Bonn zoological Bulletin 70 (2): 227-245

2021 - Wachkoo A.A. et al.

https://do1.org/10.20363/BZB-2021.70.2.227

ISSN 2190-7307

http://www.zoologicalbulletin.de

Research article

urn:|sid:zoobank.org: pub: 165FB2AA-6B1 D-446A-9191-EACFB1 8FD732

Taxonomic review of the ant genus Lepisiota Santschi, 1926

(Hymenoptera: Formicidae: Formicinae) from India

Aijaz Ahmad Wachkoo!', Himender Bharti? & Shahid Ali Akbar®*

' Department of Zoology, Government Degree College, Shopian, Jammu and Kashmir, 192303 India

? Department of Zoology and Environmental Sciences, Punjabi University, Patiala, Punjab, 147002 India

3 Division of Plant Protection; Department of Entomology, Central Institute of Temperate Horticulture, Srinagar, Jammu and

Kashmir, 191132 India

“Corresponding author: Email: kingakbarali@gmail.com

'urn:lsid:zoobank. org:author:6F 19EB1F-5 DDC-4722-BBD3-F75C29F901D9

2urn:Isid:zoobank.org:author: 5CFEBC9B-3CA9-4459-83A6-6D7B61B984B7

3urn:lsid:zoobank.org:author:5 AOAC4C2-B427-43A D-840E-7BB4F2565A 8B

Abstract. The species-rank taxonomy of the genus Lepisiota Santschi, 1926 in India is revised. Thirteen species are re-

cognized, with two described as new: L. Jayla sp. n. and L. mayri sp. n. The three previously infraspecific taxa L. integra

stat. nov., L. pulchella stat. rev. and L. wroughtonii stat. rev. are elevated to species rank. Four species or subspecies are

excluded from the Indian Lepisiota fauna: L. capensis (Mayr, 1862), L. frauenfeldi (Mayr, 1855), L. rothneyi watsonii

(Forel, 1894), and L. simplex (Forel, 1892) for issues related to previous doubtful distribution or species misidentifcation.

An identification key to the worker caste of Indian Lepisiota species is provided.

Keywords. Ants, Formicinae, key, Lepisiota, new species, new status, revived status, India.

INTRODUCTION

The formicine ant genus Lepisiota Santschi, 1926 con-

tains 138 species and subspecies, including two that are

newly described here (Sharaf et al. 2020; Bolton 2021),

and is widespread in the grasslands, savannahs or wood-

lands of the Afrotropical, Indomalayan, and Palearctic

regions (Brown 2000; Hita Garcia et al. 2013). The tax-

onomy of the genus is in a dreadful condition with the

lack of any revisionary studies for most of the zoogeo-

graphical regions of the World (Sharaf et al. 2020). How-

ever, a number of authors have published isolated species

descriptions and updated, taxonomic revisions for several

regional faunas (Sharaf et al. 2020; Bolton 2021).

The Lepisiota fauna of India 1s poorly known. Most of

the species and infraspecific taxa are poorly defined and

have been treated under different genera. Apart from the

major contributions in the late nineteenth and the early

twentieth century (Forel 1892, 1894, 1895, 1902a; Bing-

ham 1903), and subsequent isolated treatments (Muker-

jee 1930; Bharti 2002), the fauna has been neglected tax-

onomically.

In this study, we revise the species-rank taxonomy of

the ant genus Lepisiota for India. We describe the two

new species L. Jayla and L. mayri and recognize the three

poorly defined sympatrically distributed infraspecific

Received: 14.12.2020

Accepted: 08.06.2021

taxa integra, pulchella and wroughtonii, at species rank.

The data reported herein represent the first deep insight

of the Indian Lepisiota with the hope of correcting some

of the taxonomic neglect that has plagued the Indian For-

micinae (Bharti & Wachkoo 2012, 2014a, b; Wachkoo &

Bharti 2015a, b).

MATERIAL AND METHODS

The taxonomic study was conducted on a Nikon SMZ

1500 stereoscope. For digital images, MP Evolution dig-

ital camera was used on the same microscope with Au-

to-Montage (Syncroscopy, Division of Synoptics, Ltd.)

software. Later, images were cleaned with Adobe Pho-

toshop CSS.

Institutional abbreviations

MHNG = Museum of Natural History, Geneva,

Switzerland

MSNG~ = Natural History Museum, Genoa, Italy

NHMUK = Natural History Museum, London, UK

PUAC — = Punjabi University Patiala Ant Collection,

Punjab, India

Corresponding editor: R. Peters

Published: 25.06.2021

228 Aijaz Ahmad Wachkoo et al.

Zoological Survey of India, Kolkata,

India

Holotype and paratypes of both new species, and other

newly collected material have been deposited in PUAC.

One paratype of both new species will be deposited at

NHMUK. Morphological terminology for measurements

(given in millimeters) and indices found below follow

Wachkoo & Bharti (2014a, b) and Akbar et al. (2017).

Abbreviations for morphological terms and indices

used in the text

CI = Cephalic index: HW/HL x 100

EL = Maximum length of eye as measured normally

in oblique view of the head to show full surface

of eye

HL = Maximum length of head in full-face view,

measured in straight line from the anterior most

point of the median clypeal margin to a line

drawn across the posterior margin

HW = Maximum width of head in full-face view

(excluding the portion of eyes that extends past

the lateral margins of the head)

ML = Mesosomal length measured from the anterior

surface of the pronotum proper (excluding the

collar) to the posteriormost point of the propo-

deal lobes

PFL = Maximum length of the profemur from its

margin with the trochanter to its margin with the

tibia

PFW = Maximum width of the profemur

= Maximum width of the pronotum in dorsal view

Scape index: SL/HW x 100

SL = Maximum length of the antennal scape

excluding the basal neck and condyle

REL = Relative eye length index: EL/HL = 100

—

SYSTEMATIC ACCOUNTS

Genus Lepisiota Santschi, 1926

Type species: Plagiolepis rothneyi Forel, 1894

Worker diagnosis. Antenna 11-segmented; eyes

well-developed, ocelli frequently present; propodeum

armed with a pair of spines, teeth, or tubercles; petiole a

scale-like with the dorsal margin bispinose, bidentate or

emarginated; acidopore well-developed (Bolton 1994).

Synoptic list of Indian species of Lepisiota

Lepisiota annandalei (Mukerjee, 1930)

Lepisiota bipartita (Smith, 1861)

Lepisiota fergusoni (Forel, 1895)

Lepisiota integra (Forel, 1894) stat. nov.

Bonn zoological Bulletin 70 (2): 227—245

Lepisiota layla sp. n.

Lepisiota lunaris (Emery, 1893)

Lepisiota mayri sp. n.

Lepisiota modesta (Forel, 1894)

Lepisiota opaca (Forel, 1892)

Lepisiota pulchella (Forel, 1892) stat. rev.

Lepisiota rothneyi (Forel, 1894)

Lepisiota sericea (Forel, 1892)

Lepisiota wroughtonii (Forel, 1902) stat. rev.

Lepisiota annandalei (Mukerjee, 1930) (Figs 1-3)

Acantholepis annandalei Mukerjee, 1930: 156, fig. 4.

Syntype workers, Shimla [Simla], Himachal Pradesh,

India [ZSI].

Lepisiota annandalei (Mukerjee); Bolton 1995: 226; first

combination in Lepisiota.

Worker measurements. HL 0.50—0.54; HW 0.46—

0.52; EL 0.16-0.18; SL 0.48-0.56; PnW 0.31-0.35;

ML 0.60-0.68; PFL 0.38—0.43; PFW 0.11—-0.13 mm. In-

dices: CI 91-95; SI 102-108; REL 29-33 (n = 14).

Description. Head subquadrate; longer than wide,

distinctly wider posteriorly than in front; lateral margins

convex, posterior margin transverse, posterolateral cor-

ners rounded; clypeus subcarinate in the middle; ante-

rior clypeal margin complete and convex; eyes broadly

oval, weakly convex, placed at the middle-line of head,

covering one-third of lateral cephalic margin; three small

ocelli present; antennal scape surpassing posterior head

margin by about one-third its length.

In lateral view promesonotum dome like, convex;

metanotum distinctly lower than promesonotum but al-

most as high as propodeum; mesometanotum demarcat-

ed, constricted; metanotal area distinct; propodeal spines

reduced to tubercles; propodeal declivity steep.

Petiole upright with angular sides, dorsally without

distinct teeth or spines, shallowly emarginate; apical cor-

ners with acute angles pointing upward.

Overall body smooth and shiny.

Gaster covered with short, abundant, erect setae; head

and pronotum with only a few setae; pubescence on body

uniformly very fine and sparse; antennal funiculus with

appressed to subdecumbent pubescence.

Color black; antenna, mandible and tarsi light brown.

Distribution and habitat. India (Himachal Pradesh,

Jammu and Kashmir, Punjab, Sikkim, Uttarakhand, West

Bengal) (Bharti et al. 2016; present study). Although,

infrequent in collections this species seems widespread

across India; workers were mostly hand collected from

tree trunks and honey baits.

Remarks. This species has largely been misidentified

as Afrotropical L. simplex (Forel, 1892) due to its su-

perficial resemblance with the latter. The small to medi-

um-sized workers can be distinguished from the some-

what similar LZ. /unaris by their smooth and shiny body,

a dorsally emarginate petiole and few setae on the prono-

©ZFMK

Review of the ant genus Lepisiota from India 229

Figs 1-3. Worker; Lepisiota annandalei (Mukerjee, 1930). 1.

Head, frontal view. 2. Body, lateral view. 3. Body, dorsal view.

Bonn zoological Bulletin 70 (2): 227—245

tum; L. /unaris has a relatively dull and microreticulate

body, a bispinose petiole and several pairs of setae scat-

tered on the mesosoma including the propodeum.

Material examined. India: Himachal Pradesh,

Andretta, 32.0744° N, 76.5856° E, 940 m, 6 99,

12.vi.2010; Renuka, 30.6083° N, 77.4615° E, 600 m,

3 QQ, 8.v.2009; Terrace, 31.9234° N, 75.9294° E,

420 m, 2 99, 11.x.2008. Jammu and Kashmir, Mans-

ar, 32.6979° N, 75.1489° E, 690 m, 3 99, 13.vii.2009.

Punjab, Dharampur, 31.8420° N, 75.9132° E, 450 m,

5 99, 14.x.2008. Uttarakhand, Dakpathar, 30.4966° N,

77.8004° E, 750 m, 5 QQ, 20.viii.2009; Forest Rese-

arch Institute, 30.3416° N, 77.9903° E, 640 m, 6 99,

1.x.2008; Mussoorie, 30.4606° N, 78.0521° E, 1820 m,

4 OQ, 9.viii.2009; Selaqui, 30.3720° N, 77.8605° E,

670 m, 3 29, 7.vill.2009, Aijaz A. Wachkoo leg.

Lepisiota bipartita (Smith, 1861) (Figs 4-6)

Formica bipartita Smith, 1861: 33. Syntype workers,

Lebanon; Israel, Holy Land [NHMUK]. [Images of

CASENT0903167 Syntype worker examined].

Acantholepis bipartita (Smith); Roger 1863: 11; first

combination in Acantholepis.

Acantholepis bipartita (Smith); Mayr 1863: 394 as junior

synonym of Acantholepis frauenfeldi.

Acantholepis frauenfeldi var. bipartita (Smith); André

1882: 211 queen described and revived from synony-

my as variety of frauenfeldi.

Acantholepis frauenfeldi bipartita (Smith); Emery 1891:

16; Emery 1898: 126; Forel 1902b: 155; Ruzsky 1905:

463; Santschi 1917: 47; variety/subspecies of frauen-

feldi.

Acantholepis frauenfeldi var. bipartita (Smith); Kara-

vaiev 1911: 44; male described.

Lepisiota frauenfeldi subsp. bipartita (Smith); Baroni

Urbani et al. 1992: 303; first combination in Lepisiota.

Lepisiota bipartita (Smith); Collingwood & Agosti 1996:

365; status as species.

Worker measurements. HL 0.66—0.80; HW 0.57—

0.71; EL 0.19-0.25; SL 0.98-1.09; PnW 0.36-0.53;

ML 0.97-1.06; PFL 0.75—0.98; PFW 0.14—0.20 mm. In-

dices: CI 84-88; SI 148-171; REL 28-31 (n= 15).

Description. Head subrectangular; longer than wide,

equally wide anteroposteriorly; lateral and posterior

margins convex, posterolateral corners rounded; clypeus

medially subcarinate to carinate; anterior clypeal margin

complete and convex; eyes subglobulose, convex, pro-

jecting beyond cephalic lateral margins, covering one-

third of lateral cephalic margin, placed at posterior half

of head; three ocelli present; antenna long, scape surpass-

ing posterior head margin by about half its length.

In lateral view pronotum convex, mesometanotum

strongly constricted, lower than pronotum and propo-

deum, giving mesosoma a dumbbell shape; mesometa-

©ZFMK

230 Aijaz Ahmad Wachkoo et al.

i 0.2 mm

Figs 4—6. Worker; Lepisiota bipartita (Smith, 1861). 4. Head,

frontal view. 5. Body, lateral view. 6. Body, dorsal view.

Bonn zoological Bulletin 70 (2): 227—245

notum demarcated; metanotal area distinct; propodeum

armed with a pair of teeth diverging posteriorly; propo-

deal declivity steep.

Petiole upright with distinctly angular sides, dorsally

emarginate, apical corners teeth like pointing upward.

Head and mesosoma feebly shining; gaster relatively

more shiny; overall body weakly microreticulate, howev-

er, sculpturing is effaced in some specimens.

Body covered with erect setae on head, pronotum and

gaster; setae on gaster usually restricted to venter and

segmental margins; pubescence very fine and sparse,

most visible on head; almost absent on mesosoma and

gaster; antennal funiculus with appressed to decumbent

pubescence.

Head and gaster dark brown; mesosoma reddish brown;

antenna and tarsi brown.

Distribution and habitat. Greece, India (Andhra

Pradesh, Gujarat, Haryana, Himachal Pradesh, Jammu

and Kashmir, Karnataka, Kerala, Maharashtra, Megha-

laya, Punjab, Rajasthan, Uttarakhand, West Bengal),

Iran, Israel, Lebanon (Bharti et al. 2016; Dad et al. 2019;

Sharaf et al. 2020). This species is widespread across In-

dia. The specimens were collected by hand, from honey

baits and by beating vegetation.

Remarks. Lepisiota bipartita is a medium to large-

sized ant. The main feature that will enable it to be dis-

tinguished from L. integra, the species with which it is

most likely to be confused, is the shiny bicolored body,

whereas the latter species has a dull and uniformly red-

dish-brown body.

Material examined. India: Himachal Pradesh: Andret-

ta, 32.0744° N, 76.5856° E, 940 m, 1 9, 11.vi.2010, 1 9,

27.vill.2009. Jammu and Kashmir: Manda, 32.7496° N,

74.8673° E, 500 m, 1 9, 15.vii.2009, Aijaz A. Wachkoo

leg.

Lepisiota fergusoni (Forel, 1895) (Figs 7-9)

Acantholepis fergusoni Forel, 1895: 459. Syntype work-

ers, Thiruvananthapuram [Travancore], Kerala, In-

dia [MHNG]. [Images of CASENT0909883 Syntype

worker examined].

Acantholepis fergusoni Forel; Bingham 1903: 319.

Lepisiota fergusoni (Forel); Bolton 1995: 227; first com-

bination in Lepisiota.

Worker measurements. HL 0.69-0.72; HW 0.65—

0.66; EL 0.17-0.19; SL 0.69-0.70; PnW 0.42-0.45;

ML 0.95—1.05; PFL 0.59-0.61; PFW 0.16—0.17 mm. In-

dices: CI 91-94; SI 105-107; REL 24—26 (n = 5).

Description. Head subquadrate; slightly longer than

wide, wider posteriorly than in front; lateral margins

convex, posterior margin convex, with rounded postero-

lateral corners; clypeus carinate in the middle; anterior

clypeal margin complete and convex; eyes oval, weakly

convex, placed at the middle-line of head, covering about

one-third of lateral cephalic margin; three small ocelli

©ZFMK

Review of the ant genus Lepisiota from India 231

ymm

Figs 7-9. Syntype worker (AntWeb: CASENT0909883); Lepi-

siota fergusoni (Forel, 1895). 7. Head, frontal view. 8. Body,

lateral view. 9. Body, dorsal view. (Photograph courtesy of

Zach Lieberman).

present; antennal scape surpassing posterior head margin

by about one-third its length.

In lateral view promesonotum convex, metanotum low,

almost straight; pronotum flat above; mesometanotum

demarcated; metanotal area distinct; mesometanotum

constricted; propodeum armed with a pair of diverging

Bonn zoological Bulletin 70 (2): 227—245

blunt spines directed backward; propodeal declivity

Steep:

Petiole upright with angular sides, dorsally without

distinct teeth or spines, shallowly emarginate; apical cor-

ners with acute angles pointing upward.

Head distinctly reticulate-striate, opaque; mesosoma

dorsally fine microreticulate, overall opaque, laterally

relatively shiny; gaster polished smooth and shining:

propleuron relatively smoother, remainder of lateral

mesosoma longitudinally striate; mesometanotal suture

cross-ribbed.

Body abundantly covered with erect setae; pubescence

very fine and sparse; antennal funiculus with subdecum-

bent to suberect pubescence.

Head reddish-brown, mesosoma and petiole red-

dish-yellow, gaster reddish-brown with a light red-

dish-yellow patch anteriorly on first gastral tergite.

Distribution and habitat. India (Karnataka, Kerala,

West Bengal), Sri Lanka (Bharti et al. 2016; Dias et al.

2020). This species is infrequent in collections and seems

to be restricted to the coastal plains of India. The speci-

mens were collected in a leaf litter sample.

Remarks. Lepisiota fergusoni is a medium-sized ant.

The reticulate-striate head and blunt propodeal spines

directed backward allow distinction from the somewhat

similar L. pulchella which has a reticulate-punctate head

and pointed propodeal spines directed upward.

Material examined. India: Kerala: Periyar Tiger Re-

serve, Thanikkudy, 9.4997° N, 77.2665° E, 1003 m,

5 99, 15.x.2011, Shahid A. Akbar leg.

Lepisiota integra (Forel, 1894) stat. nov. (Figs 10-12)

Acantholepis frauenfeldi var. integra Forel, 1894: 411.

Syntype workers, Dharamshala, Himachal Pradesh, In-

dia [MHNG]. [Images of CASENT0909888 Syntype

worker examined].

Acantholepis frauenfeldi integra Forel; Forel 1906: 86;

Santschi 1917: 44; race/stirps of frauenfeldi.

Lepisiota frauenfeldi var. integra (Forel), Bolton 1995:

227, first combination in Lepisiota.

Worker measurements. HL 0.66—0.75; HW 0.57—

0.66; EL 0.20—-0.22; SL 0.82-0.95; PnW 0.39-0.49;

ML 1.00—1.12; PFL 0.66—0.77; PFW 0.13—0.16 mm. In-

dices: CI 84-88; SI 140-151; REL 27—30 (n= 10).

Description. Head subrectangular; longer than wide,

almost equally wide anteroposteriorly; lateral and poste-

rior margins convex, posterolateral corners rounded; cly-

peus medially subcarinate; anterior clypeal margin com-

plete and convex; eyes subglobulose, convex, projecting

beyond cephalic lateral margins, covering one-third of

lateral cephalic margin, placed at posterior half of head;

three ocelli present; antennal scape surpassing posterior

head margin by about half its length.

In lateral view pronotum convex, mesometanotum

strongly constricted, lower than pronotum and propo-

©ZFMK

232 Aijaz Ahmad Wachkoo et al.

deum, giving mesosoma a dumbbell shape; mesometa-

notum demarcated; metanotal area distinct; propodeum

armed with a pair of teeth diverging posteriorly; propo-

deal declivity steep.

Figs 10-12. Worker; Lepisiota integra (Forel, 1894) stat. nov.

10. Head, frontal view. 11. Body, lateral view. 12. Body, dorsal

view.

Bonn zoological Bulletin 70 (2): 227—245

Petiole upright, with distinctly angular sides, dorsum

emarginate with teeth-like apical corners.

Overall body dull and opaque, relatively densely and

evenly microreticulate.

Body covered with erect setae on head and pronotum;

gastral setae restricted to apical segments; pubescence

relatively more as in Lepisiota bipartita and Lepisiota

sericea, fine and sparse, most visible on head and gaster;

sparser on mesosoma; antennal funiculus with appressed

to decumbent pubescence.

Color uniformly reddish-brown.

Distribution and habitat. India (Himachal Pradesh,

Jammu and Kashmir, Madhya Pradesh, Meghalaya, Pun-

jab, Uttarakhand), Pakistan (Bharti et al. 2016; Rasheed

et al. 2019). This species seems to be general in distri-

bution in India; it was collected in non-forest as well as

forest habitats. Workers were mostly collected by beat-

ing vegetation, Winkler’s extractor, hand collecting, from

honey baits and in pitfall traps.

Remarks. Lepisiota integra is raised here to species

rank, originally described by Forel (1894) as an infra spe-

cies taxon frauenfeldi var. integra. This is a medium to

large-sized ant with a dull and uniformly reddish-brown

body and a dorsally emarginate petiole, whereas L. frau-

enfeldi (Mayr, 1855) is a relatively shiny, paler species

with mesosoma, legs and antennae orange, distinctly

contrasting the brown head and gaster, and a dorsally

dentate petiole.

Material examined. India: Himachal Pradesh:

Baijnath, 32.0527° N, 76.6500° E, 1125 m, 9 99,

17.v1.2010; Bakhra, 31.4087° N, 76.4327° E, 650 m,

21 QQ, 7.x.2008; Ghamrur, 31.6620° N, 76.0601° E,

460 m, 9 29, 1.v1.2009; Ghatti, 31.9300° N, 75.9302° E,

425 m, 1 Q, 12.x.2008; Guraldhar, 31.6670° N,

76.4684° E, 660 m, 15 99, 2.vi.2009; Jogi Panga,

31.5408° N, 76.3161° E, 600 m, 13 99, 9.x.2008; Re-

nuka, 30.6083° N, 77.4615° E, 600 m, 5 29, 8.v.2009;

Terrace, 31.9234° N, 75.9294° E, 420 m, 10 99,

24.v.2009, 20 99, 25.v.2009, 2 99, 9.vii.2010. Jammu

and Kashmir: Manda, 32.7496° N, 74.8673° E, 500 m,

8 QQ, 15.vil.2009; Mansar, 32.6979° N, 75.1489° E,

690 m, 4 99, 13.vi1.2009. Punjab: Chohal, 31.6666° N,

76.0666° E, 450 m, 17 99, 11.x.2008. Uttarakhand: Fo-

rest Research Institute, 30.3416° N, 77.9903° E, 640 m,

3 29, 11.v.2009, 3 99, 12.v.2009; Ranger’s College,

30.3225° N, 78.0445° E, 660 m, 20 99, 22.v.2010,

9 OQ, 25.v.2010, Aijaz A. Wachkoo leg.

Lepisiota layla sp. n. (Figs 13-15)

urn: |sid:zoobank.org:act:4AF8&C610-5E75-42B6-921F-E21 DED00054E

Type material. Holotype worker. India, Himach-

al Pradesh, Kotla, 31.8821° N, 75.9963° E, 500 m,

22.x.2010. Paratype, 1 worker, same data as for holotype,

Aijaz A. Wachkoo leg.

©ZFMK

Review of the ant genus Lepisiota from India 233

Figs 13-15. Holotype worker; Lepisiota layla sp. n. 13. Head,

frontal view. 14. Body, lateral view. 15. Body, dorsal view.

Bonn zoological Bulletin 70 (2): 227—245

Worker measurements. HL 0.57—0.58; HW 0.57—

0.58; EL 0.18-0.19; SL 0.64-0.65; PW 0.40-0.41;

PFL 0.50-0.51; PFW 0.12-0.13; ML 0.86—0.87 mm. In-

dices: CI 100; SI 112; REL 33 (n= 2).

Description. Head subquadrate; as long as wide, as

wide posteriorly as in front; lateral margins convex, pos-

terior margin transverse with strongly rounded postero-

lateral corners; clypeus subcarinate in the middle; ante-

rior clypeal margin complete and convex; eyes broadly

oval, weakly convex, placed at the middle-line of head,

covering about one-third of lateral cephalic margin; three

small ocelli present; antennal scape surpassing posterior

head margin by about one-third its length.

In lateral view promesonotum convex, metanotum low,

almost straight; mesometanotum demarcated; metanotal

area distinct; mesometanotum constricted; propodeum

armed with a pair of posteriorly diverging thick blunt

spines; propodeal declivity steep.

Petiole upright with angular sides, dorsally without

distinct teeth or spines, shallowly emarginate; apical cor-

ners with acute angles pointing upward.

Overall body polished, smooth and shining; mesometa-

notal suture cross-ribbed; mesometapleuron longitudi-

nally striate.

Body abundantly covered with long, erect white setae;

scape and legs with relatively shorter erect setae than

on body; antennal funiculus with dense suberect pubes-

cence.

Color black; scape and tarsi yellow brown.

Distribution and habitat. India (Himachal Pradesh).

This species seems to be rare in India: the specimens

were found in the Shivalik range of Western Himalaya.

The specimens were collected by hand from the trunk of

an Acacia tree.

Etymology. The species epithet is an Arabic noun

meaning dark beauty, in reference to the shining black

color of this species.

Remarks. Lepisiota layla is a medium-sized ant. It is

superficially similar to L. /unaris and L. modesta, but its

body is abundantly covered with long, erect white setae

while L. /unaris and L. modesta have a body covered

with usual short erect setae.

Lepisiota lunaris (Emery, 1893) (Figs 16-18)

Acantholepis lunaris Emery, 1893a: 250, plate 6,

fig. 12. Syntype workers, Colombo, Sri Lanka [Cey-

lon] [MSNG]. [Images of CASENT0905157 Syntype

worker examined].

Acantholepis capensis lunaris Emery; Forel 1895: 458;

Forel 1906: 86; Forel 1909: 395; Emery 1925: 24; Me-

nozzi 1939: 312; Chapman & Capco 1951: 209; race/

subspecies of capensis.

Acantholepis capensis subsp. lunaria Emery; Chap-

man & Capco 1951: 209; misspelled as /unaria.

Acantholepis lunaris Emery; Collingwood 1970: 379;

status as species.

©ZFMK

234 Aijaz Ahmad Wachkoo et al.

Figs 16—18. Worker; Lepisiota lunaris (Emery, 1893). 16. Head,

frontal view. 17. Body, lateral view. 18. Body, dorsal view.

Bonn zoological Bulletin 70 (2): 227—245

Acantholepis lunaris Emery; Imai et al. 1984: 9; karyo-

type described.

Lepisiota capensis subsp. /unaris (Emery); Bolton 1995:

228; first combination in Lepisiota.

Lepisiota capensis subsp. /unaris (Emery); Bolton 1995:

228; Bharti et al. 2016: 28; Bharti et al. 2017: 42; erro-

neously treated as subspecies of capensis.

Worker measurements. HL 0.56—0.63; HW 0.51-

0.57; EL 0.17-0.19; SL 0.62-0.66; PnW 0.34—0.42:

ML 0.74—0.86; PFL 0.48—0.53; PFW 0.12—0.14 mm. In-

dices: CI 89-91; SI 111-123; REL 29-31 (n=9),.

Description. Head subquadrate; slightly longer than

wide, wider posteriorly than in front; lateral and poste-

rior margins convex, posterolateral corners rounded; cly-

peus subcarinate in the middle; anterior clypeal margin

complete and convex; eyes broadly oval, weakly convex,

placed at the middle-line of head, covering one-third of

lateral cephalic margin; three small ocelli present; anten-

nal scape surpassing posterior head margin by about one-

third its length.

In lateral view promesonotum convex, metanotum low,

almost straight; mesometanotum demarcated; metanotal

area long and constricted; propodeum armed with a pair

of posteriorly diverging thick blunt spines; propodeal de-

clivity steep.

Petiole upright, with angular sides, dorsally emargin-

ate, armed with a pair of almost straight spines pointing

upward.

Head and mesosoma feebly shiny to subopaque; gas-

ter polished, smooth and shiny; overall head and meso-

somal dorsum weakly microreticulate; propleuron shiny;

mesonotum and lateral mesosoma longitudinally striate;

declivous face transversally striate.

Body covered with short, erect, abundant setae on head

and gaster, sparser on mesosoma; pubescence very fine

and sparse, most visible on head; almost absent on me-

sosoma and gaster; antennal funiculus with sparse ap-

pressed to decumbent pubescence.

Black; antenna, mandible and tarsi light brown.

Distribution and habitat. India (Himachal Pradesh,

Jammu and Kashmir), Nepal, Pakistan, Sri Lanka (Thapa

2000; Bharti et al. 2016; Rasheed et al. 2019; Dias et al.

2020). This species seems to be widespread across India,

although infrequent in the collections. The specimens

were collected both in forest and non-forest habitats,

mainly by hand collecting from the dry soil and honey

baits.

Remarks. This small to medium-sized species is sim-

ilar to many workers of the medium sized L. mayri but

the erect setae on the body are distinctly sparse, whereas

the latter species is covered with abundant erect setae and

very hairy overall.

Material examined. India: Himachal Pradesh: Bari,

31.6591° N, 76.5000° E, 520 m, 3 9 9, 6.v1i.2009; Ghatti,

31.9300° N, 75.9302° E, 425 m, 3 99, 12.x.2008; Guga,

©ZFMK

Review of the ant genus Lepisiota from India 235

31.6864° N, 76.1898° E, 600 m, 2 9°, 22.x.2008; Jas-

sur, 32.2824° N, 75.8496° E, 520 m, 6 99, 6.vi.2009;

Kotla, 31.8821° N, 75.9963° E, 500 m, 5 9 9, 22.x.2008;

Palampur, 32.1109° N, 76.5430° E, 1200 m, 4 99,

18.vi.2010; Nagabari, 32.3004° N, 75.8901° E, 420 m,

3 OQ, 18.vi.2009; Terrace, 31.9234° N, 75.9294° E,

420 m, 4 QQ, 12.x.2009. Punjab: Dunera, 32.4443° N,

75.8900° E, 520 m, 3 99, 24.vii.2010, Aijaz A. Wach-

koo leg.

Lepisiota mayri sp. n. (Figs 19-21)

urn: lsid:zoobank. org: act: FIBBB1FB-4249-4BCB-9FCF-5128382CF9AC

Type material. Holotype worker. India, Himach-

al Pradesh, Andretta, 32.0744° N, 76.5856° E, 940 m,

21.vi.2010. Paratypes: 12 workers, same data as for

holotype; 7 workers, Himachal Pradesh, Baijnath,

32.0527° N, 76.6500° E, 1125 m, 17.vi.2010; 6 work-

ers, Himachal Pradesh, Kotla, 31.8821° N, 75.9963° E,

500 m, 13.x.2008; 2 workers, Himachal Pradesh, Nahan,

760 m, 30.5596° N, 77.2960° E, 27.viii.2009; 1 work-

er, Himachal Pradesh, Renuka, 30.6083° N, 77.4615° E,

600 m, 8.v.2009; 5 workers, Himachal Pradesh, Terrace,

31.9234° N, 75.9294° E, 420 m, 24.v.2009; 3 workers,

Uttarakhand, Forest Research Institute, 30.3416° N,

77.9903° E, 640 m, 2.1x.2009; 8 workers, Uttarakhand,

Rajaji Forest Area, 30.2483° N, 77.9878° E, 660 m,

21.v.2010; 1 worker, Uttarakhand, Selaqui, 30.3720° N,

77 .8605° E, 670 m, 24.v.2010, A1jaz A. Wachkoo leg.

Worker measurements. HL 0.65—0.73; HW 0.60—

0.69; EL 0.20-0.23; SL 0.68-0.77; PnW 0.42-0.50;

ML 0.92-1.04; PFL 0.57-0.64; PFW 0.14—0.17 mm. In-

dices: CI 91-95; SI 111-116; REL 29-31 (n= 15).

Description. Head subquadrate; slightly longer than

wide, wider posteriorly than in front; lateral and posteri-

or margins convex, with rounded posterolateral corners;

clypeus carinate in the middle; anterior clypeal margin

complete and convex; eyes broadly oval, weakly convex,

placed at the middle-line of head, covering about one-

third of lateral cephalic margin; three small ocelli pres-

ent; antennal scape surpassing posterior head margin by

about one-third its length.

In lateral view promesonotum convex, metanotum low,

almost straight; mesometanotum demarcated; metanotal

area distinct; mesometanotum constricted; propodeum

armed with a pair of posteriorly diverging thick blunt

spines; propodeal declivity steep.

Petiole upright, with angular sides, dorsally emargin-

ate, armed with a pair of nearly straight spines pointing

upward.

Overall body shiny; head weakly microreticulate to

smooth without sculpture; pronotum, metanotum and

propodeal dorsum coarsely microreticulate; mesonotum

weakly microreticulate, mesometanotal suture cross-

ribbed; propleuron sometimes weakly longitudinally stri-

ate; mesometapleuron coarsely longitudinally striate.

Bonn zoological Bulletin 70 (2): 227-245

Body abundantly covered with short, erect setae; scape

and legs with dense suberect pilosity; antennal funiculus

with dense subdecumbent to suberect pubescence.

Figs 19-21. Holotype worker; Lepisiota mayri sp. n. 19. Head,

frontal view. 20. Body, lateral view. 21. Body, dorsal view.

©ZFMK

236 Aijaz Ahmad Wachkoo et al.

Black; scape and tarsi yellow brown.

Distribution and habitat. India (as L. capensis;

Arunachal Pradesh, Assam, Bihar, Goa, Haryana, Him-

achal Pradesh, Jammu and Kashmur, Jharkhand, Karnata-

ka, Kerala, Madhya Pradesh, Maharashtra, Manipur, Me-

ghalaya, Mizoram, Sikkim, Uttarakhand, West Bengal)

(Bharti et al. 2016; Dad et al. 2019). This species appears

to be widespread on the Indian subcontinent typically re-

ported as L. capensis; specimens were collected by hand,

beating vegetation and honey baiting.

Etymology. The species is named in honor of Gustav

Mayr, author of L. capensis to which it has been a hom-

onym.

Remarks. The species is similar to L. modesta with

which it shares its uniform black color and a medi-

um-sized body. The distinguishing features include a

combination of fine microreticulate sculpture on head

and mesosoma, prominent propodeal spines and abun-

dant erect setae on the body in L. mayri, versus a smooth

and shiny body, reduced propodeal spines, and sparse

erect setae in L. modesta.

Lepisiota modesta (Forel, 1894) (Figs 22—24)

Acantholepis modesta Forel, 1894: 412. Syntype work-

ers, Mussoorie, Uttarakhand, India [MHNG]. [Images

of CASENT0909892 Syntype worker examined].

Lepisiota modesta (Forel); Bolton 1995: 228; first com-

bination in Lepisiota.

Worker measurements. HL 0.60—0.67; HW 0.52-

0.63; EL 0.16—0.20; SL 0.62-0.70; PnW 0.35-0.44:

ML 0.78—-0.92; PFL 0.48—0.57; PFW 0.12—-0.15 mm. In-

dices: CI 85—93; SI 110-125; REL 26-29 (n = 12).

Description. Head subquadrate; longer than wide,

wider posteriorly than in front; lateral margins convex,

posterior margin shallowly concave, with rounded pos-

terolateral corners; clypeus carinate in the middle; anteri-

or clypeal margin complete and convex; eyes oval, weak-

ly convex, placed at the middle-line of head, covering

about one-third of lateral cephalic margin; three small

ocelli present; antennal scape surpassing posterior head

margin by about one-third its length.

In lateral view promesonotum convex, metanotum low,

almost straight; mesometanotum demarcated; metanotal

area distinct, mesometanotum constricted; propodeal

spines reduced to tubercles; propodeal declivity steep.

Petiole upright with angular sides, dorsally emarginate,

armed with spines pointing upward.

Overall body shiny; mesometanotal suture cross-

ribbed; mesometapleuron coarsely longitudinally striate

weakly so, on lateropropodeum; propodeal declivity

transversally striate.

Body abundantly covered with erect setae; pubescence

on body uniformly very fine and sparse; antennal funicu-

lus with subdecumbent to suberect pubescence.

Color black; scape, mandible and tarsi yellow brown.

Bonn zoological Bulletin 70 (2): 227-245

Distribution and habitat. India (Himachal Pradesh,

Jammu and Kashmir, Punjab, Uttarakhand) (Bharti et al.

2016; Wachkoo et al. 2020). Although this species is col-

lected infrequently it seems widespread across India. It

24 0.5 mm

Figs 22-24. Worker; Lepisiota modesta (Forel, 1894). 22. Head,

frontal view. 23. Body, lateral view. 24. Body, dorsal view.

©ZFMK

Review of the ant genus Lepisiota from India DSi

seems to have a preference for relatively higher altitudes

above 1200 m. Most of the specimens were collected un-

der stones and from honey baits.

Remarks. The medium-sized workers can be dis-

tinguished from the somewhat similar small to medi-

um-sized L. annandalei by a combination of abundant

erect setae on body, a transversally striate propodeal de-

clivity and a bispinose petiolar dorsum; L. annandalei

has sparse erect setae on the body, a smooth and shiny

propodeal declivity and an emarginate petiolar dorsum.

Material examined. India: Himachal Pradesh: Lwasa,

30.7394° N, 77.1528° E, 1200 m, 17 99, 27.viii.2009.

Jammu and Kashmir: Baderwah, 32.9832°N, 75.7097° E,

1700 m, 18 QQ, 30.vi.2009, 30 29, 1.vii.2009. Utta-

rakhand: Mussoorie, 30.4606° N, 78.0521° E, 1820 m,

37 PQ, 9.vili.2009, Aijaz A. Wachkoo leg.

Lepisiota opaca (Forel, 1892) (Figs 25—27)

Acantholepis opaca Forel, 1892: 43. Syntype workers,

Kanara, Karnataka, India [MHNG, MSNG]. [Imag-

es of CASENT0909893, CASENT0905158 Syntype

workers examined].

Acantholepis opaca Forel; Bingham 1903: 318.

Lepisiota opaca (Forel); Xu 1994: 235; first combination

in Lepisiota.

Worker measurements. HL 0.63—0.70; HW 0.58

0.65; EL 0.19-0.22; SL 0.64—-0.76; PnW 0.42-0.47;

ML 0.79-1.00; PFL 0.53—0.61; PFW 0.14—0.16 mm. In-

dices: CI 91-93; SI 111-120; REL 29-32 (n = 8).

Description. Head subquadrate; slightly longer than

wide, wider posteriorly than in front; lateral margins con-

vex, posterior margin nearly transverse to convex, with

rounded posterolateral corners; clypeus carinate in the

middle; anterior clypeal margin complete and convex;

eyes oval, weakly convex, placed at the middle-line of

head, covering about one-third of lateral cephalic mar-

gin; three small ocelli present; antennal scape surpassing

posterior head margin by about one-third its length.

In lateral view promesonotum convex, metanotum low,

almost straight; pronotum flat above; mesometanotum

demarcated; metanotal area distinct; mesometanotum

constricted; propodeum armed with a pair of posteriorly

diverging, upward directed sharp spines; propodeal de-

clivity steep.

Petiole upright, with angular sides, dorsally emargin-

ate, armed with a pair of almost straight spines pointing

upward.

Overall body smooth and shiny; malar space usually

coarsely microreticulate, remainder of head mostly shiny

with effaced sculpture; pronotum smooth and shiny, re-

mainder of mesosomal dorsal feebly microreticulate:

propleuron smooth and shiny, rest of lateral mesosoma

longitudinally striate; mesometanotal suture cross-

Bonn zoological Bulletin 70 (2): 227—245

ribbed; propodeal declivity feebly transversally striate:

sculpturing almost entirely absent in some specimens.

Figs 25-27. Worker; Lepisiota opaca (Forel, 1892). 25. Head,

frontal view. 26. Body, lateral view. 27. Body, dorsal view.

©ZFMK

238 Aijaz Ahmad Wachkoo et al.

Body abundantly covered with erect setae; pubescence

very fine and sparse, least on mesosoma; antennal funic-

ulus with subdecumbent to suberect pubescence.

Head reddish-brown, mesosoma and petiole red-

dish-yellow, gaster purplish-black, some specimens with

a light reddish-yellow patch anteriorly on first gastral

tergite; antenna, mandible and legs reddish-brown.

Distribution and habitat. China, India (Arunachal

Pradesh, Assam, Himachal Pradesh, Goa, Jammu and

Kashmir, Karnataka, Kerala, Madhya Pradesh, Maha-

rashtra, Sikkim, West Bengal), Sri Lanka (Guénard &

Dunn 2012; Bharti et al. 2016; Sheikh et al. 2019; Dias

et al. 2020). This is one of the widespread species of its

genus and relatively common in India. The specimens

were mainly collected by beating vegetation.

Remarks. Lepisiota opaca is a medium-sized ant. It

superficially resembles L. fergusoni and L. pulchella

but can be separated from both by a smooth and shiny

body, whereas L. fergusoni and L. pulchella have dull and

coarsely sculptured bodies.

Material examined. India: Himachal Pradesh:

Andretta, 32.0744° N, 76.5856° E, 940 m, 16 99,

11.v1.2010; Chanaur, 32.0545° N, 75.6503° E, 600 m,

5 9Q, 12.vi.2009, 5 9°, 20.x.2008; Ghatti, 31.9300° N,

75.9302° E, 425 m, 7 2, 12.x.2008, 6 99, 27.1x.2009,

3 QQ, 28.ix.2009; Guraldhar, 31.6670° N, 76.4684° E,

660 m, 1 Q, 2.v1.2009, 2 29, 10.vi.2009; Khatiar,

32.0057° N, 75.9388° E, 450 m, 7 99, 18.x.2008, 1 8,

3.vi.2009; Kotla, 31.8821° N, 75.9963° E, 500 m, 8 99,

13.x.2008, 4 99, 22.x.2008, 5 99, 28.v.2009; Lwasa,

30.7394° N, 77.1528° E, 1200 m, 12 99, 7.viii.2009;

Nahan, 30.5596° N, 77.2960° E, 760 m, 13 Q29,

20.vill.2009, 9 QO, 27.vili.2009; Siholi, 31.9456° N,

75.9949° E, 560 m, 1 9, 4.vi.2009; Terrace, 31.9234°N,

75.9294° E, 420 m, 36 29, 11.x.2008, 1 2, 21.x.2008,

4 29, 26.v.2009, 1 9, 13.vi.2009, 11 99, 24.1x.2009,

10 929, 25.1x.2009. Jammu and Kashmir: Manda,

32.7496° N, 74.8673° E, 500 m, 7 @Q, 15.vii.2009;

Mansar, 32.6979° N, 75.1489° E, 690 m, | 9, 12.vi1.2009,

LO" Seyi 2009: = Samba,» 32°5537°" N.S 1S 17°" F,

390 m, 2 YY, 11.vii.2009; Surinsar, 32.7009° N,

75.1512° E, 700 m, 2 99, 14.vii.2009. Punjab: Dharam-

pur, 31.8420° N, 75.9132° E, 450 m, 2 99, 14.x.2008;

Dunera, 32.4443° N, 75.8900° E, 520 m, 1 9, 23.vi.2009;

Thein Dam, 32.4426° N, 75.7305° E, 1 9, 24.vi.2009.

Uttarakhand: Dakpathar, 30.4966° N, 77.8004° E, 750 m,

1 Q, 20.viii.2009; Forest Research Institute, 30.3416° N,

77.9903° E, 640 m, 1 9, 17.vi11.2009; Mussoorie,

30.4606° N, 78.0521° E, 1820 m, 2 29, 13.vii.2009; Ra-

jaji Forest Area, 30.2483° N, 77.9878° E, 660 m, 39 99,

6.vill.2009, 4 99, 10.vili.2009, 1 9, 13.vill.2009; Sela-

qui, 30.3720° N, 77.8605° E, 670 m, 1 &, 7.vili.2009,

2 29, 24.v.2010, Aijaz A. Wachkoo leg.

Bonn zoological Bulletin 70 (2): 227—245

Lepisiota pulchella (Forel, 1892) stat. rev. (Figs 28-30)

Acantholepis opaca t. pulchella Forel, 1892: 43. Syntype

workers, Pune [Poona], Maharashtra, India [MHNG].

[Images of CASENT0909894 Syntype worker exam-

ined].

Acantholepis pulchella Forel, Dalla Torre 1893: 172;

Bingham 1903: 318; Wu & Wang 1995: 129; Zhou

2001: 168; status as species.

Acantholepis opaca subsp. pulchella Forel; Emery

1893b: 172; Forel 1894: 414; Forel 1895: 458; Emery

1925: 27; Chapman & Capco 1951: 210; Bolton 1995:

228; Guénard & Dunn 2012: 34; Bharti et al. 2016: 28;

Bharti et al. 2017: 42; subspecies of opaca.

Lepisiota opaca subsp. pulchella (Forel); Bolton 1995:

228; first combination in Lepisiota.

Worker measurements. HL 0.56—0.64; HW 0.51-

0.59; EL 0.17-0.20; SL 0.60—0.68; PnW 0.35-0.42:

ML 0.77-0.87; PFL 0.45—0.53; PFW 0.12—0.14 mm. In-

dices: CI 90-94; SI 111-119; REL 30-31 (n= 15).

Description. Head subquadrate; slightly longer than

wide, wider posteriorly than in front; lateral and poste-

rior margins convex, posterolateral corners rounded; cly-

peus subcarinate in the middle; anterior clypeal margin

complete and convex; eyes oval, weakly convex, placed

at the middle-line of head, covering about one-third of

lateral cephalic margin; three small ocelli present; anten-

nal scape surpassing posterior head margin by about one-

third its length.

In lateral view promesonotum convex, metanotum low,

feebly concave; mesometanotum demarcated; metanotal

area distinct; mesometanotum constricted; propodeum

armed with a pair of posteriorly diverging, upward di-

rected sharp spines; propodeal declivity steep.

Petiole upright, with angular sides, dorsally emargin-

ate, armed with a pair of divergent spines pointing up-

ward and outward.

Head and mesosomal dorsum opaque; gaster polished,

smooth and shining; head and mesosomal dorsum dis-

tinctly reticulate-punctate; propleuron relatively smooth

and shiny, remainder of lateral mesosoma longitudinally

striate; mesometanotal suture cross-ribbed; propodeal

declivity transversally striate.

Body abundantly covered with erect setae; pubescence

very fine and sparse, least on mesosoma; antennal funic-

ulus with subdecumbent to suberect pubescence.

Head brown, mesosoma reddish-brown to dark brown;

antenna, petiole and legs light to dark brown; gaster

black with a light reddish-yellow patch anteriorly on first

gastral tergite.

Distribution and habitat. China, India (Karnataka,

Kerala, Maharashtra) (Forel 1894; Guénard & Dunn

2012; Dad et al. 2019). This species seems to be distrib-

uted in the Western Ghats and other parts of central In-

dia. Records from Arunachal Pradesh, Assam, Himachal

Pradesh, Jammu and Kashmir, Punjab, Sikkim, Uttara-

©ZFMK

Review of the ant genus Lepisiota from India 239

khand and West Bengal (Bharti et al. 2016) seem to be a

misidentification of L. opaca and are excluded until fur-

ther evaluation. The workers were mainly collected from

honey baits and by beating vegetation.

Remarks. This medium-sized ant originally described

by Forel (1892) as an infraspecific taxon opaca r. pul-

chella was raised to species rank by Bingham (1903).

Emery (1925) without any justification reverted it back to

subspecies. It is a valid species, resembling more closely

L. fergusoni than L. opaca and 1s therefore revived here

Figs 28-30. Worker; Lepisiota pulchella (Forel, 1892) stat. rev.

28. Head, frontal view. 29. Body, lateral view. 30. Body, dorsal

view.

Bonn zoological Bulletin 70 (2): 227—245

to species level. The main characters that enable distin-

guishing it from LZ. opaca are a combination of a dull and

opaque body with a distinctly reticulate-punctate head

and mesosoma. L. opaca is smooth and shiny with super-

ficial reticulate sculpture.

Material examined. India: Karnataka: Gundlupet,

11.8132° N, 76.6858° E, 800 m, 8 99, 27.1x.2010. Ker-

ala: Salim Ali Bird Sanctuary, 10.0918° N, 76.7420° E,

118 m, 6 QQ, 10.x.2011, Shahid A. Akbar leg.

Lepisiota rothneyi (Forel, 1894) (Figs 31-33)

Plagiolepis rothneyi Forel, 1894: 415. Syntype workers,

Belgaum, Karnataka; Barrackpore [Barrakpore], West

Bengal, India [MHNG]. [Images of CASENT0909865,

CASENT0909866 Syntype workers examined].

Acantholepis rothneyi Forel; Santschi 1926: 15; first

combination in Acantholepis.

Lepisiota rothneyi Forel; Bolton 1995: 228; first combi-

nation in Lepisiota.

Worker measurements. HL 0.74—1.04; HW 0.71-

1.03; EL 0.20—-0.28; SL 0.72-0.88; PnW 0.48-0.63;

ML 0.88-1.09; PFL 0.61—0.81; PFW 0.18—0.25 mm. In-

dices: CI 95—98; SI 98-104; REL 26-27 (n = 13).

Description. Head subquadrate; about as long as wide,

wider posteriorly than in front; lateral margins convex,

posterior margin gently convex, posterolateral corners

rounded; clypeus subcarinate in the middle; anterior

clypeal margin complete and convex; eyes relatively

small, broadly oval, weakly convex, placed at the mid-

dle-line of head, covering three-tenths of lateral cephalic

margin; three small ocelli present; antennal scape sur-

passing posterior head margin by about one-fourth its

length.

Mesosoma constricted in the middle, in lateral view

promesonotum dome like, convex; metanotum distinctly

lower than promesonotum but almost as high as propo-

deum; mesometanotum demarcated; metanotal area short

but distinct; propodeum unarmed without a pair of teeth

or spines; propodeal declivity steep.

Petiole upright, with smoothly curved sides, dorsally

rounded, without a pair of teeth or spines.

Body overall smooth and shiny. Setae restricted to cly-

peus, gastral venter and segmental margins; a few setae

near lateral ocelli usually present; body covered with

Sparse pubescence; antennal funiculus with appressed to

decumbent pubescence.

Color brown to black; antenna, mandible and tarsi light

brown.

Distribution and habitat. Bangladesh, China, India

(Karnataka, Kerala, Orissa, Tamil Nadu, Uttarakhand,

West Bengal), Myanmar, Vietnam (Wheeler 1927; Han-

nan 2003; Zryanin 2011, Guénard & Dunn 2012; Bharti

et al. 2016). This species is widespread in India however,

it appears to be a forest ant, restricted to relatively un-

©ZFMK

240

disturbed areas. The workers were mostly hand collected

from tree trunks and by beating vegetation.

Figs 31-33. Worker; Lepisiota rothneyi (Forel, 1894). 31. Head,

frontal view. 32. Body, lateral view. 33. Body, dorsal view.

Bonn zoological Bulletin 70 (2): 227—245

Aijaz Ahmad Wachkoo et al.

Remarks. The medium-sized ant closely resembles

L. wroughtonii, but can be separated from it by having

a sparsely pubescent and shiny body, lacking erect setae

on the mesosomal dorsum. In contrast, L. wroughtonii is

more pubescent with few short setae on the pronotum.

Material examined. India: Uttarakhand: Forest Re-

search Institute, 30.3416° N, 77.9903° E, 640 m, 6 99,

Wwv.2009.01 2..13:v:2009, 1°). 30:vil.2009, 2 2.0"

26.v.2010; Rajaji Forest Area, 660 m, 30.2483° N,

77.9878° E, 1 Q, 5.viii.2009, 2 22, 6.vili.2009, 4 99,

11.viti.2009, 1 9, 6.1x.2009; Selaqui, 30.3720° N,

77.8605° E, 670 m, 4 29, 7.vili.2009, Aijaz A. Wachkoo

leg.

Lepisiota sericea (Forel, 1892) (Figs 34—36)

Acantholepis frauenfeldi var. sericea Forel, 1892: 41.

Syntype workers, Mussoorie, Uttarakhand, India

[MHNG]. [Images of CASENT0909885 Syntype

worker examined].

Acantholepis sericea Forel; Pisarski 1967: 408; status as

species.

Lepisiota sericea (Forel); Bolton 1995: 228; first combi-

nation in Lepisiota.

Worker measurements. HL 0.66—0.84; HW 0.59-

0.74; EL 0.20-0.25; SL 0.99-1.08; PnW 0.36-0.52:

ML 1.02—1.27; PFL 0.77-0.97; PFW 0.14—0.18 mm. In-

dices: CI 85—90; SI 146-152; REL 29-30 (n = 6).

Description. Head subrectangular; longer than wide,

slightly narrowed anteriorly; lateral and posterior mar-

gins convex, posterolateral corners rounded; clypeus me-

dially subcarinate; anterior clypeal margin complete and

convex; eyes subglobulose, convex, projecting beyond

cephalic lateral margins, covering one-third of lateral

cephalic margin, placed at posterior half of head: three

ocelli present; antennal scape surpassing posterior head

margin by about half its length.

In lateral view pronotum convex, mesometanotum

strongly constricted, lower than pronotum and propo-

deum, giving mesosoma a dumbbell shape; mesometa-

notum demarcated; metanotal area distinct; propodeum

armed with a pair of teeth diverging posteriorly; propo-

deal declivity steep.

Petiole upright, with smoothly curved sides, dorsally

rounded and narrow, without teeth or spines.

Body dull, overall feebly microreticulate, gastral sculp-

turing even feebler.

Body covered with sparse erect setae on head, pro-

notum and gaster; setae on gaster usually restricted to

venter and segmental margins; pubescence very fine and

sparse, most visible on head and gaster; almost absent on

mesosoma; antennal funiculus with appressed to decum-

bent pubescence.

Color uniformly dark brown to black; antenna, mandi-

ble and tarsi brown.

©ZFMK

Review of the ant genus Lepisiota from India 241

Distribution and habitat. Afghanistan, India (Him-

achal Pradesh, Jammu and Kashmir, Maharashtra, Ut-

36 0.5 mm

Figs 34-36. Worker; Lepisiota sericea (Forel, 1892). 34. Head,

frontal view. 35. Body, lateral view. 36. Body, dorsal view.

Bonn zoological Bulletin 70 (2): 227—245

tar Pradesh, Uttarakhand), Iran, Pakistan, Turkmenistan

(Kuznetsov-Ugamsky 1929; Pisarski 1967; Ghahari et al.

2011; Bharti et al. 2016; Rasheed et al. 2020). This spe-

cies seems widespread in India. Specimens were hand

collected, under stones, from tree trunks and the ground.

Remarks. Lepisiota sericea is a medium to large-sized

ant which can be distinguished from the very similar spe-

cies L. integra by a combination of a dark brown body,

smoothly curved sides of the petiole and a narrow, round-

ed petiolar dorsum. L. integra has a reddish-brown body,

angular sides of the petiole and an emarginate petiolar

dorsum with teeth-like apical corners.

Material examined. Himachal Pradesh: Andretta,

32.0744° N, 76.5856° E, 940 m, 9 99, 21.vi.2010; Bai-

jnath, 32.0527° N, 76.6500° E, 1125 m, 6 29, 17.vi.2010;

Lwasa, 30.7394° N, 77.1528° E, 1200 m, 6 99,

27.vill.2009; Palampur, 32.1109° N, 76.5430° E,1200 m,

7 QQ, 18.vi.2010. Jammu and Kashmir: Surinsar,

32.7009° N, 75.1512° E, 700 m, 10 99, 14.vil.2009. Ut-

tarakhand: Mussoorie, 30.4606° N, 78.0521° E, 1820 m,

7 OQ, 9.viil.2009, Aijaz A. Wachkoo leg.

Lepisiota wroughtonii (Forel, 1902) stat. rev. (Figs 37-

39)

Plagiolepis rothneyi r. wroughtonii Forel, 1902a: 292.

Syntype workers, Nilgiris, Tamil Nadu, India [MHNG].

[Images of CASENT0909869 Syntype worker exam-

ined].

Plagiolepis wroughtonii Forel; Bingham 1903: 321; sta-

tus as species.

Plagiolepis rothneyi subsp. wroughtonii Forel; Emery

1925: 23; subspecies of rothneyi.

Lepisiota rothneyi subsp. wroughtonii (Forel); Bolton

1995: 229; first combination in Lepisiota.

Worker measurements. HL 0.60—0.75; HW 0.54—

0.69; EL 0.16-0.21; SL 0.60—-0.73; PnW 0.36-0.48:

ML 0.76—0.96; PFL 0.49-0.62; PFW 0.16—0.20 mm. In-

dices: CI 82—93; SI 103-119; REL 26-28 (n = 13).

Description. Head subquadrate; longer than wide,

wider posteriorly than in front; lateral margins convex,

posterior margin gently convex, posterolateral corners

rounded; clypeus subcarinate in the middle; anterior

clypeal margin complete and convex; eyes relatively

small, broadly oval, weakly convex, placed at the mid-

dle line of head, covering three-tenths of lateral cephalic

margin; three small ocelli present; antennal scape sur-

passing posterior head margin by about one-fourth its

length.

Mesosoma constricted in the middle, in lateral view

promesonotum dome like, convex; metanotum distinctly

lower than promesonotum but almost as high as propo-

deum; mesometanotum demarcated; metanotal area short

but distinct; propodeum unarmed without pair of teeth or

spines; propodeal declivity steep.

©ZFMK

242 Aijaz Ahmad Wachkoo et al.

Petiole upright, with smoothly curved sides, dorsally

rounded, without a pair of teeth or spines.

Body overall dull covered with relatively dense pu-

bescence. Setae restricted to clypeus, gastral venter and

‘

4 a

‘ } y

3 -

Figs 37-39. Worker, Lepisiota wroughtonii (Forel, 1902) stat.

rev. 37. Head, frontal view. 38. Body, lateral view. 39. Body,

dorsal view.

Bonn zoological Bulletin 70 (2): 227—245

segmental margins; few short erect setae on pronotum

and near lateral ocelli usually present; pubescence rela-

tively dense, more so on gaster; antennal funiculus with

appressed to decumbent pubescence.

Color black; antenna, mandible and tarsi light brown.

Distribution and habitat. China, India (Himachal

Pradesh, Kerala, Tamil Nadu, Uttarakhand, West Ben-

gal), Sri Lanka (Guénard & Dunn 2012; Bharti et al.

2016; Dias et al. 2020). This species appears widespread

in India and was mainly hand collected from tree trunks,

by beating vegetation, from honey baits and occasionally

in pitfall traps.

Remarks. This medium-sized ant was originally de-

scribed by Forel (1892) as an infraspecific taxon roth-

neyi r. wroughtonii and later elevated to species rank by

Bingham (1903). Emery (1925) reverted it back to sub-

species without any justification. Based on our examina-

tion, we consider it as a valid species and elevate it to

species rank here. The main characters that will enable

it to be distinguished from L. rothneyi, the species with

which it is most likely to be confused, are a combination

of a densely pubescent and dull body with few erect se-

tae on the pronotum, whereas L. rothneyi has a sparsely

pubescent and shiny body, lacking erect setae on the me-

sosomal dorsum.

Material examined. India: Himachal Pradesh: Khatiar,

32.0057° N, 75.9388° E, 450 m, 3 99, 18.x.2008; Poanta

Sahib, 30.4384° N, 77.6239° E, 420 m, 4 99, 11.v.2009.

Uttarakhand: Assan Barrage, 30.4417° N, 77.6754° E,

750 m, 3 QQ, 21.vii.2009; Forest Research Institute,

30.3416° N, 77.9903° E, 640 m, 3 99, 1.x.2008, 2 99,

12.v.2009, 2 2, 30.vii.2009, 2 29, 20.v.2010, 4 99,

26.v.2010; Rajaji Forest Area, 30.2483° N, 77.9878° E,

660 m, 4 99, 6.vili.2009, 4 OQ, 10.vili.2009, 3 99,

25.v.2010; Ranger’s College, 660 m, 30.3225° N,

78.0445° E, 8 29, 25.v.2010, 1 9, 27.v.2010; Selaqui,

30.3720° N, 77.8605° E, 670 m, 6 9 9, 24.v.2010, 10 99,

7.vill.2009, 1 2, 5.1x.2010, Aijaz A. Wachkoo leg.

Species excluded from the Indian fauna

The following species are excluded from the Indian Lepi-

siota fauna and are considered as doubtful records and/or

misidentifications.

Lepisiota capensis (Mayr, 1862)

The nominal form of L. capensis (Mayr, 1862) does not

occur naturally in India, despite its implied presence by

the use of the name L. capensis reported by Bingham

(1903). The species referred to in the latter paper, with

abundant yellowish erect setae is clearly not L. capen-

sis, which has sparse dark (brown or black) erect setae

(see Mayr 1862, 1865; Arnold 1920) and is distributed

in southern Africa (not from the Himalayas through to

Northeastern Africa).

Bingham’s (1903) report was apparently based on an

erroneous reporting of the presence of L. capensis with

©ZFMK

Review of the ant genus Lepisiota from India

whitish setae in India and Sri Lanka (then Ceylon) by

Forel (1892, 1894), and the attribution of dense pilosity

to L. capensis served only to perpetuate this error until

today (Peter Hawkes & F. Hita Garcia, pers. comm. ).

The Indian material currently and in the past attributed

to L. capensis in fact represents a very different species,

and is accordingly described as a new species under the

name L. mayri to minimize confusion by eliminating fu-

ture use of the name L. capensis associated with Indian

species.

Lepisiota frauenfeldi (Mayr, 1855)

Forel (1885) first considered that L. frauenfeldi was Pale-

arctic in distribution and attributed Indian specimens

from Kolkata (then Calcutta), West Bengal to L. bipar-

tita. But, in his subsequent publications (Forel 1886,

1894), he argued that he had confused L. bipartita with

L. frauenfeldi and accordingly treated specimens from

Kolkata and Barrackpore as L. frauenfeldi. However,

he also mentioned its rare occurrence in India. Bingham

(1903), Rothney (1903), and subsequent Indian authors

until today continued with Forel’s (1886, 1894) unver-

ified reporting without further clarification (see Bharti

et al. 2016).

Lepisiota rothneyi watsonii (Forel, 1894)

Bharti et al. (2016) include distribution of this species

in India based on Wheeler (1921). However, Wheeler

(1921) does not mention its distribution in India and it is

therefore excluded from the known Indian fauna.

Lepisiota simplex (Forel, 1892)

This is an African species that seems unlikely to be found

in India. Forel (1894) first reported this species from

Orissa based on of its superficial resemblance to the type

material from Somalia. Bingham (1903) and subsequent

Indian authors followed this view without any further ex-

amination of Indian specimens (see Bharti et al. 2016).

Based on our examination, Indian material attributed to

L. simplex belong mostly to L. annandalei and can be dis-

tinguished by few setae restricted to pronotum, whereas

former has several pairs of setae scattered on the meso-

soma including the propodeum.

Key to Indian species of Lepisiota (workers)

1. Antennal scape long, surpassing posterior margin of

head by half its length or more (Figs 5, 11, 35) ..... 2

— Antennal scape shorter, surpassing posterior margin

of head by a third of its length or less (Figs 2, 14,

23)

2. Bicolored; head and gaster dark brown to black,

mesosoma reddish brown (Fig. 5) ........0..ccceee

Re ane ORNS Ren 2.2 cae a L. bipartita (Smith)

— Uniform reddish-brown or black-brown (Figs 11,

35)

Bonn zoological Bulletin 70 (2): 227—245

10.

Tek,

12:

243

Reddish-brown; petiolar scale with distinctly angular

sides, dorsally emarginate with teeth-like apical

corners (Figs 11-12) ......L. integra (Forel) stat. nov.

Black-brown; petiolar scale with smoothly curved

sides, dorsally rounded and narrow, without teeth

(F18S/35=36 piso te weet L. sericea (Forel)

Bicolored; mesosoma paler than gaster, mainly or

entirely reddish (Figs 8, 26, 29)

Whole body uniformly dark brown to black (Figs 2,

14, 38 )

All parts of the body shining with superficial

microreticulate sculpture at most (Figs 26—27) ........

eh rh Ee seco Roorena a cally Fay a «han L. opaca (Forel)

Body sculpture coarse, general appearance opaque

(Figs 9, 30)

Head reticulate-striate; propodeal spines blunt,

directed backward; petiole dorsally emarginate

CEE S/O) m es. ..eee, Se ee re Et, L. fergusoni (Forel)

Head reticulate-punctate; propodeal spines pointed,

directed upward; petiole dorsally bispinose (Figs 28—

SO) bs Berets has aie L. pulchella (Forel) stat. rev.

Propodeal spines indistinct; gastral pilosity restricted

to few pairs of black setae on the posterior margin of

tergites (Figs 32, 38)

Propodeal spines well-developed in the form of two

broadly-based blunt tubercles, teeth or spines; whole

gastral dorsum covered with abundant pale setae

(Figs 2, 14, 17)

Pronotum without any setae; body sparsely

pubescent, shiny (Fig. 32) .......... L. rothneyi (Forel)

Pronotum with few short setae; body fairly pubescent,

opaque (Fig. 38) ....L. wroughtonii (Forel) stat. rev.

Body abundantly covered with long, erect white

Serer Che: Neb \eeere ss. cemres zoentonec A wna L. layla sp. n.

Body covered with usual short erect setae (Figs 17,

DODO \eth ctectn oth asain ee hae teste eect ae 10

Petiole dorsally emarginate without distinct teeth

Or spines; mesosomal setae (if any) restricted to

pronotum (Figs 2-3) ....... L. annandalei (Mukerjee)

Petiole dorsally distinctly bispinose; setae present

across entire mesosomal dorsum (Figs 17, 20, 23) ..

Head and mesosoma smooth and shining; propodeal

spines in the form of two broadly-based blunt

tubercles (Figs 22—23) .........000.... L. modesta (Forel)

Head and mesosoma finely microreticulate and dull;

propodeal spines pointed (Figs 16, 20) .......00..0... 12

Antennal scape with sparse appressed to decumbent

pubescence; mesosomal setae sparse; smaller species

(HW 0.51-0.57) (Figs 16-18) ....L. lunaris (Emery)

Antennal scape with dense subdecumbent to suberect

pubescence; mesosomal setae abundant; larger

species (HW 0.60-0.69) (Figs 19-21) 00...

LAR aaah A salichad A cies ns meorina coc haa Aaa L. mayri sp. n.

©ZFMK

244 Aijaz Ahmad Wachkoo et al.

Acknowledgements. Financial assistance rendered by the Min-

istry of Environment and Forests (Grant No. 14/10/2007-ERS/

RE), Govt. of India, New Delhi is gratefully acknowledged. We

are grateful to Peter Hawkes (University of Pretoria) and Fran-

cisco Hita Garcia (Okinawa Institute of Science and Technolo-

gy) for their helpful comments and suggestions about the man-

uscript. We are also thankful to the teams of AntWeb (www.

antweb.org), antmaps (www.antmaps.org) and AntWiki (www.

antwiki.org) for their immensely useful work. We would also

like to thank the subject editor Ralph S. Peters for his valuable

input.

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https://do1.org/10.20363/BZB-2021.70.2.247

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http://www.zoologicalbulletin.de

Research article

urn:|sid:zoobank.org:pub:69EE66EA-026F-45B5-AEDA-2D7FA1133B3A

A review of the genus Leptaulax Kaup, 1868

(Coleoptera: Passalidae) from Vietnam,

with the first record of L. loebli Kon, Johki & Araya, 2003

Phu Van Pham! *, Masahiro Kon’, Nhi Thi Pham’, Nga Quynh Thi Cao‘ & Tru Hoang Vu°

"34° Institute of Ecology ang Biological Resources, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet,

Cau Giay, Hanoi, Vietnam

? Pressance Kyodai-Higashi 406, 116-3 Nishida-cho, Jodoji, Sakyo, Kyoto, 606-8417 Japan

“Corresponding author: Email: phupham.iebr@gmail.com

'urn:lsid:zoobank.org:author:09B6F968-3B83-4052-89A 1-1DB2D91933CF

2urn:Isid:zoobank.org:author:A8D964E1-7B15-4C7B-BF63-555FD9FD966B

3urn:lsid:zoobank.org:author:34BE7CCE-9EES-4F 5E-A432-4467BA0101D4

*urn:lsid:zoobank.org:author:97296616-B2E7-4108-88C0-4B2C2E92CS5BB

>urn:lsid:zoobank.org:author:9F907485-D2A5-44A3-8BB1-55B74DBCF5FB

Abstract. This overview of the passalid genus Leptaulax Kaup, 1868 in Vietnam is based on literature and examined

specimens. A total of six species is recorded, of which Leptaulax loebli Kon, Johki & Araya, 2003 is recorded for the first

time for the fauna of Vietnam. An identification key to all Vietnamese Leptaulax species is provided.

Keywords. Genus Leptaulax, new record, Vietnam.

INTRODUCTION

The genus Leptaulax Kaup, 1868 was established based

on the type species Passalus dentatus Fabricius, 1792

and placed in the subfamily Passalinae. According to

present literature, ca. 70 species have been recorded in

this genus. Most of them occur in the Oriental region,

relatively few species are known from the Australian and

Palaearctic regions (Gravely 1918; Hincks & Dibb 1935;

Boucher 2006; Iwase 1995, 1996a, 1996b, 1998a, 1998b,

1998c; Johki & Kon 2003; Johki et al. 2003; Kon et al.

2003; Kon & Bezdek 2016; Kon et al. 2017).

In Vietnam, the genus Leptaulax was first reported by

Kuwert (1891) when the author described L. saigonicus

from Saigon (Ho Chi Minh City). After more than 20

years, Gravely (1914), who studied Oriental passalids

based on the collection at the Indian Museum (Kolkata),

synonymized L. saigonicus with L. dentatus and added

two species, L. bicolor and L. cyclotaenius for Vietnam’s

fauna. Iwase (1998) described L. pacholatkoi based on

material collected from Lang Biang Mountain, Da Lat,

Lam Dong Province (Vietnam). Recently, Kon et al.

(2017) described L. niisatoi collected from Phia Oac—

Phia Den National Park in 1992 in Northern Vietnam

Received: 08.09.2020

Accepted: 09.06.2021

and this species 1s currently known only from the type

locality. Based on the passalid collections at the Museum

ftir Naturkunde Berlin (MfN) and the Institute of Ecology

and Biological Resources (IEBR), we provide an over-

view on this genus for the first time and record a further

species, L. Joebli Kon, Johki & Araya, 2003, as new for

Vietnam. An identification key to Leptaulax species re-

corded so far from Vietnam is provided.

MATERIAL AND METHODS

The specimens in this study were collected by light trap

and by hand collecting. Male genitalia were extracted

and then cleaned in 10% KOH for 24 hours. After clean-

ing, they were glued to pinned cards. The photographs of

morphological details of the species were taken with a

Nikon DS-Fi3 camera attached to a Nikon SMZ80ON ste-

reomicroscope (Fig. 1A—T) and a Canon 700D combined

with Sigma 100 mm macro lens (Fig. 1U—Y). Afterwards,

images were stacked with Combine ZP and subsequently

edited (Photoshop ver. CS6).

We follow Gravely (1914) and Neumann et al. (2013)

for the terms used in the key and diagnoses.

Corresponding editor: D. Ahrens

Published: 25.06.2021

248 Phu Van Pham et al.

RESULTS

Genus Leptaulax Kaup, 1868

Leptaulax Kaup, 1868: 11.

Type species (by original designation): Passalus den-

tatus Fabricius, 1792.

Key to Leptaulax species of Vietnam

1. The 5" visible abdominal sternite emarginated at

POStENIOP Marea CP IS. WIVES Ba vant Beceoreseg eddematgnidhe 2

— The 5" visible abdominal sternite not emarginated at

posterior mare ir (E Ie CHD s cess eee ve chs daaakunea 3

2. Pronotum strongly and densely puncate in lateral

portion; lateral striae of elytron with large punctates

eh be ret he ence L. niisatoi Kon, Johki & Pham, 2017

— Pronotum weakly and sparely punctate in lateral

portion (Fig. 1S); lateral striae of elytron finely

PUNCLALS ceca deBit conse L. pacholatkoi twase, 1998

3. Last abdominal sternite widely hairy (Fig. 1H);

pronotum with few punctures (Fig. 1R); anterior

lower tOOnibiden tate: x. 2..9 0. cre eee

EXT bigger. bit Pity L. loebli Kon, Johki & Araya, 2003

— Last abdominal sternite hairless or rugose (Fig. 1G,

I); pronotum with a few punctures on lateral portion

(Fig. 1Q, T); anterior lower tooth simple ............... 4

4. Lateral striae of elytron with normal, more or less

rounded punctures (Fig. 1L) ......0.00c cece

Pee a AT Seah sed Lan he L. bicolor (Fabricius, 1801)

— Lateral striae of elytron with more or less scalariform

PUNE UIE SCE KS es rhe antes nndys A tt geateneis 5

5. Mesosternum with very obvious, strong punctures,

metasternum with irregular punctures in the middle

(Fig. 1M); hyposotomal process with distinct

longitudial groove on ventral surface .........0.....00.....

RITE «tA ete 7 oe L. cyclotaenius Kuwert, 1891

— Méesosternum and metasternum glabrous, without

punctures (Fig. 10); hyposotomal process without

distinct groove on ventral surface ............000..ee

Phe os Ay bah asec eRe L. dentatus (Fabricius, 1792)

Leptaulax bicolor (Fabricius, 1801)

(Fig. 1A-C, I, L, T, Y)

Passalus bicolor Fabricius, 1801: 256.

Leptaulax bicolor Kaup, 1868: 14.

Type specimens in the Copenhagen Museum (Arrow

1910).

Diagnosis. Parietal ridge extending to supraorbital

ridge, anterior marginal groove of pronotum more pro-

nounced and broader in dorsal view, always with many

large, distinct punctures; lateral margin with strong punc-

tures, particularly in the anterior portion very closely

lined up in pairs; hypostomal process without longitudi-

nal groove on ventral surface; male genitalia moderately

Bonn zoological Bulletin 70 (1): 247-251

robust, basal piece longer than parameres in ventral view,

paramere projected straightly.

Material examined. | 3, 1 9 (IEBR), Bac Kan Prov-

ince, Ba Be National Park, light trap, 8.v.2014, H.T. Pham

leg.— 1 ¢, 1 9 (IEBR), Cao Bang Province, Phia Oac-

Phia Den National Park, light trap, 11.v.2011, 1160 m

a.s.l., V.T. Hoang leg. — 1 3 (IEBR) Cao Bang Province,

Phia Oac-Phia Den National Park, light trap, 7.v.2013,

V.T. Hoang leg. — 1 ¢ (IEBR) Hoa Binh Province, Hang

Kia—Pa Co Nature Reserve, light trap, 19—28.vi.2002,

1100-1200 mas.l.—5 ¢¢, 4 29 (MEN), Tonkin, Mon-

tes-Mauson (Lang Son Province, Mau Son mountain),

April—May, 2—3000 ft, H. Fruhstorfer leg.

Distribution. Australia, Cambodia, Eastern Himala-

yas, India, Indonesia (Borneo, Java, Moluccas, Sulawesi,

Sumatra), Laos, Malaysia, Myanmar, New Guinea, Phil-

ippines, Sri Lanka, Thailand (Kon et al. 2001; Neumann

et al. 2013) and Vietnam (Lang Son,) (Gravely 1914).

This species is recorded from Bac Kan, Cao Bang and

Hoa Binh provinces for the first time.

Leptaulax cyclotaenius Kuwert, 1891

(Fig. 1G, K, M, Q, V)

Leptaulax cyclotaenius Kwuert, 1891: 188.

Type specimens in the Museum Histoire Naturelle, Paris

(R. Oberthur’s collection; Arrow 1910).

Diagnosis. Body more or less small, head and side of

pronotum strongly punctures. Sides of elytra with strong

scalariform punctures; mesosternum and the middle of

metasternum distinctly punctate; hypostomal process

with longitudinal groove on ventral surface.

Material examined. 5 ¢¢, 4 99 (IEBR), Kon Tum

Province, Ngoc Linh Nature Reserve, hands, 6.111.2019,

1700 ma.s.l., PV. Pham leg. —3 44 (IEBR), Lam Dong

Province, Bi Doup-Nui Ba National Park, light trap,

3.vi.2013, T.V. Hoang leg. — 1 2 (IEBR), Lam Dong

Province, Bi Doup-Nui Ba National Park, light trap,

4.vi.2013, D.T. Tran leg. — 1 2 (MfN), Tonkin, Mon-

tes-Mauson (Lang Son Province, Mau Son Mountain),

April-May, 2—3000 ft, H. Fruhstorfer leg.

Distribution. Cambodia, India, Indonesia (Borneo,

Sulawesi, Sumatra), Laos, Malaysia, Myanmar, Thailand

and Vietnam (Lang Son) (Gravely 1914). This species is

reported from the Central Highlands of Vietnam for the

first time (Kon Tum and Lam Dong provinces).

Leptaulax dentatus (Fabricius, 1792)

(Fig. 1D-F, O, W)

Passalus dentatus Fabricius, 1792: 24.

Leptaulax dentatus Kaup, 1868: 14.

Type specimens: may be preserved in Copenhagen Mu-

seum, but not confirmed yet.

Diagnosis. Parietal ridge not extending to supraorbital

ridge; side of elytra with distinct scalariform punctuation,

but unclear and strong as L. cyclotaenius; mesosternum

and metasternum almost glabrous; hypostomal process

©ZFMK

A review of the genus Leptaulax Kaup, 1868 from Vietnam, with the first record of L. /oebli 249

W a, Y

Fig. 1. A-F. Male genitalia. G—J. 5" visible abdominal sternite and last abdominal sternite. K—L. Lateral striae of elytron.

M-P. Meso- and metasternum. Q—-T. Pronotum. U-Y. Habitus (scale bar: 0.5 cm). A-C, I, L, T, Y. Leptaulax bicolor. D-F, O,

W. L. dentatus. G, K, M, Q, V. Leptaulax cyclotaenius. H, P, R, X. L. loebli. J, N, S, U. L. pacholatkoi.

Bonn zoological Bulletin 70 (1): 247-251 ©ZFMK

250 Phu Van Pham et al.

either without longitudinal groove or with weak depres-

sion in anterior portion; male genitalia robust and cyclin-

drical in dorsal view.

Material examined. | 2 (IEBR), Bac Kan Province,

Ba Be National Park, light trap, 6.viii.2002 —1 4 (IEBR),

Gia Lai Province, KBang District, Dak Roong Commue,

Kon Ka Kinh National Park, 06.v1.2011, 1060 m, light

trap — 2 5, 1 2 (TEBR), Hoa Binh Province, Ngoc Son-

Ngo Luong Nature Reserve, hands, 18.1x.2016, P.V.

Pham leg. — 1 ¢, 1 9 (IEBR), Kon Tum Province, Dak

Ha, Dak Man, light trap, 18.vi1.2012, V.T. Hoang leg. —

1 3 (EBR), Ninh Thuan Province, Phuoc Binh National

Park, light trap, 13.xi.2008 —7 63’, 5 92 (MEN), Tonkin,

Montes-Mauson (Lang Son Province, Mau Son moun-

tain), April-May, 2—3000 ft, H. Fruhstorfer leg. — 1 9°

(IEBR), Vinh Phuc Province, Tam Dao National Park,

light trap, 16—24.vi.2011, V.T. Hoang leg. — 1 9 (IEBR),

Vinh Phuc Province, Tam Dao National Park, light trap,

18.v.2014—6 3, 1 2 (IEBR), Vinh Phuc Province, Tam

Dao National Park, light trap, 12—16.11.2014.

Distribution. Cambodia, China, Eastern Himalayas,

India, Indonesia (Borneo, Java, Lesser Sunda, Moluc-

cas, Sulawesi, Sumatra), Laos, Malaysia, Myanmar, New

Guinea, Philippines, Tatwan, Thailand and Vietnam (Ho

Chi Minh, Lang Son,) (Gravely 1914; Kon et al. 2001).

This species 1s recorded from Bac Kan, Gia Lai, Hoa

Binh, Kon Tum, Ninh Thuan and Vinh Phuc provinces

for the first time.

Leptaulax loebli Kon, Johki & Araya, 2003

(Fig. 1H, P, R, X)

Leptaulax loebli Kon, Johki & Araya, 2003: 181.

Type specimens in Muséum d’ Histoire Naturelle Geneve,

Switzerland (Kon et al. 2003).

Diagnosis. Anterior lower of mandible tooth bidentate;

last abdominal sternite with an expanded and densenly

hairy and punctate area, 5th visible abdominal sternite

not emarginated at posterior margin; hypostomal process

without longitudinal groove.

Material examined. 1 9, 1 @ (MfN), Annam, Phuc

Son (Quang Nam Province, Phuoc Son?), xi—xu, H.

Fruhstorfer leg. — 1 ¢ (IEBR), Gia Lai Province, Kon Ha

Nung, Buon Luoi, light trap, 22.vi.1980, HTHCT.

Distribution. Previously known from Laos, Myanmar

and Thailand (Kon et al. 2003; Neumann et al. 2013).

These are the first records of this species from Vietnam.

Leptaulax niisatoi Kon, Johki & Pham, 2017

Leptaulax niisatoi Kon, Johki & Pham, 2017: 97.

Type specimens in Vietnam National Museum of Nature,

Hanoi (Kon et al. 2017).

Diagnosis. Parietal rigde extending to supraorbital

ridge; hypostomal process smooth, shiny and hairless;

mesosternum impunctate; metasternum entirely hairless,

central area shiny and impunctate; lateral striae of elytron

with large and somewhat oblong punctures; pronotum

Bonn zoological Bulletin 70 (1): 247-251

strongly and densely punctate in lateral portion; posterior

margin of 5th visible abdominal sternite gently emargi-

nated.

Material examined. None.

Distribution. Currently known only from Vietnam

(Kon et al. 2017).

Leptaulax pacholatkoi Twase, 1998

(Figs SU)

Leptaulax pacholatkoi Iwase, 1998: 157.

Type specimens in Natural History Museum Vienna

(Iwase 1998c).

Diagnosis. Anterior lower tooth simple; frontal area

more distinctly transverse; pronotum weakly and sparse-

ly punctate; last abdominal sternite more or less hairy

punctures, Sth visible abdominal sternite emarginated at

posterior margin.

Materials examined. | 9° (IEBR), Lam Dong Prov-

ince, Bidoup-Nui Ba National Park, Da Chais commune,

04.vi.2013, light trap, T.V. Hoang leg. — 1 2 (IEBR), Gia

Lai Province, KBang District, Dak Roong Commue, Kon

Ka Kinh National Park, 06.v1.2011, 1060 m, light trap.

Distribution. Cambodia, Laos and Vietnam (type lo-

cality: Lam Dong) (Neumann et al. 2013).

Remarks. Currently, two subspecies are recognized,

L. pacholatkoi pacholatkoi and L. pacholatkoi mutonia-

tus. They differ by the difference of punctures in par-

ticular behind anterior margin and around lateral scar

of pronotum, the width of wrinkles at posterior border

of pronotum, and punctures and hairs on 6th abdominal

segment. The female specimen in this study is closer to

L. pacholatkoi pacholatkoi, however, its lateral scar of

pronotum is smaller and less punctate.

CONCLUSIONS

The present study added Leptaulax loebli to the Leptau-

/ax fauna of Vietnam with the result that Vietnam harbors

now six known species of this genus. Additionally, we

updated the information on the distribution for three spe-

cies: L. bicolor, L. cyclotaenius and L. dentatus in the

country. Especially, the knowledge of the distribution

range of L. cyclotaenius is extended to the Central High-

lands of Vietnam.

Acknowledgments. We are grateful to Mr. Bernd Jaeger and

Mr. Joachim Villers in the Coleoptera collection of the MfN

for their help accessing to the important collection of Passali-

dae specimens. We would like to thank Assoc. Prof. Dr. Pham

Hong Thai (Vietnam National Museum of Nature, Vietnam) for

sending the scientific literature and Dr. Dirk Ahrens (Zoological

Research Museum Alexander Koenig, Bonn, Germany) for his

©ZFMK

A review of the genus Leptaulax Kaup, 1868 from Vietnam, with the first record of L. oebli 251

helpful comments on the manuscript. This research is funded by

IEBR under grant number IEBR.ThST.7-20.

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optera: Passalidae) from Southeast Asia in the collection of

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torischen Museums in Wien 100B: 157-173

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(Coleoptera, Passalidae) from Borneo. Elytra 21: 385-390

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Kaup J (1868) Prodromus zu einer Monographie der Passali-

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https://do1.org/10.20363/BZB-2021.70.2.253

ISSN 2190-7307

http://www.zoologicalbulletin.de

Research article

urn:lsid:zoobank.org:pub:91DC41FD-F43D-4DDD-9344-A72AC99C4C4E

Reptiles and amphibians of the State of Kuwait

Zuhair S. Amr'*, Abdulrahman AI-Sirhan Alenezi?, Mohammed T. Al-Sayegh? & Mohammad A. Abu Baker*

' Department of Biology, Jordan University of Science and Technology, Irbid, Jordan

?P.O. Box 49272, Omariya, Kuwait 85153

3Science Department, College of Basic Education, The Public Authority for Applied Education and Training, Kuwait

*Department of Biology, the University of Jordan, Amman, Jordan

“Corresponding author: Email: amrz@just.edu.jo

'urn:Isid:zoobank.org:author:2ACACEB7-5479-4043-9B71-8B965C3DDDFA

2urn:Isid:zoobank.org:author:99 148B32-D697-4A22-A 1 8E-25592CA9355E

3urn:|sid:zoobank.org:author:40A 1 C46E-6EFC-4C97-9DF2-F7EFB5C9463 1

“urn:|sid:zoobank.org:author: A048D018-F6EA-4CB2-9D56-FOFB45D6E4B5

Abstract. In this study, we identified the diversity of the freshwater, marine and terrestrial herpetofauna of the State of

Kuwait. It consists of a total of 45 extant species; 44 species of reptiles and a single species of amphibian according to

recent updated taxonomic studies. All specimens of reptiles collected and held in American and European natural history

museums are documented. Four species are added to the herpetofauna of Kuwait: Chalcides ocellatus, Hydrophis lapemo-

ides, Hydrophis viperina, and Trapelus agnetae.

Key words. Kuwait, herpetofauna, diversity, systematics, amphibians.

INTRODUCTION

The first study on the reptiles of Kuwait, published by

Eissa & El Assy (1975), included 29 species. Clayton &

Pilcher (1983) featured elaborate images of the reptiles

of Kuwait covering some species. However, no localities

were mentioned for the photographed reptiles.

Additional records of reptiles in Kuwait have been

published within the past 15 years, including: the mel-

anistic whip snake Dolichophis jugularis (Al-Mohanna

et al. 2007); Murray’s Comb-fingered Gecko Stenodac-

tylus affinis, and Gulf Sand Gecko Stenodactylus kho-

barensis (Al-Sirhan 2009; Metallinou et al. 2012); the

Web-footed Sand Gecko Stenodactylus arabicus (Delima

and Al-Nasser, 2007); and the Small-spotted Desert Liz-

ard Mesalina guttulata (Al-Sirhan 2008). Al-Sirhan &

Brown (2010) gave an account on the distribution of two

species of the genus Phrynocephalus in Kuwait.

The marine turtles of Kuwait have received much at-

tention, with several studies on their taxonomy and biolo-

gy (Gasperetti et al. 1993; Al-Mohanna & Meakins 1998,

2000a, 2000b; Meakins & Al-Mohanna 2000, 2003;

Bishop et al. 2007; Bishop & Alsaffar 2008; Al Mohan-

na & George 2010; Al-Mohanna et al. 2013; Rees et al.

2013).

Other studies focused on the biology of some des-

ert species (Agama persica = Trapelus persicus and

Diplometopon zarudnyi) (Cloudsley-Thompson 1979).

Received: 29.03.2021

Accepted: 18.06.2021

Sey & Al-Ghaith (2000) examined the helminths of the

green toad and the spiny tailed lizard in Kuwait. Several

papers on the hematology of Kuwaiti lizards were pub-

lished without giving localities for the studied material

(Abdel-Fattah et al. 1974; Al-Badry 1975; Al-Badry et al.

1975; Al-Badry & Al-Sdirawi 1976). The impact of oil

pollution on body size, weight, timing of morning emer-

gence, basking and foraging behaviours and substrate

preferences of Acanthodactylus scutellatus in Kuwait

was investigated (Al-Hashem et al. 2008; Al-Hashem &

Brain 2009a & b). Al-Sayegh (2017) conducted a study

on its eco-physiological implications of conservation. A

recent account on sea snakes of the Arabian Gulf with

an illustrated key was published by Rezaie-Atagholipour

et al. (2017).

The present study documents and updates previous

records of the herpetofauna of Kuwait, adding reptile

specimens collected from Kuwait and held in American

and European natural history museums as well as recent

observations.

MATERIAL AND METHODS

Geographical Setting

The total area of the State of Kuwait is 17,818 km? of land

and about 1,000 km? of off-shore islands. It is situated in

Corresponding editor: W. Bohme

Published: 25.06.2021

254 Zuhair S. Amr et al.

46°45'0"E 47°0'0"E 47°15'0"E 47°30'0"E 47°45'0"E 48°0'0"E 48°15'0"E 48°30'0"E 48°45'0"E

29°45'0"N 30°0'0"N 30°15'0"N

29°45'0"N

29°30'0"N

29°15'0"N

Tr]

—

29°0'0"N

28°45'0"N

e Localities

____| National Border

Elevation(M)

— High : 295

= Low: -42

28°30'0"N

0 12.5 25

28°15'0"N

Sources: Esri, US Es, NOAA

46°45'0"E 47°0'0"E 47°15'0"E 47°30'0"E 47°45'0"E 48°0'0"E 48°15'0"E 48°30'0"E 48°45'0"E

Fig. 1. Map of Kuwait showing localities from which materials were recorded.

Table 1. Natural history museums curating reptiles collected from Kuwait.

Abbreviation Name of museum No. of specimens

AMNH American Museum of Natural History, New York, USA 3

BEV Laboratoire de Biogéographie et Ecologie des Vertébrés de |’Ecole Pratique des Hautes 13

Etudes, Montpellier, France

BMNH British Museum of Natural History, London, UK 87

CAS California Academy of Sciences, USA y)

CMNH Carnegie Museum of Natural History, USA 1

FMNH The Field Museum (Chicago), Illinois, USA 2

LAMC Natural History Museum of Los Angeles County, California, USA 1

LSUMZ Louisiana State University Museum of Natural Science, Louisiana, USA 17

SM Senckenberg Museum, Frankfurt, Germany 1

FLMNH Florida Museum of Natural History, Florida, USA 1

USNM Smithsonian Museum of Natural History, Washington DC, USA 9

ZFMK Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany 1

Bonn zoological Bulletin 70 (2): 253-272 ©ZFMK

Reptiles and amphibians in Kuwait

Table 2. Localities indicated in the text and their coordinates.

Locality

Abdali

Abraj Khaitan

Agriculture Research Station

Ahmed Al Jaber Air Base

Al Abatih

Al Abraq

Al Ah’madi

Al Ah’madi Governorate

Al Asimah

Al Atraf

Al Dbatyyah

Al Jahra East Outfall

Al Jahrah

Al Jalia

Al Liyah

Al Mutla

Al Nuwaiseeb

Al Ritqa

Al Salmi

Al Shlallal Farm

Al Subiya

Al Sugaihia

Al Wafrah

Al Wafrah New City

Ali Al Salem Air Force Base

Auhha Island

Benadier

Bodai’s Farm

Burgan hills

Doha area

Failaka Island

Hawalli

Jal Az Zor

Judailiyat

Kabed

Kadma

Khiran

Khwaisat

Kuwait city

Mahboula

Mina Al Zour

Mina Said (Mina Al-Zour)

30°03’41.90”

29e1 F138”

29°08. 59°82”

28°56’26.70”

29°51. 18:20”

292 17-001 O-

29705-0528

29°08’ 46.65”

29°20" 0250"

29°18°56.40”

28°56’09.70”

29°21500,387

29°20" LNG5?

28°53’02.60”

2829S 23

29°24 50/52?

28°35’00.80”

30°04’39.20”

29°08’ 33.10”

29°34’23.50”

29°35’36.48”

29°39’ 56.80”

28°33 SR 52"

28°33’48.03”

29°217°31.40”

29222'° 39,982

28°47° 46.70”

20°57 55:60"

29°03) 10347

29°18758.68”

29°26’ 40.04”

29°20’ 43.06”

29°30’ 36.46”

29°19'00.06”

29°05’ 43.23”

29°24’ 58.30”

28°39 44.67”

29°22748.10”

29°22 il 32

29°08’51.80”

28°44’ 08.60”

28°44’29.10”

47°40’59.60”

47°58’26.42”

47°39’57.70”

47°47? 38.40”

47°317°46.20”

46°58’54.50”

48°04’34.61”

48°07? 26.28”

47°58’35.43”

47°32’50.81”

48°12? 10.20”

47°43’58.93”

47°40°31.94”

48°15’51.50”

47°10°06.07”

47°40°33.27”

48°23’06.60”

47°30°54.60”

46°43’09.40”

47°54’28.20”

48°06’53.76”

47°08 18.60”

48°04’20.73”

48°04?21.77”

47°30’ 16.98”

48°26’25.66”

48°16758.30”

47°48°42.90”

48°02’29.13”

47°48 12.35”

48°16751.30”

48°01°58.73”

47°48°38.95”

47°46°52.10”

47°43’09.79”

47°44’ 36.00”

48°18°49.82”

47°40’°59.70”

47°59°25.81”

48°07715.3”

48°22°57.90”

48°23’30.50”

255

Locality N E

Mula’a Farm 29°33°12.07” 47°42733.30”

N Bubiyan Bridge 29°36’28.06” 48°09'11.00”

N Sabriya 29°52’44.62” 47°52°49.71”

NW Kubber Island 29°10°00.90” 48°28°00.23”

Qaruh Island 28°49°03.20” 48°46’35.70”

Rabiya 29°17°57.53” 47°56°35.86”

Ras Salmiya 29°20°54.31” 48°05’50.35”

Rhawdatain 29°49°30.70” 47°50°25.55”

Sabah Al-Ahmad Natural 29°34’13.14” 47°48°44.67”

Reserve

Salmiya 29°20°26.00” 48°04’17.00”

Shiggat Ai-Awazin 29°52’55.08” 47°13°32.90”

Shuaiba Industrial Area 29°02’36.79” 48°09°23.54”

Subahiya 29°04’52.00” 48°06’46.50”

Sulaibikhat Reserve 29°19°12.20” 47°52719.90”

Sulaibiya Pivot Fields 29°15°12.60” 47°45711.30”

Tulha 29°35°08.50” 47°46’59.40”

Umm AI-Maradim Island 28°40°48.46” 48°39°06.28”

Umm Niga 30°01°04.10” 47°52723.10”

Wadi AI-Tilha 29°35’00.92” 47°49°58.73”

Zayed Reserve 29°20°46.70” 47°48°23.10”

Bonn zoological Bulletin 70 (2): 253-272

the most northwestern corner of the Arabian Gulf

(Fig. 1). Kuwait is an arid country, consisting mainly of

desert land. It is bordered by Saudi Arabia to the south

and southwest and by Iraq to the north and northwest.

The landscape is relatively flat, broken only by occa-

sional low sand dunes and shallow depressions. The sur-

face rises gently from east to west reaching about 300 m

above sea level at Al-Shigaya and Al-Salmi. The eastern

part of the state, including all of the inhabited area, over-

looks the Arabian Gulf along a coastline that extends for

about 195 km. The surface consists of flat sandy plains

interspersed with some low-rise hills, some of which

reach a height of nearly 145 meters, and in northern Ku-

wait there is a series of hills such as Jal Al-Zour ridge,

the hills of Al-Marw and Al-Luiah. In the south, hills in

the form of domes are common in the regions of War-

ah and Burgan. Sand dunes areas are mostly located in

northwestern Kuwait along Al-Huwamiliyah-Al-Nimri-

tayn zone reaching Al Atraf area. Kuwait is crossed by

several dry desert wadi systems, such as Wadi Al-Batin,

extending along the Kuwaiti-Iraqi borders about 150 km,

and running along the western borders of Kuwait, with

a width in some parts reaching about 10 km and a maxi-

mum depth of about 57 meters. To the north lies the Raw-

datain Plain, consisting of flat land that slopes towards

the east and the northeast (Fig. 2).

©ZFMK

256 Zuhair S. Amr et al.

(Photo by A. Alenezi). E. Khwaisat. F. Um Al Rimam (Photo by A. Alenezi).

Kuwait includes nine islands in the Arabian Gulf, the

largest is Bubtyan Island (683 km7). The coastal strip ex-

tends for about 500 km, with a number of small bays and

lagoons. The largest are Kuwait and the Kazma Bays,

with several lagoons such as Abdullah, Boubyan, and Al

Subiya khawrs in the north, and Al Maftah and Al Amma

khawrs in the south.

Museum specimens, literature and additional records

Reptiles specimens held at American and European mu-

seums were obtained from the online catalogues data-

bases or by communication with curators of museums

known to have materials from the Middle East. We locat-

ed 12 museums keeping reptiles collected from Kuwait

Bonn zoological Bulletin 70 (2): 253-272

with a total of 137 specimens. The British Museum of

Natural History, London, and the Louisiana State Univer-

sity Museum of Natural Science, Louisiana, USA, held

the highest number of specimens (Table 1). All previous-

ly published records were extracted from published pa-

pers, and books. Additionally, new record localities from

field observations are included. Table 2 lists all localities

indicated in the text.

RESULTS

A total of 45 extant species — 44 species of reptiles be-

longing to 14 families (Chelonitidae, Dermochelyidae,

Gekkonidae, Agamidae, Trogonophidae, Scincidae,

©ZFMK

Reptiles and amphibians in Kuwait 257

Lacertidae, Varanidae, Typhlopidae, Boidae, Colubridae,

Psammophiidae, Viperidae and Elapidae) and a single

species of amphibians of the family Bufonidae — are re-

ported.

Amphibians

Family Bufonidae

This family of toads is represented by a single species,

Bufotes sitibundus. Kuwait lacks natural freshwater bod-

ies. Species of this family are dependent on rainfall, and

now they are found around agricultural areas and around

sewage treatment plants.

Bufotes sitibundus (Pallas, 1771) (Fig. 3)

Variable Green Toad

Previous records: Al Wafrah (Sey & Al-Ghaith 2000) as

Bufo viridis.

New records: Abdali, Al Jahrah Farms, Sulaibiya Piv-

ot Fields, Bodai’s Farm.

Remarks: The green toad is very common at Al Jah-

rah Nature Reserve. The taxonomy of the genus Bufotes

is still not fully resolved. Previously, populations of the

Eastern Mediterranean were referred to the variabilis

complex (Dufresnes et al. 2019).

2000; Al-Mohanna et al. 2013; Rees et al. 2018). The

Green Turtle, Chelonia mydas is considered as the most

common species in the Arabian Gulf.

Family Cheloniidae

Caretta caretta (Linnaeus, 1758)

Loggerhead Turtle

Previous records: Shuaiba Industrial Area (Al-Mohan-

na & Meakins 2000)

Remarks: The Loggerhead Turtle is a rare species

in the Kuwaiti waters. It was reported once by Al-Mo-

hanna & Meakins (2000) from the water near Shuaiba

Industrial Area. Masirah Island, Oman, hosts the largest

nesting aggregations worldwide, with thousands of indi-

vidual turtles frequenting the island for nesting on yearly

base (Ross & Barwani 1982).

Chelonia mydas (Linnaeus, 1758) (Fig. 4A)

Green Turtle

Previous records: Kuwaiti waters (Eissa & El Assy

1975), Doha area (Al-Mohanna & Meakins 2000), Umm

Al-Maradim and Qaruh islands (Al-Mohanna et al. 2013).

Fig. 3. The Variable Green Toad, Bufotes sitibundus (Photo by

A. Alenezi).

Reptiles

Marine Turtles

Five species of marine turtles in two families (Chelonii-

dae and Dermochelyidae) are known to occur in the Ku-

wait waters (Gasperetti et al. 1993; Al-Muhanna & Meak-

ins 2000b; Bishop et al. 2007; Al-Mohanna & Meakins

Bonn zoological Bulletin 70 (2): 253-272

Fig. 4. Marine turtles of Saudi Arabia. A. Eretmochelys imbri-

cata. B. Chelonia mydas (Photos by A. Al Mansi).

©ZFMK

258

Remarks: This species was found to nest around Umm

Al-Maradim and Qaruh islands (Al-Mohanna et al. 2013;

Rees et al. 2018). Gasperetti et al. (1993) included a map

showing several nesting sites in the Arabian Gulf on the

coast of Saudi Arabia and the United Arab Emirates.

Eretmochelys imbricata (Linnaeus, 1766) (Fig. 4B)

Hawksbill Turtle

Previous records: Shuaiba Industrial Area (Al-Mohan-

na & Meakins 2000)

Remarks: This is a rare species in the Kuwaiti waters.

Few individuals were found to nest in Kuwait (Rees et al.

2018). Gasperetti et al. (1993) stated that this a rare spe-

cies in the Arabian Gulf with nesting populations around

the coasts of Saudi Arabia.

Lepidochelys olivacea (Eschscholtz, 1829)

Olive Ridley Turtle

Previous records: NW Kubber Island (Bishop et al.

2007).

Remarks: So far this is the only record for Kuwait

(Bishop et al. 2007). The Olive Ridley Turtle is consid-

ered as a pan tropical turtle of Atlantic and the Indo-Pa-

cific oceans (Gasperetti et al. 1993). Tollab et al. (2015)

gave recent observations for this species in the Arabian

Gulf including records from Iran and Bahrain.

Family Dermochelyidae

Dermochelys coriacea (Vandelli, 1761)

The Leatherback Turtle

Previous records: Benadier, S Kuwait (Al-Muhanna &

Meakins 2000b).

Remarks: This is a rare species in the Arabian Gulf

and the Arabian Sea (Gasperetti et al. 1993). A dy-

ing specimen was collected from Benadier, S Kuwait

(Al-Muhanna & Meakins 2000b).

Family Gekkonidae

This family is represented by six genera (Bunopus, Cyr-

topodion, Hemidactylus, Pseudoceramodactylus, Steno-

dactylus and Trigonodactylus), including nine species.

All species reported from Kuwait are eremic species

known across Arabia. Species of the genus Stenodactylus

were revised at the molecular level, including specimens

from Kuwait (Metallinou et al. 2012).

Bunopus tuberculatus Blanford, 1874 (Fig. 5A)

Baluch Rock Gecko

Museum specimens: BMNH 1972.713-715, Kuwait,

leg. A d’A Bellairs. BMNH 1972.354-358, Kuwait, leg.

Bonn zoological Bulletin 70 (2): 253-272

Zuhair S. Amr et al.

K Blackwell. BMNH 1976.1714, 1976, Kuwait, leg. D.

McIntyre. BMNH 1978.2067, 1978, Kuwait, leg. S. M.

Eissa. BMNH 1985.1161-1164, Auhha Island.

FMNH 190984, Kuwait, leg. M. Kaiser, M. Al-Taqi, S.

Issa, & K. Behbehani.

USNM 581915, 9.4.2014, Al Jahrah, 16 km W of Ali

Al Salem Air Base.

LSUMZ-Herps 88942, 89494, 89496-97, and 89499,

2.2.2005, Mahboula (Al Ah’madi Governorate), leg. J.

Bishop. LSUMZ-Herps 89502, 13.3.2005, Al Ah’madi

Governorate, leg. J. Bishop.

Previous records: Aubha and Kubbar islands (Eissa &

El Assy 1975), Al Wafrah (Al-Khalifa et al. 2012).

New records: Al Jahrah East Outfall, around Kabed

Reserve, Khwaisat, Mina Al Zour, Sabah Al-Ahmad Nat-

ural Reserve.

Remarks: This common species in Kuwait was re-

corded from several localities (Eissa & El Assy 1975;

Al-Khalifa et al. 2012). It is widespread across the Mid-

dle East extending as far east as Pakistan (Sindaco & Jer-

eméenko 2008).

Cyrtopodion scabrum (Heyden, 1827) (Fig. 5B)

Rough Bent-toed Gecko

Museum specimens: BMNH 1972.1311, 1972, Kuwait,

leg. A. d’A Bellairs. BMNH 1985.1165, Auhha Island,

leg. A. d’A Bellairs.

USNM Amphibians & Reptiles 581744, 15.4.2014, Ali

Al Salem Air Force Base (Al Jahrah Governorate).

Previous records: Jal Az Zour, Gabed (=Kabed) (Eis-

sa & El Assy 1975).

New records: Al Jahrah Farms, Jahrah East Outfall, Al

Wafrah Farms, south Subahiya,

Remarks. The Rough Bent-toed Gecko is widespread

across Egypt and the Middle East extending as far east as

India (Sindaco & Jereméenko 2008). This is a common

species that occupies houses. More than 30 individuals

were observed at south Subahiya, a place with ample

bushes and water.

Hemidactylus flaviviridis Ruppell, 1840 (Fig. 5C)

Yellow-belly Gecko

Museum specimens: BMNH 1978.2068, 1978, Kuwait,

leg. S.M. Eissa. BMNH 1985.1166, Auhha Island.

Previous records: Kuwait (Eissa & El Assy 1975).

New records: Abdali, Al Jahrah Farms, Al Wafrah

Farms, Kabed Reserve.

Remarks: This a widespread species distributed

across Egypt to Eritrea in Africa, to the Arabian Peninsu-

la, Iraq reaching as far as India (Sindaco & Jereméenko

2008). This is the largest gecko in Kuwait. It is frequently

observed at night on walls inside houses hunting insects

and spiders. It is very common in houses, usually seen

near florescent lights on buildings. Over the years and

©ZFMK

Reptiles and amphibians in Kuwait 259

*¢4

Beet

SU ia be

Fig. 5. Some geckos of Kuwait. A. Bunopus tuberculatus (Photo by A. Alenezi). B. Cyrtodactylus scabrum (Photo by A. Alenezi).

C. Hemidactylus flaviviridis (Photo by A. Alenezi). D. Hemidactylus persicus (Photo by A. Alenezi). E. Stenodactylus affinis (Pho-

to by A. Alenezi). F. Stenodactylus doriae (Photo by A. Al Yousef). G. Trigonodactylus arabicus.

based on field observations by one of the authors (A. Al

Sirhan), it is believed that Hemidactylus flaviviridis has

displaced Hemidactylus persicus in newly developed ar-

eas in Kuwait.

Hemidactylus persicus Anderson, 1872 (Fig. 5D)

Persia Leaf-toed Gecko

Museum specimens: BMNH 1972.716, Kuwait, leg. A.

dA Bellairs.

Bonn zoological Bulletin 70 (2): 253-272

Previous records: Kuwait (Eissa & El Assy 1975)

Remarks: It seems that the record of Hemidactylus

turcicus by Eissa & El Assy (1975) was considered to be

Hemidactylus persicus by Sindaco & Jereméenko (2008)

as shown by their distribution map for H. persicus. The

distribution of the Turkish Gecko Hemidactylus turcicus

is confined to Turkey and southern Europe (Greece, Italy

and Spain) (Moravec et al. 2011). Castilla et al. (2013)

gave a detailed distribution map for H. persicus, cov-

ering S Iran, Iraq, Saudi Arabia, N Oman, United Arab

©ZFMK

260

Emirates, Bahrain, Kuwait, Qatar, Pakistan and N India,

along with its habitat preference in Qatar. This species 1s

probably becoming rare or even extinct in Kuwait. It was

not found in Kuwait despite extensive search by one of

the authors (A. Al Sirhan), but may still exist in the old

houses in Kuwait City.

Stenodactylus affinis (Murray, 1884) (Fig. 5E)

Murray’s Comb-fingered Gecko

Museum specimens: BEV.10036, 500 m W of Sulai-

bikhat Reserve, W. Kuwait City. BEV.10095-98, Mina

Said.

Previous records: Al Jahrah East Outfall, JEO (AI-Sir-

han 2009), W Sulaibikhat Reserve, Mina Said (Metalli-

nou et al. 2012).

New records: Al Dubaiya, Al-Nuwaiseeb.

Remarks: Al-Sirhan (2009) found that this species is

the dominant gecko species in the Al Jahrah East Outfall,

inhabiting areas with bushes of Nitraria retusa. This spe-

cies is distributed throughout SE Iraq and Iran (Leviton

et al. 1992).

Stenodactylus doriae Blanford, 1874 (Fig. 5F)

Middle Eastern Short-fingered Gecko

Museum specimens: BEV.10037, 13 km N-NE of Al

Jahrah.

Previous records: Kuwait (Eissa & El Assy 1975), Al

Wafrah (Al-Khalifa et al. 2012), Al Jahrah (Metallinou

et al. 2012).

New records: Al Abragq, Al Ritga.

Remarks: This is a rather common species that is

widely distributed across the arid habitats of the Middle

East (Sindaco & Jereméenko 2008).

Stenodactylus slevini Haas, 1957

Slevin’s Short-fingered Gecko

Museum specimens: BEV.10065, Sabah Al-Ahmad Nat-

ural Reserve. BEV.T1501, Ratqa, Kuwait-Iraq borders.

LSUMZ-Herps 88944, 2.2.2005, Mahboula (AI

Ah’madi Governorate), leg. J. Bishop. LSUMZ-Herps

89505, 24.7.2004, Mahboula, 0.5km SW of Fintas Tow-

ers (Al Ah’madi Governorate), leg. J. Bishop. LSUMZ-

Herps 89506, 11 9.2004, Mahboula (Al Ah’ madi Gover-

norate), leg. J. Bishop.

USNM 129877-78, 1951, S of Kuwait.

Previous records: Kuwait (Eissa & El Assy 1975),

Sabah Al-Ahmad Natural Reserve, Al Ritga (Metallinou

et al. 2012).

New records: Al-Salmi.

Remarks: Slevin’s Short-fingered Gecko was found in

sandy areas in Kuwait. It is widespread across Bahrain,

S Jordan, S Iraq, Kuwait, Qatar, NW Saudi-Arabia, W

Bonn zoological Bulletin 70 (2): 253-272

Zuhair S. Amr et al.

United Arab Emirates and Yemen (Sindaco & Jereméen-

ko 2008).

Pseudoceramodactylus khobarensis Haas, 1957

Gulf Short-fingered Gecko

Museum specimens: BEV.10039-40, Al Wafrah Farms,

20 km E of Al Wafrah.

Previous records: Al Jahrah East Outfall, Al Ritqa

Police Station, Al Subrya (Al-Sirhan 2009), Al Wafrah

(Metallinou et al. 2012).

Remarks: This species was recorded from Kuwait by

Al-Sirhan (2009). Metallinou et al. (2012) updated the

distribution map of the Gulf Short-fingered Gecko in the

eastern Arabian Peninsula and Iran. It was recorded from

Bahrain, Kuwait, Oman, Qatar, Saudi Arabia, United

Arab Emirates and S Iran. Valdeon et al. (2013) stated

that this species was found to inhabit coastal areas in Qa-

tar, but also was found on inland sabkhas.

Trigonodactylus arabicus (Haas, 1957) (Fig. 5G)

Web-footed Sand Gecko

Museum specimens: BEV.10042-43, Al Wafrah Farms,

20 km E of Al Wafrah.

Previous records: Agriculture Research Station (Deli-

ma & Al-Nasser 2007)

Remarks: This species was recorded by Delima &

Al-Nasser (2007) for the first time in Kuwait. Its distribu-

tion ranges across eastern Arabia including Saudi Arabia,

Qatar, United Arab Emirates, S Oman, Bahrain and east

to Iran (Sindaco & Jereméenko 2008).

Family Agamidae

Five species within three genera (Phrynocephalus,

Trapelus, and Uromastyx) were recorded from Kuwait.

The Egyptian Spiny-tailed Lizard, Uromastyx aegyptia,

is a species of special conservation importance.

Phrynocephalus arabicus (Anderson, 1894) (Fig. 6A)

Arabian Toadhead Agama

Previous records: Al Atraf, Kabed, Umm Niga (AI-Sir-

han & Brown 2010).

New records: Al Abatih. Al Sugaihia.

Remarks: The taxonomic status of the Ph. arabicus is

complex, requiring further evaluation of the populations

in eastern Arabia. Melnikov et al. (2014) showed that at

least three species of this complex occur within the Ara-

bian Peninsula; Phrynocephalus arabicus sensu stricto

for the southern Arabian populations, Phrynocephalus

nejdensis for the north-western Arabia populations in-

cluding southern Jordan, and Phrynocephalus macropel-

tis representing the populations in eastern coastal Arabia.

Phrynocephalus maculatus Anderson, 1872 (Fig. 6B)

©ZFMK

Reptiles and amphibians in Kuwait 261

bss

as.

Fig. 6. Agamids of Kuwait. A. Phrynocephalus arabicus. B. Phrynocephalus maculatus (Photo by A. Alenezi). C. Trapelus agnetae

(Photo by A. Alenezi). D. Trapelus persicus (Photo by A. Alenezi).

Phrynocephalus maculatus Anderson, 1872

Blacktail Toadhead Agama

Museum specimens: BMNH 1973.472, Kuwait, leg. K.

Blackwell. BMNH 1978.2070-2071 Kuwait, leg. S.M.

Eissa. BMNH 1985.1167, Auhha Island.

Previous records: Al Ah’madi and Burgan hills, Jal

Az Zour (Eissa & El Assy 1975), Khiran (Al-Sirhan &

Brown 2010).

New records: Sabah Al-Ahmad Natural Reserve.

Remarks: This is a common species associated with

vegetated mixed sand and gravel areas. Al-Sirhan &

Brown (2010) gave an account on its presence in Kuwait.

It was reported from SE Al-Samawa, Iraq on shrubs or

buried in the soil (Al-Barazengy 2014). It was also ob-

served in coastal area east to Sabah Al-Ahmad Natural

Reserve.

Trapelus agnetae (Werner, 1929) (Fig. 6C)

New records: Al Liyah Reserve, Al Ritqa, Al Salmi.

Remarks: This species is known to occur across Jor-

dan, Iraq, N Saudi Arabia and Kuwait (Arnold 1986; Sin-

daco & Jereméenko, 2008).

Trapelus persicus (Blanford, 1881) (Fig. 6D)

Museum specimens: BMNH 1978.1000, 1978, Kuwait,

leg. S.M. Eissa. BMNH 1970.1922-1926, Kuwait city,

leg. A. d’A Bellairs.

Bonn zoological Bulletin 70 (2): 253-272

LSUMZ-Herps 44766, 18.2.1983, Al Jahrah Gover-

norate, leg. J. Bishop. (listed as Trapelus blanfordi).

Previous records: Al Jalia, Al Wafrah, Gabed

(=Kabed) (Eissa & El Assy 1975), Umm Niga (Clouds-

ley-Thompson 1979).

New records: Sulaibikhat, Zayed Reserve.

Remarks: Rastegar-Pouyani (2000) synonymized

T. persicus with T: ruderatus. For nomenclature stability

Ananjeva et al. (2013) adopted 7 persicus to describe

what was originally known as 7. persicus for the Middle

East. This is a common species in Kuwait with several

records. It is found along coastal areas, but not in true

desert west or north of Kuwait.

Uromastyx aegyptia (Forskal, 1775) (Fig. 7)

Egyptian Spiny-tailed Lizard

Museum specimens: BMNH 1971.749, 1971, Kuwait,

leg. A. d’A Bellairs. BMNH 1978.2072, 1978, Kuwait,

leg. S. M. Elissa.

LSUMZ-Herps 88937 and 88939, 2.2.2005, Mahbou-

la (Al Ah’madi Governorate), leg. J. Bishop. LSUMZ-

Herps 88940, 26.10.2004, Mahboula (Al Ah’ madi Gov-

ernorate), leg. J. Bishop. LSUMZ-Herps Herps 88941,

13.3.2005, Mahboula (Al Ah’madi Governorate), leg. J.

Bishop. LSUMZ-Herps 89513-16, 13.3.2005, Mahbou-

la (Al Ah’madi Governorate), /eg. J. Bishop. LSUMZ-

Herps 89519, 2.2.2005, Mahboula (Al Ah’madi Gover-

norate), /eg. J. Bishop. LSUMZ-Herps 89520 and 89522,

©ZFMK

262 Zuhair S. Amr et al.

Fig. 7. The Egyptian Spiny-tailed Lizard, Uromastyx aegyptia (Photo by A. Ragab).

17.11.2005, Mahboula, 1km SW Fintas Towers (Al

Ah’madi Governorate), leg. J. Bishop.

Previous records: Al Mutla, Al Wafrah, Al Atraf,

Kathma, Rhawdatain (Eissa & El Assy 1975), Desert

around Kuwait City (Sey & Al-Ghaith 2000), Al Wafrah

(Al-Khalifa et al. 2012), Al Liyah (Al Jahrah Governor-

ate) and Kabed (Farwaniyah Governorate) (Al-Sayegh

2017).

New records: Al Salmi.

Remarks: The Spiny-tailed Lizard Uromastyx aegyp-

tia 1s a common species in the arid habitats of Kuwait

as recorded above. Food plants consumed by this spe-

cies were studied by Robinson (1995). Wilms & Bohme

(2007) revised the distribution and systematics of lizards

of the genus Uromastyx in Arabia, including Kuwait. Al-

Sayegh (2017) conducted a comprehensive study of the

eco-physiological (body mass, tail volume, body tem-

perature, and active hours) implications for conservation

of this species in two protected and two non-protected

locations in Kuwait. This species has a special conserva-

tion value since it is captured and traded in Kuwait and in

Saudi Arabia where its populations are declining (Aloufi

et al. 2019).

Bonn zoological Bulletin 70 (2): 253-272

Family Trogonophidae

This family is represented by a single species in Kuwait.

The Zarudny’s Worm Lizard Diplometopon zarudnyi was

reported from several localities across Kuwait.

Diplometopon zarudnyi Nikolsky 1907 (Fig. 8A)

Zarudny’s Worm Lizard

Museum specimens: AMNH 134244-46, Kuwait.

BMNH 1977.356, 1977, Kuwait, leg. A. d’A Bellairs.

BMNH 1977.770, 1977, Kuwait, leg. A. d’A Bellairs.

BMNH 1978.1313, 1978, Kuwait, leg. A. d’A Bellairs.

BMNH 1978.2080, 1978, Kuwait, leg. S. M. Eissa.

BMNH 1980.188-189, 1980, Kuwait, leg. A. d’A Bel-

lairs. BMNH 1983.1284-1285, 1983, Kuwait, leg. A. d’A

Bellairs.

FMNH (Amphibians & Reptiles) 263890, 24.7.1972,

Kuwait.

LSUMZ-Herps 44767-68, 26.7.1984, Wadi Al-Tilha

(Al Jahrah Governorate), leg. J. Bishop. - LSUMZ-Herps

88933, 2005.5.5, Mahboula (Al Ah’madi Governorate),

leg. J. Bishop. LSUMZ-Herps 89528-29, 1.5.1986, Shig-

gat Ai-Awazin (Al Jahrah Governorate), leg. J. Bishop.

LSUMZ-Herps 89530, 2.5.1986, Shiggat Ai-Awazin,

(Al Jahrah Governorate), leg. J. Bishop. LSUMZ-Herps

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Reptiles and amphibians in Kuwait 263

A. Alenezi). C. Scincus mitranus (Photo by A. Alenezi). D. Scincus scincus.

89531, 18.4.1986, Al Wafrah area (Al Ah’madi Gover-

norate), leg. J. Bishop. LSUMZ-Herps 89532, 9.3.1989,

Ras Salmiya (0.5km SSW Ras Al-Ardh), (‘Hawalli Gov-

ernorate), leg. J. Bishop.

LACM-Herps 186768, 22.5.2016, Ahmad Al Jaber Air

Base, leg. S. White.

SM 64516, 18.5.1967, 10 miles away of the Shuaiba

Coast.

FLMNH-Herp 68567, 30.11.1977, Kuwait, leg. C.

Gans.

Previous records: Kuwait (Eissa & El Assy 1975)

New records: Al! Shlallal Farm, Al Subtya.

Remarks: This is a common species in Kuwait with

several records. It is a burrowing species, distributed

throughout eastern and central Arabia, W Iran and S Iraq

(Sindaco & Jereméenko 2008).

Family Scincidae

The skinks of Kuwait are exemplified in four species rep-

resenting three genera (Ablepharus, Chalcides and Scin-

cus). Species of the genus Scincus are strictly sand-dwell-

ing species. Arnold & Leviton (1977) revised the species

of the genus Scincus in Arabia.

Bonn zoological Bulletin 70 (2): 253-272

Ablepharus pannonicus (Fitzinger, 1824) (Fig. 8B)

Asian Snake-eyed Skin

New records: Al Jahrah Farms, Abdali Farms.

Remarks: The only previously available documen-

tation of this species is based on an image depicted by

Clayton & Pilcher (1983) without a specific locality.

Specimens were found under the dry palm leaves in dif-

ferent locations. It is a common species at Al Jahra farms.

Chalcides ocellatus (Forskal, 1775)

Ocellated (Bronze) Skink

Museum specimens: BMNH 1978.2079, 1978, Kuwait,

leg. S.M. Elissa.

Remarks: This species was not listed by Eissa & El

Assy (1975). It is widely distributed through southern It-

aly to the Middle East, to North Africa and the Sahara,

eastwards to India (Sindaco & Jereméenko, 2008).

Scincus mitranus Anderson, 1871 (Fig. 8C)

Arabian Sand Skink

Museum specimens: BMNH 1976.1859, 1976, Kuwait,

leg. I. Said.

©ZFMK

264 Zuhair S. Amr et al.

Remarks: A specimen from Kuwait was depicted by

Arnold & Leviton (1977). Its distribution is confined to

south and east Arabia, including Saudi Arabia, Oman,

Qatar, United Arab Emirates, and Yemen (Arnold & Levy-

iton 1977).

Scincus scincus (Linnaeus, 1758) (Fig. 8D)

Sandfish Skink

Museum specimens: BMNH 1972.1311, 1972, Kuwait,

leg. K. Blackwell.

Previous records: Al Jalia (Eissa & El Assy 1975),

Kuwait (Arnold & Leviton 1977).

New records: Al Sugaihia, Kabed Reserve.

Remarks: Previously recorded from Kuwait (Eissa &

EI Assy 1975; Arnold & Leviton 1977). This is a sand

dwelling species distributed across eastern Arabia and

Iran (Arnold & Leviton 1977).

Family Lacertidae

Six species of lacertids occur in Kuwait, belonging to two

genera, Acanthodactylus and Mesalina. The systematics

of the species belonging to the genus Acanthodactylus

were revised by Salvador (1982) and at the molecular

level by Tamar et al. (2016). Sindaco et al. (2018) revised

the Mesalina guttulata species complex from Arabia, but

the status of Mesalina in Kuwait remains unresolved.

Acanthodactylus boskianus (Audouin, 1829) (Fig. 9A)

Bosk’s Fringe-fingered Lizard

Previous records: Al Ah’madi, Al Jalia, Al Mutla, Al

Wafrah, Kadma, Rhawdatain (Eissa & El Assy 1975)

New records: Al Abraq.

Remarks: This was a common species in Kuwait and

was recorded from several localities (Eissa & El Assy

1975). It is distributed across North Africa, the Sahara,

Sudan and Somalia to the Middle East (Salvador 1982).

Nowadays, it is not common, and was only observed at

Al Abragq (Personal observations by A. Al Sirhan).

Acanthodactylus opheodurus Arnold, 1980 (Fig. 9B)

Snake-tailed Fringe-toed Lizard

Museum specimens: BMNH 1964.97, Al Ah’ madi, leg.

D Vesey-Fitzgerald. BMNH 1972.360, 1973.101 (3 spec-

imens), Kuwait. BMNH 1970.1927, City of Kuwait, leg.

A @A Bellairs.

Previous records: Kuwait, City of Kuwait (Salvador

1982).

New records: Al Jahrah East outfall, Jal Al-Zour, Kh-

iran.

Bonn zoological Bulletin 70 (2): 253-272

Remarks: This is a rather common species in the Ara-

bian Peninsula, Jordan and Iraq (Sindaco & Jeremcéenko

2008). It is common in the sand dunes near Khiran living

besides the bushes of Lycium shawii and other halophyte

shrubs.

Acanthodactylus schmidti Haas, 1957 (Fig. 9C

Schmidt’s Fringe-fingered Lizard

Previous records: Al Ah’madi, Al Jalia, Al Mutla, Al

Wafrah, Kadma, Rhawdatain (Eissa & El Assy 1975), Al

Wafrah (Al-Khalifa et al. 2012).

New records: Al Subiya, N Bubtyan Bridge, Sabah

Al-Ahmad Natural Reserve.

Remarks: This is a sand dwelling species distributed

across the Arabian Peninsula, Jordan, SE Iraq, SW Iran

(Sindaco & Jereméenko 2008). It was collected from sev-

eral localities in Kuwait (Eissa & El Assy 1975).

Acanthodactylus hardyi Haas, 1957 (Fig. 9D)

Hardy’s Fringe-fingered Lizard

Museum specimens: BMNH 1964.123, Hawalli, leg. D.

Vesey-Fitzgerald. BMNH 1972.361, City of Kuwait, leg.

K. Blackwell. BMNH 1970.1928-1939, Kuwait city, leg.

A. dA Bellairs. BMNH 1978.2074-2076, 1978, Kuwait,

leg. S. M. Eissa.

BEV.11053, 35 km SW. of Ratqa.

CAS 190901-2, 1970, Al Asimah, leg. A. d’A. Bellairs.

LSUMZ-Herps 89533, 18.4.1986, Al Wafrah area (Al

Ah’madi Governorate), leg. J. Bishop.

Previous records: Al Ah’madi, Al Jalia, Al Mutla, Al

Wafrah, Kadma, Rhawdatain (Eissa & El Assy 1975),

Al Wafrah (Al-Khalifa et al. 2012) as Acanthodactylus

scutellatus.

New records: Al Ritga, Al Salmi, Al-Sugaihia, Kad-

ma.

Remarks: The status of this species remains unclear. It

was originally described as a subspecies of Acanthodac-

tylus scutellatus by Haas (1957). From a taxonomic point

of view, several authorities considered A. hardyi as a

member of the scutellatus group (Salvador 1982; Tam-

ar et al. 2016). This is a common species in Kuwait in-

habiting various habitats (Eissa & El Assy 1975). This

species was a subject of several studies in Kuwait that in-

vestigated changed substrate preferences (Al-Hashem &

Brain 2009a), and the effects of oil pollution on its body

size, weight, emergence, basking and foraging behaviors

(Hashem & Brain 2008; 2009a and b). This species oc-

curs in Jordan, Iraq, Saudi Arabia, Kuwait (Tamar et al.

2016).

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Reptiles and amphibians in Kuwait 265

Fig. 9. Lacertids of Kuwait. A. Acanthodactylus boskianus. B. Acanthodactylus opheodurus. C. Acanthodactylus schmidti.

D. Acanthodactylus scutellatus. E. Mesalina brevirostris. ¥. Mesalina guttulata. (All Photos by A. Alenezi).

Mesalina brevirostris Blanford, 1874 (Fig. 9E)

Blanford’s Short-nosed Desert Lizard

Museum specimens: BMNH 1970.1940-1941, Kuwait.

BMNH 1972.362-364, Kuwait, leg. K Blackwell. BMNH

1978.2077-2078, 1978, Kuwait, leg. S.M. Elissa.

Previous records: Al Ah’madi, Al Jalia, Al Mutla, Al

Wafrah, Kadma, Rhawdatain (Eissa & El Assy 1975), Al

Wafrah (Al-Khalifa et al. 2012).

New records: Al Salmi, Al Ritga, Failaka Island.

Remarks: This is a rather common species in Kuwait

inhabiting gravelly deserts. It is distributed across North

Africa, Sudan and Somalia to the Middle East (Sinda-

co & Jereméenko 2008).

Bonn zoological Bulletin 70 (2): 253-272

Mesalina guttulata (Lichtenstein, 1823) (Fig. 9F)

Small-spotted Desert Lizard

Previous records: Al-Salmi (Al-Sirhan 2008).

New records: Sabah Al-Ahmad Natural Reserve.

Remarks: This lizard was included in the herpetofauna

of Kuwait by Al-Sirhan (2008), and was encountered at

Sabah Al-Ahmad Natural Reserve. It is distributed across

North Africa to the Middle East (Sindaco & Jereméenko

2008).

Family Varanidae

One species of the family Varanidae was reported to

occur in Kuwait. Varanus griseus is widely distributed

across the Arabian Peninsula.

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266

Varanus griseus (Daudin, 1803) (Fig. 10)

Desert Monitor

Museum specimens: USNM 129879, 1951, Kuwait,

south of; along Persian Gulf strip. Voucher image,

ZFMK, 14.5. 1998, Wirral Sabriya (=N Sabrtya).

Previous records: Al Mutla, Rhawdatain (Eissa & El

Assy 1975).

New record: Kabed Nature Reserve, Tulha.

Remarks: The Desert Monitor is a common species

in the Arabian Peninsula, distributed from North Africa,

through arid parts of the Middle East, to reach as far as

India (Sindaco & Jereméenko 2008).

Zuhair S. Amr et al.

Previous records: Al Dbaiyyah, Al Jahrah, Al Jalia, Al

Wafrah, Khwaisat (Eissa & El Assy 1975).

Remarks: This obligate parthenogenetic species is

widespread with records from most of the world. It seems

to be common in Kuwait, resulting from its introduction

in plants imported in pots (Egan 2007).

Family Boidae

This family is represented by a single species. Eryx jay-

akari. It seems to be common throughout the country. In

the Arabian Peninsula, two species were recorded, Eryx

Jaculus and E. jayakari (Egan 2007).

Fig. 10. The Desert Monitor, Varanus griseus (Photo by A. Ragab).

Family Typhlopidae

This family is represented by one species, /ndotyphlops

braminus. This Asian species is widespread and has be-

come almost cosmopolitan in distribution.

Indotyphlops braminus (Daudin, 1803)

Brahminy blind Snake, Flowerpot Snake

Museum specimens: LSUMZ-Herps 88932, 15.10.2005,

Rabiya (Farwaniyah Governorate), leg. J. Bishop.

Bonn zoological Bulletin 70 (2): 253-272

Eryx jayakari Boulenger, 1888 (Fig. 11A)

Arabian Sand Boa

Museum specimens: BMNH 1972.366, Kuwait, leg. K

Blackwell. BMNH 1978.2081, Kuwait, leg. S. M. Elissa.

BMNH 1985.1168, Auhha Island.

CMNH-Herps 157129, 24.7.1972, Kuwait.

LSUMZ-Herps 89536, 22.6.2003, Al Wafrah New City

(Al Ah’ madi Governorate), leg. J. Bishop.

Previous records: Kuwait (Eissa & El Assy 1975).

New records: Al Ritga, Subtya.

©ZFMK

Reptiles and amphibians in Kuwait

: 3 y, ‘ 4 5

—y fe sae a hea) By ‘ ‘ ss , «EY A "AW

267

Fig. 11. Some snakes of Kuwait. A. Eryx javakari(Photo by A. Alenezi). B. Lytorhynchus diadema gaddi (Photo by A. Alenezi).

C. Malpolon moilensis. D. Platyceps ventromaculatus (Photo by A. Alenezi). E. Spalerosophis diadema cliffordii. F. Cerastes

gasperettii (Photo by A. Alenezi).

Remarks: The Arabian Sand Boa is distributed across

the eastern part of the Arabian Peninsula (Kuwait, Oman,

Saudi Arabia, United Arab Emirates) Iraq and SW Khuz-

estan, Iran (Egan 2007).

Family Colubridae

The family Colubridae, sensu lato, includes the highest

number of snake species with worldwide distribution. In

Bonn zoological Bulletin 70 (2): 253-272

Kuwait it is represented by four species in four genera

(Dolichophis, Lytorhynchus, Platyceps and Spaleroso-

phis).

Dolichophis jugularis (Linnaeus 1758)

Large Whip Snake

Previous records: Doha (AI-Mohanna et al. 1997)

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268

Remarks: This snake was reported from Doha, Ku-

wait (AI-Mohanna et al. 1997). In southern Iraq, it was

found in Al-Kufa district (Rhadi et al. 2017). This species

is common in Jordan, Lebanon, Palestine, Syria and Tur-

key (Amr & Disi 2011).

Lytorhynchus diadema_ gaddi

(Fig. 11B)

Crowned Leafnose Snake

(Nikolsky, 1907)

Previous records: Al Jalia, Al Jahrah, Al Wafrah (Eis-

sa & El Assy 1975), Kuwait (Meakins & Al-Mohanna

2003).

New records: AI Ritqa.

Remarks: Shafiei et al. (2015) showed that L. d. gaddi

occurs to the west of Iran with localities bordering Iraq.

This snake 1s associated with loose soil, especially sand.

Platyceps ventromaculatus (Gray, 1834) (Fig. 11D)

Platyceps ventromaculatus (Gray, 1834)

Glossy-bellied Racer

Museum specimens: BMNH 1947.3.2.30-32, Kuwait,

leg. D.V. Dickson.

USNM 581743, 4, 15.4.2014, Ali Al Salem Air Force

Base (AI Jahrah Governorate).

Previous records: Al Jalia (Eissa & El Assy 1975).

New records: Judailiyat, Kabed Reserve.

Remarks: Schatti (2006) referred to the populations of

this species inhabiting Iraq, Kuwait and Saudi Arabia as

Platyceps cf. ventromaculatus.

Spalerosophis diadema cliffordii (Schlegel, 1837)

(Fig. 11E)

Spalerosophis diadema cliffordii (Schlegel, 1837)

Clifford’s Royal Snake

Museum specimens: BMNH 1972.367, Kuwait, leg. K.

Blackwell.

Previous records: Al Atraf, Al Jalia (Eissa & El Assy

1975).

New records: Amgara, Mula’a farm, Sabah Al-Ahmad

Natural Reserve.

Remarks: The Clifford’s Royal Snake has a wide

range of distribution extending from North Africa to

western Iran, southern Turkey to the Arabian Peninsula

(Amr & Disi 2011). This is an aggressive snake inhabit-

ing arid regions.

Family Psammophiidae

This family includes two species in two genera (Rhager-

rhis and Psammophis). Both species are desert adapted

species.

Bonn zoological Bulletin 70 (2): 253-272

Zuhair S. Amr et al.

Rhagerrhis (Malpolon) moilensis (Reuss,

(Fig. 11C)

Moila Snake

1834)

Museum specimens: BMNH 1972.368-69, Kuwait, leg.

K. Blackwell. BM1978.2082, Kuwait, leg. S. M. Eissa.

LSUMZ-Herps 89508, 19.1.2005, Al Ah’ madi Gover-

norate, leg. J. Bishop.

Previous records: Al Jalia (Eissa & El Assy 1975).

New records: Sabah Al-Ahmad Reserve.

Remarks: This is a diurnal snake inhabiting gravelly

deserts. It is distributed in Algeria, Egypt, and Sudan to

Southwest Asia including southwestern Iran (Amr & Disi

2011).

Psammophis schokari (Forskal, 1775)

Forskal’s Sand Snake

Previous records: Al Jahrah, Al Wafrah (Eissa & El

Assy 1975)

Remarks: This is acommon snake inhabiting arid habi-

tats as well as temperate Mediterranean areas. Its distri-

bution extends from North Africa to India and Syria to

the Arabian Peninsula (Amr & Disi 2011).

Family Viperidae

This family includes one species of the genus Cerastes.

This genus is widespread across the deserts of northern

North Africa eastward through Arabia and to Iran, with

three species; Cerastes gasperettii, Cerastes cerastes and

Cerastes vipera (Sindaco et al. 2013).

Cerastes gasperettii Leviton & Anderson, 1967 (Fig. 11F)

Museum specimens: BMNH 1947.3.2.34, Abraj Khai-

tan, 8 miles S. of Kuwait, leg. Dr. H. Dickson. BMNH

1967.2200, Al Jahrah, leg. D. Vesey-Fitzgerald. BMNH

1972.371-372, Kuwait, leg. K. Blackwell. BMNH

1978.2085, Kuwait, leg. S.M. Eissa. BMNH 1985.1172,

Aubha Island.

USNM 129881, 1951, Kuwait, S of, along Persian Gulf

strip.

Previous records: Al Ah’madi, Al Jalia, Jal Az Zour,

Kadma (Eissa & El Assy 1975)

New records: outside Kabed Reserve, Rawdatain,

Subiya.

Remakes: This is a psammophile species known

across the sand areas of the Arabian Peninsula, south-

ern Iraq and Jordan (Gaspereti 1988; Amr & Disi 2011;

Rhadi et al. 2017). An illustrated guide to the horned vi-

per of Kuwait was published by Al-Fares and Al-Metairie

(2014) with a series of photographs.

©ZFMK

Reptiles and amphibians in Kuwait 269

Family Elapidae

Subfamily Hydrophiinae

Ten species of sea snakes have been recorded from the

Arabian Gulf (Rezaie-Atagholipour et al. 2016; Castilla

et al. 2017; Buzas et al. 2018). Within the Kuwaiti waters,

the presence of four species 1s confirmed. Rezaie-Atagho-

lipour et al. (2016) presented a comprehensive account

on the sea snakes of the Arabian Gulf including identifi-

cation keys and illustrations.

HAydrophis cyanocinctus Daudin, 1803 (Fig. 12A)

Annulated Sea Snake

Museum specimens: BMNH 1978.2083, 1978, Kuwait,

leg. S. M. Elissa.

LSUMZ-Herps 43335, 3, 20.6.1983, Kuwait, leg. J.

Bishop. LSUMZ-Herps 98341, 26.9.2004, Station BUB-

02 of Boubyan Project (Al Jahrah Governate), leg. J.

Bishop.

Previous records: Kuwait (Eissa & El Assy 1975).

Remarks: The Annulated Sea Snake is known from

the Indo-West Pacific, from the Arabian Gulf to Japan

(David and Ineich 1999). Within the Arabian Gulf, it was

recorded from the waters of all Gulf States (Castilla et al.

2017).

Hydrophis lapemoides (Gray, 1849) (Fig. 12B)

Persian Gulf Sea Snake

Museum specimens: LSUMZ-Herps 43336, 9,

20.6.1983, Kuwait, leg. J. Bishop.

Remarks: This snake occurs across the Indian Ocean,

extending from the Arabian Gulf to Malay Archipelago

(Rasmussen 1987; Rezaie-Atagholipour et al. 2016).

Within the Arabian Gulf, tt was recorded from the wa-

ters of all Gulf States (Castilla et al. 2017). This museum

record represents the first documentation of the Persian

Gulf Sea Snake from the Kuwaiti waters.

Fig. 12. Sea snakes of Kuwait. A. Hydrophis cyanocinctus (Photo by Csaba Géczy). B. Hydrophis lapemoides (Photo by Csaba

Géczy). C and D. Hydrophis platurus (Photo by Balazs Buzas -balazsbuzas.com).

Bonn zoological Bulletin 70 (2): 253-272

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270

Hydrophis platurus (Linnaeus, 1766) (Fig. 12C—D)

Yellow-bellied Sea Snake

Museum specimens: BMNH 1978.2084, 1978, Kuwait,

leg. S. M. Eissa.

LSUMZ-Herps 89540, 9, 21.6. 1995, Salmiya, Kuwait

Bay, 2.8 km W. of Ras-Al-Ardh, leg. J. Bishop. LSUMZ-

Herps 89541, 27.11.1993, Failaka Island near Auha Is-

land, leg. J. Bishop.

Previous records: Kuwait (Eissa & El Assy 1975).

Remarks: The Yellow-bellied Sea Snake occurs across

the Indo-Pacific, from east and south of Africa as far as

the western coasts of North and South America (Heat-

wole 1999). Within the Arabian Gulf, it was recorded

from the waters of all Gulf States (Castilla et al. 2017).

Hydrophis viperina (Schmidt, 1852)

Viperine Sea Snake

Museum specimens: LSUMZ-Herps 43338, 9,

24.5.1983, Kuwait, leg. J. Bishop.

Remarks: This sea snake occurs in the Indian Ocean,

extending from the Arabian Gulf to the Malay Archipel-

ago (David & Ineich 1999). Within the Arabian Gulf, it

was recorded from the waters of Bahrain, Qatar, Saudi

Arabia, and the United Arab Emirates (Castilla et al.

2017; Buzas et al. 2018). This museum record represents

the first documentation of the Viperine Sea Snake from

the Kuwaiti waters.

DISCUSSION

Eissa & El Assy (1975) reported a total of 28 species of

reptiles from Kuwait, however, three species may rep-

resent misidentifications (Uromastyx thomasi, Agama

Jayakari and Hemidactylus turcicus). The first two spe-

cies are known from Oman and eastern Arabia, while

H. turcicus occurs in Turkey and southern Europe. The

finding of D. jugularis in Kuwait by Al-Mohanna et al.

(2007) represents its most southern range of distribution

in its eastern range, and signifies a relict population. It

was recorded from Al-Kufa district, southern Iraq (Rhadi

et al. 2017).

The herpetofauna of Kuwait consists of three ele-

ments; Arabian species such as Acanthodactylus ophe-

odurus, Acanthodactylus schmidti, Cerastes gasperettii,

Diplometopon zarudnyi, Eryx jayakari and Scincus mi-

tranus, and to some extent Iranian and/or oriental species

exemplified by Platyceps ventromaculatus, Phrynoceph-

alus arabicus, Phrynocephalus maculatus, and Stenod-

actylus affinis. Many other species have a wide range of

distribution across North Africa to the Arabian Peninsula

and beyond (e.g., Acanthodactylus boskianus, Cyrtopo-

dion scabrum, Varanus griseus and Uromastyx aegyptia).

Some species represent extension range of eastern Ara-

Bonn zoological Bulletin 70 (2): 253-272

Zuhair S. Amr et al.

bian forms such as Hemidactylus persicus, Pseudocer-

amodactylus khobarensis and Trigonodactylus arabicus.

Records of reptiles from two Kuwaiti islands are of

special interest. Bunopus tuberculatus was found in both

Auhha and Kubbar islands, while Cyrtopodion scabrum,

H. flaviviridis, Eryx jayakari, Phrynocephalus macula-

tus and Cerastes gasperettii were collected from Auhha

Island. Both islands (Kubbar and Auhha) are very small

with an area of 0.11 and 0.35 km? respectively. Soorae

(2004) reported nine species of reptiles (five geckos, one

lizard and three snakes) from 13 islands in the Arabian

Gulf belonging to the United Arab Emirate (UAE). He

attributed the presence of these reptiles to anthropogenic

activities. Bourquin (2011) recorded thirteen species of

reptiles from Sir Bani Yas Island (UAE), including eight

geckos, two skinks and three snakes. The reptiles of the

Kuwaiti islands should be investigated both at the organ-

ismal and molecular levels.

Gaps in collecting areas are evident as indicated in

Fig. 1. Very few or no data are available from the west-

ern and southwestern parts of Kuwait. The urgent priority

is to conduct baseline herpetological surveys in the pro-

tected areas of Kuwait. Future studies should address the

threats facing the reptiles in this fast developing country

with urban and industrial expansion.

Acknowledgments. We extend our thanks to the IUCN-ROWA

and the Environment Public Authority of the State of Kuwait

for their support through the project “Monitoring and documen-

tation of biodiversity in Kuwait’. Our thanks are also extended

to Mr. Hanna Haddad for map preparation. The authors express

their gratitude for Mr. A. Al Mansi, Mr. Ahmad Ragab, Mr. Ab-

dul Aziz Al Yousef, Mr. Balazs Buzas and Dr. Csaba Géczy for

providing images indicated in the manuscript. Special thanks

are extended to Prof. David Warrell for improving the English.

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Bonn zoological Bulletin 70 (2): 273-279

2021 - Mabrouki Y. et al.

https://do1.org/10.20363/BZB-2021.70.2.273

ISSN 2190-7307

http://www.zoologicalbulletin.de

Research article

urn:|sid:zoobank.org:pub:69B780CD-F787-4729-8A40-5D1433FCA15A

Further records of freshwater Gastropods

(Mollusca: Hydrobiidae, Lymnaeidae, Planorbidae) from Morocco

Youness Mabrouki' *, Abdelkhaleq Fouzi Taybi? & Peter Gléer*

' Sidi Mohamed Ben Abdellah University, Faculty of Sciences Dhar El Mehraz, Biotechnology,

Conservation and Valorisation of Natural Resources laboratory, Fez, Morocco

? Université Mohammed Premier, Faculté Pluridisciplinaire de Nador,

Equipe de Recherche en Biologie et Biotechnologie Appliquées. Morocco

3Schulstr, 3, D-25491 Hetlingen, Germany

“Corresponding author: Email: younes_mab@hotmail.fr

'urn:|sid:zoobank.org:author:9FE7D 1 12-2607-4855-A BF7-9BFOBA38B8C8

2urn:Isid:zoobank.org:author:358C248 1 -FAD9-4524-A EFC-E1 8008EEO4AC

> urn:|sid:zoobank.org:author:8CB6BA7C-D04E-4586-BA 1 D-72FAFF54C4C9

Abstract. Galba oblonga (Puton, 1847), Stagnicola fuscus (Pfeiffer, 1821) and Ancylus striatus Quoy & Gaimard, 1834

are reported for the first time in North Africa, while Mercuria globulina (Letourneux & Bourguignat, 1887) is new to

Morocco. In addition, we provide new distributional data on the two Moroccan endemic and recently described species

Aghbalia aghbalensis Gloer, Mabrouki & Taybi, 2020 and Mercuria bakeri Gloer, Boeters & Walther, 2015 known previ-

ously from the type localities only, which is a key element in promoting their conservation.

Key words. New records, North Africa, additional information, endemic species.

INTRODUCTION

Thanks to its geographical position, Morocco is one of

the most interesting biogeographical regions in the West-

ern Mediterranean. It represents a contact area between

Europe and Africa and a compulsory passage for much of

the fauna between the Palaearctic and Afrotropical region

and between the Mediterranean and the Atlantic. Within

the Maghreb, Morocco possesses the most extensive lo-

tic system (FAO 2003). The precipitation that falls in the

high mountain ranges of the Rif, Middle Atlas, High Atlas

and Anti-Atlas feeds rivers generally flowing north-east-

ern to the Mediterranean, north-westward to the Atlantic

or south-eastward towards the Sahara. These ecosystems

provide a variety of ecological habitats, promoting great

biodiversity while at the same time offering significant

socioeconomic benefits (Chillasse et al. 2001; Taybi et al.

2020a).

Freshwater molluscs play a vital role in freshwater eco-

systems, as they are essential to the maintenance and bal-

ance of aquatic biota, primarily through their control of

water quality and nutrient balance through filter-feeding

and algal-grazing and, to a lesser degree, as a food source

for predators including a number of species of vertebrates

(Oscoz et al. 2014; Tachet et al. 2020). There are an es-

timated 7,000 valid species around the world and about

10,000 more await description (Darwall et al. 2005). Un-

Received: 26.10.2020

Accepted: 24.06.2021

fortunately, these organisms are one of the most threat-

ened freshwater taxa (Kay 1995).

Freshwater molluscs show a high degree of regional

endemism, with about 80% of species endemic to the

North African region (Garcia et al. 2010), which gives

them a privileged place for taxonomical and ecological

studies. Previously, only 52 species of freshwater mol-

luscs were known with certainty from Morocco (Dakki

et al. 1997). However, this list was not complete and

many new species were added recently, including species

new to science (Boulaassafer et al. 2018, 2020; Ghamizi

2020; Gloer et al. 2020a, b; Mabrouki et al. 2020a, 2021).

In order to increase knowledge on the freshwater Mal-

acofauna of Morocco, different field expeditions have

been conducted since 2014 through the northern part of

the country, including its great natural barriers such as the

Moulouya River basin and the Middle Atlas massif.

MATERIAL AND METHODS

In order to promote knowledge on the freshwater mol-

luscs of Morocco, several field expeditions have been

conducted since 2014 in the northern part of the country

(Fig. 1), with a focus on the Middle Atlas and the Mou-

louya River basin. More than 100 localities have been

investigated and most of these sampling sites were visit-

Corresponding editor: B. Huber

Published: 05.07.2021

274 Youness Mabrouki et al.

Oe i 4

: , ey, ees

1 Pt Lae el Og %

y H Ker

w/ i te - 4

BK Newer en nne= 1

oy po

O | ay 4

sy a 4°30°0"W 3°0'0"W

2/20

“ye

bad /

4°0'0"W

3°0'0"W

2°0'0"W

1°30'0"W

36°0'0"N

35°0'0"N

34°30'0"N

e Aghbalia aghbalensis ©

a Mercuria bakeri A

¢ Mercuria globulina

% Stagnicola fuscus

33°0'0"N

«x Ancylus striatus

Fig. 1. Distribution range of the recorded species in Morocco (new records black, old records white).

ed at least three times (e.g., Taybi et al. 2017; Mabrouk1

et al. 2020b for more details on the localities). The sam-

ples of benthic fauna including molluscs were collect-

ed using kick nets or entomological forceps or by hand.

The dissections and measurements of the genital organs

and the shells were carried out using a stereo microscope

(ZEISS); the photographs were made with a digital cam-

era system (Leica R8). The map was made using ArcGIS

software. Voucher specimens were deposited in the col-

lections of the authors. Identifications were made using

the identification key of Gloer (2019).

RESULTS

Galba oblonga, Stagnicola fuscus and Ancylus striatus

are new to North Africa; Mercuria globulina is reported

for the first time in Morocco. In addition, the two recently

described Moroccan endemic species Aghbalia aghbal-

ensis and Mercuria bakeri are reported for the first time

outside their type localities.

Bonn zoological Bulletin 70 (2): 273-279

Family Hydrobiidae Stimpson, 1865

Genus Aghbalia Gloer, Mabrouki & Taybi, 2020

Aghbalia aghbalensis Gloer, Mabrouki & Taybi, 2020

(Fig. 2)

Distribution. This recently described Moroccan en-

demic was known only from its type localities Aghbal

and Tiffert springs (Gloer et al. 2020a). We record it for

the first time in the springs of Ain Sfa and Guercif, in

addition to the potamal section of Moulouya River at a

place named Safsaf (Fig. 1), increasing considerably re-

cords of its distribution area, which is a key element in

promoting its conservation.

Sampling site. Guercif (07/09/2020) 33°57°33” N,

3°30 00. 8) W; Aval Safsaf (19/06/2019)

34°54’°27.53” N, 2°3878.86” W; Ain Sfa (16/06/2019)

34°45712”N 2°8’36” W.

Habitat. The species was known to occur in rheocren-

ous springs only. We document its presence in a big river

(Fig. 8). However, the section of the Moulouya where

the species was found is supplied by various freshwater

springs, which may explain its presence in this habitat.

Associated species. Theodoxus sp.; Galba truncatula

(Miller, 1774); Melanopsis praemorsa (Linnaeus, 1758).

©ZFMK

Additions to the freshwater Gastropod fauna of Morocco 275

Figs 2—7. Shells of the recorded species in Morocco. 2. Aghbalia aghbalensis. 3. Mercuria bakeri. 4. M. globulina. 5. Galba oblon-

ga. 6. Stagnicola fuscus. 7. Ancylus striatus.

Genus Mercuria Boeters, 1971

Mercuria bakeri Gloer, Boeters & Walther, 2015 (Fig. 3)

Distribution. The Moroccan endemic species was

known only from its type locality at the Tingitane Pen-

insula in northwestern Morocco (Gloer et al. 2015). We

record it for the first time in the Oriental Region of the

country at different localities (Fig. 1), increasing there-

fore its distribution range.

Sampling sites. Oued Messoussate (10/06/2020)

35°04’02.8” N, 2°54’11.6” W; Bouareg plain well 1

(28/06/2020) 35°06712.3” N, 2°50’25.8” W; Bouareg

well 2 (28/06/2020) 35°06’25.2” N, 2°50°14.8” W;

La’azib (10/07/2020) 35°03714.8” N, 2°57°27.9” W;

Aroui (14/07/2020) 35°01°07.2” N, 2°59°37.9” W.

Habitat. Previously, M. bakeri was known inhabiting

only a swampy area around a small artificial pond in the

Tingitane Peninsula. During the study period, the species

was collected from a watercourse and spring of low al-

Bonn zoological Bulletin 70 (2): 273-279

titude, it was also sampled from wells, which is a new

feature registered for the species (e.g., Fig. 9).

Associated species. Physella acuta (Draparnaud,

1805).

Mercuria globulina (Letourneux & Bourguignat, 1887)

(Fig. 4)

Distribution. This Maghrebian species was previously

known from Algeria and Tunisia only, where it was re-

stricted to the central and eastern parts of the Tell (Gloer

2019; Gloer et al. 2010). We record it for the first time

in Morocco from the Oriental Region. During the sam-

pling period, the species was collected at a place named

Leghriba, belonging to the Selouane River basin (Fig. 1).

Sampling site. Leghriba (23/07/2020) 35°04’25.3” N,

2°58’42.1” W.

Habitat. This species usually occupies springs and

small rivers. In Morocco, it was collected from a single

©ZFMK

276

well, which is also a new feature documented for the spe-

cies.

Family Lymnaeidae Lamarck, 1799

Galba oblonga (Puton, 1847) (Fig. 5)

Distribution. Whether this taxon is a good species, as

proposed by Vinarski & Kantor (2016: 314), is unclear.

Though the type locality of G. oblonga is in France, it

is listed in the West European literature as a form of

G. truncatula (Gloer 2019: 219), or as a good species

(see WORMS MolluscaBase 2019). We collected only

one empty shell so the anatomy as well as the molecu-

lar genetic could not be studied to solve the taxonomic

problem. We list this taxon here to direct attention to it

for future investigations. In Morocco, G. oblonga was

collected at Selouane (Fig. 1).

Sampling — site. Saguia

35°04536:95 N. 2°55: 23:9" W.

Habitat. In the study area, it was collected from an

irrigation canal system, which routes the waters of the

Moulouya River through the Ouled Settout reservoir

(Fig. 10).

Associated species. Stagnicola fuscus (Pfeiffer, 1821);

Physella acuta, Melanopsis praemorsa.

Selouane (21/08/20)

Stagnicola fuscus (Pfeiffer, 1821) (Fig. 6)

Distribution. The full range of this species is not

known, but is likely to be found throughout Western Eu-

rope (Gloer 2019). We report it here for the first time in

Morocco and North Africa. It was collected at Selouane

(Fig. 1), which is the new confirmed southern limit of its

distribution range in the Palearctic. The species can be

identified by the short phallotheca and the two prostate

folds by which it is separated from all other Palaearctic

Stagnicola spp.

Stagnicola fuscus is one of five European species of

the Stagnicola palustris (Miller, 1774) species complex

(Jackiewicz 1993). Stagnicola palustris was the only

known species of the genus occurring in Morocco, there-

fore, its distribution in the country and surrounding area

must be re-examined, since identification was formerly

based probably on the shells only.

Sampling — site. Saguia

35°04’36.9” N, 2°55’23.9” W.

Habitat. This species prefers the mountainous regions.

It frequents the standing waters or hydrosystems with a

very low flow, often among the algae and macrophytes;

it can also colonize artificial pipes and semi-temporary

pools as in the study area, where it was found in an irri-

gation canal system supplied by Ouled Settout reservoir

(Fig. 10).

Associated species. Galba oblonga; Physella acuta,

Melanopsis praemorsa.

Selouane (21/08/20)

Bonn zoological Bulletin 70 (2): 273-279

Youness Mabrouk‘i et al.

Family Planorbidae Rafinesque,, 1815

Ancylus striatus Quoy & Gaimard, 1834 (Fig. 7)

Distribution. Previously, the genus Ancylus Miller,

1774 was represented by only A. fluviatilis (Muller, 1774)

in Morocco, where it is supposed to be widely distributed

(Taybi et al. 2017). However, since we record for the first

time A. striatus in Morocco and North Africa, the status

and range distribution of A. fluviatilis should be re-ex-

amined. Ancylus striatus was described initially from the

Canary Islands (Quoy & Gaimard 1834), and it was re-

corded later from the Azores Island and the northeastern

Iberian Peninsula (Pfenninger at al. 2003; GBIF 2019).

During the sampling period, the species was collected

at Krouchen and Guercif, corresponding to the oriental

slope of the Middle Atlas Massif, which is part of the

Moulouya River Bassin (Fig. 1).

Sampling site. Krouchen (13/06/2014) 32°44’°49.6” N,

5°10°17” W; Guercif (07/09/2020) 33°57°35.8” N

3°30°40.4” W.

Habitat. This species 1s rheophilous, preferring mod-

erate to fast-flowing, well-oxygenized rivers and streams

(Fig. 11), usually on solid substrate and rocky bottoms.

Remark. Pfenninger et al. (2003: 2742) stated that the

Ancylus sp. (clade 3) found in the Canary islands and

Morocco as well could be Ancylus striatus, while Al-

brecht et al. (2006, Appendix I) thought it could be An-

cylus pileolus Férrussac, 1822, mentioned by Hubendick

(1972: 110), who listed A. striatus and A. pileolus (from

Scio Island) both as synonyms of A. fluviatilis. A. stria-

tus and A. pileolus are both listed as accepted species by

WORMS (2019). This problem cannot be solved here,

however, it 1s possible to understand which species is

meant since we refer to the identification key of Gloer

(2019) where A. striatus has been described and depicted.

Associated species. Anisus sp; Galba truncatula, Phy-

sella acuta, Aghbalia aghbalensis, Melanopsis praemor-

Sd.

DISCUSSION

High-quality data on species identity and distribution

are a good tool for biodiversity management and con-

servation. It allows biologists and managers to come up

with meaningful action steps and make wise decisions

to achieve biodiversity protection (Guralnick et al. 2007;

Hortal et al. 2007). Combining these data with various

environmental predictors and different parameterisation

techniques, it is possible to make predictions about the

distribution of species, especially of endemic ones (Mil-

lan et al. 2014).

The inventory list of the freshwater mollusc fauna of

Morocco is certainly still incomplete. Unfortunately, due

to the great pressure on continental aquatic ecosystems,

©ZFMK

Additions to the freshwater Gastropod fauna of Morocco 277

Figs 8-11. 8. Big section of the Moulouya River. 9. Typical well in the study area. 10. View of Ouled Settout reservoir. 11. Water-

course from the Middle Atlas.

many species will probably have disappeared before be-

ing recorded or even described. Indeed, as in most of the

Maghrebian territory, the regional aquatic ecosystems are

increasingly threatened by human activities, through wa-

ter abstraction, habitat loss and modification, industrial

effluents, domestic sewage and agricultural runoff in-

cluding fertilisers, pesticides and drainage of water (Ben-

saad et al. 2017; Mabrouki et al. 2017; Taybi et al. 2016,

2020b). Even worse, alien invasive species, potentially

able to change the current biotic interactions in benthic

communities, have been recorded recently from the hy-

dro-systems of Morocco (Mabrouki et al. 2019a, b, c,

2020c; Taybi et al.2020c, d, e), including molluscs, such

as the Asian clam Corbicula fluminea (Miller 1774) and

the New Zealand mudsnail Potamopyrgus antipodarum

(Gray, 1843) (Taybi et al. 2017; Taybi et al. 2021). These

invasive species could be a serious problem for the native

aquatic biodiversity. Therefore, monitoring the presence

and distribution of species, as well as studies improving

their biological and ecological knowledge, are of crucial

concern in promoting the conservation of the Moroccan

freshwater biodiversity.

Bonn zoological Bulletin 70 (2): 273-279

Acknowledgements. We cordially thank the editor and review-

ers, whose helpful and sincere comments have improved the

manuscript.

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BHL

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Bonn zoological Bulletin 70 (2): 281-315

2021 - Bugaj-Nawrocka A. et al.

https://do1.org/10.20363/BZB-2021.70.2.281

ISSN 2190-7307

http://www.zoologicalbulletin.de

Research article

urn:lsid:zoobank.org: pub: DAB40723-C66E-4826-A 8F7-A678A FABA 1 BC

Scale insects (Hemiptera: Coccomorpha)

in the entomological collection of the Zoology Research Group,

University of Silesia in Katowice (DZUS), Poland

Agnieszka Bugaj-Nawrocka"*, Lukasz Junkiert?, Malgorzata Kalandyk-Kolodziejezyk*? & Karina Wieczorek*

2.34 Faculty of Natural Sciences, Institute of Biology, Biotechnology and Environmental Protection,

University of Silesia in Katowice, Bankowa 9, PL-40-007 Katowice, Poland

“Corresponding author: Email: agnieszka.bugaj-nawrocka@us.edu.pl

'urn:Isid:zoobank. org:author: BSA9DF 15-3677-4F5C-A DOA-46B25CA350F6

2urn:Isid:zoobank.org:author: AF78807C-2115-4A33-AD65-9190DA612FB9

3urn:Isid:zoobank.org:author:600C5C5B-38C0-4F26-99C4-40A4DC8BBO016

*urn:Isid:zoobank.org:author:95A5CB92-EB7B-4132-A 04E-6163503ED8C2

Abstract. Information about the scientific collections is made available more and more often. The digitisation of such

resources allows us to verify their value and share these records with other scientists — and they are usually rich in taxa

and unique in the world. Moreover, such information significantly enriches local and global knowledge about biodiversi-

ty. The digitisation of the resources of the Zoology Research Group, University of Silesia in Katowice (Poland) allowed

presenting a substantial collection of scale insects (Hemiptera: Coccomorpha). The collection counts 9369 slide-mounted

specimens, about 200 alcohol-preserved samples, close to 2500 dry specimens stored in glass vials and 1319 amber inclu-

sions representing 343 taxa (289 identified to species level), 158 genera and 36 families (29 extant and seven extinct). A

significant part is the collection of an outstanding Polish coccidologist, Professor Jan Koteja. The geographical analysis

of the scale insects collected shows mainly Eurasian areas, but there is also material from North America, South America,

as well as Africa and New Zealand.

Keywords. Alpha taxonomy, biodiversity conservation, Coccoidea, Sternorrhyncha.

INTRODUCTION

There are many natural history collections, both private

and in museums, and all of them are extremely import-

ant for the knowledge of biodiversity. Information about

which specimens are in each collection is now more acces-

sible than ever. Many natural history museums and other

institutions curating scientific collections provide such

information on websites as databases. They also publish

articles from which we can learn a lot about the details of

these collections (like, for example, those devoted to dif-

ferent groups of insects: Wijesekara & Wijesinghe 2003;

Lampe et al. 2007; Notton 2007; Gonzalez et al. 2013;

Franquinho Aguiar & Carvalho 2016; Borowski & Singh

2017; Zaragoza-Caballero & Pérez-Hernandez 2017; Ca-

ballero et al. 2020; Zahiri et al. 2021).

Scale insects (Hemiptera: Coccomorpha) constitute

the superfamily Coccoidea Handlirsch, 1903 within the

infraorder Coccomorpha Rtbsaamen, 1899 and subor-

der Sternorrhyncha Amyot & Serville, 1843 (Gullan &

Cook 2007; Williams & Hodgson 2014). There are 55

families within scale insects, 35 extant (1213 genera

and 8400 species) and 20 extinct (52 extinct genera and

Received: 12.04.2020

Accepted: 24.06.2021

86 species) (Garcia Morales et al. 2016). It is a significant

group of insects due to its economic impact worldwide

— many species of these plant-feeding hemipterans are

considered serious pests of agriculture, horticulture and

forestry, and vectors of plant diseases (Gullan & Martin

2009; Garcia Morales et al. 2016).

The collection in the Zoology Research Group, Univer-

sity of Silesia in Katowice is composed mainly of scale

insect samples collected by such specialists as de Boer,

Borchsenius, Danzig, Kalandyk-Kotodziejczyk, Kawec-

ki, Koteja, Kerzhner, Kosztarab, Lagowska, Nur, Podsi-

adito, Simon and Zak-Ogaza, and is one of the largest and

most important in Poland. Thanks to the project Digital

Database of Entomological Specimens HEMIPTERON

(CEBD HEMIPTERON), this collection has been already

digitised, and the results of our work are presented here.

A major part of this scale insects collection belonged

to Professor Jan Koteja — an outstanding Polish entomol-

ogist, who made extraordinary contributions to cocci-

dology. He was born on the 17" of September 1932 in

Siemianowice Slaskie, Poland. He graduated from the

Faculty of Biology and Earth Sciences of the Jagiello-

nian University and worked at the University of Agri-

Corresponding editor: M. Espeland

Published: 05.07.2021

282 Agnieszka Bugaj-Nawrocka et al.

culture in Krakow until his retirement (Wegierek 2005).

Inspired by Professor Zbigniew Kawecki and Doctor

Barbara Zak-Ogaza, he became interested in coccoids.

Professor Koteja conducted comprehensive research on

scale insects, including faunistics, morphology, taxono-

my, palaeontology, and phylogeny (Wegierek 2005; Dz-

iedzicka & Podsiadto 2007), and is considered a pioneer

of scale insect palaeontology (Wegierek 2005; Dziedzic-

ka & Podsiadto 2007; Gullan 2008). As Gullan (2008)

emphasised, the genius of his work was manifested in the

combination of knowledge about extant scale insect spe-

cies with a thorough understanding of fossils. As a result

of enormous, careful work, he described extant and ex-

tinct taxa new to knowledge. All his species descriptions

are detailed and precisely illustrated. He passed away on

the 19" of August 2004 (Wegierek 2005; Dziedzicka &

Podsiadto 2007). His research has had a significant in-

fluence on the development of coccidology. According to

Professor Koteja’s will, his collection of scale insects, in-

cluding microscope slides, dry material, specimens pre-

served in alcohol, amber inclusions as well as his book

collection was donated to the Faculty of Biology and En-

vironmental Protection, Department of Zoology, Univer-

sity of Silesia in Katowice (DZUS), currently Zoology

Research Group, Institute of Biology, Biotechnology and

Environmental Protection, Faculty of Natural Sciences.

MATERIAL AND METHODS

Professor Koteja’s donation is the richest part of the scale

insects collection of the DZUS. However, in our collec-

tion are also deposited specimens collected by numerous

international hemipterists as well as by specialists and

students from the University of Silesia in Katowice.

Slide-mounted samples from Professor Koteja’s collec-

tion were made according to his method (Koteja 1996):

1. Specimens were placed in 10% potassium hydroxide

(KOH) or sodium hydroxide (NaOH) at room tempera-

ture for a varying period: from a few hours up to a few

days. The solution was replaced with a new 10% KOH

or 10% NaOH during maceration. In order to shorten

the procedure to a few minutes, a solution can be heat-

ed in a water bath (the solution should not be boiled).

2. Specimens were placed on watch glass (in 10% KOH

or NaOH, later in distilled water or 96% ethanol) and

gently pressed to remove all body contents. To purify

the specimens, they were placed in a saturated sodium

carbonate solution and then transferred to acetic acid.

The procedure can be repeated several times until the

specimens were cleaned;

3. Specimens were transferred to a solution of 96% etha-

nol and acetic acid. The proportions of ethanol and acid

were of little importance. A few drops of a saturated

acid fuchsin solution in 96% ethanol and acetic acid

Bonn zoological Bulletin 70 (2): 281-315

were added. Specimens were placed in this dye solu-

tion for half an hour to several hours;

4. Individuals were placed in 96% ethanol, then in 100%

ethanol for several minutes. If the specimens were

large and numerous, the procedure should be repeated;

5. Insects were transferred to clove oil or cedarwood oil,

in which they should not be stored for more than a few

days;

6. Single specimens with their ventral surface upwards

were mounted in Canada balsam under a cover glass.

The additional specimens in the collection were col-

lected by carefully combing through the host plants,

turf, and the surface layers of the soil by hand. Species

identification of all specimens collected between 2005

and 2008 was confirmed by Professor Elzbieta Podsi-

adto (Warsaw University of Life Sciences - SGGW).

The insects were preserved in 70% ethanol and mount-

ed according to the method described by Williams &

Kosztarab (1972) and further modified by Lagowska

(1996). Minor changes were made following Kalan-

dyk & Wegierek (2010):

1. Specimens were placed in 10% potassium hydroxide

(KOH) at room temperature for a varying period: from

a single day up to two weeks (the time was longer in

the case of individuals characterised by strongly scle-

rotised cuticula, e.g., some females of the family Coc-

cidae);

2. The body contents were teased out by gently pressing

the specimens with a small wooden spatula (during the

process, insects were kept in 10% KOH solution);

3. Transparent specimens were transferred to distilled

water for about 10 minutes;

4. The insects were kept in glacial acetic acid for at least

2 hours (up to 12 hours):

5. The specimens were stained in a dye solution for 1—2

hours (such a solution was obtained by adding enough

acid fuchsin to the glacial acetic acid to make the solu-

tion dark pink);

6. Individuals were transferred to 95% ethanol for

10 minutes, then to 99.9% ethanol for 5 minutes;

7. Specimens were kept in clove oil for 1—2 hours (up to

a few days);

8. Single specimens with their ventral surface upwards

were mounted in Canada balsam under a cover glass;

9. The slides were placed in a drying oven at 37°C for

about two weeks.

Identifications were carried out using a Nikon Eclipse

E600. Slide-mounted samples were scanned using a Ep-

son Perfection V850 Pro scanner with a resolution of

800 dpi and edited in PhotoScape X ver. 4.1.1. Pictures

of amber inclusions were taken with a Nikon D750 and

edited in Helicon Focus ver. 7.6.6.

The scale insects database was created in Microsoft

Excel 2017® software and Microsoft Access 2017®

software. The database has the following fields: unique

©ZFMK

Scale insects of DZUS collection 283

ID, collector, gathering method, identifier, sex or devel-

opment stage of specimens, preparation type, number of

individuals of a given record, geolocation data (locali-

ty, latitude, longitude, altitude and country ISO 3166

code), date of collection of the sample (yyyy-mm-dd),

taxonomic information (family, genus, species, author of

the species name — old and current names), associated

botanical taxa (family, genus, species, author of the spe-

cies name). All taxonomic changes (like synonymisation

that have already been proposed) have been included.

The database will be available at

A world map with the countries where the collected

material comes from was created in Microsoft Excel

2019® software.

Labels of type specimen material

All data are rewritten as they appeared on the respective

labels. Double slash (//) separates data on different la-

bels; a single slash (/) separates lines within each label:

additional comments are placed in square brackets.

RESULTS AND DISCUSSION

The collection of coccoids at the Zoology Research

Group, University of Silesia in Katowice (DZUS) in

Poland preserves more than 13.300 curated specimens

stored as 9369 slide-mounted specimens, about 200 al-

cohol-preserved samples, close to 2500 dry specimens in

glass vials and 1319 amber inclusions. In total, all the

samples mentioned above were identified as belonging

to 343 taxa (289 identified to species level, 54 identified

to genera level), 158 genera and 36 families (29 extant

and seven extinct). Among the material found in amber

inclusions, 474 specimens remain unidentified.

All slide-mounted samples are stored in wooden boxes

in specially adapted cabinets (Fig. 1). The liquid vouch-

er specimens are preserved in 70% ethanol (Fig. 2A—B),

dry samples are stored in glass vials in wooden boxes

Fig. 1. Scale insects (Hemiptera: Coccomorpha) samples deposited in the Zoology Research Group, University of Silesia in Ka-

towice (DZUS) collection. A. Specially adapted cabinets for wooden slide holders. B. A close-up on one of the shelves on which

boxes with slides are systematically placed. C. Wooden slide holder with samples curated.

Bonn zoological Bulletin 70 (2): 281-315

©ZFMK

284 Agnieszka Bugaj-Nawrocka et al.

Fig. 2. Scale insects (Hemiptera: Coccomorpha) samples deposited in the Zoology Research Group, University of Silesia in Ka-

towice (DZUS) collection. A-B. Small containers with specimens preserved in 70% ethanol. C. Glass vials with dry samples.

D. Example of stored amber inclusions.

(Fig. 2C), and amber inclusions are stored in foil bags

with a description of the sample (Fig. 2D). The scanned

specimens were additionally provided with a sticker with

a unique ID, which allows them to be easily located in the

database (Fig. 3).

Within slide-mounted specimens, 272 slides are rep-

resented by males (361 individuals), 7513 are represent-

ed by females (9612 individuals), 1334 represent larvae

(5995 individuals), 63 represent prepupae (95 individu-

als), 58 represent pupae (96 individuals), and 41 repre-

sent eggs (906 individuals). In turn, among amber inclu-

sions, 938 are represented by males (1282 individuals),

266 are represented by females (358 individuals), and

97 represent larvae (227 individuals). Professor Koteja

identified all determined species from amber inclusions.

Table 1 (Appendix I) lists all identified taxa, sex or

stage, host plants, and countries in which they were

collected. Species known only from greenhouses are

also marked. The material was collected over the years

1890-2019. It comes from many places across the world

but is dominated by material from Europe and Asia

Bonn zoological Bulletin 70 (2): 281-315

(Fig. 4). The highest diversity is represented by species

of Coccidae Stephens, 1829 (23.6%) and Pseudococci-

dae Heymons, 1915 (23.6%), followed by Diaspididae

Maskell, 1878 (14.3%) and Eriococcidae Cockerell,

1899 (11.4%). Only few species represent the remain-

ing families (Fig. 5 — only extant families are shown).

As males are collected much less frequently than females

and larvae, figure 6 present the genera and species of

male specimens in the collection. Males were recorded

from 24 genera (31 species) of plants in 16 families. On

the other hand, slide-mounted females were recorded

from 168 genera (312 species) of host plants in 65 fam-

ilies. As many as 34.4% of the slide-mounted samples

are specimens collected from grasses (Poaceae (R. Br.)

Barnh.). The species of the genera Festuca L., Agros-

tis L., Calamagrostis Adans., Poa L., Deschampsia P.

Beauv., Brachypodium L. and Elymus L. (all Poaceae)

dominate among the grasses. In the second place, with a

20.7% share, are plants from the Cyperaceae Juss., where

species of the genera Carex L. and Eriophorum vagina-

tum L. have been identified. The third place (11.5%) is

©ZFMK

Scale insects of DZUS collection 285

Fig. 3. Slides of curated scale insects (Hemiptera: Coccomor-

pha) samples deposited in the Zoology Research Group, Uni-

versity of Silesia in Katowice (DZUS) collection. A-B. Exam-

ples of how slide-mounted samples are described — data about

species, collection site, date, host plant and persons responsible

for collection and identification. C. Arranging samples in the

box — view from the unique ID label side.

taken by host plants from the Ericaceae Juss., where spe-

cies from the genera Arctostaphylos Adanson, Calluna

Salisb., Dracophyllum Labill., Empetrum L., Erica L.,

Rhododendron L. and Vaccinium L. were recorded. In

turn, the species with the highest number of record-

ed host plants are Trionymus perrisii (Signoret, 1875)

(42 plant species from 20 genera), Kaweckia glyceriae

(Green, 1921) (37 plant species from 18 genera), Par-

thenolecanium corni (Bouché, 1844) (29 plant species

from 24 genera), Phenacoccus aceris (Signoret, 1875)

(29 plant species from 16 genera), Lepidosaphes ulmi

(Linnaeus, 1758) (27 plant species from 20 genera), 7ri-

onymus aberrans Goux, 1938 (27 plant species from 16

genera), 7rionymus thulensis Green, 1931 (22 plant spe-

cies from 12 genera), Anophococcus insignis (Newstead,

1891) (21 plant species from 16 genera), Anophococcus

pseudinsignis (Green, 1921) (19 plant species from 15

genera), Phenacoccus phenacoccoides (Ben-Dov, 1994)

(19 plant species from 15 genera), Saccharicoccus is-

farensis (Borchsenius, 1949) (18 plant species from 12

genera), Atrococcus cracens Williams, 1962 (17 plant

species from 12 genera), Ceroputo pilosellae Sulc,

1898 (15 plant species from 10 genera), Heterococcus

Bonn zoological Bulletin 70 (2): 281-315

nudus (Green, 1926) (15 plant species from 13 genera),

and Lecanopsis formicarum Newstead, 1893 (15 plant

species from 9 genera).

Type specimens deposited at the Zoology Research

Group, University of Silesia in Katowice (DZUS)

The Zoology Research Group, University of Silesia in

Katowice (DZUS) collection preserves 49 type speci-

mens stored slide-mounted. Some of them have already

been synonymised, but we mention them because they

once formed the basis for describing a new taxon.

Family Coccidae Fallén, 1814

1. Exaeretopus orientalis Danzig, 1975

Exaeretopus orientalis Danzig, 1975a: Zoologicheskii

Zhurnal 54: 137-138

Type material examined. ¢ 4354 / Exaeretopus / orien-

talis Danz. / 9 type material / Carex pediformis / det. E.

Danzig // 23.VII.1970 / 132-71 / Bocm. Caan, / Apuian

/ leg. E. Jlanmur (DZUS). * 4355 / Exaeretopus / orien-

talis Danz. / 2 type material / Carex pediformis / det:

E. Danzig // 23. VII.1970 / 132-71 / Bocmorusiii Caan /

Apuiau / leg. E. JJanuur (DZUS). * 4356 / Exaeretopus

/ orientalis Danzig / 2 type material / Carex pediformis

/ det: E. Danzig // 23.VII.1970 / 132-71 / Bocmorupiit /

Caau / Apuiau / leg. E. Jlanuur (DZUS).

Type locality. Buryatia Aut. Oblast, Eastern Sayan,

Tunkinsk Ridge, Arshan [Russian Federation]

Current status. Valid species.

Remarks. Holotype (by original designation) is in

Saint Petersburg: Zoological Museum, Academy of Sci-

ence, Russia. Specimens at DZUS collection labelled as

“type material” are considered paratypes.

2. Luzulaspis americana Koteja & Howell, 1979

Luzulaspis americana Koteja & Howell, 1979: Annals of

the Entomological Society of America 72: 334-342

Type material examined. * Luzulaspis / americana /

Koteja et Howell / 2 / paratype / 10 // HHT-231-75 /

On: Unident. Grass / Echols Co. Ga. / X-2-75 / Coll. R.

Beshear (DZUS).

Type locality. Echols County, Georgia [United States

of America]

Current status. Valid species.

Remarks. Holotype (by original designation) is in

Washington: United States National Entomological Col-

lection, U.S. National Museum of Natural History, Dis-

trict of Columbia, USA.

3. Luzulaspis crassispina Borchsenius, 1959

Luzulaspis crassispina Borchsenius, 1959: Entomolog-

icheskoe Obozrenye 38: 164—175

Type material examined. * 1314 / Luzulaspis / crassi-

spina / Borchs. 9 / paratyp / Luzula sp. / det. Borchs //

©ZFMK

286 Agnieszka Bugaj-Nawrocka et al.

Fig. 4. World map showing the countries where the scale insects samples deposited in the Zoology Research Group, University of

Silesia in Katowice (DZUS) collection come from.

18.5.1957 / Junnan / Chiny / leg. Borchsentus (DZUS).

¢ 1315 / Luzulaspis / crassispina / Borchs. @ / paratyp /

Luzula sp. / det. Borchs. // 18.5.1957 / Junnan / Chiny /

leg. Borchsenius (DZUS).

Type locality. Yunnan Province, Kingtung [China|

Current status. Valid species.

Remarks. Holotype (by original designation) is in

Beiing: Institute of Entomology, Academy of Sciences,

China, and was designated from Carex sp.

4. Luzulaspis kosztarabi Koteja & Kozar, 1979

Luzulaspis kosztarabi Koteja & Kozar, 1979: Acta Zoo-

logica Academiae Scientiarum Hungaricae 25: 121-

125

Type material examined. * Tornanadaska / Carex /

1975.VIII.8. // Luzulaspis / kosztarabi / Koteja et Kozar

/ paratype / leg. M. Kosztrab / F. Kozar / 545 (DZUS).

Type locality. Tornanadaska [Hungary]

Current status. Valid species.

Remarks. Holotype (by original designation) 1s in Bu-

dapest: Hungarian Natural History Museum, Zoological

Department, Hungary.

5. Luzulaspis minima Koteja & Howell, 1979

Luzulaspis minima Koteja & Howell, 1979: Annals of the

Entomological Society of America 72: 334-342

Type material examined. * Luzulaspis / minima / Koteja

et Howell / ° / paratypes // Exaeratopus / On Carex /

Eagle Lake Mineral / King Tulare co. Calif. / 10, [illegi-

ble] / Aug. 28 1946 G.F-F. / Stanford University / Natural

History Museum (DZUS).

Bonn zoological Bulletin 70 (2): 281-315

Type locality. Tulare County, Eagle Lake, Mineral

King, California [United States of America|

Current status. Valid species.

Remarks. Holotype (by original designation) is in

Washington: United States National Entomological Col-

lection, U.S. National Museum of Natural History, Dis-

trict of Columbia, USA.

6. Luzulaspis nemorosa Koteja, 1966

Luzulaspis nemorosa Koteja, 1966: Polskie Pismo Ento-

mologiczne 36: 45—56

Type material examined. * 2587 / Luzulaspis / nemoro-

sa / 9 - before ovipos. / holotype / Luzula nemorosa /

det. J. Koteja // 11.7.1962 / Przegorzaty / Krakow / leg. J.

Koteja (DZUS). * 2591 / Luzulaspis / nemorosa / 9 - be-

fore ovipos. / paratype / Luzula nemorosa / det. J. Kote-

ja // 22.7.1962 / Przegorzaty / Krakow / leg. J. Koteja

(DZUS). * 2593 / Luzulaspis / nemorosa / 9 - before

ovipos. / paratype / Luzula nemorosa / det. J. Koteja //

22.7.1962 / Przegorzaly / Krakow / leg. J. Koteja(DZUS).

¢ 2595 / Luzulaspis / nemorosa / 9 - before ovipos. /

paratype / Luzula nemorosa / det. J. Koteja // 22.7.1962

/ Przegorzaly / Krakow / leg. J. Koteja (DZUS). * 2596

/ Luzulaspis / nemorosa / ° - before ovipos. / paratype /

Luzula nemorosa / det. J. Koteyja // 22.7.1962 / Przegorza-

ly / Krakow / leg. J. Koteja (DZUS). * 215 / Luzulaspis

/ nemorosa / 9 - young / paratype / Luzula nemorosa /

det. J. Koteja // 22.7.1962 / Przegorzaty / Krakow / leg.

J. Koteja (DZUS).

Type locality. Przegorzaly, Cracow [Poland]

Current status. Valid species.

©ZFMK

Scale insects of DZUS collection 287

Asterolecaniidae a 9

Beesoniidae u 1

Callipappidae s 1

Cerococcidae — 5

Cocca as mneeaeratasnssaccammmmamicaaans

Coelostomidiidae — 6

Conchaspididae [i 2

Cryptococcidae a 2

Dactylopiidae ai 2

cee Ts 3:

Croco? eens. 37

Kermesidae

Kerridae

Kuwaniidae

Lecanodiaspididae

Marchalinidae

Margarodidae

Micrococcidae

me ;

ee?

Matsucoccidae [INN 10

Monophlebidae [IE s

Ortheziidae Is

| #

Phenacoleachiidae

Phoenicococcidae

Peudecocdee I 75

Putoidae [6

Rhizoecidae [i 3

Steingeliidae be 1

Stictococcidae uw 1

Xylococcidae 4 |

Fig. 5. Summary of the number of species representing a given family of scale insects in the Zoology Research Group, University

of Silesia in Katowice (DZUS) collection.

Bonn zoological Bulletin 70 (2): 281-315 ©ZFMK

288 Agnieszka Bugaj-Nawrocka et al.

7. Luzulaspis pieninica Koteja & Zak-Ogaza, 1966

Luzulaspis pieninica Koteja & Zak-Ogaza, 1966: Acta

Zoologica Cracoviensia 11: 305-332

Type material examined. * 1240 / Luzulaspis / pieninica

/ 2 - holotype / Carex / ornithopoda / det. B. Ogaza J.

Koteya // 17.8.1965 / Cisowa Skala / pow. Nowy Targ /

leg. B. Ogaza J. Koteja (DZUS). ¢ 818 / Luzulaspis / pie-

ninica/ & - paratype / Carex / ornithopoda / det. B. Ogaza

J. Koteja // 8.10.1963 / Macelowa / Gora/ p. Nowy Targ

/ leg. B. J. Koteja Ogaza (DZUS). * 1110 / Luzulaspis /

pieninica / Q - paratype / Carex / ornithopoda / det. B.

Ogaza J. Koteja // 30.7.1964 / Skatka miedzy / Cisowa

i Oblaza / kolo Nowe] Biatej / pow. Nowy Targ / leg.

J. Koteja (DZUS). * 1217 / Luzulaspis / pieninica / 9 -

paratype / Carex / ornithopoda / det. B. Ogaza J. Koteja

// 17.8.1965 / Cisowa Skata / pow. Nowy Targ / leg. B.

Ogaza J. Koteja (DZUS). * 2263 / Luzulaspis / pieninica /

Q - paratype / Carex / ornithopoda / det. B. Ogaza J. Ko-

teya // 17.8.1965 / Cisowa Skala / pow. Nowy Targ / leg.

B. Ogaza J. Koteja (DZUS). * 2264 / Luzulaspis / pieni-

nica / Q - paratype / Carex / ornithopoda / det. B. Ogaza

J. Koteja // 17.8.1965 / Cisowa Skata / pow. Nowy Targ

/ leg. B. Ogaza J. Koteja (DZUS). * 2265 / Luzulaspis

/ pieninica / 9 - paratype / Carex ornithopoda / det. B.

Ogaza J. Koteja // 17.8.1965 / Cisowa Skala / pow. Nowy

Targ / leg. B. Ogaza J. Koteja (DZUS).

Type locality. Cisowa Skala in the Nowy Targ Valley

[Poland]

Current status. Synonym of Luzulaspis grandis

Borchsenius, 1952 (synonymised by Danzig (1980)).

8. Pulvinaria terrestris Borchsenius, 1953

Pulvinaria terrestris Borchsenius, 1953: Entomologich-

eskoe Obozrenye 33: 281—290

Type material examined. ¢ 1363 / Pulvinaria / terrestris /

9 paratype / Ulmus glabra / det. Borchsenius // 13.6.1938

/ Krym / okreg Jaltajski / leg. Kiri¢enko (DZUS). * 1364

/ Pulvinaria / terrestris / 9 paratype / Ulmus glabra / det.

Borchsenius // 13.6.1938 / Krym / okreg Jattajski / leg.

Kiricéenko (DZUS).

Type locality. Vagravar Megrinsk Ridge, Ayrum [Ar-

menia|

Current status. Valid species.

Remarks. Syntypes (by original designation) are in

Saint Petersburg: Zoological Museum, Academy of Sci-

ence, Russia. Specimens at DZUS collection labelled as

“paratype” are considered syntypes, because Borchse-

nius probably did not designate the Holotype.

Family Diaspididae Targioni Tozzetti, 1868

9. Odonaspis fistulata Ferris, 1921

Odonaspis fistulata Ferris, 1921: Stanford University

Publications, Biological Sciences. Palo Alto 1: 61-132

Type material examined. * Odonaspis / fistulate Ferris /

= Circulaspis / 9 / Mtd. From / TYPE MAT /R.F. Wilkey

Bonn zoological Bulletin 70 (2): 281-315

/ -*70 / Punta Pamilla, nr. San / José del Cabo, “Baja /

California, MEXICO / July, 1919 / ex Distichlis spicata /

Coll. GFFerris // VPI & SU Collection (DZUS).

Type locality. Baja California, Punta Palmilla, near

San José del Cabo [Mexico]

Current status. Change of combination by Ferris in

1938 to Circulaspis fistulata (Ferris, 1921)

Remarks. Holotype (by original designation) 1s in The

Bohart Museum of Entomology, University of Califor-

nia, California, USA. Specimen at DZUS collection la-

belled as “TYPE MAT” is considered paratype.

10. Niveaspis fenestrata Ferris, 1941

Niveaspis fenestrata Ferris, 1941: Atlas of the scale in-

sects of North America. Series 3. Stanford University

Press Palo Alto, California

Type material examined. * Niveaspis / fenestrata Ferris /

9 /Mtd. From / TYPE MAT /R.F. Wilkey / -‘70 / Tonila,

Colima, / MEXICO — 1962 / ex Bumelia sp. / Coll. GF-

Ferris MEX. #286 // VPI & SU Collection (DZUS).

Type locality. Colima, Tonila [ Mexico]

Current status. Valid species.

Remarks. Syntypes (by original designation) are in

The Bohart Museum of Entomology, University of Cali-

fornia, California, USA. The specimen at DZUS collec-

tion labelled as “T'YPE MAT” is considered syntype.

11. Velataspis cornigera Ferris, 1941

Velataspis cornigera Ferris, 1941: Atlas of the scale in-

sects of North America. Series 3. Stanford University

Press Palo Alto, California

Type material examined. « Velataspis / cornigera Ferris

/ 2 /Mtd. From / TYPE MAT / R.F. Wilkey / -‘70 / Da-

vid, PANAMA / 1938 / ex unid. shrub / Coll. GFFerris

PAN. #306 // VPI & SU Collection (DZUS).

Type locality. Chiriqui, David [Panama]

Current status. Valid species.

Remarks. Holotype (by original designation) 1s in The

Bohart Museum of Entomology, University of Califor-

nia, California, USA. The specimen at DZUS collection

labelled as “TYPE MAT” is considered paratype.

Family Eriococcidae Cockerell, 1899

12. Acanthococcus sasae Danzig, 1975

Acanthococcus sasae Danzig, 1975b: Entomologich-

eskoe Obozrenye 54: 62-81

Type material examined. « Paratypus [red label] //

143=67 / Acanthococcus / sasae Danzig / Sasa kurilensis

/ o. KyHaummp / CepHosogck / 11.V1.67 r. E. Jjanuur /

4459 (DZUS).

Type locality. Kunashir Island, Sernovodsk [Russian

Federation]

Current status. Valid species.

©ZFMK

Scale insects of DZUS collection 289

Remarks. Holotype (by original designation) is in

Saint Petersburg: Zoological Museum, Academy of Sci-

ence, Russia.

13. Rhizococcus palustris Dziedzicka & Koteja, 1971

Rhizococcus palustris Dziedzicka & Koteja, 1971: Acta

Zoologica Cracoviensia 16: 557—579

Type material examined. * 2582 / Rhizococcus / pal-

ustris / 9 / holotype / 2 / Eriophorum / vaginatum / det.

J. Koteja // 10.8.1963 / Puscizna W. / Piekielnik / pow.

Nowy Targ / Poland / leg. E. Koteja (DZUS). * 736 /

Rhizococcus / palustris / 2 paratype / Eriophorum / va-

ginatum / det. Koteya // 10.8.1963 / Puscizna W. / Pie-

kielnik / pow. Nowy Targ / leg. E. Koteya (DZUS). * 737

/ Rhizococcus / palustris / 11 paratype / Eriophorum

/ vaginatum / det. J. Koteja // 10.8.1963 / Piekielnik /

(ORAWA) / Torfowisko wys. / Puscizna Wielka / leg. J.

Koteja (DZUS). * 759 / Rhizococcus / palustris / ° pa-

ratype / 9 /Eriophorum / vaginatum / det. J. Koteja //

10.8.1963 / Puscizna W. / Piekielnik / pow. Nowy Targ /

leg. E. Koteyja (DZUS). * 2580 / Rhizococcus / palustris /

9 paratype / 9 /Eriophorum / vaginatum / det. Koteja //

10.8.1963 / Puscizna W. / Piekielnik / pow. Nowy Targ /

leg. E. Koteja (DZUS). * 2583 / Rhizococcus / palustris /

9 paratype / 9 /Eriophorum / vaginatum / det. Koteja //

10.8.1963 / Puscizna W. / Piekielnik / pow. Nowy Targ /

leg. E. Koteja (DZUS). * 2585 / Rhizococcus / palustris /

9 paratype / 2 /Eriophorum / vaginatum / det. J. Koteja

// 10.8.1963 / Puscizna W. / Piekielnik / pow. Nowy Targ

/ leg. E. Koteya (DZUS). * 2586 / Rhizococcus / palustris

/ Q paratype / Eriophorum / vaginatum / det. Koteja //

10.8.1963 / Puscizna W. / Piekielnik / pow. Nowy Targ /

leg. E. Koteja (DZUS).

Type locality. Puscizna Wielka, Piekielnik, Nowy Targ

[Poland]

Current status. Valid species.

Remarks. This species was described by Dziedzic-

ka and Koteja (1971) as Rhizococcus palustris and was

transferred to the genus Acanthococcus by Kosztarab and

Kozar (1978) (as Acanthococcus palustris). Many spe-

cies of the genus Acanthococcus were sometimes syn-

onymised with Eriococcus, because of this Dziedzicka

and Koteja (1985) replaced the synonymised name Erio-

coccus palustris with Eriococcus podhalensis as nomen

novum. Miller and Gimpel (1996) proposed a new name,

Acanthococcus dziedzickae, for the homonym Acanthoc-

occus palustris. They did not notice that Dziedzicka and

Koteja (1985) had already changed the name to Eriococ-

cus podhalensis. In the Catalogue of the Eriococcidae

by Miller and Gimpel (2000), this species was listed as

Eriococcus podhalensis. However, Kozar (2009) pro-

posed that the species Rhizococcus palustris should be

considered as appropriate. Kozar et al. (2013) kept the

name. Also, Lagowska and Golan (2020) list this species

as Rhizococcus palustris.

Bonn zoological Bulletin 70 (2): 281-315

Family Margarodidae Cockerell, 1899

14. Neomargarodes chondrillae Archangelskaya, 1935

Neomargarodes chondrillae Archangelskaya, 1935: The

carmine-producing coccids (Margarodes) of Middle

Asia and species of allied genus Neomargarodes (In

Russian). Tashkent Publishing Committee of Sciences

in Uzbek

Type material examined. ¢ Paralectotypus / design. R.

Jashenko [red label] // Neomargarodes / chondrillae /

Arch / Chondrilla brevirostris / Moiiprkym Axsip / Tede,

secku / 25.VI.31 M. Becpmuyes (DZUS).

Type locality. Akeer-Tube [Akyrtobe], near Turkes-

tan-Siberian railway [Kazakhstan |

Current status. Valid species.

Remarks. Lectotype (by original designation) is in

Saint Petersburg: Zoological Museum, Academy of Sci-

ence, Russia. Lectotype and paralectotypes were desig-

nated by R. Jashenko based on the original material used

by Archangelskaya in her first description of this species

(personal communication).

Family Pseudococcidae Cockerell, 1905

15. Mirococcus psammophilus Koteja, 1971

Mirococcus psammophilus Koteja, 1971: Polskie Pismo

Entomologiczne 41: 3-18

Type material examined. * 3224 / Mirococcus / psam-

mophilus / 9 holotype / Festuca ovina / det. J. Koteja //

4.8.1969 / 53 / Mikoszewo / distr. Nowy / Dwor Gdanski

/ Poland / leg. J. Koteja (DZUS). * 3221 / Mirococcus

psam- / mophilus / Longicoccus / Trionymus aberr. / 9 /

Festuca ovina / det. J. Koteja // 4.8.1969 / 53 / 54 / Mi-

koszewo / p. N Dwor Gdanski / leg. J. Koteja (DZUS).

¢ 3222 / Mirococcus / psammophilus / Longicoccus /

9 paratype / Festuca ovina / det. J. Koteja // 4.8.1969 /

53 / Mikoszewo / p. N. Dwor / Gdanski / leg. J. Koteja

(DZUS). * 3281 / Mirococcus / psammophilus / Longi-

coccus / 9 paratype / Festuca ovina / w p. lisc. / det. J.

Koteja // 10.8.1969 / 73 / Mikoszewo / p. Nowy Dwor /

Gdanski / leg. J. Koteyja (DZUS). * 3282 / Mirococcus /

psammophilus / Longicoccus / 8 paratype / Festuca ovi-

na / w p. lisc. / det. J. Koteja // 10.8.1969 / 73 / Miko-

szewo / pow. Nowy / Dwor Gdan- / ski / leg. J. Koteja

(DZUS). * 3283 / Mirococcus / psammophilus / Longi-

coccus / 9 paratype / Festuca ovina / w p. lisc. / det. J.

Koteja // 10.8.1969 / 73 / Mikoszewo / p. Nowy Dwor /

Gdanski / leg. J. Koteyja (DZUS). * 3284 / Mirococcus /

psammophilus / Longicoccus / 8 paratype / Festuca ovi-

na/wp. lisc. / det. J. Koteya // 10.8.1969 / 73 / Mikosze-

wo / p. Nowy Dwor / Gdanski / leg. J. Koteja (DZUS).

¢ 3285 / Mirococcus / psammophilus / Longicoccus /

9 paratype / Festuca ovina / w p. lis¢ / det. J. Koteja //

10.8.1969 / 73 / Mikoszewo / p. Nowy Dwor / Gdansk /

leg. J. Koteja (DZUS).

©ZFMK

290 Agnieszka Bugaj-Nawrocka et al.

Type locality. Mikoszewo, Nowy Dwor Gdanski [Po-

land]

Current status. Synonym of Mirococcus clarus

Borchsenius, 1949 (synonymised by Danzig & Gavri-

lov-Zimin (2014)).

16. Mirococcus festucae Koteja, 1971

Mirococcus festucae Koteja, 1971: Polskie Pismo Ento-

mologiczne 41: 3-18

Type material examined. * 1401 / Mirococcus / fes-

tucae / 2 holotype / Festuca pallens / det. J. Koteja //

10.9.1966 / 3a / Pieskowa Skala / distr. Olkusz / leg. J.

Koteja (DZUS). * 1404 / Mirococcus / festucae / 9 pa-

ratype / Longicoccus / Festuca pallens / det. J. Koteya //

10.9.1966 / 3a / Pieskowa / Skata / pow. Olkusz / leg. J.

Koteja (DZUS). * 1405 / Mirococcus / festucae / 9 pa-

ratype / Longicoccus / Festuca pallens / det. J. Koteja //

10.9.1966 / 3a / Pieskowa Skala / pow. Olkusz / leg. J.

Koteja (DZUS).

Type locality. Pieskowa Skala, Olkusz [Poland]

Current status. Valid species.

17. Dysmicoccus pauper Danzig, 1971

Dysmicoccus pauper Danzig, 1971: Entomologicheskoe

Obozrenye 50: 366-391

Type material examined. « Paratypus [red label] //

194-61 / Dysmicoccus / pauper Danzig / 31ak (kopHn) /

BuanusBoctox, Ceganka, / IIpumopcxnit kpait / 23. VIL.61

r. E. Jaauur (DZUS).

Type locality. Vladivostok, Primor’ye Kray [Russian

Federation]

Current status. Valid species.

Remarks. Holotype (by original designation) 1s in

Saint Petersburg: Zoological Museum, Academy of Sci-

ence, Russia.

18. Dysmicoccus balticus Koteja & Lagowska, 1986

Dysmicoccus balticus Koteja & Lagowska, 1986: Polskie

Pismo Entomologiczne 56: 381-388

Type material examined. * 6681a / Dysmicoccus / para-

type / 9 przdd ciala / Deschampsia / flexuosa / det. J.

Koteja/ B. Lagowska // 29.7.1969 17/ Mikoszewo / The

Baltic Coast / Poland / leg. J. Koteya (DZUS). * 6681b /

Dysmicoccus / paratype / 9 srédtutow / Deschampsia /

flexuosa / det. J. Koteja / B. Lagowska // 29.7.1969 17

/ Mikoszewo / The Baltic Coast / Poland / leg. J. Koteja

(DZUS). * 6681c / Dysmicoccus / paratype / 2 koniec

ciala / Deschampsia / flexuosa / det. J. Koteja / B. La-

gowska // 29.7.1969 17 / Mikoszewo / The Baltic Coast

/ Poland / leg. J. Koteja (DZUS). * 6682 / Dysmicoccus

/ paratype / 2 / Deschampsia / flexuosa / det. J. Koteja /

B. Lagowska // 29.7.1969 17 / Mikoszewo / The Baltic

Coast / Poland / leg. J. Koteja (DZUS). * 6683 / Dysmi-

coccus / paratype / 2 / Deschampsia / flexuosa / det. J.

Koteja/ B. Lagowska // 29.7.1969 17/ Mikoszewo / The

Baltic Coast / Poland / leg. J. Koteya (DZUS).

Bonn zoological Bulletin 70 (2): 281-315

Type locality. Mikoszewo, Nowy Dwor Gdanski [Po-

land]

Current status. Synonym of Trionymus placatus

(Borchsenius, 1949) (synonymised by Danzig (1998)).

Family Putoidae Beardsley, 1969

19. Puto orientalis Danzig, 1978

Puto orientalis Danzig, 1978: Trudy Akademii Nauk SSR

Zoologicheskogo Instituta. St. Petersburg 61: 124—132

Type material examined. « Paratypus [red label] //

67=70 / Puto / orientalis Danzig / Schizandra chinensis

/ ¥0. IIpumopse / CynyTuHcKuit 3anoBeqHUK / CKasIbI /

15.VI.69. 24 E. Jlanunr / Marepuan Becb (DZUS).

Type locality. Yakutia-Sakha AR, Khaptagay, Irkutsk

[Russian Federation]

Current status. Valid species.

Remarks. Holotype (by original designation) is in

Saint Petersburg: Zoological Museum, Academy of Sci-

ence, Russia.

Aknowledgments. The scale insects material was digitised

in the framework of the project no ND-RPSL.02.01.00-24-

01DD/17: Digital Database of Entomological Specimens HE-

MIPTERON (CEBD HEMIPTERON), co-financing from the

EU, Priority axis: II. Digital Silesia, Measure: 2.1. Support for

the development of digital public services.

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Table 1. List of scale insect species in the Zoology Research Group, University of Silesia in Katowice (DZUS) collection, including

distribution and host plants. Abbreviations: [Sex/stage:] F = female; M = male; L = larva; P = pupa or prepupa; E = egg; [Prepara-

tion type:] S = slide; D = dry; L = liquid; I = amber inclusion. “Specimens collected in Poland only in greenhouses.

Species

Sex/stage

Arnoldidae Koteja, 2008 +

Arnoldus sp. | |

Asterolecaniidae Cockerell, 1896

Asterodiaspis japonica (Cockerell, 1900)

Asterodiaspis quercicola x x

(Bouche, 1851)

Asterodiaspis variolosa (Ratzeburg, 1870) | x x

Asterodiaspis sp. xX x

*A sterolecanium epidendri x

(Bouché, 1844)

Asterolecanium sp.

Mycetococcus ehrhorni (Cockerell, 1895)

Planchonia arabidis Signoret, 1877

Pollinia pollini (Costa, 1857) x

Beesoniidae Ferris, 1950

Beesonia napiformis (Kuwana, 1914) me | | x

Callipappidae MacGillivray, 1921

Platycoelostoma compressum x

(Maskell, 1892)

Cerococcidae Balachovsky, 1942

Antecerococcus intermedius x

(Balachowsky, 1930)

Bonn zoological Bulletin 70 (2): 281-315

Geographic Host plant Preparation type

distribution

Poland | | xX

China

Poland Quercus robur

Poland Quercus robur 5 ey || De

Quercus sp.

Poland Quercus palustris «||

Quercus robur

Quercus sp.

Poland x

Poland Quercus sp.

USA Quercus sp.

Italy, Poland | Origanum vulgare x | 3c: | es

Achillea millefolium

Thymus sp.

Russian Olea europaea x

Federation

Japan | Quercus glauca x | |

New Zealand | Libocedrus bidwillii x |

©ZFMK

Species

Eriopeltis sp.

*Eucalymnatus tessellatus

(Signoret, 1873)

Eulecanium ciliatum (Douglas, 1891)

Eulecanium douglasi (Sulc, 1895)

Eulecanium franconicum

(Lindinger, 1908)

Eulecanium sericeum (Lindinger, 1906)

Eulecanium tiliae (Linnaeus, 1758)

Eulecanium sp.

Exaeretopus agropyri (Hadzibejli, 1960)

Exaeretopus orientalis Danzig, 1975

Filippia follicularis

(Targioni Tozzetti, 1867)

Bonn zoological Bulletin 70 (2): 281-315

Scale insects of DZUS collection

Sex/stage

F\|M/L/P/E

ee | ae)

Ke” ||» Meee || EX

x x

x <

Geographic

distribution

Poland

Georgia

Russian Fed-

eration

Croatia

Host plant

Agropyropsis lolium

Agrostis capillaris

Agrostis Sp.

Brachypodium pinnatum

Brachypodium sylvaticum

Brachypodium sp.

Calamagrostis epigejos

Carex sp.

Deschampsia flexuosa

Deschampsia sp.

Festuca ovina

Festuca rubra

Festuca sp.

Phleum pratense

Poa nemoralis

Poa stiriaca

Poa sp.

Brosinum sp.

Crescentia sp.

Hura sp.

Laurus sp.

Malpighia glabra

Alnus incana

Fagus sylvatica

Populus x canadensis

Quercus petraea

Quercus sp.

Ribes sp.

Calluna vulgaris

Abies alba

Crataegus sp.

Acer pseudoplatanus

Carpinus betulus

Prunus domestica

Prunus sp.

Quercus robur

Ribes sp.

Rosa sp.

Tilia platyphyllos

Tilia sp.

Acer pseudoplatanus

Aesculus hippocastanum

Calluna vulgaris

Carpinus betulus

Corylus avellana

Empetrum nigrum

Myrica gale

Salix repens subsp. are-

naria

Salix sp.

Tilia cordata

Vaccinium myrtillus

Vaccinium uliginosum

Elymus repens

Carex pediformis

Olea europaea

Pistacia terebinthus

293

Preparation type

S;D;|L {I

me] Bae

il] RK |

Ka || ge

eal re!

©ZFMK

294 Agnieszka Bugaj-Nawrocka et al.

Species Sex/stage Geographic Host plant Preparation type

FIMIL!PlE distribution Se [le ayall res oy

Lecanopsis formicarum Newstead, 1893 RPK. | OK Pox Poland Agrostis capillaris || aX:

Agrostis sp.

Brachypodium pinnatum

Bromus inermis

Dactylis glomerata

Deschampsia sp.

Elymus repens

Elymus sp.

Festuca ovina

Festuca pratensis

Festuca rubra

Festuca sp.

Koeleria glauca

Phleum pratense

Phleum sp.

Lecanopsis subterranea (Gomez-Menor x Hungary, | Festuca sp. x

Ortega, 1948) Poland Poa compressa

Lecanopsis sp. x x Poland Anthoxanthum odoratum | x

Briza media

Festuca ovina

Festuca rubra

Phleum phleoides

Poa compressa

Lichtensia viburni (Signoret, 1873) x a Poland, Slova-| Ficus benjamina KI || es

kia, Ukraine | Hedera helix

Olea europaea

Pistacia terebinthus

Luzulaspis americana Koteja & Howell, 53 USA xX

1979

Luzulaspis bisetosa Borchsentius, 1952 ell aK | KE Russian Fed- | Carex callitrichos Kea ||

eration Carex humilis subsp.

nana

Carex sp.

Luzulaspis crassispina Borchsenius, 1959 me China Luzula sp. %. || 3s

Luzulaspis dactylis Green, 1928 xX Italy, Poland | Carex sp. x || ox

Luzulaspis erianthi Rehaéek, 1954 x Cyprus Saccharum ravennae Xl He

Luzulaspis frontalis Green, 1928 Kl RO) ae | Germany, | Carex brizoides Re | ax

Great Britain,| Carex remota

Poland Elymus sp.

Festuca ovina

Luzula luzuloides

Luzulaspis grandis Borchsenius, 1952 xX % Czech Repub-| Carex ornithopoda ae || a |e

lic, Poland, | Carex sp.

Russian Fed-

eration

Luzulaspis kosztarabi Koteja & Kozar, x Hungary Carex sp. x

1979

Luzulaspis luzulae (Dufour, 1864) x | x | x | x | x |Great Britain,| Carex brizoides | Oe

Poland, Rus- | Carex sp.

sian Federa- | Luzula campestris

tion, Luzula capitata

Luzula multiflora

Luzula pilosa

Luzula sp.

Pinus sp.

Luzulaspis minima Koteja & Howell, 1979 | x USA Carex sp. xX

Bonn zoological Bulletin 70 (2): 281-315 ©ZFMK

Species

Luzulaspis nemorosa Koteja, 1966

Luzulaspis scotica Green, 1926

Luzulaspis sp.

Nemolecanium graniforme (Wunn, 1921)

Palaeolecanium bituberculatum (Signoret,

1873)

Parafairmairia bipartita (Signoret, 1874)

Parafairmairia gracilis Green, 1916

Parafairmairia sp.

*Parasaissetia nigra (Nietner, 1861)

Parthenolecanium corni (Nuzzaci, 1969)

Bonn zoological Bulletin 70 (2): 281-315

Scale insects of DZUS collection

Sex/stage

F|M/L|P/E

> x xX >. xX

x x x

x x >, x

Geographic

distribution

Bulgaria,

Czech Repub-

lic, Great Brit-

ain, Poland

Poland,

Romania,

Russian Fed-

eration

Poland

Poland, Re-

public of Mol-

dova

Kenya, Po-

land, Russian

Federation

Czech Repub-

lic, Germany,

Great Britain,

Poland, Rus-

sian Federa-

tion

Poland

Host plant

Luzula campestris

Luzula luzuloides

Carex nigra

Carex sp.

Eriophorum vaginatum

Carex sp.

Luzula campestris

Luzula luzuloides

Luzula nemorosa

Luzula pilosa

Luzula sp.

Abies alba

Crateagus sp.

Cocophagus palaelole-

cant

Agrostis capillaris

Agrostis Sp.

Carex brizoides

Carex sp.

Festuca sp.

Carex brizoides

Carex digitata

Carex ornithopoda

Carex sp.

Carex sylvatica

Eriophorum vaginatum

Carex brizoides

Betula sp.

Calluna vulgaris

Carpinus betulus

Cornus mas

Corylus avellana

Crataegus sp.

Empetrum nigrum

Euonymus europaeus

Fraxinus sp.

Genista sp.

Helianthemum nummu-

larium

Luzula pilosa

Luzula sp.

Myrica gale

Populus x canadensis

Prunus domestica

Prunus spinosa

Prunus sp.

Quercus sp.

Rhamnus cathartica

Ribes aureum

Ribes sp.

Rosa sp.

Rubus sp.

Sarothamnus scoparius

Sorbus aucuparia

Tilia platyphyllos

Tilia sp.

Vaccinium sp.

295

Preparation type

S;D;L {I

eal pe

x || Be

ie] [ok

5| ex

pal?

©ZFMK

296

Species

Parthenolecanium fletcheri (Cockerell,

1893)

*Parthenolecanium perlatum (Cockerell,

1898)

Parthenolecanium pomeranicum (Kawecki,

1954)

Parthenolecanium rufulum (Cockerell,

1903)

Parthenolecanium sp.

Phyllostroma myrtilli (Kaltenbach, 1874)

Physokermes hemicryphus (Dalman, 1826)

Physokermes piceae (Schrank, 1801)

Poaspis intermedia (Goux, 1939)

Poaspis jahandiezi (Balachowsky, 1932)

Poaspis kondarensis (Borchsenius, 1952)

Protopulvinaria pyriformis Cockerell, 1894

Psilococcus ruber Borchsenius, 1952

*Pulvinaria floccifera (Westwood, 1870)

Pulvinaria psidii Maskell, 1893

Pulvinaria salicicola Borchsentius, 1953

Pulvinaria terrestris Borchsentus, 1953

Pulvinaria vitis (Linnaeus, 1758)

Bonn zoological Bulletin 70 (2): 281-315

~ ~ KK K KK

Sex/stage

M|L{|P

Sc" |. [eee

Agnieszka Bugaj-Nawrocka et al.

Geographic

distribution

Poland

Hungary

Tajikistan

Italy, USA

Poland, Re-

public of

Korea

Croatia, Po-

land, Russian

Federation

USA

Poland

Poland,

Ukraine

Azerbaijan,

Poland, Rus-

sian Federa-

tion

Host plant

Taxus sp.

Prunus sp.

Quercus robur

Quercus sp.

Rubus sp.

Vaccinium sp.

Empetrum nigrum

Vaccinium myrtillus

Erica carnea

Vaccinium myrtillus

Vaccinium sp.

Picea abies

Dactylis glomerata

Fatsia japonica

Carex arenaria

Carex brizoides

Carex hirta

Carex sp.

Camellia L.

Cactus sp.

Gossypium Sp.

Alnus sp.

Betula pendula

Carpinus betulus

Corylus avellana

Crataegus sp.

Euonymus europaeus

Prunus sp.

Ribes aureum

Ribes uva-crispa

Ribes sp.

Salix sp.

Sorbus aucuparia

Ulmus glabra

Cydonia sp.

Populus sp.

Ribes uva-crispa

Preparation type

S;D;|L{I

x lie

x x

lee

6 |x:

x x

x ex

% || x

De | ax

©ZFMK

Species

Pulvinaria sp.

Rhizopulvinaria artemisiae (Signoret, 1873)

Rhodococcus spiraeae (Borchsentus, 1949)

*Saissetia coffeae (Walker, 1852)

*Saissetia oleae (Gomez-Menor Ortega,

1955)

Scythia craniumequinum Kiritchenko, 1938

Scythia festuceti (Sulc, 1941)

Sphaerolecanium prunastri (Boyer de

Fonscolombe, 1834)

Stotzia maxima (Borchsenius, 1957)

Toumeyella pini (King, 1901)

Vittacoccus longicornis (Green, 1916)

unknown species

Coelostomidiidae Morrison, 1928

Coelostomidia montana Green, 1929

Coelostomidia pilosa (Maskell, 1891)

Coelostomidia zealandica (Maskell, 1880)

Ultracoelostoma assimile (Maskell, 1890)

Ultracoelostoma brittini Morales, 1991

Ultracoelostoma dracophylli Morales, 1991

Bonn zoological Bulletin 70 (2): 281-315

Scale insects of DZUS collection

Sex/stage

F/M/L| P/E

xX xX xX

xX xX

Geographic

distribution

Poland

Czech Repub-

lic, Poland,

Ukraine

Armenia

Greece, Po-

land

Poland

Kazakhstan,

Ukraine

Russian Fed-

eration

Greece, Po-

land

Turkmenistan

USA

Czech Repub-

lic, Poland

Poland

New Zealand

Host plant

Alnus glutinosa

Alnus incana

Betula pubescens

Crataegus sp.

Euonymus europaeus

Populus tremula

Populus x canadensis

Prunus sp.

Ribes aureum

Ribes uva-crispa

Salix repens subsp. are-

naria

Salix sp.

Sorbus aucuparia

Dianthus sp.

Asparagus Sp.

Chlorophytum sp.

Coffea arabica

Evonymus sp.

Dracaena sp.

Pistacia lentiscus

Buxus sempervirens

Calluna vulgaris

Calluna sp.

Cycas sp.

Hibiscus syriacus

Laurus nobilis

Muhlenbekia sp.

Nerium oleander

Olea laurifolia

Pistacia lentiscus

Sideroxylon lycioides

Festuca sp.

Leymus angustus

Stipa capillata

Festuca sp.

Prunus spinosa

Ephedra sp.

Pinus sp.

Carex brizoides

Carex sp.

Dracophyllum traversii

Dacrycarpus dacry-

dioides

Muhlenbekia sp.

Pittosporum sp.

Nothofagus solandri

Dracophyllum sp.

297

Preparation type

S;D;L {I

3 al. 2

Ko| Coe

xX x

5) Se je

K| Ue

Ko| oe

©ZFMK

298

Species

unknown species

Conchaspididae Green, 1896

Conchaspis angraeci Cockerell, 1893

Conchaspis lata Hempel, 1937

Cryptococcidae Kosztarab, 1968

Cryptococcus fagisuga (Douglas, 1886)

Pseudochermes fraxini (Kaltenbach, 1860)

Dactylopiidae Signoret, 1875

Dactylopius confusus (Cockerell, 1893)

Dactylopius opuntaie (Cockerell, 1896)

Dactylopius sp.

Diaspididae Targioni-Tozzetti, 1868

*Aonidia lauri (Bouche, 1833)

Aonidiella aurantii (Maskell, 1879)

Aonidomytilus crookiae (Ferris, 1954)

*4 spidiotus nerii (Bouche, 1833)

Aspidiotus sp.

Aulacaspis rosae (Bouché, 1833)

Carulaspis juniperi (Bouché, 1851)

Chionaspis salicis (Linnaeus, 1758)

Chionaspis sp.

Circulaspis fistulata (Ferris, 1921)

Comstockaspis perniciosa (Comstock,

1881)

Diaspidiotus bavaricus (Lindinger, 1912)

Bonn zoological Bulletin 70 (2): 281-315

Agnieszka Bugaj-Nawrocka et al.

Sex/stage

F|M|L|P

xX »<

xX xX »<

»<

»< »<

xX >< xX

Geographic

distribution

Poland

Venezuela

Poland

USA

Croatia, Po-

land

Greece

USA

Croatia,

Greece, Po-

land

Russian Fed-

eration

Poland

Finland

Mexico

New Zealand

Poland

Host plant

Acalypha wilkesiana

Fagus sylvatica

Fraxinus excelsior

Opuntia sp.

Laurus nobilis

Laurus sp.

Elaeagnus pungens

Hypericum sp.

Agave americana

Aloe arborescens

Asparagus densiflorus

Brugmansia arborea

Chamaerops humilis

Citrus sp.

Euonymus japonicus

Grevillea robusta

Howea forsteriana

Kentia sp.

Nerium oleander

Rosa sp.

Rubus sp.

Cryptomeria Japonica

Juniperus communis

Sequoia sempervirens

Alnus incana

Alnus sp.

Fraxinus excelsior

Populus tremula

Rhamnus cathartica

Salix caprea

Salix repens

Salix sp.

Sorbus aucuparia

Vaccinium myrtillus

Distichlis spicata

Pyrus sp.

Calluna vulgaris

Empetrum nigrum

Preparation type

>.<

Klas 1) Se

|| P|

alee | OG

xy |,

Ko || Se

al |e

Me || axe

a | aK

©ZFMK

Scale insects of DZUS collection

Species Sex/stage

F|M|L| P/E

Diaspidiotus ostreaeformis (Curtis, 1843) xX

Diaspidiotus prunorum (Laing, 1931) x

Diaspidiotus zonatus (Frauenfeld, 1868)

Diaspidiotus sp.

*Diaspis boisduvalii Signoret, 1869 x

*Diaspis bromeliae (Kerner, 1778) x

Dynaspidiotus abietis x

(Schrank, 1776)

Dynaspidiotus sp. x

*Furchadaspis zamiae (Morgan, 1890) xX

*Hemiberlesia cyanophylli (Signoret, 1869)} x

*Hemiberlesia palmae (Cockerell, 1893) xX

*Hemiberlesia rapax (Comstock, 1881) x

*Hemiberlesia sp.

*Howardia biclavis (Comstock, 1883)

Lepidosaphes beckii (Newman, 1869) x

Lepidosaphes conchiformis (Gmelin, 1790)| x

Lepidosaphes newsteadi (Sulc, 1895)

Bonn zoological Bulletin 70 (2): 281-315

Geographic

distribution

Poland, Re-

public of Mol-

dova

Armenia

Poland

Hungary

Poland

Host plant

Alnus incana

Alnus sp.

Betula pendula

Betula sp.

Malus sp.

Pinus sp.

Populus tremula

Tilia cordata

Betula sp.

Calluna sp.

Salix caprea

Salix sp.

Quercus sp.

Cattleya sp.

Phoenix canariensis

Butia capitata

Abies alba

Picea abies

Picea sp.

Pinus sylvestris

Pinus sp.

Abies alba

Pinus sylvestris

Chrysomyphelus ficus

Butia capitata

Cordyline indivisa

Ruscus aculeatus

Chamaerops humilis

Acacia mearnsii

Acacia paradoxa

Aucuba japonica

Calluna vulgaris

Calluna sp.

Ceratonia siliqua

Euonymus japonicus

Myrtus communis

Rhododendron sp.

Bauhinia tomentosa

Ficus bengalensis

Ficus ulmifolia

Lasia spinosa

Opuntia ficus-indica

Petiveria alliacea

Strelitzia sp.

Pinus sylvestris

299

Preparation type

S;D;L {I

ge | Exe

x) || oe

Keal|| be

©ZFMK

300 Agnieszka Bugaj-Nawrocka et al.

Species Sex/stage Geographic Host plant Preparation type

FIMIL!PlE distribution eal ajal aosly

*Lepidosaphes ulmi (Linnaeus, 1758) || Greece, Betula pendula me

Montenegro, | Betula sp.

Poland Calluna vulgaris

Cerratonia siliqua

Corylus avellana

Crataegus laevigata

Crataegus sp.

Empetrum nigrum

Malus sp.

Myrica gale

Pinus sylvestris

Populus tremula

Prunus spinosa

Quercus petraea

Quercus robur

Quercus sp.

Rhamnus cathartica

Rhododendron tomen-

tosum

Rubus ideus

Rubus sp.

Salix sp.

Sorbus aucuparia

Spartium Junceum

Syringa vulgaris

Tilia cordata

Vaccinium myrtillus

Vaccinium vitis-idaea

Vaccinium sp.

Lepidosaphes sp. xX x Poland Abies alba Nem || aX

Betula sp.

Calluna vulgaris

Corylus avellana

Crataegus sp.

Tilia platyphyllos

Leucaspis lowi (Colvee, 1882) xX ae Croatia, Hun-| Pinus mugo Ke || x

gary, Poland | Pinus nigra

Pinus sylvestris

Pinus sp.

Leucaspis pini (Hartig, 1839) ae x Poland Pinus nigra || oe

Pinus sylvestris

Pinus sp.

Leucaspis pusilla (Low, 1883) x Pinus pinea x

Pinus sp.

Leucaspis sp. se Poland Pinus sylvestris XS

Pinus sp.

Melanaspis obscura (Comstock, 1881) x USA Quercus virginiana x

Quercus sp.

Niveaspis fenestrata Ferris, 1941 x Mexico Sideroxylon sp.

Parlatoria parlatoriae (Sulc, 1895) x Poland Picea abies

*Parlatoria pergandii Comstock, 1881 xX Poland Diffenbachia sp.

Parlatoria proteus (Curtis, 1843) x Poland

*Pinnaspis aspidistrae (Signoret, 1869) xX Poland Nephrolepis exaltata

Nephrolepis sp.

Prodiaspis tamaricicola (Balachowsky, x --- x

1953)

*P seudaulacaspis pentagona (Targioni xX Poland Salix sp. x

Tozzetti, 1886)

Bonn zoological Bulletin 70 (2): 281-315 ©ZFMK

Scale insects of DZUS collection

301

Species Geographic Host plant Preparation type

distribution

Rhizaspidiotus canariensis (Lindinger, Poland Aster amellus

1911)

Targionia vitis (Signoret, 1876) Croatia Quercus ilex

Velataspis cornigera Ferris, 1941 Panama

unknown species Poland x

Eriococcidae Cockerell, 1899

Acanthococcus aceris (Signoret, 1875) Poland Acer campestre AD!) Pe |e

Acer platanoides

Acer pseudoplatanus

Acer sp.

Acanthococcus azaleae (Comstock, 1881) USA Rhododendron sp. x

Acanthococcus crispus (Boyer de Fonsco- Greece

lombe, 1834)

Acanthococcus salicis (Borchsenius, 1938) Poland

Acanthococcus sasae Danzig, 1975 Sasa kurilensis

Acanthococcus ulmarius Danzig, 1975 Korea (Demo-| U/mus sp.

cratic People’s

Republic of)

Acanthococcus sp. Poland Acer platanoides Ke | Fx

Achillea millefolium

Brachypodium pinnatum

Briza media

Calluna vulgaris

Dianthus carthusianorum

Erica sp.

Fragaria vesca

Galium verum

Leontodon sp.

Picea abies

Potentilla sp.

Trifolium sp.

Vaccinium myrtillus

Vaccinium vitis-idaea

Anophococcus agropyri (Borchsenius, Poland Elymus repens ee) be | Pa

1949) Elymus sp.

Festuca sp.

Holcus sp.

Anophococcus confusus (Danzig, 1962) Poland

Anophococcus formicicola (Newstead, Croatia, Hun-| Cynodon dactylon x

1897) gary

Anophococcus herbaceus (Danzig, 1962) Poland, Rus- | Agrostis capillaris Ko OG

sian Federa- | Agrostis sp.

tion Brachypodium pinnatum

Calamagrostis arundi-

nacea

Calamagrostis epigejos

Calamagrostis villosa

Carex brizoides

Carex sp.

Eriophorum vaginatum

Festuca ovina

Koeleria glauca

Luzula campestris

Luzula luzuloides

Anophococcus inermis (Green, 1915) x Poland Calamagrostis epigejos | x | x

Calamagrostis sp.

Festuca ovina

Festuca pratensis

Bonn zoological Bulletin 70 (2): 281-315 ©ZFMK

302 Agnieszka Bugaj-Nawrocka et al.

Species Sex/stage Geographic Host plant Preparation type

FIMIL! P/E distribution (lajull ers ley

Anophococcus insignis (Newstead, 1891) se x Hungary, | Agrostis capillaris p oa lle.

Poland Agrostis sp.

Agrostis stolonifera

Anthoxanthum odoratum

Brachypodium pinnatum

Bromus sp.

Calamagrostis epigejos

Calamagrostis sp.

Carex brizoides

Dactylis glomerata

Elymus repens

Festuca ovina

Festuca sp.

Koeleria glauca

Luzula campestris

Luzula sp.

Molinia caerulea

Nardus stricta

Phalaris arundinacea

Potentilla erecta

Rubus sp.

Anophococcus pseudinsignis (Green, 1921)) x | x | x Poland Agrostis capillaris RE ll x

Ammophila arenaria

Anthoxanthum odoratum

Calamagrostis arundi-

nacea

Calamagrostis epigejos

Calamagrostis sp.

Carex sp.

Dactylis glomerata lo-

bata

Deschampsia flexuosa

Elymus repens

Festuca ovina

Festuca sp.

Luzula campestris

Luzula luzuloides

Molinia caerulea

Phalaris arudinacea

Phragmites australis

Poa pratensis

Taraxacum officinale

Anophococcus sp. xX Poland x

Balticococcus spinosus Koteja, 1988 x Poland x

Eriococcus buxi (Boyer de Fonscolombe, xX Georgia xX

1834)

Eriococcus podhalensis Dziedzicka & xX a Poland Eriophorum vaginatum | X

Koteja, 1985

Eriococcus williamsi Danzig, 1987 x Russian Fed- x

eration

Gedanicoccus gracilis Koteja, 1988 i Poland x

Gossyparia salicicola Borchsenius, 1949 xX Tajikistan

Gossyparia spuria (Modeer, 1778) ee Pet aoe | OX Poland, Re- | U/mus sp. pan ES alls

public of Mol-

dova

Greenisca brachypodii Borchsenius & Dan-| x Poland Brachypodium pinnatum | x | xX

zig, 1966 Brachypodium sp.

Calamagrostis sp.

Greenisca gouxi (Balachowsky, 1954) x Poland Brachypodium pinnatum

Greenisca sp. 52 Poland Brachypodium pinnatum

Bonn zoological Bulletin 70 (2): 281-315 ©ZFMK

Species

Kaweckia glyceriae (Green, 1921)

Kuenowicoccus sp.

Kuwanina parva (Maskell, 1897)

Neokaweckia laeticoris (Tereznikova, 1965)

Neokaweckia rubra (Matesova, 1960)

Ovaticoccus agavium (Douglas, 1888)

Rhizococcus baldonensis (Rasina, 1966)

Rhizococcus cantium (Williams, 1985)

Rhizococcus devoniensis Green, 1896

Rhizococcus greeni (Newstead, 1898)

Rhizococcus munroi (Boratynski, 1962)

Bonn zoological Bulletin 70 (2): 281-315

Scale insects of DZUS collection

Sex/stage

F/M/{|L/|P/E

xX x

% x

* x

Geographic

distribution

Poland

Japan

Poland

Italy

Russian Fed-

eration

Poland

Host plant

Agrostis capillaris

Agrostis stolonifera

Agrostis Sp.

Alopecurus pratensis

Alopecurus sp.

Anthoxanthum odoratum

Brachypodium pinnatum

Brachypodium sp.

Briza media

Bromus inermis

Bromus secalinus

Calamagrostis sp.

Corynephorus canescens

Corynephorus sp.

Dactylis glomerata

Dactylis sp.

Deschampsia caespitosa

Deschampsia flexuosa

Deschampsia sp.

Elymus repens

Elymus sp.

Festuca ovina

Festuca pratensis

Festuca rubra

Festuca sp.

Holcus lanatus

Holcus sp.

Koeleria glauca

Luzula sp.

Nardus stricta

Phleum phleoides

Phleum pratense

Phleum sp.

Poa compressa

Poa pratensis

Poa trivialis

Poa sp.

Prunus sp.

Agrostis capillaris

Corynephorus canescens

Furcraea selloa

Agrostis capillaris

Erica tetralix

Vaccinium myrtillus

Vaccinium vitis-idaea

Agrostis capillaris

Briza media

Calamagrostis sp.

Carex brizoides

Corynephorus canescens

Festuca ovina

Melica uniflora

Plantago lanceolata

Poa nemoralis

Calluna vulgaris

Vaccinium sp.

303

Preparation type

S;D;L{I

eal D2

x | Oe

| ke

©ZFMK

304

Species

Rhizococcus sp.

unknown species

Electrococcidae Koteja, 2000 +

Turonicoccus beardsleyi Koteja, 2000

Turonicoccus grimaldii Koteja, 2000

Grimaldiellidae Koteja, 2000 +

Grimaldiella resinophila Koteja, 2000

Grimaldiella sp.

Inkaidae Koteja, 1989 +

unknown species

Kermesidae Signoret, 1875

Kermes quercus (Linnaeus, 1758)

Kermes sp.

Nidularia pulvinata (Planchon, 1864)

unknown species

Kerridae Lindinger, 1937

Kerria lacca (Varshney, 1977)

Kuwaniidae MacGillvray, 1921

Hoffeinsia sp.

Kuwania pasaniae Borchsenius, 1960

Neosteingelia texana Morrison, 1927

Labiococcidae Koteja, 2000 +

Labiococcus sp.

Solicoccus sp.

Lecanodiaspididae Targioni Tozzetti, 1869

Cosmococcus albizziae Borchsenius, 1960

Lithuanicoccidae Koteja, 2008 +

Lithuanicoccus damzeni Koteja, 2008

Lithuanicoccus kosmowskae Koteja, 2008

Lithuanicoccus sp.

Marchalinidae Morrison, 1927

Marchalina hellenica (Gennadius, 1883)

Margarodidae Cockerell, 1899

Neomargarodes aristidae Borchsentus,

1949

Neomargarodes chondrillae Archangelska-

ya, 1935

Neomargarodes erythrocephalus Green,

1914

Bonn zoological Bulletin 70 (2): 281-315

Sex/stage

M|L{| P

x. | x

x |

| x

Agnieszka Bugaj-Nawrocka et al.

Geographic

distribution

Poland

France, Italy

Poland

Host plant

Brachypodium pinnatum

Carex digitata

Carex sp.

Dactylis glomerata

Festuca ovina

Luzula campestris

Luzula luzuloides

Rosa sp.

Quercus sp.

Quercus ilex

Georgia, India} Ficus carica

Poland

China

USA

Poland

China

Poland

Greece

Uzbekistan

Kazakhstan

Sudan

Quercus sp.

Carya ovata

Carya sp.

Pinus sp.

Chondrilla brevirostris

Preparation type

S;D|LI{I

Pf |x

x| |

x] |

x} |

©ZFMK

Species

Porphyrophora crithmi (Goux, 1938)

Porphyrophora polonica (Linnaeus, 1758)

Porphyrophora sophorae (Archangelskaya,

1935)

Porphyrophora violaceae Matesova &

Jashenko, 1988

unknown species

Matsucoccidae Morrison, 1927

Matsucoccus electrinus Koteja, 1984

Matsucoccus gigas (?)

Matsucoccus josephi Bodenheimer & Har-

paz, 1955

Matsucoccus larssoni Koteja, 1984

Matsucoccus pini (Green, 1925)

Matsucoccus pinnatus (Germar & Berendt,

1856)

Matsucoccus saxonicus Koteja, 1986

Matsucoccus sinensis Chen, 1937

Matsucoccus yunnansonsaus Young & Hu,

1980

Matsucoccus sp.

unknown species

Micrococcidae Silvestri, 1939

Micrococcus bodenheimeri Bytinskii-Salz,

1961

Monophlebidae Signoret, 1875

Drosicha corpulenta (Kuwana, 1902)

Drosicha stebbingii (Stebbing, 1902)

Drosicha turkestanica Archangelskaya,

193]

Eophlebus sp.

Gigantococcus maximus (Newstead, 1914)

Icerya purchasi (Maskell, 1879)

Jersicoccus sp.

Palaeophlebus sp.

unknown species

Ortheziidae Green, 1896

Arctorthezia cataphracta (Olafsen, 1772)

Arctorthezia sp.

Bonn zoological Bulletin 70 (2): 281-315

Scale insects of DZUS collection

Sex/stage

F\/M/L/P/E

Xe | x

K. | ex

Mee |e

x x

28 *

i x

Geographic

distribution

France

Mongolia,

Poland

Kazakhstan

Kazakhstan,

Poland

Israel

Poland

Germany,

Poland

China

Poland

Cyprus

Russian Fed-

eration

China, India

Kazakhstan

Poland

Ivory Coast

Greece,

Montenegro,

Tunisia

Poland

Poland, Sudan

Poland, Rus-

sian Federa-

tion

Poland

Host plant

Caragana microphylla

Potentilla bifurca

Scleranthus perennis

Sophora sp.

Pinus halepensis

Calluna sp.

Picea abies

Pinus sylvestris

Pinus massonian

Pinus thunbergii

Pinus yunnanensis

Fraxinus sp.

Vachellia nilotica

Eleagnus sp.

Populus sp.

Pittosporum tobira

Spartium Junceum

305

Preparation type

S;D;L {I

| ke

el] LG) | 9

x x

x3 || Se

Ko] oe

©ZFMK

306

Species

Newsteadia floccosa (De Geer, 1778)

Newsteadia sp.

Orthezia urticae (Linnaeus, 1758)

Orthezia sp.

Ortheziola vejdovskyi (Sulc, 1895)

Protorthezia aurea Koteja, 1987

unknown species

Phenacoleachiidae Cockerell, 1899

Phenacoleachia zealandica (Maskell, 1891)

Phoenicococcidae Stickney, 1934

Phoenicococcus marlatti (Cockerell, 1899)

Pityococcidae McKenzie, 1942

Cancerococcus apterus Koteja, 1988

Cancerococcus sp.

Pityococcus moniliformalis Vea & Grimal-

di, 2015

Pityococcus sp.

Pseudococcidae Cockerell, 1905

Antonina crawi Cockerell, 1900

Antonina graminis (Maskell, 1897)

Antonina pretiosa Ferris, 1953

Antoninella sp.

Atrococcus cracens Williams, 1962

Atrococcus paludinus (Green, 1921)

Atrococcus sp.

Balanococcus boratynskii Williams, 1962

Balanococcus singularis (Schmutterer,

1952)

Bonn zoological Bulletin 70 (2): 281-315

Sex/stage

F;/M/|L/P

a || pace’ | Pax

54 x

x *

x | x |

x| |x

BM oe lax

Rie |||, HK

x x

Agnieszka Bugaj-Nawrocka et al.

Geographic

distribution

Poland

Croatia, Po-

land, Russian

Federation

Mexico, Po-

land

Poland

Host plant

Achillea millefolium

Vaccinium myrtillus

Artemisia sp.

Urtica dioica

Citrus sp.

New Zealand | Nothofagus fusca

USA

Poland

Russian Fed-

eration

Venezuela

USA

Ukraine

Poland

Hungary

Poland

| Phoenix dactylifera

Bambusa sp.

Agropyron Sp.

Festuca sp.

Galium verum

Glechoma hederacea

Hieracium pilosella

Hieracium sp.

Leontodon hispidus

Leontodon sp.

Melampyrum pratense

Plantago media

Potentilla sp.

Taraxacum officinale

Taraxacum sp.

Trifolium pratense

Trifolium sp.

Veronica chamaedrys

Veronica sp.

Vicia sp.

Rubus sp.

Agrostis capillaris

Deschampsia caespitosa

Festuca ovina

Poa pratensis

Preparation type

S;D|L{I

xe x

x x

xe | x

x] |

x| |

%. || x

©ZFMK

Species

Boreococcus ingricus Danzig, 1960

Brevennia pulveraria (Newstead, 1892)

Ceroputo pilosellae Sulc, 1898

Chaetococcus bambusae (Maskell, 1893)

Chaetococcus sulcii (Green, 1934)

Coccura comari (Kunow, 1880)

Coccura convexa Borchsenius, 1949

Coccura suwakoensis (Kuwana & Toyoda,

1915)

Dysmicoccus angustifrons (Hall, 1926)

Dysmicoccus sp.

Dysmicoccus walkeri (Newstead, 1891)

Dysmicoccus wistariae (Green, 1923)

Ferrisia virgata (Cockerell, 1893)

Fonscolombia abdita (Borchsentus, 1949)

Fonscolombia europaea (Newstead, 1897)

Fonscolombia phenacoccoides (Kiritchen-

ko, 1932)

Bonn zoological Bulletin 70 (2): 281-315

Scale insects of DZUS collection

Sex/stage

F|M/|L| P/E

xX xX xX

xX xX

Geographic

distribution

Poland

Greece,

Poland,

Slovakia

China

Poland

Mongolia

China, Rus-

sian Federa-

tion

Poland

USA

Poland

Host plant

Carex sp.

Agrostis capillaris

Agrostis gigantea

Agrostis Sp.

Festuca ovina

Poa pratensis

Carex sp.

Fragaria sp.

Fragaria vesca

Hieracium pilosella

Hieracium sp.

Leontodon sp.

Phleum sp.

Plantago lanceolata

Plantago media

Poa sp.

Rubus sp.

Taraxacum officinale

Thymus capitatus

Thymus pulegioides

Thymus sp.

Bambusa multiplex

Bambusa sp.

Phyllostachys sp.

Festuca ovina

Festuca pallens

Rubus sp.

Artemisia sp.

Persica sp.

Rubus sp.

Hieracium pilosellae

Agrostis capillaris

Calamagrostis arundi-

nacea

Calamagrostis epigejos

Festuca rubra

Holcus lanatus

Molinia caerulea

Taxus cuspidata

Gleditsia sp.

Robinia pseudacacia

Brachypodium pinnatum

Brachypodium sp.

Bromus sp.

Corynephorus canescens

Festuca ovina

Holcus lanatus

Phleum phleoides

Phleum pratense

Festuca ovina

Festuca pallens

Koeleria sp.

Poa compressa

Corynephorus canescens

307

Preparation type

S;D;|;L {I

gy bx:

a | Bae

Kw! FE. |G

| Gok

©ZFMK

308

Species

Fonscolombia tomlinii (Newstead, 1892)

Heliococcus bohemicus (Sulc, 1912)

Heliococcus destructor Borchsenius, 1941

Heliococcus sulcii Goux, 1934

Heliococcus sp.

Heterococcus nudus (Green, 1926)

Heterococcus sp.

Kiritshenkella lianae Koteja, 1988

Lacombia dactyloni (Bodenheimer, 1943)

Metadenopus festucae Sulc, 1933

Mirococcopsis subterranea (Newstead,

1893)

Mirococcus clarus (Borchsentius, 1949)

Mirococcus festucae Koteja, 1971

*Nipaecoccus nipae (Maskell, 1893)

Paraputo angustus (Ezzat & McConnell,

1956)

Bonn zoological Bulletin 70 (2): 281-315

~ Ke KR K

Agnieszka Bugaj-Nawrocka et al.

Sex/stage

M/L

Geographic

distribution

Poland

Hungary,

Poland

Malta

Poland

Russian

Federation

Host plant

Agrostis capillaris

Festuca ovina

Poa compressa

Fagus sylvatica

Festuca sp.

Melampyrum sp.

Pinus sylvestris

Populus tremula

Quercus robur

Quercus sp.

Vaccinium myrtillus

Dianthus sp.

Thymus sp.

Hieracium sp.

Corynephorus canescens

Thymus sp.

Agropyron sp.

Agrostis capillaris

Briza media

Briza sp.

Bromus sp.

Corynephorus canescens

Danthonia decumbens

Deschampsia caespitosa

Elymus repens

Festuca cinerea

Festuca ovina

Holcus lanatus

Phleum pratense

Picea abies

Poa compressa

Poa pratensis

Setaria viridis

Agrostis capillaris

Carex nigra

Brachypodium sp.

Deschampsia caespitosa

Deschampsia sp.

Festuca ovina

Festuca sp.

Poa pratensis

Poa sp.

Dactylis glomerata

Festuca ovina

Festuca pallens

Poa compressa

Brachypodium pinnatum

Corynephorus canescens

Festuca ovina

Festuca pallens

Carludovica palmata

Chamaedorea elegans

Hedyscepe canterburyana

Howea forsteriana

Livistona chinensis

Rhapis excelsa

Bambusa sp.

Preparation type

S;D|LI{I

x | x

x. | ox

x || x

ee |e

x lie

©ZFMK

Species

Paraputo pauper (Danzig, 1971)

Peliococcopsis parviceraria (Goux, 1937)

Peliococcus calluneti (Lindinger, 1912)

Pelionella balteata (Green, 1928)

Pelionella manifecta (Borchsenius, 1949)

Phenacoccus aceris (Signoret, 1875)

Phenacoccus dearnessi King, 1901

Phenacoccus gossypii Townsend & Cock-

erell, 1898

Phenacoccus hordei (Lindeman, 1886)

Phenacoccus interruptus Green, 1923

Bonn zoological Bulletin 70 (2): 281-315

Scale insects of DZUS collection

Sex/stage

F/M/|L/|P/E

SMe | aie || Be

Xe x

* 5

x x

Geographic

distribution

Russian

Federation

Poland

USA

Poland

Host plant

Corynephorus canescens

Calluna vulgaris

Empetrum nigrum

Erica tetralix

Pinus sp.

Vaccinium myrtillus

Vaccinium uliginosum

Anthoxanthum odoratum

Calamagrostis sp.

Danthonia decumbens

Festuca pratensis

Festuca sp.

Melica sp.

Acer platanoides

Acer sp.

Aesculus hippocastanum

Alnus glutinosa

Alnus incana

Alnus sp.

Betula pendula

Betula sp.

Carpinus betulus

Carpinus sp.

Corylus avellana

Crataegus sp.

Fraxinus excelsior

Malus domestica

Malus sp.

Padus avium

Padus sp.

Poa sp.

Prunus domestica

Prunus sp.

Quercus robur

Quercus sp.

Ribes aureum

kibes uva-crispa gros-

sularia

Ribes sp.

Sorbus aucuparia

Tilia cordata

Tilia platyphyllos

Tilia sp.

Crataegus sp.

Plectranthus parviflorus

Agrostis capillaris

Anthoxanthum odoratum

Brachypodium sp.

Festuca ovina

Koeleria glauca

Leontodon sp.

Phleum phleoides

Poa cenisia

Agrostis capillaris

309

Preparation type

S;D;L {I

call Be

x | TR

©ZFMK

310 Agnieszka Bugaj-Nawrocka et al.

Species Sex/stage Geographic Host plant Preparation type

FIMIL!IPlE distribution lapel. |)

Phenacoccus phenacoccoides (Kiritchenko,| x x Poland Agrostis capillaris 6

1932) Agrostis sp.

Alopecurus pratensis

Apera sp.

Brachypodium pinnatum

Briza media

Bromus erectus

Calamagrostis sp.

Corynephorus canescens

Corynephorus sp.

Dactylis sp.

Elymus sp.

Festuca ovina

Holcus lanatus

Koeleria glauca

Koeleria sp.

Phleum phleoides

Poa compressa

Poa sp.

Phenacoccus piceae (Low, 1883) x x Poland Festuca sp. lll oe

Picea abies

Picea sp.

Phenacoccus pumilus Kiritshenko, 1936 x Slovakia Convolvulus arvensis

Phenacoccus sp. x Poland Acer platanoides X || 3s

Acer sp.

Aesculus hippocastanum

Agrostis sp.

Alnus incana

Calamagrostis sp.

Carpinus betulus

Corylus avellana

Corylus sp.

Corynephorus canescens

Crateagus sp.

Elymus sp.

Koeleria sp.

Prunus domestica

Prunus sp.

Ribes sp.

*Planococcus citri (Risso, 1813) x % Poland Brosimum utile XK, || ae

Citrus medica

Coffea arabica

Duranta erecta

Lasia spinosa

Planococcus vovae (Nasonov, 1909) x Poland Juniperus communis lex

Juniperus sp.

Pseudococcus longispinus (Targioni Toz- xX x Poland Chamaedorea oblongata | x | x | xX

zetti, 1867) Crassula arborescens

Crassula muscosa

Opuntia microdasys

Ricinus sp.

Vachellia horrida

Vitis voineriana

Vitis sp.

Bonn zoological Bulletin 70 (2): 281-315 ©ZFMK

Species

*Pseudococcus maritimus Ehrhorn, 1900

Pseudococcus sp.

Rhodania occulta Schmutterer, 1952

Rhodania porifera Goux, 1935

Rhodania sp.

Saccharicoccus isfarensis (Borchsenius,

1949)

Saccharicoccus sp.

Trabutina crassispinosa Borchsentius, 1941

Bonn zoological Bulletin 70 (2): 281-315

Scale insects of DZUS collection

F|M

Sex/stage

L.| P| E

Geographic

distribution

Poland

Hungary,

Poland

Russian

Federation

311

Host plant Preparation type

Aloe sp. x

Calluna vulgaris

Coffea arabica

Ficus elastica

Haemanthus sp.

Duranta erecta

Muhlenbekia sp.

Opuntia sp.

Umbellularia californica

Aloe sp. ‘3

Calamagrostis sp.

Fagus silvatica

Holcus lanatus

Leontodon sp.

Picea abies

Taraxacum sp.

Thymus sp.

Agrostis sp. x

Festuca sp.

Festuca ovina 4

Festuca pallens

Koeleria sp.

Festuca pallens x

Festuca sp.

Agrostis capillaris pA

Agrostis sp.

Agrostis stolonifera

Anthoxanthum odoratum

Arrhenatherum sp.

Brachypodium pinnatum

Calamagrostis epigejos

Dactylis sp.

Elymus repens

Festuca ovina

Festuca rubra

Festuca sp.

Holcus mollis

Nardus stricta

Phleum phleoides

Phleum sp.

Poa compressa

Poa pratensis

Agrostis capillaris x

Festuca ovina

Festuca pallens

Tamarix sp. x

©ZFMK

312

Species

Trionymus aberrans

(Goux, 1941)

Trionymus caricis

McConnell, 1941

Trionymus hamberdi

(Borchsentus, 1949)

Trionymus multisetiger

(Borchsenius, 1949)

Trionymus newsteadi

(Green, 1917)

Bonn zoological Bulletin 70 (2): 281-315

Sex/stage

F/M/L/P

ll x:

x x

Agnieszka Bugaj-Nawrocka et al.

Geographic

distribution

Austria, Hun-

gary, Monte-

negro, Poland

USA

Hungary,

Poland

Republic of

Korea

Poland

Host plant

Agrostis capillaris

Agrostis sp.

Agrostis stolonifera

Ammophila arenaria

Anthoxanthum odoratum

Brachypodium pinnatum

Briza media

Bromus inermis

Bromus sp.

Calamagrostis epigejos

Corynephorus canescens

Corynephorus sp.

Deschampsia caespitosa

Deschampsia sp.

Elymus repens

Festuca ovina

Festuca pratensis

Festuca rubra

Festuca sp.

Holcus lanatus

Holcus mollis

Nardus stricta

Phleum pratense

Phleum sp.

Poa compressa

Poa pratensis

Stipa capillata

Chasmanthium ornitho-

rhynchum

Agrostis capillaris

Agrostis sp.

Anthoxanthum odoratum

Calamagrostis sp.

Corynephorus canescens

Corynephorus sp.

Festuca ovina

Holcus lanatus

Koeleria glauca

Poa sp.

Dendranthema sp.

Fagus silvatica

Quercus sp.

Preparation type

S;D;|LI{I

ale’

all ee

©ZFMK

Species

Trionymus perrisii

(Goux, 1941)

Trionymus placatus

(Borchsenius, 1949)

Trionymus radicum

(Newstead, 1895)

Bonn zoological Bulletin 70 (2): 281-315

Scale insects of DZUS collection

Sex/stage

F/M/|L/|P/E

% xX

x x

Geographic

distribution

Austria, Hun-

gary, Italy, Po-

land, Russian

Federation,

Slovakia

Poland

Host plant

Agrostis canina

Agrostis capillaris

Agrostis stolonifera

Agrostis sp.

Anthoxanthum odoratum

Arrhenatherum elatius

Arrhenatherum sp.

Bambusa sp.

Brachypodium pinnatum

Bromus erectus

Bromus inermis

Calamagrostis epigejos

Calamagrostis varia

Calamagrostis sp.

Corynephorus canescens

Corynephorus sp.

Dactylis glomerata

Dactylis sp.

Danthonia decumbens

Deschampsia caespitosa

Deschampsia flexuosa

Deschampsia sp.

Elymus caninus

Elymus repens

Festuca ovina

Festuca pratensis

Festuca rubra

Festuca sp.

Holcus lanatus

Holcus mollis

Koeleria glauca

Koeleria sp.

Leontodon sp.

Lolium perenne

Nardus stricta

Phleum phleoides

Phleum pratense

Phleum sp.

Poa annua

Poa compressa

Poa pratensis

Poa sp.

Deschampsia flexuosa

Agrostis capillaris

Agrostis Sp

Festuca ovina

Festuca rubra

Festuca sp.

Holcus mollis

Phleum pratense

Poa compressa

313

Preparation type

S;';D|L{I

eal] ke

Kail PX

©ZFMK

314 Agnieszka Bugaj-Nawrocka et al.

Species Sex/stage Geographic Host plant Preparation type

FIMIL!IPlE distribution slplxili

Trionymus thulensis x 5.8 Hungary, | Agrostis canina ll ok

Green, 1931 Poland Agrostis capillaris

Agrostis stolonifera

Agrostis sp.

Alnus sp.

Anthoxanthum odoratum

Brachypodium sp.

Calamagrostis epigejos

Carex sp.

Deschampsia caespitosa

Deschampsia flexuosa

Elymus repens

Elymus sp.

Festuca ovina

Festuca rubra

Festuca sp.

Holcus lanatus

Holcus mollis

Phleum pratense

Phleum sp.

Poa cenisia

Poa sp.

Trionymus sp. xX xX Poland Agrostis capillaris RK || eat

Agrostis gigantea

Agrostis sp.

Agrostis stolonifera

Anthoxanthum odoratum

Calamagrostis arundi-

nacea

Calamagrostis epigejos

Deschampsia caespitosa

Elymus repens

Festuca ovina

Festuca pallens

Festuca rubra

Festuca sp.

Hierochloé odorata

Holcus lanatus

Phleum pratense

Poa compressa

Poa pratensis

Poa stiriaca

Poa sp.

Volvicoccus volvifer xX Hungary, Stipa capillata x

(Goux, 1945) Poland

unknown species Keg |B | eak Poland x 3

Putoidae Beardsley, 1969

Puto arctostaphyli x x USA Arctostaphylos sp. 54

Ferris, 1950

Puto caucasicus xX Russian Fed- x

Hadzibejli, 1956 eration

Puto mexicanus ee ||) | fee. | Mexico Aloe sp. Me | eK

(Cockerell, 1893)

Puto orentalis x Russian Schisandra chinensis %

Danzig, 1978 Federation

Puto pricei x USA x

McKenzie, 1960

Puto superbus ee Pe || ex: Croatia, Arrhenatherum elatius x

(Leonardi, 1907) Poland

Puto sp. Ak Poland xX

Bonn zoological Bulletin 70 (2): 281-315 ©ZFMK

Scale insects of DZUS collection

315

Species Sex/stage Geographic Host plant Preparation type

FIiMILIPple distribution I

unknown species Poland

Rhizoecidae Williams, 1969

Rhizoecus dianthi x Poland Acacia sp. xX

Green, 1926 Syzygium paniculatum

Rhizoecus vitis x Russian Vitis sp. x

Borchsenius, 1949 Federation

Ripersiella halophila x Poland x

(Hardy, 1868)

Serafinidae Koteja, 2008 +

Steingeliidae Morrison, 1927

Steingelia gorodetskia KP | PS S| oe Poland Luzula luzuloides x? || &

Nasonov, 1908

unknown species xX Poland xX

Stictococcidae Lindinger, 1913

Stictococcus intermedius x Central A fri- x

Newstead, 1917 can Republic

Xylococcidae Pergande, 1898

Xylococculus betulae x USA x

(Pergande in Hubbard & Pergande, 1898)

Xylococcus filiferus x x Poland Tilia sp. Re ||

Loéw, 1882

Xylococcus japonicus x mt Japan x

(Oguma, 1926)

Xylococcus sp. eS EP RE Oe Japan x

unknown species Poland x

unknown family

Bonn zoological Bulletin 70 (2): 281-315 ©ZFMK

BHL

Blank Page Digitally Inserted

Bonn zoological Bulletin 70 (2): 317-331

2021 - Volynkin A.V.

https://do1.org/10.20363/BZB-2021.70.2.317

ISSN 2190-7307

http://www.zoologicalbulletin.de

Research article

urn:|sid:zoobank.org: pub: 8B4F0351-C374-41D2-BC14-CAE2CDCE4ECF

Contribution to the knowledge of the genus Thumatha Walker, 1866

(Lepidoptera: Erebidae: Arctiinae) from Africa

with descriptions of four new species

Anton V. Volynkin

The African Natural History Research Trust (ANHRT), Street Court Leominster, Kingsland, HR6 9QA, UK

Altai State University, Lenina Avenue 61, RF-656049, Barnaul, Russia

Email: anton@anhrt.org.uk

urn:Isid:zoobank.org:author:25EA 1403-9502-42BE-A 820-96ACS5 1 CE862E

Abstract. The present paper contains descriptions of four new species of the genus 7iumatha Walker, 1866: 7? ngewo

sp. n. (Liberia, Guinea and Ivory Coast), 7. smithi sp. n. (Zambia), 7? kuehnei sp. n. (Zambia) and 7) jiwundu sp. n. (Zam-

bia). The hitherto unknown female of 7? punctata Kthne, 2010 is illustrated and described. Thumatha punctata is reported

for the first time from Zambia and 7: /unaris Durante, 2007 is reported for the first time from Cameroon and South Sudan.

Adults together with male and female genitalia of the new and similar species are illustrated.

Key words. Lithosiini, Nudariina, Zambia, Cameroon, South Sudan.

INTRODUCTION

Thumatha Walker, 1866 is a distinctive lithosiine genus

belonging to the subtribe Nudariina of the Lithosiini dis-

tributed widely from the A frotropics through the Palaearc-

tic and Oriental Regions to Australia. Despite the fact that

several species of the genus were described during the

last two decades (Holloway 2001; Durante 2007, 2009;

Kuhne 2007, 2010), the taxonomy of the genus, especial-

ly its African members, has remained poorly studied and

the genus is in need of revision. In the course of studying

the Lithosuni collection of the African Natural History

Research Trust (Leominster, UK), four unidentified spe-

cies of Thumatha collected in Zambia, Liberia, Guinea

and Ivory Coast were found. The comparison of the gen-

italia structures of these species with other similar taxa

in the genus has confirmed their specific distinctness and

they are described below as species new to science. Ad-

ditionally, the illustration and diagnosis of the hitherto

unknown female of 7iumatha punctata Kihne, 2010 are

provided in the present paper.

MATERIAL AND METHODS

Abbreviations of the depositories used

ANHRT = African Natural History Research Trust,

Leominster, UK

= private collection of L. Kthne,

Potsdam, Germany

LKP

Received: 26.05.2021

Accepted: 06.07.2021

MEN = Museum of Natural History, Berlin (Museum

fur Naturkunde), Berlin, Germany

OUMNH = Oxford University Museum of Natural

History, Oxford, UK

= The Bavarian State Collection of Zoology

(Zoologische Staatssammlung Munchen),

Munich, Germany

ZSM

Other abbreviations

AV = genitalia slide prepared by A.V. Volynkin

HT = holotype

PT = paratype

The genitalia were dissected and mounted in euparal on

microscope slides. The photos of adults were taken us-

ing a Nikon D3100/AF-S camera equipped with a Nikkor

18-55 mm lens while the photos of genitalia were taken

using the same camera attached to a microscope with an

LM-scope adapter. All pictures were processed using the

Adobe Photoshop CC 2018 software.

RESULTS

Genus Thumatha Walker, 1866

Thumatha Walker, 1866 (Walker 1866: 1900). Type spe-

cies: Thumatha fuscescens Walker, 1866, by monotypy.

= Pelobrochis Lucas, 1892 (Lucas 1892: 250). Type spe-

cies: Scaeodora rava Lucas, 1890 (a junior synonym of

Thumatha fuscescens), by monotypy.

Corresponding editor: M. Espeland

Published: 17.07.2021

318

= Dictenus Butler, [1897] (Butler 1896: 846). Type spe-

cies: Dictenus inconstans Butler, [1897], by monotypy.

= Nudaridia Hampson, 1900 (Hampson 1900: 420). Type

species: Nudaria muscula Staudinger, 1887, by origi-

nal designation.

= Thumata auct. (Draudt 1914; Witt et al. 2011, an incor-

rect spelling).

Diagnosis. The members of the genus are small or

medium sized moths with a relatively wide forewing

and ochreous or brownish coloration. The male anten-

na is bipectinate with short branches or serrate while the

female antenna is serrate or ciliate. The male genitalia

of Thumatha are similar to those of certain members of

the Asura/Miltochrista generic complex namely the Ja-

van Thamoma Volynkin, 2020 (illustrated by Volynkin

(2020)) from which Thumatha differs in the relatively

short and swollen uncus bearing a dorsal crest (an au-

tapomorphic feature) whereas the uncus of 7hamoma is

markedly more elongate, lacking a dorsal crest and only

slightly swollen subapically. The ventral costal protru-

sion is fused with the base of the ventro-distal process of

the valva in both genera but in Thumatha it is wider and

originates from the distal section of the costa whereas in

Thamoma it is narrower and projects subdistally while

the distal section of the costa is thin and protrudes dis-

tally. Additionally, in Thumatha, the ventral costal pro-

trusion lacks a process or (in certain species) bears only

a small setose bulge whereas that of Thamoma bears a

large, heavily sclerotized stick-like process directed dor-

so-distally with the outer edge of its base fused with the

ventro-distal process of the valva (an autapomorphic fea-

ture of the genus). The female genitalia of the genus are

characterized by the presence of sclerotized areas in the

posterior section of the corpus bursae whereas the corpus

bursae of Thamoma is rugose posteriorly.

Distribution. Sub-Saharan Africa (including Mada-

gascar), temperate and southern Eurasia, Sundaland and

Australia.

The species composition of the genus Thumatha

— africana Kihne, 2007 (Kenya, Tanzania)

— brunnea Kithne, 2007 (Rwanda)

— fuscescens Walker, 1866 (Australia, Indochina, India,

Sri Lanka)

= rava Lucas, 1890

— inconstans (Butler, [1897]) (Malawi, South Africa)

= inconstans vat. limbatula Strand, 1912

= inconstans vat. delimbatula Strand, 1912

= thumathaeformis Strand, 1912

— infantula (Saalmiuller, 1880) (Madagascar, Democratic

Republic of Congo, Sudan, Nigeria, Guinea)

— insularis Durante, 2009 (Indonesia: Java)

—jiwundu sp. n. (Zambia)

— kakamegae Kihne, 2007 (Kenya, Uganda)

Bonn zoological Bulletin 70 (2): 317-331

Anton V. Volynkin

— kuehnei sp. n. (Zambia)

— lunaris Durante, 2007 (Nigeria, Cameroon, South Su-

dan)

— monochroa Zolotuhin, 1996 (SE Kazakhstan)

— muscula (Staudinger, 1887) (southern Russian Far

East, Japan)

= inouei Okano, 1958

—ngewo sp. n. (Guinea, Liberia, Ivory Coast)

— ochracea (Bremer, 1861) (southern Russian Far East,

Japan)

= japonica Okano, 1957

— orientalis Holloway, 2001 (Malaysia: Borneo)

— punctata Kuthne, 2010 (Namibia, Zambia)

— senex senex (Hubner, [1808]) (Europe, West and East

Siberia, NE Kazakhstan)

= rotunda Haworth, 1809

= karvajszkyi Didszeghy, 1923

= tramontana Dannehl, 1929

— senex syriaca (Daniel, 1939) (Turkey)

— smithi sp. n. (Zambia)

Description of the new species

Thumatha smithi sp. n.

(Figs 1-2, 19-21, 34)

urn: lsid:zoobank.org:act: BSACEB15-846C-470B-85 7C-D56309072261

Type material

Holotype (Figs 1, 19). 3, “Zambia, 1400m, Hill-

wood, Ikelenge (Miombo / Riverine forest mosaic)

11°16'02" S, 24°18'59" E, 23-30.x1.2019 MV Light Trap,

Bashford, M., Miles, W., Mulvaney, L., Smith, R. Leg.

ANHRT:2019.25”, ANHRT unique number: 00204263,

gen. prep. No.: AV6383 (ANHRT).

Paratypes. ZAMBIA: | ©, same data as for holotype,

ANHRT unique number: 00204264 (ANHRT); 1 9,

1400 m, Hillwood, Ikelenge, 11°16'02”" S, 24°18'59" E,

23-30.x1.2019, MV Light Trap, Bashford, M., Miles, W.,

Mulvaney, L., Smith, R. leg. ANHRT:2019.25, ANHRT

unique number: 00204317 (ANHRT); 1 <, the same lo-

cality, but 30.iv—11.v.[20]14, Light Trap, leg. Smith, R.,

Takano, H., Chmurova, L., Smith, L., ANHRT unique

number: 00200974, gen. prep. No.: AV4498 (ANHRT);

1 Q, the same locality, but 9-16.v.[20]15, Light Trap,

leg. Smith, R., Takano, H. & Aristophanous, M.,

ANHRT:2017.13, ANHRT unique number: 00039466,

gen. prep. No.: AV5646 (ANHRT),; 1 3, 1147m, Lukwak-

wa, West Lunga NP, 12°39'40" S, 24°26'13" E, 28-29.

iv.[20]14, Light Trap, leg. Smith, R., Takano, H., Chmur-

ova, L., Smith, L., ANHRT unique number: 00201063

(ANHRT),; 1 4, 1437m, Kapishya Hot Springs, Shiwa

N’gandu Estate, 11°10'13" S, 31°36'00" E, 1-111.2016,

M.T. Harvey Coll., Smith, R., Takano, H. leg., ANHRT:

2017.29, ANHRT unique number: 00070216, gen. prep.

No.: AV5645 (ANHRT).

©ZFMK

Four new Thumatha from Africa 319

ZANBIA 1400m orn |

l (Mobe / Rivet forest ye ZAMBIA 1400m

Slide 11°1802'S, 24°18'59°E TI'IBOeS oe BSUE

ANHRTUK AV6383 # a ANHRTUK 23-30 fo pets tT rap

00204263 a Volynkin Muvaney L. $0 L., Smith, R. Leg. 00204317 Mulvaney, L So Smith, R'Leg.

T. smithi sp. n., HT 2 T. smithi sp. n., PT 2

a ZAMBIA. 1340

ae ae PPE. Y Slide Ist 1"5t5#" F630 20"

ANHRTUK “AV3314 97 steveoot rae SEDEI8'S9" ANHRTUK! AV5650 P ii arin ellie

00200972, =| A. Volynkin ta 00010323} A.Volynkin — | ANHRT:201712

T. kuehnei sp. n., HT 3 T. kuehnei sp. n., PT &

Slide

“AV4268 57

A. Volynkin

T. kakamegae 3 T. kakamegae

Slide |

utaré ZSM Arct. |

Sele 3 [Pep 2020-207 ¢

LIN| A. Volynkin

— WANDA

T. brunnea, PT & 10mm T. brunnea 2

Figs 1-8. Thumatha spp., adults. Depositories of the specimens: 1-4. = ANHRT; 5. = MFN; 6—7. = LKP (photo by L.

Kuhne); 8. = ZSM.

Bonn zoological Bulletin 70 (2): 317-331 ©ZFMK

320

LIBERIA

pee se , Foya el

10)

i. rion 10°1636°W a

9.xi, 2017"

Slide

ANHRTUK AV4595 9”

00020068 =A. Volynkin _

A. Volynkin

AV6389 7

Anton V. Volynkin

LIBERIA 140m

Sinoe County, Krahn-Bassa

1 @) Reserve, J River,

Slide 7 aye hig eo Town

ANHRTUK AV5649 Q i a 2018 Stn Trap

001 01 929 ena

A.Volynkin | aner dota

CAMEROUN

8 km N, d'Edea

30.111 - leIV.1970

CGC. Herbulot

Slide

ZSM Arct.

2020-206 £

A.Volynkin |

T. lunaris ©

14 15 ANHRTUK 16 ANHRTUK

00137429 Katto ese ey aah Noe 00059351 Tr Svan Me (Monee

b Bye | oe s850% Beira Slide | etbes eae

Dauerpraparat AV6387 oe aS. mil AV5079 o” fepiave ibs ‘3 ve

aE No. 688 A. Volynkin | Sia 59842 A. Volynkin HRY 201840

T. punctata, HT ah T) punctata ‘ah T. punctata a

ZAMBIA 1340m

Jiwundu Feb (Miombo /

lide | RM

ANHRTUK ference ” ice ian

00057085 a. volynkin Sa, S4 098 Og

T. punctata 3

Figs 9-18. Thumatha spp., adults. Depositories of the specimens: 9-1, 15-18. = ANHRT; 12. =

ZSM: 14 = MEN.

Bonn zoological Bulletin 70 (2): 317-331

Jrds Svamp (\ionoo!

Nair aoe

ANHRTUK “AVE651 see Leteles rp, Arua, M

00056935 A. Volynkin oto

T. punctata &

10 mm

OUMNH; 13.

©ZFMK

Four new Thumatha from Africa 321

Diagnosis. Thumatha smithi (Figs 1—2) 1s superficially

reminiscent of 7? punctata (Figs 14-18) but can easily

be distinguished by the somewhat wider forewing, the

olive brown forewing ground color (it is ochreous brown

in T. punctata), the paler and more indistinct forewing

markings and the absence of a discal spot on the hind-

wing which is distinct in 7! punctata. Additionally, the

head of 7) smithi is ochreous yellow whereas it is brown

with a suffusion of black scales in 7! punctata. The male

genitalia of the new species (Figs 19-21) are most sim-

ilar to those of the externally dissimilar 7’ kakamegae

(Figs 22-23). In the male genital capsule, the uncus of

7: smithi is somewhat less dilated medially than in 7° ka-

kamegae, the valva is less dilated distally with costa

markedly less concave medially, the medial costal pro-

cess 1S conspicuously shorter and narrower basally and

the ventro-distal process of the valva is narrower basally.

The aedeagi of the two species display no differences. In

the vesica of T. smithi, the main chamber is large and pro-

truding distally whereas it is very short in 7. kakamegae.

The medial diverticulum of the new species is less elon-

gate and more rounded than that of 7’ kakamegae bearing

fewer cormuti. The female genitalia of 7) smithi (Fig. 34)

differ from those of 7! kakamegae (Fig. 35) in the nar-

rower 8" abdominal segment, the narrower antrum, the

longer sclerotized posterior section of the corpus bursae

(in proportion to the length of the anterior section), and

the somewhat more elongate appendix bursae.

Description

External morphology of adults (Figs 1—2). Forewing

length 9.0 mm in males and 10.5 mm in males. Sexual

dimorphism limited: Female somewhat larger than male

with forewing having more elongate apex and more in-

distinct markings than in male. Antenna yellow, bipecti-

nate in male and ciliate in female. Head ochreous yellow.

Thorax brown with ochreous yellow collar. Forewing

ground color olive brown. Markings indistinct, dark

brown. Subbasal and costal area with intense brown suf-

fusion. Discal spot almost round in male and falcate and

indistinct in female. Subterminal line sinuous, interrupt-

ed constituting small spots on veins. Cilia olive brown.

Hindwing and cilia pale ochreous with slight brown suf-

fusion along costal margin. Abdomen ochreous yellow.

Male genitalia (Figs 19-21). Uncus elongate, swol-

len, dilated distally with small claw-shaped tip and short

dorsal crest. Tuba analis membranous, scaphium thin

and weakly sclerotized. Tegumen approximately half as

long as valva, its arms moderately wide, fused in distal

quarter. Vinculum ca. 1.6 x longer than tegumen, saccus

short (ca. 4 of vinculum length), moderately sclerotized,

U-shaped. Juxta weakly sclerotized, wide, shield-like

with shallow, rounded medio-distal depression. Valva

lobe-like, dilated distally. Costa narrow, convex medial-

ly and curved ventrad subapically, its distal section thin,

fused with distal membranous lobe of valva dorsally. Me-

Bonn zoological Bulletin 70 (2): 317-331

dial costal process elongate and thin, tapered and slightly

curved distally, rounded apically. Ventral protrusion of

costa projecting subapically, dilated ventrally, fused with

base of ventro-distal process of valva, bearing small, se-

tose bulge medio-distally. Ventro-distal process of valva

triangular with rounded tip. Distal membranous lobe of

valva wide, almost triangular with rounded apex. Saccu-

lus narrow, short (ca. 1.4 x shorter than ventral margin of

valva), weakly setose dorsally. Aedeagus dilated distally

with short and narrow coecum. Distal quarter of aedea-

gus serrulate. Main chamber of vesica broadly conical

and rounded distally. Medial diverticulum globular, as

large as main chamber, bearing three short, stout cornuti

of wide base. Distal diverticulum twice as narrow as me-

dial one, bearing an elongate cluster of numerous short,

acute triangular cornuti of various sizes. Vesica ejacula-

torius constricted basally, weakly granulated subbasally,

directed anteriad.

Female genitalia (Fig. 34). Papillae anales trapezoi-

dal with rounded corners, weakly setose. Apophyses an-

teriores and posteriores moderately long and thin, equal

in length. Antrum short and wide with wide but shallow

medial depression ventrally and convex lateral margins,

moderately sclerotized and granulated. Posterior section

of corpus bursae heavily sclerotized, equal in length with

anterior one, asymmetrical with left side longer and more

heavily sclerotized than right one. Anterior section of

corpus bursae membranous, teardrop-shaped. Appendix

bursae erected postero-ventrally, broadly conical, moder-

ately sclerotized, directing postero-laterally on right side.

Distribution. The species is to date only known from

Zambia.

Etymology. The species is dedicated to Mr Richard

Smith, founder and director of the African Natural Histo-

ry Research Trust, and one of the collectors of the holo-

type who has organized and undertaken numerous ento-

mological expeditions to Sub-Saharan Africa.

Thumatha kuehnei sp. n.

(Figs 3-4, 24—25, 36)

urn:lsid:zoobank.org:act: E57F 7EF F-03F7-417B-AD2D-79ABDA80B60C

Type material

Holotype (Figs 3, 24), 4, “Zambia, 1400m, Hillwood,

Ikelenge, 11°16'02" S, 24°18'59" E, 17—24.111.[20]13

Light Trap leg. Smith, R. & Takano, H.”, ANHRT unique

number: 00200972, gen. prep. No.: AV3314 (ANHRT).

Paratypes. ZAMBIA: 1 <4, 1400 m, Hillwood, Ike-

lenge (Miombo/Riverine forest mosaic) 11°16'02" S,

24°18'59" E, 23-30.x1.2019 MV Light Trap, Bashford, M.,

Miles, W., Mulvaney, L., Smith, R. leg. ANHRT:2019.25,

ANHRT unique number: 00204231, gen. prep. No.:

AV6384 (ANHRT); 4 3, 1340 m, Jiwundu Swamp,

11°51'54' S', 25°33'20" E, 21-24.x1.[20]14, Light Trap,

leg. Smith, R. & Takano, H., ANHRT:2017.12, ANHRT

©ZFMK

322 Anton V. Volynkin

a. nae TE

T. smithi sp. n., HT

Zambia, Hillwood, Ikelenge, slide AV6383

T. smithi sp. n., PT

Zambia, Hillwood, Ikelenge, slide AV4498

T. smithi sp. n., PT

Zambia, Kapishya Hot Springs, slide AV5645

Figs 19-21. Thumatha smithi sp. n., male genitalia. The specimens dissected are deposited in ANHRT.

Bonn zoological Bulletin 70 (2): 317-331 ©ZFMK

Four new Thumatha from Africa 323

unique numbers: 00008718, 00010323, 00010381,

00200973, gen. prep. Nos.: AV3315, AV5650 (ANHRT).

Diagnosis. Thumatha kuehnei (Figs 3—4) is superfi-

cially similar to 7’ kakamegae (Figs 5-6) but differs in

its brown head (it is ochreous yellow in 7’ kakamegae)

and the more diffuse forewing markings. The new spe-

cies is also reminiscent of 7! brunnea (Figs 7-8) which

is so far only known from males, but is distinguished by

its paler forewing ground color. Compared to 7. kakame-

gae (Figs 22—23), the male genital capsule of 7’ kuehnei

(Figs 24—25) has a dorso-ventrally more dilated uncus,

distally less dilated valva with somewhat narrower ven-

tro-distal process. Additionally, the medial costal process

of the valva is somewhat shorter (in proportion to the val-

va length) and directed more or less distally in the new

species whereas that of 7’ kakamegae is directed rather

dorso-distally. The aedeagus of 7? kuehnei is more dilat-

ed distally than that of 77 kakamegae. In the vesica of

the new species, the main chamber is markedly longer

and wider than in 7’ kakamegae (in proportion to the ae-

deagus dimensions), the medial diverticulum is broad-

er bearing fewer, somewhat more robust cornuti, and

the distal diverticulum is reduced and replaced by two

short but robust cornuti (whereas in 7’ kakamegae, the

distal diverticulum is elongate bearing a row of numer-

ous smaller cornuti). The female genitalia of 7? kuehnei

(Fig. 36) differ from those of 7’ kakamegae (Fig. 35) in

the larger papillae anales (in proportion to the ovipos-

itor), the somewhat narrower antrum, and the laterally

more heavily sclerotized appendix bursae. Compared

to 7: brunnea (Figs 37-38), the female genitalia of the

new species have a wider, funnel-like antrum (whereas

in 7: brunnea, it has almost parallel lateral margins and

triangular lateral processes directed posteriorly), a mark-

edly wider posterior sclerotized section of the corpus

bursae, and a conspicuously wider and longer appendix

bursae.

Description

External morphology of adults (Figs 3-4). Forewing

length 11.0 mm in males and 13.5 mm in males. Sex-

ual dimorphism limited: Female somewhat larger than

male with forewing having more pointed apex. Antenna

yellow, bipectinate in male and ciliate in female. Head

ochreous brown. Thorax ochreous brown. Patagia dark

brown. Tegula dark brown basally and ochreous brown

distally. Forewing ground color ochreous brown, mark-

ings dark brown. Costal half of subbasal area with in-

tense dark brown suffusion. Antemedial line thick, dif-

fuse, its anal half inwardly oblique. Medial line wide,

diffuse, somewhat inwardly oblique between cell and

anal margin. Discal spot small, round, diffuse. Postme-

dial line irregularly zigzagged, thin, diffuse. Subtermi-

nal line thin and diffuse, irregularly zigzagged, dilated

into an amorphous costal patch. Terminal line interrupted

constituting indistinct spots between veins. Cilia ochre-

Bonn zoological Bulletin 70 (2): 317-331

ous brown. Hindwing and cilia paler than forewing, pale

ochreous yellow. Medial transverse line indistinct, wavy,

comprised of slight dark brown suffusions. Abdomen

ochreous brown with admixture of brown scales medi-

ally.

Male genitalia (Figs 24—25). Uncus elongate, swollen

but somewhat flattened laterally, strongly dilated medi-

ally, tapered distally with tiny claw-like tip. Tuba ana-

lis membranous, scaphium thin and weakly sclerotized.

Tegumen 1.7 x shorter than valva, its arms moderately

wide, fused in distal third. Vinculum ca. 1.2 x longer than

tegumen, saccus short (ca. 7/s of vinculum length), mod-

erately sclerotized, U-shaped. Juxta weakly sclerotized,

wide, shield-like with a wide but shallow medio-distal

depression. Valva lobe-like, strongly dilated distally.

Costa narrow, convex medially and curved ventrad sub-

apically, its distal section thin, fused dorsally with distal

membranous lobe of valva. Medial costal process long,

apically rounded, distally gradually tapering with wide

triangular base somewhat protruding beyond the costa

dorsally. Ventral protrusion of costa projecting from sub-

apical section of costa, wide, dilated ventrally, fused with

base of ventro-distal process of valva, bearing a small,

setose bulge medio-distally. Ventro-distal process of val-

va triangular with rounded tip. Distal membranous lobe

of valva wide, short with convex outer margin. Sacculus

narrow, short (ca. 1.5 x shorter than ventral margin of

valva), weakly setose dorsally. Aedeagus dilated distally

with short and narrow coecum and serrulate ventro-dis-

tal section protruding distally. Main chamber of vesica

broad, curved dorsad. Medial diverticulum large, sack-

like, bearing 3—5 short, wide-based cornuti. Distal diver-

ticulum reduced, replaced with two short, acute, wide-

based cornuti. Vesica ejaculatorius constricted basally,

weakly granulated subbasally, directed anteriad.

Female genitalia (Fig. 36). Papillae anales trapezoi-

dal with rounded corners, weakly setose. Apophyses

elongate and thin, apophyses posteriores ca. twice as

long as apophyses anteriores. Antrum short, wide, fun-

nel-shaped, moderately sclerotized and granulated. An-

terior section of corpus bursae moderately sclerotized,

strongly asymmetrical, its right side equal in length with

antrum, left side very short. Posterior section of corpus

bursae ca. half as long as anterior one, asymmetrical with

left side more heavily sclerotized than right side with

convex lateral margin. Anterior section of corpus bursae

globular, membranous. Appendix bursae positioned pos-

tero-ventrally, broadly conical, moderately sclerotized

with membranous tip, directed postero-laterally on right

side.

Distribution. The species is known from north-west-

ern Zambia.

Etymology. The species is named after Mr Lars Kthhne

(Potsdam, Germany), a renowned lepidopterist and au-

thor of the related species 7. kakamegae and T: brunnea.

©ZFMK

324 Anton V. Volynkin

T. kakamegae, PT T. kakamegae

Kenya, Kakamega Forest, slide AV4267 Uganda, Kibale National Park, slide AV4268

T. kuehnei sp. n., HT T. kuehnei sp. n., PT

Zambia, Hillwood, Ikelenge, slide AV33 14 Zambia, Hillwood, Ikelenge, slide AV6384

Figs 22-25. Thumatha spp., male genitalia. Depositories of the specimens dissected: 22—23. =ANHRT; 24—25. = MEN.

Bonn zoological Bulletin 70 (2): 317-331 ©ZFMK

Four new Thumatha from Africa 325

Thumatha ngewo sp. n.

(Figs 9-10, 26-29, 39)

urn: lsid:zoobank. org: act: 20EBC3 10-B334-4521-8617-F C946 EA4A958

Type material

Holotype (Figs 9, 26). 3, “Liberia, 530m, Lofa County,

Foya Proposed Protected Area, 7°56'36"” N 10°16'36" W,

10-19.x1.2017, MV Light Trap (125w), Aristopha-

nous, M., Safian, Sz., Simonics, G. & Smith, L. leg.,

ANHRT:2017.33”, ANHRT unique number: 00020068,

gen. prep. No.: AV4595 (ANHRT).

Paratypes. LIBERIA: 7 3, 2 29, 140 m, Sinoe

County, Krahn-Bassa Reserve, Juboe River, 7.5 km SW

Pellokon Town, 5°39'4" N, 8°39'4" W, 14—20.1.2018, MV

Light Trap, Geiser, M., Safian, Sz., Simonics, G. leg.,

ANHRT:2017.33, ANHRT unique numbers: 00099497,

00099512, 00099526, 00100802, 00101894—00101896

($3), 00089715, 00101929 (2), gen. prep. Nos.:

AV5648 (3), AV5649 (2) (ANHRT). GUINEA:

2 Jo, 540-600 m, 619 km ESE of Conakry, Nzerekore

Region, Prefecture de Lola, Ziela env., x.2017, 7°42' N,

8°21' W, Local collectors leg., ANHRT:2020.6, ANHRT

unique numbers: 00192371, 00192372, gen. prep. No.:

AV6386 (ANHRT). [IVORY COAST: 174 m, Tai NP,

Tai Research Station, 05°49'59.8" N, 07°20'32” W, 5-10.

vui.2015, Light Trap, Aristophanous, M., Moretto, P., Ru-

zzier, E. leg., ANHRT:2017.14, ANHRT unique number:

00108987, gen. prep. No.: AV6385 (ANHRT).

Diagnosis. The new species (Figs 9-10) is vaguely

reminiscent externally of 7) /unaris (Figs 12-13) but

is distinguished by the larger size, the paler forewing

ground color, the more intense blackish brown subbas-

al suffusion of the costa and the darker blackish brown

subapical spots. The male genital capsule of 7’ ngewo

(Figs 26-29) is similar to that of 7? /unaris (Fig. 31)

but differs in the distally more dilated valva, the medi-

ally more convex costa, the more angular apex and the

somewhat more elongate distal lobe of the valva. The

ventro-distal process of the valva of the new species is

tapered distally and pointed apically whereas it is broader

distally and rounded apically in 7: /unaris. Additionally,

the juxta of 7’ ngewo is wider than in 7. /unaris and the

saccus 1s much broader and more rounded. The aedeagus

of 7’ ngewo 1s conspicuously dilated distally whereas in

T. lunaris it has a more or less uniform width along its

length. The vesica of the new species is basally wider

than in 7: /unaris. The cornuti of 7’ ngewo are short with

a wide base whereas they are spine-like in 7 /unaris. In

the female genitalia of 77 ngewo (Fig. 39), the antrum

is a wide funnel-shape whereas in 7. /unaris (Fig. 40) it

is markedly narrower (in proportion to the 8" abdomi-

nal segment) with almost parallel lateral margins, a wide

medio-distal depression ventrally and triangular lateral

protrusions directed posteriorly. The anterior section of

the ductus bursae 1s reduced in the new species whereas

it is present in 7! /unaris. Additionally, the posterior scle-

Bonn zoological Bulletin 70 (2): 317-331

rotized section of the corpus bursae of 7’ ngewo is lon-

ger (in proportion to the length of the anterior section),

more swollen postero-laterally and the appendix bursae

is more elongate and basally wider than that of 7. /unaris.

Description

External morphology of adults (Figs 9-10). Fore-

wing length 9.0 mm in males and 12.0 mm in males. Sex-

ual dimorphism limited: Female somewhat larger than

male, its forewing wider with more pointed apex. Anten-

na dark brown, bipectinate with short branches in male

and ciliate in female. Head and thorax brown with ad-

mixture of black scales. Forewing ground color ochreous

brown, markings blackish brown. Antemedial line dif-

fuse, smoothly curved medially and angled outwards at

anal margin. Subbasal area with intense blackish brown

suffusion between vein Cu and costa. Medial line wide

forming an amorphous patch between costa and vein Cu,

curved inwards in cell, narrow and indistinct medially

and posteriorly. Discal spot small but distinct, elliptical

in male and more or less round in female. Postmedial line

thin and indistinct with a loop-like curve anteriorly and

inwardly oblique posteriorly. Subterminal line interrupt-

ed forming a large trapezoid spot on the costa, a small-

er elliptical spot opposite cell, and series of indistinct

tiny spots on veins posteriorly. Terminal line interrupted

constituting small indistinct spots between veins. Cilia

ochreous brown with admixture of dark brown scales

opposite the spots of terminal line. Hindwing and cilia

ochreous brown, somewhat paler than forewing. Postme-

dial transverse line consisting of weak brown suffusion

in the male. Abdomen brown with admixture of black

scales medially and distally.

Male genitalia (Figs 26-29). Uncus elongate, swol-

len, dilated distally with small claw-shaped tip and short

dorsal crest. Tuba analis membranous, scaphium thin

and weakly sclerotized. Tegumen half as long as valva,

its arms moderately wide, fused in distal third. Vincu-

lum ca. 1.5 x longer than tegumen, saccus short (ca.

of vinculum length), moderately sclerotized, U-shaped.

Juxta weakly sclerotized, shield-like with triangular me-

dio-distal depression. Valva lobe-like, dilated distally,

costa narrow, convex medially and strongly angled ven-

trad subapically, its distal section thin, fused with distal

membranous lobe of valva dorsally. Medial costal pro-

cess elongate, finger-shaped, somewhat curved distally

with rounded tip. Ventral protrusion of costa projecting

subapically, dilated ventrally, fused with base of ven-

tro-distal process of valva, bearing small, setose bulge

medio-distally. Ventro-distal process of valva triangular

with rounded tip. Distal membranous lobe of valva wide,

almost triangular with rounded apex. Sacculus narrow,

short (ca. half as long as ventral margin of valva), setose

dorsally. Aedeagus dilated distally with short and nar-

row coecum. Main chamber of vesica almost globular,

bearing a cluster of short, triangular cornuti ventrally and

©ZFMK

326 Anton V. Volynkin

T. ngewo sp. n., HT T. ngewo sp. n., PT

Liberia, Lofa County, Foya, slide AV4595 Liberia, Sinoe County, Krahn-Bassa Reserve, slide AV5648

T. ngewo sp. n., PT T. ngewo sp. n., PT

Guinea, Nzerekore Region, slide AV6386 Ivory Coast, Tai National Park, slide AV6385

Figs 26—29. Thumatha ngewo sp. n., male genitalia. The specimens dissected are deposited in ANHRT.

Bonn zoological Bulletin 70 (2): 317-331 ©ZFMK

Four new Thumatha from Africa 327

latero-subbasally. Distal chamber of vesica somewhat

narrower than main chamber, curved dorsad, bearing

a cluster of short triangular cornuti subapically. Vesica

ejaculatorius constricted basally, weakly granulated sub-

basally, directed anteriad.

Female genitalia (Fig. 39). Papillae anales trapezoidal

with rounded corners, weakly setose. Apophyses anteri-

ores and posteriores elongate and thin, equal in length,

apophyses anteriores somewhat dilated apically. Ductus

bursae represented only by antrum, anterior section re-

duced. Antrum short and wide, funnel-like, moderately

sclerotized and granulated. Posterior section of corpus

bursae somewhat narrower than anterior section, asym-

metrical with left side more heavily sclerotized and hav-

ing more convex lateral margin than right side. Anterior

section of corpus bursae ovoidal, membranous. Appen-

dix bursae positioned postero-laterally on right side of

posterior section of corpus bursae, conical, sclerotized

basally and membranous apically.

Distribution. The species is known from Guinea, Li-

beria and Ivory Coast.

Etymology. Ngewo is the supreme god in the tradi-

tional beliefs of the Mende people inhabiting Sierra Le-

one and Liberia.

Thumatha jiwundu sp. n.

(Figs 11, 30)

urn. |sid:zoobank. org: act:62DB7CF4-BB9A-4924-A D58-5648593F7D00

Type material

Holotype (Figs 11, 30). 4, “Zambia, 1340m, Jiwundu

Swamp (Miombo / Riverine forest mosaic), 11°51'54" S,

25°33'20" E, 29.x-4.xi.2018, LepiLED Light Trap,

Aristophanous, M., Derozier, V., Laszlo, G., Oram, D.

leg., ANHRT:2018.40”, ANHRTUK unique number:

00056957, gen. prep. No.: AV5082 (ANHRT).

Diagnosis. Thumatha jiwundu (Fig. 11) can easi-

ly be distinguished from the closely related 7’ /unaris

(Figs 12—13) by its somewhat larger size (the male wing-

span is 17.5 mm versus 15.0—17.0 mm in 7: /unaris), the

black head, tegulae and patagia (brown in 7° /unaris), the

much darker forewing with indistinct transverse lines

due to the intense black suffusion (in 7: /unaris, the fore-

wing is pale brown with more distinct brown markings)

and the larger, round discal spot on the forewing which

is dot-like in 7. /unaris. The male genital capsule of the

new species (Fig. 30) is very similar to that of 7: /unaris

(Fig. 31) but differs in the somewhat shorter uncus and

tegumen, the longer vinculum (both in proportion to the

valva length), the more or less rectangular saccus (it is

U-shaped in 7: /unaris), and the somewhat less convex

costal margin of the valva. The aedeagus of 7) jiwundu

is more dilated distally than in 7: /unaris. The subbasal

chamber of the vesica of 7: jiwundu is broader than in

T: lunaris (in proportion to the aedeagus width) and lack-

Bonn zoological Bulletin 70 (2): 317-331

ing spinulose scobination whereas that of 7: /unaris bears

a cluster of spinules distally.

Description

External morphology of adult (Fig. 11). Forewing

length 9.0 mm in holotype male. Male antenna dark

brown, bipectinate with short branches. Head black.

Thorax dark brown, tegulae and patagia black. Forewing

ground color brown with intense black suffusion along

costa. Transverse lines indistinct consisting of narrow

areas of black suffusion, darker at costa. Discal spot dif-

fuse, round. Terminal line interrupted constituting spots

between veins. Cilia brown with admixture of black

scales between veins. Hindwing ground color and cilia

pale ochreous brown. Medial transverse line diffuse, con-

stituting very slight suffusion of black scales. Abdomen

brown with admixture of black scales subapically

Male genitalia (Fig. 30). Uncus elongate, swollen, di-

lated distally with small claw-shaped tip and short dor-

sal crest. Tuba analis membranous, scaphium thin and

weakly sclerotized. Tegumen half as long as valva, its

arms moderately wide, fused in distal third. Vinculum ca.

1.3 x longer than tegumen, saccus short (ca. 4 of vincu-

lum length), moderately sclerotized, somewhat tapered

distally with rectangular tip. Juxta weakly sclerotized,

shield-like with elongate and somewhat tapered apex.

Valva lobe-like, slightly dilated medially and distally

with narrow, medially slightly convex costa, its apex

thin, fused with distal membranous lobe of valva dor-

sally. Medial costal process elongate, curved, somewhat

tapered distally, rounded apically. Ventral protrusion of

costa projecting subapically, dilated ventrally, fused with

base of ventro-distal process of valva, bearing a small, se-

tose bulge medio-distally. Ventro-distal process of valva

narrowly triangular with rounded tip. Distal membranous

lobe of valva wide but short with convex outer margin.

Sacculus narrow, short (ca. half as long as ventral margin

of valva), weakly setose dorsally. Aedeagus narrow with

short and rounded coecum, somewhat dilated distally, its

distal section with a cluster of spines directed distally ex-

tending over the base of vesica. Main chamber of vesica

short, elliptical with three short spine-like cornuti later-

ally. Distal chamber of vesica somewhat narrower than

main chamber, tubular, curved anteriad bearing a cluster

of short spine-like cornuti distally. Vesica ejaculatorius

weakly granulated subbasally and medially, directed an-

teriad.

Female unknown.

Distribution. The species is known only from

north-western Zambia.

Etymology. The species is named after its type locali-

ty, Jiwundu Swamp.

©ZFMK

328 Anton V. Volynkin

T. jiwundu sp. n., HT T. lunaris

Zambia, Jiwundu Swamp, slide AV5082 South Sudan, White Nile, slide AV6389

T. punctata, HT T. punctata

NE Namibia, E Capriyi, slide 688 Kiihne Zambia, Jiwundu Swamp, slide AV5079

Figs 30—33. Thumatha spp., male genitalia. Depositories of the specimens dissected: 30, 33.=ANHRT; 31. =OQUMNH;:

32 = MEN.

Bonn zoological Bulletin 70 (2): 317-331 ©ZFMK

Four new Thumatha from Africa 329

34 35 36

T. smithi sp. n., PT T. kakamegae, PT T. kuehnei sp. n., PT T. brunnea, HT

Zambia, Hillwood, Ikelenge, W Kenya, Kakamega Forest, Zambia, Jiwundu Swamp, Rwanda, Akagera River,

slide AV5646 (after Kiihne 2007) slide AV5650 slide 438 Kiihne

38 39 io 40 41

T. brunnea T. ngewo sp. n., PT T. lunaris I. punctata

Rwanda, Butare, Liberia, Sinoe County, Cameroon, Littoral Region, Zambia, Jiwundu Swamp,

slide ZSM Arct. 2020-207 Volynkin slide AV5649 slide ZSM Arct. 2020-206 Volynkin slide AV5651

Figs 34—41. Thumatha spp., female genitalia. Depositories of the specimens dissected: 34, 36, 39, 41. = ANHRT;

35. = LKP (after Kthne 2007); 37. = LKP (photo by L. Kthne); 38, 40. = ZSM.

Bonn zoological Bulletin 70 (2): 317-331 ©ZFMK

330 Anton V. Volynkin

Thumatha lunaris Durante, 2007

(Figs 12-13, 31, 40)

Thumatha lunaris Durante, 2007 (Durante 2007: 86,

figs 3-6, 15-17, 20, 22).

Type locality. “Nigeria, Rivers State, Port Harcourt”.

Material examined. CAMEROON: 1 9&2, Camer-

oun, 8 km N d’Edea, 30.11i—-1.1v.1970, C. Herbulot, gen.

prep. No.: ZSM Arct. 2020-206 (prepared by Volynkin)

(ZSM). SOUTH SUDAN: 1 3, 998, Feb. 11 1912, White

Nile, Lat. 6°6' N, Malek, G.B. Longstaff / K.998 Malek

11.2.12/1912 4221, gen. prep. No.: AV6389 (OUMNH).

Remark. The species was described from southern N1-

geria and 1s currently known only from its type locality.

The male specimen from South Sudan and the female

specimen from Cameroon are similar to the paratypes il-

lustrated by Durante (2007: figs 3-6). The detailed com-

parison with 7! jiwundu and T: punctata is provided under

the diagnosis sections of these species.

Distribution. The species is known from southern

Nigeria (Durante 2007), south-western Cameroon and

South Sudan (new country records).

Thumatha punctata Kithne, 2010

(Figs 14-18, 32-33, 41)

Thumatha punctata Kithne, 2010 (Kthne 2010: 442 (de-

scription), 453 (male genitalia), pl. 27, fig. 25 (adult)).

Type locality. “Namibia, E of Caprivi Katima Mulilo”.

Type material examined

Holotype (Figs 14, 32). 3, blue label “Namibia-Exp.

ZMB 1992, E. Capriyi: Katima Mulilo, 17°29' S/24°17'E,

lux, 3.—8.JII.[19]92, leg. W. Mey” / red label “Holotypus

Thumatha punctata by L. Kuthne” / “L. Kthne Dauer-

praparat No. 688” (MEN).

Additional material examined. ZAMBIA: 3 <3,

1 2, 1340m, Jiwundu Swamp (Miombo / Riverine for-

est mosaic), 11°51'54" S, 25°33'20" E, 29.x—4.xi.2018,

Actinic Light Trap, Aristophanous, M., Dérozier, V.,

Laszlo, G., Oram, D. leg., ANHRT:2018.40, gen. prep.

Nos.: AV5079, AV5081, AV5647 (33), AV5651 (QQ)

(ANHRT), 2 do, 2 29, 1340m, Jiwundu Swamp,

11°51'54" §S, 25°33'20" E, 21-24.xi.[20]14, Light

Trap, leg. Smith, R. & Takano, H., ANHRT:2017.12,

gen. prep. Nos.: AV4497, AV4512 (33), AV4492 (9)

(ANHRT); 1 male, 1189m, Kalambo Falls, 30 km North

of Mbala, Northern Province, 08°35'50" S, 31°14'26" E,

9-11.v.2019, MV Light Trap, Dérozier, V., Laszlo, G.,

Miles, W. leg., ANHRT:2019.12, gen. prep. No.: AV6387

(ANHRT).

Diagnosis. The forewing length is 7.5-10.0 mm in

males and 11.5—12.5 mm in males. Males of the species

(Figs 14-17) significantly vary in size within the same

population. The species is externally reminiscent of

T. smithi, T: lunaris and T. ngewo but can easily be distin-

Bonn zoological Bulletin 70 (2): 317-331

guished by the distinct black discal spot on the hindwing

which is absent in its congeners. The male genital cap-

sule (Figs 32—33) is most similar to those of 7) /unaris

(Fig. 31) and 7. jiwundu (Fig. 30) but in 7? punctata, the

costa 1s more convex subapically, the medial costal pro-

cess 1s shorter and distally thinner, and the ventro-distal

process of the valva is markedly shorter. The aedeagus of

7. punctata is thicker with an area of short dentation api-

cally compared to 7. /unaris and T. jiwundu, which bear

a dense cluster of elongate spines directed distally in the

distal section of the aedeagus. The vesica of 7? punctata

is broader than in 7. /unaris and T: jiwundu, lacking a

small medial cluster of cornuti which is present in the

other two species, and having a wider distal cluster of

more numerous and more robust but short cornuti. The

female genitalia of 7. punctata (Fig. 41) differ from those

of 7. Junaris (Fig. 40) in the somewhat longer apophyses

anteriores (in proportion to the ovipositor), the shorter

and more rectangular antrum having a shallower me-

dio-ventral depression, and the longer and more heavily

sclerotized posterior section of the corpus bursae.

Description of female

External morphology of adult (Fig. 18). Female larg-

er than male with serrate antenna (bipectinate with short

branches in male) and more elongate forewing having

less convex costal margin and more pointed apex. Head

and thorax dark brown with admixture of black scales.

Forewing ground color brown. Forewing markings

black, more diffuse than in male. Sub-costal area with

three wide spots of black suffusion subbasally, medially

and subapically. Discal spot round, distinct. Subterminal

line indistinct, zigzagged. Terminal line diffuse, inter-

rupted, in the form of small spots between veins. Cilia

brown. Hindwing and cilia paler than forewing, ochreous

brown. Discal spot semilunar with diffuse margins. Ab-

domen brown

Female genitalia (Fig. 41). Papillae anales rectangular

with rounded corners, setose. Apophyses anteriores and

posteriores elongate and thin, equal in length. Antrum

relatively wide, rectangular, its ventral margin with wide

but shallow posterio-medial depression. Anterior section

of ductus bursae somewhat longer than antrum, tapered

anteriorly, sclerotized. Posterior section of corpus bursae

somewhat longer than ductus bursae, heavily sclerotized

with wide rounded lateral protrusion on the left side. An-

terior section of corpus bursae ca. twice as long as pos-

terior section, teardrop-shaped, membranous. Appendix

bursae reduced.

Distribution. The species is known from north-east-

ern Namibia (Kihne 2010) and Zambia (new country

record).

Acknowledgements. I express my sincere thanks to the fol-

lowing colleagues for their kind assistance during the visits

to collections under their care: Dr Wolfram Mey (MEN), Dr

©ZFMK

Four new Thumatha from Africa 331

Axel Hausmann, Dr Wolfgang Speidel and Mr Ulf Buchsbaum

(ZSM), and Dr James Hogan (OUMNH). I am also grateful to

Mr Lars Ktshne (Potsdam, Germany) for providing me with pic-

tures of specimens of 7) brunnea and T: kakamegae from his

private collection.

The following collaborative partners and authorities are

thanked for the diverse administrative and technical assistance

provided during the field work. Guinea: Mr Mamadou Diawara

(Guineé Ecologie), Colonel Layaly Camara and Mr Cece Papa

Konde (Ministere de |’Environment et aux Eaux et Foréts),

Mr Jamison Suter (Societe des Mines de Fer de Guineé —

Responsibilite Environmentale et Social). Ivory Coast: Scien-

tific Research in Céte d’Ivoire was authorised by the Ministere

de l’Enseignement Supérieur et de la Recherche Scientifique.

The Office Ivoirien des Parcs et Réserves (OIPR) and the

Société de Développement des Foréts (SODEFOR) are thanked

for authorising access to protected forests and providing export

permits. Liberia: Mr Mike Doryen, Mr Darlington Taugben and

Mr Kederick F. Johnson (Forestry Development Authority).

Zambia: Ms Rhoda Kachali (Department of National Parks and

Wildlife - ZAWA, Lusaka), Ms Claire Mateke and Ms Martha

Imakando (Livingstone Museum, Livingstone).

The author declares that to the best of his knowledge he con-

forms to the national regulations and meets with the conditions

and requirements of International Conventions concerning col-

lecting/export and handling of the specimens presented in this

Article.

REFERENCES

Butler AG (1896) On two collections of lepidoptera made by

Mr R. Crawshay in Nyasa-land. Proceedings of the general

meetings for scientific business of the Zoological Society of

London 1896: 817-850

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Draudt M (1914) Arctiidae. Tiger moths (Miltochrista to Ste-

nosia). Pp. 135-223 in: Seitz A (ed.) Die Gross-Schmetter-

linge der Erde: eine systematische Bearbeitung der bis jetzt

bekannten Gross-Schmetterlinge. Volume 10. Alfred Kernen,

Stuttgart

Durante A (2007) I lepidotteri del Delta del Niger (settimo

contributo). Genus Thumatha Walker (Arct., Lithos.), con

descrizione della nuova specie Thumatha lunaris. Thalassia

salessina 30: 81-92

Durante A (2009) Individuazione del gruppo fuscescens Walker

all’interno del genere Thumatha Walker (Lepidoptera, Arcti-

idae, Lithosiinae). Lambillionea 109: 123—127

Hampson GF (1900) Catalogue of the Arctiadae (Nolinae,

Lithosianae) in the British Museum. Catalogue of the Lepi-

doptera Phalaenae in the British Museum 2: 1-590

Holloway JD (2001) The Moths of Borneo, part 7. Family Arc-

tiidae, subfamily Lithosiinae. Malayan Nature Journal 55:

279-486

Kuthne L (2007) Beschreibung neuer Flechtenbararten aus Af-

rika nebst taxonomische Anmerkungen (Arctiidae: Lithosi-

inae). Esperiana Memoir 3: 353-394

Kuhne L (2010) Taxonomische Ergebnisse der Bearbeitung der

Nachtfalterfauna des stidlichen Afrikas (Lepidoptera, Noc-

tuoidea). Esperiana Memoir 5: 433-456

Lucas TP (1892) On twenty new Species of Australian Lepi-

doptera. Proceedings of the Linnean Society of New South

Wales. Series 2 7 (2): 249-266

Volynkin AV (2020) Thamoma, a new genus for Miltochrista

vetusta Piepers & Snellen, 1904 from Java (Lepidoptera, Er-

ebidae, Arctiinae). Ecologica Montenegrina 35: 138-143

Walker F (1866) Catalogue of Lepidoptera Heterocera. Seventh

Series. List of the Specimens of Lepidopterous Insects in the

Collection of the British Museum 35: 1535-2040

Witt TJ, Speidel W, Ronkay G, Ronkay L, Laszl6 GM (2011)

Subfamilia Arctiinae. Pp. 81—217 in: Witt TJ, Ronkay L (eds)

Lymantriinae and Arctiinae, including Phylogeny and Check

List of the Quadrifid Noctuoidea of Europe. Noctuidae Euro-

paeae. Volume 13. Entomological Press, Soro

©ZFMK

BHL

Blank Page Digitally Inserted

Bonn zoological Bulletin 70 (2): 333-338

2021 - Volynkin A.V. & Laszlo G.M.

https://do1.org/10.20363/BZB-2021.70.2.333

ISSN 2190-7307

http://www.zoologicalbulletin.de

Research article

urn:|sid:zoobank.org:pub: CE2FA6EF-F33B-4542-844D-992 169C85D2E

Kruegerilema, a new genus for a new species endemic to S40 Tomé Island

(Lepidoptera: Erebidae: Arctiinae: Lithosiini)

Anton V. Volynkin' * & Gyula M. Laszl6

' 2The African Natural History Research Trust (ANHRT), Street Court Leominster-Kingsland, HR6 9QA, UK

‘Altai State University, Lenina Avenue, 61, RF-656049, Barnaul, Russia

“Corresponding author: Email: anton@anhrt.org.uk

'urn:Isid:zoobank.org:author:25EA 1403-9502-42BE-A820-96ACS51CE862E

*urn:lsid:zoobank.org:author:5|0CAA370-A89A-488E-A635-1B2D6FD71001

Abstract. The present paper contains a description of the new genus Kruegerilema gen. nov. which is erected for the new

species Kruegerilema smithi sp. nov. endemic to SAo Tomé Island. The diagnostic comparison is made with the genus

Brunia Moore, 1878. Adults together with male and female genitalia of the new and similar genus are illustrated.

Key words. Lithosiina, Sub-Saharan Africa.

INTRODUCTION

The recently published generic classification of the Afro-

tropical Lithosiina (footman moths) contains 98 genera

from mainland Africa (Kruger 2015) with an additional

new genus described subsequently from South Africa

(Kruger 2016). Although these publications are undoubt-

edly fundamental in the classification of the subtribe, they

inevitably do not provide a perfect overview of the group

due to the complexity of the Afrotropical Lithosiina. Un-

resolved taxonomic problems are especially prevalent in

poorly studied regions such as West Africa and the Congo

Basin with the generic assignment of several groups of

Lithosiina still uncertain.

In the course of identifying the Lithosiini housed in

the African Natural History Research Trust, Leominster,

large series of a peculiar, dark brown Lithosiina species

from Sao Tomé Island was found. As no reference of

the occurrence of any Lithosiina in Sao Tomé could be

traced, the species was in the first instance believed by

the authors to be an undescribed Brunia Moore, 1878.

Having compared the genitalia morphology of the new

taxon with the Oriental type species of Brunia and its

Afrotropical synonym Ovenna Birket-Smith, 1965, sev-

eral remarkable autapomorphies were discovered sug-

gesting that the species belongs to a distinct monophylet-

ic lineage that not only represents a hitherto undescribed

species but also a new genus, the descriptions of which

are provided herein.

Received: 15.06.2021

Accepted: 09.07.2021

MATERIAL AND METHODS

Abbreviations of the depositories

ANHRT = African Natural History Research Trust,

Leominster, UK

NHMUK = Natural History (formerly BMNH)

Museum, London, UK

Other abbreviations

AV = genitalia slide prepared by A.V. Volynkin

HT = holotype

PT = paratype

The genitalia were dissected and mounted in euparal on

microscope slides. The photos of adults were taken us-

ing a Nikon D3100/AF-S camera equipped with a Nikkor

18-55 mm lens while the photos of genitalia were tak-

en by the same camera attached to a microscope with an

LM-scope adapter. All images were processed using the

Adobe Photoshop CC 2018 software.

RESULTS

Kruegerilema gen. nov.

urn: lsid:zoobank. org: act: BS 1F553E-2A 78-424 5-S8E41-FFE66D56737F

Type species. Kruegerilema smithi sp. nov.

Diagnosis. The new genus (Figs 1-2) is external-

ly similar to the genus Brunia Moore, 1878 (=Ovenna

Birket-Smith, 1965) (Figs 3-4) but distinguished by a

Corresponding editor: M. Espeland

Published: 17.07.2021

334

number of diagnostic features. The sexual dimorphism

is limited with the female having a wider forewing and

a smaller head (in proportion to the body size) than the

male whereas in Brunia, the sexual dimorphism is sub-

stantial: males have an intense dark terminal suffusion

of the forewing, while in the female the ground colour is

monotonous but with a paler costa which may be bright

yellow in certain species.

The male genital capsule of Kruegerilema (Figs 5—6) is

similar to that of Brunia (Fig. 7) (the type species, Brunia

antica (Walker, 1854) is illustrated by Holloway (2001)

and Bucsek (2017)) in the relatively short and thick uncus,

the V-shaped vinculum, the relatively narrow dorsal part

of the valva curved ventrad and the presence of a short

ampulla (processus momenti sensu Birket-Smith (1965)).

The new genus is distinguished from Brunia by certain

autapomorphic features such as the highly modified un-

cus with a subbasal ventral process and the presence of

a heavily sclerotized, wide subscaphium, in contrast to

Brunia’s unmodified uncus and tuba analis lacking a sub-

scaphium. Additionally, the scaphium of Kruegerilema is

relatively thick and distally extended to a wide, densely

dentate area whereas the scaphium 1s membranous along

its full length in Brunia. The distal saccular process of the

new genus is simple, relatively narrow and elongate with

smooth margins while it is considerably shorter (in pro-

portion to the dorsal part of the valva), wider and com-

plex, bearing additional processes or strong dentation

in Brunia. The aedeagus of Kruegerilema is relatively

narrow, somewhat tapered distally and having a narrow

carinal cluster of short spines whereas the aedeagus of

Brunia 1s markedly wider, dilated distally with a longitu-

dinally rugose and expandable scobinated apical section

(an autapomorphic feature of the genus). The vesica of

Kruegerilema 1s shorter than the aedeagus, bearing finely

granulated areas only, whereas in Brunia it is markedly

wider and longer than the aedeagus and armed with a few

large, robust thorn-like or lanceolate flattened cornuti

which 1s a further autapomorphic feature of the genus.

In the female genitalia (Fig. 8), the long and complex

postvaginal plate bearing a heavily sclerotized, rounded

swollen postero-medial protrusion is an autapomorphic

character whereas the postvaginal plate of Brunia (Fig. 9)

is short (in proportion to the length of the 8" sternite), un-

modified and plate-like.

Description

Adult (Figs 1—2). Forewing length 10-13 mm in males

and 12—15 mm in females. Sexual dimorphism limited:

female somewhat larger with smaller head and wider

forewing than in the male. Antenna densely ciliate in the

male, weakly ciliate in the female. Body brown. Fore-

wing narrow, elongate with costal margin convex sub-

apically and anal margin convex subbasally. Forewing

ground colour and cilia brown. Discal spot small, dark

Bonn zoological Bulletin 70 (2): 333-338

Anton V. Volynkin & Gyula M. Laszlo

brown, round. Hindwing monotonous greyish brown.

Cilia brown.

Male genitalia (Figs 5—6). Uncus short, thick, distally

curved ventrad, apically pointed, subbasally with a short,

narrow, apically rounded process projecting ventrad. Teg-

umen shorter than valva, weakly sclerotized with wide

arms fused in distal third. Tuba analis wide (width ca. 2/3

of tegumen length). Scaphium thin, bearing an extensive,

densely dentate terminal area. Subscaphium wide, heav-

ily sclerotized, dorso-ventrally flattened, plate-like with

deep triangular incision basally. Vinculum somewhat

longer than tegumen with narrow, V-shaped saccus. Juxta

weakly sclerotized, shield-like with shallow depression

distally. Valva elongate and narrow, costal margin slightly

concave proximally, convex distally, dorsal section dis-

tally tapered, curved ventrad, apically pointed. Ampulla

(processus momenti) short, tubercle-like, weakly setose.

Sacculus narrow (ca. half of valva width) with weakly

setose dorsal surface. Distal saccular process long, heav-

ily sclerotized with a claw-shaped, dorsally curved and

apically pointed distal section. Aedeagus tubular, some-

what shorter than length of tegumen-vinculum complex,

coecum short and rounded, distal section slightly tapered

bearing a narrow lateral cluster of short spines apically.

Vesica shorter than aedeagus, its main chamber sack-like,

somewhat dilated distally, granulated basally and distally

but membranous medially. Ventral subbasal diverticulum

broad, semi-globular, granulated. Dorsal subbasal diver-

ticulum elongate, conical and apically rounded, weakly

granulated. Smaller medial diverticulum short, conical

and apically rounded, scobinated. Larger medial divertic-

ulum globular, membranous. Vesica ejaculatorius located

subbasally, directed dorsad.

Female genitalia (Fig. 8). Papilla analis trapezoidal

with rounded corners, weakly setose. Apophyses me-

dium long, very thin, equal in length. Postvaginal plate

complex, composed of a sclerotized, proximally round-

ed, quadrangular and swollen anterior plate, a narrow

bar-like, very strongly sclerotized posterior plate con-

junct with a conspicuous, round medio-distal protrusion,

and two wide, quadrangular lateral plates. Ostium bursae

equal in width with anterior section of postvaginal plate.

Ductus bursae membranous, slightly longer than anterior

section of postvaginal plate, somewhat tapered anterior-

ly. Corpus bursae subdivided into two sections. Posterior

section heavily sclerotized with evenly convex lateral

margins. Anterior section membranous, ca. 1.7 times lon-

ger and ca. two times wider than posterior one, globular,

bearing two small round signa, each with short medial

thorn-like protrusion. Appendix bursae small, wide but

very short, heavily sclerotized, positioned postero-later-

ally.

Etymology. The generic name is an aggregate of

the Latin transcription of the surname Kruger and the

generic name Ei/lema Hubner, [1819]. The gender is

feminine. The new genus 1s dedicated to the memory of

©ZFMK

Kruegerilema gen. nov. for new species from Sao Tomé 335

SAO TOME 1321m

Bom Successo, Antenna

Slide Dl eieerar ete pe ie)

ANHRTUK | AV5804 67 | 2% cia 2018 wi li sh Trap

001 04657 A. Volynkin : Mies, W. Tumer, Clive R., leg.

Kruegerilema smithi gen. & sp. n., HT 3

CORO Wack Falave)

ANHRTUK Peco % ora0 16s N, a evs

00078352 \Saton or “simone G nim

. 4 Volynkin | ANHRT 2018

Brunia agonchae &

“e TOME 482m

: es gaa (montane

forest-plantation

0°14'41.7°N? 6° a 69.8°E

ANHRTUK z1311 208 | “p18 MY Ugh Trap

00019774 Wie,

Kruegerilema smithi gen. & sp. n., PT 2

Slide CAMEROON 900m

North Region, Wack i eel

ANHRTUK | AV5966 P SSH IRN

00078416 A Volynkin ok pret e o s Simonics, G. Leg.

Brunia agonchae &

10 mm

Figs 1-4. Lithosiina spp., adults. 1. Kruegerilema smithi sp. nov., holotype, 3, Sao Tomé. 2. Ditto, paratype 2, Sdo Tomé. 3. Brunia

agonchae (Pl6tz, 1880) (= Eilema subgriseola Strand, 1912, the type species of Ovenna, a junior synonym of Brunia), 3, Camer-

oon. 4. Ditto, 2, Cameroon. The specimens are deposited in coll. ANHRT.

the late Dr Martin Kriger (1963-2019), a distinguished

expert in African Geometridae and Lithosiina taxonomy

and author of the generic classification of Afrotropical

Lithosiini.

Species composition. The new genus is monotypic.

Kruegerilema smithi sp. nov.

Figs 1—2, 5-6, 8

urn: lsid:zoobank. org: act:47 DOF 1EF-0F0A-4A 0D-A4DF-B1EF9S8BF60E

Type material

Holotype (Figs 1, 5). @, “Sao Tome, 1321m, Bom

Successo, Antenna (plantation—dry scrub forest),

0°16'31''N, 6°36'14""E, 22—25.1.2018, MV Light Trap,

Aristophanous, M., Lima, D., Miles, W., Turner, Clive

R., leg. ANHRT:2018.1”, unique number: ANHRTUK

00104657, gen. prep. No.: AV5804 (ANHRT).

Paratypes (651 specimens in total). SAO TOME.

140 specimens of both sexes, same data as for holotype,

gen. prep. No.: AV5805 (2); 92 specimens of both sexes,

Bonn zoological Bulletin 70 (2): 333-338

same data but collected by actinic light trap; 1 <', Neves,

Praia das Furnas, Mucumbli Lodge (plantation-dry scrub

forest), 60 m, 0°26'56.1" N, 6°32'22.7" E, 25—27.1.2018,

MV Light Trap, Aristophanous, M., Miles, W., Turner,

C.R. leg., ANHRT:2018.1; 1 ¢, 1 9, Guadalupe, An-

tenna (dry coastal scrub forest), 254 m, 0°22'49.8" N,

6°38'47.4" E, 31.1-2.11.2018, Actinic Light Trap, Aristo-

phanous, M., Miles, W., leg., ANHRT:2018.1; 1 4, 1 9,

same data but collected by MV light trap; 284 specimens

of both sexes, Roca Bombaim (montane / forest-planta-

tion), 482 m, 0°14'41.7" N, 6°37'59.8" E, 27-31.1.2018,

MV Light Trap, Aristophanous, M., Miles, W., leg.,

ANHRT:2018.1; 7 specimens of both sexes, same data

but collected by actinic light trap; 45 specimens of both

sexes, Antenna Bom Successo, 1303 m, 00°16'33.7" N,

6°36'19.7" E, 19.x.2016, MV Light Trap, Turner, C.R. &

Tasane, T. leg., ANHRT:2017.21, gen. prep. No.: AV5969

(2); 15 specimens of both sexes, same site and collectors

but collected at 20.x.2016; 20 specimens of both sex-

es, same site and collectors but collected at 28.x.2016;

2 3d, 1 , same site and collectors but collected at

©ZFMK

336 Anton V. Volynkin & Gyula M. Laszlo

ventral subbasal

process of uncus

main chamber of vesica

larger medial

wo diverticulum

ventral subbasal diverticulum smaller medial diverticulum

Kruegerilema smithi gen. & sp. n., HT

Sao Tomé and Principe, Sao Tomé Isl., slide AV5804 Volynkin

distal dentate area

of scaphium scaphium

subscaphium

Kruegerilema smithi gen. & sp. n., PT

Sao Tomé and Principe, Sao Tomé Isl., slide AV5958 Volynkin

Brunia agonchae

Cameroon, North Region, slide AV5965 Volynkin

Figs 5—7. Lithosiina spp., male genitalia. 5. Kruegerilema smithi sp. nov., holotype, Sao Tomé. 6. Ditto, paratype, Séo Tomé.

7. Brunia agonchae (= Eilema subgriseola Strand, 1912), Cameroon. The specimens dissected are deposited in coll. ANHRT.

Kruegerilema gen. nov. for new species from S40 Tomé 337

29.x.2016; 8 specimens of both sexes, Ponta Furada,

186 m, 0°14'01.8" N, 6°28'15.5" E, 24.x.2016, MV Light

Trap, Turner, C.R. & Tasane, T. leg.;2 43’, 1 2, Mucum-

bli Lodge, Praia das Furnas, Neves, 60 m, 00°26'56.1" N,

6°32'22.7" E, 16-17.x.2016, MV, plantation with rem-

nant native dry scrub forest, C.R. Turner & T. Tasane leg.,

ANHRT:2017.21, gen. prep. No.: AV5958 (3); 1 2, same

site and collectors but collected at 30.x.2016; 9 speci-

mens of both sexes, Ponta Figo, 635 m, 00°17'12.7" N,

6°34'14.5" E, 27.x.2016, primary and secondary forest

edge along steep mountain track slope, MV, C.R. Turn-

er & T. Tasane, leg., ANHRT:2017.21 (ANHRT). | 9, St.

Thome, X.—XI.[18]99 (Mocquerys), Rothschild Bequest

B.M. 1939-1; 1 3, St. Thome, XII.[18]99 — 1.[19]00,

(Mocquerys), Rothschild Bequest B.M. 1939-1, gen.

prep. No.: Arctiidae 5638; 3 63,9 99, Sao Thome

I[sland], 18.xi.1932, W.H.T. Tams, B.M. 1933-39; 2 ¢4,

1 9, Sao Thome I[sland], 19.x1.1932, W.H.T. Tams, B.M.

1933-39, 2 3, 18. 26., Sao Thome, 10.1.—24.1.[19]26,

Edge of virgin forest, T.A. Barns (NHMUK).

Diagnosis and description. The characterisation of

the new species is provided under the description of the

genus Kruegerilema.

postero-medial protrusion

of postvaginal plate

vii "Se .

ostium bursae postvaginal plate

posterior section of ————_

corpus bursae

anterior section of

corpus bursae é

Kruegerilema smithi gen. & sp. n., PT

Sao Tomé and Principe, Sao Tomé Isl., slide AV5805 Volynkin

Distribution. The new species is endemic to SAo Tomé

Island (the Democratic Republic of Sao Tomé and Prin-

cipe).

Etymology. The new species is dedicated to Mr Rich-

ard Smith, founder and director of the African Natural

History Research Trust, who, through organising and

undertaking numerous entomological expeditions to

Sub-Saharan Africa has enabled the discovery of numer-

ous new species to science.

Acknowledgements. We express our sincere thanks to Dr Al-

berto Zilli and Mr Geoff Martin (NHMUK) for their kind assis-

tance during the visits to their institution. Mr Arlindo de Car-

valho (General Directorate for the Environment) is thanked for

granting necessary permits to the ANHRT team for sampling

insects in Sao Tomé and Principe.

The Authors declare that to the best of their knowledge they

conform to the national regulations and meet with the condi-

tions and requirements of International Conventions concern-

ing collecting/export and handling of the specimens presented

in this article.

postvaginal plate

ostium bursae

Brunia agonchae

Cameroon, North Region, slide AV5966 Volynkin

Figs 8—9. Lithosiina spp., female genitalia. 8. Kruegerilema smithi sp. nov., paratype, Sdo Tomé. 9. Brunia agonchae (= Eilema

subgriseola Strand, 1912), Cameroon. The specimens dissected are deposited in coll. ANHRT.

Bonn zoological Bulletin 70 (2): 333-338

©ZFMK

338 Anton V. Volynkin & Gyula M. Laszlo

REFERENCES

Birket-Smith J (1965) A revision of the West African Eilemic

moths, based on the male genitalia (Lep., Arctidae, Litho-

sinae; incl. gena. Crocosia, Eilema, Lithosia, Pelosia, Phry-

ganopsis a.0.). Haile Sellassie I University. Papers from the

Faculty of Sciences, Series C (Zoology) 1: 1-161

Bucsek K (2017) Contribution to the study of the genus Brunia

Moore, 1878 (Erebidae, Arctiinae, Lithosiini). Entomofauna

38 (8): 141-176

Holloway JD (2001) The Moths of Borneo, part 7. Family Arc-

tiidae, subfamily Lithosiinae. Malayan Nature Journal 55:

279-486

Bonn zoological Bulletin 70 (2): 333-338

Kruger M (2015) Generic classification of Afrotropical footman

moths sensu stricto (Lepidoptera: Erebidae: Arctiinae: Litho-

siini (partim.)). Transvaal Museum Monograph. Vol. 15. Dit-

song National Museum of Natural History, Pretoria

Kruger M (2016) Revision of Campter Kriger and description

of Campteropsis, a related new genus from South Africa

(Lepidoptera: Erebidae: Arctiinae: Lithosiini). Annals of the

Ditsong National Museum of Natural History 6: 91-108

©ZFMK

Bonn zoological Bulletin 70 (2): 339-349

2021 - Kehlmaier C.

https://do1.org/10.20363/BZB-2021.70.2.339

ISSN 2190-7307

http://www.zoologicalbulletin.de

Research article

urn:|sid:zoobank.org:pub: DA4E1 AOC-723E-4A6F-A214-955E8BBCB81F

DNA barcoding reveals an unexpected diversity in Old World Vermileonidae

(Insecta: Diptera)

Christian Kehlmaier

Senckenberg Natural History Collections Dresden, Museum of Zoology, K6nigsbriicker LandstraBe 159,

D-01109 Dresden, Germany

Email: kehlmaier@Qweb.de

urn:|sid:zoobank.org:author:3BD2A 72C-A88E-4B2F-A22C-7CF522DE6F4D

Abstract. Bayesian Inference and Maximum Likelihood were used to analyse the DNA sequences of the mitochondrial

COI gene of 69 wormlion specimens (Diptera: Vermileonidae). Most of the studied individuals belong to Old World taxa

and belong to four genera, 1.e., Vermileo (n=53), Lampromyia (n=13), Vermipardus (n=2), and Leptynoma (n=1). Species

of the genus Vermileo were resolved into two distinct groups, with the analysed Nearctic species placed in a distinct grou-

ping from the Palaearctic and Afrotropic taxa. Old World Vermileo species were divided into a western and eastern species

group. Eight unnamed genetic lineages within the Palaearctic members of the genus Vermileo were identified, separated

by a p distance of at least 0.038, and are regarded as putative species. Vermileo cylindraceus (Costa, 1844) stat. rev. is

reinstated as a valid species.

Key words. Diptera, Vermileonidae, wormlions, systematics, COI, DNA barcoding.

INTRODUCTION

The family Vermileonidae (Insecta: Diptera) originated

about 150 Ma ago in the Upper Jurassic, and thus rep-

resents an old lineage of brachyceran flies (Wiegmann

et al. 2011). Being nowadays absent from Antarctica,

Australia and South America, 63 described extant spe-

cies placed in 11 genera are distributed globally, predom-

inantly in arid habitats of mainly subtropical to temperate

climates (Nagatomi et al. 1999; Kehlmaier 2014; Swart

et al. 2015; Woodley & Swart 2017; Carles-Tolra & Cues-

ta-Segura 2020; Bueno et al. unpubl.). Whereas adults

are nectar feeding, their larvae are predators; hence their

common name ‘wormlions’. Similar to many Myrmele-

ontidae larvae known as antlions (Insecta: Neuroptera),

wormlions construct small pitfalls in fine grained soil or

sand at rain-protected sites, waiting for prey at the deep-

est point of their trap.

The discovery of the genus Vermileo Macquart, 1834,

the type genus of the family, is a story of travelling among

the old Europe. The first published mention of this genus

as “Formica-Vulpes” (literally antfox), to differentiate it

from the “Formica-Leo” (antlion larvae), was presented

in 1706 by an anonymous person (probably Monsieur

Carré; Anonymous 1706). The text briefly described the

observations made by a friend of his on the morphology,

trap building and preying of the Vermilio larvae based on

material collected in the “French countryside’. Howev-

er, it was not until the mid-18" century when its entire

Received: 04.12.2020

Accepted: 21.07.2021

life-cycle was discovered. The French scientist René An-

toine Ferchault de Réaumur (1683-1757) had searched in

vain for wormlion larvae during a long time in the vicini-

ty of Paris, and he had already abandoned hope (Réaumur

1753) when he received a parcel from his friend Rebory

(Curé de la Palud, diocese de Riez en Provence; currently

La Palud-sur-Verdon) on 11" February 1751 with strange

“worms” found on one of his walks. Being intrigued by

these little creatures and wondering who else could work

on them, Réaumur offered part of the material to the

Queen of Sweden Luise Ulrike of Prussia (1720-1782),

who was a renowned patron of science and art. Luise UI-

rike of Prussia commissioned Carl De Geer (1720-1778)

with the study of the larvae. By the time the material ar-

rived to Stockholm from Paris by stagecoach, all of the

material had perished except a single larva. Nevertheless,

De Geer (1752) was able to rear the insect until the hatch-

ing of the imago, closely observing and documenting its

development and eventually presenting a detailed and

accurate illustrated description of its life-cycle. In the

year to follow, Réaumur (1753) published his own de-

tailed account on the matter. English translations of De

Geer (1752) and Réaumur (1753) are included in Wheel-

er (1930).

With the exception of Kehlmaier (2014), taxonomic

work on the family has been exclusively based on mor-

phology. The present paper focuses on the molecular

variability of the mitochondrial cytochrome c oxidase

subunit I (COZ) gen, the so-called DNA barcode sensu

Corresponding editor: X. Mengual

Published: 27.08.2021

340 Christian Kehlmaier

Hebert et al. (2003a, b), in the wormlion genera Lampro-

myia Macquart, 1835 and Vermileo. Whereas the former

genus has partly been studied using molecular characters

(Kehlmaier 2014), no molecular analyses have been con-

ducted for the latter so far.

MATERIAL AND METHODS

Larval and adult material was collected and sent in by

numerous colleagues, who are mentioned by name in

the acknowledgements. Upon reception, living larvae

were individualised and placed in film cases half filled

with dry sand taken from local forest habitats. The sand

was sieved with an ordinary colander and deep-frozen at

-80°C for two weeks beforehand. Larvae were fed once

or twice a week with collected ants, mainly Lasius niger

(Linnaeus, 1758), or flightless Drosophila hydei Stur-

tevant, 1921 obtained from a pet shop. Vermileonidae

larvae and adults selected for molecular work were deep

frozen and pinned or submerged in pure ethanol, and re-

ceived a label with a unique DNA voucher number. If

not otherwise stated, all material is stored at the Senck-

enberg Natural History Collections Dresden. All voucher

details including ENA accession numbers are included

in Table 1.

Morphological species identification were carried out

using the works of Stuckenberg (1998) and Kehlmaier

(2014) for Lampromyia, and Stuckenberg (1965), Na-

gatomi et al. (1999) and Carles-Tolra & Cuesta-Segu-

ra (2020) for Vermileo (also see Discussion below), in

combination with the study of the primary type material

for Leptis cylindracea Costa, 1844 and Vermileo niloti-

cus Edwards, 1935. Whereas the latter 1s deposited at the

Natural History Museum (London, United Kingdom) and

could be taken on loan, the former is in the collection of

the Centro Musei delle Scienze Naturale e Fisiche (Na-

ples, Italy) and two syntypes could only be studied from

dorsal habitus photos. Costa (1844a: 85) collected the

type material nearby Camaldoli (Naples), and mentioned

additional non-type material from the Lago del Bagno on

the island of Ischia, about 30 km from the city of Naples

(Costa 1844b: 117). Part of the material included in the

current study is from the vicinity of these localities and

therefore considered conspecific.

Genomic DNA was extracted either from single legs,

the apical portion of the abdomen, or from the apical

part of larvae using the innuPREP DNA Mini Kit of An-

alytik Jena AG (Jena, Germany). CO/ sequences were

amplified using the primer pair LCO-1490 (5'-GGT-

CAACAAATCATAAAGATATTGG-3') and HCO-2198

(5'-TAAACTTCAGGGTGACCAAAAAATCA-3')

(Folmer et al. 1994), which targets a 658 bp long frag-

ment of the 5'-end of the mitochondrial CO/ gen. PCR

conditions were an initial 94 °C for 4 min, 35 cycles at

94 °C for 30 s, 50 °C for 30 s, 72 °C for 45 s, and a final

Bonn zoological Bulletin 70 (2): 339-349

elongation at 72 °C for 10 min. Each PCR was performed

with 1—5 ul of DNA extraction in a 20 ul volume (1 ul

of each primer at 10 pmol, 1 wl of dNTP-mix at 10 nmol

each dNTP, and 1 unit of Taq polymerase (Bioron DFS

Taq, Ludwigshafen, Germany), 2 wl PCR buffer 10~ incl.

Mg(Cl,, ultra-pure H,O). PCR products were visualised

on a 1% agarose gel and purified using the ExoSAP-IT

PCR Product Cleanup Reagent (Applied Biosystems,

Foster City, CA, USA; 1:20 dilution; modified protocol:

30 min at 37 °C followed by 15 min at 80 °C). For cycle

sequencing, the same forward and reverse primers were

used in a total reaction volume of 10 ul (2 pl sequencing

buffer, 1 ul premix, | pl primer at 5 pmol concentration,

0.5—6 pl of DNA template, ultra-pure H,O) with 25 cy-

cles at 96 °C for 10 s, 50 °C for 5 s and 60 °C for 4 min

using the ABI PRISM Big Dye Terminator v.3.1 Cycle

Sequencing Kit (Applied Biosystems). For subsequent

purification, Performa DTR V3 96 Well Short Plates

(Edge Biosystems, Gaithersburg, MD, USA) were used.

Sequencing was done at the Senckenberg Natural History

Collections Dresden using an ABI 3130XL or ABI 3730

Genetic Analyser.

Sequences were checked by eye for base-calling errors

and aligned in BioEdit 7.0.5.2 (Hall, 1999). No internal

stop codons, frame shifts or missing triplets were detect-

ed. The final dataset was trimmed to 637 bp and con-

sisted of 69 ingroup and four outgroup sequences. The

sequences for the outgroup taxa (three Rhagionidae and

one Xylophagidae, which was constrained as the root

for the inferred trees) were downloaded from GenBank.

Maximum Likelihood and Bayesian Inference were used

to infer the evolutionary relationships of these taxa based

on COI sequences. Best fitting partitioning schemes and

models of sequence evolution were obtained with Parti-

tionFinder 2 (Lanfear et al. 2016) and the implemented

Bayesian Information Criterion. Maximum Likelihood

(ML) analyses were conducted with RAxML (Stamatakis

2014) using the GTR+G model as recommended by Sta-

matakis (2016: 59f), and a codon-partitioned scheme. As

a preliminary step, five independent searches were per-

formed using different starting conditions together with

the fast bootstrap algorithm to explore the robustness of

the results by comparing the best trees (not figured here).

Then, 1,000 non-parametric thorough bootstrap repli-

cates (BP) were calculated and plotted on the cladogram

inferred from the Bayesian analysis (Fig. 1). Bayesian

analysis (BA) was performed using MrBayes 3.2.6 (Ron-

quist et al., 2012) with two parallel runs (each with four

chains) and default parameters using a codon partitioned

data set and the substitution models SYM+G (codon po-

sition 1), F81+I (codon position 2), and GTR+G (codon

position 3). The chains ran for 10 million generations,

with every 1000th generation sampled. Run parameters

were checked with Tracer ver. 1.7.1 (Rambaut et al.

2018). Branch support was assessed by the Bayesian pos-

terior probabilities (BPP) and indicated in Fig. 1. Where-

©ZFMK

DNA barcoding reveals an unexpected diversity in Old World Vermileonidae 341

as BP measures the robustness of character support, BPP

evaluates the most likely phylogenetic tree for the given

data and models of evolution. Although objective thresh-

olds for strong, moderate, and weak support values are a

matter of dispute (Zander 2004), BP of 80-89 and BPP

of 0.90-0.94 are regarded as moderate, and BP > 90 and

BPP > 0.95 as strong support values in the current study.

Uncorrected p distances were calculated with MEGA ver.

10.0.5 (Kumar et al. 2018).

RESULTS

New DNA barcodes of Vermileonidae were generated for

52 specimens of Vermileo, four specimens of Lampro-

myia, two specimens of Vermipardus Stuckenberg, 1960,

and one specimen of Leptynoma Westwood, 1876. Nine

sequences of Lampromyia and one of Vermileo were add-

ed from the International Nucleotide Sequence Database

Collaboration (INSDC). No molecular data is currently

available for the rest of Vermileonidae genera, namely

Alhajarmyia Stuckenberg, 2003, /salomyia Stuckenberg,

2002, Namaquamyia Stuckenberg, 2002, Perianthomyia

Stuckenberg, 1996, Vermilynx Stuckenberg, 1995, Vermi-

ophis Yang, 1979, and Vermitigris Wheeler, 1930.

The topologies of both phylogenetic analyses are large-

ly congruent with moderate to strong (BA) or weak to

moderate (ML) statistical support values (Fig. 1). The

most interesting finding of the current dataset is the dis-

covery of eight putative new species (named sp. A to sp.

H) among the Palaearctic members of the genus Ver-

mileo. These putative new species show clear differenc-

es in the CO/ sequence when compared with the other

known Vermileo species.

The Nearctic representatives of Vermileo were resolved

together as the sister group of the remaining wormlion

taxa, but not in the same clade as the other Vermileo spe-

cies. Old World Vermileo species were placed as the sis-

ter lineage to an unsupported clade containing Lampro-

myia, Leptynoma and Vermipardus. Within this grouping,

the Palaearctic Lampromyia taxa were resolved together

(BPP = 0.99; BP < 70), whereas the single Afrotropical

Lampromyia species clusters with Vermipardus without

statistical support. The unnamed Vermipardus species

from Madagascar represents the first mention of the

genus from this island. Old World Vermileo members

(BPP = 0.96; BP < 70) were resolved into two groups,

each comprising several described and putative species

from western (Fig. 1 blue colour) and eastern (Fig. 1

green colour) parts of the Palaearctic Region.

Uncorrected p distances are summarised in Appen-

dix I. Within the Palaearctic Lampromyia, the largest

intraspecific p distance was found in L. canariensis Mac-

quart, 1838 (0.025), whereas the smallest interspecific

p distance was between L. pallida Macquart, 1835 and

L. lecerfi Séguy, 1930 (0.038) and the largest interspecif-

Bonn zoological Bulletin 70 (2): 339-349

ic p distance was between L. iberica Stuckenberg, 1998

and L. canariensis (0.131). The two Nearctic Vermileo

species have a p distance from Old World Vermileo of

0.141—0.188, with the minimum p distance between them

of 0.129. The western and eastern Old World clades of

Vermileo are separated by 0.100 to 0.148 p distance.

Within the western Vermileo clade, the largest interspe-

cific p distance was found between /. vermileo (Lin-

naeus, 1758) / Vermileo sp. G (0.124), and the smallest

interspecific p distance was between V. immaculatus

Carles-Tolra & Cuesta-Segura, 2020 and Vermileo sp.

G (0.050). Within the eastern Vermileo clade, the largest

interspecific p distance was between Vermileo sp. A and

Vermileo sp. C (0.121), and the smallest interspecific p

distance was found between Vermileo sp. E and Vermileo

sp. F (0.038). The maximum intraspecific p distance for

an Old World Vermileo species does not exceed 0.015,

with the exception of V. ater Stuckenberg, 1965 (0.036).

DISCUSSION

Although phylogenetic reconstruction based on a single

mitochondrial gene fragment is not adequate to establish

conclusions on evolutionary relationships, it still has the

potential to identify aspects in the systematics of a group

that need to be scrutinised more thoroughly.

Lampromyia

Stuckenberg (1998) scored 20 morphological charac-

ters for his cladistic analysis of Lampromyia and iden-

tified three species-groups. The pi/osula-group occurs in

southern Africa and comprises L. flavida (Engel & Cuth-

bertson, 1937), L. pilosula Engel, 1929 and L. rebecca

Stuckenberg, 1996. The canariensis-group 1s present on

the Canary Islands and has three members, L. canarien-

sis, L. fortunata Stuckenberg, 1971 and L. hemmingseni

Stuckenberg, 1971. Finally, the cylindrica-group occurs

in the West Mediterranean Basin and is further divided

into two subgroups, namely the cylindrica-subgroup

(with L. cylindrica (Fabricius, 1749), L. funebris Dufour,

1850, L. lecerfi and L. nigripennis Séguy, 1930) and the

pallida-subgroup (with L. iberica and L. pallida). Kehl-

maier (2014) described an additional species from the

pallida-subgroup, L. bellasiciliae Kehlmaier, 2014. The

current molecular analyses supported the Palaearctic spe-

cies groups and subgroups (Fig. 1), with the exception

that L. hemmingseni was resolved with weak to moderate

support (BPP = 0.94; BP < 70) as the sister taxon of the

pallida-subgroup. The future addition of its proposed sis-

ter taxon L. fortunata to the dataset (the only Palaearctic

Lampromyia currently missing) might provide additional

evidence.

The placement of Lampromyia sp. from Lesotho, a

member of the pi/osula-group, with two Vermipardus

species has no statistical support. A possible reason for

©ZFMK

342

0.99/74

0.96/<70

0.99/76

1/97/0.5%

1/100/0.8% {

1/100/0% |

1199/0.2% |

Christian Kehlmaier

CK478

CK512

CK513

CK527

CK536

CK539

CK547

CK557

CK558

CK576

CK766

CK769

CK934

Vermileo vermileo

Italy, Grosseto

Spain, Cabrils

France, Nantes

Croatia, Mljet

Croatia, Basca Voda

Italy, Roma

Italy, Sardinia

Italy, Pesaro

Italy, Arco

France, La Ciotat

Germany, Ettlingen

Vermileo balearicus CK480 Spain, Menorca

Vermileo nigriventris CK482 Portugal, Viseu

Vermileo nigriventris CK509 Portugal, Avintes

Vermileo nigriventris CK768 Spain, Burgos

Vermileo sp. D CK540 Italy, Sardinia

Vermileo sp. D CK559 Italy, Sardinia

Vermileo cylindraceus CK479 Italy, Naples

Vermileo cylindraceus CK528 Italy, Ischia

Vermileo cylindraceus CK764 Italy, Vivaro

Vermileo cylindraceus CK767 Italy, Capri

100/0% | Vermileo immaculatus CK507 Malta, Mellieha

1/98

Vermileo sp. G CK942 Italy, Sicily

Vermileo sp. G CK935 Italy, Sicily

Vermileo ater CK497 Greece, Crete

Vermileo ater CK930 Greece, Crete

Vermileo ater CK683 Greece, Crete

Vermileo ater JPTOO6 Greece, Crete

Vermileo ater CK758 Greece, Crete

Vermileo immaculatus CK941 Malta, Mellieha

1/100/0%

1/94/3.6%

1/96

175

1/85

Vermileo sp. C CK489 Greece, Macedonia

Vermileo sp. C CK526 North Macedonia, Belitsa

Vermileo sp. C CK760 Albania, Librazhd

Vermileo sp. E CK551 Greece, Rhodes

4197 Vermileo sp. E CK552 Greece, Rhodes

Vermileo sp. F CK549 Greece, Samos

Vermileo sp. F CK770 Greece, Kos

1/99/0.3%

Tal 00M). 3%

1/94

1/96

0.99/<70

0.9/<70

corner 3 : :

Vermileo sp. F CK759 Bulgaria, Blagoevgrad

+HOCO% Vermileo sp. H CK905 Iran, Dodan

Vermileo sp. H CK907 Iran, Sepidap

Vermileo sp. H CK908 Iran, Dagasyave

0.99/97/0.9%

1192

Vermileo sp. A CK556 Israel, Tel Aviv

Vermileo sp. A CK580 Israel, Dan

Vermileo sp. A MN411056 Israel, Fureidis

Vermileo niloticus CK546 Egypt, Wadi Degla

Vermileo sp. B CK487 Cyprus, Dora

Vermileo sp. B CK932 Cyprus, Paphos

Lampromyia cylindrica LM993305 Tunisia, Bizerte

Lampromyia funebris LM993300 Spain, Burgos

Lampromyia funebris CK943 Spain, Lleida

Lampromyia lecerfi CK763 Morocco, Tiznit

Lampromyia nigripennis LM993304 Morocco, Azrou

Lampromyia pallida LM993301 Tunisia, Carthago

1/99/0.2% | Lamopromyia iberica LM993299 Spain, Lleida

Lampromyia iberica CK563 Spain, Lleida

Lampromyia bellasiciliae LM993298 Italy, Sicily

Lampromyia hemmingseni LM993303 Spain, Lanzarote

Lampromyia canariensis CK913 Spain, La Gomera

Lampromyia canariensis LM993302 Spain, Tenerife

Lampromyia sp. MN411011 Lesotho, Bokong

1/99/0.6%

0.91/<70

0.94/<70

1/100/2.5%

Vermipardus sp. CK553 South Africa, Port Elizabeth

Vermipardus sp. CK936 Madagascar, Ambalavao

1/100/0%

1/100/0.2% {

Chrysopilus sp. AY165706

Coenomyia ferruginea MG968060

0.09

Symphoromyia sp. AY165652

Rhagio mystaceus GU013655

Leptynoma hessei CK484 South Africa, Robertson

Vermileo comstocki CK613 USA, Nevada

Vermileo comstocki CK616 USA, Nevada

Vermileo opacus CK491 USA, California

Vermileo opacus CK492 USA, California

Fig. 1. Phylogenetic tree of a Bayesian analysis of CO/ barcoding data of Vermileonidae. Depicted are statistical support values for

ingroup nodes of posterior probabilities (BPP) > 0.9 and bootstrap values (BP) > 70. For lineages representing known species or

putative species, the maximum intraspecific genetic p distance is given in %. DNA voucher numbers follow the species name and

more information is provided in Table 1. For those sequences retrieved from GenBank, the GenBank accession number is given

instead of the DNA voucher number. The blue colour indicates the western Palaearctic Vermileo, the green colour indicates the

eastern Palaearctic/Afrotropic Vermileo, and the red colour highlights the Nearctic Vermileo.

this unexpected placement is the length of the sequence

MN411011 (with only 242 base-pairs). A first-hand

morphological re-examination of the heavily damaged

voucher (head missing, terminalia detached and stored

Bonn zoological Bulletin 70 (2): 339-349

separately (Torsten Dikow in /itt.), see habitus photo at

http://v3.boldsystems.org/index.php/Public_Record-

View? processid=ASILO359-17) is necessary, but cur-

rently not possible. The specimen was last identified by

©ZFMK

DNA barcoding reveals an unexpected diversity in Old World Vermileonidae 343

Table 1. Material and sequence accession numbers.

Field DNA | Sequence accession

Taxon Locali Sex

Me code voucher number

Old World Vermileo

GREECE, Crete, Rethymno, Municipal Gar-

Vermileo ater den, 35.365628° N, 24.472422° E, leg. M. Auer, larva | Ret20 | CK930 0U230768

6.vil.2017.

GREECE, Crete, Maroulas, 4 km SE Perivolia, leg.

V. Michelsen, 14.v1.1999, coll. ZMUC.

GREECE, Crete, Iraklio, at window inside Natural

History Museum of Crete, leg. C. Schmidt, v.2001.

GREECE, Crete, Chania, Thymia, 35.4106° N 3

24.0440°E, leg. & coll. J. Pohjoismaki, 9.vi.2019.

GREECE, Crete, Iraklio, Krasi, 35.234241° N

Vermileo ater 25.467538° E, vii.2013, leg. C. Tummler, v1.2013, Q — CK758 0U230771

ex larva 5.111.2015.

Vermileo ater 3 = CK497 0U230769

Vermileo ater CK683 O0U230770

Vermileo ater JPTO0O6 | JPTOO6 | BOLD:ADZ6857

SPAIN, Balear Islands, Menorca, Ferreries, Barranc

de Cala Santa Galdana, leg. M.J. Ebejer, 7. VI.2008.

ITALY, Ischia, 40°43’56” N, 13°55’°15” E, leg. C.

Schmidt, 19.V.2011.

ITALY, Capri, Anacapri-Faro, 40°32’3” N,

Vermileo cylindraceus | 14°12’3” E, leg. M. Mende, 29.1x.2012, ex larva 3 Cap1 CK767 0U230774

12.vi.2013.

ITALY, Isola di Vivara, 40.743843° N,

Vermileo cylindraceus | 13.993472° E, leg. C. Schmidt, ca. 20.1v.2014, ex 3 Viv2 CK764 0U230775

larva 10.vi.2014

ITALY, Napoli, Posillipo, 40°48’24”’ N,

Vermileo cylindraceus | 14°11°31” E, leg. C. Schmidt, 18. 1v.2009, ex larva 3 Nap9 | CK479 0U230776

10.vi.2009.

MALTA, Mellieha, Santa Marija, ca. 35°57’20”N

Vermileo immaculatus | 14°22’02” E, leg. P. Gatt, 24.v11.2010, ex larva Q MTI1 CK507 0U230777

8.iv.2011.

MALTA, Mellieha, Santa Marija, ca. 35°57’20” N,

Vermileo immaculatus | 14°22’02”E, leg. P. Gatt, 24.vi1.2010, ex larva 3 MT2 CK941 0U230778

7.1v.2011.

PORTUGAL, Viseu, Silgueiros, Povoa Dao,

Vermileo nigriventris | 40°33’ N, 7°57’ W, leg. J. Almeida, 2.vi1i.2009, ex 3 Por4 CK482 0U230779

larva 8.vi.2010.

PORTUGAL, Avintes, 41°6’ N, 8°33’ W, leg. R.

Andrade, 13.11.2009.

SPAIN, Burgos, Barbadillo del Mercado, leg. U.R. 9

Liders, 23.vii1.2012, ex larva 22.v.2013.

EGYPT, Cairo, Wadi Degla, leg. leg. H. Badrawy, 3

6.x1.2010, ex larva 14.iv.2011.

CROATIA, Dubrovnik-Neretva, Mljet, Babino

Vermileo vermileo polje (Sutmiholjska beach), leg. J. Podlesnik, larva | Bab13 | CK527 0U230783

15.vui.2011.

CROATIA, Dalmatia, Baska Voda, leg. D. Devetak, 3

vill.2006, ex larva 10.v1.2007.

FRANCE, Nantes, Faculté des Sciences et

des Techniques de Nantes, 47°14’19,62” N,

1°33°17,60” E, leg. A. Lequet, 29.iv.2010, ex larva $

12.vi.2010.

Vermileo balearicus CK480 O0U230772

Vermileo cylindraceus larva | Iscl CK528 0U230773

Vermileo nigriventris larva | PT6 CK509 0U230780

Vermileo nigriventris Urs1 CK768 0U230781

Vermileo niloticus — CK546 O0U230782

Vermileo vermileo — CK536 OU230784

Vermileo vermileo F10 CK513 OU230785

344

Christian Kehlmaier

Taxon

Old World Vermileo

Vermileo vermileo

Vermileo sp. A

Vermileo sp. B

Vermileo sp. C

Vermileo sp. D

Locality

FRANCE, Bouches-du-Rhéne, La Ciotat, leg. H.

Dumas, 4.viii.2011, ex larva 27.v.2012.

GERMANY, Baden-Wirttemberg, Ettlingen,

48°57°24.8” N, 8°23’03.1” E (48.956962,

8.384270), leg. K. Grabow, v.2018, ex larva

31.v.2019.

ITALY, Pesaro & Urbino, Pesaro, Collina Baratoff,

leg. M. Paglialunga, 27.vi1.2011.

ITALY, Trentino-Alto Adige, Arco, Botanical Gar-

den, x.2013, leg. M. Auer, ex larva 29.1v.2014

ITALY, Toscana, Grosseto, Portiglione, Cala Violi-

na, 10°46’30” E, 42°51’°26” N, leg. C. Kehlmaier,

19.iv.2008, ex larva 21.v.2008.

ITALY, Roma, Lazio, Monte Mario, leg. A. Alfonsi

via D. Badano vii.2010, ex larva 14.11.2011.

ITALY, Sardinia, Sassari, Alghero, 40°32717.84” N,

8°19°31.81” E, leg. D. Badano, 28.vili.2010, ex

larva 23.1.2011.

ITALY, Sardinia, Sassari, Parco del Monserrato,

40°42’34.59” N, 8°32’54.50” E, leg. D. Badano,

15.x.2010, ex larva 4.1v.2011.

ITALY, Sardinia, Sassari, Platamona, leg. D. Bada-

no, 28.x1.2010, ex larva 21.11.2011.

SPAIN, Cabrils, leg. M. Carles-Tolra, 3.v.2009.

ISRAEL, Tel-Aviv, Ramat-Gan, 32°05’00.3” N,

34°48’38.8” E (32.083415 34.810771), leg. A. Nir,

26.v1.2010, ex larva 20.11.2011.

ISRAEL, Dan, Bet Ussishkin, leg. L. Bodner via A.

Freidberg, 22.x1.2011.

ISRAEL, Nahal Tut, 5 km East of Fureidis, leg. W.

Mathis &. A. Freidberg, 19.v.1980.

CYPRUS, 6 km N Dora, 34°48‘07* N,

32°45‘29* E, leg. & coll. M.J. Ebejer, 27.iv.2002,

ex larva 18.V.2002.

CYPRUS, Paphos, Apostolou Pavlou Avenue, leg.

K. Grabow, 6.iv.2018, ex larva 30.iv.2018.

ALBANIA, Librazhd, Pérrenjas, 41.06687° N,

20.54218° E, leg. J. Dils (K15007), 19.vii.2015.

GREECE, Macedonia, Lake Kerkini, Sidirochori,

41°14.33’ N, 23°01.25’ E, leg. & coll. M.J. Ebejer,

7.vi.2010, ex larva 22.vi1.2010.

NORTH MACEDONIA, near Belitsa, 41° 40.69’ N,

21° 12.54’ E, leg. D. Devetak & V. Klokoéovnik,

16.ix.2011.

ITALY, Sardinia, Sassari, Alghero, Spiaggia del

Lazzaretto, leg. D. Badano, 7.x1.2010, ex larva

20.11.2011.

ITALY, Sardinia, Sassari, Alghero, 40°32717.84” N,

8°19°31.81” E, leg. D. Badano, 28.vili.2010, ex

larva 27.1.2011.

Bonn zoological Bulletin 70 (2): 339-349

Sex

larva

Field

code

C105

LAZ1

Algl

AL12

Bell8

Lazzal

Alg2

DNA

voucher

CK934

CK576

CK766

CK478

CK539

CK557

CK526

CK559

CK540

Sequence accession

number

OU230786

0U230787

0U230788

O0U230789

0U230790

0U230791

0U230792

0U230793

0U230794

0U230795

0U230796

0U230797

MN411056

(as Vermileo ver-

mileo)

0U230798

0U230799

O0U230800

0U230801

O0U230802

0U230803

0U230804

©ZFMK

DNA barcoding reveals an unexpected diversity in Old World Vermileonidae

345

Taxon

Old World Vermileo

Vermileo sp. E

Vermileo sp. F

Vermileo sp. G

Vermileo sp. H

New World Vermileo

Vermileo comstocki

Vermileo opacus

Lampromyia

Lampromyia bellasi-

ciliae

Lampromyia canar-

1ensis

Lampromyia canar-

1ensis

Locality

GREECE, Rhodes Island, Rhodes city, leg. R.A.

Pantaleoni, 9.xi.2010, ex larva 17.11.2011.

GREECE, Rhodes Island, Rhodes city, leg. R.A.

Pantaleoni, 9.xi.2010, ex larva 19.111.2011.

BULGARIA, Blagoevgrad, Campus of the

American University in Bulgaria, 42°00’47” N,

~ | 23°05°45” E, leg. C. Schmidt, 3. viii.2013, ex larva

28.v.2014.

GREECE, Kos Island, SW Kefalos, leg. M. Auer,

vi.2015, ex larva 16.v1.2015.

GREECE, Samos Island, Potamia, 37°47.730 N,

26°40.404 E, leg. G. Stahls, x1.2010, ex larva

3.iv.2011.

ITALY, Sicily, Gorghi Tondi (TR), 37°36’34.82” N,

12°39°24.54” E (37.609672° N, 12.656817° E),

leg. R.A. Pantaleoni via D. Badano, 10.[X.2010, ex

larva 16.iv.2011.

ITALY, Sicily, Trapani, Mazara del Vallo, Ris-

erva Naturale Lago Preola e Gorghi Tondi,

37.611795° N, 12.654338° E, leg. A. Miller,

8.111.2016, ex larva 25.v.2016.

IRAN, Kermanshah, Dodan, 34°00’ N, 46°12’ E,

Malaise trap, leg. M. Zardouei, 5.vii.2016.

IRAN, Kermanshah, Sepidap, 34°59’ N, 46°14’ E,

Malaise trap, leg. M. Zardouei, 20.v1.2016.

IRAN, Kermanshah, Dagasyave, 35°02’ N,

46°12’ E, Malaise trap, leg. M. Zardouei,

20.vi.2016.

USA, Nevada, Zephyr Cove, Camping resort,

39°00’22” N, 119°56’42” W, 1900 m, leg. M.

Hauser, 20.vitl.2011, ex larva 27.v.2012.

USA, Nevada, Zephyr Cove, Camping resort,

39°00’22” N, 119°56’42” W, 1900 m, leg. M.

Hauser, 20.vitl.2011, ex larva 27.v.2012.

USA, California, Los Angeles County, La Crescen-

ta, Eagle Canyon, 34°14’45” N, 118°13°56” W,

2500ft, leg. S.D. Gaimari, M.S. & E.S. Anderson,

10.—17.VI.2007.

USA, California, Los Angeles County, La Crescen-

ta, Eagle Canyon, 34°14’45” N, 118°1356” W,

2500ft, leg. S.D. Gaimari, M.S. & E.S. Anderson,

10.—17.VI.2007.

ITALY, Sicily, Palermo, 38.148594° N,

13.364795° E, leg. F. Sacco, 19.x1.2012, ex larva

7.vui.2013.

SPAIN, Canary Islands, Tenerife, Fasnia, 28.24° N,

16.44° W, leg. G. Pefia Tejera, 26.vi.2012.

SPAIN, Canary Islands, La Gomera, Hermigua,

El Cerrillal, leg. J. Jakobitz, 30.x1.2016, ex larva

vi.2017.

Bonn zoological Bulletin 70 (2): 339-349

Sex

Field

code

Rho2

Rhol

Bul5

KOS7

SAM1

Sicl

Sicll

US7

US3

US-M

US-W

$1z19

DNA

voucher

CK551

CK552

CK759

CK770

CK549

CK942

CK935

CK905

CK907

CK908

CK613

CK616

CK491

CK492

CK699

CK635

CK913

Sequence accession

number

O0U230805

O0U230806

O0U230807

0U230808

O0U230809

0U230810

0U230811

0U230812

0U2308 13

0U230814

0U230815

0U230816

0U230817

0U230818

LM993298 (Kehl-

maier 2014)

LM993302 (Kehl-

maier 2014)

0U230819

©ZFMK

346 Christian Kehlmaier

Tascon Locality ai Field DNA | Sequence accession

code | voucher number

Lampromyia |]

Lampromyia cylin- TUNISIA, 6 km N Bizerte, Cap Blanc, 37.33° N, 9 _ CK494 LM993305 (Kehl-

drica 9.83° E, leg. M. Hauser, 18.vi.2000. maier 2014)

. | SPAIN, Burgos, Ura, 42.014348° N, 3.552813° W, LM993300 (Kehl-

Lampromyia fumeDriss 15 ip tryders O3yiil 20lIker larva LI DOTS: | re | MP | ER OOS I ce 014)

SPAIN, Lleida, Bell-Iloc d‘Urgell, 41°37°43” N,

Lampromyia funebris | 0°46’59” E, leg. R.M. Batlle, viti.2012, ex larva Q Ram3 | CK943 0U230820

10.vi.2013.

SPAIN, Canary Islands, Lanzarote, Playa de Papa-

Lampromyia hem- | av. 28 84°'N, 13.78° W. leg. C. Schmid-Beger | @& | —= | @Kag3 2) /MO22303(Kehh

mingseni cs maier 2014)

5 111.2008.

Lie Se SPAIN, Lleida, Timoneda d’Alfés, 10 km SE Llei- LM993299 (Kehl-

See. 7 SPAIN, Lleida, Timoneda d’Alfés, 10 km SE Llei-

Lampromyiaiberica | 4, ten RM. Batlle, 4.ix.2010, exlarva7.iv2011. | ¢ | RBI | CK563 UE eat

MOROCCO, Tiznit, Tiznit, 29.69582° N,

Lampromyia lecerfi 9.72542° W, leg. J. Dils (K15002), iv.2015, ex 3 MAI10 | CK763 O0U230822

larva 11.v1.2015.

Lampromyia ni- MOROCCO, South of Azrou, 33.32° N, 5.27° W, ioe _ CK504 LM993304 (Kehl-

gripennis leg. A.K. Hundsdorfer, vi.2006. maier 2014)

, TUNISIA, Carthago, 36.86° N, 10.33° E, leg. D. LM993301 (Kehl-

Lampromytapallida: | 76 eek 29.y-2010.ex larva 272010: 3 | TUNI | CK481 maier 2014)

Lampromyia sp. LESOTHO, Bokong, leg. L. Bevis, 26.xii.1946, Bi il|| = ad ae ane

Leptynoma

SOUTH AFRICA, Western Cape, Riverhuis Farm,

Leptynoma hessei 14 km NW Robertson, 33°46’40” S, 19°45°35” E, 3 — CK484 0U230823

leg. M. Hauser, 29.1x.2004.

Vermipardus

MADAGASCAR, Anja Reserve, W Ambalavao,

VOIR GT AHO SE eset Rasolondalagi xi 201 .cmlarva 29 1019 | t= || aall: | CRe36 OUesea8

SOUTH AFRICA, Eastern Cape Province, 70 km

Vermipardus sp. N Port Elizabeth, near Addo Elephant National 3 Addo2 | CK553 O0U230825

Park, leg. U. Fritz, 22.x1.2010, ex larva 13.11.2011.

Outgroup

Chrysophilus sp. No data available. ? — — AY 165706

CANADA, Ontario, Pukaskwa, Pukaskwa National

Rhagio mystaceus Park, Coastal Trail Boreal forest, 48.56°N 86.23°W, ? — — GU013655

leg. J. Cossey et al., 2.vii.2008.

Symphoromyia sp. No data available. vi — — AY 165652

USA, Virginia, Berkeley County, Sleepy Creek

peed ba fener: Wildlife Area, 39.52° N, 78.1503° W, leg. N.E. ? = = MG968060

& Woodley, 22.v.2004.

S.W. Bromley before 1955, and Vermipardus was erected

as a subgenus within Lampromyia only some years later

by Stuckenberg (1960).

Vermileo

As mentioned earlier, the present study indicates the

existence of eight putative new species of the genus

Bonn zoological Bulletin 70 (2): 339-349

Vermileo in the Palaearctic Region. Prior to this study,

the genus comprised twelve named species worldwide

plus an apparently undescribed species from Costa Rica

(Woodley 2009). The known Palaearctic Vermileo spe-

cies are: V. ater, V. balearicus Wheeler, 1930, V. immacu-

latus, V. nigriventris (Strobl, 1906) and V. vermileo. In the

Afrotropical Region there is only one species, V. nilot-

©ZFMK

DNA barcoding reveals an unexpected diversity in Old World Vermileonidae 347

icus. Six species are known from the Nearctic: K com-

stocki Wheeler, 1918, V. dowi Wheeler, 1931, V. fasci-

pennis (Williston, 1895), V. opacus (Coquillett, 1904),

V. tibialis (Walker, 1852), and V. willetti DeLeon, 1938.

Based on the current dataset, the species V. cylindraceus

(Costa, 1844) stat. rev., formerly considered synonymous

to V. vermileo (see Bezzi 1900), is reinstated as a valid

species.

Wheeler (1918, 1930, 1931, 1934), Edwards (1935),

Stuckenberg (1965), Nagatomi et al. (1999), and

Carles-Tolra & Cuesta-Segura (2020) where the most

recent authors commenting on Vermileo taxonomy; the

latter describing a Vermileo species from Malta and re-

instating V. balearicus as a valid species — the proposed

synonymy of V. balearicus and V. immaculatus with

V. vermileo by Ebejer & Gatt (2021) is not corroborated

by the molecular data presented in the present study. Re-

liable identification keys for the Old World fauna of this

genus do not exist, neither has there ever existed one.

In his Palaearctic revision, Lindner (1925) only included

V. vermileo into his identification key to the Rhagioni-

dae (wormlions had subfamily status within Rhagionidae

at that time) and left V balearicus and V. nigriventris as

subspecies of vermileo. Nagatomi et al. (1999) pro-

vided the only other identification key of Old World Ver-

mileo, which includes V. ater, V. nigriventris, V. niloticus,

and V. vermileo, using differences of the male postab-

domen mentioned by Stuckenberg (1965). Unfortunate-

ly, the first couplet of Nagatomi et al. (1999) is already

misleading, as the numbers of the couplets to follow on

are interchanged. The lack of a proper identification key

has repeatedly caused authors to identify their material

based on general body coloration. That way, V. vermileo

has been cited from regions where the genus is represent-

ed by different, partly undescribed taxa, e.g., Bulgaria

(Popov 1968), Egypt (Hafez et al. 1956a, 1956b), Greece

(Papp & Soltész 2019), Malta (Ebejer 1995). Morpho-

logical descriptions of the putative new species as well

as an identification key will be published elsewhere in

the future.

Both analyses resolved the Nearctic species V. com-

stocki and V. opacus as the sister group of the remaining

studied taxa of the family. Although there is no statistical

support, present results agree with the observations made

by Nagatomi et al. (1999), who pointed out that the struc-

ture of male genitalia of the Palaearctic Vermileo species

clearly differs from the male genitalia of the Nearctic

V. comstocki. Earlier, Pechuman (1938) commented on

two morphological characters that possibly help to sepa-

rate New World from Old World species on a generic lev-

el. The New World Vermileo were last studied by Pechu-

man (1938) and DeLeon (1938), and there currently are

six described species from Cuba, Jamaica, Mexico, and

USA. As the Palaearctic V. vermileo represents the type

species of Vermileo Macquart, 1834, and the first species

from the New World (V. tibialis) was originally described

Bonn zoological Bulletin 70 (2): 339-349

in the monotypic genus Pheneus Walker, 1852, this name

would be resurrected from synonymy in case that the

phylogenetic placement for the Nearctic Vermileo re-

ceives further evidence and stronger support.

As highlighted in the present study by the number of

putative new species, being apparently undescribed, the

species richness of the Vermileo in the Old World has

been greatly underestimated, especially in the Medi-

terranean Basin. The group with western Vermileo taxa

comprises seven species (V. balearicus, V. cylindraceus,

V. immaculatus, V. nigriventris, V. vermileo, Vermileo sp.

D and Vermileo sp. G) and their geographical distribu-

tion ranges from Portugal to southern Croatia, whereas

the group with eastern Vermileo representatives consists

of eight species (V. ater, V. niloticus, Vermileo sp. A, Ver-

mileo sp. B, Vermileo sp. C, Vermileo sp. E, Vermileo sp.

F and Vermileo sp. H) and they occur from Albania to

western Iran. Preliminary morphological studies of male

terminalia confirm this partition. Old World species of

Vermileo look alike superficially due to the absence of

striking coloration and the lack of conspicuous apomor-

phic features (e.g., on the head, legs or sternites). At the

same time, general body coloration and wing venation

can vary considerably within a single species. Therefore,

stable morphological characters are scarce, especially in

females. Promising features being the coloration of the

small lateral thoracic sclerites, and the shape of male

tergite 9 which displays species specific lateroapical ap-

pendages in the eastern Palaearctic/Afrotropic Vermileo

species group.

The current knowledge does not reflect the true distri-

bution range of the individual species or groups as in-

tensive sampling in the western Palaearctic has not been

done, with important geographical gaps in Corsica and

Turkey. However, when looking at the distribution of the

individual known or putative species it is striking that

seven out of 15 (47%) Palaearctic species of Vermileo

are limited to islands. Keeping in mind the old age of the

family and the unsteady geological history of the Medi-

terranean Basin (e.g., Mediterranean salinity crisis), one

tends to ask whether the observed island taxa indeed con-

stitute true endemism, 1.e., species that evolved on the

island due to reproductive isolation, or might be relicts of

formerly much wider distributed species.

Acknowledgements. I am very grateful to the many people

supporting this study over the past ten years by collecting and

sending Vermileonidae larvae and adults or granting access to

the institutional collections in their care (in alphabetical order):

Jorge Almeida, Rui Andrade, Markus Auer, Davide Badano,

Haitham Badrawy, Ramon M. Battle, Miguel Carles-Tolra,

Du8San Devetak, Jozef Dils, Hélene Dumas, Martin J. Ebejer,

Magdi El-Hawagry, Amnon Freidberg, Uwe Fritz, Paul Gatt,

Karsten Grabow, Martin Hauser, Jens Jakobitz, André Lequet,

Urs R. Ltiders, Nicola Maio (Centro Musei delle Scienze Natu-

rale e Fisiche, Napoli, Italy), Michael Mende, Benham Mota-

medinia, Anke Miller, Assaf Nir, Marco Paglialunga, Rober-

©ZFMK

348 Christian Kehlmaier

to A. Pantaleoni, Thomas Pape (Natural History Museum of

Denmark, Copenhagen), Jan Podlesnik, Jaakko Pohjoismaki,

Gianni Raffone, Harin’Hala (Rin’ha) Rasolondalao, Christian

Schmidt, Gunilla Stahls, Claudia Tummler, and Nigel Wyatt

(Natural History Museum, London, United Kingdom).

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[last accessed 10 Aug. 2021]

Bezzi M (1900) Zur Synonymie und Verbreitung des Psam-

morycter vermileo Deg. Wiener Entomologische Zeitung

19 (2/3): 56-57

Carles-Tolra M & Cuesta-Segura AD (2020) Vermileo immacu-

latus sp. n.: anew vermileonid species from Malta (Diptera:

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APPENDIX I

(electronic supplement, available at www. bonnzoologicalbulletin.de)

Uncorrected pairwise distance matrix. Blue: western Palaearc-

tic species of Vermileo. Green: eastern Palaearctic/Afrotropic

species of Vermileo. Red: Nearctic species of Vermileo.

©ZFMK

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2021 - Gebicki C. et al.

https://do1.org/10.20363/BZB-2021.70.2.351

ISSN 2190-7307

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Research article

urn:|sid:zoobank.org:pub:C2125016-AEF6-4F7E-83D9-5718C378B405

Youngeewa simillima sp. nov. (Hemiptera: Cicadellidae: Mileewinae)

from Eocene Baltic Amber

Cezary Gebicki', Marcin Walcezak”" & Marzena Zmarzly*

‘Faculty of Science & Technology, Jan Diugosz University in Czestochowa, Al. Armii Krajowej 13/15,

PL-42-200 Czestochowa, Poland

?3Zoology Research Team, Institute of Biology, Biotechnology and Environmental Protection, Faculty of Natural Sciences,

University of Silesia in Katowice, Bankowa 9, PL-40-007 Katowice, Poland

“Corresponding author: Email: marcin.walczak@us.edu.pl

'urn:lsid:zoobank.org:author:}6FC4935A-D82E-45BE-BD77-8E9EFEB9BFDD

> urn:Isid:zoobank.org:author:046FOD52-7FBA-4D7D-9F79-A EBF7F2C98B8

3urn:Isid:zoobank.org:author:E11DDOB8-412A-41 EE-A8CC-4114A2CDI1F54

Abstract. The paper describes Youngeewa simillima sp. nov. from Eocene Baltic amber as a new species within the tribe

Mileewini Evans, 1947 (Mileewinae Evans, 1947). Due to its poor condition, the specimen is only tentatively placed in the

genus Youngeewa Gebicki & Szwedo, 2001, differing from the other known species of the genus, Youngeewa bicolorata

Gebicki & Szwedo, 2001, mainly by the elements of wing venation.

Key words. Youngeewa, Cicadellidae, Mileewinae, new species, Baltic amber.

INTRODUCTION

A leafhopper specimen in Baltic amber is described as a

new species in the present work. The described species 1s

the third known fossil representative of the Mileewinae

Evans, 1947 subfamily so far. Previously, Youngeewa bi-

colorata Gebicki & Szwedo, 2001 and Eomileewa eri-

dani Gebicki & Szwedo, 2001 (Mileewinae: Mileewini)

have been described from Eocene Baltic amber (Gebicki

& Szwedo 2001). Specimens of these two species were

better preserved than the new species described here and

had well-preserved features for the subfamily. The speci-

men described here has details of its head obscured and is

placed in the subfamily based mainly on its short clavus,

an important diagnostic feature of the tribe Mileewini Ev-

ans, 1947, and it is tentatively placed in the genus Young-

eewa Gebicki & Szwedo, 2001. Three other features char-

acteristic of the extant Mileewini of the Old world have

not been confirmed, i.e., vein RP (=Sc+R or R,, ,) extend-

ing to the anterior rather than outer margin of the wing

and the marginal vein reaching the wing margin before

A, and fold between PCu and A,. Two other differences

concerning the width and shape of the forewing apex are

variable in Mileewini (see Dietrich et al. 2017; He et al.

2021). Mileewinae Evans, 1947 are an evolutionary old

auchenorrhynchan group originating from the Early Cre-

taceous central Gondwanaland (Nielson & Knight 2000).

The first representatives of the subfamily were estimated

to have originated around 126 MA, based on molecular

Received: 25.01.2021

Accepted: 27.08.2021

analysis and incorporating Youngeewa Gebicki & Szwe-

do, 2001 as a calibration point (Dietrich et al. 2017).

Mileewini seem to be closely related and probably de-

rived from the subfamily Cicadellinae Latreille, 1825 and

are also similar in wing venation to the subfamily Typhlo-

cybinae Kirschbaum, 1868 (Balme 2007; Dietrich et al.

2010), especially to the extinct Protodikraneurini Gebicki

& Szwedo, 2006 (Gebicki & Szwedo 2006) and Alebrini

McAtee, 1926 but representatives of these taxa have an

acute hind basitarsomere unique to all Typhlocybinae.

MATERIAL AND METHODS

A male was examined. Only external structures were ex-

amined using a stereoscopic microscope Olympus SZX9

Nikon Eclipse. Photographs were taken with the Canon

Eos camera with extension rings. All the equipment is

available at the University of Silesia, Faculty of Natural

Sciences, Institute of Biology, Biotechnology and Envi-

ronmental Protection. All photographs and drawings were

digitalised and worked on with CorelDRAW software.

Research funding was provided by the Dean of the Fac-

ulty of Natural Sciences, Institute of Biology, Biotechnol-

ogy and Environmental Protection, University of Silesia

in Katowice.

The holotype Y. simillima Gebicki & Walczak sp. nov.

bears labels; quoting the labels of specimen is done in the

following way: (/) 1s used to divide data in different rows

Corresponding editor: M. Espeland

Published: 29.10.2021

352 Cezary Gebicki et al.

on the label, (;) is used to divide data on different labels, RA = Radius anterior vein (RA1)

([]) is used for the author’s comments. RP = Radius posterior vein

Abbreviations for morphological terms

RESULTS

A =Anal vein (A1)

CuA = Cubitus anterior vein (CuA1 / CuA2) Youngeewa simillima Gebicki & Walczak sp. nov.

CuP = Cubitus posterior vein (Figs 1-4)

M =Media vein urn: lsid:zoobank.org:act:0C8A599B-570B-4DE4-99E2-183BE988E0F3

MA = Media anterior vein

MP =Media poserior vein Type material. Holotype 3 [written in all caps], Young-

PCu = Postcubitus vein eewa simillima / Gebicki & Walczak / sp. nov. [red label,

Sc =Subcosta vein written in bold]; Eocen Baltic amber / No. 5008 / coll.

R — =Radius vein (R / R2) Carsten Grohn.

Fig. 1. Youngeewa simillima Gebicki & Walczak sp. nov. A. Head and pronotum from the dorsal side. B. Face and right leg of the

first pair, right leg of the second pair, and foot of third pair of legs. C. Face and fore legs. D. Specimen from the ventral side.

Youngeewa simillima sp. nov. from Eocene Baltic Amber 353

Diagnosis. In general outline the species is close to

Y. bicolorata but differs in the following characteristics:

straight lateral margins of genae (no deep incision below

the eye), apex of rostrum reaching beyond bases of middle

coxae, veins SctRA, and transverse r-m and m-cu on fore

wings are arranged in one line (as in Fig. 3C), and the dis-

tance between distal terminations of veins CuA, and CuP

on hind wings is only 2.1 x as long as that between MP+C-

uA and CuA, (measured on hind wing marginal vein), (in

Y. bicolorata 2.3 x). Hind wing cell delimited by veins R

and M is somewhat broader starting from transverse vein

r-m to the wing apex than the cell delimited by veins MA

and MP+CuA.. Fore wings distinctly lighter than those

in Y. bicolorata and lacking any clear colour pattern.

Description. Total body length 3.70 mm (sex: male), de-

tailed data showing the dimensions of selected parts of

the body are included in Table 1.

Head. Coronal part of head strongly convex especially

at apex, broad, “delta’’-like in outline. Ocelli, along with

entire surface of head, completely invisible, covered by

a felt-like sheath (Fig. 1A). Coronal and frontal sutures

invisible. Antennae slightly longer than head, located

in distinct antennal pits in front of the eyes at inner eye

margin (Figs 1B—C, 3A). Scapus short and cylindrical,

pedicellus relatively narrow, strongly elongated, many

times longer than first segment. Antennal flagellum about

3x narrower than pedicellus. Lateral margins of facial

part of head almost straight, lacking a clear-cut incision

under eyes. Frontoclypeus convex, distinctly broader in

apical part, strongly tapered distally. In subapical part

of frontoclypeus there is an arcuate concavity located

in place of epistomal suture (Figs 1B, 3A). Anteclypeus

almost square in shape, small, about 5 < narrower than

postclypeus at its widest point and separated from it by a

deep transclypeal suture. Indistinct, 3-segmented rostrum

reaching beyond bases of middle coxae (Fig. 1D).

Pronotum. Shorter than vertex with indistinct lateral

carina (Figs 1A, 2A, 3B).

Wings. Fore wings relatively narrow and elongate, ev-

idently reaching beyond the abdomen, at apex nearly as

wide as corium and slightly rounded. Transverse veins

of three apical cells (2-4) arranged in a single some-

what slanting line. Inner apical field longest, and larg-

est, delimited by CuA,, which is slightly “S”-like; apical

fields 1 and 3 narrow, rectangular in outline; outer field

q , _ .

(o:5smnm' at)

Fig. 2. Youngeewa simillima Gebicki & Walczak sp. nov. A. Specimen from the dorsal side. B. Hind wing (ventral view). C. Left

mid leg. D. Right mid and hind leg. E. Hind leg tarsus in large magnification.

Bonn zoological Bulletin 70 (2): 351-357

©ZFMK

354 Cezary Gebicki et al.

0.5 mm 0.5 mm

+R+

C pis ae ae Sc+RAy

CuA 1

marginal vein

* Pale |

SS ae

0.5 mm

Fig. 3. Youngeewa simillima Gebicki & Walczak sp. nov. (reconstruction). A. Head from the ventral side. B. Head and pronotum

from the dorsal side. C. Fore wing. D. Hind wing.

Youngeewa simillima sp. nov. from Eocene Baltic Amber 355

0.5 mm

Fig. 4. Youngeewa simillima Gebicki & Walczak sp. nov. (reconstruction). A. Right fore leg from the ventral side. B. Right mid

leg from the ventral side. C. Left hind leg from the dorsal side. D. Right hind leg from the ventral side. E. Hind leg tarsus in large

magnification.

356 Cezary Gebicki et al.

Table 1. Biometric data of Youngeewa simillima Gebicki &

Walczak sp. nov. [*data difficult to measure]

Male Size [mm]

head length in midline 0.41 mm

head width probably 0.7 mm*

frons length in midline 0.69 mm

frons width probably 0.73 mm*

antennae length 0.99 mm

pronotum length in midline 0.38 mm

abdomen length in midline 1.74 mm

profemur 0.58 mm

protibia 0.63 mm

protarsus 0.31 mm

mesofemur 0.53 mm

mesotibia 0.78 mm

mesotarsus 0.31 mm

metafemur 1.04 mm

metatibia 1.49 mm

metatarsus 0.82 mm

fore wing lenght 2.95 mm

fore wing width 0.88 mm

hind wing length 2.62 mm

hind wing width 1.08 mm

is smallest, triangular in shape. Distances between main

stems of longitudinal veins R, M and Cu nearly equal.

Clavus reaching halfway along the wing, with two dis-

tinct, almost parallel veins PCu and A, running close to

each other. Corium and costal cell lacking any transverse

veins (Figs 1D, 3C). Fore wings slightly darker than hind

wings, with darker colouration along the fields adjacent

to main longitudinal veins (Fig. 2A).

Hind wings almost as long as fore wings. Veins R and

M parallel in distal part and connected by a transverse

vein r-m, which is located somewhat before nodus. Cos-

tal margin (lacking a distinct, deep incision) similar to

that in species representing the genus Mileewa and to

Y. bicolorata. Marginal vein reaching nodus; in proxim-

ity to the costal margin it runs parallel and joins internal

wing margin outside vein A,. Membranous wing mar-

gin is narrowest between termination of veins MA and

MP+CuA, whereas it is broadest at internal margin op-

posite termination of vein PCu (Figs 2B, 3D).

Legs. Fore legs. Profemur slightly longer than pro-

tibia. Protibia with quite large setae along inner margin.

Segments of protarsus small, third segments longest

(Figs 1B—C, 4A). Middle legs. With sparse fine setae

along inner margin of mesotibia (Figs 2C—D, 4B). Hind

legs. Spines on hind metatibia long, distinctly longer than

metatibia thickness. Apex of metafemur with macrochae-

tae in pattern 2+1+1 (Figs 1D, 2D, 4C—D). Basal segment

of metatarsus longest and at apex expanded into a pro-

Bonn zoological Bulletin 70 (2): 351-357

cess with two spines, outer margin with five sharp spines

and two elongate marcochaetae, which are situated near

the apex on the inner side. The middle segment with three

blunt apical spines and a single sharp lateral one. Hind

pretarsus shortest (Figs 2E, 4E).

Male. Copulatory block relatively short, details not

clearly visible in examined specimen.

Etymology. Species name originates from the Latin

word “simillima” due to the fact that the species present-

ed here is very similar to the Youngeewa bicolorata de-

scribed earlier.

Remarks. Substantial areas of head including rostrum,

parts of prothorax and some parts of legs covered with

white, felt-like substance which blurs the details of the

structure. Accordingly, it is impossible to determine the

location of the ocelli, which constitutes an important tax-

onomic feature. Right fore- and hind wings are well-vis-

ible and their venation is clear. In the male the apex of

abdomen is partly covered by the left hind wing, which

makes analyses difficult. There are no animal syninclu-

sions. For reasoning of the taxonomic placement of the

new species, see introduction and species diagnosis.

DISCUSSION

The species of the genus Youngeewa are close to many

extant Mileewini (Mileewinae Evans, 1947) species in

having a large dome-shaped head, a convex clypeus (also

in Cicadellinae Latreille, 1825), reduction of forewing

venation, a distinct fore wing appendix and shortened

fore wing clavus. These features are also found in the

South American genus Orsalebra Young, 1952 (belong-

ing to the Typhlocybine tribe Alebrini, see Introduction).

However, in this genus the marginal vein of the forewing

does not reach the apex of vein RP, which is typical of

most Mileewi, but reaches only the apex of vein M; in

most Alebrini the marginal vein is fused with the apical

wing margin. Reduced fore wing venation in Mileewinae

is typical of Typlocybinae as well as some Cicadellinae

and Nirvanini Baker, 1923 (Evacanthinae Metcalf, 1939

sensu Dietrich 2004). Hind wing venation is similar in

the above mentioned tribes while veins R and M running

parallel along their entire length (not only in their apical

part) are found in Protodikraneurini Gebicki & Szwedo,

2006 (extinct Typhlocybinae) and in the extant neotrop-

ical mileewine genus Amahuaka Melichar, 1926, as well

as in South American representatives of the tribe Tun-

gurahualini Dietrich, 2011 of the subfamily Mileewinae

(Dietrich 2011; Krishnankutty & Dietrich 2011).

Acknowledgments. We would like to thank Jacek Szwedo

from the University of Gdansk for the loan of the material and

©ZFMK

Youngeewa simillima sp. nov. from Eocene Baltic Amber BOT

Krzysztof Kudia from the University of Silesia in Katowice for

graphic corrections.

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ISSN 2190-7307

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Scientific note

urn:Isid:zoobank.org:pub: EAE5584B-2FB7-4885-9405-17E40029D91F

Two nomenclature changes in Thelastomatoidea

(Nematoda: Chromadorea: Rhabditida)

Filippo Ceccolini' & Fabio Cianferoni*”

'? Zoology, “La Specola”, Natural History Museum, University of Florence, Via Romana 17, I-50125 Florence, Italy

? Research Institute on Terrestrial Ecosystems (IRET), National Research Council of Italy (CNR), Via Madonna del Piano 10,

I-50019 Sesto Fiorentino (Florence), Italy

“Corresponding author: Email: fabio.cianferoni@cnr.it

'urn:|sid:zoobank.org:author:C 1 A7BA9F-4772-4DA7-9492-9D41C11COBB4

?urn:Isid:zoobank.org:author:20A9DD72-7 1 5C-4195-90FB-88CF864BAF54

Abstract. The following replacement names in Nematoda are proposed: Hexasinghiella Ceccolini & Cianferoni nom. nov.

pro Singhiella Rao, 1958 and Coronocephaloides Ceccolini & Cianferoni nom. nov. pro Coronocephalus Cordeira, 1981.

Further, three new combinations (comb. nov.) are made.

Key words. combinatio nova, Hystrignathidae, nomen novum, replacement name, Travassosinematidae.

Thelastomatoidea is a superfamily of Nematoda includ-

ing four families (Bezerra et al. 2021). Two genus-group

names in this superfamily are actually junior homonyms,

since the same names have already been used by differ-

ent authors in Trilobita and Hemiptera. Accordingly, new

names are needed to replace them (ICZN 1999, Art. 60).

REPLACEMENT NAMES

I. Rao (1958: 47) established the genus name Singhiella

to accommodate the new Indian species S. singhi, para-

sitic of the mole cricket Gryllotalpa africana Palisot de

Beauvois, 1805 and included it in the nematode family

Oxyuridae. Presently, the genus is accepted (Spiridon-

ov & Guzeeva 2009; Morffe & Hasegawa 2017) and it is

accommodated in the family Travassosinematidae (IRM-

NG 2021).

Unfortunately, this generic name was already preoccu-

pied, since Sampson (1943: 211) used the same name for

anew genus of Hemiptera Aleyrodidae with 7rialeurodes

bicolor Singh, 1931 as type species. Currently, Singhiella

Sampson 1s valid and includes about 30 species (Jensen

2001; Chen & Ko 2007; Martin & Mound 2007; Push-

pa & Sundararaj 2012). Therefore, Singhiella Rao is a

junior homonym of Singhiella Sampson and, since no

synonym is available, a new substitute name is needed

according to ICZN (1999 Art. 60.1, 60.2). The following

new replacement name is proposed herein: Hexasinghiel-

la Ceccolini & Cianferoni, 2021 nom. nov.

Received: 15.06.2021

Accepted: 31.08.2021

Etymology. The new name, from the Greek word €€ (hex

= six), refers to the number of distinct lips. Feminine gen-

der.

Systematics

Family Travassosinematidae Rao, 1958

Genus Hexasinghiella Ceccolini & Cianferoni nom. nov.

Species Hexasinghiella singhi (Rao, 1958) comb. nov. =

Singhiella singhi Rao, 1958 (type species)

II. The genus group name Coronocephalus appears to

have been erected several times. The first one to use this

name was Grabau (1924: 438) who established the En-

crinuridae trilobite subgenus Coronocephalus with the

type species Encrinurus (Coronocephalus) rex Grabau,

1924. Currently, the name is valid as a genus of Trilobita

(Jell & Adrain 2003; IRMNG 2021).

Subsequently, the name was used by Cordeira (1981)

for a genus of Nematoda Hystrignathidae and by Orlo-

va-Bienkowskaja (1995: 58) for a subgenus of the crus-

tacean Cladocera genus Simocephalus Schoedler, 1858.

The name proposed by Orlova-Bienkowskaja was re-

placed with Crownocephalus by Ozdikmen & Akbulut

(2010), but Coronocephalus Cordeira is still accepted

(Adamson & Van Waerebeke 1992; Nguyen & Smart Jr

2004; Gardufio-Montes de Oca & Oceguera-Figueroa

2020) and it is also a junior homonym of Coronoceph-

alus Grabau. Since no synonym is available, according

to ICZN (1999 Art. 60.1, 60.2), a new replacement name

is necessary. We propose herein the new substitute name

Coronocephaloides Ceccolini & Cianferoni nom. nov.

Corresponding editor: B. Huber

Published: 29.10.2021

360

Etymology. The new name means similar to Corono-

cephalus, the original name proposed by Cordeira (1981).

Masculine gender.

Systematics

Family Hystrignathidae Leidy, 1850

Genus Coronocephaloides Ceccolini

nom. nov.

Species Coronocephaloides ovilamelatus (Cordeira,

1981) comb. nov. = Coronocephalus ovilamelatus

Cordeira, 1981

Coronocephaloides ovipunctatus (Cordeira, 1981)

comb. nov. = Coronocephalus ovipunctatus Cordeira,

1981 (type species)

& Cuianferoni

Acknowledgement. We would like to thank two anonymous

reviewers for their suggestions.

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tomatoidea, Oxyurida of invertebrate hosts. II. Hystrignathi-

dae. Systematic Parasitology 22 (2): 111-130

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Mokievsky V, Pefia Santiago R, Sharma J, Smol N, Tchesun-

ov A, Venekey V, Zhao Z, Vanreusel A (2021) Nemys: World

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cies of whitefly (Hemiptera: Aleyrodidae) from Tatwan with

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cidas. PhD Thesis, Instituto de Ciéncias Biomédicas, Univer-

sidade de Sao Paulo

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lecular phylogeny of Thelastomatoidea (Nematoda) with the

description of a new genus and two new species of Hystri-

gnathidae associated with bess beetles (Coleoptera: Passali-

dae) from Oaxaca, Mexico. Journal of Parasitology 106 (5):

679-688

Grabau AW (1924) Stratigraphy of China. Part 1. Palaeozoic

and Older. Geological Survey, Ministry of Agriculture and

Commerce, Beijing

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(1999) International Code of Zoological Nomenclature. 4"

Edition. The International Trust for Zoological Nomencla-

ture, London

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era (2021) Online at https://www.irmng.org at VLIZ [last ac-

cessed 7 May 2021]

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lobites. Memoirs of the Queensland Museum 48 (2002) (2):

331-353.

Jensen AS (2001) A cladistic analysis of Dialeurodes, Massil-

ieurodes and Singhiella, with notes and keys to the Nearctic

species and description of four new Massilieurodes species

(Hemiptera: Aleyrodidae). Systematic Entomology 26 (3):

279-310

Martin JH, Mound LA (2007) An annotated check list of the

world’s whiteflies (Insecta: Hemiptera: Aleyrodidae). Zoot-

axa 1492 (1): 1-84

Morffe J, Hasegawa K (2017) Morphological and molecular

characterization of Travassosinema claudiae n. sp. (Oxyurid-

omorpha: Travassosinematidae) from the Japanese millipede

Parafontaria laminata (Attems, 1909) (Polydesmida: Xysto-

desmidae). Zootaxa 4282 (1): 166-178

Nguyen KB, Smart Jr GC (2004) Taxonomy of Insect Parasitic

Nematodes. Pp. 795-878 in: Chen ZX, Chen S & Dickson

DW (eds) Nematology: Advances and Perspectives Vol II.

Tsinghua University Press, Beijing

Orlova-Bienkowskaja MJ (1995) Revision of the group Simo-

cephalus (serrulatus) (Crustacea, Daphniiformes, Daphnii-

dae). Zoologicheskii Zhurnal 74 (8): 57—71 [in Russian]

Ozdikmen H., Akbulut N (2010) Crownocephalus nom. nov., a

new name for the preoccupied generic name Simocephalus

(Coronocephalus) Orlova-Bienkowskaja, 1995 (Cladocera:

Daphniidae). Munis Entomology and Zoology 5 (1): 305-306

Pushpa R, Sundararaj R (2012) The genus Singhiella Sampson

from India (Aleyrodidae: Hemiptera) with description of two

new species. Oriental Insects 46 (1): 19-29

Rao PN (1958) Studies on the nematode parasites of insects and

other arthropods. Arquivos do Museu Nacional do Rio de Ja-

neiro 46: 33-84

Sampson WW (1943) A generic synopsis of the hemipterous

superfamily Aleyrodoidea. Entomologica Americana 23 (3):

173-223

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127-134

©ZFMK

Bonn zoological Bulletin 70 (2): 361-371

2021 - Litvinchuk S.N. et al.

https://do1.org/10.20363/BZB-2021.70.2.361

ISSN 2190-7307

http://www.zoologicalbulletin.de

Research article

urn:|sid:zoobank.org:pub: E9F7E470-654D-46CC-944F-93 1 34F826D2E

Morphological differentiation between diploid and polyploid species of

ereen toads (Anura: Bufonidae: Bufotes) in Central Asia

Spartak N. Litvinchuk’", Anton O. Svinin? & Tatjana N. Dujsebayeva*

' Institute of Cytology, Russian Academy of Sciences, Tikhoretsky Pr. 4, 194064, Saint-Petersburg, Russia

? Institute of Environmental and Agricultural Biology (X-BIO), University of Tyumen, Lenina str. 25, 265003, Tyumen, Russia

3 Institute of Zoology of the Republic of Kazakhstan, al-Farabi Av., 93, Almaty 050060, Kazakhstan

“Corresponding author: Email: litvinchukspartak@yandex.ru

'urn:Isid:zoobank.org:author:95327D0B-6BF0-447D-805C-11D266D68D88

urn:|sid:zoobank. org:author:54523608-3BD7-4A 99-BE63-D 1A AD2E715C5

3urn:Isid:zoobank.org:author:58A 3FE39-EC70-4F2C-B889-B2D 1 809ECE3E

Abstract. We examined morphometric variation in green toads of the genus Bufotes using univariate and multivariate

statistics in order to identify diagnostic characters. This study describes patterns of variation in two diploid (B. sitibundus

and B. perrini) and a tetraploid (B. pewzowi7) species and triploid hybrids of B. perrini and B. pewzowi distributed in Ka-

zakhstan and Kyrgyzstan (14 localities). Since body proportion characters were highly correlated, we divided them into

each other and obtained 190 indices. Using multivariate analyses we selected among them five most valuable. Discrimi-

nant analysis showed 95% of specimens correctly classified. The greatest importance for discrimination of diploid and

tetraploid species had two indices related to the parotoid gland size, distance between nostrils and length of foot. Between

diploid species the biggest differences were observed by two indices related to length of foot, radius bone size, and diame-

ter of the eye. Based on these features we obtained two multiplicative indices that allow us to reliably identify 85-94% of

specimens of each species. Hybrids usually had intermediate values of the indices compared to parental species (B. perrini

and B. pewzowi), mostly closer to the latter. The application of these diagnostic indices will allow non-invasive identifica-

tion of individuals of these diploid and tetraploid species during field research and study of museum specimens.

Key words. Amphibia, polyploidy, morphological variation, external characters, Kazakhstan, Kyrgyzstan.

INTRODUCTION

Polyploidy is an important evolutionary phenomenon.

The process of increase in the number of chromosome

sets and its consequences concerning morphological and

ecological differentiation of polyploids and their ances-

tors, phylogenetic origin and genome evolution of poly-

ploids, role of polyploidy in diversification have attracted

much attention (Soltis & Soltis 2012).

Eurasian green toads of the genus Bufotes Rafinesque,

1815 are one of the most famous examples of polyploid

Speciation among amphibians (Litvinchuk et al. 2016).

The genus includes ten diploid and five polyploid species

(Dufresnes et al. 2019). Based on phylogenetic recon-

structions, diploid species of the genus can be divided into

two main groups. The B. /atastii group consists of three

Species restricted to Iran, Iraq, Pakistan and the Indian

Himalayas. The B. viridis group includes seven species

distributed in North Africa, Europe, and the western part

of Asia. Some species from these two groups interbred

and gave origin to three triploid and two tetraploid spe-

cies which inhabit Central Asia and the Himalaya (Stock

Received: 03.02.2021

Accepted: 21.10.2021

et al. 2001, 2006; Litvinchuk et al. 2012; Betto-Colliard

et al. 2018; Dufresnes et al. 2019). In the B. viridis group,

Species are morphologically quite similar and usually can

only be identified using genetic characteristics (Stock

et al. 2006; Dufresnes et al. 2019). Variation of their mor-

phological characters is poorly studied.

Two diploid and one tetraploid species of green toads

are distributed in the northern part of Central Asia (Ka-

zakhstan and Kyrgyzstan). The variable toad, B. sitibun-

dus (Pallas, 1771), represents a diploid species that inhab-

its Western Asia, the Arabian Peninsula, the Caucasus, the

left bank of Volga River, and the southern Ural in Russia,

as well as western and northern Kazakhstan (Fig. 1). The

Perrin’s toad (B. perrini Mazepa, Litvinchuk, Jablonski,

and Dufresnes, 2019) is a recently described diploid spe-

cies that is distributed in northeastern Iran, northern Af-

ghanistan, Turkmenistan, Uzbekistan, northern Kyrgyz-

stan, and southern Kazakhstan (Fig. 1). The Pewzow’s

toad, B. pewzowi (Bedriaga, 1898) 1s a tetraploid species

of hybrid origin (Dufresnes et al. 2019) whose maternal

species is B. perrini and whose paternal species is anoth-

er diploid species B. /atastii (Boulenger, 1882). The spe-

Corresponding editor: W. Bohme

Published: 29.10.2021

362 Spartak N. Litvinchuk et al.

cies inhabits mostly mountainous regions in the east of

Central Asia and the Altai Mountains in Siberian Russia

(Fig. 1). In Kazakhstan and Kyrgyzstan ranges of B. per-

rini and B. pewzowi are parapatric and in some places are

slightly overlapping. Here the species can hybridize with

formation of a triploid offspring (Borkin et al. 1986a,

2001, 2007; Castellano et al. 1998, 2003; Odierna et al.

2004; Litvinchuk et al. 2006).

Analyses of morphological features in various green

toads were carried out by many researchers (Peters 1971;

Pisanets 1977; Hemmer et al. 1978; Pisanets & Szczer-

bak 1979; Borkin et al. 1986b, 2007; Ataev 1987; Stu-

gren & Tassoula 1987; Castellano et al. 1998, 2000; Tok

1999; Tosonoglu 1999; Sartaeva & Vashetko 2005; Kurt-

up et al. 2006; Lada 2012). However, diagnostic morpho-

logical characters which allow determining most of these

species were not revealed (Dufresnes et al. 2019). This

prevents the successful identification of species during

field research and study of museum specimens. There-

fore, the aim of the present paper was to find morpho-

logical differences which allow distinguishing species of

the genus Bufotes, which inhabit Kazakhstan and Kyr-

gyzstan.

MATERIAL AND METHODS

During the period from 1994 to 2001, a total of 209 adult

males of B. sitibundis (n=13), B. perrini (n=64), B. pew-

zowi (n=113) and hybrids of B. perrini and B. pewzowi

(n=19) from 14 localities were sampled in Kazakhstan

and Kyrgyzstan (Table 1; Fig. 1). Among them, 54 spec-

imens of all three species were studied for the first time.

Other 155 specimens of B. perrini, B. pewzowi and their

hybrids were previously analyzed by Castellano et al.

(1998, 2000). In the study, only adult males were used,

since they are in the spawning water bodies for a long

time, and therefore they are much easier to catch than

Table 1. Localities, coordinates, altitude, ploidy level, year of collection and sample size (N) of various species of green toads of

the genus Bufotes.

Locality Country Coordinates Altitude _Ploidy Year N

B. sitibundus

1 Atyrau Kazakhstan 47.233° N, 51.933° E -20 2n 2001 13

B. perrini

2 Tulek Kyrgyzstan 43.109° N, 74.092° E 758 2n 1994-1995 16

3. Kok-jar Kyrgyzstan 42.810° N, 74.641° E 935 2n 1995, 1997 20

4 Koi-Tash Kyrgyzstan 42.684° N, 74.670° E 1350 2n 1994, 1998 2

5 Kopa Kyrgyzstan 43.417° N, 75.783° E 750 2n 1994, 1995,1998 23

6 Kapchagai Kazakhstan 43.966° N, 77.362° E 554 2n 1997 3

B. perrini x B. pewzowi

3. Kok-jar Kyrgyzstan 42.810° N, 74.641° E 935 3n 19951997 5

4 Koi-Tash Kyrgyzstan 42.684° N, 74.670° E 1350 3n 1994 1]

6 Kapchagai Kazakhstan 43.966° N, 77.362° E 554 3n 1997 3

B. pewzowi

3.“ Kok-jar Kyrgyzstan 42.810° N, 74.641° E 935 An 1995 1

4 Koi-Tash Kyrgyzstan 42.684° N, 74.670° E 1350 An 1994, 1998 10

6 Kapchagai Kazakhstan 43.966° N, 77.362° E 554 4n 1997 2

7 ~~ Kizilkum Kazakhstan 42.065° N, 67.703° E 210 4n 1995 7

8 Jabagly Kyrgyzstan 42.433° N, 70.467° E 1100 An 1995 1]

9 ~~ Karaol Kazakhstan 45.842° N, 74.802° E 344 4n 1995 11

10 Zhidely Kazakhstan 45.300° N, 75.200° E 370 An 1995 3

11 Ayakoz Kazakhstan 48.000° N, 80.417° E 700 An 1997 10

12 Alma-Ata Kazakhstan 43.250° N, 76.956° E 846 An 1994-1995 28

13. Big Lake Kazakhstan 43.052° N, 76.988° E 2509 An 1994 12

14 Issyk-Kul Kyrgyzstan 42.200° N, 76.669° E 1750 An 1994 18

Morphology of green toads in Central Asia 363

Russia

Pakistan

Fig. 1. Distribution and localities of Bufotes sitibundus (blue range and rose circle), B. perrini (yellow range and dark blue circles),

and B. pewzowi (green range and red circles). Localities, where triploid hybrids of B. perrini and B. pewzowi were found together

with parental species, are designated by red stars. Numbers for localities are given in Table 1. Photo credit: LSN and DTN (male

of B. pewzowi from Kazakhstan in the right corner; male of B. perrini from Kazakhstan in the lower left corner; and male of B. si-

tibundus from Kazakhstan in the upper left corner).

females, which are breeding in water bodies for a very

short time.

Animals were anaesthetized in 1% solution of tricaine

methane-sulfonate (MS-222) and measured to the near-

est 0.01 mm with a digital caliper. Voucher specimens

are deposited in herpetological collections of the Depart-

ment of Life Sciences and Systems Biology of Universi-

ty of Turin (Turin, Italy) and the Institute of Zoology of

the Republic of Kazakhstan (Almaty, Kazakhstan). We

measured twenty body proportion characters following

Castellano & Giacoma (1998): snout-vent length (SVL);

length of the head which was measured from the tip of

snout until the posterior edge of the mandible (LHEAD);

width of the head (distance between the posterior edg-

es of mandibles; WHEAD); minimum distance between

the nostrils (INTNOS); distance between nostrils and

the tip of snout (NOSTIP); minimum distance from the

nostril opening to the anterior corner of the eye (NOS-

EYE); minimum distance from the eye to the tympanum

(EYETYM); horizontal diameter of the eye (DEYE),; ver-

tical diameter of the tympanum (DTYM); length of paro-

toid glands (LPAR); distance between the elbows when

animal is kept with humerus at 90° degree angle with

respect to the body axis (WGRASP); length of the ra-

dio-ulna (RADUL); length of the hand (LHAND); length

of the first finger of the hand (LIFING); length of the

femur (LFEM); length of the tibia (LTIB); length of the

Bonn zoological Bulletin 70 (2): 361-371

tarsus (LTARS); length of the foot (LFOOT); minimum

distance from the distal extremity of the inner metatarsal

tubercle to the web between the third and fourth finger

(WEB), and length of the metatarsal tubercle (LMET).

Since all characters were highly correlated with each

other (n=209; r=0.55—0.98; P<0.05), we divided them

into each other and obtained 190 ratios (indices). Then,

for all indices the natural logarithm conversion was made.

Using the method of principal components, we selected

the 118 most variable indices (factor loadings more than

|0.035|) on the first two axes (30.6% and 15.5% of vari-

ance). This reduction in the number of indices allowed

us to use the discriminant analysis (species as a group-

ing variable), which was applied in order to find most

valuable indices for differentiation of the three studied

species. Based on the results, five most distinguishing

indices (NOSTIP/INTNOS, LFOOT/RADUL, LFOOT/

LPAR, LPAR/INTNOS, and WEB/DEYE) were select-

ed and used for all subsequent calculations. Additionally,

to estimate position of hybrid samples we performed the

second discriminant analysis with populations (n>3) as a

grouping variable. All these analyses were performed us-

ing Past ver. 4.03 (https://past.en.lo4d.com/). To perform

univariate analyses of variance, we applied the one-way

ANOVA for comparison of means and the Sheffe test for

post-hoc comparisons (P<0.001) which were calculated

in Statistica ver. 8.0.

©ZFMK

364 Spartak N. Litvinchuk et al.

DA axis 2

sitibundus

perrini

DA axis 1

Fig. 2. Discriminant analysis biplot (species as a grouping variable) of morphometric characters and indices for males of Bufotes

sitibundus (2n), B. perrini (2n) and B. pewzowi (4n).

To obtain karyotypes for each specimen, venous blood

was incubated 100-200 ul for 4 days at 25°C in MEM

(Minimal Eagle Medium, GIBCO) with 20% calf serum

and 3% Phytohaemogglutinin M. Chromosome prepara-

tions were produced by conventional air-drying method,

using KCI 0.075M as a hypotonic solution. Standard

staining method was performed using Giemsa 5% in

phosphate buffer pH 7 (Castellano et al. 1998).

RESULTS

The one-way ANOVA analysis showed significant differ-

ences among species for all five selected indices (F=57.6,

df=10, p < 0.001). The post-hoc comparisons revealed

that B. pewzowi significantly differed from both diploid

species by relatively higher values of the index LFOOT/

LPAR and relatively lower values of indices NOSTIP/

INTNOS and LPAR/INTNOS (Table 2). Additionally,

Table 2. Variability (X+SD; range) of snout-vent length and morphometric indices in three species of green toads and triploid

hybrids between Bufotes perrini and B. pewzowi. N = sample size.

Character sitibundus perrini perrini x pewzowi ~— pewzowi

N 13 64 19 M3

SVL 67.5+8.1 87.0+7.1 qo Sty 2 71.5+6.4

(57.0-80.1) (70.1—100.0) (64.8—87.3) (49.4—-84.8)

NOSTIP/AINTNOS 1.34+0.11 1.2740.08 1.20+40.09 LAgt0.12

(1.14-1.53) (1.09-1.51) (1.04—1.36) (0.76—1.43)

LFOOT/RADUL 1.86+0.08 1.6840 .06 1.80+40.05 1.82+40.07

(1.74-1.97) (1.52-1.83) (1.71-1.89) (1.65—2.00)

LFOOT/LPAR 1.7540,13 1.62+0.13 2.08+0.15 2.19+0.21

(1.50-2.00) (1.36—2.04) (1.81-2.41) (1.51-2.65)

LPAR/INTNOS 4 3640.37 4 44+0.44 3.53+0.43 3.40+0.53

(3.77-4.91) (3.51-5.62) (2.82-4.55) (2.11—5.20)

WEB/DEYE 3.01+0.23 2.78+0.18 2.96+0.20 2.94+0.23

(2.79-3.45) (2.48-3.23) (2.58-3.30) (2.42-3.53)

MII 2.51+£0.39 2.7840.43 1.724£0°33 1.59+0.41

(1.93-3.28) (1.72-3.82) (1.17—2.48) (0.83-3.44)

MI2 5.74+0.59 4 6840.40 5.32+0.43 5.35+0.48

(4.87-6.75) (3.95-5.82) (4.65-5.95) (4.34-6.55)

Bonn zoological Bulletin 70 (2): 361-371

©ZFMK

Morphology of green toads in Central Asia 365

DA axis 2

DA axis 1

Fig. 3. Plot of centroids for males of Bufotes perrini (red circles), B. pewzowi (blue) and triploid hybrids between B. perrini and

B. pewzowi (green) in the space of the first and second canonical discriminant axes. Numbers for localities see in Table 1.

B. pewzowi differed from B. perrini by having a small-

er body length and higher values of indices LFOOT/

RADUL and WEB/DEYE. Among diploids, B. perrini

differed from B. sitibundus by having a larger body length

and lower values of indices WEB/DEYE and LFOOT/

RADUL. Hybrids had usually intermediate values of the

indices as compared to their parental species (B. perrini

and B. pewzowi). Among B. pewzowi populations, toads

from Kizilkum obviously differed from other popula-

tions by indices LFOOT/LPAR and LPAR/INTNOS that

related to the parotoid gland size (Table 3).

In the discriminant analysis, the first axis included

89% of variation and the second 11%. All three species

100

SVL

1 1

perrini x pewzowi

pewzowi

. 1 1

sitibundus perrini

i 1

sitibundus perrini

formed distinct clusters with some overlap (Fig. 2). In

general, the overall correct classification rate was 94.7%,

where 93.8% were for B. perrini (two individuals were

confused with B. sitibundus and two with B. pewzowi),

94.7% for B. pewzowi (two individuals were confused

with B. perrini and four with B. sitibundus), and 100%

for B. sitibundus. In the discriminant analysis with pop-

ulations as a grouping variable (Fig. 3), centroids of

samples of B. perrini and B. pewzowi formed separate

clusters. Two hybrid triploid populations were close to

B. pewzowi, but a triploid population from Koi-Tash (lo-

cality 4) was in intermediate position between B. perrini

and B. pewzowi.

1 1

perrini x pewzowi

pewzowi

1 '

sitibundus perrini

1 1

perrini x pewzowi

pewzowi

Fig. 4. Variability (violin plots) of SVL, MI1 and MI2 indices in males of Bufotes sitibundus (2n), B. perrini (2n), B. pewzowi (4n),

and triploid hybrids between B. perrini and B. pewzowi.

Bonn zoological Bulletin 70 (2): 361-371

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366

Spartak N. Litvinchuk et al.

Table 3. Variation (X+SD) of snout-vent length and morphometric indices in populations of green toads of the genus Bufotes. N =

size. Numbers for localities see in Table | and Fig. 1.

NOSTIP/ LFOOT/ LFOOT/ LPAR/ WEB/

Locality N SVL MI1 MI2

INTNOS RADUL LPAR INTNOS' DEYE

B. sitibundus

1 Atyrau 13. 67.5+8.1 1.3440.11 1.8640.08 1.7540.13 4.3640.37 3.0940.23 2.5140.39 5.74+0.59

B. perrini

2 Tulek 16 87.448.2 1.2540.08 1.6440.06 1.6040.09 4.4540.32 2.6640.14 2.8140.33 4.3740.29

3. Kok-jar 20 84.247.3 1.2840.09 1.7240.05 1.6440.13 4.5040.48 2.8740.20 2.7840.46 4.95+0.42

4 Koj-Tash 2 89.8+1.5 1.3140.10 1.67+0.03 1.5340.09 5.08+0.76 2.9040.13 3.3440.69 4.84+0.13

5 Kopa 23 88.6+7.3 1.2540.06 1.6840.06 1.6040.13 4.3140.43 2.7840.14 2.7340.43 4.66+0.31

6 Kapchagai 3 = 88.942.0 1.3540.04 1.6740.02 1.7340.19 4.5240.39 2.7040.13 2.6540.50 4.53+0.17

B. perrini x B. pewzowi

3. Kok-jar 5 74.946.2 1.18+0.06 1.8440.05 2.0940.11 3.4440.29 2.9640.24 1.6540.21 5.44+0.48

Koi-Tash 11 73.643.5 1.2040.10 1.7740.05 2.1140.14 3.4340.33 2.9940.15 1.6440.25 5.30+0.38

6 Kapchagai 3 82.1445 1.2140.10 1.8340.05 2.0440.16 3.6840.53 2.9240.26 1.8340.41 5.34+0.51

B. pewzowi

3.‘ Kok-jar 1 SBeS) 1.14 1.89 251 3.10 3.04 1-23 55

4 Koi-Tash 10) 73.145.7 1.1840.07 1.8240.09 2.1140.20 3.6340.35 2.9840.21 1.7440.29 5.42+0.54

6 Kapchagai 2 78.747.6 1.22+0.13 1.77+0.03 2.1340.26 3.6140.09 2.7340.02 1.7140.25 4.84+0.10

7 ~~ Kizilkum 71.943.7 1.16+40.08 1.8640.08 1.6940.12 4.4740.62 2.7040.15 2.68+0.54 5.04+40.36

8 Jabagly Me = “733338 SI 28£0:08" 179008 8 29.2950 1 Sie «3.592034. 9D S020 13" “1632023. ss: 232037

9 Karaoi 11 =62.743.8 1.1840.10 1.8240.08 2.1240.10 3.3440.30 2.6140.12 1.5840.15 4.75+0.38

10 Zhidely 3 59.7£11.3 1.2140.07 1.7740.08 2.2140.14 3.5440.02 2.7440.29 1.6140.11 4.84+0.59

11 Ayakoz 10 68.342.1 0.9840.15 1.8640.06 2.36+0.17 2.6440.38 3.0040.24 1.1240.18 5.61+40.53

12. Alma-Ata 28 74.744.7 1.22+0.11 1.8040.07 2.2440.18 3.514+0.40 3.0940.18 1.5840.28 5.57+40.42

13. Big Lake 12 77.045.8 1.1640.09 1.8340.06 2.12+0.08 3.4240.31 2.9640.11 1.6240.19 5.42+0.28

14. Issyk-Kul 18 68.9449 1.1140.1 1.8440.06 2.2940.09 3.0040.16 2.9540.18 1.3140.11 5.42+0.36

The greatest impact to discrimination of diploid and

tetraploid species had the indices LPAR/INTNOS and

LFOOT/LPAR (Table 2, Fig. 2). We divided the first in-

dex by the second and obtained the first multiplicative

index MI] = LPAR?/((INTNOSxLFOOT). A comparison

of diploids and polyploids with use of this index showed

93% of correct identifications in total. Most males of

B. perrini (94%) and B. sitibundus (85%) have values

of more than 1.90, and vice versa, values of B. pewzowi

(94%) and triploid hybrids (100%) were lower than that

(Figs 4-5).

The indices LFOOT/RADUL and WEB/DEYE

demonstrated the biggest differences between diploid

species (Table 2; Fig. 2). We multiplied these indices by

each other and obtained the second multiplicative index

MI2 = (LFOOTxWEB)/((RADULxDEYE). In total, 86%

of identifications using the index were correct. Most in-

dividuals of B. perrini (88%) have values of the index

Bonn zoological Bulletin 70 (2): 361-371

below 1.27 and most B. sitibundus (77%) have values

above that number (Figs 4—5).

DISCUSSION

Previous studies of the morphometric variation in am-

phibians suggest that climatic and ecological conditions

are correlated with each other and thus may drive differ-

ences in overall body shape (e.g., Castellano & Giacoma

1998; Amor et al. 2011). Based on previous studies of

morphological variation in green toads, several general

trends can be identified. For example, diploid species can

be divided by body length (SVL) into two main groups

which were also revealed in phylogenetic reconstructions

(Dufresnes et al. 2019). Representatives of the B. viridis

group are characterized by relatively large body length,

while members of the B. /atastii group are usually small.

©ZFMK

Morphology of green toads in Central Asia 367

C) B. sitibundus

C) B. perrini

@ B. perrini < pewzowi

MI2

—ee

density

Fig. 5. Distribution of indices MI1 and MI2 in males of Bufotes sitibundus (2n), B. perrini (2n), B. pewzowi (4n), and triploid hy-

brids between B. perrini and B. pewzowi.

Polyploid species of hybrid origin occupy an intermedi-

ate position. Our findings support this regularity. In Ka-

zakhstan and Kyrgyzstan, adult males of diploid species

of the B. viridis group have usually larger average SVL

(80-87 mm in B. perrini and 68-80 mm in B. sitibundus)

as compared to polyploid B. pewzowi (62-73 mm). In

other regions, maximum values of average body length

of males of diploid toads were 83 mm for B. sitibundus

and 82 mm for B. perrini, whereas the maximum value of

average body length of tetraploid B. pewzowi was 71 mm

(Table 4).

The study on Israeli green toads has demonstrated that

variation of body size was partly a function of climate

humidity (Nevo & Schneider 1976). In amphibians, vol-

ume to surface ratios in larger species would help con-

serve water in unpredictable environments, and therefore

larger species will have greater tolerance to desiccation

(Bidau et al. 2011). For the Central Asian green toads this

suggestion 1s only partially true. As arule, B. perrini with

its largest body size inhabits very dry deserts. Interme-

diate B. sitibundus live in more moist steppes. Howev-

er, B. pewzowi with smallest SVL as a rule inhabits both

relatively moist mountain regions as well as dry moun-

tains (Eastern Tien-Shan) and deserts of premontane and

intermontane depressions (Stock et al. 2001, 2006) with

Bonn zoological Bulletin 70 (2): 361-371

extreme continental climate and maximal humidity defi-

ciency (Murzayev 1966; Vilesov et al. 1986).

Several mechanisms ensure the resistance of amphib-

ians to land conditions among which the skin plays an

important role. Unlike diploid green toad species, the

skin of tetraploid B. pewzowi contains unusual mucous

glands (“mosaic glands”), which along with typical mu-

copolysaccharide components, produce a high amount of

proteins (Fedotovskikh et al. 2020). The predominance

of protein secretion is considered a characteristic of the

terrestrial amphibians (Fujikura et al. 1988; Duellman &

Trueb 1994) and we cannot exclude that the “proteiniza-

tion“ of mucous glands enhances the protective proper-

ties of the toad’s skin and provides additional resistance

to desiccation without increasing the overall size of the

animals.

The size of the parotoid glands is probably also asso-

ciated with the described phenomenon, since the mosaic

glands were found directly in the parotoid and the dorsal

skin. In our study, B. perrini had the longest (and larg-

est) parotoids, B. sitibundus had intermediate ones, while

B. pewzowi had the shortest. Similar regularities in the

gland size variability among green toad species were pre-

viously observed by other authors (Eiselt & Schmidtler

1973; Pisanets 1977; Andrén & Nilson 1979; Borkin et al.

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368

Spartak N. Litvinchuk et al.

Table 4. Variation (X+SD) of snout-vent length in adult males of three species of green toads of the genus Bufotes (N>3). N =

sample size.

Country N X+SD (range) References

B. sitibundus

Kazakhstan 13 67.5+8.1 (57.0—80.1) Present paper

Kazakhstan 3 80.1+0.5 (79.7—80.6) Borkin et al. (2007)

Russia (Ural) 29 61.5 Toporkova (1981)

Russia (Dagestan) 26 67.2+1.1 (54.7—75.6) Pysanets (2014)

Russia (Dagestan) 90 70.6 Khonyakina (1980)

Russia (Chechnya) 12 53.046.5 Pisanets (1977)

Russia and Georgia 65.0 (55.0-75.0) Hemmer et al. (1978)

Georgia 9 (52.0-86.6) Tarkhnishvili & Gokhelashvili

(1999)

Armenia 18] 70.2 Melkumyan & Pisanets (1987)

Azerbaijan 12 56.3+3.9 Melkumyan & Pisanets (1987)

Turkey 67.9+5.7 (60.0—76.0) Flindt & Hemmer (1968)

Turkey 5 75.4 Altunisik et al. (2015)

Turkey 57 62.6 (47.9-82.8) Altunisik & Ozdemir (2015)

Turkey 50 70.0 Kurtup et al. (2006)

Turkey 54 69.2 Kurtup et al. (2011)

Turkey 90 64.1 Altunisik et al. (2021)

Israel 85 75.2 (60.0-88.0) Nevo & Schneider (1976)

Egypt (Sinai) 16 83.3 (71.0-93.0) Nevo & Schneider (1976)

Iran (Cheshmeh-ye Sefied) 5 65.6+2.7 (62.0-69.0) Andren & Nilson (1979)

Iran (Central Zagros) 26 72.7+3.8 Ashkavandi et al. (2012)

Iran, Iraq and Israel 25 67.1 (57.0-78.0) Eiselt & Schmidtler (1973)

Iran (Kerman Province) ? 70.0 (60.0—80.0) Hemmer et al. (1978)

Tran i; 75.9+8.6 (63.1—88.8) Dufresnes et al. (2019)

B. perrini

Kazakhstan 64 87.0£7.1 (70.1-100.0) — Present paper

Kazakhstan ? 80.0 (55.3—95.5) Sartaeva & Vashetko (2005)

Uzbekistan, Turkmenistan, ke (45.0-80.0) Hemmer et al. (1978)

Afghanistan and Iran

Uzbekistan and Turkmenistan fe) 81.9+5.3 (76.1—-86.6) Dufresnes et al. (2019)

Turkmenistan 7 71.1+7.0 Pisanets (1977)

Turkmenistan 12] 77.2 (56.7-91.3) Pisanets & Mezhzherin (1996)

B. pewzowi

Kazakhstan 113 71.5+6.4 (49.4—84.8) Present paper

Kazakhstan 1 (68.0—77.8) Bassalaeva et al. (1998)

Kazakhstan 2 64.2 (48.3-77.0) Sartaeva & Vashetko (2005)

Kazakhstan 1] 62.1 (55.1-67.5) Borkin et al. (2007)

Kyzgyzstan 2 (45.0-60.0) Andrushko (1951)

Bonn zoological Bulletin 70 (2): 361-371

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Morphology of green toads in Central Asia

Table 4. Continued.

369

Country N X+SD (range) References

B. pewzowi

Kyrgyzstan 5 72.8 (56.4—82.7) Pisanets & Szczerbak (1979)

Mongolia, China, Kazakhstan, Kyrgyzstan, ? (50.0-80.0) Hemmer et al. (1978)

Uzbekistan, and Tajikistan

Mongolia ? (?—78.0) Peters (1971)

Mongolia ? (?-71.9) Borkin et al. (1986b)

Mongolia 5 67.5+2.8 (65.3—72.3) Dufresnes et al. (2019)

China 42 66.1 (51.6—-76.5) Fei et al. (1999)

China 7 57.7+5.0 (50.1-63.0) Stock et al. (2001)

China 3 65.4+6.0 (59.4—71.5) Dufresnes et al. (2019)

Uzbekistan isi 64.2 (48 .3-77.0) Sartaeva & Vashetko (2005)

Uzbekistan 13 69.1+4.8 Pisanets (1977)

Tajikistan 7 Plsl29 Pisanets (1977)

Tajikistan 8 Goe7671 Pisanets (1977)

Tajikistan 45 68.1 (57.1—79.2) Pisanets & Szczerbak (1979)

Tajikistan and Uzbekistan 20 68.4+8.1 (47.6—80.7) Kondratova et al. (2020)

Uzbekistan 12 64.6 (57.1-60.0) Pisanets & Szczerbak (1979)

1986b; Ataev 1987; Pisanets & Mezhzherin 1996; Cas-

tellano et al. 1998; Fei et al. 1999; Dufresnes et al. 2019).

In our study, this trend is reflected by the indices associ-

ated with the length of the parotoids (LPAR/INTNOS,

LFOOT/LPAR and MI1), which were the most useful

indices for discriminating diploid and polyploid species.

However, the length of the parotoids can be affected by

other environmental factors. For example, among diploid

green toads lotic breeders, such as B. surdus (Boulenger,

1983) and B. /uristanicus (Schmidt, 1952), have relative-

ly smaller parotoids, as compared to lentic species (other

eight species). Probably, small parotoids help to improve

the streamlining of these toads. However, all green toad

species in Central Asia are lentic breeders and therefore,

this regularity should not affect the size of their parotoids

here.

It should be noted, that previously Castellano et al.

(1998) concluded that the best criteria for discriminating

the Central Asian green toad species are their body size

and the relative dimensions of their head and limbs. Ac-

cording to our data, the multiplicative indices MI1 and

MI2 were most informative for the distinction between

three local green toad species. Use of these indices al-

lows to identify 85-94% of specimens of each species.

However, our findings need to be confirmed in a broader

study covering a larger number of samples (especially

B. sitibundus). It is possible that an addition of such char-

acters as the form of parotoid glands to the analysis will

increase success in the species identification.

Bonn zoological Bulletin 70 (2): 361-371

Acknowledgments. The reported study was funded by the Rus-

sian Foundation for Basic Research according to the research

project N° 20-04-00918 and by the Science Committee of the

Ministry of Education and Science of the Republic of Kazakh-

stan to the Program No. BR10965224. We are very grateful

to S. Castellano and G. Odierna for valuable help with the re-

search and to F. Letoutchaia for comments.

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©ZFMK

BHL

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Bonn zoological Bulletin 70 (2): 373-375

2021 - Casiraghi A.

https://do1.org/10.20363/BZB-2021.70.2.373

ISSN 2190-7307

http://www.zoologicalbulletin.de

Scientific note

urn:|sid:zoobank.org: pub:3B5488F7-326A-4276-B266-3 BCA 937DF656

First record of the invasive grapevine aphid Aphis illinoisensis Shimer, 1866

(Hemiptera: Aphididae) in mainland Italy

Alice Casiraghi

Instituto Valenciano de Investigaciones Agrarias. (IVIA). Unidad de Entomologia, Centro de Proteccion Vegetal y Biotecnologia.

Ctra. Moncada-Naquera Km. 4,5. E-46113 Moncada, Valencia, Spain

Email: casiraghi_alia@gva.es

urn:|sid:zoobank.org:author:905837A0-9C77-4D57-A895-B19DD9FF790B

Abstract. Apterous and winged viviparous females of the Nearctic Aphis illinoisensis Shimer, 1866 have been recorded

for the first time in mainland Italy (Rome) on branches of Vitis labrusca L. in 2021, thanks to photos published in a Fa-

cebook garden group.

Key words. Invasive species, citizen science, grapevine pest, new record, Vitaceae.

Garden and insect groups on Facebook are typically ded-

icated to questions about plant health and pest problems,

but sometimes they can result in being a source of in-

formation about exotic or invasive species detection and

expansion (Seat 2015, Martinovic et al. 2019, Vuji¢ et al.

2021). In our case, a photograph of aphids on fox grape

Vitis labrusca L. taken on 17" September 2021 in a pri-

vate garden in the urban area of Colli Aniene (Rome, It-

aly) showed the first record of the Nearctic species Aphis

illinoisensis Shimer, 1866 (Hemiptera: Aphididae) in

mainland Italy.

The colony, composed by both apterous and winged fe-

males and their nymphs, was not attended by ants (Fig. 1).

Apterous females are characterized by a shiny dark red-

dish brown colour, wholly black hind tibiae and dark,

long and divergent siphunculi. Winged females are simi-

lar, and nymphs typically present a lighter reddish-brown

colouration with black siphunculi (Blackman & Eastop,

2021). The identification was confirmed by aphidologist

Nicolas Pérez Hidalgo (Departamento de Artropodos,

Museo de Ciencias Naturales de Barcelona, Spain).

Unfortunately, specimens could not be collected nor

more and detailed photos be taken, since the colony has

been immediately fumigated after the identification and

up to date no other winged female colonized the areas.

Contact with the author of the photograph has been kept

open in order to await new developments.

The invasive American grapevine aphid is native to

North America (Blackman & Eastop, 2021), but it is

also widespread in Central and South America (Ramirez

Guevara & Zuluaga 1995, Ortego et al. 2004, Trejo-Loyo

et al. 2004, Etienne 2005, Zamora Mejias et al. 2010),

where it presents a dioecious holocycle, host-alternating

Received: 01.10.2021

Accepted: 03.11.2021

between Viburnum prunifolium L. (Adoxaceae) and Vi-

taceae plants (Blackman & Eastop 2021). In 2002 A. illi-

noisensis was first recorded in the Palearctic, more specif-

ically in Turkey (Remaudiere et al. 2003). Since then, the

species has been reported throughout the Mediterranean

Basin and Middle East, 1.e., Crete, Greece (Tsitsipis et al.

2005), Cyprus (Kocadal & Ulusoy 2006), Spain (Pérez

Hidalgo et al. 2011), Israel (Barjadze & Ben-Dov 2011),

Tunisia (Kamel-Ben Halima & Mdellel 2010), Algeria

(Laamari & Coeur d’Acier 2010), Montenegro (Petro-

vic-Obradovic et al. 2010), Libya (Havelka et al. 2011),

Malta (Mifsud & Pérez Hidalgo 2011), Egypt (El-Gantiry

et al. 2012), Saudi Arabia (Hussain et al. 2015) and Iran

(Heidarnia & Derakhshan 2018). In Italy, this species was

documented from the island of Sicily, in the province of

Catania (Cocuzza & Barbagallo 2011). More recently,

it has been recorded even in Central Europe: Slovenia

(Seljak 2021) and France (Mouttet & Balmes 2021). On

the Eurasian continent A. i/linoisensis seems to develop

anholocycles on Vitis plants and no sexual morphs have

been recorded (Blackman & Eastop 2021). Its expansion

appears to be a product of the movement of winged alatae

(Havelka et al. 2011).

The grapevine aphid is considered invasive because of

the potential damage caused to the wine and grape pro-

duction. It has been recorded as a vine pest that can de-

crease the developing of young shots, leaves and fruits

and produce leaf necrotic spots (McGrew & Still 1979,

Cocuzza & Barbagallo, 2011), but its impact in Europe

does not appear to be as important as the one produced

by the grape phylloxera Daktulosphaira vitifoliae (Fitch,

1855) (Hemiptera: Phylloxeridae) (Kamel-Ben & Mdel-

lel 2010, Havelka et al. 2011). For the moment, because

Corresponding editor: R. Peters

Published: 05.11.2021

374 Alice Casiraghi

Fig. 1. Colony of Aphis illinoisensis from Rome (Italy) with

winged and apterous female morphs on Vitis labrusca. The pho-

to was taken by the Italian citizen Sara Bonanno in her garden

in the neighbourhood of Colli Aniene and posted in a Facebook

garden group about plant pests.

of its occasional records, A. i/linoisensis can be consid-

ered a minor pest of vines in Italy.

Once again, citizen science allows specialists to gain

more information about the real distribution of exotic and

invasive species.

Acknowledgments. The author is sincerely thankful to Dr.

Nicolas Pérez Hidalgo who confirmed aphid identification and

to Sara Bonanno who decided to share their photos and geo-

graphical information. She also wants to thank the reviewers

for their work and effort to improve this little scientific note.

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Bonn zoological Bulletin 70 (2): 377-382

2021 - Volynkin A.V.

https://do1.org/10.20363/BZB-2021.70.2.377

ISSN 2190-7307

http://www.zoologicalbulletin.de

Research article

urn:|sid:zoobank.org:pub: BE8963B5-EB9A-48A5-95 15-E2D3C95F285C

Juxtilema, a new genus for a new species from Zambia

(Lepidoptera: Erebidae: Arctiinae: Lithosiini)

Anton V. Volynkin

The African Natural History Research Trust (ANHRT), Street Court Leominster-Kingsland, HR6 9QA, United Kingdom

Altai State University, Lenina Avenue, 61, RF-656049, Barnaul, Russia

Email: anton@anhrt.org.uk.com

urn:Isid:zoobank.org:author:25EA 1403-9502-42 BE-A820-96ACS5 1CE862E

Abstract. The present paper contains a description of the new genus Juxtilema gen. nov. which is erected for the new

species Juxtilema smithi gen. et sp. nov. from Zambia. The diagnostic comparison is made with the genera Lophilema

Aurivillius, 1910 and Oedipygilema Kruger, 2015. The new genus is characterised by anautapomorphic feature in the male

genitalia, 1.e., the structure of the juxta which is heavily sclerotised, extremely elongate, cylindrical and evenly curved,

bearing a densely setose area apically and medially, and strongly connected to the bases of the sacculi. Adults together

with male and female genitalia of the new and similar genera are illustrated.

Key words. Lithosiina, Sub-Saharan Africa.

INTRODUCTION

Despite recent publications devoted to the generic classi-

fication of the Afrotropical Lithosiina, or footman moths

(Kruger 2015, 2016; Volynkin & Laszl6 2021), the gener-

ic assignment of a number of groups remains uncertain.

In the course of identifying the Lithosiini housed in the

African Natural History Research Trust, Leominster, a se-

ries of an unknown Lithosiina species from Zambia was

found. Although it displays a typical ‘eilemoid’ pattern

common not only for Afrotropical but also Asiatic and

European members of the subtribe Lithosiina, the genital

structures of both sexes of the species are considerably

different from those of all other Afrotropical genera sug-

gesting that it belongs to a distinct monophyletic lineage

that not only represents a hitherto undescribed species but

also a new genus, the descriptions of which are provided

herein.

Abbreviations of the depositories

ANHRT = African Natural History Research Trust,

Leominster, UK

NHMUK = Natural History Museum, London, UK

(formerly BMNH)

Other abbreviations

AV = genitalia slide prepared by A.V. Volynkin

HT = holotype

PT = paratype

Received: 10.09.2021

Accepted: 12.11.2021

The genitalia were dissected and mounted in euparal on

microscope slides. The photos of adults were taken using

a Nikon D3100/AF-S camera equipped with a Nikkor,

18-55 mm lens while the photos of genitalia were tak-

en by the same camera attached to a microscope with an

LM-scope adapter. All images were processed using the

Adobe Photoshop ver. CC 2018 software.

RESULTS

Juxtilema gen. nov.

urn: lsid:zoobank.org:act: FC 103268-29B 3-497 F-A09A-A386461BOF 15

Type species: Juxtilema smithi gen. et sp. nov.

Diagnosis

The type species of the new genus (Figs 1—2) displays a

characteristic ‘eilemoid’ pattern and is externally similar

to members of genera such as Manulea Wallengren (illus-

trated by Dubatolov & Zolotuhin (2011) and Witt et al.

(2011)), Mimelilema Kriger, Gracililema Kriger and

Pseudotigrioides Kriger (illustrated by Kruger (2015)).

Nevertheless, the male genitalia ground plan of Juxtilema

gen. nov. is most similar to that of the genus Lophilema,

members of which (Figs 5—8) considerably differ exter-

nally from the new genus in the limited sexual dimor-

phism (strongly expressed in Lophilema) and the absence

of a medial androconial area on the upper side of the

male forewing. The male genital capsule structure of Jux-

tilema gen. nov. (Fig. 9) is similar to that of Lophilema

Corresponding editor: M. Espeland

Published: 21.11.2021

378 Anton V. Volynkin

1 ZAMBIA 14/m

Lukwakwa, Wes

Cryptosepalum oe

ANHRTUK =e eae reE LED LightT

AVG466 7 rey |geeee Meola

00074156 A. Volynkin | ANHRT: 301840" ii

Juxtilema smithi gen. & sp. n., HT @

3 ZAMBIA 1365m

dan Fisher's Farm, =

ong Riverine for

Slide

ANHRTUK | avea61. 67 eae

cect

00132516 a Volynkin _| Munaney. te

Bashford, M

Oedipygilema aposema &

"ae

AV3189 o”

A. Volynkin |

Lophilema mbulu, HT 3

) :

NHMUK 010915952

Lophilema polioplaga, HT &

Z ZAMBIA 147m

Lukwakwa, West Lunga NP.,

um netses pee

ANHRTUK AV6458 g 9-15.xi, 2018 mee

Juxtilema smithi gen. & sp. n., PT

4 ZAMBIA 1147m

Lukwakwa, enag in NP.,

ANHRTUK | avieace eel

RTUK | Av6462 £. ee

00056737 | A volynkin | $3322,S, Oram.

Slide

AV4585 ¢ |

+ _A.Volynkin

Lophilema mbulu, PT @

i Slide .-—Ss TANZANIA 1679m

Mount Meru, Arusha N.P.

AV3373 = $03°14°51"E36°50"38”

. 18-24.vii.12 Light Trap

A. Wolynkin jeg. smith & Takano,

Lophilema polioplaga °

Juxtilema gen. nov. for a new species from Zambia 379

(Figs 11-12) in the elongated saccus with thin margins, a

strongly elongated intravincular corema and the distally

protruding medial section of the sacculus (similar to only

certain species of the genus, e.g., L. mbulu Volynkin,

2019). However, the male genital capsule of the new ge-

nus is distinguished from that of Lophilema by the heavi-

ly sclerotised, extremely elongate, cylindrical and evenly

curved juxta which is strongly connected to the bases of

the sacculi and bearing a densely setose area apically and

medially (an autapomorphic feature), whereas the juxta

of Lophilema is weakly sclerotised, short, dorso-ventral-

ly flattened and bears basal and distal concavities. Addi-

tionally, compared to Lophilema, the uncus of Juxtilema

gen. nov. is setose, thick and triangular in cross-section (it

is smooth, slender and laterally flattened in Lophilema),

the processus distalis plicae is present, the distal saccu-

lar process is smooth (densely setose in Lophilema), and

the anellus bears triangular sclerotised protrusions which

are absent in Lophilema. Among Afrotropical Lithosiina

genera, an extremely elongated and apically setose juxta

is also known in the genus Oedipygilema (Figs 3—4), but

in the latter it is dorso-ventrally flattened, straight and

has two longitudinal latero-ventral ribs connected prox-

imally by a membrane (Fig. 10). In addition, the genital

capsule of Oedipygilema differs from that of Juxtilema

gen. nov. in the presence of a strongly elongated and

thin basal costal process, the lack of a processus dista-

lis plicae, the short and apically well-sclerotised saccus

lacking a corema, and the longer, thinner and laterally

flattened uncus (it is relatively short, thick and triangular

in cross-section in Juxtilema gen. nov.). The phallus of

Juxtilema gen. nov. is longer and thicker (in proportion

to the genital capsule size) compared to Lophilema and

Oedipygilema. In the vesica, the presence of a sclerotised

dentate distal patch is characteristic for the new genus.

The female genitalia of Juxtilema gen. nov. (Figs 13-14)

are characterised by the ventro-laterally positioned os-

tium bursae, a character considered here as autapomor-

phic, whereas in other Afrotropical Lithosiina genera

including Lophilema (Figs 16, 17) and Oedipygilema

(Fig. 15) the ostium bursae is positioned medio-ventrally.

Description

External morphology of adults (Figs 1—2). Forewing

length 13-15 mm in males and 14-16 mm in females.

Sexual dimorphism limited: female slightly larger than

male and having somewhat more convex forewing costal

margin with slightly narrower costal stripe. Antennae of

both sexes sparsely ciliate with somewhat shorter ciliae in

females. Head deep yellow. Thorax plumbeous-grey, pa-

tagia deep yellow. Forewing ground colour plumbeous-

grey with wide deep yellow costal stripe, thin deep yel-

low terminal line and cilia. Hindwing monotonous pale

ochreous-yellow. Abdomen pale yellow proximally and

ochreous-yellow distally.

Bonn zoological Bulletin 70 (2): 377-382

Male genitalia (Fig. 9). Uncus weakly setose, thick,

triangular in cross-section, tapered distally with short

claw-shaped tip. Tuba analis with thin and weakly scle-

rotised scaphium. Arms of tegumen wide and fused in

posterior two-thirds. Vinculum ca. twice longer than teg-

umen, with well-sclerotised but thin arms connected by

thin and weakly sclerotised commissure distally. Intra-

vincular area wide, membranous and with long expand-

able medial corema bearing numerous long androconial

scales. Dorsal section of valva short, apically rounded and

setose, processus distalis plicae represented as transverse

sclerotised crest, setose dorsally. Sacculus heavily scle-

rotised and flattened with strongly ventro-distally pro-

truding, rounded and sparsely setose distal section. Distal

saccular process originating from the dorsal side of dis-

tal section of sacculus, slender, smoothly down-curved,

distally tapered and apically rounded. Juxta extremely

long, reaching the base of uncus, heavily sclerotised, tu-

bular, smoothly up-curved, with ventrally heavily setose

distal half, basally fused with basal section of sacculus.

Anellus with two short, triangular, swollen protrusions

dorsally. Phallus somewhat down-curved medially, dilat-

ed distally, with elongate and apically rounded coecum

ca. twice narrower than main tube. Vesica sack-like with

short membranous basal diverticulum, short scobinated

subbasal diverticulum, two short but broad membranous

lateral diverticula and short granulated distal diverticu-

lum bearing sclerotised dentate patch. Vesica ejaculatori-

us originates subbasally and directed laterally.

Female genitalia (Figs 13-14). Papilla analis trap-

ezoid with rounded corners, weakly setose. Apophyses

thin, apophysis anterioris ca. twice shorter and somewhat

thinner than apophysis posterioris. Osttum bursae posi-

tioned ventro-laterally, surrounded by asymmetrical scle-

rotised fold ventrally and laterally. Ductus bursae short,

dorso-ventrally flattened and weakly sclerotised. Corpus

bursae sack-like with weakly gelatinous posterior section

and membranous anterior section bearing elongate and

narrow Serrulate signum laterally. Appendix bursae short

but broad, conical, apically rounded, membranous, posi-

tioned postero-laterally on right side.

Etymology

The genus name is an aggregation of the word ‘juxta’ and

the generic group name Ei/ema, referring to the extreme-

ly elongated juxta of the type species.

©ZFMK

380 Anton V. Volynkin

Juxtilema smithi gen. & sp. n., HT Oedipygilema aposema

NW Zambia, West Lunga NP, Lukwakwa, slide AV6466 S Zambia, Chingola, slide AV6461

Lophilema mbulu, HT Lophilema polioplaga, HT

Tanzania, Mbulu Mts., slide AV3189 Kenya, slide BMNH Arctiidae 756

Figs 9-12. Lithosiina spp, male genitalia, depositories of the specimens. 9-11. ANHRT. 12. NHMUK (©The Trustees of NHMUK).

Juxtilema gen. nov. for a new species from Zambia 381

13 14

Juxtilema smithi gen. & sp. n., PT Juxtilema smithi gen. & sp. n., PT

NW Zambia, West Lunga NP, Lukwakwa, slide AV6458 C Zambia, Mutinondo Wilderness Area, slide AV6456

5 16 17

Oedipygilema aposema Lophilema mbulu, PT Lophilema polioplaga

NW Zambia, West Lunga NP, slide AV6462 Tanzania, Mbulu Mts, slide AV4585 Tanzania, slide AV3373

Figs 13-17. Lithosiina spp., female genitalia, specimens are deposited in ANHRT.

382 Anton V. Volynkin

Juxtilema smithi gen. et sp. nov.

urn: lsid:zoobank.org:act:A5C20E13-2F0B-49C9-A 1 7A-OSCSAF7832F7

(Figs 1-2, 9, 13-14)

Type material

Holotype (Figs 1, 9). 6, “Zambia, 1147m, Lukwak-

wa, West Lunga NP. (Cryptosepalum forest/Dambo)

12°39°40”S, 24°26713”E, 9-15.xi.2018 LepiLED Light

Trap, Aristophanous, M., Derozier, V., Laszlo, G., Oram,

D. Leg. ANHRT:2018.40” / “ANHRTUK 00074156” /

“Slide AV6466¢ A. Volynkin” (ANHRT).

Paratypes. ZAMBIA: 23 33, 3 99, the same data

as in the holotype, MV, LepiLED & Actinic light traps,

unique numbers: ANHRTUK 00056731, 00057112,

00057113, 00057130, 00057131, 00058739, 00058740,

00058744, 00059195, 00059382, 00061633—00061637,

00061639, 00073763, 00073783, 00074155, 00074157—

00074159, 00074189-00074192, gen. prep. Nos.:

AV6421, AV6467 (3:3), AV6458 (9): 1 J, 3 PY, the

same locality but 4—8.x1.2013, Light Trap, Smith, Takano

& Oram leg., unique numbers: ANHRTUK 00201092—

00201095: 5 ¢¢, 9 22, 1400 m, Hillwood, Ikelenge

(Miombo / Riverine forest mosaic), 11°16’02” S,

24°18°59” E, 23-30.x1.2019, LepiLED & Actinic light

traps, Bashford, M., Miles, W., Mulvaney, L., Smith, R.

leg., unique numbers: ANHRTUK 0010761 7—00107620,

00132358, 00202054—00202057, 00202059, 00202061,

00202119, 00202120, 00204138, gen. prep. No.: AV6459

(2); 1 @, the same locality but 7—10.xii.2019, Bash-

ford, M., Miles, W., Mulvaney, L. leg., unique number:

00132465, gen. prep. No.: AV6468; 3 3.3’, 3 29, 1340 m,

Jiwundu Swamp, 11°51°54” S, 25°33’20” E, 21-24.

x1.2014, Light Trap, Smith, R. & Takano, H. leg., unique

numbers: ANHRTUK 00010320, 00010341, 00010379,

00057210, 00201096, 00201097; 1 9, 1346 m, Kambishi

School, 11°54’42” S, 25°28’50” E, 10—13.x1.2017, MV

light trap, Carter, M., Lloyd, A., Miles, W., Oram, D.,

Smith, R. leg., unique number: ANHRTUK 00118606;

1 9, 1179 m, Greystone, Kitwe, Copperbelt Province,

12°55’°50” S, 28°14’29” E, 19-20.x.2013, Light Trap,

Smith, R. & Takano, H. leg., unique number: ANHR-

TUK 00201080, gen. prep. No.: AV6473; 18 3, 1 9,

1460m, Mutinondo Wilderness Area, Mpika, Northern

Prov., 12°27°06” S, 31°17°30” E, 14-17.11.2019, MV

Light Trap, Dérozier, V., Mulvaney, L., Takano, H. leg.,

unique numbers: ANHRTUK 00139538, 00139539,

00139628, 00139629, 00139671, 00191201, 00192182—

00192188, 00192190-00192192, 00219813, 00219814,

00219828, gen. prep. Nos.: AV6427, AV6469, AV6470

(33), AV6456 (2) (all in ANHRT).

Bonn zoological Bulletin 70 (2): 377-382

Diagnosis

See the diagnosis for the genus.

Description

See the description for the genus.

Distribution

The new species is known from Zambia.

Etymology

The new species is dedicated to Mr Richard Smith,

founder and director of the African Natural History Re-

search Trust, who, through organising and undertaking

numerous entomological expeditions to Sub-Saharan

Africa has enabled much of the recent advances in our

taxonomic understanding of the Afrotropical Lithosiina.

Acknowledgements. The author expresses his sincere thanks

to Dr Alberto Zilli and Mr Geoff Martin (NHMUK) for their

kind assistance during the visits to their institution. The follow-

ing collaborative partners are thanked for the diverse adminis-

trative and technical assistance provided during the field work:

Ms Rhoda Kachali (Department of National Parks and Wildlife

— ZAWA, Lusaka), Ms Claire Mateke and Ms Martha Imak-

ando (Livingstone Museum, Livingstone). The author declares

that to the best of his knowledge he conforms to the national

regulations and meets with the conditions and requirements of

International Conventions concerning collecting/export and

handling of the specimens presented in this Article.

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