abbott

1860, 1861; Gould 1859; Schlegel 1866; Sclater 1860,

1861, 1862.

Pedestal label: [two hands] Anas Cat n°4 / Dafita spinicauda /

€Vieillot} / 9 / Abbott coll: / Frank, 1860 / Falkland eilanden.

Pedestal underside: [one hand] Anas spinicauda, Vieillot / Cat

N° / 9 / pooten en bek (unreadable), het mannetje (unread-

able) / Cap. Abbot / Frank 1860 / Oost Falkland.

Schlegel 1866 (Tome VI, 31: 38-39): Anas spinicauda, Cat 4,

@ adulte, iles Malouines, voyage du Captaine Abbot, acquis

en 1860.

Remark: The collection site on the label, underside and in

Schlegel 1866 differ. The label, underside, and Schlegel

1866 all identify the specimen as female, but only the latter

identifies it as an adult. The bird was acquired through the

Amsterdam-based dealer Gustav Adolph Frank and arrived

on 30 April 1860 (see Kelp Goose RMNH.AVES.230433 as

for C.C. Abbott). Only Schlegel’s handwriting appears under

the base of the pedestal.

Yellow-billed Teal Anas flavirostris flavirostris Vieillot

1816 [RMNH.AVES.231104]. Loc: East Falkland (ca.

51° 41' 40" S, 57° 51' 10" W). Date: in either 1858 or

1859. Col: C.C. Abbott. Age/sex: adult 4. Acq: via John

Gould and Gustav Adolph Frank Sr. Status: extant, ac-

cessed 18 December 2012. Tax: mounted. Pub: Abbott

1860, 1861; Gould 1859; Schlegel 1866; Sclater 1860,

1861, 1862.

Pedestal label: [two hands] Anas Cat n°1 / Nettion / Anas flavi-

rostris / €Vieillot} / 3 / Abbott coll: / Frank, 1860 / Falkland

eilanden.

Pedestal underside: [one hand] Anas flavirostris Vieillot /

Cat 3 / Querquedula anaustirostra Philippi & Landbeck /

(1863) (unreadable) / 3 / oogen zwart, snavel geel / zwart in

/ een lijn op de bovenkaak zwart / pooten roodkleurig groen-

achtig / (Abbot) / Capt Abbot / (frank 1860 / Falkland.

Schlegel 1866 (Tome VI, 31: 59): Anas flavirostris, Cat 1, ¢

adultes, iles Falkland, voyage de Mr. Abbot, acquis en 1860.

Remark: The collection site on the label, underside and in

Schlegel 1866 differ. The label, underside, and Schlegel

1866 all identify the specimen as male, but only the latter

identifies it as an adult. Only one name source is mentioned,

Phillippi & Landbeck on the underside. Its catalogue num-

ber is 3, and wrongly labelled at the label and in Schlegel

1866. The bird was acquired through the Amsterdam-based

dealer Gustav Adolph Frank and arrived on 30 April 1860

(see Kelp Goose RMNH.AVES.230433 as for C.C. Abbott).

Only Schlegel’s handwriting appears under the base of the

pedestal.

Yellow-billed Teal Anas flavirostris flavirostris Vieillot

1816 [RMNH.AVES.231105]. Loc: Falklands (ca. 51°

Al' 0" S, 59° 10' 0" W). Date: in either 1858 or 1859.

Col: C.C. Abbott. Age/sex: adult 2. Acq: via John Gould

and Gustav Adolph Frank Sr. Status: extant, accessed 18

December 2012. Tax: mounted. Pub: Abbott 1860, 1861;

Gould 1859; Schlegel 1866; Sclater 1860, 1861, 1862.

Pedestal label: [two hands] Anas Cat n°2 / Nettten / flavirostris

/€Vieillot}/ 9 / Abbott coll: / Frank 1860 / Falkland eilanden.

Pedestal underside: [one hand] Anas flavirostris Vieillot / Cat

4/9 / Capt Abbot / Frank 1860 / Falkland.

©ZFMK

[52 Justin J. F. J. Jansen

Schlegel 1866 (Tome VI, 31: 59): Anas flavirostris, Cat 2, Fe-

melle adultes, iles Falkland, voyage de Mr. Abbot, acquis en

1860.

Remark: The collection site on the label, underside and in

Schlegel 1866 differ. The label, underside, and Schlegel

1866 all identify the specimen as female, but only the latter

identifies it as an adult. At the underside cat 4 is mentioned,

this is incorrect, it’s 2. The bird was acquired through the

Amsterdam-based dealer Gustav Adolph Frank and arrived

on 30 April 1860 (see Kelp Goose RMNH.AVES.230433 as

for C.C. Abbott). Only Schlegel’s handwriting appears under

the base of the pedestal.

Yellow-billed Teal Anas flavirostris flavirostris Vie-

illot 1816 [RMNH.AVES.231106]. Loc: Falkland

(ca. 51° 41' 0" S, 59° 10' 0" W). Date: —. Col: —. Age/

sex: adult. Acq: via Gustav Adolph Frank Sr. Status: ex-

tant, accessed 18 December 2012. Tax: mounted. Pub:

Schlegel 1866.

Pedestal label: [two hands] Anas Cat n°3 / Nettion / flavirostris

/ Vieillot / ad. / Frank 1861 / Falkland eilanden.

Pedestal underside: [one hand] Anas flavirostris / Vieillot / Cat

N°5 / Frank / 1861 / Falkland.

Schlegel 1866 (Tome VI, 31: 59): Anas flavirostris, Cat 3, Indi-

vidu des iles Falkland, 1861.

Remark: The collection site on the label, underside and in

Schlegel 1866 differ. The label only notices the ageing. The

bird was acquired through the Amsterdam-based dealer Gus-

tav Adolph Frank (see Kelp Goose RMNH.AVES.230433).

Only Schlegel’s handwriting appears under the base of the

pedestal.

PODICIPEDIDAE

White-tufted Grebe Rollandia rolland rolland (Quoy &

Gaimard, 1824) [RMNH.AVES.107453]. Loc: Falklands

(ca. 51° 41' 0" S, 59° 10' 0" W). Date: —. Col: —. Age/

sex: adult 4. Acq: via Gustav Adolph Frank Sr. Status:

extant, accessed 18 December 2012. Tax: mounted. Pub:

Schlegel 1867. (Figs 1, 2)

Pedestal label 1: [one hand] Cat n°1 / Podiceps rollandi / Gould

/ & / Falkland eilanden.

Pedestal label 2: [one hand] Podiceps Rollandii / 3. ad: Cat:

1 / Malouines.

Tag: [one hand] Podiceps rollandii 3 / Cat. N°1 / Malouines.

Pedestal underside: [one hand] Podiceps rollandii 3 / Less

Uran pl 36 / Cat N° 1 / Grébe roland / Malouines.

Schlegel 1867 (Tome VI, 33: 42): Podiceps rollandii, Cat 1,

Males adultes, Malouines.

Remark: The collection site on the label, underside and

in Schlegel 1867 differ. One label has adult and the other

has male, the latter applies also for the underside. The bird

was acquired through the Amsterdam-based dealer Gustav

Adolph Frank (see Kelp Goose RMNH.AVES.230433). One

specimen is missing from HMS Beagle (Steinheimer 2004).

Only Schlegel’s handwriting appears under the base of the

pedestal.

White-tufted Grebe Rollandia rolland rolland (Quoy &

Gaimard, 1824) [RMNH.AVES.107454]. Loc: Falklands

Bonn zoological Bulletin 67 (2): 145-169

Fig. 1. White-tufted Grebe Rollandia rolland rolland (RMNH.

AVES.107453), collected Falklands, by C.C. Abbott (photo-

versity Center, Leiden).

(ca. 51° 41' 0" S, 59° 10' 0" W). Date: —. Col: —. Age/

sex: adult 4. Acq: via Gustav Adolph Frank Sr. Status:

extant, accessed 18 December 2012. Tax: mounted. Publ:

Schlegel 1867.

Pedestal label 1: [one hand] Cat n°2 / Podiceps rollandi / Gould

/ / Frank / Falkland eilanden.

Pedestal label 2: [one hand] Podiceps Rollandii / ¢ ad: Cat 2

/ Malouines.

Pedestal underside: [one hand] Podiceps rollandii 3 / Uranie

pl 36 / Cat N°2 / Grébe rolland / Par Frank / Malouines. Het

oude 9 (unreadable / (unreadable) / oude ¢.

Schlegel 1867 (Tome VI, 33: 42): Podiceps rollandii, Cat 2,

Males adultes, Malouines.

Remark: The collection site on the label, underside and in

Schlegel 1867 differ. The label, underside, and Schlegel

1866 all identify the specimen as male, but only the latter

identifies it as an adult, but also at the underside it is identi-

fied as adult 9. The bird was acquired through the Amster-

dam-based dealer Gustav Adolph Frank (see Kelp Goose

RMNH.AVES.230433 as for C.C. Abbott). One specimen is

missing from HMS Beagle (Steinheimer 2004). Only Schle-

gel’s handwriting appears under the base of the pedestal.

White-tufted Grebe Rollandia rolland rolland (Quoy &

Gaimard, 1824) [RMNH.AVES.107455]. Loc: East Falk-

land (ca. 51° 41' 40" S, 57° 51' 10" W). Date: in either

1858 or 1859. Col: C.C. Abbott. Age/sex: adult 2. Acq:

via John Gould and Gustav Adolph Frank Sr. Status: ex-

©ZFMK

Specimen labelling errors: birds collected on the Falkland Islands prior to 1861 153

Fig. 2. White-tufted Grebe Ro/landia rolland rolland (RMNH.AVES.107453), Pedestal underside (photograph by Justin Jansen, 18

tant, accessed 18 December 2012. Tax: mounted. Pub:

Abbott 1860, 1861; Gould 1859, Schlegel 1867; Sclater

1860, 1861, 1862.

Pedestal label 1: [one hand] Cat n°3 / Podiceps rollandi / Gould

/ 9 / Capt Abbot / Stanley / 1859 / O. Falkland eilanden.

Pedestal label 2: [one hand] Podiceps Rollandii / 2 ad: Cat: 3.

ile orientale des / Malouines / M' Abbot Stanley / 1859.

Tag: [one hand] Podiceps rollandii / 9° Cat. N° 3 / Capt. Abbot

Stanley / 1859 / Oost Falkland.

Pedestal underside: [one hand] Podiceps rollandii / 2 / Cat N°

3 / Capt. Abbot Stanley / 1859 / Oost Falkland.

Schlegel 1867 (Tome VI, 33: 42): Podiceps rollandii, Cat 3,

Femelle adulte, absolument semblable au ¢ adulte; ile orien-

tale des Malouines. Voyage de Mr. Abbot Standley, acquise

en 1859.

Remark: The collection site on the label, underside and in

Schlegel 1867 differ. The labels, tag, underside, and Schle-

gel 1867 all identify the specimen as female, but only the

latter and one of the labels identifies it as an adult. The bird

was acquired through the Amsterdam-based dealer Gustav

Adolph Frank (see Kelp Goose RMNH.AVES .230433 as for

C.C. Abbott). Only Schlegel’s handwriting appears under the

base of the pedestal.

Southern Silvery Grebe Podiceps occipitalis occipital-

is Garnot, 1826 [RMNH.AVES.107612]. Loc: Falkland

(ca. 51° 41' 0" S, 59° 10' 0" W). Date: —. Col: —. Age/

sex: @. Acq: via Gustav Adolph Frank Sr. Status: extant,

Bonn zoological Bulletin 67 (2): 145-169

accessed 18 December 2012. Tax: mounted. Pub: Schle-

gel 1867.

Pedestal label 1: [two hands] P. o. occipitalis Garnot. Cat n°1

/ Podiceps calipareus / Less. / & / Frank / Falkland eilanden.

Pedestal label 2: [one hand] Podiceps occipitalis / 4 Cat: 1 /

Malouines.

Tag: [one hand] Podiceps occipitalis. cat 1. Frank. Malouines.

Pedestal underside: [one hand] Podiceps occipitalis / eatipa-

rens- / Less. Coquille pl 4 / Cat. 1 / Grebe aux belles joues

/ p. Frank / Malouines.

Schlegel 1867 (Tome VI, 33: 41): Podiceps occipitalis, Cat 1,

3 au plumage parfait, Malouines.

Remark: The collection site on the label, underside and in

Schlegel 1867 differ. The labels, tag, underside, and Schlegel

1867 all identify the specimen as male, but only the latter

identifies it as an adult. The bird was acquired through the

Amsterdam-based dealer Gustav Adolph Frank (see Kelp

Goose RMNH.AVES.230433). One specimen is missing

from HMS Beagle (Steinheimer 2004). Only Schlegel’s

handwriting appears under the base of the pedestal.

Southern Silvery Grebe Podiceps occipitalis occipital-

is Garnot, 1826 [RMNH.AVES.107609]. Loc: Falklands

(ca. 51° 41' 0" S, 59° 10' 0" W). Date: November—De-

cember 1822. Col: voyage La Coquille, R.P. Lesson /

P. Garnot. Age/sex: adult 9. Acq: April 1825 received

in exchange with the Paris Museum. Status: extant, ac-

©ZFMK

154 Justin J. F. J. Jansen

cessed 18 December 2012. Tax: mounted. Pub: Lesson &

Garnot 1826—1830; Schlegel 1867.

Pedestal label 1: [two hands] P.o. occipitalis Garnot Cat: n°2 /

Podiceps calipareus / Less. / 2° April 1825 Falkland eilanden.

Tag: [one hand] P occipitalis 9 / Cat No2 / April 1825 / Mal-

ouines.

Pedestal underside: [two hands] Podiceps occipitalis | Po=

; / 2 / Less. / Grebe aux belles Joues / Les

Coquille pl 45 / Cat 2 / Avril 1825 Avril 1825. / Malouines.

Schlegel 1867 (Tome VI, 33: 41): Podiceps occipitalis, Cat

2, Femelle au plumage parfait, absolument semblable aux

males en parure, tuee en Avril 1825, Malouines.

Remark: The collection site on the label, underside and in

Schlegel 1867 differ. The label, tag, underside, and Schlegel

1867 all identify the specimen as female, but only the latter

identifies it as an adult. Its origin could be traced from one

of the acquisition books in MNHN, that clearly shows the

exchange of this bird to Temminck in April 1825.

Southern Silvery Grebe Podiceps occipitalis occipitalis

Garnot, 1826 [RMNH.AVES.107610]. Loc: East Falk-

land (ca. 51° 41' 40" S, 57° 51' 10" W). Date: in 1858.

Col: C.C. Abbott. Age/sex: adult 4. Acq: via John Gould

and Gustav Adolph Frank Sr. Status: extant, accessed 18

December 2012. Tax: mounted. Pub: Abbott 1860, 1861;

Gould 1859, Schlegel 1867; Sclater 1860, 1861, 1862.

Pedestal label 1: [two hands] P. o. occipitalis Garnot Cat n°3 /

Podiceps calipareus / Less. / ¢ / Kapt: Abbot / Stanley 1859

/ Oost-Falkland.

Pedestal label 2: [one hand] Podiceps occipitalis / 1859 &.

Cat: 3. Iie orient des, // Voy. Abbot Stanly / Malouines.

Tag: [one hand] Podiceps occipitalis / 3 Cat 3 / Capt Abbot

Stanley / 1859 / Oost-Falkland.

Pedestal underside: [two hands] Podiceps occipitalis / 3 / Cat

3 / Capt. Abbot Stanley / 1859 / Oost-Falkland.

Schlegel 1867 (Tome VI, 33: 41): Podiceps occipitalis, Cat 3,

S au plumage parfait, ile orientale des Malouines, voyage de

Mr. Abbot Stanley, acquis en 1859.

Remark: The collection site on the label, underside and in

Schlegel 1867 differ. The labels, tag, underside, and Schlegel

1867 all identify the specimen as male, but only the latter

identifies it as an adult. The bird was acquired through the

Amsterdam-based dealer Gustav Adolph Frank (see Kelp

Goose RMNH.AVES.230433 as for C.C. Abbott).

SPHENISCIDAE

King Penguin Aptenodytes patagonicus J.F. Miller, 1778

[RMNH.AVES.107287]. Loc: —. Date: —. Col: —. Age/

sex: adult. Acq: from the Paris museum. Status: extant,

accessed 18 December 2012. Tax: mounted. Pub: Schle-

gel 1867.

Pedestal label: [one hand] Cat n°1 / Aptenodytes patachonica /

Forster / 1835 / Falkland eilanden.

Pedestal underside: [one hand] Aptenodytes patachonica / Cat

Nol / Paris 1835 / Malouines.

Schlegel 1867 (Tome VI, 33: 5-6): Adulte, Malouines, voyage

de /’Astrolabe et de la Zelee.

Remark: The collection site on the label, underside and in

Schlegel 1867 differ. The ageing is only present in Schle-

gel 1867. Either the date 1835 is incorrect or the expedition

Bonn zoological Bulletin 67 (2): 145-169

where it is collected. Only Schlegel’s handwriting appears

under the base of the pedestal.

If voyage de /’Astrolabe et de la Zelee is correct: Possibly

collected at Ciudad del Rey Don Felipe or elsewhere in the

straits of Patagonia, Chile (not the Falklands). Col: collect-

ed at the expeditions with the Astrolabe and Zélée, Jacques

Bernard Hombron or Honore Jacquinot. Date: December

1836—January 1837. Sex/age: Adult. Acquisition: from the

Paris museum in 1842.

If 1835 is correct: Loc: Falklands (51°45’S 59°00’W). Col:

voyage La Coquille, P. Lesson / P. Garnot. Date: November—

December 1822. Sex/age: adult. Acquisition: June 1835 re-

ceived in exchange with the Paris Museum.

King Penguin Aptenodytes patagonicus J.F. Mill-

er, 1778 [RMNH.AVES.107286]. Loc: Falklands

(ca. 51° 41' 0" S, 59° 10' 0" W). Date: November—De-

cember 1822. Col: voyage La Coquille, P. Lesson / P.

Garnot. Age/sex: adult. Acq: June 1835 received in ex-

change with the Paris Museum. Status: extant, accessed

18 December 2012. Tax: mounted. Pub: Lesson & Gar-

not 1826-1830; Schlegel 1867.

Pedestal label: [one hand] Cat n°.2 / Aptenodytes patachonica

/ Forster / Falkland eilanden.

Pedestal underside: [one hand] Spenicurus Pennantii / Cat n°

2 / Iles Malouines.

Schlegel 1867 (Tome VI, 33: 3): Spenicurus pennantii, Cat 2,

Adulte, Iles Malouines, acquis en 1835.

Remark: The collection site on the label, underside and in

Schlegel 1867 differ. The ageing is only present in Schlegel

1867. Notable is the lack of the note appearing in Schlegel

1867 that it was purchased in 1835, this is missing from both

label as underside. Only Schlegel’s handwriting appears un-

der the base of the pedestal.

King Penguin Aptenodytes patagonicus J.F. Miller, 1778

[—]. Loc: Falklands (ca. 51° 41'0" S, 59° 10' 0" W). Date:

—. Col: —. Age/sex: adult. Acq: —. Status: non-extant, not

found. Tax: —. Pub: Schlegel 1867.

Schlegel 1867 (Tome VI, 33: 3): Spenicurus pennantii, Cat 2,

Adulte, Malouines.

Remark: Little information is available.

Gentoo Penguin Pygoscelis papua papua (J.R. For-

ster, 1781) [RMNH.AVES.107252]. Loc: Falkland (ca.

51° 41' 0" S, 59° 10’ 0" W). Date: —. Col: —. Age/sex:

adult. Acq: via Gustav Adolph Frank Sr. Status: extant,

accessed 18 December 2012. Tax: mounted. Pub: Schle-

gel 1867.

Pedestal label: [one hand] Cat n°l / Pygoscelis papua / (For-

ster) / Falkland eilanden.

Pedestal underside: [one hand] Spheniscus papua / Apteno-

dytes papuaensis / Cat 1 / Manchot (unreadable) Sonner. Pl

115 / (unreadable) parfaite / Par Frank / Iles Falkland / Ma-

louines.

Schlegel 1867 (Tome VI, 33: 5): Sphenicurus papua, Cat 1,

Adulte, Malouines.

Remark: The collection site on the label, underside and in

Schlegel 1867 differ. The ageing is only present in Schlegel

©ZFMK

Specimen labelling errors: birds collected on the Falkland Islands prior to 1861 155

1867. The bird was acquired through the Amsterdam-based

dealer Gustav Adolph Frank (see Kelp Goose RMNH.

AVES.230433). Only Schlegel’s handwriting appears under

the base of the pedestal.

Gentoo Penguin Pygoscelis papua papua (J.R. Forster,

1781) [RMNH.AVES.107253]. Loc: Possibly collected

at Ciudad del Rey Don Felipe or elsewhere in the straits

of Patagonia, Chile (not the Falklands). Date: December

1836—January 1837. Col: collected at the expeditions

with the Astrolabe and Zé/ée, Jacques Bernard Hombron

or Honore Jacquinot. Age/sex: adult 9. Acq: from the

Paris museum in 1842. Status: extant, accessed 18 De-

cember 2012. Tax: mounted. Pub: Blanchard et al. 1854;

Schlegel 1867.

Pedestal label: [one hand] Cat n°2 / Pygoscelis papua / (For-

ster) / 9 / Falkland eilanden.

Pedestal underside: [two hands] Spheniscus papua / Apteno-

dytes papuaensis 2 / 9 / Sonnerat / Cat 2 / Voyage Astrolabe

/ Voy Astrolabe a Zelee / Malouines / Malouines.

Schlegel 1867 (Tome VI, 33: 5): Sphenicurus papua, Cat 2,

Femelle adulte, Malouines, voyage de /’Astrolabe et de la

Zelee.

Remark: The collection site on the label, underside and in

Schlegel 1867 differ. The sexing of the specimen is noted in

all sources, the ageing however is only present in Schlegel

1867. The label, underside, and Schlegel 1867 all identify

the specimen as female, but only the latter identifies it as an

adult. Two name sources are mentioned, Forster on the label

and Sonnerat on the underside. Arrived in 1842 from the Par-

is Museum (part on exchange between Temminck and Paris).

Chinstrap Penguin Pygoscelis antarcticus (J.R. Forster,

1781) [RMNH.AVES.107250]. Loc: Possibly collected

at Ciudad del Rey Don Felipe or elsewhere in the straits

of Patagonia, Chile (not the Falklands). Date: December

1836—January 1837. Col: collected at the expeditions

with the Astrolabe and Zélée, Jacques Bernard Hombron

or Honore Jacquinot. Age/sex: adult. Acq: from the Paris

museum in 1842. Status: extant, accessed 18 December

2012. Tax: mounted. Publ: Blanchard et al. 1854; Schle-

gel 1867.

Pedestal label: [one hand] Cat n°l / Pygoscelis antarctica /

(Forster) / Falkland eilanden.

Pedestal underside: [one hand] Sphenicurus antarctica | MH-

er-pt4é / (unreadable crossed) / Cat 1 / (unreadable crossed) /

M Paris Astrolabe et de Zelee / (unreadable) / Regiones Ant-

arctiques / Malouines.

Schlegel 1867 (Tome VI, 33: 5—6): Adulte, Malouines, voyage

de /’Astrolabe et de la Zelee.

Remark: The collection site on the label, underside and in

Schlegel 1867 differ. Only in Schlegel 1867 the specimen is

aged. Arrived in 1842 from the Paris Museum (part on ex-

change between Temminck and Paris Museum). Only Schle-

gel’s handwriting appears under the base of the pedestal.

Macaroni Penguin Eudyptes chrysolophus J.F. von

Brandt, 1837 [RMNH.AVES.107247]. Loc: Falkland

(ca. 51° 41' 0" S, 59° 10' 0" W). Date: —. Col: —. Age/

Bonn zoological Bulletin 67 (2): 145-169

sex: adult. Acq: via Gustav Adolph Frank Sr. Status: ex-

tant, accessed 18 December 2012. Tax: mounted. Pub:

Schlegel 1867.

Pedestal label: [two hands] Eudyptes chrysolophus (Brandt)

Cat. n°1 / Catarrhactes chrysolophus / Brandt / 3 / Frank /

Falkland Eilanden.

Pedestal underside: [three hands] Spheniscus diamematus /

chrysotopins / 5 adult / Gould Proc an 1837 p 3 / Apteno-

dytes cirhata Miller pl (unreadable) / Cat 1 / pr Frank / (line

unreadable) / Bec rouge (two unreadable words) / Iles Falk-

land.

Schlegel 1867 (Tome VI, 33: 8): Adulte, Malouines.

Remark: The collection site on the label, underside and in

Schlegel 1867 differ. The label, underside, and Schlegel 1867

all identify the specimen as male, but only the latter identi-

fies it as an adult. The bird was acquired through the Am-

sterdam-based dealer Gustav Adolph Frank (see Kelp Goose

RMNH.AVES.230433).

Fjordland Penguin Eudyptes pachyrhynchus G.R.

Gray, 1845 [RMNH.AVES.107258]. Loc: Falkland

(ca. 51° 41' 0" S, 59° 10' 0" W). Date: —. Col: —. Age/

sex: adult. Acq: via Gustav Adolph Frank Sr. Status: ex-

tant, accessed 18 December 2012. Tax: mounted. Pub:

Schlegel 1867.

Pedestal label: [two hands] Eudyptes pachyrhynchus Cat. n°2

/ GR. Gray / Catarrhactes pachyrhynchus | (G.R. Gray) /

Falkland eilanden.

Pedestal underside: [one hand] (unreadable) Gould / Spheni-

curus chrysocome / wnreadabte / Cat 2 / Gould Proc 1857 p

310. / (unreadable)/ ile Falkland.

Pedestal underside sticker: Un trés vieil et bel individu de la

Nouv. Zél., et appartenant a M. Frank est d’une taille un peu

plus forte, le bec est plus grand et les teintes foncées sont

d’un noir plus intense, Schlegel.

Schlegel 1867 (Tome VI, 33: 6-7): Adulte, Malouines.

Remark: The collection site on the label, underside and in

Schlegel 1867 differ. Ageing only present in Schlegel 1865,

note the different Latin names. The bird was acquired through

the Amsterdam-based dealer Gustav Adolph Frank (see Kelp

Goose RMNH.AVES.230433).

Fjordland Penguin Eudyptes pachyrhynchus G.R. Gray,

1845 [RMNH.AVES.107249]. Loc: Beauchene Island,

Falkland (ca. 52° 53' 11" S, 59° 12' 13" W). Date: —. Col:

—. Age/sex: adult. Acq: via Gustav Adolph Frank Sr. Sta-

tus: extant, accessed 18 December 2012. Tax: mounted.

Pub: Schlegel 1867.

Pedestal label: [two hands] Eudyptes pachyrhynchus Cat n°.3

/ (GR. Gray) / Catarrhactes pachyrhynchus / (G.R. Gray) /

Beauchene / Frank, 1861 / Falkland eilanden.

Pedestal underside: [three hands] Spheniscus chrysolophus /

(unreadable) in Gould / Cat n°2 / ile Beauchesne / Frank 1861

/ Malouines.

Pedestal underside sticker: Bought from Beauchene / island

by the sealers / Frank 1861 [two hands].

Schlegel 1867 (Tome VI, 33: 7): Spheniscus chrysolophus, Cat

2, Albinos, rapporte de l’ile Beachene, Malouines, 1861.

Remark: The collection site on the label, underside and in

Schlegel 1867 differ. The origin is only documented at the

label (Frank). The bird was acquired through the Amster-

©ZFMK

156 Justin J. F. J. Jansen

dam-based dealer Gustav Adolph Frank (see Kelp Goose

RMNH.AVES.230433).

Magellanic Penguin Spheniscus magellanicus J.R. For-

ster, 1781 [RMNH.AVES.107261]. Loc: Falkland (ca.

51° 41' 0" S, 59° 10’ 0" W). Date: —. Col: —. Age/sex:

adult. Acq: via Gustav Adolph Frank Sr. Status: extant,

accessed 18 December 2012. Tax: mounted. Pub: Schle-

gel 1867.

Pedestal label: [one hand] Cat n°2 / Spheniscus demrerstts /

magellanicus / (Forster) / Frank / Falkland eilanden.

Pedestal underside: [one hand] Spheniscus demersus / Cat N

4 / Nov spec! = Apt. demersa var C. Vieillot / Finsch / ex

Pernetti Voy aux iles Malouines II p 17 / Pernetti, P. / Frank

/ Iles Malouines.

Schlegel 1867 (Tome VI, 33: 10-12): Spheniscus demersus,

Cat 4, Individu au passage a la livrée parfaite, Malouines.

Remark: The collection site on the label, underside and in

Schlegel 1867 differ. Frank is missing from Schlegel. Finsch

thought it was a new species for science. The bird was ac-

quired through the Amsterdam-based dealer Gustav Adolph

Frank (see Kelp Goose RMNH.AVES.230433). Only Schle-

gel’s handwriting appears under the base of the pedestal.

Magellanic Penguin Spheniscus magellanicus J.R. For-

ster, 1781 [RMNH.AVES.107262]. Loc: Falkland (ca.

51° 41' 0" S, 59° 10’ 0" W). Date: —. Col: —. Age/sex:

Adult. Acq: via Gustav Adolph Frank Sr. Status: extant,

accessed 18 December 2012. Tax: mounted. Pub: Schle-

gel 1867.

Pedestal label: [one hand] Cat n°3 / Spheniscus demrerstts /

magellanicus / (Forster) / Frank / Falkland eilanden.

Pedestal underside: [one hand] sp nov. (unreadable) Finsch /

Black-footed Penguin (unreadable) / Spheniscus demersus /

Cat N°.5 / Via Frank / Malouines.

Schlegel 1867 (Tome VI, 33: 10-12): Spheniscus demersus,

Cat 5, Individu au plumage imparfait, mais offerant la taille

des adultes, Malouines.

Remark: The collection site on the label, underside and in

Schlegel 1867 differ. Notable if difference in Latin names

at the label and underside, the latter is in line with Schlegel

1867. The bird was acquired through the Amsterdam-based

dealer Gustav Adolph Frank (see Kelp Goose RMNH.

AVES.230433).

PROCELLARIIDAE

Slender-billed Prion Pachyptila belcheri Mathews, 1912

[RMNH.AVES.107095]. Loc: Falkland (ca. 51° 41' 0" S,

59° 10' 0" W). Date: February 1860. Col: —. Age/sex: -.

Acq: via John Gould and Gustav Adolph Frank Sr. Sta-

tus: extant, accessed 18 December 2012. Tax: mounted.

Pub: Schlegel 1863b.

Pedestal label: [one hand] Cat.n°2 / Prion ariel / Gould / Feb-

ruari 1860 / Frank 1863 / Falkland eilanden.

Pedestal underside: [one hand] Procellaria ariel Gould / Fr.

turtur Kuhl, Dser, p 143, pl&8 / Cat N°2 / February 1860 /

Falkland Island / Frank / 1863.

Schlegel 1863b (Tome VI, 22: 18): Individu des Mers de Il’ Aus-

tralie, obtenus en 1863 de Mr Gould.

Bonn zoological Bulletin 67 (2): 145-169

Remark: The collection site on the label, underside and in

Schlegel 1867 differ, and two birds are involved (Schle-

gel 1863b: 22). Notable is the lack of catalogue number in

Schlegel 1863b. The bird was acquired through the Amster-

dam-based dealer Gustav Adolph Frank (see Kelp Goose

RMNH.AVES.230433). A specimen is missing from HMS

Beagle (Steinheimer 2004). Only Schlegel’s handwriting ap-

pears under the base of the pedestal.

PHALACROCORACIDAE

Rock Shag Phalacrocorax magellanicus J.F. Gmelin,

1789 [RMNH.AVES.107973]. Loc: Magdalena Island

(ca. 51° 41' 0" S, 59° 10' 0" W). Date: —. Col: —. Age/

sex: adult 4. Acq: via Gustav Adolph Frank Sr. Status:

extant, accessed 18 December 2012. Tax: mounted. Pub:

Schlegel 1863a.

Pedestal label: [two hands] Phalacrocorax magellanicus /

Graculus magellanicus / m. Cat N° 1 / Malouines.

Pedestal underside: [one hand] Graculus / Carbo magellan-

icus 3 / Forster voy / Cat No 1 / Frank / Ile Magdalena /

Malouines.

Schlegel 1863a (Tome VI, 21: 21): Graculus magellanicus, Cat

1, ¢ adulte, Malouines.

Remark: The collection site on the label, underside and in

Schlegel 1863a differ. Ageing only appears in Schlegel 1853a

as aged and sexed. Due to the remark of Magdalena Island

(52°55’S 70°35’W) on the pedestal underside, the known ex-

pedition that visited this place was the voyage de / Astrolabe

et de /a Zelee and could be the source. The bird was acquired

through the Amsterdam-based dealer Gustav Adolph Frank

(see Kelp Goose RMNH.AVES.230433). Only Schlegel’s

handwriting appears under the base of the pedestal.

Rock Shag Phalacrocorax magellanicus J.F. Gmelin,

1789 [RMNH.AVES.107972]. Loc: Magdalena Island

(ca. 51° 41' 0" S, 59° 10' 0" W). Date: —. Col: —. Age/

sex: adult 2. Acq: via Gustav Adolph Frank Sr. Status:

extant, accessed 18 December 2012. Tax: mounted. Pub:

Schlegel 1863a.

Pedestal label: [two hands] Phalacrocorax magellanicus /

Graculus magellanicus / f. Cat N° 2 / Malouines.

Tag: [one hand] Graculus magellanicus / Cat: No 2 9 / Frank.

Malouines.

Pedestal underside: [one hand] Graculus / Carbo magellani-

cus / Forster voy / Cat N° 2 / Frank / Malouines.

Note Pedestal label: [one hand] Ile Magdalena / (unreadable) /

le brun / (unreadable) orange / (unreadable).

Schlegel 1863a (Tome VI, 21: 21): Graculus magellanicus, Cat

2, Femelle adulte, Malouines.

Remark: The collection site on the label, underside and in

Schlegel 1863a differ. The tag, and Schlegel 1863a all iden-

tify the specimen as female, but only the latter identifies it as

an adult. Due to the remark of Magdalena Island (52°55’S

70°35’W) on the pedestal underside, the known expedition

that visited this place was the voyage de /’Astrolabe et de la

Zelee and could be the source. The bird was acquired through

the Amsterdam-based dealer Gustav Adolph Frank (see Kelp

Goose RMNH.AVES.230433). Only Schlegel’s handwriting

appears under the base of the pedestal.

©ZFMK

Specimen labelling errors: birds collected on the Falkland Islands prior to 1861 157

Rock Shag Phalacrocorax magellanicus J.F. Gmelin,

1789 [RMNH.AVES.107971]. Loc: East Falkland (ca.

51° 41' 40" S, 57° 51' 10" W). Date: in 1860. Col: C.C.

Abbott. Age/sex: 3. Acq: via John Gould and Gustav

Adolph Frank Sr. Status: extant, accessed 18 December

2012. Tax: mounted. Pub: Abbott 1860, 1861; Gould

1859; Schlegel 1863a; Sclater 1860, 1861, 1862.

Pedestal label: [two hands] Phalacrocorax magellanicus /

Graculus magellanicus / m. Cat N°3 / voy. Cap. Abbot / 1861

/ Malouines.

Tag: [one hand] Graculus magellanicus / 3 Cat.3 / Abbot /

Frank 1861 / Oost-Falkland.

Pedestal underside: [one hand] Graculus magellanicus | & /

Cat N°3 / Abbot / Frank 1861 / Oost-Falkland.

Schlegel 1863a (Tome VI, 21: 21): Graculus magellanicus,

Cat 3, Male, habit de passage, ile orientale des Malouines,

voyage du captaine Abbot, acquis en 1861.

Remark: The name giving of the location of collecting is dif-

fers. The label, underside, and Schlegel 1866 all identify the

Specimen as male, but is not present at the label. The bird

was acquired through the Amsterdam-based dealer Gustav

Adolph Frank and arrived on 21 April 1861 (see Kelp Goose

RMNH.AVES.230433 as for C.C. Abbott). Only Schlegel’s

handwriting appears under the base of the pedestal.

CHIONIDAE

Snowy Sheathbill Chionis albus J.F. Gmelin, 1789

[RMNH.AVES.223734]. Loc: —. Date: —. Col: —. Age/

sex: —. Acq: —. Status: extant, accessed 11 February 2013.

Tax: skin. Pub: -.

Tag: [one hand] Kat.Nol. Ad / Chionis alba (Gml) — Cat. Br.

M. XXIV. p 710 / Tem.Pl.col.509 (not Type) / “Falkland Is-

lands” / No origin! / “Nouv.Zelande ‘Temmik” / “Chatham

Isl. Schlegel.

Tag (Backside): [one hand] Chionis vaginalis, Forst. -vaginalis

/ alba, Sull, — bec en foureau blanc / Tem. Pl. Col 509. Nouv

Zelande “old / label written by Temminck”.

Pedestal underside: removed.

Remark: Due to two different locations mentioned on the tag,

is uncertain if the specimen was collected at either “New

Zealand”, “Chatham Islands” or the “Falklands”. The bird

was skinned after first been mounted, the information on the

pedestal underside and the label is not contained. Possibly

this was transcribed on the tag by Otto Finsch.

HAEMATOPODIDAE

Magellanic Oystercatcher Haematopus leucopodus

Garnot, 1826 [RMNH.AVES.224449]. Loc: Falkland

(ca. 51° 41'0" S, 59° 10' 0" W). Date: —. Col: —. Age/sex:

adult. Acq: via Gustav Adolph Frank Sr. Status: extant,

accessed 18 December 2012. Tax: mounted. Pub: Abbott

1860, 1861; Gould 1859; Schlegel 1865; Sclater 1860,

1861, 1862.

Pedestal label: [one hand] Haematopus luctuosus / ad. Cat: 1

/iles Malouines.

Bonn zoological Bulletin 67 (2): 145-169

Pedestal underside: [one hand] Haematopus (unreadable and

crossed) / (3 words unreadable and crossed) / /uctuosus Cuv

/ Frank / Cat. N°1 / Malouines.

Schlegel 1865 (Tome IV, 29:74): Haematopus luctuosus, Cat 1,

Adulte, iles Malouines.

Remark: The collection site on the label, underside and in

Schlegel 1865 differ. Both the underside and Schlegel 1866

identify the specimen as female, but only the latter identi-

fies it as an adult. The bird was acquired through the Am-

sterdam-based dealer Gustav Adolph Frank (see Kelp Goose

RMNH.AVES.230433), note the source is only present at the

underside. Only Schlegel’s handwriting appears under the

base of the pedestal.

Magellanic Oystercatcher Haematopus leucopodus

Garnot, 1826 [RMNH.AVES.224449]. Loc: East Falk-

land (ca. 51° 41' 40" S, 57° 51' 10” W). Date: in either

1858 or 1859. Col: C.C. Abbott. Age/sex: adult. Acq: via

John Gould and Gustav Adolph Frank Sr. Status: extant,

accessed 18 December 2012. Tax: mounted. Publ: Abbott

1860, 1861; Gould 1859; Schlegel 1865; Sclater 1860,

1861, 1862.

Pedestal label: [one hand] Haematopus luctuosus / ad. Cat: 2.

Ile Orient des / Malouines / Mr Abbot Stanley / 1860.

Pedestal underside: [one hand] Haematopus luctuosus / Cat.

N°2 / Capt / Abbot Stanley / 1860 / Oost-Falkland.

Schlegel 1865 (Tome IV, 29:74): Haematopus luctuosus, Cat

2, Adulte, ile orientale des Malouines, voyage du Capitaine

Abbot Stanley, acquis en 1860.

Remark: The collection site on the label, underside and in

Schlegel 1865 differ. The name Abbott was transcribed in-

correctly at label, underside and in Schlegel. Both the label

and Schlegel 1865 identify the specimen as adult, it lacks

at the underside. The bird was acquired through the Amster-

dam-based dealer Gustav Adolph Frank and arrived on 30

April 1860 (see Kelp Goose RMNH.AVES.230433 as for

C.C. Abbott). Only Schlegel’s handwriting appears under the

base of the pedestal.

CHARADRIIDAE

Two-banded Plover Charadrius _falklandicus

Latham, 1790 [RMNH.AVES.223949]. Loc: Falkland

(ca. 51° 41' 0" S, 59° 10' 0" W). Date: —. Col: —. Age/

sex: adult. Acq: via Gustav Adolph Frank Sr. Status: ex-

tant, accessed 18 December 2012. Tax: mounted. Pub:

Schlegel 1865.

Pedestal label: [one hand] Charadrius falklandicus / Ad Cat 1

/ Malouines.

Pedestal underside: [one hand] Charadrius falklandicus / Lath

/ (unreadable words) / pr Frank Cat N1 / Malouines.

Schlegel 1865 (Tome IV, 29: 36): Charadrius falklandicus, Cat

1, Adulte, Malouines.

Remark: The ageing however is absent on the underside.

Handwriting from Temminck indicates an old specimen,

possible secured due to one of the exchanges with the Pa-

risian Museum. The bird was acquired through the Amster-

dam-based dealer Gustav Adolph Frank (see Kelp Goose

RMNH.AVES.230433), note the source is only present at the

©ZFMK

158 Justin J. F. J. Jansen

underside. Only Schlegel’s handwriting appears under the

base of the pedestal.

Two-banded Plover Charadrius falklandicus Latham,

1790 [RMNH.AVES.223954]. Loc: East Falkland (ca.

51° 41' 40" S, 57° 51' 10" W). Date: in either 1858 or

1859. Col: C.C. Abbott. Age/sex: adult 3. Acq: via John

Gould and Gustav Adolph Frank Sr. Status: extant, ac-

cessed 18 December 2012. Tax: mounted. Pub: Abbott

1860, 1861; Gould 1859; Schlegel 1865; Sclater 1860,

1861, 1862.

Pedestal label: [one hand] Charadrius falklandicus / 3 ad. Cat:

2 / Abbot Stanley / 1860 / Ile orient des / Malouines.

Pedestal underside: [one hand] Charadrius / falklandicus / 3 /

Cat N°2 / Capt / Abbot Stanley / 1860 / Oost Falkland.

Schlegel 1865 (Tome IV, 29: 36): Charadrius falklandicus, Cat

2, & adult, ile orientale des Malouines, voyage du Capitaine

Abbot Stanley, acquis en 1860.

Remark: The collection site on the label, underside and in

Schlegel 1865 differ. The name Abbott was transcribed in-

correctly at label, underside and in Schlegel. The label, un-

derside, and Schlegel 1866 all identify the specimen as male,

but only label and Schlegel identifies it as an adult <. The

bird was acquired through the Amsterdam-based dealer Gus-

tav Adolph Frank (see Kelp Goose RMNH.AVES.230433 as

for C.C. Abbott). Only Schlegel’s handwriting appears under

the base of the pedestal.

Two-banded Plover Charadrius falklandicus Latham,

1790 [RMNH.AVES.223955]. Loc: East Falkland (ca.

51° 41’ 40" S, 57° 51’ 10" W). Date: in either 1858 or

1859. Col: C.C. Abbott. Age/sex: adult 2. Acq: via John

Gould and Gustav Adolph Frank Sr. Status: extant, ac-

cessed 18 December 2012. Tax: mounted. Publ: Abbott

1860, 1861; Gould 1859; Schlegel 1865; Sclater 1860,

1861, 1862.

Pedestal label: [one hand] Charadrius falklandicus / 1860. 9

ad Cat: 3 / Ile orient des / Abbot Stanley / Malouines.

Pedestal underside: [one hand] Charadrius / falklandicus / 9

/ Cat N° 3 / eye black / Capt. Abbot / Stanley / 1860 / Oost

Falkland.

Schlegel 1865 (Tome IV, 29: 36): Charadrius falklandicus, Cat

3, 9 adult, ile orientale des Malouines, voyage du Capitaine

Abbot Stanley, acquis en 1860.

Remark: The collection site on the label, underside and in

Schlegel 1865 differ. The name Abbott was transcribed incor-

rectly at label, underside and in Schlegel. The label, under-

side, and Schlegel 1866 all identify the specimen as female,

but only the label and Schlegel identifies it as an adult 9°. The

bird was acquired through the Amsterdam-based dealer Gus-

tav Adolph Frank see Kelp Goose RMNH.AVES.230433 as

for C.C. Abbott). Only Schlegel’s handwriting appears under

the base of the pedestal.

Rufous-chested Plover Charadrius modestus M.H.C.

Lichteinstein, 1823 [RMNH.AVES.226062]. Loc: East

Falkland (ca. 51° 41' 40" S, 57° 51' 10" W). Date: in

either 1858 or 1859. Col: C.C. Abbott. Age/sex: adult 3.

Acq: via John Gould and Gustav Adolph Frank Sr. Sta-

tus: extant, accessed 18 December 2012. Tax: mounted.

Bonn zoological Bulletin 67 (2): 145-169

Publ: Abbott 1860, 1861; Gould 1859; Schlegel 1865:

Sclater 1860, 1861, 1862.

Pedestal label: [one hand] Morinellus modestus / 3 Cat: 1 / Ile

Orients des / M' Abbot Stanley / 1860 / Malouines.

Pedestal underside: [one hand] Morinellus modestus / 3 / Cat

N°1 / Capt. / Abbot Stanly / 1860 / Oost Falkland.

Schlegel 1865 (Tome IV, 29: 48): Morinellus modestus, Cat 1,

3S au plumage parfait, ile orientale des Malouines, voyage de

Mr. Abbot Stanley, acquis en 1860.

Remark: The collection site on the label, underside and in

Schlegel 1865 differ. The name Abbott was transcribed in-

correctly at label, underside and in Schlegel. The label, un-

derside, and Schlegel 1865 all identify the specimen as male,

but only the latter identifies tt as an adult. The bird was ac-

quired through the Amsterdam-based dealer Gustav Adolph

Frank (see Kelp Goose RMNH.AVES.230433 as for C.C.

Abbott). Only Schlegel’s handwriting appears under the base

of the pedestal.

Rufous-chested Plover Charadrius modestus M.H.C.

Lichteinstein, 1823 [RMNH.AVES.226063]. Loc: East

Falkland (ca. 51° 41' 40" S, 57° 51' 10" W). Date: in

either 1858 or 1859. Col: C.C. Abbott. Age/sex: adult &.

Acq: via John Gould and Gustav Adolph Frank Sr. Sta-

tus: extant, accessed 18 December 2012. Tax: mounted.

Publ: Abbott 1860, 1861; Gould 1859; Schlegel 1865:

Sclater 1860, 1861, 1862.

Pedestal label: [one hand] Morinellus modestus / 2 Cat: 2 / ile

Orients des / Mr. Abbot Stanley / 1860 / Malouines.

Pedestal underside: [one hand] Morinellus modestus / 2. Vat.

N° 2/eye black / Capt. / Abbot Stanly / 1860 / Oost falkland.

Schlegel 1865 (Tome IV, 29: 48): Morinellus modestus, Cat 2,

° au plumage parfait, ile orientale des Malouines, voyage de

Mr. Abbot Stanley, acquis en 1860.

Remark: The collection site on the label, underside and in

Schlegel 1865 differ. The name Abbott was transcribed in-

correctly at label, underside and in Schlegel. The label, un-

derside, and Schlegel 1865 all identify the specimen as fe-

male, but only the latter identifies it as an adult. The bird

was acquired through the Amsterdam-based dealer Gustav

Adolph Frank (see Kelp Goose RMNH.AVES.230433 as for

C.C. Abbott). Only Schlegel’s handwriting appears under the

base of the pedestal.

Southern Lapwing Vanellus chilensis chilensis (Molina,

1782) [RMNH.AVES.31529]. Loc: East Falkland (ca.

51° 41' 40" S, 57° 51' 10" W). Date: in either 1858 or

1859. Col: C.C. Abbott. Age/sex: —. Acq: via John Gould

and Gustav Adolph Frank Sr. Status: extant, accessed

18 December 2012. Tax: mounted. Publ: Abbott 1860,

1861; Gould 1859; Schlegel 1865; Sclater 1860, 1861,

1862.

Pedestal label: [two hands] occidentalis Cat 7 / Vanellus eay=

ennensis / ad. cat. 9 / Ile orientale des / Mr. Abbot Stanley /

1860 / Malouines.

Pedestal underside: [one hand] Vanellus cayennensis / Capt. /

Cat. N°9 / Abbot Stanley / 1860 / Oostfalkland.

Schlegel 1865 (Tome IV, 29: 57-58): Vanellus cayennensis, Cat

9, Adulte a tarses seulement longs de 2 pouces, ile orientale

des Malouines, voyage de Mr Abbot Stanley, acquis en 1860.

©ZFMK

Specimen labelling errors: birds collected on the Falkland Islands prior to 1861 159

Remark: The collection site on the label, underside and in

Schlegel 1865 differ. The name Abbott was transcribed in-

correctly at label, underside and in Schlegel. The bird was ac-

quired through the Amsterdam-based dealer Gustav Adolph

Frank (see Kelp Goose RMNH.AVES.230433 as for C.C.

Abbott). Only Schlegel’s handwriting appears under the base

of the pedestal.

THINOCORIDAE

Least Seedsnipe Thinocorus rumicivorus rumicivorus

Eschscholtz, 1829 [RMNH.AVES.87538]. Loc: East

Falkland (ca. 51° 41' 40" S, 57° 51' 10" W). Date: in

either 1858 or 1859. Col: C.C. Abbott. Age/sex: 3. Acq:

via John Gould and Gustav Adolph Frank Sr. Status:

extant, accessed 1] February 2013. Tax: mounted (syn-

type). Publ: Abbott 1860, 1861; Gould 1859; Sclater

1860, 1861, 1862.

Pedestal label: not present.

Pedestal underside: [one hand] Cat. 5 / 4 / 1860 / Capt / Abbot

Stanley / Oost Falkland.

Remark: The bird was acquired through the Amsterdam-based

dealer Gustav Adolph Frank and arrived on 30 April 1860

(see Kelp Goose RMNH.AVES.230433 as for C.C. Abbott).

Only Schlegel’s handwriting appears under the base of the

pedestal.

SCOLOPACINAE

Hudsonian Godwit Limosa_ haemastica Linnae-

us, 1758 [RMNH.AVES.31529]. Loc: East Falkland

(ca. 51° 41' 40" S, 57° 51' 10" W). Date: in either 1858

or 1859. Col: C.C. Abbott. Age/sex: adult 4. Acq: via

John Gould and Gustav Adolph Frank Sr. Status: extant,

accessed 18 December 2012. Tax: mounted. Publ: Abbott

1860, 1861; Gould 1859; Schlegel 1864; Sclater 1860,

1861, 1862.

Pedestal label: [one hand] Limosa hudsonica / 1860 3 Cat: 1 /

Capt."* Abbot. Ile Or des Malouines.

Pedestal underside: [one hand] Limosa hudsonica / Cat 1/4 /

Cap / Abbot Stanley / Oost Falkland.

Schlegel 1864 (Tome V, 27: 22—23): Limosa hudsonica, Cat 1,

S au plumage parfait de noces, ile orientale des Malouines,

voyage du Capitaine Abbot, acquis en 1860.

Remark: The collection site on the label, underside and in

Schlegel 1864 differ. The name Abbott was transcribed in-

correctly at label, underside and in Schlegel. The label, un-

derside, and Schlegel 1864 all identify the specimen as male,

but only the latter identifies it as an adult. The bird was ac-

quired through the Amsterdam-based dealer Gustav Adolph

Frank (see Kelp Goose RMNH.AVES.230433 as for C.C.

Abbott). Only Schlegel’s handwriting appears under the base

of the pedestal.

White-rumped Sandpiper Calidris fuscicollis Vie-

illot, 1819 [RMNH.AVES.223070]. Loc: Falklands

(ca. 51° 41' 0" S, 59° 10' 0" W). Date: in either 1858 or

1859. Col: C.C. Abbott. Age/sex: —. Acq: via John Gould

and Gustav Adolph Frank Sr. Status: extant, accessed

11 February 2013. Tax: mounted. Publ: Abbott 1860,

Bonn zoological Bulletin 67 (2): 145-169

1861; Gould 1859; Schlegel 1864; Sclater 1860, 1861,

1862.

Pedestal label: [two hands] 7ringa Bonapartei / Cat. +0 6 /

Stanley 1860 / Malouines.

Pedestal underside: [two hands] 7ringa Bonapartei / Cat N° 10

/ Capt / Abbott Stanley / 1860 / Oost Falkland.

Schlegel 1864 (Tome V, 27: 43-43): Tringa bonapartei, Cat

10, Individu dans la livrée d’hiver. Tués dans l’ile orientale

des Malouines voyage du Capitaine Abbot Stanley, acquis en

1860.

Remark: The collection site on the label, underside and in

Schlegel 1864 differ. The name Abbott was transcribed in-

correctly at label, underside and in Schlegel. The bird was ac-

quired through the Amsterdam-based dealer Gustav Adolph

Frank (see Kelp Goose RMNH.AVES.230433 as for C.C.

Abbott).

White-rumped Sandpiper Calidris fuscicollis Vie-

illot, 1819 [RMNH.AVES.220882]. Loc: Falklands

(ca. 51° 41' 0" S, 59° 10' 0" W). Date: in either 1858 or

1859. Col: C.C. Abbott. Age/sex: Adult. Acq: via John

Gould and Gustav Adolph Frank Sr. Status: extant, ac-

cessed 11 February 2013. Tax: mounted. Publ: Abbott

1860, 1861; Gould 1859; Schlegel 1864; Sclater 1860,

1861, 1862.

Pedestal Label: [two hands] 7ringa Bonapartei / Cat. 7 /

Stanley 1860. Malouines.

Pedestal underside: [two hands] 7ringa / Bonapartei / Cat

N° 11 / Abbot / 1860 / Iles Falkland.

Schlegel 1864 (Tome V, 27: 43-43): Tringa bonapartei, Cat

11, Individu dans la livrée d’hiver. Tués dans l’ile orientale

des Malouines voyage du Capitaine Abbot Stanley, acquis en

1860.

Remark: The collection site on the label, underside and in

Schlegel 1864 differ. The name Abbott was transcribed in-

correctly at label, underside and in Schlegel. The bird was ac-

quired through the Amsterdam-based dealer Gustav Adolph

Frank (see Kelp Goose RMNH.AVES.230433 as for C.C.

Abbott).

White-rumped Sandpiper Calidris fuscicollis Vie-

illot, 1819 [RMNH.AVES.220883]. Loc: Falklands

(ca. 51° 41' 0" S, 59° 10' 0" W). Date: in either 1858 or

1859. Col: C.C. Abbott. Age/sex: —. Acq: via John Gould

and Gustav Adolph Frank Sr. Status: extant, accessed 11

February 2013. Tax: mounted. Publ: Abbott 1860, 1861;

Gould 1859; Schlegel 1864; Sclater 1860, 1861, 1862.

Pedestal Label: [two hands] Tringa Bonapartei / Cat. 42 8. /

Stanley 1860 / Malouines.

Pedestal underside: [one hand] 7ringa Bonapartei / Cat. N 12.

/ Capt Abbott / Stanley / 1860. / OostFalkland.

Schlegel 1864 (Tome V, 27: 43-43): Tringa bonapartei, Cat

12, Individu dans la livrée d’hiver. Tués dans l’ile orientale

des Malouines voyage du Capitaine Abbot Stanley, acquis en

1860.

Remark: The collection site on the label, underside and in

Schlegel 1864 differ. The name Abbott was transcribed in-

correctly at label, underside and in Schlegel. The bird was ac-

quired through the Amsterdam-based dealer Gustav Adolph

Frank (see Kelp Goose RMNH.AVES.230433 as for C.C.

Abbott). Only Schlegel’s handwriting appears under the base

of the pedestal.

©ZFMK

160 Justin J. F. J. Jansen

South American Snipe Gallinago paraguaiae magel-

lanica Vieillot, 1816 [RMNH.AVES.220457]. Loc: East

Falkland (ca. 51° 41' 40" S, 57° 51' 10" W). Date: in ei-

ther 1858 or 1859. Col: C.C. Abbott. Age/sex: adult. Acq:

via John Gould and Gustav Adolph Frank Sr. Status: ex-

tant, accessed 18 December 2012. Tax: mounted. Publ:

Abbott 1860, 1861; Gould 1859; Schlegel 1864; Sclater

1860, 1861, 1862.

Pedestal label: [one hand] Gallinago paraguaiae / ad: Cat. 6

Ile orientalis des / Cap: Abbot Stanley. / Malouines / 1860.

Pedestal underside: [two hands] Gallinago paraguaiae / Scol-

opax magellanica / King / Cat. 6. /Capt. / Abbot Stanley /

1860 / Oost Falkland.

Schlegel 1864 (Tome V, 27: 11-12): Gallinago paraguaiae, Cat

6, Adultes, ile orientale des Malouines, voyage du Capitaine

Abbot Stanley, acquis en 1860.

Remark: The collection site on the label, underside and in

Schlegel 1864 differ. Only the underside identifies the spec-

imen as adult. The bird was acquired through the Amster-

dam-based dealer Gustav Adolph Frank (see Kelp Goose

RMNH.AVES.230433 as for C.C. Abbott).

South American Snipe Gallinago paraguaiae magel-

lanica Vieillot, 1816 [RMNH.AVES.220458]. Loc: East

Falkland (ca. 51° 41' 40" S, 57° 51' 10” W). Date: in ei-

ther 1858 or 1859. Col: C.C. Abbott. Age/sex: adult. Acq:

via John Gould and Gustav Adolph Frank Sr. Status: ex-

tant, accessed 18 December 2012. Tax: mounted. Pub:

Abbott 1860, 1861; Gould 1859; Schlegel 1864; Sclater

1860, 1861, 1862.

Pedestal label: [two hands] Gallinago paraguaiae / Ad: Cat 7 /

Abbot Stanley / Ile or: des: Malouines / 1860.

Pedestal underside: [two hands] Gallinago paraguaiae / Scol-

opax magellanica / King / Cat. 7 / Capt. / Abbot Stanly / 1860

/ Oost Falkland.

Schlegel 1864 (Tome V, 27: 11-12): Gallinago paraguaiae, Cat

7, Adultes, ile orientale des Malouines, voyage du Capitaine

Abbot Stanley, acquis en 1860.

Remark: The collection site on the label, underside and in

Schlegel 1864 differ. The name Abbott was transcribed in-

correctly at label, underside and in Schlegel. Only Schlegel

1864 identifies the specimen as adult. The bird was acquired

through the Amsterdam-based dealer Gustav Adolph Frank

(see Kelp Goose RMNH.AVES.230433 as for C.C. Abbott).

LARIDAE

Brown-hooded Gull Chroicocephalus maculipennis

M.H.C. Lichtenstein, 1823 [RMNH.AVES.207906].

Loc: East Falkland (ca. 51° 41' 40" S, 57° 51' 10" W).

Date: in 1858. Col: C.C. Abbott. Age/sex: adult sum-

mer-plumage. Acq: via John Gould and Gustav Adolph

Frank Sr. Status: extant, accessed 18 December 2012.

Tax: mounted. Publ: Abbott 1860, 1861; Gould 1859;

Schlegel 1863c; Sclater 1860, 1861, 1862.

Pedestal label: [one hand] Cat n°8 / Larus glaucodes / Meyen /

Kapt. Abbot / 1859 / O. Falkland eilanden.

Bonn zoological Bulletin 67 (2): 145-169

Pedestal underside: [one hand] Larus glaucotis / Cat. N°1 /

Abbot Stanley / 1859 / Oostfalkland.

Schlegel 1863c (Tome VI, 23: 42): Larus glaucotis, Cat 1,

Adulte, plumage de noces, iles Falkland, voyage du Capi-

taine Abbot, acquis en 1859.

Remark: The collection site on the label, underside and in

Schlegel 1863c differ. The name Abbott was transcribed in-

correctly at label, underside and in Schlegel. Only Schlegel

1863c identifies the specimen as adult. Erroneously at the la-

bel are mentioned Catalogue number 8 (=1). The bird was ac-

quired through the Amsterdam-based dealer Gustav Adolph

Frank (see Kelp Goose RMNH.AVES.230433 as for C.C.

Abbott). Only Schlegel’s handwriting appears under the base

of the pedestal.

Brown-hooded Gull Chroicocephalus maculipennis

M.H.C. Lichtenstein, 1823 [RMNH.AVES.207908].

Loc: Falkland (ca. 51° 41' 0" S, 59° 10' 0" W). Date:

—. Col: — Age/sex: adult 3 summer-plumage. Acq: via

Gustav Adolph Frank Sr. Status: extant, accessed 18 De-

cember 2012. Tax: mounted. Publ: Schlegel 1863c.

Pedestal label: [one hand] cat: n°9 / Larus glaucodes / Meyen /

Frank / Falkland eilanden.

Pedestal underside: [one hand] Larus glaucotis / Cat N°2 / Par

Frank / Falkland.

Schlegel 1863c (Tome VI, 23: 42): Larus glaucotis, Cat 2, 3 au

plumage de noces, iles Falkland.

Remark: The collection site on the label, underside and in

Schlegel 1863c differ. Only Schlegel 1863c identifies the

specimen as adult <. Erroneously at the label is mentioned

Catalogue number 9 (=2). The bird was acquired through the

Amsterdam-based dealer Gustav Adolph Frank (see Kelp

Goose RMNH.AVES.230433). Only Schlegel’s handwriting

appears under the base of the pedestal.

Brown-hooded Gull Chroicocephalus maculipennis

M.H.C. Lichtenstein, 1823 [RMNH.AVES.207910].

Loc: Falkland (ca. 51° 41' 0" S, 59° 10' 0" W). Date: —.

Col: —. Age/sex: adult 4 summer-plumage. Acq: —. Sta-

tus: extant, accessed 18 December 2012. Tax: mounted.

Publ: Schlegel 1863c.

Pedestal label: [one hand] Cat n°10 / Larus glaucodes / Meyen

/ 6 / Falkland eilanden.

Pedestal underside: [one hand] Larus glaucotes 3 / Meyen /

(unreadable 3 words) / Cat N°3 / Iles Falkland.

Schlegel 1863c (Tome VI, 23: 42): Larus glaucotis, Cat 3, 3 au

plumage de noces, iles Falkland.

Remark: The collection site on the label, underside and in

Schlegel 1863c differ. The label and Schlegel 1863c identify

the specimen as male.

Only Schlegel’s handwriting under the base of the pedestal.

Erroneously at the label is mentioned Catalogue number 11

(=4). Only Schlegel’s handwriting at the base of the pedestal.

Brown-hooded Gull Chroicocephalus maculipennis

M.H.C. Lichtenstein, 1823 [RMNH.AVES.207912].

Loc: Falkland (ca. 51° 41' 0" S, 59° 10' 0" W). Date:

—. Col: —. Age/sex: adult 2 summer-plumage. Acq: via

Gustav Adolph Frank Sr. Status: extant, accessed 18 De-

cember 2012. Tax: mounted. Publ: Schlegel 1863c.

©ZFMK

Specimen labelling errors: birds collected on the Falkland Islands prior to 1861 161

Pedestal label: [one hand] Cat n° 11 / Larus glaucodes / Meyen

/ Falkland eilanden.

Pedestal underside: [two hands] g/aucotus / Larus / glaucotes

/ Cat N° 4/ Meyen / (unreadable)/ Frank / Mateutres / Ma-

louines.

Schlegel 1863c (Tome I, 32: 42): Larus glaucotis, Cat 4, Fe-

melle adulte, iles Falkland.

Remark: The collection site on the label, underside and in

Schlegel 1863c differ. Only Schlegel 1863c identifies the

specimen as adult. Erroneously at the label is mentioned Cat-

alogue number 11 (=4). Temminck and Schlegel handwriting

at the base of the pedestal.

Dolphin Gull Leucophaeus scoresbii Traill, 1823

[RMNH.AVES.206067]. Loc: East Falkland (ca.

51° 41' 40" S, 57° 51' 10" W). Date: in either 1858 or

1859. Col: C.C. Abbott. Age/sex: adult plumage. Acq:

via John Gould and Gustav Adolph Frank Sr. Status: ex-

tant, accessed 18 December 2012. Tax: mounted. Publ:

Abbott 1860, 1861; Gould 1859; Schlegel 1863c; Sclater

1860, 1861, 1862.

Pedestal label: [one hand] cat: n° 1. / Larus scoresbii / Traill. /

Capt. Abbot / 1860 / Falkland eil.

Pedestal underside: [two hands] scoresbii Traill. / Larus hae-

matorhynchus / Cat N°1 / feet a beak coral red / pupil white /

Capt Abbot / 1860 / Oost-Falkland.

Schlegel 1863c (Tome VI, 23: 33-34): Larus scoresbyi, Cat 1,

Adulte, iles Falkland, voyage du Captaine Abbot, 1860: bec

et pieds rouge de corail, iris de l’oeil blanc (Abbot).

Remark: The collection site on the label, underside and in

Schlegel 1863c differ. The name Abbott was transcribed

incorrectly at label, underside and in Schlegel. Only Schle-

gel 1863c identifies the specimen as adult. The bird was ac-

quired through the Amsterdam-based dealer Gustav Adolph

Frank and arrived on 30 April 1860 (see Kelp Goose RMNH.

AVES.230433 as for C.C. Abbott).

Dolphin Gull Leucophaeus scoresbii Traill, 1823

[RMNH.AVES.206069]. Loc: East Falkland (ca.

51° 41' 40" S, 57° 51' 10" W). Date: in either 1858 or

1859. Col: C.C. Abbott. Age/sex: Immature . Acq: via

John Gould and Gustav Adolph Frank Sr. Status: extant,

accessed 18 December 2012. Tax: mounted. Publ: Abbott

1860, 1861; Gould 1859; Schlegel 1863c; Sclater 1860,

1861, 1862.

Pedestal label: [one hand] cat. n°.2 / Larus scoresbii / Traill / 3

/ Capt Abbot / Frank 1861 / Falkland eil.

Pedestal underside: [two hands] scoresbii Traill. / Larus hae-

matorhynchus (unreadable) / 3 / Jeugdig kleed in / overgang

naar het vol- / wassen kleed / Cat. N°2. / Reis van / Capt.

Abbot / Frank 1861 / O. Falkland.

Schlegel 1863c (Tome VI, 23: 33-34): Larus scoresbyi, Cat 2,

3 en passage, meme origine que le No 1.

Remark: The collection site on the label, underside and in

Schlegel 1863c differ. The name Abbott was transcribed in-

correctly at label, underside and in Schlegel. The bird was ac-

quired through the Amsterdam-based dealer Gustav Adolph

Frank and arrived on 12 May 1861 (see Kelp Goose RMNH.

AVES.230433 as for C.C. Abbott).

Bonn zoological Bulletin 67 (2): 145-169

Kelp Gull Larus dominicanus dominicanus M.H.C.

Lichtenstein, 1823 [RMNH.AVES.207350]. Loc: East

Falkland (ca. 51° 41' 40" S, 57° 51' 10" W). Date: in

either 1858 or 1859. Col: C.C. Abbott. Age/sex: adult

S summer-plumage. Acq: via John Gould and Gustav

Adolph Frank Sr. Status: extant, accessed 18 December

2012. Tax: mounted. Publ: Abbott 1860, 1861; Gould

1859; Schlegel 1863c; Sclater 1860, 1861, 1862.

Pedestal label: [one hand] cat: n° 1. / Larus fuscus / domin-

icanus Licht. / 6 / Capt: Abbot / 1860 / Oost Falkland eil.

Pedestal underside: [one hand] Larus dominicanus / Cat N° 1

3 / Lect livid, Beak greenish yellow / Capt Abbot Stanley /

1859 / Oost Falkland.

Schlegel 1863c (Tome VI, 23: 12—13): Larus dominicanus, Cat

1, ¢ adultes, iles Falkland, voyage du Capitaine Abbot Stan-

ley, acquis en 1860: pieds d’un verdatre, passant par-ci par-la

au jaune.

Remark: The collection site on the label, underside and in

Schlegel 1863c differ. The name Abbott was transcribed

incorrectly at label, underside and in Schlegel. The label,

underside, and Schlegel 1863c all identify the specimen as

male, but only the latter identifies it as an adult. Disagree-

ment at the label and stand on the date of acquisition. The bird

was acquired through the Amsterdam-based dealer Gustav

Adolph Frank and arrived on 30 April 1860 (see Kelp Goose

RMNH.AVES.230433 as for C.C. Abbott). Only Schlegel’s

handwriting appears under the base of the pedestal.

Kelp Gull Larus dominicanus dominicanus M.H.C.

Lichtenstein, 1823 [RMNH.AVES.207352]. Loc: East

Falkland (ca. 51° 41' 40" S, 57° 51' 10" W). Date: in

either 1858 or 1859. Col: C.C. Abbott. Age/sex: adult 9.

Acq: via John Gould and Gustav Adolph Frank Sr. Sta-

tus: extant, accessed 18 December 2012. Tax: mounted.

Pub: Abbott 1860, 1861; Gould 1859; Schlegel 1863c;

Sclater 1860, 1861, 1862.

Pedestal label: [one hand] Cat n° 2 / Larus fuscus / domini-

canus Licht. / 2 / Capt Abbot / 1860 / Oost Falkland eil.

Pedestal underside: [one hand] Larus dominicanus / 9 / Cat

N° 2. / 1860 / Capt Abbot / 1860 / Oost Falkland.

Schlegel 1863c (Tome VI, 23: 12—13): Larus dominicanus, Cat

2, Femelle adultes, iles Falkland, voyage du Capitaine Abbot

Stanley, acquis en 1860: pieds d’un verdatre, passant par-ci

par-la au jaune.

Remark: The collection site on the label, underside and in

Schlegel 1863c differ. The name Abbott was transcribed

incorrectly at label, underside and in Schlegel. The label,

underside, and Schlegel 1863c all identify the specimen as

male, but only the latter identifies it as an adult. The bird

was acquired through the Amsterdam-based dealer Gustav

Adolph Frank and arrived on 30 April 1860 (see Kelp Goose

RMNH.AVES.230433 as for C.C. Abbott). Only Schlegel’s

handwriting appears under the base of the pedestal.

South American Tern Sterna hirundinacea_ Les-

son, 1831 [RMNH.AVES.209487]. Loc: Falkland

(ca. 51° 41' 0" S, 59° 10' 0" W). Date: —. Col: —. Age/

sex: Summer plumage. Acq: —. Status: extant, accessed

18 December 2012. Tax: mounted. Pub: Schlegel 1863c.

©ZFMK

162 Justin J. F. J. Jansen

Pedestal label: [one hand] Cat n° 1. / Sterna hirundinacea /

Lesson / Falkland eilanden.

Pedestal underside: [two hands] Sterna meridionalis / Sterna /

Peale / Cat n° 1 / Malouines / Malouines.

Schlegel 1863c (Tome VI, 24: 15): Sterna meridionalis, Cat 1,

Individu au plumage de noces, iles Malouines.

Remark: The collection site on the label, underside and in

Schlegel 1863c differ. Two name sources are mentioned,

Lesson on the label and Peale on the underside.

CATHARTIDAE

Turkey Vulture Cathartes aura jota (Molina,

1782) [RMNH.AVES.191314]. Loc: East Falkland

(ca. 51° 41' 40" S, 57° 51' 10" W). Date: in either 1860

or 1861. Col: C.C. Abbott. Age/sex: adult 2. Acq: via

John Gould and Gustav Adolph Frank Sr. Status: extant,

accessed 18 December 2012. Tax: mounted. Pub: Abbott

1860, 1861; Gould 1859; Schlegel 1862c:; Sclater 1860,

1861, 1862.

Pedestal label: [one hand] Cat: n°l / Rhinogryphus aura / °

falklandicus / (Sharpe) / Abbot colle.1861 / Falkland-eilan-

den.

Pedestal underside: [one hand] 19 Cathartes aura / Length

from tail (unreadable) / tip of tail: 2 feet 3 inches / Broadth

from tip to tip of wing, 5’2 / head & legs: pinkish flesh color

/ eye: dark brown) / (Capt. Abbot) / Cat N°1 / reis van / Capt

Abbot / Frank 1861 / Oost Falkland.

Schlegel 1862c (Tome VI, 10: 3-4): Cathartes aura, Cat 1, Fe-

melle adulte, ile orientale des Malouines, voyage du Capi-

taine Abbot, 1861.

Remark: The collection site on the label, underside and in

Schlegel 1862c differ. The name Abbott was transcribed in-

correctly at label, underside and in Schlegel. Only Schlegel

1862c identify the specimen as adult. The bird was acquired

through the Amsterdam-based dealer Gustav Adolph Frank

and arrived on 30 April 1860 (see Kelp Goose RMNH.

AVES.230433 as for C.C. Abbott). Only Schlegel’s hand-

writing appears under the base of the pedestal.

ACCIPITRIDAE

Cinereous’ Harrier Circus cinereus _ Vieillot,

1816 [RMNH.AVES. 191467]. Loc: Falklands

(ca. 51° 41' 0" S, 59° 10' 0" W). Date: in either 1858 or

1859. Col: C.C. Abbott. Age/sex: adult 4. Acq: via Gus-

tav Adolph Frank Sr. Status: extant, accessed 18 Decem-

ber 2012. Tax: mounted. Pub: Abbott 1860, 1861; Gould

1859; Schlegel 1862d; Sclater 1860, 1861, 1862.

Pedestal label: [one hand] Circus cinereus /m. Cat N°1 / par le

Capitaine / Abbot 1860 / Malouines.

Pedestal underside: [one hand] 21 Circus cinereus / 4 / Catal.

N° 1/eye golden yellow / feet bright yellow / beak green, tip

black / reis van / Capit. Abbot / (frank 1860) / falkland-eil.

Schlegel 1862d (Tome II, 13: 5-6): Circus cinereus, Cat 1, ¢

au plumage parfait, ile orientale des malouines, recueilli par

le Captaine Abbot, abtenu en 1860.

Remark: The collection site on the label, underside and in

Schlegel 1862d differ. The name Abbott was transcribed in-

Bonn zoological Bulletin 67 (2): 145-169

correctly at label, underside and in Schlegel. Only Schlegel

1862d identifies the specimen as adult @. The bird was ac-

quired through the Amsterdam-based dealer Gustav Adolph

Frank and arrived on 30 April 1860 (see Kelp Goose RMNH.

AVES.230433 as for C.C. Abbott). Only Schlegel’s hand-

writing appears under the base of the pedestal.

Cinereous Harrier Circus cinereus Vieillot, 1816

[RMNH.AVES.191468]. Loc: East Falkland (ca.

51° 41' 40" S, 57° 51' 10" W). Date: in either 1858 or

1859. Col: C.C. Abbott. Age/sex: adult 2. Acq: via John

Gould and Gustav Adolph Frank Sr. Status: extant, ac-

cessed 18 December 2012. Tax: mounted. Pub: Abbott

1860, 1861; Gould 1859; Schlegel 1862d; Sclater 1860,

1861, 1862.

Pedestal label: [one hand] Circus cinereus / f. Cat N°2 / pr Mr.

Abbot / 1860 / Malouines.

Pedestal underside: [one hand] 11 Circus cinereus / N° 2 / 9

in kleur = ¢ (Abbot) / reis van / Abbot / Frank 1860 / Oost

Falkland.

Schlegel 1862d (Tome II, 13: 5-6): Circus cinereus, Cat 2, Fe-

melle au plumage parfait, meme origine; (la femelle adulte

parfaitement semblable par ses teintes au vieux male: note du

Capitaine Abbot).

Remark: The collection site on the label, underside and in

Schlegel 1862d differ. The name Abbott was transcribed in-

correctly at label, underside and in Schlegel. Both the under-

side and Schlegel 1862d identify the specimen as female, but

only the latter identifies it as an adult. The bird was acquired

through the Amsterdam-based dealer Gustav Adolph Frank

(see Kelp Goose RMNH.AVES.230433 as for C.C. Abbott).

Only Schlegel’s handwriting appears under the base of the

pedestal.

Cinereous Harrier Circus cinereus __ Vieil-

lot, 1816 [RMNH.AVES.191469]. Loc: Falkland

(ca. 51° 41' 0" S, 59° 10' 0" W). Date: February — March

1820. Col: L’Uranie Joseph Paul Gaimard / Jean René

Constant Quoy. Age/sex: adult 4. Acq: via exchanged

with Paris Museum. Status: extant, accessed 18 De-

cember 2012. Tax: mounted. Pub: Freycinet et al. 1824;

Schlegel 1862d.

Pedestal label: [one hand] Circus cinereus / m. Cat N°3 / voy.

Quoy & Gaimard, / du Musee de Paris / Malouines.

Pedestal underside: [one hand] 3 / Circus cinereus / Falco his-

trionius G / 7 / Malouines.

Schlegel 1862d (Tome II, 13: 5-6): Circus cinereus, Cat 3, ¢

adulte Malouines, voyage de Quoy et Gaimard, du Musee

de Paris.

Remark: The underside, and Schlegel 1862d all identify the

specimen as male, but only the latter identifies it as an adult.

Only one name source is mentioned, Quoy & Gaimard on

the label and Schlegel 1862d. Only Schlegel’s handwriting

appears under the base of the pedestal.

Cinereous Harrier Circus cinereus Vieillot, 1816

[RMNH.AVES.191473]. Loc: Falkland (ca. 51° 41'0"S,

59° 10' 0" W). Date: March 1833 or March 1834. Col:

C. Darwin. Age/sex: young @. Acq: via Gustav Adolph

©ZFMK

Specimen labelling errors: birds collected on the Falkland Islands prior to 1861 163

Frank Sr. Status: extant, accessed 11 February 2013.

Tax: mounted. Publ: Gould 1839; Schlegel 1862d; van

Grouw & Steinheimer 2008.

Pedestal label: [one hand] Circus cinereus / m. Cat N°7. / voy:

Darwin. / 4 Janvier 1837 / Malouines.

Pedestal underside: [one hand] N°7 / Circus cinereus / 20 /

Jong 4 / reis / van / Darwin / frank 1860 / 4 January 1837 /

Oost. Falkland-eiland.

Schlegel 1862d (Tome II, 13: 5-6): Circus cinereus, Cat 7,

au premier plumage, tue le 4 Janvier 1837, ile orientale des

Malouines, voyage C. Darwin.

Remark: The collection site on the label, underside and in

Schlegel 1862d differ. The underside and Schlegel 1862d

identify the specimen as young male but lacks at the label.

Only Schlegel’s handwriting appears under the base of the

pedestal.

This bird is discussed by van Grouw & Steinheimer (2008),

as the missing bird from 4 January 1837, however, all notes

have been subsequently added and no original data is avail-

able on or from the specimen. Possibly at one stage the

original label was present. Darwin donated at 4 January

1837 mammals and 450 birds of HMS Beagle expedition

to the Zoological Society of London (e.g. to John Gould to

described them), 34-39 specimens that are still in existence

from the Society arrived in 1839, 1841 and 1856 at the Brit-

ish Museum. The Society’s labels recorded donor’s name, the

acquisition date and Darwin’s specimen’s numbers (e.g. C.

Darwin Esq, Jan 4 1837) (Steinheimer 2004).

Another notable remark we find on the underside as the

specimen was reportedly purchased from Frank in 1860.

The other 5 from 7 birds from HMS Beagle expedition in

Naturalis arrived in 1863 (Steinheimer 2004, van Grouw &

Steinheimer 2008). The largest numbers of birds from The

Falklands were those that arrived in 1859-1863 collected by

C.C. Abbott (see this paper). At 30 April 1860, a large load

arrived from Abbott via Frank at Naturalis, and amongst the

specimens were 3 Circus.

Cinereous Harrier Circus cinereus Vieillot, 1816

[RMNH.AVES.191476]. Loc: East Falkland (ca.

51° 41' 40" S, 57° 51’ 10" W). Date: in either 1858 or

1859. Col: C.C. Abbott. Age/sex: young @. Acq: via John

Gould and Gustav Adolph Frank Sr. Status: extant, ac-

cessed 18 December 2012. Tax: mounted. Pub: Abbott

1860, 1861; Gould 1859; Schlegel 1862d; Sclater 1860,

1861, 1862.

Pedestal label: [one hand] Circus cinereus / Cat N°8 / pr. Mr. le

Capitaine / Abbot / 1860 / Malouines.

Pedestal underside: [one hand] 22 Circus cinereus / No.8 / 3

jeugdig kleed / Abbot / Frank 1860 / Oost Falkland.

Schlegel 1862d (13: 5-6): Circus cinereus, Cat 7, Male, Ma-

louines, recueilli par le Capitaine Abbot, abtenu en 1860.

Remark: The collection site on the label, underside and in

Schlegel 1862d differ. The name Abbott was transcribed in-

correctly at label, underside and in Schlegel. Both the un-

derside and Schlegel 1862d identify the specimen as male,

but only the former identifies it as young. The bird was ac-

quired through the Amsterdam-based dealer Gustav Adolph

Frank and arrived on 30 April 1860 (see Kelp Goose RMNH.

AVES.230433 as for C.C. Abbott). Only Schlegel’s hand-

writing appears under the base of the pedestal.

Bonn zoological Bulletin 67 (2): 145-169

Variable Hawk Geranoaetus polyosoma polyosoma

(Quoy & Gaimard, 1824) [RMNH.AVES.191227]. Loc:

Falklands (ca. 51° 41'0" S, 59° 10' 0" W). Date: in either

1858 or 1859. Col: C.C. Abbott. Age/sex: adult 3’. Acq:

via John Gould and Gustav Adolph Frank Sr. Status: ex-

tant, accessed 18 December 2012. Tax: mounted. Pub:

Abbott 1860, 1861; Gould 1859; Schlegel 1862d; Sclater

1860, 1861, 1862.

Pedestal label: [one hand] Buteo polyosoma/ m. Cat N°1 / voy.

Cap. Abbot / Falkland.

Pedestal underside: [two hands] 30 polyosoma / 3 erythrono-

tus / eye Yellow, feet yellow / Cat N°1 / reis van / Capt Abbot

/ Frank 1860 / Falklands.

Schlegel 1862a (Tome II, 6: 12-13): Buteo polyosoma, Cat 1,

S au plumage parfait, mais le gris du dos montrant encore

des traces tres sensibles de la teinte rousse de I’ habit de pas-

sage; ile orientale de Falkland, voyage du Capitaine Abbot,

obtenu en 1860: iris de l’oeil et pieds jaunes (Abott).

Remark: One generic and two specific names are cited, with

the primary binomial on the label agreeing with Schlegel

1862d. The name Abbott was transcribed incorrectly at la-

bel, underside and in Schlegel. The underside, and Schlegel

1862d identify the specimen as male, but only the latter iden-

tifies it as an adult. For information on the Amsterdam-based

dealer Gustav Adolph Frank and C.C. Abbott see: Kelp

Goose RMNH.AVES.230433.

Variable Hawk Geranoaetus polyosoma polyosoma

(Quoy & Gaimard, 1824) [RMNH.AVES.191228]. Loc:

Falklands (ca. 51° 41' 0" S, 59° 10' 0" W). Date: —. Col:

—. Age/sex: 2. Acq: via Gustav Adolph Frank Sr. Status:

extant, accessed 18 December 2012. Tax: mounted. Pub:

Schlegel 1862d.

Pedestal label: [one hand] Buteo polyosoma / f. Cat N°2 / pr

Mr. Frank / Falkland.

Pedestal underside: [two hands] 46. polyosoma/ B. erythrono-

tus / Falco tricolor 9° d’Orbigny / voy.pl3.fig2 / Cat N°2 / pr

Mr Frank / Malouines et Chill.

Schlegel 1862a (Tome II, 6: 12-13): Buteo polyosoma, Cat 2,

Femelle, habit de passage, a manteaux et scapulaires d’un

roux rouygeatre uniforme; iles Falkland, pa Mr. Frank.

Remark: The collection site on the underside differs from

that in Schlegel 1862d. Only one name source is mentioned,

d’Orbigny on the underside. Three generic and three specif-

ic names are cited, with the primary binomial on the label

agreeing with Schlegel 1866. For information on the Am-

sterdam-based dealer Gustav Adolph Frank see: Kelp Goose

RMNH.AVES.230433.

Variable Hawk Geranoaetus polyosoma polyosoma

(Quoy & Gaimard, 1824) [RMNH.AVES.191229]. Loc:

Falkland (ca. 51° 41' 0" S, 59° 10’ 0" W). Date: —. Col:

—. Age/sex: 3. Acq: via Gustav Adolph Frank Sr. Status:

extant, accessed 18 December 2012. Tax: mounted. Pub:

Schlegel 1862d.

Pedestal label: [one hand] Buteo polyosoma / m. Cat N°3 / pr.

Mr. Frank / Falkland.

Pedestal underside: [two hands] 34. polyosoma/ B. erythrono-

tus 2 / 3 / Quoy / Cat N°3 / Frank / Falkland.

©ZFMK

164 Justin J. F. J. Jansen

Fig. 3. Charles Compton Abbott label, front (Variable Hawk Geranoaetus polyosoma, NHMUK 1955.6-20.2413; photograph by

Fig. 4. Charles Compton Abbott label, back (Variable Hawk Geranoaetus polyosoma, NHMUK 1955.6-20.2413; photograph by

Schlegel 1862a (Tome II, 6: 12-13): Buteo polyosoma, Cat 3, | Remark: Sexing is absent at the label, underside identifies J

male, habit de passage moins avance que celui du NO 2; roux as female, and Schlegel 1862d indetfies the bird as 3. For

rougeatre uniforme; iles Falkland, par Mr Frank. information on the Amsterdam-based dealer Gustav Adolph

Frank see: Kelp Goose RMNH.AVES.230433.

Specimen labelling errors: birds collected on the Falkland Islands prior to 1861 165

Variable Hawk Geranoaetus polyosoma polyosoma

(Quoy & Gaimard, 1824) [RMNH.AVES.191231]. Loc:

Falkland (ca. 51° 41'0" S, 59° 10' 0" W). Date: —. Col: -.

Age/sex: adult 4. Acq: via Gustav Adolph Frank Sr. Sta-

tus: extant, accessed 18 December 2012. Tax: mounted.

Pub: Schlegel 1862d.

Pedestal label: [one hand] Buteo polyosoma / m. Cat N°5 / pr.

Mr. Frank / Falkland.

Pedestal underside: [two hands] 38 / polyvosoma / Buteo eryth-

ronotus / King / jeune / Cat N°5 / pr. Frank / Malouines.

Schlegel 1862a (Tome II, 6: 12-13): Buteo polyosoma, Cat 5,

& au premier plumage, iles Falkland, par Mr. Frank.

Remark: The collection site on the label, underside and in

Schlegel 1862d differ. Both the underside and label identify

do not sex the bird, but Schlegel 1862d identifies it as an adult

&. Only one name source is mentioned, King on the under-

side. For information on the Amsterdam-based dealer Gustav

Adolph Frank see: Kelp Goose RMNH.AVES.230433.

Variable Hawk Geranoaetus polyosoma polyosoma

(Quoy & Gaimard, 1824) [RMNH.AVES.191232]. Loc:

East Falkland (ca. 51° 41' 40" 8, 57° 51' 10" W). Date: in

either 1860 or 1861. Col: C.C. Abbott. Age/sex: adult 3.

Acq: via John Gould and Gustav Adolph Frank Sr. Sta-

tus: extant, accessed 18 December 2012. Tax: mounted.

Pub: Abbott 1860, 1861; Gould 1859; Schlegel 1862d;

Sclater 1860, 1861, 1862.

Pedestal label: [one hand] Buteo polysoma / m. Cat N°6 / voy.

Cap. Abott / Falkland.

Pedestal underside: [two hands] 40 polyosomoma / 3 / Cat

N°6 / Reis van / cap. Abbot / Frank 1862 / Oost—Falkland.

Schlegel 1862a (Tome II, 6: 12-13): Buteo polyosoma, Cat 6,

S au premier plumage, iles Falkland, voyage du Capitaine

Abbot.

Remark: The collection site on the label, underside and in

Schlegel 1862d differ. The name Abbott was transcribed in-

correctly at label, underside and in Schlegel. Both the under-

side and Schlegel 1862d identify the specimen as male, but

only the latter identifies it as an adult. For information on

the Amsterdam-based dealer Gustav Adolph Frank see: Kelp

Goose RMNH.AVES.230433. Note the difference in latin

names on label and underside.

FALCONIDAE

Striated Caracara Phalcoboenus australis J.F. Gmelin,

1788 [RMNH.AVES.193332]. Loc: East Falkland (ca.

51° 41' 40" S, 57° 51' 10" W). Date: in either 1858 or

1859. Col: C.C. Abbott. Age/sex: adult ¢. Acq: via John

Gould and Gustav Adolph Frank Sr. Status: extant, ac-

cessed 18 December 2012. Tax: mounted. Pub: Abbott

1860, 1861; Gould 1859; Schlegel 1862b; Sclater 1860,

1861, 1862.

Pedestal Label: [one hand] Polyborus australis. / Cat N°1. /

voy. Cap Abbot. Falkland.

Pedestal underside: [one hand] 43 / Polyboris australis / 3 /

Feet & bare parts of the head / bright yellow / Cat N°1 / reis

van / Capt. Abbot / (Frank 1860) / Oost Falkland.

Bonn zoological Bulletin 67 (2): 145-169

Schlegel 1862b (Tome II, 9: 3-4): Polyborus australis, Cat 1,

& au plumage parfait, ile orientale de Falkland, voyage du

capitaine Abbot, obtenu en 1860.

Remark: The collection site on the label, underside and in

Schlegel 1862b differ. The name Abbott was transcribed in-

correctly at label, underside and in Schlegel. Both label as

underside lack ageing, in Schlegel 1862b the bird is aged.

The bird was acquired through the Amsterdam-based deal-

er Gustav Adolph Frank and arrived on 30 April 1860 (see

Kelp Goose RMNH.AVES.230433 as for C.C. Abbott). Only

Schlegel’s handwriting appears under the base of the ped-

estal.

Striated Caracara Phalcoboenus australis J.F. Gmelin,

1788 [RMNH.AVES.193333]. Loc: Falklands (ca. 51°

41’ 0" S, 59° 10' 0" W). Col: voyage La Coquille, R.P.

Lesson / P. Garnot. Date: November—December 1822.

Age/sex: adult. Acq: received in exchange with the Paris

Museum. Status: extant, accessed 11 February 2013. Tax:

mounted. Pub: Lesson & Garnot 1826-1830; Schlegel

1862d.

Pedestal label: [one hand] Polyboris australis. / Cat N°2. / voy.

de Coquille. Falkland.

Pedestal underside: [two hands] Polyborus australis / Falco

novae Zelandia/ Lath /37./ Coquille Pl. / Cat. N° 2 / Caraca-

ra funébre. / Temm.P1.Col.192. / Ost-Falkland / van-Bterren

/ Patagonia.

Schlegel 1862b (Tome II, 9: 3-4): Polyborus australis, Cat 2,

Individue au plumage parfait, iles Falkland, voyage de /a Co-

quille.

Remark: The collection site on the label, underside and in

Schlegel 1862b differ. Both label as underside lack ageing,

in Schlegel 1862b the bird is aged.

Striated Caracara Phalcoboenus australis J.F. Gme-

lin, 1788 [RMNH.AVES.193334]. Loc: Falkland

(ca. 51° 41' 0" S, 59° 10' 0” W). Date: —. Col: —. Age/sex:

adult. Acq: via Gustav Adolph Frank Sr. Status: extant,

accessed 1] February 2013. Tax: mounted. Publ: Schle-

gel 1862d.

Pedestal Label: [one hand] Polyborus australis. / Cat N°3. / pr.

Mr. Frank. Falkland.

Pedestal underside: [one hand] 29 / Polyborus australis /

Falco novae Zeelandiae Lath / Caracara funebre Tem /

PI.Col.192 / Cat N°3 / pr. Frank, Matetines / Falkland.

Schlegel 1862b (Tome II, 9: 3-4): Polyborus australis, Cat 3,

Individue au premier plumage, iles Falkland, par Mr. Frank.

Remark: The collection site on the label, underside and in

Schlegel 1862b differ. Both label as underside lack ageing, in

Schlegel 1862b the bird is aged. For information on the Am-

sterdam-based dealer Gustav Adolph Frank see: Kelp Goose

RMNH.AVES.230433. Two specimens are missing from

HMS Beagle (Steinheimer 2004). Only Schlegel’s handwrit-

ing appears under the base of the pedestal.

FURNARIIDAE

Blackish Cinclodes Cinclodes antarcticus Garnot,

1826 [RMNH.AVES.168495]. Loc: East Falkland (ca.

©ZFMK

166 Justin J. F. J. Jansen

51° 41' 40" S, 57° 51' 10" W). Date: in either 1858 or

1859. Col: C.C. Abbott. Age/sex: —. Acq: via John Gould

and Gustav Adolph Frank Sr. Status: extant, accessed 18

December 2012. Tax: mounted. Pub: Abbott 1860, 1861;

Gould 1859; Sclater 1860, 1861, 1862.

Pedestal label: [one hand] Cinclodes antarcticus (Garn.) / Voy.

Capt. Abbot Stanley. East Falkland / 1860.

Pedestal underside: [one hand] Capt. / Abbot Stanley / 1860 /

Oost Falkland.

Remark: The name giving of the location of collecting is

different in underside and label. Only Schlegel’s handwrit-

ing under the base of the pedestal. The name Abbott was

transcribed incorrectly at the underside and label. The bird

was acquired through the Amsterdam-based dealer Gustav

Adolph Frank and arrived on 30 April 1860 (see Kelp Goose

RMNH.AVES.230433 above).

TROGLODYTIDAE

Grass Wren Cistothorus platensis falklandicus Chap-

man, 1934 [RMNH.AVES.168365]. Loc: East Falkland

(ca. 51° 41' 40" S, 57° 51' 10" W). Date: in either 1858 or

1859. Col: C.C. Abbott. Age/sex: —. Acq: via John Gould

and Gustav Adolph Frank Sr. Status: extant, accessed 18

December 2012. Tax: mounted. Pub: Abbott 1860, 1861;

Gould 1859; Sclater 1860, 1861, 1862.

Pedestal label: [one hand] Cistothorus platensis (Lath.) /

Frank, 1860 / East Falkland.

Pedestal underside: [one hand] Frank / 1860 / Oost-Falkland.

Remark: The collection site on the label and underside differ.

The bird was acquired through the Amsterdam-based dealer

Gustav Adolph Frank and arrived on 30 April 1860 (see Kelp

Goose RMNH.AVES.230433 above). Only Schlegel’s hand-

writing appears under the base of the pedestal.

TURDIDAE

Austral Thrush TJurdus falcklandii falcklandii Quoy &

Gaimard, 1824 [RMNH.AVES.170710]. Loc: Falkland

(ca. 51° 41' 0" S, 59° 10' 0" W). Date: before 1850. Col:

—. Age/sex: —. Acq: —. Status: extant, accessed 18 Decem-

ber 2012. Tax: mounted. Pub: -.

Pedestal label 1: [one hand] Turdus falklandicus Quoy &

Gaimard / Avant 1850 / Falkland Isl.

Pedestal label 2: [two hands] Turdus falklandicus / Q et G. /

Patria? / Falkland Isl.

Pedestal underside: [one hand] Turdus falklandicus / Quoy et

Gaimard.

Remark: Handwriting is Temminck’s and therefore an old

specimen, most likely received in exchanges with the Pari-

sian Museum. One specimen is missing from HMS Beagle

(Steinheimer 2004).

Bonn zoological Bulletin 67 (2): 145-169

ICTERIDAE

Long-tailed Meadowlark Leistes loyca falklandica

(Leverkthn, 1889) [RMNH.AVES.168495]. Loc: East

Falkland (ca. 51° 41' 40" S, 57° 51' 10" W). Date: in ei-

ther 1858 or 1859. Col: C.C. Abbott. Age/sex: —. Acq: via

John Gould and Gustav Adolph Frank Sr. Status: extant,

accessed 18 December 2012. Tax: mounted. Pub: Abbott

1860, 1861; Gould 1859; Sclater 1860, 1861, 1862.

Pedestal label: [one hand] 7rupialis militaris (L.) / Cat. /

(Sclat. Ib. 1884: p. 23) / Acquis du Capt / Abbot Stanley 1860

/ Falkland occ.

Pedestal underside: [two hands] Trupialis militaris (L.)/3/ 3

/ Kapt. / Abbot Stanley / 1860 / Oost Falkland.

Remark: Only Schlegel’s handwriting under the base of the

pedestal. The name Abbott was transcribed incorrectly at la-

bel and in underside. The collection site on the label and un-

derside differ. Sexing lacks at the label. The name Abbott was

transcribed incorrectly at the underside and label. The bird

was acquired through the Amsterdam-based dealer Gustav

Adolph Frank and arrived on 30 April 1860 (see Kelp Goose

RMNH.AVES.230433).

EMBERIZIDAE

White-bridled Finch Melanodera melanodera melan-

odera (Quoy & Gaimard, 1824) [RMNH.AVES.165592].

Loc: East Falkland (ca. 51° 41' 40" S, 57° 51' 10" W).

Date: in either 1858 or 1859. Col: C.C. Abbott. Age/sex:

9. Acquisition: via John Gould and Gustav Adolph Frank

Sr. in 1860. Status: extant, accessed 18 December 2012.

Tax: mounted. Pub: Abbott 1860, 1861; Gould 1859;

Sclater 1860, 1861, 1862.

Pedestal label: absent.

Pedestal underside: [one hand] 9 / Capt / Abbot Stanley /

1860 / Oost Falkland.

Remark: The name Abbott was transcribed incorrectly at

the underside. The bird was acquired through the Amster-

dam-based dealer Gustav Adolph Frank and arrived on 30

April 1860 (see Kelp Goose RMNH.AVES.230433 above).

Only Schlegel’s handwriting appears under the base of the

pedestal.

White-bridled Finch Melanodera melanodera melan-

odera (Quoy & Gaimard, 1824) [RMNH.AVES.165582].

Loc: Falkland (ca. 51° 41' 0" S, 59° 10' 0" W). Date: —.

Col: —. Age/sex: 3. Acq: —. Status: extant, accessed 18

December 2012. Tax: mounted. Pub: -.

Pedestal label: [one hand] Chlorospiza Gray / Emberiza melan-

dera / Beagle pl 32. Uranie alt pl. / 3 / Malouines.

Pedestal underside: no text.

Remark: Handwriting is Temminck’s and therefore an old

specimen, most likely received in exchanges with the Pari-

sian Museum.

©ZFMK

Specimen labelling errors: birds collected on the Falkland Islands prior to 1861 167

DISCUSSION

Original labels

I found no original labels on Falkland specimens from

the French expeditions (both in Naturalis as in MNHN).

However, the information (albeit only for some) is re-

corded in a number of books and papers in the MNHN

archives; most of this information is also found on the

pedestal underside, with a summary of the information

from the pedestal underside being transcribed on the la-

bels (where the information was first recorded — in the

books and papers or under the pedestals — is uncertain).

The Falkland specimens in the MNHN were apparent-

ly treated in the same manner as were earlier specimens

from the Baudin expedition (389 specimens examined)

(Jansen 2018), which also arrived in France as skins.

Usually only single specimens, representing the male,

female, and young (if present and recognisable) of each

species collected on the Baudin expedition were kept,

with duplicates used for exchanges and donations (Jan-

sen 2016, 2018). When mounted, the known information

was transcribed on the pedestal underside by the muse-

um’s taxidermists, as it was for the 25 Falkland speci-

mens examined in the MNHN. The information found on

the pedestal underside does not have either the original

collector, the date of collection, or the specific collecting

locality within the Falklands.

For the Falkland birds collected by the British expe-

ditions, I found a number of specimens with original

labels. Of 69 birds examined collected by McCormick

on the Erebus and Terror expedition, 38 had an origi-

nal label (some of McCormick’s birds however are at-

tributed to the Admiralty). Of 3 birds examined from the

Rattlesnake expedition, only one had an original label;

one of those without an original label was collected by

MacGillivray. Of 15 birds examined from HMS Beagle

(collected by Covington/Darwin and Fitzroy and donated

by Burnett), none had original labels. However, most of

Abbott’s Falkland birds in the NHMUK still have their

original labels attached: of 16 examined, only one did not

have an original label or annotations from Abbott.

Original collector

We find on the labels attached to the Naturalis specimens

the following data regarding sources, including collectors

and expeditions: C.C. Abbott (46), Frank Merchants from

Amsterdam (21), no data (11), La Coquille (5), L’Astro-

labe (2), HMS Beagle (1), L’Uranie (1) and MNHN (1).

Of these, 3 birds (two from La Coquille and one with no

data) are not indisputably from the Falklands.

For 54 Naturalis birds (60%) — 46 birds collected by

Abbott and 7 birds from the French expeditions and

HMS Beagle — we can more or less determine the origi-

nal collector and date of collection. Similar data for the

Bonn zoological Bulletin 67 (2): 145-169

remaining 35 Naturalis birds (40%), however, is not pos-

sible to reconstruct. Based on information from Abbott

(Gould 1859, Abbott 1860, 1861) and data found in the

journals of the French expeditions and HMS Beagle, we

know the approximate period when collecting occurred

(odd is that the HMS Beagle bird is the sole bird with

a full date collecting date on it, and one of the very few

birds in Naturalis collected prior to 1850 with such a

dataset). However, precise data, including exact location,

date of collection, and specific collector, is beyond the

scope of this paper.

For the birds purchased from the Frank Merchants in

Amsterdam, we find 21 birds in Naturalis that have Frank

recorded as their source. Obviously, they were not the

collectors of these specimens, as no bird was collected at

all by the Frank family in the Falklands (there is no doc-

umented Falkland Islands visit from these Amsterdam

based merchants).

Did the Naturalis archives provide any additional help?

We only find notes in the Naturalis archives referring to

Falkland birds received from Frank to the Rijksmuseum

van Natuurlijke Historie (forerunner of Naturalis) from 3

May 1854 (Procellaria), 16 June 1856 and 30 April 1860.

These notes typically describe the acquired specimens

only vaguely.

Transcription errors and omissions

All birds in NHMUK from Abbott have recorded on their

original labels the month and year when collected. Other

details — in particular notations on bare parts — are also

present on some of the Abbott labels in the NHMUK.

However, older specimens (distinguished by having

smaller labels attached, while specimens collected later

have larger labels, with “Captain Abbott, Falkland Is-

lands Detachments” printed on the reverse) note only the

Species, sex, and month of collecting (a portion of Abbott

birds have the printed labels attached to them).

The 46 Abbott birds in Naturalis (51.6% of all (89)

Falkland birds in Naturalis) once had original labels

attached like those attached to the Abbott birds in the

NHMUK. However, of these 46 Naturalis specimens,

only 14 (29.8%) include some details on bare parts (all

noted on the pedestal underside); the remainder (70.2%)

lack such details. Of these 14 specimens, 8 have such

notes transcribed in English, while in the remaining 6,

the notes are translated into Dutch; the original contexts

of the latter are therefore missing, and no notes in the

original English can be traced. These notes are found

only on the pedestal underside and not in any other of the

available sources, such as Schlegel’s publications, new

tags, or pedestal labels. If we compare labels, pedestal

undersides and Schlegel’s various publications to each

other, we find no consistency in either the way Abbott

was addressed (e.g. either as “Mr.” or “Captain”, “Cat.”,

“Cap.”, “Capt.” and variations) or how his name was

©ZFMK

168 Justin J. F. J. Jansen

spelled (e.g. “Abbot”, “Abbott”, and “Abott”); more-

over, “voyage” or “Voy.” (e.g. “voyage du Capitaine

Abbot” and “Voy. Cap. Abbot”), “Stanley” (e.g. “Abbot

Stanley”, “Cat. Abbot Stanley”, “Voy: Abbot Stanly”

and “Voy. Abbot Stanley”), and “col” (e.g. “Abbot col’)

were also recorded. Neither the month or year when each

specimen was collected / arrived is recorded on any of

the 46 Abbott birds in Naturalis, as they are on all of the

NHMUK specimens.

CONCLUSIONS

Coenraad Jacob Temminck and Hermann Schlegel, both

collection managers of birds at the Rijksmuseum van

Natuurlijke Historie (now Naturalis), discarded all the

original labels at the time when original information and

precise data was not appreciated or as valued as it is today

(Steinheimer 2010); moreover, the data that was available

was imprecise and treated poorly, with the information

not being transcribed precisely and suffering alteration

over time. However, as shown in results, the damage can

be controlled for a large number of specimens by analysis

of the available original material and interpretation of the

information supplied by various sources.

Further research into diaries, letters and acquisition

books and the actual specimens should be considered in

the future to establish the correct labeling of the speci-

mens (for example to sort out the possible mix-up from

L’Uranie and La Coquille specimens). Also, the condi-

tion (looks fair to good in most specimens) of the mounts

/ skins can be examined as they could be subject to x-ra-

diation (Jansen & Steinheimer 2017).

Acknowledgments. I’m grateful for Robert Prys-Jones for

encouraging me to submit this paper and to Igor Fadeev for

his help. Pepijn Kamminga and Steven van der Mije gave me

access to the collections and helped me with the registration

numbers that were missing from my files. Hein van Grouw

(NHMUK) helped with missing register numbers of some of

the specimens and for granting me access on several occasions

over the years.

REFERENCES

Abbott CC (1861) Notes on the Birds of the Falkland Islands.

Ibis (3): 149-167

Abbott CC (1860) The Penguins of the Falkland Islands. Ibis

(2): 336-338

Darwin CR (1839) Narrative of the surveying voyages of His

Majesty’s Ships Adventure and Beagle between the years

1826 and 1836, describing their examination of the southern

shores of South America, and the Beagle’s circumnavigation

of the globe. Journal and remarks 1832-1836. John Murray,

London

Darwin CR (1859) On the origin of species by means of natural

selection, or the preservation of favoured races in the strug-

gle for life. John Murray, London

Bonn zoological Bulletin 67 (2): 145-169

Darwin CR (1871) The Descent of Man, and Selection in Rela-

tion to Sex. John Murray, London

de Freycinet LCD (ed.) 1824 Voyage autour du monde: entre-

pris par ordre du roi ... exécuté sur les corvettes de S. M.

l’Uranie et la Physicienne, pendant les années 1817, 1818,

1819 et 1820. Chez Pillet, Paris

de Freycinet LCD (ed.) (1837) Voyage autour du Monde...

exécuté sur les corvettes de L. M. «l’Uranie» et «La Physi-

cienne,» pendant les années 1817, 1818, 1819 et 1820. Paris

Gould J (1839) Part 3 (2): Birds. Pp. 17—32, pls. 11—20. In: Dar-

win C. (ed.) The zoology of the Voyage of H.M.S. Beagle,

under the command of Capt. Fitzroy, R.N., during the years

1832 to 1836, part 6. London

Gould J (1859) List of Birds from the Falklands, with Descrip-

tions of the Eggs of some of the species, from specimens col-

lected principally by Captain C. C. Abbott, of the Falkland

Islands Detachment. Proceedings of the Zoological Society

of London 27: 93-98

van Grouw H, Steinheimer FD (2008) Charles Darwin’s lost

Cinereous Harrier found in the collection of the National Mu-

seum of Natural History Leiden. Zoologische Mededelingen

Leiden 82 (48): 595-598

del Hoyo J, Collar NJ (2014) HBW and Birdlife Internation-

al Illustrated Checklist of the Birds of the World. Volume 1:

Non-Passerines. Lynx Edicions, Barcelona

del Hoyo J, Collar NJ (2016) HBW and Birdlife Internation-

al Illustrated Checklist of the Birds of the World. Volume 2:

Passerines. Lynx Edicions, Barcelona

Jansen JJFJ (2016) Towards the resolution of long-standing is-

sues regarding the birds collected during the Baudin expedi-

tion to Australia and Timor (1800-1804): the discrepancy in

the number of birds collected and their subsequent handling.

Journal of the National Museum (Prague). Natural History

Series 185 (2): 7-19

Jansen JJFJ (2018) The Ornithology of the Baudin expedition

(1800-1804). Privately published, Grave

Jansen JJFJ, van der Mije SD (2015) Review of the mounted

skins and skulls of the extinct Falkland Islands wolf, Dusicy-

on australis, held in museum collections. Archives of Natural

History 42 (1): 91-100

Jansen JJFJ, Steinheimer FD (2017) Research into the authen-

ticity of ‘Piwi Drepanis coccinea (G. Forster, 1781) skins

from Cook’s 3rd voyage: what taxidermy can add to the dis-

cussion. Bulletin of the British Ornithologists’ Club 137 (4):

246-260

Lanjouw J, Stafleu FA (1954) Index herbariorum; a guide to the

location and contents of the world’s public herbaria; Pt. II,

1; Collectors A-D. International Bureau for Plant Taxonomy

and Nomenclature of the International Association for Plant

Taxonomy, Utrecht

Lesson RP, Garnot P (1826-1830) Voyage autour du monde en-

trepris par ordre du government sur la corvette la Coquille

pendant les années 1822, 1823, 1824 et 1825. Paris

MacGillivray W (1852) Narrative of the voyage of H.MS.

Rattlesnake, commanded by the late Captain Owen Stanley

during the years 1846—50, including discoveries and surveys

in New Guinea, the Louisiade Archipelago, etc: to which is

added Mr. E.B. Kennedy’s expedition for the exploration of

the Cape York Peninsula. London

Mees GF (1953) The white-eyes of the Aroe Islands (Aves,

Zostreopidae). Zoolgische Mededelingen 32 (2): 25-30

Mees GF, Fisher CT (1986) On the type specimens of birds

from Lifu, described by E. L. Layard in 1878. Bulletin of the

British Ornithologists’ Club 106: 153-156

©ZFMK

Specimen labelling errors: birds collected on the Falkland Islands prior to 1861 169

Rasmussen PC, Prys-Jones RP (2003) History vs mystery: the

reliability of museum specimen data. Bulletin of the British

Ornithologists’ Club 123: 66—94

Schlegel H (1862a) Muséum d’ Histoire naturelle des Pays-Bas.

— Revue méthodique et critique des collections déposées

dans cet établissement. Tome II. 6: Buteones— August 1862.

E.J. Brill, Leiden

Schlegel H (1862b) Muséum d’ Histoire naturelle des Pays-Bas.

— Revue méthodique et critique des collections déposées dans

cet établissement. Tome II. 9: Polybori — November 1862.

E.J. Brill, Leiden

Schlegel H (1862c) Muséum d’ Histoire naturelle des Pays-Bas.

— Revue méthodique et critique des collections déposées

dans cet établissement. Tome II. 10: Vultures — November

1862. E.J. Brill, Leiden

Schlegel H (1862d) Muséum d’Histoire naturelle des Pays-

Bas. — Revue méthodique et critique des collections déposées

dans cet établissement. Tome II. 13: Circi— December 1862.

E.J. Brill, Leiden

Schlegel H (1863a) Muséum d’ Histoire naturelle des Pays-Bas.

— Revue méthodique et critique des collections déposées

dans cet établissement. Tome VI. 21: Pelecani — July 1863.

E.J. Brill, Leiden

Schlegel H (1863b) Muséum d’ Histoire naturelle des Pays-Bas.

— Revue méthodique et critique des collections déposées dans

cet établissement. Tome VI. 22: Procellaridae — July 1863.

E.J. Brill, Leiden

Schlegel H (1863c) Muséum d’ Histoire naturelle des Pays-Bas.

— Revue méthodique et critique des collections déposées

dans cet établissement. Tome VI. 32: Lari — August 1863.

E.J. Brill, Leiden

Schlegel H (1863d) Muséum d’Histoire naturelle des Pays-

Bas. — Revue méthodique et critique des collections déposées

dans cet établissement. Tome VI. 24: Sternae — September

1863. E.J. Brill, Leiden

Schlegel H (1864) Muséum d’ Histoire naturelle des Pays-Bas.

— Revue méthodique et critique des collections déposées

dans cet établissement. Tome V. 27: Scolopaces — November

1864. E.J. Brill, Leiden

Schlegel H (1865) Muséum d’ Histoire naturelle des Pays-Bas.

— Revue méthodique et critique des collections déposées

dans cet établissement. Tome IV. 29: Cursores — March 1865.

E.J. Brill, Leiden

Schlegel H (1866) Muséum d’ Histoire naturelle des Pays-Bas.

— Revue méthodique et critique des collections déposées

dans cet établissement. Tome VI. 31: Anseres — May 1866.

E.J. Brill, Leiden

Schlegel H (1867) Muséum d’ Histoire naturelle des Pays-Bas.

— Revue méthodique et critique des collections déposées

dans cet établissement. Tome VI. 33: Utrinatores — April

1867. E.J. Brill, Leiden

Sclater PL (1860) Catalogue of the. Birds of the Falkland Is-

lands. Proceedings of the Zoological Society of London 28:

382-391

Sclater PL (1861) Additions and Corrections to the list of the

Birds of Falkland Islands. Proceedings of the Zoological So-

ciety of London 29: 45—47

Sclater PL (1862) Catalogue of a collection of American birds.

N. Trubner and Co., London

Sharpe RB (1906) The history of the collections contained in

the natural history departments of the British Museum. Birds.

Trustees of the British Museum, London

Bonn zoological Bulletin 67 (2): 145-169

Steinheimer FD (2004) Charles Darwin’s bird collection and

ornithological knowledge during the voyage of H.M.S. Bea-

gle, 1831-1836. Journal of Ornithology 145: 300-320

Steinheimer FD (2010) Data-basing historical specimens — a

science of its own. proceedings of the 5th International Meet-

ing of European Bird Curators: Natural History Museum Vi-

enna, August 29th—31st, 2007, Vienna, Austria: 113-119

APPENDIX I

Birds as noted as collected on the Falklands on L’Ura-

nie from the lists of specimens that arrived in the

MNHN (archives Laboratory MNHN, October 1820,

unknown ammount arrived in 1821, Freycinet (also

indicated as Quoy & Gaimard))

‘goose’ Chloephaga 6, Chiloe Wigeon 3, “ducks” 3,

‘grebe’ 2+4, ‘heron’ 4, ‘pigeon’ 4, ‘rail’ 1, King Pen-

guin 1, ‘penguin’ Eudyptes 1, ‘penguin’ 1, ‘albatros’ 3,

‘petrel’ 3, Magellanic Cormorant 6, Snowy Sheatbill 1,

South American Snipe 1, Kelp Gull 1, ‘tern’ 4, Turkey

Vulture 4, Cinereous Harrier 2, Variable Hawk 2, Caraca-

ra 1, Blackish Cinclodes 4, Austral Thrush 7, Long-tailed

Meadowlark 1—2, White-bridled Finch 1.

Obviously the ‘pigeons’ are a mistake, as they don’t

occur on the Falklands (therefore the number 1s corrected

from 69 to 65).

Still present in MNHN (verified through a quick check)

Upland Goose 2, Mallard 1, Crested Duck 1, Yel-

low-billed Teal 2, White-tufted Grebe 3, Common Div-

ing-petrel Pelecanoides urinatrix 1, Rock Shag 3, South

American Snipe 2, Brown Skua Stercorarius antarcti-

cus 2, Black-crowned Night-heron 2, Sterna ssp 1, Tur-

key Vulture, Cinereous Harrier 2, Variable Hawk 1, Stri-

ated Caracara 4, Austral Thrush 1, White-bridled Finch 2.

APPENDIX II

Birds as noted as collected on the Falkland on La Co-

quille from the lists of specimens that arrived in the

MNHN (archives Laboratory MNHN, April 1825,

Garnot & Lesson)

Variable Hawk 5, Austral Thrush 1, Cinclodes 2, Plo-

ver 3, Oystercatcher 2, Grebe 2, Petrel 1, Skua 2, Cormo-

rant 1, Goose 1, Duck 4.

Still present in MNHN (verified through a quick check)

Kelp Goose 2, Falkland Steamer Duck 1, Southern Sil-

very Grebe 1, Rock Shag 2, Snowy Sheatbill 1, Mag-

ellanic Oystercatcher 1, Southern Lapwing 2, Striated

Caracara 1, Austral Thrush 1.

©ZFMK

BHL

i

Blank Page Digitally Inserted

Bonn zoological Bulletin 67 (2): 171-174

2018 - Smith P.

https://doi.org/10.20363/bzb-2018.67.2.171

ISSN 2190-7307

http://www.zoologicalbulletin.de

Scientific note

urn:|sid:zoobank.org:pub:E29AA 1 F1-9CA8-4E55-81 DF-CADAE8F6201C

Azara’s spinetails (Aves: Furnariidae).

The identity of No. 236 Chicli and No. 237 Cégogo

Paul Smith!

"FAUNA Paraguay, Encarnacion, Dpto. Itaptia, Paraguay; www,faunaparaguay.com

? Para La Tierra, Centro IDEAL, Mariscal Estigarribia 321 c/ Tte. Capurro, Pilar, Dpto. Neembuct, Paraguay;

www.paralatierra.org

E-mail: faunaparaguay@gmail.com

urn:|sid:zoobank.org:author:F43 14C5F-3 A6B-4067-AD97-908D6C5866BD

Abstract. The descriptions of birds provided by Félix de Azara (1742—1821) from his time in Paraguay constituted some

of the first and most detailed descriptions of the South American avifauna from the period. Whilst many formed the basis

for scientific names still in use today, others have remained inconclusively identified. In this paper the identities of two of

these descriptions referring to spinetails of the genus Synallaxis are elucidated. There has been considerable disagreement

amongst previous authors as to the correct identity of two of these descriptions. No. 236 Chicli is shown to refer to Soo-

ty-fronted Spinetail Synallaxis frontalis and No. 237 Cogogo to Pale-breasted Spinetail S. albescens.

Key words. Pale-breasted Spinetail, Sooty-fronted Spinetail, Synallaxis frontalis, Synallaxis albescens.

The ornithological magnum opus “Apuntamientos para

la historia natural de los paxaros del Paraguay y Rio de

la Plata” (1802-1805) by the Spanish military engineer

Félix de Azara (1742-1821) was one of the first serious

attempts to document the avifauna of the Southern Cone

of South America produced by a researcher living in the

study area. Based in Asuncion, Paraguay, Azara was a

meticulous student of the fauna, geography and culture of

the land he now called home, a broad area he called “Par-

aguay and La Plata”, which includes modern day eastern

Argentina, Paraguay, extreme southern Brazil and Uru-

guay. Working in close conjunction with the Jesuit priest

Padre Noseda, who resided in the town of San Ignacio,

Misiones department, he produced simple, but highly de-

tailed descriptions of 448 different “species”, over half of

which were unknown to science at that time. A detailed

chronology of his movements, life and work is provided

by Contreras (2010).

Despite over two centuries having passed since the pub-

lication of Azara’s work, not all of Azara’s descriptions

have been conclusively associated with known species

and a series of authors have attempted to identify and ap-

ply them to known taxa (Sonnini in Azara 1809; Hartlaub

1847; Burmeister 1861; von Berlepsch 1887; Bertoni

1901; Laubmann 1939; Pereyra 1945; Smith 2018; Smith

et al. 2018). Despite this some species remain unidenti-

fied, whilst others have been the subject of disagreement

between authors. In this paper I provide identifications

for two of Azara’s spinetails for which there has been no

previous consensus in the literature. Note that Azara’s

Received: 09.02.2018

Accepted: 28.11.2018

measurements are given in inches (= 25.4 mm), lines

(= 2.21 mm) and varas (= 83.6 cm).

No. 236 CHICLI (Azara Volume 2: 266)

“Le doy este nombre, porque lo canta con claridad en

tono alto y agudo, oyéndose de muy léjos, y repitiéndolo

de modo, que las pausas no duran mas que los cantares.

No dudo que la hembra es silenciosa, y que no difiere del

macho, pues seguramente he visto y tenido cien indivi-

duos idénticos, todos en el Paraguay. Es estacionario y

solitario sin abundar. Habita entre los caraguatas 6 aloes

y matorrales espesos, sin subir jamas a dos varas de al-

tura, ni dexarse ver a descubierto. Esta en movimiento

continuo, sin salir a los campos, ni internarse en bosques

grandes, y sin duda come arafias é insectos. Sus vuelos se

reducen a pasar de un matorral a otro inmediato: y aun-

que no conoce esquivez, es dificil verle en sus guaridas,

y al oirle se figurara el que no lo conozca que esta en lo

alto del arbol 6 matorral, quando se halla al pie entre la

ramazon 0 caraguatas.

Longitud 6 4 pulgadas: cola 3: braza 7. El angosto

tupé es pardo, y sobre la cabeza, el encuentro, cobijas

y cola, de un bermellon algo obscurecido en la cola. De

esta al cogote pardo acanelado, y los remos como la cola.

Baxo de la cabeza hay una mancha de plumas negras,

con las puntas casi blancas, que tapan lo negro quando

estan bien ordenadas. De alli al pecho pardo blanquizco,

como la ceja y el costado de la cabeza. Baxo el cuerpo

Corresponding editor: T. Topfer

Published: 06.12.2018

172

mas blanquizco : sus costados como el lomo, pero mas

claros; y las tapadas de canela clara y linda.

Remos 19, céncavos y endebles, el quarto mayor:

la cola 10 plumas agudas, débiles y en escalera: la de

afuera 24 lineas mas corta: pierna 13: tarso 10, verdoso

aplomado: dedo medio 6 ‘2: pico 5, muy comprimido por

los costados, recto con la puntita poco corva, que excede

a la mitad inferior, obscuro encima, blanquizco debaxo,

y el iris color de cafia roxiza.”

(My translation)

“T give it this name because it sings it with clarity in a

high and sharp tone, being audible from far away, and

repeating it ina way that the pauses between notes are of

lesser duration than the notes themselves. I dont doubt

that the female is silent, and doesnt differ from the male;

in fact I have surely seen and had hundreds of identical

individuals, all in Paraguay. It is resident and solitary,

but not abundant. It lives amongst the bromeliads or al-

oes, and in dense scrub, without ever rising to a height of

more than 1.67 metres, nor does it let itself be seen in the

open. It is continuously on the move, without ever com-

ing out into the fields or entering the great forests, and

without a doubt it eats spiders and insects. Its flight is re-

duced to passing from one patch of scrub to the next: and

although it isnt shy, it is difficult to see it in its haunts,

and upon hearing it the inexperienced observer will think

that it is high in the tree or scrub, when in fact it is at the

base amongst the bushes and bromeliads.

Length 158.8 mm: tail 76.2 mm: wing span 177.8 mm.

The thin forehead is grey, and the top of the head, the

wing bend, coverts and tail are bright reddish, slightly

darker on the tail. From the forehead to the nape brown-

ish-cinnamon, and the flight feathers like the tail. Under

the head there is a spot of black feathers with the tips

white, which cover the black when they are well ordered.

From there to the breast brownish-grey, as with the su-

percilium and sides of the head. Under the body more

greyish : the sides as the back, but paler; the underwing

an attractive pale cinnamon.

Flight feathers 19, concave and weak, the fourth is lon-

gest: the tail with 10 sharp, weak feathers, increasing in

size towards the centre pair: the outermost 53 mm short-

er: leg 28.7 mm: tarsus 22.1 mm, greenish-grey: mid-toe

14.4 mm: bill 11.1 mm, very compressed at the sides,

straight with a slightly hooked tip, that is longer than the

lower mandible, dark grey above, pale grey below, with

the iris the color of red rum.”

Associated with Synallaxis ruficapilla Vieillot, 1819 by

Hartlaub (1847) and Burmeister (1861), Synallaxis fron-

talis frontalis von Pelzeln, 1859 by Bertoni (1901) and

Laubmann (1939) and Synallaxis albescens albescens

Temminck, 1823 by Pereyra (1945). The description of

Bonn zoological Bulletin 67 (2): 171-174

Paul Smith

the voice, habits, measurements, forehead and tail colour

indeed identify it as Sooty-fronted Spinetail Synallaxis

frontalis von Pelzeln, 1859, a species that 1s abundant

and widespread in Paraguay (Guyra Paraguay 2005).

Though Pereyra (1945) noticed that the measurements

and throat colour were more consistent with S. frontalis,

he seems to have erred when he stated that the voice in-

dicated S. albescens. Apparently unfamiliar with the spe-

cies in the field and unduly convinced by the similarity

of Azara’s call description to the common Argentinian

name Pujiui or Chicli that he attributed to this species

(today the former being a generic name for any Synallax-

is spinetail in Argentina) he opted for S. albescens. In fact

Azara’s name Chicli (which is based on the call note) is

a near perfect rendition of that of S. frontalis, and quite

unlike the typical wee-bidget call given by Paraguayan

S. albescens.

No. 237 COGOGO (Azara Volume 2: 268)

“Noseda y yo encontramos bastantes por septiembre y

octubre en las cercanias del pueblo de S. Ignacio guazu

del Paraguay; y observamos, que aunque iban solos, las

hembras no estaban léjos, ni diferian en medidas y colo-

res. El canto, peculiar del macho, no es brillante, pero

puede pasar entre los adocenados, aunque tiene poca

variedad, y lo repite con mucha frequencia. Habita los

matorralitos, mantillas y pillas de leita de los campos,

sin internar en los bosques grandes, ni subir a las man-

tillas altas y aparentes; y desde el momento que se posa

empieza a corretear lo interior y mas baxo de la escoba 6

matorranito, prefiriendo al parecer los secarrones y que

tienen al pies mas broza que verdura. Tanto por su con-

tinua inquietud, como por el cuidado con que se esconde

huyendo del lugar 6 costado donde le puedan ver, es muy

dificil matarlo. Sus vuelos son por lo comun cortos ; pero

a veces los prolonga para buscar matorrales lejanos, con

mayor velocidad de la que parece compete a sus alas.

Vive de los insectos que pilla en dichos matorralitos y en-

tre el pasto muy alto y cerrado. Le llamo Cogogo, porque

me aseguran que asi canta en tiempo que no es de amor,

con la gravedad y fuerza que un pdaxaro grande; pero no

salgo fiador de que esto sea verdad.

Longitud 5 7% pulgadas : cola 2 "/2, braza 6 12. Viendo

que este paxaro no diferia del anterior, maté a4 un Chicli

y a un Cogogo en el mismo dia para compararlos, bus-

cando a uno y otro en sus respectivos domicilios, diri-

giéndome por su voz, a fin de evitar toda equivocacion.

Los exdminé con todo mi cuidado, y hallé que las formas

del todo y de cada una de las partes eran absolutamen-

te idénticas, sin que la mayor perspicacia pudiese notar,

sino que el Chicli tiene el ala ménos tendida. Por lo que

hace a colores y el lugar que ocupan, son identicos, sin

exceptuar los del pico, iris y tarso, pues aunque con la

mayor atencion se nota que el Cogogo no tiene tan en-

©ZFMK

Azara’s spinetails 173

Table 1. Mean and range measurements of Azara’s descriptions compared with morphometrics of mist-netted individuals of the two

proposed species (P. Smith unpublished data). Synallaxis albescens three individuals from Encarnacion, Itapua department (Octo-

ber 2005), Laguna Blanca, San Pedro department (November 2005) and Parque Nacional Teniente Enciso, Boqueron department

(July 2006); Synallaxis frontalis four individuals from Encarnacion, Itapua department (August and October 2005).

[no 257 Cages | Synalanis abescens 3) _[ No. 236 Chicli| Synaltanis Fontalis (=)

cendida la cola, sobre la cabeza, y cobijas; y que el color

sobre el cuerpo es poquito ménos acanelado, y por deba-

xo algo mas blanquizco: estas diferencia son tan peque-

fias que caben sobradamente en la misma especie, y solo

se advierten en el cotejo inmediato, sin poderse expresar

con palabras; de modo que pueden decirse nulas, y que

es imposible distinguir los paxaros por otros caracteres

que la voz y costumbres; pues aunque la magnitud sea

algo diferente, es tan corta cosa, que cabe en la propria

especie, y aun en el modo de medir.

Remos 19, poco céncavos, no muy deébiles, el prime-

ro bastante corto, y el tercero, quarto y quinto iguales:

cola 10 plumas en todo como la del Chicli, y en ambos

la central agudisima, las demas no tanto: el resto como

en el anterior.”

(My translation)

“Noseda and I found many during September and Octo-

ber close to the town of San Ignacio Guazu in Paraguay,

and we observed that, although they went alone, the fe-

males were not far away, and they didnt differ in form or

colour. The song, unique to the male, is not brilliant, but

can pass as mediocre, although it shows little variety and

is repeated frequently. It inhabits scrub, bushes and wood

piles in the fields, without entering the great forests, nor

climbing the tallest most conspicuous bushes ; and from

the moment it lands it begins to hop around the interior

and lowest bush, preferring it would seem the drier parts

and with the base more brushy than leafy. It is extremely

difficult to kill, as much for its constant restlessness as for

the care it takes to hide itself by fleeing from the place it

can be seen. Its flights are typically short ; but occasion-

ally they are prolonged to find more distant patches of

scrub, with greater speed than it seems from how it beats

its wings. It lives on insects that it finds in the aforemen-

tioned scrub and amongst the high, dense grass. I call it

Cogogo, because they assure me that this is how it sings

Bonn zoological Bulletin 67 (2): 171-174

outside of the reproductive season, with the gravity and

strength of a large bird; but I cannot personally vouch

Jor this being true.

Length 143.8 mm : tail 74.1 mm, wing span 165.1 mm.

Seeing that this bird did not differ from the previous spe-

cies, I killed a Chicli and a Cégogo on the same day to

compare them, looking for each one in their respective

haunts, being guided by the voice so to avoid mistakes. I

examined them with all my care, and found that the struc-

ture of both was absolutely identical, and with only the

most careful attention was it noticeable that the Chicli

has the wing less elongated. In terms of colour and the

parts of the body where they occur, they are identical,

except for the bill, iris and tarsus, and with the great-

est of attention it can be noted that the Cogogo does not

have the colour of the tail, top of the head and coverts

so brightly coloured; and that the colour of the upper-

parts is a little less cinnamon, and below it is somewhat

paler : these differences are so slight that they fit within

the variation of a single species, and they are only obvi-

ous in direct comparison, without being able to express

them in words; to the point where it could be said they

are non-existent and that it impossible to distinguish the

species by characters other than the voice and their be-

haviour; you see although the size is somewhat different,

it is so minor a difference that it is consistent with the

same species, and even with the method of measurement.

Flight feathers 19, slightly concave, not very weak,

the first very short, and the third, fourth and fifth equal:

tail 10 feathers, as in the Chicli, and in both the central

feathers are very sharp, the others not so much: the rest

is as for the previous species.”

The identity of this form has been much debated, with

Sonnini in Azara (1809) stating that it was a hitherto

unknown species, Hartlaub (1847) noting only that the

bird was not mentioned by Vieillot, Burmeister (1861)

associating the description with Synallaxis fulginiceps

©ZFMK

174 Paul Smith

(= Leptasthenura fulginiceps d’Orbigny & Lafresnaye,

1837 which does not occur in Paraguay), von Berlepsch

(1887) and Bertoni (1901) identifying it only so far as

Synallaxis sp., Laubmann (1939) preferring not to offer

an identification and Pereyra (1945) opting for Synallaxis

jrontalis frontalis von Pelzeln, 1859.

Azara notes that with Padre Noseda they found “sever-

al during September and October close to the town of San

Ignacio Guazu” (Misiones department). He describes

this species as extremely similar to the “Chicli”, differ-

ing in voice, habitat, paler underparts, slightly smaller

size (Tab. 1) and less “cinnamon” upperparts including

the tail, crown and coverts which are less “brightly-co-

loured.” These differences are clear enough to identify

this bird as Pale-breasted Spinetail Synallaxis albescens

Temminck, 1823 in that area of Paraguay, where it oc-

curs sympatrically with S. frontalis the only other species

with which it may be confused (Guyra Paraguay 2005,

Narosky & Yzurieta 2006).

Azara states that he uses the name Cogogo because he

has been assured that this is how the bird sings in the

non-breeding season “with the gravity and strength of a

large bird”. However he clarifies that he cannot vouch

for the veracity of that statement, indicating that he never

heard the bird produce such a call. The description of the

call is possibly due to confusion by locals with Chotoy

Spinetail Schoeniophylax phryganophila Vieillot, 1817,

a superficially similar bird that occurs in the same areas

and habitats, and with a call that may be rendered in such

a way. Regardless Azara’s admission that the information

is secondhand and the fact that no Paraguayan Synallaxis

is known to change its call during the non-breeding sea-

son is enough to discard it from consideration.

Acknowledgements. The author would like to express his ad-

miration for the work of Félix de Azara upon which this paper

is based. Thanks to Clara Rojas for assistance with the meaning

of some older Spanish words. The anonymous librarians behind

the online Biodiversity Heritage Library Project made some of

the older, more obscure references cited here available to a wid-

er readership. It would not have been possible without them.

The support of the Pronii Project of CONACYT Paraguay is

gratefully acknowledged.

REFERENCES

Azara F de (1805) Apuntamientos para la Historia Natural de

los Paxaros del Paraguay y Rio de la Plata. Tomo 2. Imprenta

de la Viuda de Barra, Madrid, Spain

Bonn zoological Bulletin 67 (2): 171-174

Azara F de (1809) Voyages dans |’Amérique Meéridionale. Oi-

seaux. Vol. 3. Dentu, Paris, France

Berlepsch H von (1887) Appendix. Systematisches Verze-

ichniss in der Republik Paraguay bisher beobachteten Vo-

gelarten. Journal fiir Ornithologie 35: 113-134

Bertoni A de W (1901) Aves nuevas del Paraguay. Continuacion

a Azara. Anales Cientificos Paraguayos 1: 1-216

Burmeister H (1861) Reise durch die La Plata-Staaten mit be-

sonderer Rticksicht auf die physische Beschaffenheit und den

Culturzustand der Argentinischen Republik, Zweiter Band.

HW Schmidt, Halle

Contreras JR (2010) Félix de Azara. Su Vida y su Epoca.

Tomo 2. Calidad Grafica, Zaragoza, Spain

Guyra Paraguay (2005) Atlas de las aves de Paraguay. Guyra

Paraguay, Asuncion, Paraguay

Hartlaub CJG (1847) Systematischer Index zu Don Félix du

Azara’s Apuntamientos para la Historia Natural de los Pax-

aros del Paraguay y Rio de la Plata. Schiinemann, Bremen

Laubmann A (1939) Die Vogel von Paraguay. Band 1. Strecker

und Schroder Verlag, Stuttgart

Narosky T, Yzurieta D (2006) Guia para la identificacion de las

aves de Paraguay. Vazquez Mazzini, Buenos Aires

Pelzeln A von (1859) Uber neue Arten der Gattungen Synallax-

is, Anabates und Xenops in der kaiserlichen ornithologischen

Sammlung, nebst Ausztigen aus Johann Natterer’s nachge-

lassenen Notizen tber die von ihm in Brasilien gesammelten

Arten der Subfamilien: Furnarinae und Synallaxinae. Sitzu-

ngsberichte der Kaiserlichen Akademie der Wissenschaften,

Mathematisch-Naturwissenschaftlichen Classe 34: 99-134

Pereyra JA (1945) La obra ornitologica de Don Félix de Azara.

Biblioteca Americana, Buenos Aires

Smith P, Pacheco JF, Bencke GA, Aleixo A (2018). Se-

nior synonyms for three Neotropical birds described by

Vieillot based on Azara (Passeriformes: Thraupidae,

Tyrannidae, Tityridae). Zootaxa 4433: 141-150.

https://do1.org/10.11646/zootaxa.4433.1.8

Smith P (2018). The identity of Sylvia ruficollis Vieillot, 1817

and Azara’s No. 240 “Cola aguda cola de canela obscura”

(Aves; Furnariidae). Ardea 106: 79-83

Temminck CJ (1823) Nouveau recueil de planches coloriées

d’ oiseaux, pour servir de suite et de complément aux planch-

es enluminées de Buffon. Dufour & d’Ocagne, Paris, France

Vieillot LP (1817) Nouveau dictionnaire d’histoire naturells,

appliquée aux arts, a agriculture, a l’économie rurale et do-

mestique, a la médicine, etc. par una société de naturalistes

et d’agriculteurs. Nouvelle édition. Tome 7. Deterville, Paris,

France

Vieillot LP (1819) Nouveau dictionnaire d’histoire naturells,

appliquée aux arts, a agriculture, a l’économie rurale et do-

mestique, a la médicine, etc. par una société de naturalistes et

d’agriculteurs. Nouvelle édition. Tome 32. Deterville, Paris,

France

©ZFMK