LIB Leibniz Institute for the Analysis

== of Biodiversity Change

Bonn zoological Bulletin 71 (2): 87-97

2022 Jiaranaisakul K. & Constant J.

museum

: KOENIG

https://do1.org/10.20363/BZB-2022.71.2.087

ISSN 2190-7307

http://www.zoologicalbulletin.de

Research article

urn:|sid:zoobank.org: pub: 58EF7D30-DF39-402E-B497-94D6FD2D2C5D

New records of Penthicodes \anternfly species from Thailand and Malaysia

and nomenclatural notes on the genus

(Hemiptera: Fulgoromorpha: Fulgoridae)

Kawin Jiaranaisakul ©, & JérOéme Constant©?

'Rabbit in the Moon Foundation, 399, Village No. 3, Suan Phueng, Ratchaburi, 70180, Thailand

‘Department of Entomology, Faculty of Agriculture at Kamphaeng Saen, Kasetsart University,

Kamphaeng Saen Campus, Nakhon Pathom, 73140, Thailand

?Royal Belgian Institute of Natural Sciences, O.D. Taxonomy & Phylogeny — Entomology, Vautier Street 29,

B-1000 Brussels, Belgium

“Corresponding author: Email: kawin2127@gmail.com

'urn:Isid:zoobank.org:author: E70 BBC6F-D963-4EA9-9109-933DD4C83CC8

*urn:|sid:zoobank.org:author:6E6072A 1 -9415-4C8D-8E60-2504444DB290

Abstract. Penthicodes (Penthicodes) farinosus (Weber, 1801), P. (Ereosoma) bimaculatus (Schmidt, 1905) and P. (Er-

eosoma) caja malayanus Constant, 2010 are formally recorded from Thailand for the first time; the second one is also

recorded from Peninsular Malaysia for the first time, based on photographs. Updated distribution maps and the first host

plant records are provided for these species. The genus name Penthicodes Blanchard, 1845 must be treated as masculine

in gender according to the International Code of Zoological Nomenclature and an updated list of the species-group names

in Penthicodes is provided accordingly.

Key words. Aphaeninae, Fulgoroidea, new record, food plant, Indochina, Sunda.

INTRODUCTION

The lanternfly genus Penthicodes Blanchard, 1845 be-

longs to the subfamily Aphaeninae with Penthicodes

(Penthicodes) farinosus (Weber, 1801) as the type spe-

cies. Members of the genus are widely distributed in

Southeast Asia (Constant 2010; Bourgoin 2022). Al-

though the article 30.1.4.4 of the International Code of

Zoological Nomenclature (ICZN 1999) states that com-

pound genus-group names ending tn the suffix -odes are

to be treated as masculine unless originally otherwise

stated, Penthicodes was continuously treated as fem-

inine in all previous studies (Metcalf 1947; Bourgoin

2022). Currently, 12 species are recognized in the genus,

among which ten species belong to the subgenus Erveo-

soma Kirkaldy, 1906 and two species to the subgenus

Penthicodes Blanchard, 1845 (Constant 2010; Bourgo-

in 2022). The species of the subgenus Ereosoma were

reviewed by Constant (2010) who also provided an il-

lustrated identification key to all species of the genus

Penthicodes. Recently, five species of the genus were

recorded from Thailand: P. (Ereosoma) atomaria (Weber,

1801), P. (Ereosoma) caja (Walker, 1851), P. (Ereosoma)

pulchellus Guérin-Méneville, 1838, P. (Ereosoma) varie-

gatus Guérin-Méneville, 1829, and P (Ereosoma) war-

leti Constant, 2010 (Constant 2010; Jiaranaisakul et al.

Received: 10.01.2022

Accepted: 01.09.2022

2018). A sixth species, Penthicodes (Ereosoma) bimac-

ulatus (Schmidt, 1905) was mentioned from Thailand by

Hutacharern et al. (2007) but this record was not substan-

tiated by a reference or voucher specimen data. It was

probably based on a misidentification of P (Ereosoma)

variegatus, a species closely resembling P. quadrimacu-

latus Lallemand, 1963, with the latter being erroneously

treated by Nagai & Porion (1996) as a junior synonym of

P. (Ereosoma) bimaculatus (Constant 2010). Hence the

record of Hutacharern et al. (2007) was regarded as un-

reliable (Jiaranaisakul et al. 2018). All species so far re-

corded from Thailand belong to the subgenus Ereosoma.

Recent surveys of the lanternfly fauna in Thailand led to

the discovery of specimens of P. (Ereosoma) bimaculatus

and P. (Penthicodes) farinosus, providing the first record

in Thailand for these species, while photographic records

additionally provided the first data of the former species

in Peninsular Malaysia. Host plants are also recorded for

the first time for these two species in Thailand, with an

additional host plant record from Borneo for P. (Penthi-

codes) farinosus. In the present paper, we provide the first

substantiated records of P. (Ereosoma) bimaculatus and

P. (Penthicodes) farinosus in Thailand, the first records

from Malaysia for the former species, and the first host

plant data for both species as well as distribution maps

and diagnostic characters to recognize both species. The

Corresponding editor: R. Peters

Published: 11.09.2022

88 Kawin Jiaranaisakul & Jér6me Constant

distribution of these two species 1s discussed. An updated

list of the species names in the genus 1s also provided.

MATERIAL AND METHODS

The male genitalia were removed from the tip of the ab-

domen, then soaked in 10% potassium hydroxide solu-

tion (KOH) for one day, and put in glycerin for detailed

observation. The pygofer was separated from the abdo-

men and the aedeagus dissected with a needle blade for

examination, then stored in glycerin for preservation.

Photographs were taken with a Canon EF 100 mm f/2.8

Macro USM lens attached to a Canon EOS 7D mark II

digital camera. The species were identified using the il-

lustrated key in Constant (2010). The distribution maps

were produced with SimpleMappr (Shorthouse 2010).

Institutional abbreviations

LTRS = = Lam Takhong Research Station, Nakhon

Ratchasima, Thailand

THNHM = Thailand Natural History Museum, Pathum

Thani, Thailand

RESULTS

Taxonomy

Family Fulgoridae Dumeril, 1820

Subfamily Aphaeninae Blanchard, 1847

Tribe Aphaenini Blanchard, 1847

Genus Penthicodes Blanchard, 1845

Blanchard (1845) did not give the gender of the genus

name in the original description nor did he associate it

to a species name from which the gender could be ex-

trapolated. According to the International Code of Zoo-

Fig. 1. Illustration of Lystra dimidiata in Hope (1843).

Bonn zoological Bulletin 71 (2): 87-97

logical Nomenclature (ICZN 1999), article 30.1.4.4 “A

compound genus-group name ending in the suffix -ifes,

-oides, -ides, -odes, or -istes 1s to be treated as masculine

unless its author, when establishing the name, stated that

it had another gender or treated it as such by combining it

with an adjectival species-group name in another gender

form’. Hence, the correct names of the 17 taxa in Penthi-

codes are as follows:

Penthicodes (Ereosoma) Kirkaldy, 1906

Penthicodes (Ereosoma) astraea (Stal, 1864)

Penthicodes (Ereosoma) atomaria (Weber, 1801)

Penthicodes (Ereosoma) bimaculatus (Schmidt, 1905)

Penthicodes (Ereosoma) caja (Walker, 1851)

Penthicodes (Ereosoma) caja caja (Walker, 1851)

Penthicodes (Ereosoma) caja malayanus Constant,

2010

Penthicodes (Ereosoma) celebicus Constant, 2010

Penthicodes (Ereosoma) pulchellus

Guerin-Méneville, 1838

Penthicodes (Ereosoma) quadrimaculatus

Lallemand, 1963

Penthicodes (Ereosoma) rugulosus (Stal, 1870)

Penthicodes (Ereosoma) variegatus

Guerin-Méneville, 1829

Penthicodes (Ereosoma) warleti Constant, 2010

Penthicodes (Penthicodes) Blanchard, 1845

Penthicodes (Penthicodes) farinosus (Weber, 1801)

Penthicodes (Penthicodes) farinosus aerugineus

(Stal, 1870)

Penthicodes (Penthicodes) farinosus farinosus

(Weber, 1801)

Penthicodes (Penthicodes) farinosus leucosticticus

(White, 1845)

Penthicodes (Penthicodes) farinosus niasensis

Schmidt, 1923

Penthicodes (Penthicodes) farinosus tullia

(Breddin, 1901)

Penthicodes (Penthicodes) nicobaricus (Stal, 1869)

Notes. (1) The species-group epithets astraea, atomaria,

caja and tullia are not adjectives and therefore indeclin-

able (D. Yanega pers. com., Sept. 2021).

(2) Hope (1843) described Lystra dimidiata Hope,

1843 from “Silhet” (currently Sylhet in Bangladesh,

24°54'"N 91°52'E), which was later synonymized under

Penthicodes (Penthicodes) farinosus by Nagai & Pori-

on (1996). However, the specimen illustrated by Hope

(1843) (Fig. 1) actually seems to be a specimen of Pen-

thicodes (Penthicodes) farinosus leucosticticus (White,

1845) (see illustration in Nagai & Porion 1996: pl. 4,

fig. 61). The latter subspecies is well characterized by its

hind wings pale blue basally with numerous small black

spots arranged in rows and is only known from the Phil-

ippines. There is no record of any P. farinosus specimen

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New records of Penthicodes from Thailand and Malaysia with notes on the genus 89

Fig. 2. Penthicodes (Ereosoma) bimaculatus (Schmidt, 1905), @. A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus,

lateral view. D. Head and thorax. E. Perpendicular view of frons.

Bonn zoological Bulletin 71 (2): 87-97 ©LIB

90 Kawin Jiaranaisakul & Jér6me Constant

Fig. 3. Penthicodes (Ereosoma) caja malayanus Constant, 2010, 9. A. On unidentified plant. B. Showing the disc of hind wing.

matching Hope’s illustration from Sunda region or conti-

nental Southeast Asia, which represent the area between

the Philippines and Bangladesh (Fig. 4; Nagai & Porion

1996: pl. 4, figs 62, 64, 67, 70); instead, all specimens

examined from Sunda and the continent, lack the rows of

black spots on the hind wings. Hence, we consider that

the specimen collection data given by Hope is probably

erroneous, and propose to remove P. (Penthicodes) fari-

nosus from the fauna of Bangladesh.

(3) The subspecies given for P. (Penthicodes) farinosus

follow Nagai & Porion (1996).

Penthicodes (Ereosoma) bimaculatus (Schmidt, 1905)

(Figs 2, SA—D, 6A—C, 7A, 9A)

Diagnosis. The external morphology of the species is

close to P. (Ereosoma) celebicus and P. (Ereosoma) ru-

gulosus that are known from Indonesia (Sulawesi) and

Philippines, respectively. They share the characters of

a white patch along the sutural margin on nodal line of

cross-veins and irregular black-brown markings on teg-

mina, but the disc of the hind wing is red with orange

apically in P. (Ereosoma) bimculatus. The male genitalia

of this species differ from other species by these charac-

ters: anal tube short with lateral curved expansion direct-

ed ventrally (Fig. 5B, D); gonostyli slightly elongated,

round at apex in lateral view.

Material examined. Thailand: 2 @9, Yala

Prov., Betong Dist., 24.VIII 2018, M. Naiduang-

chan & P. Pawangkhanant leg., THNHM-I-24962

Bonn zoological Bulletin 71 (2): 87-97

and THNHM-I-24963 (THNHM); 1 ©, Yala Prov.,

Betong Dist., 3.VIII 2020, P. Pawangkhanant leg.,

THNHM-I-24961 (THNHM).

Material examined from photographs. Ma-

laysia: 1 specimen (Fig. 6A), Selangor, Semenyih,

2.9474° N, 101.8451° E, 24.V.2020, T. Eng Wah; 1

specimen (Fig. 6B), Selangor, Sungai Tua, 3.25233° N,

101.67538° E, 10.XII.2016, T. Eng Wah; 1 specimen

(Fig. 6C), Selangor, Ulu Yam, 3°25'59" N, 101°39'27" E,

9.V1.2012, H.P. Guek.

Distribution. Borneo, Indonesia (Sumatra), West Ma-

laysia (new country record), Singapore and Thailand

(Constant 2010; Bourgoin 2022).

Biology. Specimens (male and female) of P. (Ereoso-

ma) bimaculatus have been confirmed from Thailand and

Peninsular Malaysia. This species is confined to Sunda-

land. So far, the northern limit in its distribution range

is in the southernmost part of Thailand. The male spec-

imen was collected from southern Thailand on a trunk

of Shorea cf. curtisii (Dipterocarpaceae) together with a

cockroach (Fig. 7A).

Penthicodes (Ereosoma) caja malayanus Constant,

2010

(Figs 3, 8)

Diagnosis. The species differs from other species by its

tegmina with large dark-brown markings, ground colour

of tegmina and membrane brown; white patch along su-

tural margin and costal margin on nodal line of cross-

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New records of Penthicodes from Thailand and Malaysia with notes on the genus 91

Fig. 4. Penthicodes (Penthicodes) farinosus (Weber, 1801), &. A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, lateral

view. D. Head and thorax. E. Perpendicular view of frons.

Bonn zoological Bulletin 71 (2): 87-97 ©LIB

92 Kawin Jiaranaisakul & Jér6me Constant

Fig. 5. Male genitalia. A-D. Penthicodes (Ereosoma) bimaculatus (Schmidt, 1905) (modified from Constant 2010). A. Pygofer

(Py), anal tube (An) and gonostyli (G), left lateral view. B. Anal tube, dorsal view. C. Pygofer and gonostyli, posteroventral view.

D. Anal tube, posterior view. E-H. Penthicodes (Penthicodes) farinosus (Weber, 1801). E. Pygofer, anal tube and gonostyli, left

lateral view. F. Anal tube, posterior view. G. Pygofer and anal tube, dorsal view. H. Pygofer and gonostyli, posteroventral view.

veins. The subspecies malayanus is is very easy to rec-

ognize from P. caja caja by the disc of hind wings red as

opposed to orange in P. caja caja.

Material examined. Thailand: 1 9, Nakhon Ratchasi-

ma Prov., Mu Si, 21.[II.2020, W. Khaikaew leg. (LTRS).

Distribution. Malaysia (Constant 2010) and Thailand

(new country record).

Biology. The specimen was found on an unidentified

plant in dry evergreen forest.

Penthicodes (Penthicodes) farinosus (Weber, 1801)

(Figs 4, 5E-H, 6D, 7B—D, 9B)

Diagnosis. The species is easy to differentiate from the

other Penthicodes species by the vertex wider than long

Bonn zoological Bulletin 71 (2): 87-97

in dorsal view with two patches of white waxy secretion.

Only P. (Penthicodes) nicobaricus (Stal, 1869) from An-

daman and Nicobar Islands shows waxy patches on the

vertex but its vertex 1s as wide as long in dorsal view

(Constant 2010; Constant & Mohan 2017).

Material examined. Thailand: 3 ¢¢, 2 99,

Yala Prov., Betong Dist., 3.VHI.2020, A. Aksorn-

neam, P. Pawangkhanant & T. Ruangsuwan leg.,

THNHM-I-24964-THNHM-I-24968 (THNHM); 1 8,

Phang Nga Prov., Thai Mueang Dist., Khanim Waterfall,

8°29°48.7” N, 98°17°01.3” E, 50 ma.s.1., 20.X.2021, K.

Jiaranaisakul leg. (THNHM).

Material examined from photographs. Malaysia

(Borneo): 1 specimen (Fig. 6D), Sarawak, Mulu National

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New records of Penthicodes from Thailand and Malaysia with notes on the genus 93

~—

soma) bimaculatus, Selangor, Ulu Yam, 9.V1.2012. © H.P. Guek. D. Penthicodes (Penthicodes) farinosus (Weber, 1801), Sarawak,

Mulu National Park, 27.II 2009, on Ailanthus integrifolia Lam. (Simaroubaceae). © Fletcher & Baylis.

Bonn zoological Bulletin 71 (2): 87-97 ©LIB

94 Kawin Jiaranaisakul & Jér6me Constant

Fig. 7. Live specimens from Thailand. A. Penthicodes (Ereosoma) bimaculatus (Schmidt, 1905), Yala, Betong, 3. VII 2020, tended

by cockroach. © P. Pawangkhanant. B. P. (Penthicodes) farinosus (Weber, 1801), Yala, Betong, 3. VIII 2020, on A/stonia scholaris

(L.) R.Br. (Apocynaceae). © P. Pawangkhanant. C. P. (Penthicodes) farinosus, Yala, Betong, 3. VIII 2020, showing hind wings.

Bonn zoological Bulletin 71 (2): 87-97 ©LIB

New records of Penthicodes from Thailand and Malaysia with notes on the genus

des caja malayanus

ad Penth

ICO

‘

4 7...

be, “A

"bres, he

Fig. 8. Distribution map of Penthicodes (Ereosoma) caja malayanus Constant, 2010.

©LIB

Bonn zoological Bulletin 71 (2): 87-97

96 Kawin Jiaranaisakul & Jér6me Constant

% haba es Akon *

SQLS

Say

160 240 km

Oo _Penthicodes farinosus

Sal, SN

NES My, 2

160 240 km

Fig. 9. Distribution map of Penthicodes Blanchard, 1845 spp. in the Malay Peninsula. A. Penthicodes (Ereosoma) bimaculatus

(Schmidt, 1905). B. Penthicodes (Penthicodes) farinosus (Weber, 1801).

Park, 27.11.2009, on Ailanthus integrifolia Lam. (Sima-

roubaceae), W.K. Fletcher & D.M. Baylis.

Thailand: 1 specimen (Fig. 7D), Yala Province, Beto-

ng, 31.XII.2020, K. Saechan.

Distribution. Borneo, Indonesia (Java, Sumatra), West

Malaysia, Myanmar, Philippines (Distant 1901; Lalle-

mand 1963; Seidel & Wessel 2013; Bourgoin 2022) and

Thailand (new country record).

Note. The record from Tavoy (currently Dawei, Myan-

mar), far north of the Isthmus of Kra, by Distant (1906)

is regarded as doubtful and further studies are needed for

confirmation.

Biology. Penthicodes (Penthicodes) farinosus has been

found only in the southernmost part of Thailand. A/sto-

nia scholaris (L.) R.Br. (Apocynaceae) is a host plant of

P. (Penthicodes) farinosus in Thailand (Fig. 7B), while

the species was recorded on Ailanthus integrifolia Lam.

in Borneo (W.K. Fletcher & D.M. Baylis pers. com.,

2020; Fig. 6D).

Bonn zoological Bulletin 71 (2): 87-97

DISCUSSION

Three species and one subspecies of the genus Penthi-

codes were recently added to the fauna of Thailand, as

well as several new host plant records (Jiaranaisakul

et al. 2018; Jiaranaisakul & Constant 2021; present

study). Additionally, the nomenclature of the genus had

to be adapted to a correct masculine treatment of the ge-

nus instead of the feminine treatment erroneously applied

by all authors for more than 150 years.

Seven species of the genus Penthicodes are now re-

corded from Thailand, representing 58.33% of the to-

tal number of species in the genus. Unfortunately, their

life-history and biology remain still very poorly docu-

mented, for example, eggs and nymphs are still unknown.

We hope that cooperation with citizen-scientists and lo-

cal authorities will help fill those gaps in the knowledge

of this iconic insect group.

Acknowledgements. We thank Akrachai Aksornneam (Thai-

land), Mali Naiduangchan (Thailand), Parinya Pawangkhanant

(Thailand), Thiti Ruangsuwan (Thailand), Dr Teo Eng Wah

©LIB

New records of Penthicodes from Thailand and Malaysia with notes on the genus 97

(Vincent) (Malaysia), W.K. Fletcher & D.M. Baylis (Malaysia)

and Hock Ping Guek (Malaysia) for their valuable specimens,

photographs of live specimens and habitat and data; Mado

Berthet (Royal Belgian Institute of Natural Sciences) for her

help with the male genitalia drawings of P. (Penthicodes) fa-

rinosus, Prof. Thierry Bourgoin (MNHN) for his comments.

The first author would like to thank Dr. Weeyawat Jaitrong

(THNHM) for his careful reading and useful correction of an

early version of the manuscript; Wuttikrai Khaikaew and the

staff of Lam Takhong Research Station (LTRS) for providing

a specimen of P. (Ereosoma) caja malayanus. A very special

thanks is due to Dr Doug Yanega (University of California Riv-

erside, California, USA) for attracting our attention on the gen-

der issue in the treatment of Penthicodes.

REFERENCES

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ridae). Belgian Journal of Entomology 49: 1-24

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BHL

Blank Page Digitally Inserted

LIB Leibniz Institute for the Analysis

=— of Biodiversity Change

Bonn zoological Bulletin 71 (2): 99-103

2022 Dinh T'S. et al.

forsewunas

museum

: KOENIG

ISSN 2190-7307

http://www.zoologicalbulletin.de

https://do1.org/10.20363/BZB-2022.71.2.099

Scientific note

urn:|sid:zoobank.org:pub: BSA8E7E9-48 1 1-480C-8C 13-2239ECB31CE5

New country record of 7rimerodytes yapingi (Guo, Zhu & Liu, 2019)

(Squamata: Natricidae) from Laos

with the first description of a male specimen and expanded diagnosis

Tuong Sy Dinh’, Vilay Phimpasone’, Hong Bich Ha’, Truong Quang Nguyen‘, Thomas Ziegler’ &

Vinh Quang Luu®

'26 Faculty of Forest Resources and Environmental Management, Vietnam National University of Forestry,

Xuan Mai, Chuong My, Hanoi, Vietnam

?Faculty of Forestry, National University of Laos, Dong Dok Campus, Vientiane, Lao PDR

>College of Forestry Biotechnology, Vietnam National University of Forestry, Hanoi, Vietnam

* Institute of Ecology and Biological Resources, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet,

Hanoi 10072, Vietnam

* Graduate University of Science and Technology, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet, Cau Giay,

Hanoi 10072, Vietnam

°AG Zoologischer Garten K6ln, Riehler Strasse 173, D-50735 Cologne, Germany

° Institute of Zoology, University of Cologne, Ziilpicher Strasse 47b, D-50674 Cologne, Germany

“Corresponding author: Email: vinhlq@vnuf.edu.vn

'urn:lsid:zoobank.org: author: A0589B58-E392-4796-860 1-3 17BOB555EAB

2 urn:Isid:zoobank.org:author:333E61C1-B43B-44BF-BFB4-BC8FE2F56C09

3urn:lsid:zoobank.org:author:6827C355-EB28-4699-94 1 1-65A D02ED3030

*urn:|sid:zoobank.org:author: 22872A6-1C40-461F-AAOB-6A 20EE0O6A DBA

>urn:|sid:zoobank.org:author:57 16DB92-5FF8-4776-ACC5-BF6FA8C2E1BB

Surn:|sid:zoobank.org:author:052B40C7-8AFB-43A5-AF22-7 13DA7B5BBDD

Abstract. Trimerodytes yapingi (Guo, Zhu & Liu, 2019), a species previously known only from Yunnan Province, China,

is reported for the first time from Laos based on two specimens from Houaphan Province. The newly collected specimens

(a couple) from Laos slightly differ from the female holotype from China in body size, number of ventrals and subcaudals

as well as by a low genetic divergence of 1.1% (Cytb gene). In addition, we provide the first description of a male speci-

men and an expanded diagnosis of the species.

Key words. New record, genetic divergence, 7rimerodytes, Houaphan Province.

INTRODUCTION

Sinonatrix yapingi Guo, Zhu, & Liu, 2019 was original-

ly described from Yunnan Province, China, by Guo et al.

(2019) based on the adult female holotype. Subsequently,

this species was placed in the genus 7rimerodytes Cope,

1895 by Ren et al. (2019). The species is characterized

by having dorsal scales in 19-19-17 rows, moderately

keeled except four outermost rows; prefrontal single;

ventral scales 149; subcaudals 55, paired; cloacal plate

divided; body with 30 black bands; ventral surface cream;

and subcaudal surface gray (Guo et al. 2019). As a re-

sult of our field work in 2020, two individuals of a water

snake were found in Houaphan Province, Laos. Morpho-

logical examination and genetic analysis revealed these

snakes to be conspecific with 7’ yapingi Guo, Zhu, & Liu,

2019, a species previously known only from China. Thus,

Received: 02.06.2022

Accepted: 01.09.2022

we herein report the first country record of this poorly

known species for Laos and provide the first description

of a male specimen as well as an expanded morphologi-

cal diagnosis. Field survey was conducted in Phong Song

Village, Xon District, Houaphan Province, Laos in March

2020.

MATERIAL AND METHODS

The specimens were anaesthetized and euthanized with

ethyl acetate, subsequently fixed in approximately 85%

ethanol, and then transferred to 70% ethanol for perma-

nent storage in the collections of the Vietnam National

University of Forestry (VNUF), Hanoi, Vietnam and the

National University of Laos (NUOL). A tissue sample

was preserved separately in 95% ethanol.

Corresponding editor: W. Bohme

Published: 14.09.2022

100

Tuong Sy Dinh et al.

Measurement and meristic character abbreviations

SVL =

TaL

tail length

Head scales

InN

SuprAo

Lor:

PreOc

PostOc

Atem

Ptem

SL/orbit

IL/ 1% chin shield

Body scales

DSR

KI/Sm =

Ven

total length

snout-vent length

internasal

prefrontal

frontal

supraocular

loreal scale

preocular scale

postocular scales

anterior temporal scales

posterior temporal scales

supralabial scales

scale order of supralabials

touching the orbit

infralabial scales

number of infralabials touching

the first chin shield

dorsal scale row

keeled or smooth

ventral scales

precloacal scale

subcaudal scales

99 |

100

Total genomic DNA was extracted from liver samples

using the animal DNA isolation Kit (QlAamp DNA Mini

Kit, Germany). The total DNA purity and integrity were

tested by Scandrop 3830A-0341 spectrophotometer (An-

alytik Jena, Germany) and then diluted to a concentra-

tion of 20 ng/ul. The mitochondrial cytochrome b (Cytb)

gene region was amplified using primers L14919/

H16064 (Guo et al. 2012). All double-stranded products

were sequenced by 1*t BASE (Malaysia), and then edit-

ed manually using Bioedit ver. 7.0.5.2 (Hall 1999). The

new sequences were added to a dataset including 11 cytb

sequences from seven species of 7rimerodytes for the

subsequent analyses. Natrix natrix Linnaeus, 1758 was

selected as outgroup. Phylogenetic trees were performed

using maximum likelihood (ML) on MEGA ver. 7.0 (Ku-

mar et al. 2015) software with 1000 bootstrap replicates.

Genetic distances among species were calculated using

MEGA ver. 7.0 (Kumar et al. 2015). The Cytb nucleotide

sequence matrix contained 988 characters without inser-

tions or deletions. The Maximum Likelihood analysis

produced identical topologies (Fig. 1). The two newly

collected specimens have the same Cytb nucleotide se-

quence (GenBank accession numbers: ON603619-20).

In the phylogenetic tree, all samples of the genus 7ri-

merodytes clustered in a monophyletic group, and can

be divided into four clades (A—D). Trimerodytes per-

carinatus (Boulenger, 1899) formed a mono-typic clade

(clade C), this clade was supported by a bootstrap val-

ue of 100. Trimerodytes annularis (Hallowell, 1856)

and 7: balteatus (Cope, 1895) were grouped in clade D

(Bootstrap 76). Clade B included 7rimerodytes yunnan-

Trimerodites annularis (JQ687437 1)

Trnmerodytes annularis (MNS824 70.7)

Tnmerodytes balteatus (MNOT? 775.7)

Timerodytes percarinatus (M N58 2500. 7)

Timerodytes percarinatus (MN58 2457.7)

Timerodytesyunnanens's (IMN582 458.7}

Trimerodites yunnanensis (MN582 467.7)

Tnmerodytes aequifasciatus (JQ68 7430. 7)

Trimerodytes aequ fasciatus (M N53 2462.1)

100

Timerodytes praemaxillans (OK315847.7)

Trimerodyte s yaping! (MK627 916.7)

| Trimerodytes yapingi (VNUF R.2020.14)

97 | Trimerodytes yapingi (NUOL R.2020.15)

D 76

100 |

B |

100

om0

Natnx natnx (LL999947)

Fig. 1. Maximum Likelihood phylogeny of the genus 7rimerodytes Cope, 1895. The numbers at the branches are confidence values

based on Felsenstein’s bootstrap method (based on 1000 bootstrap replicates).

Bonn zoological Bulletin 71 (2): 99-103

©LIB

New country record of 7rimerodytes yapingi from Laos 101

Fig. 2. Trimerodytes yapingi (Guo, Zhu & Liu, 2019) (NUOL R.2020.15), adult male (preserved) from Laos. A. Dorsal view.

B. Ventral view. Photos: Tuong S. Dinh.

ensis Rao & Yang, 1998 and T. aequifasciatus (Barbour,

1908) with a bootstrap value of 72. Within clade A, Trim-

erodytes praemaxillaris Angel, 1929 and T: yapingi Go,

Vietnam

~ Sen

Fig. 3. Distribution of 7rimerodytes yapingi (Guo, Zhu & Liu,

2019). 1 = type locality in Yunnan Province, China; 2 = new

record from Houaphan Province, Laos.

Bonn zoological Bulletin 71 (2): 99-103

Zhu & Liu, 2019 were placed together with a bootstrap

value of 66. The two new samples from Laos were placed

with 7. yapingi from China with a high bootstrap value of

100, and a small genetic distance (1.1%).

Redescription of Trimerodytes yapingi (Guo, Zhu &

Liu, 2019) with the first record of the male specimen

Jingdong Water Snake (Fig. 2)

Specimens examined (n=2). One adult female, VNUF

R.2020.14 (field number: HP.14) and one adult male,

NUOL R.2020.15 (field number: HP.15), collect-

ed on March 22" 2020 in a stream (20°187142” N/

103°20°322” E, at an elevation of 1,086 ma.l.s.) of Phon

Song Village within Nam Et — Phou Louey National

Protected Area by Vilay Phimpasone and Oanh Van Lo

(Fig. 3). Expanded diagnosis: As the original description

of Trimerodytes yapingi was based on one adult female

only (Guo et al. 2019) we herein expand the diagnosis of

this species as follows: 1) Body size large (TL from 635

to 795 mm); 2) dorsal scales in 19-19-17 rows, mod-

erately keeled except outer two to four scale rows; 3)

prefrontal single; 4) ventral scales 149-156, subcaudals

55-65, paired; 5) cloaca divided; 6) body with 30 black

bands; 7) belly milk-white, without speckles or bands; 8)

reduction of dorsal scale rows from 19 to 17 anterior to

87" ventral scale in the male, and to the 90-97" ventral

scale in females; 9) reduction of caudodorsal scale rows

from 8 to 6 anterior to 20" subcaudal in the male, and

anterior to 12—24" subcaudals in females; as well as from

6 to 4 anterior to 34" subcaudal in the male, and 31-38"

subcaudals in females. Description of the male specimen:

Body stout, cylindrical. SVL 485 mm, TaL 150 mm, TL

©LIB

102 Tuong Sy Dinh et al.

\ : ge

(Cas fg x .

af, Nae ef ee

a a we

AGE Te

we 7 Ny

Fig. 4. Habitat of Trimerodytes yapingi (Guo, Zhu & Liu, 2019) in Phon Song Village, Xon District, Houaphan Province, north-

eastern Laos. Photo: V. Phimpasone.

635 mm, TaL/TL 0.23; HL 20.38 mm, HW 10.43 mm;

elongated and narrow head, longer than twice its width,

snout blunt; internasals much longer than wide, posterior-

ly wider, shorter than prefrontal; prefrontal single, nearly

two times as wide as long, extending downward on both

sides and in contact with loreals, preoculars, and nasals;

frontal shield-like, 1.5 < as long as wide; loreal 1/1,

separated from the eye; preocular 1/1, postoculars 3/3;

anterior temporals 2/2, posterior temporals 3/3; supral-

abials 8/8, fourth touching the eye, sixth largest; infral-

abials 9/10, first to fifth (both sides) in contact with chin

shields; dorsal scales in 19-19-17 rows, 11 rows moder-

ately keeled except two or four outermost scale rows at

each side which are smooth; ventrals 151; cloacal plate

divided; subcaudals 65, paired. Coloration in life: Dorsal

surface of body grayish brown, with many black irreg-

ular bands on vertebral line. There are many V-shaped

speckles on both sides of the body flank, nearby ventral

scales, connecting and constituting W-shaped speckles.

Dorsal surface of head dark gray without marking, ven-

tral head white, infralabials dark anteriorly and white

posteriorly. Ventral side yellowish white, ventral surface

of tail dark white. Coloration in preservative: Dorsal sur-

face dark-brown. Ventral side white cream, ventral sur-

face of tail dark white. Ecological notes: The specimens

were found at 13h10 in a stream within the evergreen

forest at an elevation of 1,086 m a.s.l. The relative tem-

perature was about 27.8°C and the humidity 44%. The

surrounding habitat was mixed evergreen forest of hard-

woods and shrub and vines (Fig. 4). Distribution. This is

the first country record of Trimerodytes yapingi for Laos.

Bonn zoological Bulletin 71 (2): 99-103

Elsewhere, this species is known from Yunnan, China

(Fig. 3). The specimens from Laos slightly differ from

the the original description by having more ventrals (156

versus 149+1), more subcaudals (62—65 versus 55), body

length shorter than reported for the holotype (SVL 485

versus 635 mm; TaL 150-155 mm versus 160 mm). The

first discovered male of the species has more subcaudals

(65) compared with the females (55 and 62). Our record

of Trimerodytes yapingi from Laos is approximately

500 km distant from the type locality in Yunnan, China.

It is probable that the species will also be reported from

northern Vietnam in the future. The new record in Laos

was reported from lower elevation (1,086 m a.s.l.) com-

pared with that in the original description (1,500 maz.s.1.).

Whereas the type specimen was collected in a rice field

near the evergreen forest, the specimens from Laos were

found within disturbed evergreen forest.

Acknowledgements. We thank S. Sitthivong (NUOL, Vien-

tiane), and O.V. Lo (Center for Nature Conservation and Devel-

opment, Hanoi) for their assistance during field work in Houa-

phan Province.

REFERENCES

Angel F (1929) Liste des reptiles et batraciens du Haut-Laos

recueillis par M. Delacour. Descriptions d’un genere de es-

peces et d’une variete d’ophidiens. Bulletin du Muséum Na-

tional d’Histoire Naturelle 2: 75-81

Barbour T (1908) Some new reptiles and amphibians. Bulletin

of the Museum of Comparative Zoology at Harvard College

51 (12): 315-325

©LIB

New country record of 7rimerodytes yapingi from Laos 103

Boulenger GA (1899) On a collection of reptiles and batra-

chians made by Mr. J.D. La Touche in N.W. Fokien, Chi-

na. Proceedings of the Zoological Society of London 1899:

159-172

Cope ED (1895) On a collection of Batrachia and Reptilia from

the island of Hainan. The Proceedings of the Academy of

Natural Sciences of Philadelphia 46: 423-428

Guo P, Liu Q, Xu Y, Jiang K, Hou M, Ding L, Pyron RA, Bur-

brink FT (2012) Out of Asia: Natricine snakes support the

Cenozoic Beringian Dispersal Hypothesis. Molecular Phylo-

genetics and Evolution 63: 825—833

Guo P, Zhu F, Liu Q (2019) A new member of the genus Sinon-

atrix (Serpentes: Colubridae) from western China. Zootaxa

4623 (3): 535-544

Hall TA (1999) BioEdit: A User-Friendly Biological Sequence

Alignment Editor and Analysis Program for Windows 95/98/

NT. Nucleic Acids Symposium Series 41: 95—98

Hallowell E (1857) Notes on the reptiles in the collection of the

museum of the Academy of Natural Sciences. The Proceed-

Bonn zoological Bulletin 71 (2): 99-103

ings of the Academy of Natural Sciences of Philadelphia 8

(4): 146-153

Kumar S, Stecher G, Tamura K (2015) MEGA7: Molecular

Evolutionary Genetics Analysis version 7.0. Molecular Biol-

ogy and Evolution 30: 2725-2729

Linnaeus C (1758) Systema Naturae per regna tria naturae, se-

cundum classes, ordines, genera, species, cum characteribus,

differentiis, synonymis, locis. Editio decima, reformata. Lau-

rentius Salvius, Holmiae

Rao DQ, Yang DT (1998) A new species of Sinonatrix (Ser-

pentes: Colubridae) of China with preliminary survey on Sin-

onatrix. Russian Journal of Herpetology 5 (1): 70-73

Ren JL, Wang K, Guo P, Wang YY, Nguyen TT, Li JT (2019) On

the generic taxonomy of Opisthotropis balteata (Cope,1895)

(Squamata: Colubridae: Natricinae): Taxonomic revision of

two natricine genera. Asian Herpetological Research 10 (2):

105-128

©LIB

BHL

Blank Page Digitally Inserted

LIB. Leibniz Institute for the Analysis

== of Biodiversity Change

Bonn zoological Bulletin 71 (2): 105-108

2022 Drohvalenko M. & Fedorova A.

museum

: KOENIG

https://do1.org/10.20363/BZB-2022.71.2.105

ISSN 2190-7307

http://www.zoologicalbulletin.de

Scientific note

urn:|sid:zoobank.org:pub: AAA BACA0-585E-42B8-8568-42A218E782BB

The first evidence of triploidy among Pelophylax esculentus (Linnaeus, 1758)

(Anura: Ranidae) in the Chornobyl Exclusion Zone

Mykola Drohvalenko! & Anna Fedorova*”

'2 Department of Zoology and Animal Ecology, V. N. Karazin Kharkiv National University, maidan Svobody 4, Kharkiv, Ukraine

“Corresponding author: Email: anna,fedorova@karazin.ua

'urn:Isid:zoobank.org:author: DSCSAACE-2A 0E-4137-B99B-46083135CA73

2 urn:Isid:zoobank.org:author:5 DE735D6-891B-47A B-BB07-60BC76145368

Abstract. Pelophylax esculentus complex hemiclonal systems are a unique evolutionary object due to intricate mecha-

nisms of their sustainability at genomic, gamete, and developmental levels. Chornobyl Exclusion Zone (Ukraine) presents

a no less unique object of nature evolution under the unprecedented radiation impact and human pressure decrease. By

measuring erythrocyte lengths, we report the first findings of triploid hybrid frogs P esculentus from two localities in

Chornoby! Exclusion Zone, where diploid hybrids only were reported before. The presence of triploids implies high com-

plexity of the local water frog population systems, enabling new research opportunities.

Key words. Water frog, hybrid, population system, erythrocyte.

Hybridogenetic species complexes are great models for

studying the evolution of reproduction due to hybrids’

capacity to reproduce themselves. The Pelophylax es-

culentus complex is one of the most studied (Dedukh &

Krasikova 2021). It consists of pool frog Pelophylax

lessonae (Camerano, 1882), marsh frog Pelophylax ridi-

bundus (Pallas, 1771), and their hemiclonal hybrid, edi-

ble frog Pelophylax esculentus (Linnaeus, 1758), whose

range approximately coincides with the range of pool

frogs (Hoffmann et al. 2015). Hybrids that transmit paren-

tal genomes clonally are presented as both sexes and two

ploidies (Tunner 1974; Berger 1977; Plotner 2005; Jakob

2007). Hybrids typically coexist with parental and oth-

er hybrid forms in mixed population systems, leading to

formation of a new hybrid generation. Each such system

has evolved the mechanisms to maintain its composition,

like different ontogenetic strategies and selective survival

(e.g., Berger 1973; Hoffmann et al. 2015; Shabanov et al.

2015). It is caused by the variety of ways different forms

contribute to reproduction, mainly the types of gametes:

not only haploid L and R (from hybrids and parental

species), but also occasionally diploid LL, RR, and LR

gametes from hybrids (Dedukh et al. 2013; Pruvost et al.

2015). The most complicated mechanisms are in the sys-

tems with triploids since they include the highest number

of forms (parental species, 2n and 3n hybrids) balancing

with each other (Pruvost 2013; Mikuliéek et al. 2015). In

Ukraine, such complex systems were known to be wide-

spread in the eastern part only, in Siverskyi Donets river

basin (Borkin et al. 2004; Shabanov et al. 2020).

Chornobyl Exclusion Zone per se possesses many oppor-

Received: 06.03.2022

Accepted: 01.09.2022

tunities in herpetology, at least due to long-term decreased

anthropogenic pressure. Its radioactive contamination

is already known for affecting amphibians (Gashchak

et al. 2009), though some species remain thriving under

its impact (Burraco et al. 2021a; Burraco 2021b). Radi-

ation also affects amphibian populations. For instance,

the changes in local evolution were described for Hyla

orientalis Bedriaga, 1890 in the Zone, though without

obvious harmful consequences (Car et al. 2022). How-

ever, the impact of some registered effects is not so clear.

Water frogs of the P. esculentus complex from the area

of the Chornobyl] fallout impact in Belarus (Briansk re-

gion) were shown to have reduced genome size ( Vinogra-

dov & Chubinishvili 1999). Despite the found influence

on their genomes (Rozanov et al. 1990; Vinogradov et al.

1990), all the previously examined water frogs from the

Zone were diploids. Therefore, it was assumed that only

diploid population systems exist here (S. Litvinchuk, St.

Petersburg, pers. comm.). Herein, we report the evidence

on the previously unrecorded presence of triploid P. escu-

lentus in the Chornobyl Exclusion Zone.

The amphibian survey in the Chornobyl Exclu-

sion Zone was carried out on 9-12 of August 2021 in

collaboration with and legal permission from Chor-

nobyl Radiation and Ecological Biosphere Reserve

(http://zapovidnyk.org.ua) and aimed to explore the

Pelophylax populations within the Zone. The region

of the study included: the Prypiat River (in Chornobyl;

51.272483, 30.244840) and its floodplain (51.341432,

30.199178), the Uzh River (51.273852, 29.741894) and

Corresponding editor: W. Bohme

Published: 15.09.2022

106

its floodplain (51.256711, 30.222522), isolated old me-

lioration channels (51.245596, 30.169778) and those

in Ilia River valley (51.278037, 29.808185), and the

bypass channel of Chornobyl nuclear plant’s cooling

pond (51.396829, 30.141884). The frogs were caught by

hands, dip-net, and using flashlights at night. Species and

sex were identified morphologically. The presence of vo-

cal sacs and nuptial pads indicated males, while species

were identified by body coloration, metatarsal tubercle

shape, and hindlimbs proportions. Juveniles had weakly

expressed species- and sex-specific features so they were

described as a homogenous group. Snout-vent length

(SVL) was measured via scale photographing and later

measured on photographs. The blood samples were taken

from each frog by cutting the fingertips; wounds were

treated with the antibiotic Vetbicilin-3 (Basalt, Ukraine).

We prepared the air-dried blood smears and photographed

and measured ~50 erythrocytes for each frog using a

Leica DFC3000 G camera with Leica LASX Software.

Triploid water frogs have about 1.5-fold more DNA in

nuclei than diploids, so the ploidy of P. esculentus frogs

Mykola Drohvalenko & Anna Fedorova

can be estimated by mean erythrocytes length (Ogielska

et al. 2004; Bondareva et al. 2012). It is also known that

the exact cut-off value of erythrocyte length for triploids

should be separately estimated for a particular population

system since it varies among them (though often about

26 um) (Drohvalenko et al. 2019). Techniques used in

the capture and sampling sought to minimize animal suf-

fering according to Directive 2010/63/EU (protection of

animals used for scientific purposes). All the frogs were

released after the sampling.

The total catch was 52 frogs: 1 tadpole (late devel-

opment stage), 31 juveniles, and 19 adults (10 females,

9 males). Adult hybrids were found in Prypiat and Uzh

floodplains, in Ilia River, old melioration channels, and

bypass channel of cooling pond.

The distribution of collected frogs by SVL and eryth-

rocytes lengths are shown in Fig.1. Adults are visually

divided into two groups without any transient state. The

vast majority (with a mean of 21.54 um, 95% CI [21.06,

22.02]) belong to diploids. Two spike values exceeding

the considered cut-off value putatively mean the triploidy

A Juveniles Adults

ae 3n? ¢

|_|

= Fay =

& & 3n

aca a] I | || 4g? UT a hh a a i id Maasai a

> @ © Juveniles

BA & ® Males

”

a 24

Sy & @ Females

o © 4 Oo

= oe ¢ - e o

22 ©

2 oe? % ® ie) e # a

LU @

20 30 40 50 60 70 80

SVL, mm

B Mean (95% Cl) Median (IQR) Min Max

Juveniles 22.16 (21.39, 22.93) 21.81 (20.94, 22.58) 19.94 27.76

Putatively 2n juveniles 21.48 (21.03, 21.93) | 21.50 (20.59, 21.93) 19.94 23.33

Putatively 3n juveniles 25.87 (23.57, 28.17) 25.70 (25.06, 26.52) 24.35 27.75

Adults 2n 21.54 (21.06, 22.02) 21.69 (20.93, 22.16) 19.38 23.08

Adults 3n 26.86 (24.6, 29.12) 26.86 (26.77, 26.94) 26.68 27.03

Fig. 1. A. The distribution of frogs from the Zone by their mean erythrocyte lengths vs body length (SVL). Red dash line marks

putative cut-off value for adult triploids; dash circle marks juveniles of doubtful ploidy. Boxplots denote means (middle line), IQRs

(boxes) and extreme values (whiskers). B. The sample parameters for different groups of studied frogs.

Bonn zoological Bulletin 71 (2): 105—108

©LIB

The first evidence of triploidy among Pe/ophylax esculentus in the Chornobyl Exclusion Zone 107

of these individuals (male from bypass channel and fe-

male from Prypiat floodplain, with 27.03 and 26.67 um,

respectively). Although the gap between small-cell and

large-cell individuals is obvious, more individuals should

be examined to specify the erythrocyte length ranges for

diploids and triploids, as well as their possible overlap-

ping.

The distribution of erythrocyte sizes for juveniles looks

more intricate. As there is no gap but rather a continuous

set of cell lengths from small to large, we cannot confi-

dently define the cut-off value for juveniles, so we treat-

ed them on the graph as a single group (mean 22.16 um,

95% CI [21.39, 22.93]). It is known for certain systems

(Drohvalenko et al. 2021) that cell length distribution

could vary much between adults and juveniles, widely

overlapping in the last. Worth noting that the juveniles

with the largest cells (dash circle Fig. 1; mean 25.87 um,

95% CI [23.57, 28.17]) originated from the bypass chan-

nel, where the triploid male was assumingly found. We

thus can carefully suggest that these juveniles are a new

generation of triploids. At the same site, there were also

caught three individuals with tails remnants (still in the

course of metamorphosis), which SVL weren’t measured

— they, however, had the smallest erythrocytes: 19.79,

19.71, and 18.80 um. No significant differences were

found between different ages, neither among diploids

(Student’s t-test p=0.354) nor triploids (Mann-Whitney’s

p= 0.533).

Due to the late summer survey, the catch of frogs ap-

peared relatively small and insufficient to claim neither

the exclusive presence of triploids in two localities only

nor the exact composition of the studied systems. Nev-

ertheless, every presence of hybrids P. esculentus prob-

ably means the presence of the parental species, which

they depend on — and triploids presence implies the pres-

ence of diploid hybrids, producing 2n-gametes (Biriuk

et al. 2016). In certain locations, the presence of some

forms could be suggested by habitat preferences (like

P. lessonae in swampy channels; [Pysanets, 2014]) or

behavioral features (like P. ridibundus possibly migrat-

ing to bypass channel from Prypiat at breeding; [Wells,

2007]). The local absence of any form could indeed be

just an artifact.

The features of triploid P. esculentus from the Chor-

nobyl Exclusion Zone are planned to be examined using

precise methods. Their very presence here would mean

the existence of more complicated hemiclonal popula-

tion systems there than was supposed before. At least

the L-E-R system with triploids (in the bypass channel)

is already more complex than the most complex system

studied in Ukraine (Meleshko et al. 2014). They could

become new models of interspecies evolution due to a

combination of their inherent hemiclonality phenomenon

and external radiation influence. The absence of previous

data on triploids in the Zone could possibly be explained

by samples too small to record non-numerous triploids.

Bonn zoological Bulletin 71 (2): 105-108

Or, we can hypothesize the recent emergence of triploids

due to local population systems evolution — in that case,

the role of radiation in this phenomenon is yet to be stud-

ied.

Our next step is to focus on obtained material deep-

ly. As the collection of new bigger samples has become

quite complicated due to ongoing war and following haz-

ardous aftermath, the application of advanced techniques

available for ploidy and genomes identification (karyolo-

gy, microsatellite analysis, FISH) is scheduled.

Acknowledgments. We want to thank the Chornobyl] Radiation

and Ecological Biosphere Reserve scientists, without whose

kind assistance the whole research wouldn’t have been per-

formed: Denys Vyshnevskyi and Sergii Domashevskyi. We also

thank Dmytro Shabanov for his help with improving the man-

uscript. And we are enormously grateful to all the defenders

of Ukraine, whose efforts made this paper possible to finalize.

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=— of Biodiversity Change

: KOENIG

Bonn zoological Bulletin 71 (2): 109-118

2022 Pertegal C. et al.

https://do1.org/10.20363/BZB-2022.71.2.109

forsewunas

museum

Research article

urn:|sid:zoobank.org:pub:43 EEF244-4D7E-435F-A7CC-52341 D9E7FBD

Description of a new trapdoor spider species from southern Spain

that exhibits an as yet unknown defence strategy

(Araneae: Mygalomorphae: Nemesiidae)

Cristian Pertegal ©., , liigo Sanchez Garcia ©. Rafael Molero-Baltanas ©? & Stephen Knapp”!

!3 Departamento de Zoologia, Universidad de Cordoba, Edificio C-1, Campus de Rabanales, E-14071 Cordoba, Spain

?Zoobotanico de Jerez. c/ Madreselva s/n, E-11408 Jerez de la Frontera, Spain

*Casa Athene, km 1.2 CA 4300, La Muela, Vejer, E-11150, Cadiz, Spain

*Corresponding author: Email: cristianpertegal@hotmail.es

'urn:|lsid:zoobank. org:author:4 1 9DAFF5-D5E7-4F8D-B4D8-6F29EE3CCOE3

?urn:lsid:zoobank.org:author:87A9A384-5201-4D09-AFOC-84E1C602001B

3urn:lsid:zoobank.org:author:826D3806-9744-4C48-8600-5EE8E034A 13F

*urn:Isid:zoobank.org:author: 5F404D7C-0302-499C-990C-005CBB321DB6

Abstract. The description of the new trapdoor spider species Nemesia shenlongi sp. n. is provided, together with details

of its habitat and burrow structure. The spider uses a round ball made of soil particles and saliva to plug part of its burrow

and leave an isolated chamber for protection. The ball differs from all known ‘burrow-blocking structures’ produced by

Nemesiidae, for example, Nemesia manderstjernae Koch, 1871, and N. fagei (Frade & Bacelar, 1931), in that it is not

packed in silk and is not attached to the burrow wall. The new species is compared with morphologically similar Neme-

Sia spp. as well as with N. fagei.

Keywords. Ecology, Nemesiidae, phenology, taxonomy, trapdoor spiders, western Mediterranean.

Descripcion de una especie nueva de arafa trampera del sur de Espaiia que presenta una estrategia de defensa hasta

ahora desconocida (Araneae: Mygalomorphae: Nemesiidae)

Resumen. Se aporta la descripcién morfologica de la nueva especie de arafia trampera Nemesia shenlongi sp. n. junto a

detalles de su habitat y la estructura de su madriguera. Esta arafia utiliza una bola que fabrica con particulas de suelo y

saliva para bloquear parte de su madriguera y dejar una camara aislada donde protegerse. La bola difiere de todas las ‘es-

tructuras de bloqueo de madrigueras’ conocidas que la familia Nemesiidae fabrica, por ejemplo, Nemesia manderstjernae

Koch, 1871, y N. fagei (Frade & Bacelar, 1931), en que no esta envuelta en seda y no esta unida a la pared de la madri-

guera. La nueva especie se compara con otras especies morfologicamente similares, asi como con N. fagei, que muestra

ISSN 2190-7307

http://www.zoologicalbulletin.de

un comportamiento similar.

Palabras clave. Arafias tramperas, ecologia, fenologia, Mediterraneo occidental, Nemesiidae, taxonomia.

INTRODUCTION

Trapdoor spiders conceal their burrow with a hinged cov-

er at the entrance. Their life takes place almost entirely

inside these burrows, where they feed, develop and breed:

only the mature males leave the safety of their nest to

look for females to mate (Buchli 1969). Among trapdoor

spiders, the genus Nemesia Audouin, 1826 currently con-

tains 68 named species, mostly distributed in the western

Mediterranean basin (World Spider Catalog 2022). His-

torically the identification of Nemesia species has been

a challenge because the descriptions of some species are

imprecise and ambiguous, based on a single sex, even on

a single specimen; moreover, the type material of some

taxa is lost. Interspecific differences are very subtle in

many cases (Bond et al. 2006) and the hidden life in a

Received: 01.06.2022

Accepted: 08.09.2022

burrow make the study of these spiders even more diffi-

cult. Some of the recent publications incorporate data on

the behaviour and type of burrows built by each species

(Decae & Huber 2017; Luis de la Iglesia 2019; Luis de

la Iglesia et al. 2021; Calvo 2020, 2021). The structure of

the nest could help with species level identification (Mora

2015: 31, 76, 304), as some of the Nemesia species build

very characteristic burrows. Most informative in this

respect are those species that construct special, presum-

ably defensive, devises inside their burrows (Moggridge

1873, 1874; Frade & Bacelar 1931). Nemesia fagei is so

far the only Iberian species for which such a devise has

been formally reported, although several unpublished re-

ports exist in which Iberian Nemesia species are indicated

to construct internal burrow plugging devises. Nemesia

fagei constructs a silk packed, bullet-shaped clay plug

Corresponding editor: B. Huber

Published: 15.09.2022

110 Cristian Pertegal et al.

that is attached to the internal silk lining of the burrow

(see Frade & Bacelar 1931 for a description of the plug

and the associated defence behaviour) Here we describe

a second Iberian species that, in contrast to all known

types of burrow-plugging devices, constructs a burrow

plug that is not packed in silk and is not attached to the

silken burrow lining.

In this respect, the here described burrow-plugging de-

vice appears unique within fossorial mygalomorph spi-

ders as they are known today.

MATERIAL AND METHODS

The specimens of the new species were collected in the

area of Torrecera, Jerez de la Frontera, Spain. Males were

collected by dropping traps with water and salt as pres-

ervation medium, while females were collected directly

by searching their burrows. Specimens collected by both

methods were preserved in 70% alcohol and deposited in

the collections of the Museo Nacional de Ciencias Na-

turales (MNCN) and the personal collection of the first

author (CPC).

Photographs were taken with a Finepix S camera, a

Nikon D3300 camera coupled with an adapter to the ste-

reomicroscope Euromex SB.1903-P and a Bresser ocular

camera. Morphological characters proposed in previous

works were considered for description (Decae et al. 2007;

Isaia & Decae 2012; Decae & Huber 2017; Zontein 2017;

Luis De La Iglesia 2019; Calvo 2020, Luis De La Igle-

sia et al. 2021), including the copulatory organs of both

sexes. The spermathecae of females were observed by

cleaning the epigastric area with microdissection scissors

and a hypodermic needle. The copulatory organs of spec-

imens of both sexes were extracted to generate an image

by photo stacking. In addition, the epiandric area, the ra-

tio length/width of the tibia of the palpus (Fig. 1) and the

ratio number of epiandric fusillae/carapace length were

studied in males, and the corresponding means and stan-

dard deviations of these metrics were calculated.

Spine formulae of palps and legs are indicated by the

number of spines on left member segment and the num-

ber of spines on the same segment of the right member,

both numbers separated by a dot. Variations are given

with the same formula but between parentheses.

Abbreviations of body and legs parts

AER = anterior eye row, length

ALE = anterior lateral eye, length

AME = anterior median eye, length

Bl = body, length

Bul = bulb, length (Fig. 1)

Cal = caput, length

Ch = caput, height

Cl = carapace, length

Bonn zoological Bulletin 71 (2): 109-118

Clyl clypeus, length

Cw = carapace, width

Cyl = cymbium, length

18) = eye, length

Eml = embolus, length (Fig. 1)

Ewl = embolus, width 1 (Fig. 1)

Ew2 = embolus, width 2 (Fig. 1)

Ew3 = embolus, width 3 (Fig. 1)

Fel = femur, length

LI = labium, length

Lw = labium, width

Ml = maxillae, length

Mel = metatarsus, length

MTF4 ratio = relative lengths of metatarsus, tibia

and femur of leg IV

Mw = maxillae, width

Pal = patella, length

PER = posterior eye row, length

PLE = posterior lateral eye, length

PLS = posterior lateral spinnerets

PME = posterior median eye, length

PMS = posterior median spinnerets

POP = pattern of the deep black pigmentation

on the ocular process

PSP = prolateral spines of patella

RSP = retrolateral spines of patella

Sl = sternum, length

Sw = sternum, width

Tal = tarsus, length

Aa: = thorax, height

Til = tibia, length (Fig. 1)

Tiw = tibia of the palp, width (Fig. 1)

RESULTS

Taxonomy

Nemesia shenlongi sp. n.

urn: |sid:zoobank.org:act: 400COD68-3 0E3-4F 08-AFD5-26181F CBBABF

Figs 24

Holotype

SPAIN: Torrecera, Jerez de la Frontera, 36.5968, -5.9686,

alt. 15 m., 6, MNCN 20.02/20440, 21.ix.2021.

Paratypes

SPAIN: Torrecera, Jerez de la Frontera, 36.5968,

-5.96864, alt. 15 m, 3 Gd (CPC), 21.ix.2021; 5 od

(CPC), 26.1x.2021; 1 2 (CPC), 1.xii.2019; 2 99 (CPC),

6.1x.2020; 1 9 MNCN 20.02/20441, 20.viii.2020; 1 9

(CPC), 4.1v.2021.

Etymology. The species is named after the wish-granting

mythological dragon, Shenlong, that can be summoned

by collecting all seven Dragon Balls.

©LIB

Description of a new trapdoor spider species from southern Spain 111

Fig. 1. a. Nemesia uncinata (Bacelar, 1933), copulatory bulb. b. N. uncinata, male palpal tibia. Abbreviations: see Material and

methods.

Diagnosis. Within Nemesia, N. shenlongi sp. n. fits

in a morphologically well-defined species group con-

taining N. dorthesi (Thorell, 1875), N. santeugenia (De-

cae, 2005), N. santeulalia (Decae, 2005), N. uncinata

(Bacelar, 1933), and N. valenciae (Kraus, 1955) charac-

terized by filiform posterior median spinnerets, females

with sac-like or potato-shaped spermathecae and males

with dorsal spines on the tibiae and metatarsi, and glo-

bose and ornamented bulbs (Zonstein 2017, 2019). Males

of N. shenlongi sp. n. demonstrate a very characteristic

morphology never described before, possessing the con-

ical embolus with a flat duckbill-shaped tip (Fig. 2).

The spermathecae in N. shenlongi sp. n. resemble two

mounds divided by a deep depression (Fig. 2). In males

of other species of this group, the basic part of the em-

bolus is fairly cylindrical and curved towards their apical

half, whereas in N. shenlongi sp. n. this area is flat and

wide. On the other hand, the spermathecae in the new

species may resemble those of N. santeugenia, N. santeu-

lalia, and N. uncinata. The morphology of the receptacles

of N. uncinata resembles two rounded mounds with the

glandular tissue distributed from the apical zone to the

middle of the spermatheca. The spermathecae of the oth-

er two species are potato-shaped in their distal half and

Bonn zoological Bulletin 71 (2): 109-118

differ from each other by the section in the middle of

the receptacle of N. santeulalia. Despite this, the over-

all shape of the spermatheca of N. shenlongi sp. n. is

more elongated at the end than in N. uncinata and lacks

a sectioned area as in N. santeugenia and N. santeulalia.

The burrow of N. shenlongi sp. n. could be considered

an important diagnostic character, because none of other

congeners was previously known to build a flask-shaped

burrow provided with a spherical plug of soil, in addition

to the typical thin flexible trapdoor.

Description

Male (holotype)

General habitus. Prosoma brown with dark pilosity on

the edges of the cephalic area, the fovea and the posterior

area. Opisthosoma yellowish with a dark cardiac mark

and five to six chevrons dorsally, and light yellow with

several dark dots ventrally (Fig. 3).

Prosoma. Clypeus dark with seven bristles and white

pubescence. POP broken between AME. Caput slight-

ly elevated. Crest zone light brown with seven bristles.

Shallow fovea. Chelicerae black, with two stripes of

whitish pubescence. Cheliceral rastellum of six spikes.

©LIB

112 Cristian Pertegal et al.

) :

Fig. 2. Nemesia. shenlongi sp. n., male and female copulatory organs. a. Retroventral, ventral and proventral aspects of bulb.

b. Prodorsal, dorsal and retrodorsal aspects of bulb. ¢. Ventral view of spermathecae. Scale bars: 0.5 mm.

Bonn zoological Bulletin 71 (2): 109-118 ©LIB

Description of a new trapdoor spider species from southern Spain 113

Smooth fang ridge. Cheliceral furrow with six promar-

ginal denticles. Sternum yellowish brown with circular to

oval sigilla, the two first pairs of sigilla marginal and the

third pair submarginal. Maxillae dark brown with some

probasal strong bristles, but in other males two to seven

cuspules. Labium and coxae dark brown (Fig. 3).

Palp and legs I-IV. Cymbium with dorsal patch of

18-19 spines in holotype, 19-27 in other males. Bulb

globular and embolus with duckbill-shaped tip (Fig. 3).

Palp tibia ratio 2.35+0.14. Palpal tibia dorsally with api-

cal patch of 13 spines (11-17 in other males). Clasper

hooked. Clasper field with scopula. Metatarsus I slightly

curved. Palp and legs femora black, the other segments

brown. Scopula extends from distal tibiae to tarsi I-I;

on legs HI-IV present only on tarsi. MTF4 ratio: tibia>

femur> metatarsus, usually tibia> metatarsus> femur.

Prolateral and retrolateral spination. Leg I, prolat-

eral side Pa: 2(1). 2(1); Ti: 3(4). 3(2-4); Met: 4(2). 4(3)

and retrolateral side Pa: 2(1). 2(1); Ti: 3. 3; Met: 3(1-2).

2(1-3). Leg II, prolateral side Pa: 2(1—3). 2; Ti: 3(2-4).

4(2—3-5): Met: 4. 3(2-4) and retrolateral side Pa: 1(2).

1(0); Ti: 3. 3(2); Met: 3(2-4). 3(2). Leg III, prolateral

side Pa: 2(0—3). 2(1); Ti: 533-4). 4(3); Met: 3(4). 3(4) and

retrolateral side Pa: 1.1; Ti: 3(2).3(2); Met= 3.3(4). Legs

IV, prolateral side Pa: 3(2). 3(2); Ti: 6(7—8). 9(5—7-8);

Met: 3(4—5). 3(4—5) and retrolateral side Pa: 1.1; Ti: 5(3-

4—7). 3(4-6-9); Met: 4(3-5—9). 5(3-4-6-7).

Patellar spine formulae, PSP [p: 1. 1; I: 2. 2; I: 2. 2; III:

DOP LVS. S|. Rok pp O0akr 2: 2 lie) elie iol seg ToT.

1]. Variations (n=8) PSP [p: 1. 1; I: 2. 2(1); Il: 1—-2-3.2;

I: 26). 2: IV: 32): 3@)). RSP [p: 0,01: 1Q).1@)3

1@)21(0)2 Ia eV sd) |:

Opisthosoma. In the holotype, epiandrous fusillae: 65;

trapezoid-triangular epiandric area of 1.23 mm? (Fig. 3);

density: 52.85 epiandrous fusillae/mm/’; ratio carapace

length/number of epiandrous fusillae 0.11. The remain-

ing males, epiandrous fusillae: 6346; epiandric area:

0.9940.12 mm/?; density: 64.12+6.24 epiandrous fusillae/

mm’; ratio carapace length/number of epiandrous fusillae

8.71+1.16. Spinneret morphology: PLS: basal segment

longer than middle segment+ distal segment. PMS cone-

shaped.

Measurements (mm). BI: 16.25 (14.09-16.75); dorsal

area of prosoma: Cal: 4.47 (3.95-4.88); Ch: 2.2 (1.85-—

2.43); Cl: 7.64 (7.68-8.07),; Cw: 6.12 (4.89-6.38); Th:

1.74 (1.48-1.94); Clyl: 0.33 (0.19-0.31); ocular group:

AER: 0.21 (1.07—1.15); PER: 1.09 (1.14—1.2); ALE: 1.16

(0.28-0.34); PLE: 0.29 (0.2—0.32); AME: 2.23 (0.18—

0.31); PME: 0.19 (0.16—0.27); El: 0.17 (0.54—0.59); ven-

tral area of prosoma: Mal: 2.16 (1.88—2.49); Maw: 1.35

(1.16—1.41); Lal: 0.67 (0.6—0.79); Law: 1.27 (1-1.27); SI:

3.92 (3.0-3.92); Sw: 2.91 (2.14-2.61); Palp: Bulb: 1.78

(1.58—1.86); Eml: 0.93 (0.82-1); Ew1: 0.61 (0.57-0.69);

Ew2: 0.24 (0.22-0.25); Ew3: 0.4 (0.40.49); Cyl: 1.61

(1.18—1.61); Til: 2.83 (2.42—2.9); Tiw: 1.27 (1.03-1.27);

Pal: 2.0 (1.62—2.07); Fel: 3.7 (3.11-3.95); TOTAL: 10.14

Bonn zoological Bulletin 71 (2): 109-118

(9.02-10.14); Leg I: Tal: 2.6 (2.55—2.97); Mel: 4.08

(3.48-4.11); Til: 4.18 (3.52-4.39); Pal: 3.54 (2.92-3.54):

Fel: 5.77 (5.21-6.18); TOTAL: 20.17 (18.23—21.03); Leg

II: Tal: 2.54 (2.18-2.69); Mel: 4.25 (3.64-4.25); Til: 3.97

(3.54-3.97); Pal: 3.04 (2.89-3.27); Fel: 5.03 (5.03—5.83);

TOTAL: 18.83 (17.57—19.64); Leg II: Tal: 2.43 (2.43-

3.05); Mel: 4.62 (4.36—5.06); Til: 3.74 (3.23-3.79); Pal:

2.75 (2.25—2.75); Fel: 4.75 (3.85-—5.05); TOTAL: 18.29

(16.96-19.2); Leg IV: Tal: 3.23 (2.81-3.23); Mel: 6.54

(5.97—7.05); Til: 6.58 (6.43—7.1); Pal: 3.88 (3.14—3.88):;

Fel: 6.57 (5.61-6.57); TOTAL: 26.8 (24.44—27.29).

Female

General appearance. Alive specimens light to dark

brownish grey. Carapace entirely covered with whitish

pubescence; black medal-shaped pubescence present on

cephalic area and around fovea, with orange brownish

background in alcohol (Fig. 4b). Opisthosoma light yel-

lowish, dorsally with darker brownish cardiac mark and

three to seven chevrons; ventrally with several darker

dots.

Prosoma. Clypeus dark coloured with a row of six mar-

ginal setae. POP black. Caput elevated. Crest zone light-

ly pigmented with a row of five to seven bristles. Fovea

shallow. Chelicerae as in holotype. Cheliceral rastellum

of six to seven strong spikes. Fang ridge smooth. Cheli-

ceral furrow with six to eight promarginal teeth. Maxillae

dark brown, three to seven cuspules. Labium of same co-

lour as maxillae. Sternum as in holotype (Fig. 4c).

Palp and legs I-IV. Legs and palps dark brown on

proximal segments to orange brownish on distal ones.

Scopula extends from tarsus to distal tibia on palp and

legs I-II; present only on tarsi of legs III-IV. MTF4 ratio

usually tibia> femur> metatarsus, but Tibia> metatarsus>

femur in one specimen.

Prolateral and retrolateral spination. Palp, prolater-

al side: Pa: 2.2; Ti: 2.2; leg I, prolateral side Pa: 2(1).2(1);

Ti: 3(2).3(2). Leg II, prolateral side Pa: 2(1).2; Ti: 3.3(2);

Met: 1(2).1(0). Leg III, prolateral side Pa: 2(1).2(1); Ti:

2(3).2(3); Met: 3(4).3 and retrolateral side Ti: 2.2; Met:

3.3(4). Legs IV, prolateral side Met: 2.2, retrolateral Ti: 3

(2).2(3); Met: 3.3.

Patellar spine formulae, variations (n=5) PSP [p: 2(3).

2; I: 2(1). 2(1); Il: 2(1).2; MH: 2(0-1). 2(1); TV: 2. 2]. RSP

var [p: 0.0; I: 0.0; If: 0.0; TI: 0.0; TV: 0.0].

Opisthosoma. Spinneret morphology: PLS: as in

male. Maculae present on basal segment, sometimes

poorly visible. PMS: cone-shaped (Fig. 4). Spermathe-

cae: mound-shaped with deep depression between both

mounds (Fig.4).

Measurements (mm). Bl: 16.84—20.87; dorsal area of

prosoma: Cal: 4.7—5.33; Ch: 2.21-2.91; Cl: 7.68-8.07;

Cw: 5.8-6.31; Th: 1.17—1.63; Clyl: 0.23—-0.39; ocular

group: AER: 1.21—1.36; PER: 1.3—1.42; ALE: 0.34—0.45:;

PLE: 0.26—0.38; AME: 0.18—0.31; PME: 0.14—0.24; EI:

0.61—0.73; ventral area of prosoma: Mal: 2.41—2.99;

©LIB

114 Cristian Pertegal et al.

Fig. 3. Nemesia shenlongi sp. n., 3, holotype (MNCN 20.02/20440). a. Body, dorsal. b. Maxillae, labium and sternum, ventral.

c. Bulb, proventral. d. Palp, prolateral. e. Opisthoma, ventral. f. Metatarsus I, with the clasper area marked with an arrow, prolateral.

g. Epigastric area with epiandrous fussillae, ventral. Scale bars: a= 5 mm; b, d, f-g = 1 mm; c = 0.5; e=2 mm.

Bonn zoological Bulletin 71 (2): 109-118 ©LIB

Description of a new trapdoor spider species from southern Spain 115

Fig. 4. Nemesia shenlongi sp. n., 2, paratype (MNCN 20.02/20441). a. Living specimen. b. Carapace, dorsal. e. Maxillae; labium

and sternum, ventral. d. Spinnerets, ventral. e. Spermathecae, ventral. Scale bars: b = 5 mm; c-d = 1 mm; e = 0.5 mm.

Bonn zoological Bulletin 71 (2): 109-118 ©LIB

116 Cristian Pertegal et al.

ek , Strap door P

Fig. 5. a. Habitat where Nemesia shenlongi sp. n. was found. b. First section of a burrow. c. Detail of the ball blocking the chamber.

d. Ball. Scale bar: 5 mm.

Bonn zoological Bulletin 71 (2): 109-118 ©LIB

Description of a new trapdoor spider species from southern Spain 117

Entrance

a b

Entrance

Fig. 6. Comparison of burrows. a. Burrow of N. shenlongi sp. n. b. Burrow of N. shenlongi sp. n. with the security chamber closed

in presence of predator. c. Burrow of N. fagei (Frade & Bacelar, 1931) based on the figure by Decae (1996).

Maw: 1.36—1.77; Lal: 0.75—0.93; Palp: Tal: 1.99-2.6;

Til: 2.15—2.68; Pal: 1.87—2.31; Fel: 3.08-4.16; TOTAL:

9.52-11.67; Leg I: Tal: 1.82—2.22; Mel: 2.87—3.15; Til:

3.48-3.89; Pal: 3.14-3.52; Fel: 5.11-5.57; TOTAL:

16.94-17.95; Leg II: Tal: 1.81-2.06; Mel: 2.73—2.97;

Til: 3.13—3.37; Pal: 2.69-3.35; Fel: 4-5; TOTAL: 15.24—

16.53; Leg III: Tal: 1.87—2.06; Mel: 3.12—3.27; Til: 2.29-

2.62; Pal: 2.44—2.7; Fel: 3.87-4.1; TOTAL: 13.77-14.45;

Leg IV: Tal: 2.02—2.44; Mel: 4.73-5.14; Til: 5.51-6.3;

Pal: 3.51-3.87; Fel: 4.94-5.49; TOTAL: 20.95—22.88.

Ecology

This species was found in an open ‘dehesa’ formed by

mastic, Pistacia lentiscus L., wild olive, Olea europaea

var. sylvestris (Mill.) Lehr, and fan palm, Chamaerops

humilis L., with a poor grass layer on the soil. Neme-

sia shenlongi sp. n. shares its habitat with other myga-

lomorph spiders such as Macrothele calpeiana (Walck-

enaer, 1805), spiders belonging to the genus Ummidia

(Thorell, 1875) and /beresia (Decae & Cardoso, 2005),

N. uncinata, and other unidentified Nemesia species. The

nest is covered with a thin flexible trapdoor in the en-

trance and a silk-linked flask-shaped burrow about 18 cm

deep. In addition, this spider builds a sphere reminiscent

of a marble of soil particles bound with saliva which

it uses to close the narrow area of the burrow as a safe

room (Fig. 5b—d). All found specimens tried to close their

burrow with particles of soil and silk when the ball was

removed. The spiders usually demonstrated a defensive

behaviour when removed from their nests.

Bonn zoological Bulletin 71 (2): 109-118

Males were collected between the 21" and 26" of Sep-

tember of 2021. No specimens were collected in the pit-

fall traps during the preceding season. Only the female

collected on 1*' of December of 2019 had been seen with

offspring.

DISCUSSION

The copulatory organ morphology of both sexes and the

novel protection system described above — the duck-

bill-shaped embolus of males, the mound-shaped sper-

mathecae with deep depression between both and the

marble-shaped plug — allow to distinguish this species

from other species of the genus Nemesia.

Several species of Nemesia build a supplementary

structure to protect itself as an internal door in its bur-

row or a plug (Moggridge 1873, 1874; Frade & Bacelar

1931). The internal doors are integrated in the wall of

burrow with silk and could be closed in case of danger,

for example in N. manderstjernae and N. meridionalis

(Costa, 1835) (based on figures of Moggridge 1873 and

1874), meanwhile the plugs are an auxiliary piece used to

block the nest. Nemesia fagei is the only previously de-

scribed species known to build a supplementary structure

to plug the entrance to the burrow. In N. fagei the plug is

bullet-shaped rather than round, and it is attached with

silk to the wall of the burrow at a short distance from

the entrance, which facilitates obstructing the entrance

(Fig. 6c). Nemesia shenlongi sp. n. makes a marble-like

©LIB

118 Cristian Pertegal et al.

plug that the spider usually lets loose at the bottom of the

burrow. In case of danger, the spider takes the ball and

plugs the narrow part of the burrow, leaving the tubular

area free and the wider part completely isolated and pro-

tected (Fig. 6a—b). Despite having a similar strategy of

protection, the morphology of N. fagei shows important

differences being compared with N. shenlongi sp. n., es-

pecially in the structure of the copulatory organs.

Acknowledgements. We want to thank the editorial team of

the journal for their work, the Delegacion Territorial de Medio

Ambiente y Ordenacion del Territorio de Cadiz for the authori-

sation (Ref. GB- / JMLV) and the curator of the MNCN, Dr.

Begofia Sanchez, for her kindness in depositing the type spec-

imens in the museum collection. Finally, the first author would

like to thank Irene Reina Alfonso, David and Victor Pertegal

Pérez for their patience and encouragement, and Arthur Decae,

Sergei Zonstein, José Antonio Luis de la Iglesia and Marta Cal-

vo for their advices.

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lis Costa, 1835 (Araneae, Myglomorphae, Nemesiidae), and

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and J. barbara in the Iberian Peninsula (Araneae: Nemesi-

idae). Arachnology 18 (2): 156-171.

https://doi.org/10.13156/arac.2018.18.2.156

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R (2021) Iberesia castillana (Frade & Bacelar, 1931), rede-

scription of the male, first description of the female, and a

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ae, Nemesiidae). Revista Ibérica de Aracnologia 38: 163-185

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©LIB

LIB Leibniz Institute for the Analysis

== of Biodiversity Change

Bonn zoological Bulletin 71 (2): 119-137

2022 Arkhipova K.I. et al.

museum

: KOENIG

https://do1.org/10.20363/BZB-2022.71.2.119

Research article

urn:|sid:zoobank.org: pub: C6BE4D70-494B-43 1 4-9E98-FO4BE61A2D17

Identity of certain stoneflies (Insecta: Plecoptera) from Ukraine and Poland,

with notes on available museum material from XIX—XX centuries

Khrystyna I. Arkhipova”', David Muranyi”?, Wieslaw Krzeminski©? & Roman J. Godunko®**

'4 State Museum of Natural History, National Academy of Sciences of Ukraine, Teatralna 18, 79008 Lviv, Ukraine

? Department of Zoology, Eszterhdzy Karoly Catholic University, Leanyka u. 6, Eger H-3300, Hungary

3 Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Slawkowska str. 17, 31016 Krakow, Poland

‘Biology Centre of the Czech Academy of Sciences, Institute of Entomology, Branigovskd 31, 37005 Ceské Budéjovice,

Czech Republic

*Department of Invertebrate Zoology and Hydrobiology, University of Lodz, Banacha 12/16, 90 237 Lédz, Poland

“Corresponding author: Email: roman.hodunko@biol.uni.lodz.pl; godunko@seznam.cz

'urn:|sid:zoobank.org:author:E52FE784-02 16-4F3B-8945-3719AE82FA AO

2 urn:lsid:zoobank.org:author:A 1905072-B418-4EC8-B19A-BOAC579319FE

> urn:|sid:zoobank.org:author:78COE530-D134-45C0-B79C-0155E3ED248F

*urn:lsid:zoobank.org:author:3 125D70F-AD47-4A D9-9B20-772FA7087B1E

Abstract. Available stonefly materials collected in Ukraine during late nineteenth and early twentieth century are enu-

merated and analysed from the National Museum, Prague (Czech Republic), State Museum of Natural History, National

Academy of Sciences of Ukraine (former Muzeum Dzieduszyckich), Lviv (Ukraine), and Institute of Systematics and

Evolution of Animals, Polish Academy of Sciences, Krakow (Poland). Occurrence of two autumnal representatives of

the Leuctra prima species group, 1.e., L. autumnalis Aubert, 1948 and L. carpathica Kis, 1966 is confirmed based on old

specimens, as well as presence of Perla pallida Guérin-Méneville, 1843 ‘Type 2’ sensu Sivec & Stark (2002). A lectotype

is designated for Arcynopteryx carpathica Klapalek, 1907, a junior synonym of A. dichroa (McLachlan, 1872). Notes on

the wing venation variability of Oemopteryx loewii (Albarda, 1889) are presented. Additionally, [soperla sudetica (Ko-

lenati, 1859), Dinocras megacephala (Klapalek, 1907), Perla bipunctata Pictet, 1833, Brachyptera braueri (Klapalek,

1900), Brachyptera monilicornis (Pictet, 1841), Brachyptera seticornis (Klapalek, 1902), Oemopteryx loewii (Albarda,

1889), Rhabdiopteryx harperi Vingon & Muranyi, 2009, Taeniopteryx nebulosa (Linnaeus, 1758), and Nemurella pic-

tetii Klapalek, 1900 are reported for Krakow collection for the first time. Corrections of the previous determinations are

performed. The occurrence in Ukraine of further two species, 1.e., /soperla sudetica (Kolenati, 1859) and Siphonoperla

transsylvanica (Kis, 1963), is confirmed based on recently collected material.

Keywords. Carpathians, Dziedzielewicz, Klapalek, faunistic, Leuctra, Arcynopteryx, Oemopteryx, lectotype, paralecto-

ISSN 2190-7307

http://www.zoologicalbulletin.de

type.

INTRODUCTION

The stonefly (Plecoptera) fauna of Ukraine received ac-

tive exploration during the last decades of the 19th and

early 20th century, due to the activity of Jozef Dziedziele-

wicz (*1844—F1918) and his long-term cooperation with

Franti8ek Klapalek (* 1863-71919), and other entomolo-

gists of the former Austro-Hungarian Empire (for more

information see Zhiltzova 1966; Godunko & Ktonows-

ka-Olejnik 2003; Godunko & Klymyshyn 2004; Diakiv

2010). Working in different cities of Western Ukraine

and travelling with the aim to collect Neuroptera, J. Dz-

iedzielewicz takes a part of support also from the side of

V. Dzieduszycki that established and built the Muzeum

Dzieduszyckich in Lviv. Then, J. Dziedzielewicz influ-

enced by him returned to Lviv and worked part time as

scientific secretary at the Museum. In 1915 J. Dziedziele-

wicz took the part of the collection of aquatic insects with

Received: 11.02.2022

Accepted: 01.09.2022

him to Myslenice village (Poland) and it was also trans-

ferred later to the Akademia Umieyetnosci w Krakowie [=

Polish Academy of Art and Sciences in Krakow] in 1918,

after his death. Dziedzielewicz’ collecting activity con-

cerned mostly to the Ukrainian Carpathians and several

other regions of former Galicja [[ammunna (Halychyna)

in Ukrainian]. However, some part of the material was

also collected in the Western Carpathians within the pres-

ent territory of Poland. The results of these early studies

were discussed in a detailed faunistical summary pub-

lished by the end of WWI (Dziedzielewicz 1918).

The old collections of the stoneflies in three museums

studied contain the material from the second half of the

19th and first decades of the 20th centuries. The aquatic

insect collections housed at the Museum of Natural His-

tory of the Institute of Systematics and Evolution of An-

imals, PAS in Krakow (further ISEA) are only a part of

the larger entomological collections formerly housed at

Corresponding editor: R. Peters

Published: 28.09.2022

120 Khrystyna I. Arkhipova et al.

the Museum of Physiographical Commission of the Pol-

ish Academy of Art and Sciences in Krakow (see above).

Such collections as documentary material were deposited

obligatorily by scientists (including J. Dziedzielewicz),

who had in that way to account for money granted to

them for the conduction of studies.

As aresult of cooperation between Dziedzielewicz and

Klapalek, numerous dried pinned specimens from the

Ukrainian Carpathians replenish the Klapalek’s collec-

tion which 1s now deposited in the National Museum in

Prague (further NMP).

Another part of the specimens collected by J. Dz-

iedzielewicz is housed at the State Museum of Natural

History NAS Ukraine in Lviv (further SMNH). The first

reinvestigation of the stoneflies collection in SMNH was

held in 2002—2003 and some results were published by

Godunko & Ktonowska-Olejnik (2003). However, the

second part of J. Dziedzielewicz’ plecopteran collection

housed in ISEA remained unidentified. The resolving of

some dispute questions in the field of identification and

some changes in taxonomy predetermined the revision

and identification of the specimens from both of these

collections.

Due to confirm the identity of old Plecoptera speci-

mens, we investigated the collections where these mate-

rials were traced, namely in NMP, SMNH and ISEA. The

available museum material from Ukraine and partly from

Poland is enumerated below, with notes and figures on

some important specimens and diagnoses of certain taxa,

completed with recent collecting that confirm identity of

lacking old specimens.

MATERIAL AND METHODS

The historical specimens examined are stored dry in the

Department of Entomology, Natural History Museum

(NMP) and Natural Museum (ISEA), and preserved in

75% ethanol in the Laboratory of Entomology (SMNH)

(for more information see Arkhipova et al. 2018). Iden-

tifications of material discussed below have been per-

formed by us in 2002—2003 and 2012-2016.

Drawings were made with the aid of a drawing tube

applied to a Nikon SMZ800 microscope. Photos were

made with a Keyence VHX 5000 digital microscope and

a Nikon D70s camera. Some drawings were made using

a stereo microscope Olympus SZX7 and a microscope

Olympus BX41, both equipped with drawing attachment.

Specimen terminalia were cleared in KOH and stored in

a microvial with glycerine pinned beneath the specimen.

Recently collected specimens are preserved in 75% etha-

nol and stored in the Collection of Smaller Insect Orders,

Department of Zoology, Hungarian Natural History Mu-

seum (HNHM).

Bonn zoological Bulletin 71 (2): 119-137

Morphological terminology primarily follows Graf

et al. (2012), Muranyi (2011), Ravizza & Vincon (1998)

and Sivec & Stark (2002).

Distributional data were compiled from literature in-

formation from various sources referenced in Plecoptera

Species File (PSF) (DeWalt et al. 2018). Distributional

information of old specimens analysed based on original

labels; for materials from SMNH and ISEA the original

labels cited without changes.

Abbreviations used in the text

For geographical regions

BM = Beskid Makowski

CCA Ciscarpathians

CHOR = _ Chornohora Range

GOR = _ Gorgany Range

GP = Great Polissya

ISP, = Little Polissya [also as ‘Male Polissya’ |

PS = Pogorze Slaskie

MAP = Matopolska Range

PH = Podilska Hight

ROZ = _ Roztochchia Range

SAB =_ Sadecki Beskid

SG = Slask Gorny

ok = ‘Tatry

WPOD =_ Western Podillya

UC = Ukrainian Carpathians

For collectors

CU = collector unknown

JD = J. Dziedzielewicz

FS = F Schille

JF = J. Fudakowski

KJ = K. Jelski

PD = P. Dzieduszycki

VL = V. Lazorko

JW = J. Werchratski

Abbreviations from original labels [geographical and

administrative terms]

dol. [dolina] = valley

Gub. = Gubernya [province]

p. or pot. [potok] = stream

localities

Ag [Angielow]| = Anheliv village (Lvivska

Region, Ukraine)

Kol. or Kt. [Kotomyja] = Kolomyya town (Ivano-

Frankivska Region, Ukraine)

Kosciel. = Koscielisko (Tatry, Poland)

Krak = Krakow (Poland)

Mik. [Mikuliczyn] = Mykulychyn (Ivano-

Frankivska Region, Ukraine)

Molodyatyn village (Ivano-

Frankivska Region, Ukraine)

MI [Miodiatyn] =

©LIB

Identity of certain stoneflies from Ukraine and Poland with notes on available museum material from XTX—XX centuries 121

Rebrow [Rebrowacz] = Rebrovach Mt. (Ukrainian

Carpathians)

Wad. = Wadowice (Poland)

Wot [Wotyn] = Volyn Region (Ukraine)

Wr. = Warwarynci (Ternopilska

Region, Ukraine)

Zak. = Zakopane (Poland)

Others

Dz. or Dziendz. = J. Dziedzielewicz

Ogr. bot. [ogrod botaniczny| = botanical garden

okol. [okolicy] vicinity

w. [wakacje] = collected holiday time

sex unknown

(damaged specimens)

RESULTS AND DISCUSSION

Commented species list

Family Leuctridae Klapalek, 1905

Genus Leuctra Stephens, 1836

Leuctra albida Kempny, 1899

Material: [SEA: POLAND: “Rytro”, “30/13”, SAB

(FS),12:

Leuctra fusca (Linnaeus, 1758)

Material. [SEA: UKRAINE: “Mikuliczyn. 1 do 5. VIII.

1904”, “126/19”, GOR (JD), 19.

Remarks. Widespread species in the Carpathians re-

gion, mentioned by Dziedzielewicz (1918) from locality

cited above.

Leuctra autumnalis Aubert, 1948

Figs 14

Material. NMP: UKRAINE: “Pod Turkul, 10.[X.1908”

[under Turkul Mt.|], CHOR (JD), 44, 299 [as L. sigin-

fera Kny, det. Klapalek; one male and one female termi-

nalia KOH boiled, pinned beneath the specimen in mi-

crovial]; “Pozyzewska [Pozhyzhevska Mt.], 9.09.1908”,

CHOR (JD), 19 [as L. siginfera Kny, det. Klapalek;

terminalia KOH boiled, pinned beneath the specimen in

microvial].

Diagnosis. Male tergum VIII with large median pro-

cess having short but widely separated digits; tergum IX

median sclerite with slight medial subdivision; tergum X

with paired spike-like posterior process by the postero-

medial hollow. Female subgenital plate with posterior

lobes not divided, posterior margin rounded; sternum IX

with slight, V-shaped anteromedial hollow.

Remarks. The closely related species L. signifera

Kempny, 1899 was described from Gutenstein, Lower

Bonn zoological Bulletin 71 (2): 119-137

Austria, and for several decades all autumnal members

of the L. prima group were reported under this name

(Mosely 1932). The first related species that was distin-

guished is L. autumnalis, originally described from Swit-

zerland (Aubert 1948). Its occurrence in the Carpathians

was proved in the 60ies, reported simultaneously from

Romania (Kis 1963a), Slovakia and the Czech Repub-

lic (RauSer 1965), and Ukraine (Zhiltzova 1964). Pres-

ence of L. signifera in the Carpathians was questioned

(Kis 1966; Zhiltzova 1968) and it was not reported since

but seems to be restricted to the Eastern Alps and its

foothills in Austria, Hungary and Slovenia (Ravizza &

Vingon 1998; Graf & Weinzierl 1999; Kovacs 2006).

Contrary, L. autumnalis is a widespread Alpine-Carpath-

ian species extending also to the Jura (Ravizza & Vincon

1998; Ravizza 2002). The present Ukrainian specimens

of the NMP were originally identified as L. signifera by

F. Klapalek (based on the handwrite of the det. label), but

not enumerated in publication before.

Leuctra carpathica Kis, 1966

Figs 5-8

Material. NMP: UKRAINE: “Worochtensky, 7.09.1908”

[Vorokhtenskyi stream], CHOR (JD), 34 [as L. sig-

infera Kny, det. Klapalek; one terminalia KOH boiled,

pinned beneath the specimen in microvial); “Pozyzews-

ka, 9.09.1908”, CHOR (JD), 1¢ [as L. siginfera Kny,

det. Klapalek; terminalia KOH boiled, pinned beneath

the specimen in microvial ].

Diagnosis. Male tergum VIII with small median pro-

cess having long but closely set digits; tergum LX median

sclerite less widened and with deep medial subdivision;

tergum X lack posterior process by the posteromedial hol-

low. Female subgenital plate with posterior lobes divided

but set close, posterior margin rounded; sternum IX with

deep, U-shaped anteromedial hollow.

Remarks. The species was described from the South-

ern and Inner Eastern Carpathians of Romania (Kis 1966),

previously confused with L. signifera (Bogoesco & Tab-

acaru 1960). Subsequently, it was reported from several

further Carpathian localities in Romania (Kis 1974) but

still lacking from the Transylvanian Alps (Muranyi &

Kovacs 2015). Its presence in the Ukrainian Carpathi-

ans was first reported by Zhiltzova (1977). The species

seems to be lacking from the Northern Carpathians but

present in the extreme nortwestern ranges of the Eastern

Carpathians in Hungary (Andrikovics & Muranyi 2001),

Poland (Sowa 1970) and Slovakia (Ziak 2016). Report

from the Eastern Alps (Graf & Weinzierl 1999) refers

to another species of not yet resolved identity (W. Graf

pers. com.; Muranyi 2006), whereas report from Mon-

tenegro (Kacanski & Baumann 1981) probably refers to

the Dinaric L. jahorinensis Ka¢anski, 1972 or L. mal-

cor Muranyi, 2007 (Muranyi 2007, 2008). The present

Ukrainian specimens of the NMP originally identified as

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hee Khrystyna I. Arkhipova et al.

4%, 4

©...

“

Figs 1-4. Terminalia of Leuctra autumnalis Aubert, 1948. 1. 3, Ukraine [Turkul Mt., Chornohora Range]. 2. 3, Austria [St. Os-

wald]. 3. 9, Ukraine [Pozhyzhevska Mt., Chornohora Range]. 4. 2, Austria [St. Oswald]. The specimens are deposited in NMP

and HNHM collections. Scale bar: 1 mm.

Bonn zoological Bulletin 71 (2): 119-137 ©LIB

Identity of certain stoneflies from Ukraine and Poland with notes on available museum material from X-X—XX centuries 123

L. signifera by F. Klapalek (based on the handwrite of the

det. label); the specimens from Vorokhta village were re-

ported by Dziedzielewicz (1918) as “L. signifera Kemp”.

Leuctra major Brinck, 1949

Material. [SEA: UKRAINE: “31/7 Zall.”, “87/1” (CU),

19; POLAND: “Tatry 8/2”, “147/10”, T (JD), 14 [both

specimens were located in the box near label: “cylindri-

cal. DeGe” & “major Brinck’’].

Remarks. This species mentioned from Tatry Mts.

(Zakopane) and Krzeszowice in Poland, and Iva-

no-Frankivska Region (Ukraine) as “Leuctra cylindrica.

De Geer” by Dziedzielewicz (1891).

Leuctra spp.

(damaged and incomplete specimens)

Material. JSEA: UKRAINE: “Tartarow. Chomiak.

15.7 1903”, GOR (JD), 134; “9.VII.1904. Chomiak.

Mikuliczyn”, “125/19”, GOR (JD), 19: “Kolomyja.

17. XI. 1900”, “78/14”, CCA (JD), 19: “Mykietynice.

Stanislawow”, “86/1”, CCA (JD); 14: “20.VII.1909.

Chomiak Mikuliczyn”, “125/15”, GOR (JD), 2¢¢;

“(Chomiak) Mikuliczyn. 9.VII.1904”, “125/19”, GOR

(JD), 42 2; “(Gorgan Bor.). Mikuliczyn. 13. VII.1904”,

“127/19”, GOR (JD), 134; “Worochta. Okolice.

24.10.1909. na Ssniegu.”, “134/24” [probably Leuctra

fusca (Linnaeus, 1758)], CHOR (JD), 14; “Worochta.

Okolice. 27.10.1908”, “134/24”, CHOR (JD), 24.3: “8/5

KEZ. “15/5? CCA(ID 22 OO: Sha” 9218 CGA OD).

283; POLAND: “Rytro”, “30/13”, SAB (FS), 299:

“Rytro”, “30/13”, SAB (FS), 24; “Rytro”, “30/13”,

SAB (FS), 14; “Rytro”, “29/15”, SAB (FS), 14; “Babia

Gora. Zawoja. 3.VIII.1909”, T (JD), 14: “27/7, Zall”,

“88/1” (CU), 19: “Myslenice. 13. VIII. 1909”, “131/24”,

MAP (JD), 14; “19/8 Zak.”, “87/1”, T (CU), 13; others:

“T16/5" (EU), 32:9"

Family Capniidae Klapalek, 1905

Genus Capnia Pictet, 1841

Capnia nigra (Pictet, 1833)

Material. SMNH: UKRAINE: “Kolomyja 4. Prut.”,

CCA (JD), 14,19.

Remarks. Some specimens that previously were iden-

tified as C. nigra (Godunko & Ktonowska-Olejnik 2003),

belong to Capnia vidua Klapalek, 1904, namely all mate-

rial reported from the Chornohora Mts.

Capnia vidua Klapalek, 1904

Material. SMNH: UKRAINE: — “Foreszczynka.

15.11.1909. Czarnohora.”, CHOR (JD), 1¢; “Czarnoho-

ra. Foreszcezynka. 1.IV.1910.”, CHOR (JD), 744, 229;

Bonn zoological Bulletin 71 (2): 119-137

“Czarnohora. Foreszczynka. 1.1V.1910.”, CHOR (JD),

24.4; “Czarnohora. Foreszcezynka. 1.1V.1910.”, CHOR

(JD), 12?; “Chomiak. Btotek. 26.V.1909”, GOR (JD),

12, [this specimen was identified as Capnia atra Mor-

ton, 1896 by Godunko & Ktonowska-Olejnik (2003)];

“Koltomyja 3.4 Prut”, CCA (JD), 16.

Remarks. Four additional specimens from SMNH

collection reported as C. vidua by Godunko & Ktonows-

ka-Olejnik (2003) in fact are damaged representatives of

the order Raphidioptera labeled as: 1: “3/8 Wol.”, “36.

Bepxpau.”, GP (JW); 13: “10.V.52.”, “Bproxosyun”,

ROZ (VL); 14: “Tligurore Ky3bmuuneus 15.V1.08. coll.

JIa30pKo”, GR (VL).

Family Perlidae Latreille, 1802

Genus Agnetina Klapalek, 1907

Agnetina elegantula (Klapalek, 1905)

Material. SMNH: POLAND: “Krakow. Wisla. VIII”,

MAP (? KJ), 19.

Remarks. A single specimen from SMNH collection

was wrongly identified as Marthamea vitripennis (Bur-

meister, 1839) and published in Dziedzielewicz (1918);

no any records for Ukraine.

Genus Dinocras Klapalek, 1907

Dinocras megacephala (Klapalek, 1907)

Material. NMP: UKRAINE: “Worochta, 06.1911” [Vo-

rokhta village], leg. Lokay, det. I. Sivec 1999, CHOR,

14 [as P. marginata, det. Klapalek?; lacks terminalia].

ISEA: UKRAINE: “26/5 K?’, “138/4”, CCA (JD), 19.

Remarks. There were no published mentions about the

presence of D. megacephala in old collections and con-

tributions. In 2010—2014 we sampled larvae in the Chor-

na Tysa River near Rakhiv town and the exuviae in the

Bystrycya Nadvirnyanska River near Maksymets village

that were similar to D. megacephala by the description

(Diakiv 2010). However, for confirmation of these re-

cords, the adults should have been collected. The present

determination of male and female from both collections

listed above is a first reliable record of D. megacephala

from the Ukrainian Carpathians and Ukraine at all. Ad-

ditionally, it was found in Poland for the Tatry, Pieniny,

Bieszczady and Kotlina Nowotarska Mts. (Fiatkowski &

Kittel 2002).

Genus Perla Geoffroy, 1762

Perla abdominalis Burmeister, 1839

Material. NMP: UKRAINE: “Worochta, Rebrowacz”

[ Vorokhta village, Rebrovach Mt.], leg. A. Stoeckel, 13;

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124 Khrystyna I. Arkhipova et al.

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Figs 5-8. Terminalia of Leuctra carpathica Kis, 1966. 5. 3, Ukraine [Pozhyzhevska Mt., Chornohora Range]. 6. 3, Romania

[Tusa]. 7. 3, Ukraine [Pozhyzhevska Mt., Chornohora Range], lateral view. 8. 9, Romania [Tusa]. The specimens are deposited in

NMP and HNHM collections. Scale bar: 1 mm.

Bonn zoological Bulletin 71 (2): 119-137 ©LIB

Identity of certain stoneflies from Ukraine and Poland with notes on available museum material from XITX—XX centuries 125

“Worochta” [Vorokhta village], leg. A. Stoeckel, CHOR,

SMNH: UKRAINE: “Kotomyja 16.V.1899.”, CCA

(JD), 12; “Kolomyja. 19.V.99.”, CCA (JD), 14; “17.-6.-

1907 Worochta.”, “K-628 II-945”, “Perla abdominalis”’,

CHOR (JD), 13; “Kotomyja. Prut. 30.-V.”, CCA (JD),

299; “Kniazdwor. 31.V.”, CCA (JD), 14; “Kolomyja.

Pryt. 4.VI.”, CCA (JD), 12.

ISEA: UKRAINE: “Kotomyja, may’, “128/10”, CCA

(JD), 14; “5/5 KY’ “143/4”, CCA (JD), 14; POLAND:

“Rytro” “25/13”, SAB (FS), 19.

Remarks. Godunko & Ktonowska-Olejnik (2003)

reported this species for SMNH collection under the

synonymic name P. burmeisteriana Claassen, 1936.

The specimen from the Tatry Mts. were reported by Dz-

iedzielewicz (1918); detailed records from the Polish

Carpathians listed by Fiatkowski & Kittel (2002).

Perla bipunctata Pictet, 1833

Material. ISEA: UKRAINE: “Mikuliczyn 5.9.1903”,

“83/20”, GOR (JD), 19; “Mikuliczyn 5-10. VIII 1904”,

”116/19”, GOR (JD), 12; “25/5 Mt.”, “143/4”, CCA (JD),

12; “31/5 Stobidka lesna”, “36/7”, CCA (JD), 12; PO-

LAND: “Babia gora Zawoja” “153/29”, “Perla alpicola,

Klap.”, T (JD), 14; “Babia gora Zawoja 23.VIII.1909”

“153/29”, “Perla alpicola, Klap.”, T (JD), 12.

Remarks. There are no mentions about P. bipunctata

at the territory of the Ukrainian or Polish Carpathians till

1935 (Despax 1935). However, there are specimens of

P. bipunctata from the Ciscarpathians, Gorgany Range

and Tatry Mts. in ISEA collection. At the previous exam-

ination of SMNH collection one specimen was identified

as Perla sp. (Godunko & Ktonowska-Olejnik 2003).

Perla grandis Rambur, 1842

Material. SMNH: POLAND: “Babia Gora Zawoja.

23.VII.1909.”, T (JD), 12.

Remarks. From the Ukrainian part of the Carpath-

ians the species known only from the Tysa river-basin

(Kovacs et al. 2008a, 2008b). In Polish Carpathians that

was found also in the Bieszczady and Western Beskid

Mts. (Fiatkowski & Kittel 2002).

Perla marginata (Panzer, 1799)

Material. NMP: UKRAINE: “Worochta, VI.1911” [Vo-

rokhta village], leg. Lokay, det. F. Klapalek, vid. I. Sivec

1999, CHOR, 106'3'7, 29 9, [one male has a label: P. bar-

cinonensis, det. Klapalek].

SMNH: UKRAINE: “Kotomyja 16.V.1899.”, CCA

(JD), 19; “Mikuliczyn 3.7. Zeniec 1903”, GOR (JD),

19; “Tartarow. 6.7 Chomiak 1903”, GOR (JD), 1; “Ta-

tarow. (Blotek.) 28.-6.-1905. Dz.”, GOR (JD), 14, “Mi-

kuliczyn. (Okolice) 11.-7.-1906.”, GOR (JD), 13’; “ Ko-

Bonn zoological Bulletin 71 (2): 119-137

tomyja. 4.-VII.”, CCA (JD); 14: “Lanezyn. Prut. 8.VI.”,

CCA (JD), 14; “Kolomyja. 6.VII.”, “Perla maxima,

Scop.”, “3”, CCA (JD), 134; “Kotomyja. Prut. 13.-VII.”,

“1” CCA (JD), 12; “Kotomyja. Prut. 15-20.-VII.”, “6.”,

“Perla maxima, Scop.”, “2”, CCA (JD), 19; “Mikuliczyn

Zeniec. 20.VII.”, GOR (JD), 14; “Mikuliczyn okolice.

VII’, GOR (JD), 23'3; “Mikuliczyn okolice”, GOR

(JD), 13; “Prut”, CU, (JD), 1 larval skin; POLAND: “Ta-

try. 1891”, T (JD), 14; “Tatry”, T (JD), 14; “Tatry. VII.

VII.”, T (JD), 13; 2 larval skins, 19, all without labels.

ISEA: UKRAINE: “Ag”, “19/3”, CCA (JD), 14; “11/7

Dora.”, “37/7”, CCA (JD), 14; “5/8. Mt.”, “105/5”, CCA

(ID), 334; “29/9 Mlodiatyn (Kotomyjskie)”, “38/7”,

CCA (JD), 1: [all listed above specimens earlier were

identified as “Dinocras cephalotes” with respective label

in the entomological box]; POLAND: “Myslenice, Stro-

za 18 VIII 1909”, “151/24”, “Perla marginata Panz.”,

MAP (JD), 23°; “Rytro”, “25/13”, SAB (FS), 19; oth-

ers. “20/7 Klen” [the origin of the acronym is unknown],

“75/14” (CU), 19; “8/9 Wr.”, “141/4”, CCA (CU), 2206;

“142/4”, (CU), 13; “77/14”, (CU), 12.

Remarks. According to Sivec & Stark (2002), P. mar-

ginata is a West European species and does not occur

East of the Alps. The Ukrainian specimens in the NMP

are conspecific with the taxon depicted as P. marginata

by Kis (1974) from Romania, to confirm their identity

needs further studies. Some specimens of P. margina-

ta were identified previously as D. cephalotes (Curtis,

1827) by Godunko & Ktonowska-Olejnik (2003). We

also identified the specimen from Matopolska Region

[Myslenice]. Perla marginata is reported to be wide-

spread all over the Carpathians (Kis 1974; Teslenko &

Zhiltzova 2009; Ziak 2016).

Perla pallida Guérin-Méneville, 1943 ‘Type 2’ sensu

Sivec & Stark 2002

Figs 9-10

Material. NMP: UKRAINE: “Worochta, 06.1911” [Vo-

rokhta village], leg. Lokay, det. I. Sivec 1999, CHOR, 19

[as P. marginata, det. Klapalek]; “Worochta, 8.09.1908”,

JD, CHOR, 16 [as P. dacica, det. Klapalek].

SMNH: UKRAINE: “Chomiak. Blotek. 19.-6.-1907.”,

GOR (JD), 14; “Worochta. Okolice. 1.-8.-1907”, “Perla

dacica, Klap.! Det. Klapalek.”, CHOR (JD), 14; “Miku-

liczyn (Prut) 6. VIII.1909.”, GOR (JD), 19.

ISEA: POLAND: “Mikuliczyn 10.7.1903”, “116/19”,

GOR (JD), 1°.

Remarks. The presence of P. pallida in the Carpath-

ians was first reported by the description of P. dacica

Klapalek, 1907 from Romania. Later this taxon was re-

garded as a subspecies (Kis 1963a), then a synonym of

P. pallida (Kis 1974). In Ukraine, the species was first

reported by Dziedzielewicz (1918), probably on the ba-

sis of the NMP specimens. During the latest revision of

the genus, P. pallida was treated as a species complex

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126 Khrystyna I. Arkhipova et al.

with four ‘Types’ distinguished on the basis of egg mor-

phology (Sivec & Stark 2002). The Carpathian popula-

tions were first studied on the basis of eggs from Mara-

mures, Romania, confirming the presence of “Type 2’

(Muranyi 2006). Herein we confirm the same ‘Type 2’

from Ukraine, having eggs with distinct hexagonal FCIs

and collar made up of two narrow rows (Fig. 10). Perla

pallida is widespread all over the Carpathians.

Perla spp.

(damaged and incomplete specimens)

Materials. [SEA: UKRAINE: “5/8 Mi.”, “102/5”, CCA

(JD), 19: “Mykielynce Stanislawow”, “73/1”, CCA

(CU), 19; “13/6 KP’, “101/5", CCA (JD), 19; “15.

VIII.1904, Mikuliczyn”, “118/19”, GOR (JD), 14: “9/7

Mik.”, “104/5”, GOR (JD), 14: “Swidowa. Czortkow.

Kiel.”, “39/7”, CCA (JD), 192: others: “139/4” (JD), 1

exuvium; “76/14” (CU), 12;

Family Perlodidae Klapalek, 1909

Genus Arcynopteryx Klapalek, 1904

Arcynopteryx dichroa (McLachlan, 1872)

=Arcynopteryx carpathica Klapalek, 1906

Figs 11-13

Materials. NMP: UKRAINE: “Chomiak, pot. Bara-

ni, 6.07.1905” [Khomyak Mt., Baranii stream], det. F.

Klapalek, GOR (JD), 14,29 9 [the male was labelled as

lectotype and the females as paralectotypes by P. Zwick

in 1980 without further designation according to the re-

quirements of the International Code of Zoological No-

menclature (ICZN); male terminalia cleared, pinned be-

neath the specimen in microvial, one female have only the

wings remained]; “Chomiak, pot. Barani, 26.06.1905”,

det. F. Klapalek, GOR (JD), 244’, 19 [labelled as para-

lectotypes by P. Zwick in 1980 without further designa-

Bonn zoological Bulletin 71 (2): 119-137

Figs 9-10. Perla pallida Guérin-Méneville, 1943 “Type 2’ sensu Sivec & Stark 2002, 2, Ukraine [Vorokhta]. 9. Habitus of the

specimen in NMP collection. 10. Dissected eggs. Fig. 9 without scale. Scale bar: 10 = 0.5 mm.

tion according to the requirements of ICZN; one male

terminalia cleared, pinned beneath the specimen in mi-

crovial]; “Chomiak, Blotek, 9.07.1904”, det. F. Klapalek,

GOR (JD), 12 [wrongly labelled as paralectotype by P.

Zwick in 1980; not belongs to syntype series according

to the requirements of ICZN]; “Chomiak, 27.06”, det.

F. Klapalek, GOR (JD), 14, 12 [both lacks abdomen];

“Worochta, potok nad Rebrowaczem” , 25.06.1908”

[Vorokhta village, stream up to Rebrovach Mt.], det. F.

Klapalek, GOR (JD), 19°; “Worochta, 06.1911”, leg. Lo-

kay, det. F. Klapalek, GOR (JD), 14, 19; “Worochta”,

leg. Stoeckel, det. F. Klapalek, GOR (JD), 53'3..

SMNH: UKRAINE: “Danzerz. 20. VII — 1906. zrodta

przy drodze.”, CHOR (JD), 1; “Worochta. Okolice. p.

pod. Rebrow. 25.-6.-1907.”, CHOR (JD), 19; “Czarno-

hora. Kozty. 8.-7.-1907.”, CHOR (JD), 14; “Woroch-

ta. Okolice. 30.-7.-1907.”, CHOR (JD), 19; Worochta.

Okolice. pot. z Rebrow. 15.-6.-1908.”, CHOR (JD), 19;

“Chomiak [Blotek] 5.VI 1909”, GOR (JD), 14; “Tatry.

1892. Dziedz.”, “dovrensis Klapalek”, T (JD), 343.

ISEA: UKRAINE: “Chomiak. pot. Barani. 26.-6.-

1905. Dz.”, “82/20”, GOR (JD), 2¢'4 [both labelled as

paralectotypes by Kh.I. Arkhipova & R.J. Godunko in

October, 2019].

Remarks. Arcynopteryx carpathica is the fourth oldest

name presently classified as a junior synonym of A. di-

chroa (DeWalt et al. 2018). The synonymy under A. com-

pacta (McLachlan, 1872) was stated by Brinck (1949),

and inherited to A. dichroa when Teslenko (2012) clari-

fied the misunderstanding in the using of the two names.

In the present contribution we accepted synonymy of

A. dichroa and A. carpathica established by DeWalt et al.

(2018). In a previous contribution on SMNH collection

(Godunko & Ktonowska-Olejnik 2003) A. dichroa was

mentioned under the synonymic name A. compacta (for

more details see Teslenko 2012).

In 1980 Peter Zwick investigated the collection of

Frantisek Klapalek housed in NMP. The specimens of

©LIB

Identity of certain stoneflies from Ukraine and Poland with notes on available museum material from XIX—XX centuries 127

A. carpathica were selected, and part of them belong-

ing to the type series [syntypes according to ICZN, Arti-

cle 73.2.] were labelled as lectotype and paralectotypes,

without further designation according to the requirements

of ICZN. Later, Zwick (1982) published designation of

types for several species of the family Nemouridae only,

and mention of the specimen labelled as the lectotype of

A. compacta is lacking in his paper.

The quantity of syntype specimens were not reported

in the original description. However, Klapalek (1906)

mentioned many males and females. In addition to the

NMP collection, two males discovered in ISEA also be-

long to the type series and present paralectotypes accord-

ing to Article 74.1.3. of the ICZN (see also below). At the

same time, 14 and 39 of A. carpathica listed by Sar-

tori et al. (1990) from the Musée de Zoologie, Lausanne

as paratypes and labelled as “Tchécoslovaquie, Carpates,

Chomiak, Barani, 3-4.VII. 1905. Dziedziedlewicz leg.

A sec. Ex coll. Klapalek!” are not syntypes, because the

date of material sampling does not match the information

in the original description [see ICZN, Articles 73.2., 74].

Extensive molecular studies on the Holarctic A. di-

chroa suggested strong population differentiation be-

tween mountain ranges (Theissinger et al. 2012). Further

investigations may result in reinstatement of some of the

fifteen names treated as junior synonyms of A. dichroa.

For the Carpathian and Bulgarian populations that were

included in one lineage during the studies of Theissinger

et al. (2012), there are two available names: A. carpath-

ica and the older Dictyopteryx transsylvanica Klapalek,

1901. There are 14 and 19 syntypes available also of the

latter in the NMP (ROMANIA, Transylvania, leg. Strobl:

according to the original description, they were collected

at the Iezer Lake, Rodna Mts. on 27.07.1896). Thought it

would be useless to illustrate genital characters without

knowledge on distinctive features that may separate these

presently cryptic species, herein we include figures on

Figs 11-13. Type series of the Carpathian Arcynopteryx Klapalek, 1904. 11. A. carpathica Klapalek, 1906, lectotype and paralec-

totypes. 12. Dictyopteryx transsylvanica Klapalek, 1901, syntypes. 13. A. carpathica Klapalek, 1906, lectotype. The specimens are

deposited in NMP collection [entomological box]. Without scales.

Bonn zoological Bulletin 71 (2): 119-137

©LIB

128 Khrystyna I. Arkhipova et al.

the habitus of the type series due to recognition of the

available types (Figs 11-13).

Thus, in order to preserve stability of nomenclature in

the future we designate the lectotype and paralectotypes

of A. carpathica herein [ICZN, Article 74], using syntype

series housed in NMP and ISEA [according to ICZN, Ar-

ticles? 322) |

Lectotype [ICZN, Article 74, Recommendations

73C, 74C, 74D]: 13 labelled as “Chomiak, pot. Bara-

ni, 6.07.1905”; coll. J. Dziedzielewicz; det. F. Klapalek,

housed in the collection of the National Museum in

Prague (Figs 11, 13).

Type locality [ICZN, Articles 74 and 76.2., Recom-

mendation 74E]: UKRAINE, Ivano-Frankivsk Region,

Gorgany Range, Khomyak Mt, Baranii stream.

Paralectotypes [ICZN, Article 74, Recommenda-

tion 74F]: 243, 19 labelled as “Chomiak, pot. Barani,

26.06.1905”; 229 labelled as “Chomiak, pot. Barani,

6.07.1905” (all specimens in collection of NMP); 204

labelled as “Chomiak. pot. Barani. 26.-6.-1905. Dz.”,

“82/20” (both specimens in collection of ISEA).

Remarks. Arcynopteryx dichroa is a Holarctic species

that is known from all the river-basins of the Ukrainian

Carpathians in Chornohora, Gorgany and Rivna Ranges

and in Poland from the Western Beskid Range, Bieszcza-

dy Ranges and Tatry Mts. (Dziedzielewicz 1918; Despax

1933; Zhiltzova 1966, 1968; Fiatkowski & Kittel 2002).

Genus Diura Billberg, 1820

Diura bicaudata (Linnaeus, 1758)

Material. SEA: UKRAINE: “Chomiak. Pot. Barani.

26.-6.-1905. Dz.”, “82/20”, GOR (JD), 23'4; “Chomiak,

pot. Barani. 26.6.1905. Dz.”, GOR (JD), 14; POLAND:

“Babia Gora. Zawoja. 27. VII.1909”, “154/24”, T (JD),

1s “1891 Tatty”, 125/10", TE GID). 1h: “Tatry:8/2,

“125/10”, T (JD), 24'S [three last specimens were locat-

ed in the entomological box under label “Dictvogenus

Klap. alpinus Pict.’ |.

Remarks. Diura bicaudata 1s mentioned from the

Tatry Mts. in old publications as “Dictyopterygella

Klap.”, that were collected by F. Klapalek in 1904 (see

e.g. Dziedzielewicz, 1918). Additionally, there are also

specimens from the Gorgany Range within the Ukrainian

Carpathians. The species was also found in the Nesam-

ovyte Lake (Chornochora Range) (Diakiv 2011). In Po-

land it was found also in the Western Sudety, Western

Beskid and Bieszczady Ranges (Fialkowski & Kittel

2002).

Bonn zoological Bulletin 71 (2): 119-137

Genus Isogenus Newman, 1833

TIsogenus nubecula Newman, 1833

Material. SMNH: UKRAINE: “Jaryszow. Gub. Podol-

ska. 30.IV. zeb P. hr. Dzieduszycki”, WPOD (PD), 13;

“Wertelka pod Zaloice. 26.V.”, (JF), 12.

ISEA: UKRAINE: “12/5 Lancorona”’, “137/4”, BM

(JF), 14; “Kotomyja”, “127/10”, CCA (JD), 1°: others:

21/9" (CU), eo CW), 1G.

Remarks. There were only two specimens in the col-

lection of SMNH from the Western Podillya (Khmel-

nytska Region) mentioned by Dziedzielewicz (1918).

During the revision it was identified also in the collec-

tion of ISEA from the Beskid Makowski Range and from

Kolomyya town. The last one site was mentioned by

Dziedzielewicz (1918). The species seems to be extinct

in Western and Central Europe from the second half of

the XX century, but there are some frequent founds in

Eastern Europe (Fochetti & Tierno de Figueroa 2006).

That is why J. nubecula belongs to the category EN [en-

dangered] in the Carpathian Red List (Pawlowski 2003).

In 2006 I. nubecula was found for the first time for the

Transcarpathians in Ukraine from the Tysa, Teresva and

Rika rivers (Kovacs et al. 2008b; larval records).

Genus Perlodes Banks, 1903

Perlodes intricatus (Pictet, 1841)

Material. SMNH: UKRAINE: “Poturzyca przy Bugu’,

“Klapalek angusticeps”, LP (JD), 1°.

ISEA: UKRAINE: “Chomiak. Pot. Bogdan. 14.-

7.-1905. Dz.”, “80/’, GOR (JD), 12; “Kotomyja”,

“127/10”, CCA (JD), 19; “25/4 Kt”, “134/4”, CCA

(JD), 14; POLAND: “Kosciel. VIII’, “72/1”, T (JD),

12; “Koscielisko. Tatr. 1891. Dziendz.”, “124/10”, T

(JD), 12; “Koscielisko”, “124/10”, T (JD), 19; “Rytro”

“24/13”, SAB (FS), 22 9; “Rytro”, “24/”, SAB (FS), 19;

“Wad” “71/1”, PS (CU), 19; others: “24/13”, SAB (FS),

Oe 2h DIAL.

Remarks. In the paper published by Godunko &

Ktonowska-Olejnik (2003), material from SMNH is

mentioned as Perlodes intricata (Pictet, 1841).

The specimen mentioned as Perlodes dispar (Rambur,

1842) in the previous contribution on SMNH collection

(Godunko & Ktonowska-Olejnik 2003) belongs to P. in-

tricatus. That was the single specimen, mentioned by Dz-

iedzielewicz (1918) as P. dispar. We have identified fur-

ther specimens from Koscielisko [village in Malopolska

Province], although there was no mention about findings

of P. intricatus from the Tatry Mts. in J. Dziedzielewicz’

papers. In the Ukrainian Carpathians, besides the Chor-

nohora Range, P. intricatus 1s recently reported from the

Gorgany Range (Diakiv 2011).

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Identity of certain stoneflies from Ukraine and Poland with notes on available museum material from XTX—XX centuries 129

Perlodes microcephalus (Pictet, 1833)

Material. SMNH: UKRAINE: “Chomiak. pot. Bogdan.

11.-7.-1905. Dz.”, “2”, GOR (JD), 19; “Chomiak. pot.

Barani. 10.-6.-1907.”, GOR (JD), 19; “Chomiak. Blotek.

1.VI.1909.”, “Perlodes (Dictyopteryx) microcephala

Pict.”, GOR (JD), 19; “Kolomyja. 30.V.”, CCA (JD),

19; “Kotomyja. Prut.”, CCA (JD), 12.

ISEA: UKRAINE: “Chomiak. pot. Bogdan. 18.-7.-

1905. Dz.”, “80/20”, GOR (JD), 192; “Chomiak. pot.

Bogdan. 5.-7-.1905. Dz.”, “80/20”, GOR (JD), 19;

“Chomiak. pot. Baran1. 26.-6.-1905. Dz.”, “80/20”, GOR

(JD), 14; “2/4 KY’ “133/4”, CCA (JD), 14; “8/5 KY’,

“100/5”, CCA (JD), 13; “3/6 Mt.”, “99/5”, CCA (JD),

12; POLAND: “Myslenice. Stanistawow’, “70/1”, MAP

(JD), 49°: others. “98/5” (CU) 204; “133/4”, CCA

(JD), 22 9; “133/4, CCA” (JD), 33'S.

Remarks. There were determined specimens of P. mi-

crocephalus from ISEA collection. The findings belong

to the region of Gorgany and Ciscarpathians as those in

the Ukrainian part of the collection. By the latter inves-

tigations, P. microcephalus was found in the Ukrainian

Beskydy (Diakiv 2011) and in most of the Polish Car-

pathians (Fiatkowski & Kittel 2002).

Genus Isoperla Banks, 1906

Tsoperla grammatica (Poda, 1761)

Material. SMNH: UKRAINE: “Mikuliczyn. Polanica

5.VHI.1888”, GOR (JD), 12; “Kotomyja 12.V.1899.”,

CCA (JD), 13; “Tatarow. (Blotek.) 10.-7.-1905. Dz”,

GOR (JD), 14; “Mikuliczyn (Zeniec) 23.VII’”, GOR

(JD), 19; “Mikuliczyn (Zeniec).”, GOR (JD), 12; “Kolo-

myja Prut.”, CCA (JD), 322; “Kotomyja”, CCA (JD),

13; “Kotomyja. 30.V.”, CCA (JD), 14, 19; “Kolomyja.

5.VI.”, CCA (JD), 14; POLAND: “Tatry”, T (JD), 12.

ISEA: UKRAINE: “Czarnohora. Dancerz.

12.1X.1909”, “139/29”, CHOR (JD), 12; POLAND: “

Myslenice Oklejna. 19. VII.1909”, “166/24”, MAP (JD),

13; “Rytro”, “26/15”, SAB (FS), 14; others: “88/20”

(CU). LO. 13971 072 CID) ee 138/10: (CU), 1

Remarks. There were determined specimens from

the Chornohora Range within Ukrainian Carpathians

(Diakiv, 2010; 2011). Dziedzielewicz (1918) reported

about the presence of /. grammatica all over the Carpath-

ians as “Chloroperla grammatica Scop.”

Tsoperla rivulorum (Pictet, 1841)

Material. SMNH: UKRAINE: “Mikuliczyn. (Zeniec).”,

GOR (JD), 192; POLAND: “Babia Gora. Zawoja.

23.VII.1909.”, T (JD), 12.

Remarks. Earlier J. rivulorum was found also in the

Prut river-basin (Dziedzielewicz, 1891) and then it was

found on the other side of the Chornohora Range, 1.e., in

Bonn zoological Bulletin 71 (2): 119-137

the Keveliv stream (Tysa river-basin) during our investi-

gations (Diakiv 2011). In Poland the species is found in

the Western Sudety and Western Beskid Regions (Fial-

kowski & Kittel 2002). Due to distributional pattern of

this species, which is confirmed for the Alps but scarcely

known from Central and Eastern Europe, the question

about systematic position of all material mentioned as

I. rivulorum (e.g., in the Carpathians) should be clarified

in the future.

Isoperla sudetica (Kolenati, 1860)

Figs 14-16

Material. HNHM: UKRAINE: Zakarpatska Region,

Mizhhirskyi district, Krasna Mts., spring at a forest edge

in the upper valley of Kvasovec Stream, 48°22.79’ N,

23°42.86’ E, 1250 maz.s.l., 21.05.2002, leg. D. Murany1:

13.

ISEA: POLAND: “1891. Tatry”, “137/10”, T (JD), 1°.

Diagnosis. General coloration yellowish with dark

brown markings; head with horseshoe-shaped dark mark-

ing connecting ocelli, pronotum with large rugosities.

Vesicle of sternite VIII well developed, wider than long.

Both medial and lateral penial armatures with spike-like

scales; medial armature weakly notched medially, width

and length ratio 1.3.

Remarks. The species was described from the Czech

portion of the Eastern Sudetes (Kolenati 1860). Later it

was found also in the German and Polish areas of the

whole Sudetes (Reusch & Weinzierl 1999; Fiatkows-

ki & Kittel 2002), as well as from several localities in

the Czech, Slovak, Polish and Romanian areas of the

Carpathians (Kis 1974; Fiatkowski & Kittel 2002; Bo-

jkova & Soldan 2013; Ziak 2016).

Isoperla sudetica was reported for the Prut river-ba-

sin (Dziedzielewicz 1908). Nevertheless, no other speci-

mens in the collections studied, except the female imago

listed above. This specimen was reported from the Tatry

Mts. by Dziedzielewicz (1895) as “Chloroperla sudetica

Kol.”. Additionally, this author expressed some doubts

because he couldn’t distinguish /. sudetica from I. gram-

matica (cited as “Chloroperla grammatica Scop.”) be-

cause of the taxonomic doubts in that time. In Ukraine,

Dziedzielewicz (1918) reported the species from Cho-

miak Mt. and the Chornohora Range, but no subsequent

data were published. The species was included in the lat-

est checklist of the Ukrainian Carpathians (Diakiv 2011),

but excluded even from the monograph of the former

Soviet Union (Teslenko & Zhiltzova 2009). Occurrence

of the species out of the Sudetes and the Carpathians is

questionable. The single Alpine data from Austria could

not be proved (Graf 1999), and the species was recently

reported from southern Serbia, being the single Balkan

report of the species (Petrovic et al. 2014).

A related, yet unnamed taxon was reported and brief-

ly illustrated from the nearby Ignis Plateau, Maramures,

©LIB

130 Khrystyna I. Arkhipova et al.

te.

Figs 14-16. [soperla sudetica (Kolenati, 1860), 9 [Poland] and 3 [Ukraine]. 14. Head and pronotum. 15. Male terminalia, ventral

view. 16. Penial armatures. The specimens are deposited in ISEA and HNHM collections. Scale bars: 14-15 = 1mm; 16 =0.5 mm.

Romania (Muranyi 2006). Our Ukrainian specimen mor-

phologically agrees with /. sudetica specimens from the

Northern Carpathians, and differs from the unnamed tax-

on from the Ignis by paler coloration, and less widened

medial penial armature with width and length ratio below

1.5 (Figs 14-16).

Tsoperla spp.

(damaged and incomplete specimens)

Material. JSEA: UKRAINE: “Czarnohora. Dan-

cerz.12.-IX.1909”, “139/24”, CHOR (JD), 2¢4; “Tar-

tarow. (Blotek.) 8.-7.-1905. Dz”, “86/20”, GOR (JD),

1°; “Tatarow. (Blotek.) 29.-6.-1905. Dz.”, “86/20”,

GOR (JD), 1?; “Jablonica za Tartarowem. 18.-7.-1905.

Dz.”, “87/20”, CHOR (JD), 1?; “Chomiak. pot. Bogdan.

18.-7.-1905. Dz”, “85/20”, GOR (JD), 12; POLAND:

“Tatry 8/27, “139/10”, TD), 1?y “1891. Tatry.: Dzier,

“137/10”, T (JD), 12; “dol. Bialego.”, “136/10”, T (CU),

1?; “Babia Gora. Zawoyja. 25. VII. 1909.”, “138/24”, T

(JD), 12; “Myslenice. Stréze. 18.VII.1909.”, “146/24”,

MAP (JD), 1°.

Also there are representatives of Perlodidae spp. that are

impossible to identify due to damages:

ISEA: UKRAINE: “Chomiak. pot. Barani. 3.-7.-1905.

Dz.” “82/20”, GOR (JD), 1?; “Chomiak. pot. Barani. 26.-

6.-1907. Dz.” “82/20”, GOR (JD), 1?; POLAND: “Ry-

tro”, “24/13”, SAB (JD), 1°.

Bonn zoological Bulletin 71 (2): 119-137

Family Chloroperlidae Okamoto, 1912

Genus Siphonoperla Zwick, 1967

Siphonoperla neglecta (Rostock, 1881)

Material. SMNH: UKRAINE: “Chomiak. (Blotek.) 13.-

7.-1905. Dz.”, GOR (JD), 14; “9.VI.1904. Chomiak

Mikuliczyn”, GOR (JD), 192; “Chomiak. [Btotek] 14. VI

1909”, GOR (JD), 23'¢; “20.7. Zeniec. Mikuliczyn”,

GOR (JD), 14; “24.7. Mikuliczyn pot. Zeniec”, GOR

(JD), 13.

ISEA: 13: “148/17” (CU).

Remarks. The species was mentioned by Godunko &

Ktonowska-Olejnik (2003) from the Prut basin based on

investigation the specimens from SMNH collection (see

above). Besides the Prut river-basin, S. neglecta is also

known from Tysa river-basin (Zhiltzova 1968). In the

Polish Carpathians, S. neglecta is known from the West-

ern Beskid and Tatry Mts. (Fiatkowski & Kittel 2002).

Siphonoperla transsylvanica (Kis, 1963)

Figs 17, 19

Materials. HNHM: UKRAINE: Zakarpatska Region,

Tyachivskyi district, Krasna Mts., right tributary of

Luzanka River, 48°21.90’N 23°42.60’°E, 870 m as.l.,

22.05.2002, leg. D. Muranyi: 14, 29 9; Mizhhirskyi dis-

trict, Krasna Mts., Kolochava, Tereblya River above the

village, 48°25.35’N 23°41.58’E, 545 maz.s.1., 23.05.2002,

leg. D. Muranyi: 22 9, 1 exuvium.

SMNH: UKRAINE: “20. VII.1909. p. Bogdan ad Cho-

miak. Mikuliczyn 1909.”, GOR (JD), 19; “Stobddka les-

na 25.5”, CCA (JD), 14; “ Stobodka lesna 4.6.”, CCA

(JD), 13.

Remarks. The species was described from the Ro-

manian Carpathians (Kis 1963b), and soon reported

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Identity of certain stoneflies from Ukraine and Poland with notes on available museum material from XIX—XX centuries 131

also from the Eastern Carpathians of Slovakia (RauSer

1964) and Hungary (Ujhelyi 1969). Similar to Leuctra

carpathica, the species seems to be lacking from the

Northern Carpathians (Ziak 2016) but is widespread in

the Eastern and Southern Carpathians (Kis 1974). Out of

the Carpathians, it is known from the Bulgarian parts of

the Stara Planina (Braasch & Joost 1971), eastern Bosnia

and Herzegovina (Kacanski 1973), eastern and northern

Macedonia (Ikonomov 1986) and was recently reported

from western Serbia (Petrovic¢ et al. 2014). Due to mor-

phological similarity and allopatric distribution, S. trans-

sylvanica was Classified as a subspecies of S. torrentium

(Pictet, 1841) by Zwick (1971). Later, genetic analyses

proved rather high distance between S. torrentium tor-

rentium and two of its subspecies, thus S. transsylvanica

was reinstated as separate species (Weiss et al. 2012).

Siphonoperla transsylvanica had never been reported

from Ukraine; however, the vicariant S. torrentium was

reported several times (Dziedzielewicz 1918; Zhiltzova

1968; Diakiv 2010). Concerning their distribution, and

the lack of S. torrentium in eastern Slovakia (Ziak 2016)

and the whole of Romania (Kis 1974), previous Ukrainian

data must rely on S. transsylvanica. According to Zwick

(1971) and Weiss et al. (2012), morphological distinction

of the two taxa relies on the structure of penial scales

and length of titillators, while the original description

mentioned mostly coloration differences and the shape

of the male epiproct (Kis 1963b). Herein we illustrate the

color pattern differences between our Ukrainian S. trans-

sylvanica specimens and a S. torrentium torrentium pop-

ulation from the central Slovakian Polana Mts. (Hrifova,

Slatina Stream): our S. transsylvanica specimens lack

any dark pattern between the ocelli and the mesosternal

furcal pit is pale, while the S. torrentium specimens have

small dark markings around posterior ocelli and the fur-

casternal pit is distinct, dark brown (Figs 17—20).

Genus Xanthoperla Zwick, 1967

Xanthoperla apicalis (Newman, 1836)

Material. SMNH: UKRAINE: “Kolomyja 25.6.”, CCA

(JD) 13; “6. Kotomyja”, CCA (JD), 14; “Koltomyja w

Czerwceu”, CCA (JD), 14; “6 Kt.”, CCA (JD), 303.

Remarks. The specimens listed above are the only

ones for the Ukrainian Carpathians from old collections.

All investigators just cited Dziedzielewicz (1891, 1883,

1877) and Majewski (1885) records. By the latter inves-

tigations it was found in some regions of the Polish Car-

pathians (Fiatkowski & Kittel 2002). Larvae of X. api-

calis recently were found in the Beskids of Ukrainian

Carpathians (Diakiv 2011).

Chloroperlidae spp.

(damaged and incomplete specimens)

Material. JSEA: UKRAINE: “25/6 Kt”, “150/4”,

CCA (JD), 14; “Ag”, “21/3”, CCA (JD), 299; “Ag”,

“21/3”, CCA (JD), 14; “Mikuliczyn. 1 do 5. VIII. 1904”,

“124/19”, (JD), 14; “Tartarow. 1902.31.7”, “146/17”

GOR (JD), 19; “Tartarow. 1902.26.7. potudn. stoki Cho-

miaka”, “147/17”, GOR (JD), 13); “Tartarow. 15.7. Cho-

miak. 1903”, “123/19”, GOR (JD), 19; “Tartarow. 15.7.

Chomiak.”, “123/19”, GOR (JD), 192; “potudn. stoki

Chomiaka” “147/17”, GOR (JD), 14; POLAND: “Wad”,

“80/1”, PS (CU), 14; “Tatry”, “140/10”, T (JD), 14; oth-

ers: “3/7 Dr/?”, “109/5” (CU), 14; “8/6 Sh.” “148/4”

(CU), 36:6; (CU), 12.

Figs 17—20. Female adults of Siphonoperla transsylvanica (Kis, 1963) and S. torrentium torrentium (Pictet, 1841). 17. S. transsyl-

vanica, Ukraine [Tereblya], head and pronotum. 18. S. torrentium torrentium, Slovakia [Hrinova], head and pronotum. 19. S. trans-

sylvanica, Ukraine [Tereblya], meso- and metasternum. 20. S. torrentium torrentium, Slovakia [Hrinova], meso- and metasternum.

The specimens are deposited in NMP and HNHM collections. Scale bar: 1 mm.

Bonn zoological Bulletin 71 (2): 119-137

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132 Khrystyna I. Arkhipova et al.

Family Taeniopterygidae Klapalek, 1905

Genus Brachyptera Newport, 1849

Brachyptera braueri (Klapalek, 1900)

Material. ISEA: POLAND: “Rytro”, “28/13”, SAB

(FS), 33.

Remarks. Dziedzielewicz (1918) reported B. braueri

only for the Prut River near Kolomyya. The species be-

longs to the rare taxa in Ukraine and is known for the

country only from the beginning of 20" century. In the

Polish Carpathians the species 1s known only from the

Western Beskid (Fiatkowski & Kittel 2002).

Brachyptera monilicornis (Pictet, 1841)

Material. [SEA: POLAND: “28/13”, SAB (FS), 1¢.

Remarks. There is no mention about B. monilicornis

in J. Dziedzielewicz’ papers. In the Ukrainian Carpath-

ians this species is known for the upper part of the Prut

river-basin (Diakiv 2011). For Poland, Fiatkowski & Kit-

tel (2002) reported B. monilicornis for the Western Sude-

ty and Tatry Mts.

Brachyptera seticornis (Klapalek, 1902)

Material. SEA: UKRAINE: “23/4 Kf’, “157/4”, CCA

(JD), 19; “19/3 K¥’, ”157/4”, CCA (JD), 234; “8/5

KY’, “112/5”, CCA (JD), 19 [with ball of eggs on the

abdomen]; “Mykitynce, Stanistawow”’, “84/1”, CCA

(JID), 13; “Tartarow.1902.28.7. pod Chomiakiem”,

“151/17”, GOR (JD), 12; “Chomiak. pot. Barani. 29.-

7.-1905. Dz.”, GOR (JD), 19; “145/10”, T (JD), 12;

“Forzeszczynka. 15. II. 1909. Czarnohora.”, “135/24”,

CHOR (JD), 244; “Czarnohora. Forzeszczynka. 15. III.

1909”, “135/24”, CHOR (JD), 14’; POLAND: “1891 Ta-

try”, “145/10”, T (JD), 14; “Rytro”, “28/13”, SAB (FS),

14; “144/10” (CU), 19; “156/4” (CU), 343; “144/10”

(CU), 174, “159/4” (CU), 19.

Remarks. These specimens of B. seticornis, that have

been mentioned in the old literature (see e.g. Dziedziele-

wicz 1908), are confirmed for the collection. It was not

mentioned from the Dnister river-basin. In Poland it is

known from the Western Sudety and Western Beskid

Ranges, and Tatry Mts. (Fiatkowski & Kittel 2002). One

of these specimens is preserved with the ball of eggs on

the abdomen.

Brachyptera spp.

(damaged and incomplete specimens)

Material. ISEA: UKRAINE: “20/4, Kt”, “157/4”, CCA

(JD), 12; “Mykietyrice. Stanislawow”, “26/3 M”, “84/1”,

Bonn zoological Bulletin 71 (2): 119-137

CCA (JD), 12; POLAND: “Krak. 1/4”, “z.” (CU), 1?;

others. “144/10” (CU), 1?; “74/8” (CU), 1?.

Remarks. The specimen from “Mykietynce.

Stanistawow” can be B. trifasciata as this species was

mentioned from this region in papers (Dziedzielewicz

1891). Others specimens could belong to any species

of Brachyptera genus mentioned by J. Dziedzielewicz,

e.g., B. braueri, B. monilicornis, B. risi (Morton, 1896),

B. seticornis and/or B. trifasciata.

Genus Oemopteryx Klapalek, 1900

Ocemopteryx loewii (Albarda, 1889)

Figs 21-27

Material. [SEA: UKRAINE: “8/5 Ki.”, “114/5”, CCA

(JD), 29 9; “20/4 Kt.”, “158/40”, CCA (JD), 19; “112/5”,

CCA (JD), 1°; others: “146/10” (JD), 29 9 (Fig. 27).

Remarks. This species was considered to be extinct

for almost 100 years on the territory of Ukraine. Accord-

ing to Fiatkowski & Kittel (2002), the reliable records of

O. loewii from Poland (Beskid Zachodni: Beskid Sadec-

ki: Rytro) were published by Dziedzielewicz (1918) and

Wojtas (1962), but their information about presence of

the species in the Pieniny Range within Polish Carpathi-

ans is erroneous (Fiatkowski & Kittel 2002).

The precious specimens from Kolomyya_ (Iva-

no-Frankivska Region: Ukraine) were listed in the paper

of Dziedzielewicz (1918). Besides these specimens, the

finds from vicinity of Lviv (Kryvchytsi was a village at

far vicinity of the city; now it is vicinity of Lviv) are men-

tioned in the same paper. It 1s interesting that O. /oewii is

known mostly from larger lowland rivers (Soldan et al.

1998; Zhiltzova 2003). Nevertheless, both sampling sta-

tions mentioned by J. Dziedzielewicz are characterized

by the presence of the middle-sized cismountains part of

river (e.g., Prut River near Kolomyya town), and only

small streams around of Kryvchyts1.

Among the specimens studied by us, the wings have

some variability in venation. Most of the key characters

of the forewing venation are well preserved. However, Sc

is not so short as it described in Zhiltzova (2003), e.g., do

not reach the half of the wing length. Among the speci-

mens in ISEA collection, Sc reach 2/3 of the wing length

(Figs 23, 24). Two specimens have an additional c—r vein

instead of one (Figs 21, 22, 25, 26), and a single specimen

is characterized by the presence of RS branch divaricates

medially (Figs 23, 24). Additionally, one specimen has

four branches instead of three, and three anterior branch-

es divaricate (Figs 25, 26). Cu, has one additional branch,

however, the shape of this furcation differs from those

depicted by Zhiltzova (2003). The pterostigma between

C and R is obvious, that has not been mentioned before

(Figs 22, 24, 26).

©LIB

Identity of certain stoneflies from Ukraine and Poland with notes on available museum material from X-X—XX centuries 133

Figs 21-27. Oemopteryx loewii (Albarda, 1889), ° 2, Ukraine. 21, 23, 25. Forewings of separate specimens. 22, 24, 26. Body, fore-

and hind wings of separate specimens. 27. Original labels. The specimens are deposited in ISEA collections. Without any scales.

Genus Rhabdiopteryx Klapalek, 1902

Rhabdiopteryx harperi Vingon & Muranyi, 2009

Material. JSEA: UKRAINE: “Chomiak. Btotek.

2.V1.1909”, “136/24”, GOR (JD), 23'4, “85/1”, (CU),

re,

Remarks. The species was identified for the first time

from the old collections. Previously, this species was

published as Rhabdiopteryx alpina Kuhtreiber, 1934

from the Carpathians (see Vingon & Muranyi 2009). In

the Polish Carpathians, R. harperi is known from the

Bonn zoological Bulletin 71 (2): 119-137

Bieszczady, and in the Ukrainian Carpathians from the

Tysa river-basin.

Genus Taeniopteryx Pictet, 1841

Taeniopteryx nebulosa (Linnaeus, 1758)

Material. [SEA: POLAND: “Krakow 10/IV 86 Ogr.

bot”, “K. Jelski”’, “K. Jelski”, MAP (KJ), 14; “Oswiecim

5 Il nad Sota”, SG (CU), 14.

Remarks. 7aeniopteryx nebulosa is known from the

Prut and Dnister river-basins in Ukraine, and from the

©LIB

134 Khrystyna I. Arkhipova et al.

Western Sudety, Western and Eastern Beskid Ranges,

and Tatry Mts. in Poland (Fialkowski & Kittel 2002).

Taeniopteryx spp.

(damaged and incomplete specimens)

Material. ISEA: POLAND: “Wad”, “82/1”, PS (CU), 22:

others: “143/10”, (JD), 22; “144/10” (CU), 12; “156/4”

(CU), 12; “143/10” (CU), 12; (CU), 43.4.

Family Nemouridae Newman, 1853

Genus Amphinemura Ris, 1902

Amphinemura triangularis (Ris, 1902)

Material. ISEA: POLAND: “Myslenice. 11.VIII.1909”,

“125/24”, MAP (JD), 16.

Remarks. There was no mention about any of species

of the genus Amphinemura in J. Dziedzielewicz’ papers.

Most probably in his contributions Amphinemura was

identified as another genus/species.

Genus Nemoura Latreille, 1796

Nemoura cinerea (Retzius, 1783)

Material. ISEA: UKRAINE: “Tatarow. (Blotek.) 16.-

7.-1905. Dz.”, “91/’, GOR (JD), 192; “Podole okol Ra-

kutowej Kamieniecki”, WPOD (? JD), 14; POLAND:

“Krakow L1O/IV 86 Ogr. bot” “K. Jelski”, MAP (KJ),

12; “Jordanow 7.8.1921”, BM (JF), 19; “Jordanow

98.1921”, BM (JF) 12; “Kobierzyn 26.V.1921” MAP

GE) $12:

Remarks. This species is widespread all over Polish

and the Ukrainian Carpathians.

Nemoura spp.

(damaged and incomplete specimens)

Material. ISEA: UKRAINE: “22.6.1902 Biatohorszc-

Za. 8 ASST ROZ. ID) VO: RR Wd? CCA:

(JD), 12 [very damaged specimen], “Mykietyrce.

Stanislawow’”, “89/1”, CCA (JD), 1?; “Ag”, “22/3”, CCA

(JD), 12; POLAND: “Krak? Jablonski”, MAP (PJ), 1°29

[very damaged specimen]; “Kobierzyn 26.V.1921” MAP

(JF), 19; others: “152/4”, (CU), 13;

Genus Nemurella Kempny, 1898

Nemurella pictetii Klapalek, 1900

Material. [SEA: UKRAINE: “1.6.1902. Hotosko w.”,

“155/17”, “Nemurella Kemp. picteti Klap.”, “incospicua,

Pict.”, PH (JD), 33:4; “Hotosko, 1902.15.9” “155/17”,

“Nemurella Kemp. picteti Klap.”, “incospicua, Pict.”,

PH (JD), 1c. )

Bonn zoological Bulletin 71 (2): 119-137

Remarks. These specimens were collected near Lviv

City by J. Dziedzielewicz, but in his paper he mentioned

N. pictetii from the Svydivets Range. Despax (1935)

pointed it for the Chornohora Range within the Ukrainian

Carpathians. This species is considered to be ubiquist

(Zhiltzova 2003). In the Ukrainian Carpathians N. pic-

tetii was found in Tysa, Prut (Despax 1935; Zhiltzova

1966, 1967) and Seret river-basins in small streams with

gravel or sand bottom and in the alpine lakes (Diakiv

2011). Martynov et al. (2017) reported this species from

Eastern Ukraine in Starobilsk slope-elevated, West-Do-

netsk slope-elevated and Donetsk elevated physiographic

areas.

Genus Protonemura Kempny, 1898

Protonemura praecox (Morton, 1894)

Material. ISEA: UKRAINE: “Mikuliczyn. 1902.13.8.”,

“154/17”, GOR (JD), 10.

Remarks: There are no mentions about P. praecox in J.

Dziedzielewicz’ papers. Probably the species was identi-

fied as another taxon.

Protonemura spp.

(damaged and incomplete specimens)

Material. ISEA: POLAND: “Rytro”, “32/13”, SAB

(FS), 19; “118/5” (CU), 12.

Nemouridae spp.

(damaged and incomplete specimens)

Material. ISEA: UKRAINE: “22.6.1902. Biatohorszc-

ze.”, “153/17”, CCA (JD), 3?; “92 Miodiatyn”, “149/10”,

CCA (JD), 1?; “24/5 Kt”, “119/5”, CCA (JD), 1?; “12/7

Kt.”, “75(8)”, CCA (JD), 19; “25/5 K.”, “162/4”, (CU), 1?;

“dol. Biatego”, “148/10”, (?JD), 12; “Mykietynce. Sta-

nislawow’, “89/1”, CCA (JD), 3°; “Sopow’”, “150/10”,

CCA (CU), 3?; “Myslenice Oklejna. 14.VIII.1909”,

“127/27”, MAP (JD), 1?; “Ag”, “23/3”, CCA (JD), 4?;

“Jablonica za Tartarowem. 8.-7.-1905. Dz.”, “93/20”,

CHOR (JD), 1?; “Mikuliczyn.1902.21.7.”, “152/17”,

GOR (JD), 12; “Mikuliczyn. 1902.24.7.”, “152/17”,

GOR (JD), 1?; “Mikuliczyn. 1902.25.7.”, 154/17”, GOR

(JD), 12; “Chomiak. pot. Bogdan. 19.-7.-1905. Dz.”,

“92/20”, GOR (JD), 12; “Chomiak. (Barania.) [zro-

dto].5.-7.-1905. Dz.”, “94/20”, GOR (JD), 1?; “Woroch-

ta. Okolice. 13.-7.-1907”, CHOR (JD), 1?; “Czarnoho-

ra. Kozmieska. 2.-X.1909”, “133/24”, CHOR (JD), 1?:

“Czarnohora. Kozmieska. 2.-X.1909”, “124/24”, CHOR

(JD), 1?; POLAND: “Tatry 8/2”, “152/10”, T (JD), 1?:

Tatey. 8/2". 154/075: Te). 2052618: Zaks SO2 Ar

T (CU), 1?; “Rytro”, “31/13”, SAB (FS), 1?; “Rytro”,

“29/13”, SAB (FS), 1?; “161/4”, 12; “Wad”, “90/1”, PS

(CU), 12; “Wad”, “93/1”, PS (CU), 12; “Babia Gora. Za-

©LIB

Identity of certain stoneflies from Ukraine and Poland with notes on available museum material from XITX—XX centuries 135

woja. 25.VII.1909”, T (JD), 12; others: “151/10” (CU),

19: ARIS? (CU): 12“ 163/47. (CU) a 12 107/52 (CU),

12; “12/3 Wz.”, “160/4”, (CU), 12; “16/8 Bramka”,

SON? (CU). 12.

Further specimens of Plecoptera from Ukraine in the

collection of NMP

There are more than hundred Nemouridae, a few dozens

Leuctra and several /soperla specimens from Ukraine

left unrevised in the collection of the NMP. Most of these

are lacking identification labels, and are pinned after se-

ries of common species like N. cinerea, or simply stored

among unidentified materials. Since the F. Klapalek col-

lection arrived to the NMP in chaotic order after World

War II (Muranyi & Li 2015), most of the old Ukrainian

specimens should be originally identified and readily

published by J. Dziedzielewicz or F. Klapalek. Re-identi-

fication of these pinned specimens will need considerable

further efforts, and since no possible types are included,

we postponed this work.

After revising materials of the old collections, species,

whose occurrence in Ukraine has been questioned, have

been confirmed. The revised material contains informa-

tion about localities that were not mentioned in the old

publications. The brief remarks on these findings based

on redetermined collections are listed in the article.

Acknowledgements. We are very indebted to Pavel Chvojka

(NMP) and Andrzej Palaczyk (ISEA) for help during works

in museum collections. The research was supported by the

SYNTHESYS Project and FP7 “Capacities” Program (CZ-

TAF-3636) for DM; RJG acknowledges the Czech Academy

of Sciences for institutional support RVO: 60077344 and the

Alexander von Humboldt Foundation (Bonn, Germany) for Re-

search Fellowship for Experienced Researchers.

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BHL

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LIB. Leibniz Institute for the Analysis

=— of Biodiversity Change

Bonn zoological Bulletin 71 (2): 139-176

2022 Hazmi IR & Wagner Th.

forsewunas

museum

: KOENIG

https://do1.org/10.20363/BZB-2022.71.2.139

Research article

urn:lsid:zoobank.org:pub: 7FFF8A 8D-1640-4565-A9DE-239D1CD7FS50A

Revision of Monolepta Chevrolat, 1836 from the Sundaland area

(Coleoptera: Chrysomelidae: Galerucinae)

Izfa Riza Hazmi ©' & Thomas Wagner”

‘Center for Insect Systematics, School of Environmental and Natural Resource Sciences, Faculty of Science and Technology,

Universiti Kebangsaan Malaysia, 43600 Bangi, Malaysia

?Universitat Koblenz-Landau, Institut fiir Integierte Naturwissenschaften - Biologie, Universitatsstr. 1, D-56070 Koblenz

“Corresponding author: Email: thwagner@uni-koblenz.de

54. contribuion to the taxonomy, phylogeny and biogeography of the Galerucinae

'urn:lsid:zoobank.org:author:CB54FDOA-CF4C-4972-8C6F-F2743F BEBB8B

2urn:Isid:zoobank.org:author:6A F65C9C-246B-41CA-B326-2D93C427CCD1

Abstract. Species of Monolepta Chevrolat, 1836 currently known from the Sundaland area are taxonomically revised.

The region covers the states of Malaysia, Singapore, Brunei and Indonesia including the Wallacea-Subregion (without

Irian Jaya on New Guinea). About 2300 specimens, mainly from museal collections, from this area are checked, all

relevant literature is considered and the type material is examined. Out of 72 species originally described in Monolepta

and occurring in this region, only 13 valid species (including five new synonymies) can remain in Monolepta when

phylogenetical aspects are considered: M. bifasciata (Hornstedt, 1788) (Chrysomela bifasciata Hornstedt, 1788; = Cryp-

tocephalus multicolor Gmelin, 1790; = Crioceris quadrinotata Fabricius, 1801; = Luperodes latefascia Motschulsky,

1858; = M. parvonotata Jacoby, 1886, syn. nov.; = M. mustaphai Mohamedsaid, 1997, syn. nov.; = M. entimauensis

Mohamedsaid, 1998, syn. nov.); — M. flavofasciata Jacoby, 1889.— M. mentaweiensis (Jacoby, 1896) (= M. basalis Jaco-

by, 1884; = M. hageni Weise, 1916, syn. nov.). — M. jacobyi Weise, 1908. — M. marginicollis Jacoby, 1896. — M. militaris

Jacoby, 1896. — M. orientalis Jacoby, 1889 (= M. konbiriensis Duvivier, 1891). —M. putri Mohamedsaid, 2001.—M. rubra

(Gyllenhal, 1808) (= Luperodes javanensis Jacoby, 1887). — M. rufipennis Jacoby, 1899. — M. signata (Olivier, 1808)

(Galeruca signata Olivier, 1808; = Crioceris neglecta Sahlberg, 1823: 72; = Luperodes quadripustulatus Motschulsky,

1858; = Luperodes hieroglyphicus Motschulsky, 1858; = M. elegantula Boheman, 1859: 183; = Luperodes dorsalis Mot-

schulsky, 1866: 415; = Luperodes quadriguttata Fairmaire, 1887: 333; = M. picturata Jacoby, 1896; syn. nov.; = M. sim-

plex Weise, 1913). — M. tiomanensis Mohamedsaid, 1999. — M. zonula Weise, 1916. Other species originally described

in Monolepta occuring in the region are still or need to be transferred to other genera. Eight species are newly described:

M. bruneiensis sp. nov., M. empatbulat sp. nov., M. hitam sp. nov., M. kuninghitam sp. nov., M. marginicolloides sp. nov.,

M. mohamedsaidi sp. nov., M. ranuensis sp. nov., M. sulawensis sp. nov. An identification key for the 21 species, a check-

list of the current taxa of Monolepta from Sundaland, and a list of species originally described in Monolepta occurring in

this region that still are or need to be transferred to other genera are provided.

Key words. Taxonomy, nomenclature, new species, redescription, new synonym, lectotype, distribution maps, identifica-

tion key, Oriental Region.

ISSN 2190-7307

http://www.zoologicalbulletin.de

INTRODUCTION

Monolepta Chevrolat, 1836 is the most species rich group

in Galerucinae, and has a wide distribution in tropical and

subtropical regions of the world. Wilcox (1973) has listed

600 species worldwide. At least 260 nominal species are

recorded from the Oriental Region, and about 72 species

occur in Sundaland area (Kimoto 1990; Mohamedsaid

2001, 2004, 2005). Particularly in the Oriental Region,

the group has been studied by several authors (Maulik

1936; Gressitt & Kimoto 1963; Kimoto 1989; Mohamed-

said 2004). For many years, the limitation and taxonomic

status of Monolepta were uncertain. Maulik (1936) for

instance realized this problem but decided to maintain the

Received: 19.01.2022

Accepted: 21.09.2022

status quo and provided two sets of keys for Monolepta;

one for species with a pronotal depression and another

one without pronotal depression. Wilcox (1973) has also

indirectly commented about the inhomogeneous classifi-

cation and systematic of Monolepta and some other gen-

era in “Monoleptites” and proposed to revise this group

in the future.

Despite so many species that have been subsequently

described in Monolepta, a revision, taking all described

Species into account, has never been done so far. A revi-

sion of the Afrotropical taxa was started in Wagner (1999),

and more recent data were provided in Nie et al. (2017),

where 708 species are listed. With the revision on the

genotype, M. bioculata (Fabricius, 1781), a new concept

Corresponding editor: D. Ahrens

Published: 28.09.2022

140 Izfa Riza Hazmi & Thomas Wagner

of the genus taking phylogenetic aspects into account,

was given (Wagner 2004, Stapel et al. 2008). As a result

of the revision, many changes in number of species and

allocation of species to genera in Afrotropical taxa was

published, and out of 180 species originally described in

Monolepta from Africa, only 50 valid species remained,

40 species were treated as, mainly new, synonyms, and

90 species needed to be transferred to other genera, due

to their phylogenetic position. On the other hand, 50 new

species were described (Wagner 2007, 2017).

The ongoing revision of Afrotropical taxa (overviews

in Wagner 1999, 2004, 2017) was not only the first step

taken in revising Monolepta, on a larger geographical

scale, but as well the first step in consequently describing

the genitalic characters for all taxa of Monolepta. The

previous authors have, with very few exceptions, de-

pended on external morphology and colouration pattern

only, but in particular the colouration pattern is highly

variable within many species, and in most cases not suit-

able for species identification.

A revision of Monolepta on a worldwide scope is a

gigantic task. Revision of species from a selected geo-

graphical area can be a step in the right direction. The

Oriental Region consists of the Indian subcontinent in-

cluding Pakistan, south of the Pamir and the Himalayas,

Myanmar (formerly Burma), Indochinese and Malay

Peninsulas, Indonesia including Malay Archipelago (for-

merly the East Indies) to Timor and Sulawesi (formerly

Celebes), the Philippines, Taiwan, southern China from

the Tsinling Mountains and the Tibetan Scarp and south-

ern Japan. Some recent local contributions with focus

to the fauna of Monolepta were published from Taiwan

(e.g., Lee 2009, 2018, 2020), the Philippines (Medvedev

2005), Vietnam (Nguyen & Gomez-Zurita 2017), and

China (Lei et al. 2021).

Due to the high number of Monolepta from the entire

Oriental Region, we focused on the species of “true”

Monolepta known from the Sundaland in this work. This

area comprises Malaysia (Peninsular Malaysia, Bor-

neon states), Brunei, Singapore and Indonesia without

Irian Jaya, that definitely belongs to the Australian Re-

gion, while Sulawesi and all other islands of Wallacea,

as intermediate area between the Oriental and Australian

Region (Lohmann et al. 2011, Mohammedsaid 2009) are

included here. There are 72 species originally described

in Monolepta recorded in this region.

In the course of our revision, only 13 valid species

can remain in Monolepta in the sense of the generotype

M. bioculata (Wagner 2007), next to 13 synonyms recog-

nized by previous authors (see also Wagner & Bieneck

2012), we found further five new synonyms (for these

taxa. These 13 taxa of “true” Monolepta are herein rede-

scribed, further eight are newly described. A key to spe-

cies, current checklist of valid species of Monolepta from

Sundaland, and a list of species from this region that need

to be transferred to other genera, are provided.

Bonn zoological Bulletin 71 (2): 139-176

MATERIALS AND METHODS

A total of 2375 labelled specimens from several major

collections have been examined in this study. Fresh ma-

terial 1s also included, but the number is far more little

than dried museal specimens. The acronyms of collec-

tions and institutions that are involved in this study are

explained below and responsible curators are given in

parentheses.

The Natural History Museum, London (BMNH; M.

Barclay, M. Geiser); Brigham Young University collec-

tion, Provo, Utah (BYUC; Shawn Clark); Collection of

Jan Bezdék, Brno, Czech Republic (CJB); Collection

of Haruo Takizawa, Japan (CTJ); Instituto de Investi-

gacao Cientifica Tropical, Lissabon, Portugal (ICTZ;

L. Mendes); Institute Royal des Sciences Naturelle de

Belgique, Brussel (IRSN; P. Limbourg); Museo Civico

di Storia Naturale, Genova (MCGD; R. Poggi); Museum

of Comparative Zoology, Harvard University (MCZH);

Museum fiir Naturkunde, Berlin (MNHU; J. Frisch, J.

Willers); Swedish Museum of Natural History Stock-

holm (NHRS; J. Bergsten, B. Viklund); Naturkundemu-

seum Erfurt (NME; M. Hartmann); Nationaal Natuurhis-

torisch Museum, Leiden (RMNH:; F. van Assen); Centre

for Insect Systematics, UKM, Malaysia (UKM; R. Yu-

sop); Zoological Institute St. Petersburg (ZISP; A. Kire-

jtshuk); Zoological Museum of Kiel University (ZMUK;

M. Kuhlmann).

A standard set of figures is given for each species, scale

line in figures is 1 mm, different scales for colouration

pattern and genital structures and basal antennomeres.

These include illustrations of the colouration (dorsal

view), including the right antenna, where black coloura-

tion 1s indicated by black, yellow colouration by white,

red colouration by light grey, and brown by dark grey

shading. The antennomeres, usually of one male and

one female, dorsal, ventral and lateral view of the medi-

an lobe including endophallic structures, spermathecae

of up to three females, and bursa-sclerites (if applica-

ble) are figured. Measurements were made for external

characters. Absolute measurements are total length from

the clypeus to apex of the elytron, length of the elytron,

maximal width of both elytra (usually in the middle or

posterior third of the elytra), and width of the pronotum.

Relative measurements are length to width of the pro-

notum, maximal width of both elytra to length of the el-

ytron, length of the second to third antennomeres, and

length of the third to fourth antennomeres. A number of

specimens measured are given in the description under

“total length”. Further materials examined are listed, and

all label data are re-written. Material is arranged by the

states in alphabetical order, and in chronological order

within states. For location data, geographical coordinates

were given in degree and minute. These coordinates were

mostly taken from Google Earth. The distribution maps

have been produced by ArcGis.

©LIB

Revision of Monolepta Chevrolat, 1836 from the Sundaland area 141

Jan Bezdék generously provided us with photographs

of some type specimens from MCGD and NHRS he did

during visits in those collections. Labels of those speci-

mens are cited literally, but the photographs are not given

here.

REDESCRIPTION OF SPECIES

Monolepta bifasciata (Hornstedt, 1788)

Figs 1-6

Chrysomela bifasciata Hornstedt, 1788: 3 (transferred to

Monolepta; Jacoby 1884a: 53).

= Cryptocephalus multicolor Gmelin, 1790: 1712 (syn.

by Weise 1924: 166).

= Crioceris quadrinotata Fabricius, 1801: 460 (syn. by

Weise 1924: 166).

= Luperodes latefascia Motschulsky, 1858: 104 (syn. by

Maulik 1936: 408)

= Monolepta parvonotata Jacoby, 1886: 97; syn. nov.

= Monolepta mustaphai Mohamedsaid, 1997: 204;

syn. nov.

= Monolepta entimauensis Mohamedsaid, 1998: 248:

syn. nov.

Type material. Chrysomela bifasciata. Type material is

not available to us. We adopt Weise’ s (1924) statement

that Crioceris quadrinotata Fabricius, 1801 is a junior

synonym of this species, which type material is available.

Cryptocephalus multicolor. Type material is not avail-

able to us. We adopt Weise’ s (1924) statement that this

Species is a junior synonym of M. bifasciata (Hornstedt,

1788).

Crioceris quadrinotata. Syntype: 1 ex. “C. 4-notata

Sumatra Daldorff / Type” (ZMUK; Fig. 0).

Luperodes latefascia. Lectotype: 3, “Luperodes late-

fascia Motsch., Ind. Or” (ZISP; Fig. 1). Paralectotypes,

4 ex., same data as lectotype (ZISP). All five syntypes

were originally glued tightly together on one card. The

lectotype, the male paralectotype and one female para-

lectotype are now each on one card, two female paralec-

totypes together on one card (Wagner & Bieneck 2012:

Palle

Monolepta parvonotata. Syntype: °, “parvonotata Jac.

(Monolepta) / Type 18453 / Sumatra, Mt. Singgalang,

Luglio 1878, O. Beccari / 1% Jacoby Coll.” (MCZH;

Fig. 2). Type locality: 0°23’ S/100°19' E. Jacoby gave

no details on specimen numbers and next to the one in

MCZH, there are further types in BMNH and MCGD.

There are two specimens from the Jacoby collection in

BMNH labelled ’co-type”. These are invalid types since

they are not from the type locality.

Monolepta mustaphai. Holotype: 3 “Sabah, Pulau

Banggi, 8-12. Mei 1996, Salleh, Zaidi, Ismail & Sham /

Holotype Monolepta mustaphai n. sp. des. Mohammed-

Bonn zoological Bulletin 71 (2): 139-176

Figs 0-1. Type material of Monolepta bifasciata (Hornstedt,

1788). I. 1. Crioceris quadrinotata Fabricius, 1802, syntype

(ZMUK). 1. Luperodes latefascia Motschulsky, 1858, lecto-

type, & (ZISP).

said 1997” (UKM: Fig. 3). Paratypes. 1 3,2 2, same data

as holotype (UKM). Type locality: 7°16' N/117°09' E.

Monolepta entimauensis. Holotype: & “Sarawak: Lan-

jak Entimau, 28-29 Feb. 1992, Zaidi / Holotype Mono-

lepta entimauensis n. sp., des. Mohamedsaid 1998”

(UKM: Fig. 4). Paratypes: 3 4, 4 9, same data as holo-

type (UKM). Type locality: 1°11’ N/111°51' E.

SADA: Pulau Man

A Mel i996 2 sa

Salleh. Amiditsmail& Shen

Tae Sy re

Figs 2—4. Type material of Monolepta bifasciata (Hornstedt,

1788) IH. 2. Monolepta parvonotata Jacoby, 1886, syntype,

Q (MCZH). 3. Monolepta mustaphai Mohammedsaid, 1997,

holotype, 4 (UKM). 4. Monolepta entimauensis Mohammed-

said, 1998, holotype, 3 (UKM).

©LIB

142 Izfa Riza Hazmi & Thomas Wagner

Note. Weise (1924: 166) also synonymized Monolepta

rubrosignata Boheman, 1859: 182 with M. bifasciata. As

photos of a syntype from NHRS (“Manilla / Kinb. / Type

/ rubrosignata Bhm.,) clearly indicate, this species is not

a Monolepta and belong most likely to Nadrana Baly,

1865.

Further material examined. — Australia. | ex., N.

Queensland, 4.XII.1938, Redlynch (BMNH); 2 ex., Qid.

Tully, Mission Beach, 23.VIII.1976, D. C. Geijskes

(RMNH). — Bangladesh. 1 ex., Bengal, Mandar,

23°41' N/90°21' E, VII.91, Cardon (IRSN); 2 ex., Bel-

gaum, 15°51' N/74°30' E (BMNH); 3 ex., Bengal, Sarda,

23°41' N/90°21' E, F. W. C (BMNH). — Cambodia. 3 ex.,

Prek Toal (Tonie Sap Lake), 12°55' N/104°03' E,

27.V.2003, Light trap, J. Constant et al. (IRSN). — East

Timor. 29 ex., Raimundo, Timor, No 549 (ICTZ). — In-

dia. 7 ex., Jawanlagiri, Ayur, Denkanikota, North Salem,

12°34' N/77°47' E, T.—VII.1930, F.R.I. Sandal, Insect

Survey (BMNH); 1 ex., Shimoga, 1865 ft., 13°55' N/

75°34' E, 25.V.1936, P. S. Nathan (BMNH); 10 ex., Nil-

giri Hills, Cherangode, 11°25' N/76°30' E, II.1950, Susai

Nathan P. (IRSN); 2 ex., Coimbatore, 11°01' N/76°58' E,

2.[X.1950, P. Susai Nathan (IRSN); 17 ex., S. Coorg-Am-

manit, 3100 ft., 12°25' N/75°44' E, 1.—X1.1952, P. S. Na-

than (IRSN); 4 ex., Anamalai Hills, Chinchona, 3500 ft.,

V.1964, P. Susai Nathan (RMNH); 1 ex., Anamalai Hills,

Cinchona 350 ft., [1V.1967, P. S. N (MNHU). — Indone-

sia. | ex., Java, Batavia, 6°12' S/106°50' E, Grabowsky

(MNHU); 2 ex., O. Borneo, Pajau River, Myéberg

(NHRS); 3 ex., Flores, 8°03' S/120°09' E, Wallace

(BMNH); 4 ex., O. K. Sumatra, Dolokmerangir, coll. V.

Eldik (RMNH),; 10 ex., Java, Propeoek-Tegal, [X.1909,

Valek Lucassen (RMNH); 1 ex., Palembang, Sumatra,

2°59' S/104°45' E, M. Knappert (RMNH); 2 ex., Java,

Fig. 5. Monolepta bifasciata (Hornstedt, 1788). A. Colour pat-

terns. B. Basal antennomeres. C. Median lobe, a. dorsal, b. lat-

eral, c. ventral, without endophallic structures. D. Spermathe-

cae. E. Bursa-sclerites, a. dorsal, b. ventral.

Bonn zoological Bulletin 71 (2): 139-176

Delanggoe, W. Grippeling; N. C. Pilpers (RMNH):

16 ex., Sumatra, Deli, 3°35’ N/98°39' E, d Bury (RMNH);

1 ex., Semarang, 6°58’ S/110°25' E, Coll. Veth (RMNH);

1 ex., Bandong, 6°54' S/107°36' E, Coll. Veth (RMNH);

1 ex., Lombok, 8°39' S/116°19' E, Coll. Veth (RMNH);

1 ex., Celebes, 1°50' S/120°31' E, Coll. Veth (RMNH);

8 ex., Soerol, 0°35' S/101°20' E, Coll. Veth (RMNH);

1 ex., Maero Laboe, Coll. Veth (RMNH); 1 ex., Soepa-

jang, 0°27' S/100°54’ E, Coll. Veth (RMNH); 2 ex., N. O.

Sumatra, Tandjong Morawa, Serdang, 0°35' S/101°18' E,

Dr. B. Hadgen (RMNH); | ex., Rawas, 1°11’ S/132°13'E

(RMNH); 1 ex., Nias, 1°07' N/97°31' E, J. D. Pasteur

(RMNH),; 8 ex., Sumatra, Padang, 0°57' S/100°21' E, J.

D. Pasteur (RMNH); 2 ex., Serdang, Sumatra,

0°55' S/102°4' E, Schag (RMNH); 3 ex., Boengamas,

Palembang, 2°59' S/104°45' E, J. C. Hasselt (RMNH);

1 ex., Batavia, 6°12' S/106°50' E, Semme Link (RMNH);

1 ex., Soekaranda, Kampong, 0°37' N/94°29' E, J. C. van

Hasselt (RMNH); 3 ex., Java, Gedeh (MNHU); | ex., S.

O. Borneo, Grabowsky (MNHU); 2 ex., Sumatra, Pagh-

erang Pisang, X.1890.—III.1891, E. Modigliani (BMNH);

4 ex., Sumatra, Balighe, X.1890—XII.1891, E. Modigli-

ani (BMNH); | ex., Sumatra, Si Rambe, 6°11'S/106°48' E,

XII.1890.—-1.1891, E. Modigliani (BMNH); 2 ex., Java

occident, 4000 ft., 1893, H. Fruhstorfer (BMNH); 29 ex.,

Sumatra, Manna, 4°30’ S/102°58' E, 1902, M. Knappert,

Coll. Veth (RMNH); 2 ex., Malabar, Fry Coll. 27.V.1903,

light trap, J. Constant et al. IRSN); 1 ex., G. Oerangan,

Java, III.1906, Drescher (MNHU); 1 ex., N. O. Sumatra,

Prov. Langkat, 1906, E. Heinze (MNHU); 1 ex., W. Su-

matra, Kambang, 23.—24.XI.1908, Schoede (MNHU);

10 ex., Java, Bantam, Preanger, de Vos, [X.1909, Valck

Lucassen (RMNH); 3 ex., Sumat., Matur. Fort De Kock,

X.1913, Edw. Jacobson, Coll. Veth (RMNH); 1 ex., Su-

matra, Sungai Penok, Korinchi Valley, 2600 ft., III.1914

(BMNH); | ex., Mt. Banahao, 20.VI.1914, G. Boettcher

(MNHU); 2 ex., Java, Preang, Tjigembong, VIII.1915, B.

Corporaal (RMNH); 3 ex., Medan, 3°30' N/98°37' E,

1918, F. C. van Heurn (RMNH); 3 ex., Java, 1921, Madi-

oen, Paroembangan, A. E. Kerkhoven (RMNH); 2 ex.,

Sumatra, Fort de Kock, 920 m, III.1921, E. Jacobson

(BMNH); 1 ex., Java, Pelabokan Patoe, 1923, A. E.

Kerkhoven (RMNH),; | ex., Sipora Island, West Sumatra,

2°13' S/99°40' E, X.1924, C. B. K & N. S (BMNB);

2 ex., Sumatra exp., 1927, Rensch (MNHU); 1 ex., Sun-

da—Exp., Semongkat Batos-Lanteh-Gbg., N—Hang,

8°36' S/117°212' E, 400 m, 10.1.1927, Rensch (MNHU);

9 ex., Sunda, Exp. Lombok, Narwada, 14.11.1927,

Rensch (MNHU); 2 ex., O. Soembawa, Dormpoe, 24.—

25.V.1927, Rensch (MNHU); 1 ex., Sunda-Inseln, West

Flores, VI.1927, Rensch (MNHU); 3 ex., Sunda, Exp. O.

Flores, Geli Moetoe, 14.—20. VII.1927, Rensch (MNHU);

4 ex., Sunda-Ins., W-Flores, Rana Mese, 20.—30. VI. 1927,

Rensch (MNHU); | ex., M. Flores, Rana Mese, 20.—30.

XI1.1927, Rensch (MNHU); 13 ex., West Lombok, Nar-

mada, 8°34' S/116°11' E, 14-20.]I1.1927, Rensch

©LIB

Revision of Monolepta Chevrolat, 1836 from the Sundaland area 143

(MNHU); 2 ex., Lombok Segare, 8°22’ S/116°10' E,

5.VI.27, Rensch (MNHU); 2 ex., Fraserpet, Coorg., F. R.

I. Sandal, H.1930, Insect Survey (BMNH),; 6 ex., Java, G.

Oengaran, 31.[X.1933, P. H. V. Doesburg (RMNH);:

2 ex., Ambon Island, 3°40' S/128°10' E, 24.11.1964, A.

M. R. Wegner (RMNH); 6 ex., North Sumatra, Alas Val-

ley, Gumpang, 3.77°N /97.5°E, 11.V1.1972, J. Krikken

(RMNH); 25 ex., North Sumatra, Toba plateaue, Ti-

gadolok, Holzweg Drei, 2°50' N/99°3' E, 20.VI.1972, J.

Krikken (RMNH); 1 ex., North Sumatra, Bivouac Two,

Mt. Bandahara, 3°44' N/97°43E, 5—10.VII.1972, J.

Krikken (RMNH); 3 ex., North Sumatra, Alas Valley,

Kutatjane, Tanah Merah; Balelutu, 3°31' N/97°47' E,

3-9. VIII.1972, J. Krikken (RMNH); 7 ex., Java, Sema-

rang, 6°58’ S§/110°25' E, 1973, P. H. V. Doesburg, E. R

Jacobson (RMNH); 1 ex., Taronggo, 1°45’ S/121°40' E,

27 —30.111.1980, M. J. D. Brendell (BMNH); 28 ex., Su-

lawesi Tengah, Nr. Morowali, Ranu Lakes,

6°14' S/106°49' E, 27.1.—28 IV.1980, M. J. D. Brendell

(BMNH); 1 ex., North Central Sulawesi, 1°25' S/

121°22' E, V.1980 (BMNH); | ex., Indonesia, Sulawesi

Utara, Dumoga Bone NP, site 14 1140 m, 7—8.III.1985,

J. D. Holloway (BMNH); 1 ex., N. Sulawesi, Dumoga

Bone NP between base camp and Mopuya, 1°50' S/

120°31' E, 16. VHI.1985, J. Huijbregts (RMNH); | ex.,

Indonesia, Sumatra Utara, 15 km S. Brastagi,

3°22' N/98°34' E, 21.11.1992, B. Gustafsson et. al

(NHRS); 1 ex., Sumatra Utara, Aekpopo alt. 1600 m,

13. JI1.1995, Maruyama (CTS); 1 ex., Irian Jaya, Wamena

S, 1700 m, 4°12' S/139°01' E, 1.1999, A. Weigel (NME);

2 ex., Java (MNHU); | ex., Indonesia, Sumatra Utara,

Silalahi, 4.V.1999, S. Tsuyuki (CTJ); 14 ex., Sumatra,

Riau, Bukit Tigapuluh NP, 0°50' S/102°26' E, 1.2000, J.

Bezdék (CJB). — Malaysia. 11 ex., Borneo, Long Na-

vang, 3°35' N/116°38' E, Mjoberg (NHRS); 1 ex., Malac-

ca, 2°12' N/102°15' E, Coll. Chapuis (IRSN); 3 ex., Per-

ak, 4°48' N/100°48' E, Sharp Coll. (BMNH); 16 ex.,

Penang, Bowring (BMNH); | ex., Nord-Borneo, Coll.

Waterstradt (MNHU); 1 ex., Kelantan, 5°06' N/101°53’ E,

Coll. Veth (RMNH); 9 ex., Malakka, Perak,

2°12' N/102°15' E (MNHU); 1 ex., W. Sarawak, Quop,

1°37’ N/110°24' E, HI.1914, G. E. Bryant (BMNH);

1 ex., Malaya Peninsula, Barnam R., A. M. Lea & wife

(BMNH),; 1 ex., Sarawak, Puak, 3°48’ N/114°26' E,

3.V.1914, G. E. Bryant (BMNH); 2 ex., Selangor-Pah-

ang, 3°30' N/101°31' E, 1.1915 (BMNH); 2 ex., Kuala

Lumpur, Maxwell road, 3°08' N/101°41' E, 10.1.1916

(BMNH); 1 ex., Pahang, Kuala Tahan, 4°22' N/102°24' E,

XI1.1921, F. N. Chasen Coll. (BMNH); 1 ex., Kuantan,

3°49' N/103°19’ E, 10.11.1922, G. H. Corbett et al.

(BMNH); 1 ex., Pahang, F. M. S. Sungai Tembeling,

4°18' N/102°36' E, XI.1922 (BMNH); | ex., Kedah-Per-

ak, 5°52' N/100°31' E, III.1928 (BMNH); 2 ex., Pahang,

Bentong, 3°31' N/101°54' E, 22.X1I.1931 (BMNH);

1 ex., Perak, F. M. S. Larut Hills, 3700 ft., 5N°/100°53' E,

11.1932, H. M. Pendlebury (BMNH); 3 ex., Sarawak, foot

Bonn zoological Bulletin 71 (2): 139-176

of Mt. Dulit, junction of rivers Tinjar & Lejok,

3°20' N/114°8' E, 9.VUI.1932 (BMNH); 4 ex., Kuala

Lumpur, VI.1935, ex. F. M. S. (BMNH); 16 ex., Pahang,

F. M. S. Cameron Highland, 500 ft., 4°30' N/101°28' E,

VI-VIH.1935, H. M. Pendlebury (BMNH); | ex., Pahang

nr. Karak, Chintamani, 3°24' N/102°02' E, VUHI.1935

(BMNH); 5 ex., Pahang, F. M. S. Fraser Hill, 4200 ft.,

3°46' N/101°45' E, VI.1936, H. M. Pendlebury (BMNH);

1 ex., Pahang F. M. S., Pekan, 3°30' N/103°23' E, 25.

IV.1939 (BMNH); 1 ex, Kuala _ Kangsar,

4°46' N/100°56' E, 9.X1.1943 (BMNH); | ex., Sarawak,

Bario, 3°44' N/115°28' E, 2.XII.1965, Coll. G. H. L.

Rotschild (BMNH); 1 ex., Sabah, Mt. Trus Madi, 1800

ft., 5°33’ N/116°31' E, 18.—28.VHI.1977 (BMNH); 1 ex.,

Perak, Bukit Larut, 4°47’ N/100°45' E, 9.1X.1986, Sal-

leh & Ismail (UKM); 1 ex., Pahang, Cameron Highland,

Tanah Rata, 4°28' N/101°22' E, 25.1X.1986, Ismail &

Md. Nor (UKM); 1 ex., Negeri Sembilan, Johol,

2°52' N/102°15' E, 1.1.1987, T. K. Philips (BYUC);

2 ex., Sabah, Keningau area Tenom, alt. m 1230,

5°15' N/116°19' E, 23.X1.1987, Krikken & Rombaut

(RMNH); 1 ex., Pahang, T. Bera, 3°19' N/102°27' E,

1.-—3.X1.1990, Ruslan (UKM); 1 ex., Perak, Banding,

5°32' N/101°19' E, 29. —30. VIL.1991, Ismail et. al (UKM);

3 ex., Sabah, Pulau Banggi, 7°16’ N/117°9' E, 8—

12.V.1996, Salleh, Zaidi, Ismail & Sham (UKM); 3 ex.,

Pahang prov. Kuala Tahan, 4°19’ N/102°20' E, 6-9.

XI1.1999; 5.—9 111.2007, P. Kocarek et al. (CJB); 4 ex.,

West Pahang, 35 km see Ipoh, 1500 m, Cameron High-

lands, Tanah Rata, 4°30' N/101°28' E, 21.-24.IV.2001,

M. Riha (CJB); 1 ex., W. Perak, 1200 m, 25 km, N. E.

Ipoh, Banjaran Titiwangsa Mts. Koribu,

4°56' N/101°38' E, 6.-12.V.2001, M. Riha (CJB); 1 ex.,

Perak, Cameron highlands, 4°22' N/101°20' E, V.2011,

Sipek & Vondraéek (CJB). — Papua-New Guinea.

16 ex., Papua, Kokoda, 8°53' S/147°44' E, 1X.1933,

1,200—1,300 ft., L. E. Cheesman (BMNH); | ex., Mafu-

lu, 4000 ft., 1.1934, L. E. Cheesman (BMNH); 23 ex.,

Neth., New Guinea Exp. Star Range, 5°00S' /140°50' E,

1260 m, IV.—IX.1959, C. v. Heyningen (RMNH); | ex.,

Madang Dist., 5°14' S/145°48' E, Finisterre Mts., Dam-

anti 3550 ft., 2—11.X.1964 (BMNH); 2 ex., SW Sokopa,

Araban, 200 m, 4°35' S/152°07' E, II.2000, A. Weigel

(NME). -— Myanmar. 9 ex., Burma, Rangoon,

16°48' N/96°0' E, V.10—33, H. L. Andrewes (BMNH). —

Philippines. 5 ex., N. Luzon, Semper, 17°36' N/118°12'E,

Coll. Chapuis (IRSN); 1 ex, Los Banos,

14°10' N/121°13' E, P. I. Baker (NHRS); 7 ex., Phillipp.,

Jacoby Coll. (BMNH); 3. ex., Luzon, Jagor,

16°33' N/121°15' E, 58402 (MNHU),; 26 ex., Philippine,

16°33' N/121°15' E, Luzon (MNHU); 1 ex., Mindoro

(BMNH); 6 ex., Philippine, Mindanao, 7°51' N/124°51'E

(MNHU); 11 ex., Luzon Benguet, La_ Trinidad,

16°33' N/121°15' E, V.1914, G. Roettcher (MNHU);

1 ex., Philippine Island, 1919 (BYUC); 1 ex., Luzon,

Benguet, Kabayan, 16°33’ N/121°15' E, 21.X1.1997, L.

©LIB

144 Izfa Riza Hazmi & Thomas Wagner

F., Mey et al. (MNHU). — Singapore. | ex., Singapore,

1°21’ N/103°49' E, 97-74, H. N. Ridley (BMNH). — Sri

Lanka. 1 ex., Bogawantalawa, 10°07' N/88°24' E, II.—

IV.1882 (BMNH); 3 ex., Ceylon, 7°18' N/80°36' E,

11.1884, Mus. Columbo (MNHU); 1 ex., Ceylon, He-

naratgoda, XII.1889, H. P. Green (BMNH); 1 ex., Putta-

lam, 8°02' N/79°50' E, 1899, W. Horn (MNHU); 1 ex.,

Ceylon, Polonnarowa, 7°55' N/81°0' E, 27.11.1906, H.

Schoede (MNHU); 4 ex., Ceylon, Diyatalawa; Kandy,

5°15' N/100°29' E, VI.-IX.1908, G. E. Bryant; T. B. F

(BMNH). — Thailand. 9 ex., Bohol, Semper,

9°50! N/124°10' E, Coll. Chapuis (IRSN); 2 ex., Tringa-

no (BMNH); 5 ex., Japanoeli, A.L.v.H, Coll. Veth

(RMNH); 12 ex., Restit. 1885, Coll. Chapuis (IRSN):

1 ex., Wellesley Prov., 1904-105, H. N. Ridley (BMNH);

2 ex., Penin. Siam, Patalung Trang, 7°25' N/99°54' E,

3.V.1924, I. H. N. Evan (BMNH); 1 ex., Getassan,

1100 m, 1973, P. H. V. Doesburg (RMNH); 2 ex., Chum-

phon Prov., Pha To env., V.1998, Pradek & Sigut (CJB).

Redescription

Total length. 3.70—5.10 mm (mean 4.40 mm; n= 10).

Head. Varied from pale yellow to reddish, brownish

or black (Figs 1-5A), some specimens with black vertex

and yellow frons (Figs 2b, 4b, SAb). Antenna slender,

yellow, last antennomere usually brownish. Second and

Fig. 6. Distribution of Monolepta bifasciata (Hornstedt, 1788).

Bonn zoological Bulletin 71 (2): 139-176

third antennomere elongated, third slightly longer; ratio

length of second to third antennomere 0.75—1.05 (mean

0.89); ratio length of third to fourth antennomere 0.43—

0.57 (mean 0.50; Fig. 5B).

Thorax. Pronotum very finely punctuated, yel-

low to yellowish-red. Pronotal width 1.15—1.65 mm

(mean 1.40 mm), ratio length to width 0.61—0.65 (mean

0.63). Scutellum yellow, brownish or black, meso- and

metathorax pale yellow to black. Elytron yellow, bas-

al quarter to third including humerus, and correspond-

ing lateral margins in about 80 % of material examined

dark brown to black (Figs 1b, 4b, SAb), others pale red-

dish-brown (Figs 3b, 5Aa), rarely black base strongly

reduced (Fig. 2b). Elytron furthermore with transverse

black to reddish-brown band, usually paler than at elytral

base in the apical quarter (Fig. 1b) that does not reach the

lateral margin, sometimes reduced to a circular spot, but

rarely small (Fig. 2b) or absent (Fig. 3b). Very rarely ely-

tron almost entirely yellow. Elytral length 2.90-4.00 mm

(mean 3.45 mm), maximal width of both elytra together

2.00—2.80 mm (mean 2.40 mm), ratio of maximal width

of both elytra together to length of elytron 0.68—-0.72

(mean 0.70). Legs yellow to pale brownish.

Abdomen. Pale yellow to yellowish-red or brown.

Male genitalia. Median lobe lanceolate, becomes little

narrow towards apex (Figs 5C). Tectum long, broad. Me-

ody] | | |

©LIB

Revision of Monolepta Chevrolat, 1836 from the Sundaland area 145

dian spiculae brush-like of similar size (Figs 5Ca, Cb),

lateral spiculae broad, lobe-like with curved base, ventral

spiculae absent.

Female genitalia. Nodulus of spermatheca very small,

cornu long and median part curved, evenly sclerotised

(Fig. 5D). Two strongly different parts of bursa sclerites,

dorsal pair with strong spines (Fig. 5Ea), ventral part

elongated, with smaller spines at outer margin (Fig. 5Eb).

Diagnosis. In most specimens with characteristic co-

lour pattern, possessing yellow elytra with two transverse

reddish to blackish bands, where the subapical one is not

reaching the elytral outer margins (can be reduced to

spots in some specimens, and rarely completely yellow).

Transverse elytral bands also occur in M. orientalis and

M. flavofasciata, but in these bands are much wider or bi-

colorous (Figs 20, 23), and the apical one 1s reaching the

outer margin. The genitalic characters are most similar

to M. rubra, M. rufipennis and M. kuninghitam sp. nov.

since all four species lacs the ventral spiculae in medi-

an lobes (Figs 5C, 8C, 37C, 51C) and the spermathecae

(Figs 5D, 8D, 37D, 51D) possess a very small nodulus.

In term of the body size, M. rubra (4.50-6.00 mm), and

M. rufipennis (4.75-6.35 mm) are on average bigger

than M. bifasciata (3.70—5.10 mm) and M. kuninghitam

sp. nov. (3.70—4.35 mm), the ratio length to width of pro-

notum, in M. bifasciata and M. rubra is almost the same

(mean: 0.63 vs 0.64) while in M. kuninghitam sp. nov.

the pronotum is very narrow (0.71). In the few aberrant

coloured specimens, only the genitalia allow a clear allo-

cation to species.

Distribution. One of the most abundant species of

Monolepta with very wide distribution throughout the

Oriental Region, from India, southern China (literature

data) up to the Australian Region in New Guinea and

even North-East Australia (Fig. 6).

Monolepta rubra (Gyllenhal, 1808)

Figs 7-9

Crioceris rubra Gyllenhal, 1808: 272 (Chevrolat 1836:

407).

= Luperodes javanensis Jacoby, 1887: 234 (syn. by

Bowditch 1925: 253).

Type material. Crioceris rubra. Syntypes: 2 ex. “Crioce-

ris 81. 107 / rubra Gyllenh. Synon. Inf. 34 / Ind. Or. [?]

Lund.” and another ex with “Java Mellenb.” (NHRS).

Luperodes javanensis. Lectotype: “Batavia / Luperodes

javanensis Jac. Type / Type 18105 / Sythoff, Batavia,

Java” (RMNH). Paralectotype: 1 ex. same data as lecto-

type (MCZH; Fig. 7). Type locality: 6°10' S/106°S1' E.

Jacoby mentioned “two specimens” from one location in

his original publication. Both are available, and we des-

ignate a lectotype here to fix the name on a single spec-

imen.

Further material examined. — Indonesia. 2 ex., Nierst,

Depok, 6°23' S/106°48' E, 1899, Coll. Veth (RMNH);

Bonn zoological Bulletin 71 (2): 139-176

Fig. 7. Type material of Monolepta rubra (Gyllenhal, 1808).

— Luperodes javanensis Jacoby, 1887, paralectotype (MCZAH).

33 ex., Java, Proepoek Tegal, 6°52' S/109°08' E, [X.1909,

Valck Lucassen (RMNH); 4 ex., Java, Slawi Tegal,

6°59' S/109°08' E, [X.1909, Valck Lucassen (RMNH):

3 ex., Soemba Northcoast, 9°41' S/119°58' E, IV.1930,

W. C. van Heurn (RMNH); 5 ex., Java, Semarang,

6°58' S/110°25' E, E. R Jacobson (RMNH); 1 ex., Ban-

dong, 6°54’ S/107°36' E, Coll. Veth (RMNH),; 2 ex., West

Java, Krawang ds Randas-dengklok, 6°18' S/107°17' E,

2.X1.1951, L.V.L (RMNH). — Malaysia. | ex., Sarawak

(BMNH); 2 ex., Borneo, German Mission, 56358, Martin

Schmidt (MNHU).

Redescription

Total length. 4.50—6.00 mm (mean 5.31 mm; n= 10).

Head. Entirely reddish-brown to blackish brown. An-

tenna slender, pale yellowish-brown, terminal antenno-

mere usually dark-brown (Fig. 8A). Third antennomere

slightly longer than second; ratio length of second to third

Fig. 8. Monolepta rubra (Gyllenhal, 1808). A. Colour pattern.

B. Basal antennomeres. C. Median lobe, a. dorsal, b. lateral,

c. ventral, without endophallic structures. D. Spermathecae.

E. Bursa-sclerites, a. dorsal, b. ventral.

©LIB

146 Izfa Riza Hazmi & Thomas Wagner

antennomere 0)..75—1.00 (mean 0.83); ratio length of third

to fourth antennomere 0.38—0.56 (mean 0.42; Fig. 8B).

Thorax. Pronotum finely punctuated, entirely red-

dish-brown to dark-brown, surface strongly convex.

Pronotal width 1.40-1.75 mm (mean 1.58 mm), ratio

length to width 0.63—0.65 (mean 0.64). Scutellum, meso-

and metathorax dark brown. Elytron uniformly reddish

to dark brown, rarely more yellowish, sometimes with

fine darker outer margins (Fig. 7b). Elytral length 3.65—

4.30 mm (mean 3.94 mm), maximal width of both elytra

together 2.50—3.00 mm (mean 2.77 mm), ratio of max-

imal width of both elytra together to length of elytron

0.68—0.72 (mean 0.70). Legs fulvous.

Abdomen. Dark brown.

M. rubra

@ 1-2

@ 3-5

@ 6-33

M. marginicollis

m i-2

M 3-4

M@ o5-1

HE iiz-22

Fig. 9. Distribution of Monolepta rubra (Gyllenhal, 1808) and

Monolepta marginicollis, Jacoby, 1896.

Male genitalia. Median lobe lanceolate, slender, be-

comes slightly narrow towards apex. Tectum pointed.

Few long and slender median spiculae, lateral spiculae

club-shaped, lobe-liked at apex, (Figs 8Ca, Cb), ventral

spiculae weakly sclerotized and hardly visible.

Female genitalia. Spermatheca with very small nod-

ulus, middle part and cornu long, curved (Fig. 8D), two

pairs of bursa sclerites with strong spines (Fig. 8E).

Diagnosis. Monolepta rubra is characterized by an en-

tirely brownish-red to dark brown colouration. Monolep-

ta rufipennis and M. sulawensis sp. nov. are most similar

in colouration (Figs 37A, 60A). Total length of M. rubra

(4.50-—6.00 mm) is similar to M. rufipennis, while M. su-

lawensis sp. nov. is much smaller (3.70-4.25 mm), the

pronotum of M. rubra is less broad (0.64—0.68) than in

the other two species (0.60—0.64). M. rufipennis can be

distinguished also by the contrast between black head

and pronotum to the reddish elytra. Median lobe is slen-

der and lacks the ventral spiculae in M. rubra and M. ru-

fipennis, and both species are surely closely related, but

the outer shape of the median lobe is much more slender

in M. rubra (Figs 8Ca, 37Ca).

Distribution. Known from Java and adjacent islands

(Fig. 9), and Borneo.

Bonn zoological Bulletin 71 (2): 139-176

Monolepta signata (Olivier, 1808)

Figs 10-15

Galeruca signata Olivier, 1808: 665 (Jacoby 1889: 229).

= Crioceris neglecta Sahlberg, 1823: 72; repated in

1829: 29 (syn. by Maulik 1936: 410).

= Luperodes hieroglyphicus Motschulsky, 1858: 104

(syn. by Wagner & Bieneck 2012: 210).

= Luperodes quadripustulatus Motschulsky, 1858: 105

(syn. by Maulik 1936: 410).

= Monolepta elegantula Boheman, 1859: 183 (syn. by

Weise 1913: 229).

= Luperodes dorsalis Motschulsky, 1866: 415 (syn. by

Wagner & Bieneck 2012: 210).

= Luperodes quadriguttata Fairmaire, 1887: 333 (syn.

by Weise 1924: 169).

= Monolepta picturata Jacoby, 1896: 292; syn. nov.

= Monolepta simplex Weise, 1913: 229 (syn. by Weise

1924: 168).

Type material. Galeruca signata. Type material is not

available to us.

Crioceris neglecta. Syntype: ’Ind. Or. / neglecta Sahl-

berg n.sp. guttata Gyll:” At least on syntype is available

in NHRS.

Luperodes hieroglyphicus. Lectotype: @, “Luperodes

hieroglyphicus Motsch., Ind. Or” (ZISP; Fig. 10). Para-

lectotypes, 5 ex. same data as lectotype (ZISP). All six

syntypes were originally glued tightly together on one

card. The lectotype and one female paralectotype have

been remounted, the other four paralectotypes are left on

the original card (Wagner & Bieneck 2012: 210).

Luperodes quadripustulatus. Lectotype: 3, “Lupero-

des 4pustulatus Motsch., Ind. Or’ (ZISP; Fig. 11). Para-

lectotypes, 2 ex. same data as lectotype (ZISP). These

two paralectotypes have been originally glued together

with the choosen lectotype on the original card (Wag-

ner & Bieneck 2012: 210). A further type, not stated as

paralectotype in Wagner & Bieneck (2012) is in NHRS.

Monolepta elegantula. Syntype: Malacca / Kinb. /

type / elegantula Bhm.” At least on syntype is available

in NHRS. There is one specimen labelled ’Co-type” in

BMNH, invalid type from Nias Island near Sumatra,

while the species is only described from ”Malacca”.

Luperodes dorsalis. Holotype: 3, ”Luperodes dosalis

Motsch. Ceylon” (ZISP; Fig. 12) (Wagner & Bieneck

20:12: 210).

Luperodes quadriguttata. Type material is not avail-

able to us. We adopt Weise’ s (1924) statement that this

species is a junior synonym of M. signata (Olivier, 1808).

Monolepta picturata. Lectotype: “M. picturata Jacoby

1896 / Type / Toungoo India / Monolepta picturata Jac.

/ Jacoby Coll. 1909-28a” (BMNH; Fig. 13). — Paralecto-

type: 1 ex., same data as lectotype (BMNH). Type local-

ity: 18°56' N/96°25' E. Jacoby gave no details on speci-

men number and there are two specimens in his collecion

in BMNH available. We herein designate a lectotype here

©LIB

Revision of Monolepta Chevrolat, 1836 from the Sundaland area 147

Figs 10-13. Type material of Monolepta signata (Olivier,

1808). 10. Luperodes hieroglyphicus Motschulsky, 1858, lecto-

type, ¢ (ZISP). 11. Luperodes quadripustulatus Motschulsky,

1858, lectotype, 4 (ZISP). 12. Luperodes dorsalis Mot-

schulsky, 1866, holotype, ¢ (ZISP). 13. Monolepta picturata

Jacoby, 1896, lectotype (BMNH).

to fix the name on a single specimen. Jacoby was aware

about the similarity to M. signata ”A plainly marked

Species, resembling somewhat in colouration M. signata

Oliv., but easily distiguished by the number, position and

shape of the elytral pale spots”. To that time, colouration

was an important diagnostic character.

Monolepta simplex. Syntype: 1 ex. ”P. Princesa Pala-

wan Baker / simplex m.” (NHRS).

Further materials examined. — Bangladesh. | ex.,

Dacca, 23°42’ N/90°22' E, 7.1X.1945, D. Leston, B.M.

1945-86 (BMNH). — Cambodia. 1 ex., Peal-Leap,

11°42' N/103°02' E, V.1950 (BMNH). — China. 3 ex.,

China (BMNH); 2 ex., Fukien (MNHU); 3 ex., Chusan

Is., 30°10' N/122°24' E, J. J. Walker (BMNH); 4 ex.,

Kanton, 23.4°N/113.5°E, 1.-—30.X.1916, H. Weigold

(MNHU); 1 ex., Yunnan, 25°02’ N/102°42' E, 1.

VIII.1922. Prof J. W. Gregory (BMNH); 2 ex., SW. Chi-

na, Yunnan, 4.VIII.1922, Prof. J.W. Gregory (BMNH):

2: 6xm -<Anoys. 35500 NAT 04°11". E1923) 25 TE ieht

Bonn zoological Bulletin 71 (2): 139-176

(BMNH); 3. ex., China-Yunnan, Lijian — env.,

25°02' N/102°42' E, 10.VHI.1995, J. Scheider (CJB);

10 ex., Yunnan, Menghai, 21°58’ N/100°28' E, 6.10.

111.1999, River, P. Grootaert (IRSN). — Hong Kong.

5 ex., Hong Kong, 22°22' N/113°56' E, Walker Coll.

(BMNH); | ex., Hong Kong, 22°23' N/114°6' E, Stimp-

son (RMNH); 2 ex., Hong Kong, Tai Lung Farm, [X.—

XI1.1965, I. W. B. Thornton (BMNH); | ex., Hong Kong,

22°23' N/114°6' E (BMNH). — India. 32 ex., C. Almora,

Ranikhet, Kumaon, W. Almora, 29°35' N/79°39' E, 5.—

8.16, H. G. C. (BMNH); 14 ex., U. Gumti Val., W. Almo-

fasDn, 29°35’ N/79°39"-E, VIlT1917,1V.1919, OG, C,

Champion Coll. (BMNH); 9 ex., Assam, Sudiya, Doherty,

27°50' N/95°40' E, Fry Coll. (BMNH),; 4 ex., Assam, Pat-

kai Mt., 21°05' N/91°13' E, Doherty (BMNH); 2 ex.,

Malabar, 10°15’ N/75°14' E, Doherty, Fry Coll. (BMNH):

14 ex., India, Baly Coll. (BMNH); 1 ex., Nilgiri Hills,

11°25' N/76°30' E, T. V. Champbell (BMNH); 6 ex.,

Haldwani Div., Kumaon, 29°13' N/79°31' E, H. G. C.

(BMNH); 2 ex., Dahra Dun, 30°18' N/78°2' E (BMNH);

14 ex., Nilgiri Hills, 11.35°N/76.42°E, H. E. Andrewes

(BMNH); 2 ex., Sikkim, Gopaldhara, Rungbong Vall., H.

Steven (BMNH); 2 ex., Travancore, Wallardi, 37°N/

144.89°E, Collection J. Achard (BMNH); | ex., Taple-

jung District, Sengu 27°21' N/87°40' E, c 6200' (BMNH);

1 ex., Assam, 26°12' N/92°56' E, Doherty (BMNH);

1 ex., Deli, 3°35' N/98°39' E, S. V. L (RMNH); 1 ex.,

Punjab, Kangra Vall., 1903-37 (BMNH); 1 ex., Himala-

ya, Chaubattia, Almora District, 6-7000 ft., 1920,

29°35' N/79°39' E, S. R. Archer (BMNH); 3 ex., Bel-

gaum, 15°50' N/74°34' E, Andrewes Bequest, B.M.

1922-221 (BMNH); 1 ex., Punjab, Simla, 31°6' N/

77°10' E, E. C. Ansorgee (BMNH); 7 ex., Bengal, Sarda,

F.W.C. (BMNH); 3 ex., Bengal, Mandar, 21°41' N/

87°33' E, VU.1891, P. Cardon, Coll. Duvivier (IRSN);

1 ex., Bhowali, Nr. Naini Tal, 4500 ft., 29°22’ N/ 79°31'E,

21.X.1934 (BMNH); 4 ex., Assam, Mishmi Hills, Delai

Valley, Taphlogam, 28°14' N/95°59' E, 7.X1.1936

(BMNH); 4 ex., Coimbatore, 11°1' N/76°58' E, VI.1950,

P. Susai Nathan (IRSN); 13 ex., Nilgiri Hills, Cheran-

gode, 3500 ft., 11°25' N/76°30' E, X.1950, P. Susai Na-

than (IRSN); 21 ex., Annamalai Hills Cinchona,

10°02' N/77°07' E, 3500ft, V.1952, IV.1964, V.1968, P.

Susai Nathan (RMNH); 45 ex., S. Coorg Ammanit, 3100

ft, 12°25' N/75°44' E, XI.1950-II.1952, P. Susai Nathan

(IRSN); 24 ex., Kodaikanal Pulney Hills, 6500 ft,

10°12’ N/77°30' E, V.1953, P. S. Nathan (IRSN); 1 ex.,

Bengal, 21°56' N/88°51' E, Sunderbans, F.W.C, H.G.

Champion Coll. B.M. 1953-156 (BMNH); 2 ex.,

Arunachal Pr. 8 km S Jamiri, Sessa vicinity, 28°14" N/

94°4' E, 26.V—4.V1.2005, L. Dembicky (BMNH). — In-

donesia. | ex., Java, Tengger, 6°58’ S/111°17' E, 4000,

Coll. Duvivier (IRSN); 1 ex., Sumatra, Mjoberg, Kota

Tajne (NHRS); 5 ex., Sumatra, Myjoberg, Brastagi,

1°17' S/102°40' E (NHRS); 6 ex., Sumatra, Myjoberg,

Medan, 3°35' N/98°E (NHRS); 2 ex., Sumatra, Mjoberg,

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148 Izfa Riza Hazmi & Thomas Wagner

Tjinta Radja (NHRS); 1 ex., Sumatra, Mjoberg, Perda-

gangan (NHRS); | ex., Java, 7°36’ S/110°42' E (NHRS);

5 ex., Java, K.R.H. (RMNH); 1 ex., W. Java (RMNH);

4 ex., Java, Batavia, 6°12' S/106°50' E, Sythoff (RMNH);

2 ex., Bandar Baroe, Sumatra, 5°15' N/96°04' E, J.J.d.V

(RMNH); 1 ex., Java, Preange, Tjigembong, 6°42' S/

110°56' E, J. B. Corporaal (RMNH); 15 ex., N.O. Suma-

tra, Tandjong Morawa Serdang, 0°35' S/101°18' E, Dr. B.

Hadgen (RMNH); 40 ex., Nias, 1°07' N/97°31' E, J. D.

Pasteur (RMNH); 1 ex., Java, Kraksakun (RMNH):

14 ex., Java, Ardja-Sari, Preanger, 7°03' S/107°38' E, P.

F. Sitjhorf (RMNH); 1 ex., Java, G. Ocengaran,

7°11' S/110°20' E, v. Doesberg (RMNH); 1 ex., Kedir1,

7°49' S/112°0' E, Java de Vos (RMNH); 1 ex., Java,

Ameyer (RMNH); 1 ex., Sumatra, Brastagi,

3°22' N/98°34' E, Mjoberg (NHRS); 1 ex., Java Orient,

Montes Tengger, 4000 ft, 1890, H. Fruhstrofer (RMNH);

1 ex., Sumatra, Siboga, 1°44' N/98°46' E, X.1890-

11.1891, E. Modiglianii (BMNH); 3 ex., Sumatra, Pagh-

erang Pisang, X.1890-III.1891, E. Modigliani (BMNH);

31 ex., Sumatra, Manna, 4°30' S/102°58' E, 1902, M.

Knappert; Museum Leiden, ex. Collection J.J. de Vos tot

Nederveen Cappel (RMNH); 1 ex., Sumatra, Nias,

1°07' N/97°31' E, Fry Coll. (BMNH); 11 ex., Nias Island,

Schreiber, 1°07' N/97°31' E, Sharp Coll. (BMNH); 1 ex.,

Slawi Tegal, Java, 6°59' S/109°08' E, 1909, Valck Lucas-

sen (RMNH); 1 ex., Sumatra, Soekaranda, Dohrn, Jaco-

by Coll. (BMNH); 3 ex., Java, Nongkodyadjar, 7°38' S/

112°54' E, Jan. 1911, E. Jacobson (RMNH); 1 ex., Java,

Mount Salak, 7°46' S/112°56' E, 400 m, 1921, L. G. E.

Kalshoven (IRSN); 3 ex., Java, Kerkhoven, 1921, Pa-

noembang, ardayasari (RMNH); | ex., Java, 24.XII.1925

(IRSN); 1 ex., Central Java, 7°09' S/110°08' E, 4.11.1927,

Fig. 14. Monolepta signata (Olivier, 1808). A. Colour pattern.

B. Basal antennomeres. C. Median lobe, a. dorsal, b. lateral,

c. ventral, without endophallic structures. D. Spermathecae.

E. Bursa-sclerites, a. dorsal, b. ventral.

Bonn zoological Bulletin 71 (2): 139-176

2093 m, Rensch (MNHU),; 3 ex., C. Java, Kedae Pagar,

Saemang, 800 m, 2.11.1932, D. V. L (RMNH); 1 ex.,

Java, Lembang, 6°49’ S/107°37' E, V.1933, v. Doesburg

(RMNH); | ex., Tangk-Prahoe, 6. VI.1933, P. H. V. Does-

burg (RMNH); 1 ex., Java, G. Kawi, 7°55' S/112°27' E,

VII.1934, v. Doesberg (RMNH); 1 ex., Bremi, Probul-

ingga, 7°58' S/113°29' E, 1000 m, X1.1934, W. C. v. Heu-

ton, ex. Collection, S. J. van Ooststroom, rec. 1982

(RMNH); 1 ex., 23.VII.1938, H. M. Pendlebury

(BMNH); | ex., G. Gede, Pontiek, 6°46’ S/106°56' E

1485 m, 2.V.1948, Dr. L. Kalshoven (RMNH); 6 ex., W.

Java, Tjibadas, 7.19°N/107.36°E, V.1950, v. Ooststroom,

S. J. van Ooststroom, rec. 1982 (RMNH); 2 ex., West

Java, Puntjak, 6°39' S/106°56' E, 1400 m, 27.1X.1953, J.

v. der Vocht (RMNH); 1 ex., N. Sumatra, Susuk,

3°09’ N/98°21' E, 29.V.1994, Muruyama (CTJ); 1 ex.,

Java Central, Bandar, 7°09' S/110°08' E, 550 m, at light,

20.1.1998, R. Cervenka (CJB); 1 ex., N. Sumatra, Silalahi

alt. 1600 m, nr D. Toba, 2°19’ N/98°43' E, 2.V.1998,

Muruyama (CTJ). — Malaysia. 4 ex., Malacca,

2°19' N/102°20' E, Coll. Duvivier, Coll. Chapuis (IRSN);

1 ex., Kuala Lumpur, 3°08' N/101°41' E (BMNH),; 1 ex.,

Bengal Occ. Betana, 1890, Coll. Duvivier (IRSN); 1 ex.,

Sarawak, Bidi, 2°33' N/113°01' E, 28.1.1909, C. J.

Brooks (BMNH), | ex., Malay Penin., Selangor F. M. S.,

Gombak Valley, 3°17' N/101°38' E, 17.X.1921, H. M.

Pendlebury (BMNH); 2 ex., Pahang, F. M. S. Kuala Tah-

an, 4°22' N/102°24' E, 19.X1.1922, H. M. Pendlebury

(BMNH); | ex., South China Sea, Pulau Tioman, Seda-

gong, 900' , 2°47’ N/104°10' E, V.1927, N. Smedly

(BMNH); 1 ex., Malay Penin., Kedah Perak,

5°53' N/100°31' E, 23.11.1928 (BMNH); | ex., Perak, F.

M. S. Larut Hills, 4500 ft., 4°47' N/100°45’ E, 20.11.1932,

H. M. Pendlebury (BMNH); | ex., Malay Penin., Pah-

ang, Fraser’ s Hill, 3°46’ N/101°45' E, 25.X.1933, Pine

trees (BMNH),; 3 ex., Pahang, F. M. S. Cameron High-

lands, 4800 ft., 4°29’ N/101°23' E, 22. VI.1935 (BMNH):

1 ex., Tapah, 4°12' N/101°15' E, 27.11.1974, Y. Kiyoya-

ma (BMNH); 1 ex., Cameron Highlands, 4°29' N/

101°23' E, 6.1.1982 (BMNH); 1 ex., Pulau Tioman,

2°47' N/104°10' E, Tekek, 24.11.1987, T. E. Leiler

(NHRS); 1 ex., N. Borneo Exp., Sabah, Interior zone,

5.1°N/115.59°E, 16 km of Tenom Agric. Res. Station

along Sg. Pegalan, 5.1°N/115.59°E, 200, 23.X1.1987, J.

Huisman & R. de Jong (RMNH); | ex., Malaysia, Fraser

Hill, 3°46' N/101°45' E, 21.11.1991, RM exped. (NHRS);

2 ex., Perak, Banding, 4°47' N/101°11' E, 2.X1.1991, Is-

mail et al. (UKM); 1 ex., Kedah, Langkawi, Lubuk

Semilang, 6°21’ N/99°47' E, 8-—10.XII.1992, Zabidi

et al. (UKM); 3 ex., Perak, Pangkalan Hulu, 5°42' N/

100°59' E, XI.1991-IT.1993, Ismail et al. (UKM); Pah-

ang, Tasik Bera, Pos Iskandar, 3°07' N/102°36' E, 4.—

8.V.1993, Sham et al. (UKM); 1 ex., Pahang, Pulau Tio-

man, 2°47' N/104°10' E, 18.—21.1X.1999, Ismail & Sham

(UKM); 1 ex., W-Perak, Bamjaran Titi Wangsa Mts.,

5°58' N/101°20' E, 1200 m, V.2001, P. Cechovsky

©LIB

Revision of Monolepta Chevrolat, 1836 from the Sundaland area 149

(NME); 19 ex., W-Kelantan, Gunung Basor,

5°30' N/101°45' E, 1700 m, V.2016, P. Cechovsky

(NME). — Myanmar. 3 ex., Tenasserim, Javoy, Fry Coll.

(BMNH); | ex., U. Burma, Kyauktau, 20°51' N/92°58' E,

Sehwebo Dn. HGC (BMNH); 1 ex., Birmania, Shwego

Myo, X.1885, Fea (IRSN); 1 ex., Carin, Cheba,

19°13' N/96°35' E, 1.1888, I. Fea (IRSN); 1 ex.,

Tharrawaddy, Burma, 17°39' N/95°48' E, 1902.294

(BMNH); 1 ex., Burma, Toungoo, 18°56’ N/96°25' E,

1902.294 (BMNH); 1 ex., Burmah, 21°54' N/95°57' E,

Andrewes Bequest (BMNH); 1 ex., Upper Burma, Se-

ingkhu Valley, 28.5°N/97.35°E, 1926, P. Kingdon Ward

(BMNH); | ex., Upper Burma, Mali Kha Valley, 1926, P.

Kingdon Ward (BMNH); 4 ex., Tenasserim, Malvedaung,

30 mS. of YE, 15.-25.X1.1934 (BMNH); 6 ex., Burma,

Mishmi Hills, 28°14' N/95°59' E, 1935, M. Steele

(BMNH); 1 ex., Upper Burma, Hkamti Long, [V—V.1935

(BMNH); 6 ex., Upper Burma, Nam Tai Valley,

27°42' N/97°54' E, 2. VIII.1938, R. Kaulback (BMNH).

— Nepal. | ex., British-Nepal Exped., 1950 (BMNH);

1 ex., West Nepal, Silgarhi-Doti, Kali Gad,

29°16' N/80°59' E, 26.VII.1953, J. B. Tyson (BMNH);

1 ex., Rimi, 29°07' N/82°34' E, 1000 ft., 28.[X.1952

(BMNH),; | ex., Pokhara, 3000 ft., 28°15' N/83°58' E,

12.1V.1954, J. Quinlan (BMNH); 1 ex., Pokhara, Basun-

ghara Parc, 800 m, 28°12' N/83°50' E, IX.2014, F.

Creutzberg (NME). — Philippines. 1 ex., Palawan, P.

Princess, 9°26' N/118°22' E, Baker (NHRS); 4 ex., Lu-

zon, 16° N/121° E, 38765 (MNHU). — Singapore. 4 ex.,

Singapore, 1°21' N/103°49' E, C.J. Saunders (BMNH).

— Sri Lanka. 3 ex., Ceylon (MNHU), 4 ex., Colombo on

Coast level, 6°56' N/79°50' E, 7.-27. IV.1882, G. Lewis

(BMNH); 1 ex., Kandy, 7°17’ N/80°38' E, 1546-1727 ft.,

17.—23.11.1882, G. Lewis (BMNH); 1 ex., Halupahani,

Haldummulle, 6.76°N/80.88°E, 1904 (BMNH); 3 ex.,

Ceylon, T. B. Fletcher, 1909 (BMNH); 3 ex., Ceylon, A.

Rutherford, 1914, T. B. Fletcher (BMNH); 4 ex., Ham-

bantona, 6°07' N/81°07' E, IX.1890, H. P Green

(BMNH),; 1 ex., Ceylon, Kalawewa, 8°01' N/80°33' E,

14.11.1953 (BMNH); 1 ex., Dickoya 20 km WSW, Nu-

wara Eliya, 6°52' N/80°36' E, 28 111.1973, G. Benick

(MNHU),; 2 ex., Sri Lanka NWP Kur. D Pannala, 2.

11.1974, P. I. Perrson (NHRS); 1 ex., Kitulgalle,

6°59’ N/80°25' E, 1700 ft., 17.-20.1.1982 (BMNH):

1 ex., S. Sri Lanka near Tissamaharama, 90 km near NE

of Matara, 6°17' N/81°17' E, 14.01.1994, Z. Kajval

(CJB). — Thailand. 4 ex., Siam, Renong, Doherty

(BMNH); 1 ex., C. Siam, 150 m, Kwae Nov River. Exp.

Niki, 14°19’ N/98°57' E, 23. IV.—5.V.1946, J. K. Jonkers

(RMNH); 2 ex., Thailand, 25 km NM v. Lan-Sak,

15°27' N/99°34' E, [X.1990 (MNHU); 2 ex., N. Thailand,

Mae Hong San env. Ban Huai Po, 18°44’ N/97°52' E,

1700 m, 24.—30.VI.1993, J. Schneider (CJB); 3 ex., Loe

Chiang Khan, 17°50' N/101°45' E, 11.11.1999, P. Groot-

aert (IRSN); 2 ex., Pattaya, 12°53' N/100°53' E, III.2016,

R. Mazrozis (NME). — Vietnam. 5 ex., Coll. Duvivier;

Bonn zoological Bulletin 71 (2): 139-176

Coll. Chapuis (IRSN); 5 ex., Barway, Cardon, Coll. Du-

vivier (IRSN); 2 ex., Berhampur (BMNH); 1 ex., Da-

laen-saen nr Nong-po, Walker Coll. 93-18 (BMNH);

1 ex., Jad. Bov, Coll. Veth (RMNH); 1 ex., Annam, Phuc-

Son, Nov—Dez, H. Fruhstorfer (RMNH); 2 ex., Central

Tonkin Chiern-Hoa, Aug. Sept., H. Frihstorfer (RMNH):

2 ex., MHaut-Tonkin, Madon, Riviere Claire,

19°45’ N/107°45' E (URSN); 1. ex., Tonkin,

19°45' N/107°45' E, Coll. Mandon (IRSN); 2 ex., Hanoi,

21°1' N/105°51' E (IRSN); 1 ex., Saigonh (IRSN); );

4 ex., Soereol, 7.1878 (RMNH); 6 ex., Hanoi, 1903, Coll.

Veth (RMNH); 8 ex., Tonkin, Hoa Binh, Hanoi, II.—

VI.1917; VII.1918, R. V. de Slavaza (BMNH); 1 ex.,

Tonkin, Bao Ha, X.—XII.1923, H. Stevens (BMNH):

3 ex., Tonkin, Thai-Nien Basin of Fleuve Rouge, 1924,

H. Stevens (BMNH); 1 ex., Vietnam, Prov. Lao-Cai,

22°20' N/104°E, 1900 m, 26.1X.1963, Mai (MNHU);

1 ex., Vietnam N., Huong Son Prov. Ha Son Bin,

21°27' N/105°59' E, 26.— 29.1V.1991, J. Strand (CJB);

2 ex., Cao Bang Prov., Pia Oac NR, 22°38' N/105°53' E,

V.2014, A. Weigel (NME).

Redescription

Total length. 3.50—5.25 mm (mean 4.47 mm; n= 10).

Head. Very finely punctuated, pale brown, red-

dish-brown or red. Labrum dark brown and mandible

partly black. Antenna slender and extending to the mid-

dle of the elytra, dark brown to black (Fig. 14Aa), usu-

ally only three basal antennomeres pale yellow or red-

dish (Figs 10b—13b, 14Ab). Second antennomere usually

schorter than third; ratio length of second to third anten-

nomere 0.67—1.00 (mean 0.77); ratio length of third to

fourth antennomere 0.38—0.50 (mean 0.45; Fig. 14B).

Thorax. Pronotum finely punctuated, pale yellow to

brown-reddish. Pronotal width 1.15—1.55 mm (mean

1.35 mm), ratio length to width: 0.60—0.63 (mean 0.61).

Scutellum brown to black, meso- and metathorax black.

Elytron brown to black, varied on number of yellow

spots, from usually one beyond humerus up to three.

Elytral length 2.90-3.60 mm (mean 3.32 mm), maxi-

mal width of both elytra together 2.00—2.80 mm (mean

2.40 mm), ratio of maximal width of both elytra together

to length of elytron 0.72—0.75 (mean 0.73). Coxae pale

yellow to brown, base of femur yellow, outer parts black,

also apical part of tibiae and all tarsi brown to black

(Figs 10b—13b).

Abdomen. Pale yellow to brown.

Male genitalia. Median lobe parallel-sided, becomes

narrow towards apex. Median spiculae consist of a group

of long and slender structures, apical median spiculae

slightly curved. Lateral spiculae c-shaped, Ventral spic-

ulae not visible from dorsal view, short, slightly curved

and slender (Fig. 14Cb).

Female genitalia. Spermatheca with spherical nod-

ulus, cornu long and curved (Fig. 14D). Dorsal part of

©LIB

150 Izfa Riza Hazmi & Thomas Wagner

M. signata

@ ic-31

Fig. 15. Distribution of Monolepta signata (Olivier, 1808).

bursa sclerite with saw-like spines, ventral part longer,

slender, with fine spines (Fig. 14E).

Diagnosis. This species shows a high variety in co-

louration pattern. Characteristic are the brownish to black

elytron with circular humeral and praeapical yellow spots

(Figs 10b-13b, 14A). Monolepta zonula, M. empatbu-

lat sp. nov. and some specimens of MZ mohamedsaidi

sp. nov. resemble this species in terms of colouration,

particularly regarding the yellowish spot on elytron, but

all three have a black head, while head in M. signata is

always pale brown, reddish to yellowish. Total length of

M. signata varied from 3.50—5.25 mm, quite similar to

M. zonula (3.50—4.90 mm) while M. empatbulat sp. nov.

(3.25-—3.80 mm) and M. mohamedsaidi sp. nov. (3.25—

4.00 mm) are on average smaller. The genitalic char-

acters are quite different among these four species, and

specimens can be clearly differentiated by a check of the

male genitalia (Figs 14C, 25C, 47C, 56C).

Distribution. Probably the most abundant species of

Monolepta in the Oriental Region from Pakistan (liter-

ature data) and southern China to Java and Bali, but not

known from Wallacea (Fig. 15).

Bonn zoological Bulletin 71 (2): 139-176

fa ie al

maa | SS

Spel it

Monolepta jacobyi Weise, 1908

Figs 16-18

Monolepta jacobyi Weise, 1908: 326.

Replacement name for Monolepta basimarginata Jaco-

by, 1884a: 54 a junior homonym of Galleruca basimargi-

nata Boisduval, 1835 a species from New-Guinea, trans-

ferred to Monolepta by Weise (1908).

Type material. Monolepta basimarginata. Lectotype:

“Rawas, 5.78 / Monolepta basimarginata Boisd. ?”

(RMNH,; Fig. 16). Type locality: 2°31' S/102°54' E. Jaco-

by mentioned several locations (1.e., specimens) in his

> BOEL ‘eSIa ny

1Agooel e}dajouoy

Fig. 16. Type material of Monolepta jacobyi Weise, 1908. —

Monolepta basimarginata Jacoby, 1884, lectotype (RMNH).

©LIB

Revision of Monolepta Chevrolat, 1836 from the Sundaland area |e |

original publication and we herein designate a lectotype

to fix the name on a single specimen. The specimens has

a location label from the type locality, the species name

with “Boisd. ?” is in Jacoby’ s handwriting, but not on

usual light blue paper he used for type labels. Seems so

that Jacoby was not completely convinced about his de-

scription.

a |

i > |

Fig. 17. Monolepta jacobyi Weise, 1908. A. Colour pattern.

B. Basal antennomeres. C. Median lobe, a. dorsal, b. lateral,

c. ventral, without endophallic structures. D. Spermathecae.

E. Bursa-sclerites, a. dorsal, b. ventral.

Further material examined. — Indonesia. 14 ex., Java,

Ardja-Sari, 6.91°S/107.67°E, Preanger (RMNH); 24 ex.,

Sumatra, 4°27' S/102°59' E, M. Knappert (RMNH); 3 ex.,

Java, Malang, 7°58' S/112°37' E, A. Koller (RMNH);

1 ex., Java, Piep (RMNH); | ex., Sumatra, Palembang,

2°59' N/104°45' E, M. Knappert (RMNH); 5 ex., Sumatra,

Solok, 0°48' S/100°38' E, P. O. Stolz (RMNH); 4 ex., Su-

matra, Padang, 0°58' S/100°37' E, J. D. Pasteur (RMNH);:

1 ex., Palembang, Boengamas, 2°59' S/104°45' E, G.

van Hassen (RMNH); 1 ex., W. Java (RMNH); 4 ex.,

Java, 7°35' S/110°42' E, Baly Coll. (BMNH); 1 ex., Su-

matra, Sir S. Raffles (BMNH); 3 ex., Sumatra, Sibolan-

git, 3°18’ N/98°35' E, Mjoberg (NHRS); 2 ex., Medan,

3°35' N/98°40' E, Mjoberg (NHRS); 1 ex., Lombok,

8°39' S/116°19’ E, Carl Auriv (NHRS); 1 ex., Java, Desa

Kembangan, 6°14' S/106°48' E, Drescher (MNHU); 2 ex.,

Sumatra (MNHU); 1 ex., Spjg., 0°27' S/100°54' E, X.1877

(RMNH); 2 ex., Sumatra, Pagherang-Pisang, X.1890.—

II1.1891, E. Modigliani (BMNH); 2 ex., Java occident.,

Mons Gede 4000’ , 6°46’ S/106°57' E, VIII.1892, H.

Fruhstorfer (RMNH); 3 ex., Java occident., Pengalengan

4000' , 6°12' S/106°56' E, 1893, H. Fruhstorfer (RMNH);

1 ex., Java occident., Sukabumi, 6°55' S/106°55' E,

1893, H. Fruhstorfer (RMNH); 2 ex., Mentawei Sipo-

Bonn zoological Bulletin 71 (2): 139-176

ra, 2°10' S/99°41' E, V—V1.1894, Modiglianii (MNHU):

12 ex., Borneo Exped.; Borneo, Poelau Sibau, VI.1894,

Dr. J. Bultirofer (RMNH); 3 ex., Java, Banjoewangi,

8°12' S/114°22' E, 1910, Mac Gillavry (RMNH); 5 ex.,

Java, Nongkodjadyar, 1.1911, E. Jacobson (RMNH); 4 ex.,

Sumatra, Siolak Daras, Korinchi Valley, 3100 ft., HI.1914

(BMNH); 1 ex., Java, 1921, Kerkhoven (RMNH); | ex.,

West Sumatra, Siberut Island, 1°22' S/98°54' E, [X.1924.,

C.B.K. & N.S (BMNH); 13 ex., Sipora Island, West Su-

matra, 2°10' S/99°41' E, X.1924, C. B. K. & N.S; H. H.

Karny (BMNH); | ex., Indonesia, W. Java, Poentjak-pas,

ca. 1000 m, Onderneming “Tyiliwoeng”, VI.1932, W. C.

van Heurn (RMNH); | ex., Java, Moeria, 16. VII.1933,

P. H. V. Doesburg (RMNH); 1 ex., East Java, Soeraba-

ja and surroundings, 7°17' S/112°44' E, 1938, W.C. van

Heurn (RMNH); 7 ex., N. Sulawesi, Mt. Ambang nr

Kotamobagu, 0°44" N/124°18' E, 20 km E of alt. m. ca.

1000, 28 —29.V.1985, J. Huijbregts (RMNH); 25 ex., Su-

lawesi Utara, Danau Modat, 1200 m near Kotamobagu,

0°43' N/124°27' E, VIII.1985 (BMNH); | ex., C Sulawesi,

Lore Lindu NP Dongi Dongi Shelter, 1°31' S/120°11' E,

alt. 940 m, 3.-9.XII.1985, J. Krikken (RMNH); 1 ex.,

Sumatra, Umg. Prabal, 1050 mm, 2°47' N/98°56' E,

VIII.1992, U. Buchsbaum (NME); 6 ex., S Sumatra,

Lampung Prov., Bukit Barisan Selatan, 5°4' S/104°4' E,

7-17.11.2000, Liwa, J. Bezdék (CJB); 1 ex., Indone-

sia, Bali, 8°24' S/115°11' E, 19-21 111.2007, B. H. Izfa

(UKM). — Malaysia. 1 ex., Perak, 4°48’ N/100°48’ E,

Doherty (BMNH); 1 ex., Malay Penin., Selangor F.

M. S., Kuala Lumpur, Batu Caves, 3°15’ N/101°40' E,

TX.1921, H. M. Pendlebury (BMNH); 5 ex., Perak, F. M.

S. Jor camp, 3°54’ N/101°34' E, VUI.-IX.1922, E. Sei-

mun (BMNH); 12 ex., Perak, F. M. S. Batang Padang,

Jor Camp, 3°54' N/101°34' E TI.1924, F. M. Pendlebury

(BMNH); 1 ex., Malaya, Terengganu, Jerangau Es-

tate, 4°54’ N/103°11' E, 7.111966, Dept. of Agriculture

(BMNH); 1 ex., S. W. Sabah nr Long Pa Sia (East) ca.

1000 m, 5°20' N/117°10' E, 25.XI.—7.XII.1987, C. v.

Achterberg (RMNH); 2 ex., Perak, Tapah, Lata Iskandar,

4°12’ N/101°15’ E, 3.-4.IV.1990, 15.1X.1995, Ismail &

Ruslan (UKM); 1 ex., Borneo, Sarawak, Kuching, Bako

National Park, 1°43’ N/110°28' E, 5.V.1999, P. Vortruba

(CJB); 3 ex., Kelantan, 90 km N of Gua Musnag, Mt.

Basor 1770 m, III.2015, P. Cechovsky (CJB). — Philip-

pines. 1 ex., Philli. Island (BMNH); 1 ex., Los Banos,

14°10’ N/121°14' E, P. I. Baker (NHRS); 1 ex., Luzon,

16°33' N/121°15' E, Semper (IRSN); 1 ex., Coll. Duvi-

vier (IRSN); 1 ex., Manila 14°35’ N/120°59' E (MNHU);

4 ex., Luzon, Id. Nueva Viscaya, Sta. Fe. Dalton Pass,

16°35' N/121°15' E, 900 m, 8—12.VI.1991, Rolland A.

Muller (RMNH); 2 ex., N. Luzon, Mts. Prov. Chatel,

17°02' N/121°03' E, 24. [X.-14.X.1988, Cerny & Schint-

Imeister (MNHU); 10 ex., N. Luzon, Ifugao Banaue

vic., 16°54' N/121°06' E, 22. 1X —16.X.1988, Cerny &

Schintlmeister (MNHU). — Singapore. 2 ex., Singapore,

1°21’ N/103°49' E, C. J. Saunders (BMNH).

©LIB

152 Izfa Riza Hazmi & Thomas Wagner

Redescription

Total length. 6.40—7.70 mm (mean 6.86 mm; n = 10).

Head. Finely punctuated, entirely pale yellow to

brown. Labrum and mandible brown. Antenna slender,

extending beyond the middle of elytra, entirely yellow

(Fig. 17A) or terminal antennomere with brownish tip

(Fig. 16b). Second and third antennomere approximately

of the same length; ratio length of second to third an-

tennomere 0.80—1.00 (mean 0.92); ratio length of third

to fourth antennomere 0.36—0.45 (mean 0.40; Fig. 17B).

Thorax. Pronotum sparsely punctuated, entire-

ly yellow, surface slightly convex. Pronotal width

2.00—2.25 mm (mean 2.12 mm), ratio length to width

0.59-0.61 (mean 0.60). Scutellum yellow. Meso- and

metathorax black. Elytron yellow, basal margin black,

sometimes extend to the lateral margin as up to anteri-

or half; finely punctuated. Elytral length 5.25-6.00 mm

(mean 5.61 mm), maximal width of both elytra together

3.50-4.00 mm (mean 3.79 mm), ratio of maximal width

of both elytra together to length of elytron 0.66—0.70

(mean 0.68). Legs yellow to pale brownish.

SN =

oO \

M. jacobyi CO) XS e a x

@ 1-2 Qo \e

— @ 3-5 ~ Bs v5

@ 6-10 ~% a |-

@ 1-17 e Owe), . a

7G Geol

@ is-2

— BS

I . - A

Fig. 18. Distribution of Monolepta jacobyi Weise, 1908.

Abdomen. Yellow to yellowish-brown.

Male genitalia. Median lobe long and slightly conical

towards apex. Median spiculae long and slender, later-

al spiculae club-shaped with lobe-like apex (Fig. 17Ca),

ventral spiculae weakly sclerotised, hardly visible. Ven-

tral groove very broad and continuously convergent to

the orifice (Fig. 17Cc).

Female genitalia. Spermatheca with spherical nodu-

lus, median part long and curved cornu (Fig. 17D). Bursa

sclerites strongly sclerotized; dorsal and ventral bursa

sclerites almost of the same size, both carrying strong

spines (Fig. 17E).

Diagnosis. Monolepta jacobyi is one of the largest

Monolepta species of Sundaland (total length 6.40—

7.70 mm). This yellowish species with narrow black el-

ytral base at humeri looks similar to the much smaller

Bonn zoological Bulletin 71 (2): 139-176

M. kuninghitam sp. nov. (total length 3.70-4.35 mm).

The genitalic characters are very different between both

species (Figs 17C—E, 51C—E). The combination of large

size, uniform yellow dorsal colouration with narrow

black base at humerus is very characteristic for M. jaco-

byi.

Distribution. This abundant species is mainly known

from the Sundaland area, eastwards to Sulawesi (Fig. 18).

Monolepta orientalis Jacoby, 1889

Figs 19-21

Monolepta orientalis Jacoby, 1889: 227.

= Monolepta konbirensis Duvivier, 1891: 47 (syn. by

Maulik 1936: 407).

Type material. Monolepta orientalis. Syntypes: “In-

dia / Monolepta orientalis Jac. / 1%* Jacoby Coll. / Type

18437” (MCZH,; Fig. 19); “Bhamo, Birmania, Fea VHI. /

orientalis Jac. / Monolpet aorientalis Jac.” (MCGD).

Monolepta konbirensis: Type material is not available

to us, Maulik (1936) mentioned three specimens in coll.

Duvivier in BMNH. We adopt Maulik' s (1936) state-

ment that it is a junior synonym of Monolepta orientalis

Jacoby, 1889.

Fig. 19. Type material of Monolepta orientalis Jacoby, 1889,

syntype (MCZAH).

Further material examined. - _ Bangladesh.

2 ex., Bengal (BMNH); 18 ex., Bengal, Mandar,

23°41' N/90°21' E, VII.1891, P. Cardon (IRSN). — In-

dia. 1 ex., India (IRSN); 1 ex., Kanara (BMNH); 2 ex.,

Khasia Hills, 25°34' N/91°39' E (BMNH); 1 ex., Bom-

bay, 18°54' N/73°05' E (BMNH); 1 ex., Nandidrug, S.

India, T. V. C (BMNH); 2 ex., South India, Pondicherry

State, 10°55’ N/79°50' E, Karikal (MNHU); | ex., South

Mysore, 12°16' N/76°38' E, H. E. Andrewes (BMNH);

19 ex., Nilgiri Hills, 11°25' N/76°30' E, G. F. Hampson,

H. E. Andrewesi, A. K. Weld Downing (BMNH); 4 ex.,

Travancore, Wallardi, Collection J. Achard (BMNH);

1 ex., Nilgiri Hills, 11°25' N/76°30' E, A. K. Weld Dow-

ning (BMNH); 5 ex., Belgaum, 15°51' N/74°30' E, 1891

(BMNH); | ex., Deolali, 19°28’ N/74°37' E, 1.1.1922,

Maj. J. E. M. Boyd (BMNH); 28 ex., South India, S. Co-

©LIB

Revision of Monolepta Chevrolat, 1836 from the Sundaland area i353

Fig. 20. Monolepta orientalis Jacoby, 1889. A. Colour pattern.

B. Basal antennomeres. C. Median lobe, a. dorsal, b. lateral,

c. ventral, without endophallic structures. D. Spermathecae.

E. Bursa-sclerites, a. dorsal, b. ventral.

org-Ammatti, 3100 ft., 12°25' N/75°44' E, II.—X1.1952, P.

S. Nathan (IRSN); 1 ex., Anamalai Hills, Cinchona 3500

ft., V.1967, P.'S.N. (MNHU); 2 ex., South India, Karnata-

ka State, Chikmagalur, 13°18' N/75°46' E, V.1982, 4000

ft., T. R. S. Nathan (CJB); 1 ex., North Indien, 600 m

Uttar Pradesh, Shiwalik Kette, 13 km SW Dehra Dun,

27°34' N/80°05' E, 20. VIII.1985, J. Schulze (MNHU);

1 ex., India, Kerala, 27 km S. Calicut, Univ. Calicut

Botanical Garden, 11°15' N/75°46' E, 6.X.1985, N. F.

Johnson, D. C. Darling (CJB); 1 ex., N. India, Chandi-

gar, 30°43’ N/76°46' E, VII.1986, K. Werner (MNHU);

1 ex., Pune, 18°31' N/73°51' E, IL-IX.1997, H. V.

Ghate (BMNH). — Indonesia. | ex., Batavia, Tanjong

Priok, 6°07' S/106°52' E, 90-80 (BMNH). — Malaysia.

1 ex., Malay Penin. (BMNH); | ex., Island of Penang,

5°15' N/100°29' E, Baker (NHRS); 1 ex., Malay Penin.,

West Coast, Langkawi Is., 6°21' N/99°47' E, 28.1V.1928

(BMNH); 1 ex., Kedah, Pulau Langkawi, Teluk Datai,

6°23' N/99°42' E, 20.V.1992, Ismail et al. (UKM); 2 ex.,

Perlis, Taman Negeri Wang Klian, 6°40' N/100°11' E,

29 IX.—4.X.1999, Zaidi et al. (UKM). — Myanmar.

1 ex., N. Toungoo, L. Burma, 18°56' N/96°25' E, XII.26,

H. G. C. (BMNH). — Nepal. 3 ex., Chitwan Nat. Pk.,

700' , 27°29' N/84°24' E, 3.-6.V1.1983, Sauraha, at MV

light (BMNH). — Sri Lanka. 3 ex., Konbi (MNHU);

7 ex., Band., Horn (MNHU); 3 ex., Ceylon, 30388

(MNHU),; 1 ex., Weligama, 5°58’ N/80°26' E, 1899,

W. Horn (MNHU); 10 ex., Kandy, 7°18’ N/80°36' E,

VI-IX.1907/1916, G. E. Bryant (BMNH); 2 ex., Fra-

serpet, Coorg., VII.1930, F. R. I. Sandal, Insect Survey

(BMNH); 4 ex., Ayur, North Salem, VI—II.1930/31, F.

R. I. Sandal, Insect Survey (BMNH).

Bonn zoological Bulletin 71 (2): 139-176

Redescription

Total length. 4.10—5.80 mm (mean 4.68 mm, n= 10).

Head. Very finely punctuated, entirely brownish-yel-

low. Labrum and mandible black. Antenna slender, ex-

tending almost to the middle of elytra, entirely yellow,

terminal antennomere usually blackish (Fig. 20A). Sec-

ond and third antennomere approximately of the same

length; ratio length of second to third antennomere 0.85—

1.00 (mean 0.97); ratio length of third to fourth antenno-

mere 0.30—0.44 (mean 0.37; Fig. 20B).

Thorax. Pronotum finely punctuated, entirely yellow,

rarely yellowish-brown. Pronotal width 1.20-1.80 mm

(mean 1.52 mm), ratio length to width 0.55—0.59 (mean

0.58). Scutellum brownish. Meso- and metathorax red-

dish to brownish. Elytron reddish-brown with a broad

yellowish transverse band beyond the middle, bounded

above and below by a narrow black band, and another

narrow black band at base. In few specimens, elytron

black with one transverse brown-reddish band at the bas-

al half. Elytral length 2.90-4.50 mm (mean 3.68 mm),

maximal width of both elytra together 2.10—-3.50 mm

(mean 2.73 mm), ratio of maximal width of both elytra

together to length of elytron 0.70—0.78 (mean 0.74). Legs

yellow to yellowish-brown, middle and hindfemur dark

brown, often also hind-taris darker.

Abdomen. Brownish-red.

Male genitalia. Median lobe broad, becomes sig-

nificantly narrowed towards apex. Few strong, slightly

curved median spiculae, lateral spiculae broad, slightly

s-shaped curved (Fig. 20Ca), ventral spiculae short and

strong (Fig. 20Cb).

Female genitalia. Spermatheca with small nodulus,

long, slender and curved cornu (Fig. 20D). Dorsal bursa

sclerites with spines, ventral one only slightly longer than

dorsal spiculae, only with undulate margin (Fig. 20E).

Diagnosis. Monolepta orientalis can be easily distin-

guished by the specific colouration pattern on the elytra

(Figs 19b, 20A) it is, together with size, and some exter-

nal measurements, somewhat similar to M. flavofasciata

(Figs 22a, 23A). In doubtful cases, species delimitation

can be easily done by the significant differences in geni-

tal patterns of both sexes (Figs 20C—E, 23C—E).

Distribution. Widely distributed in the north-west-

ern part of the Oriental Region, in particular in India

(Fig. 21).

Monolepta flavofasciata Jacoby, 1889

Figs 21-23.

Monolepta flavofasciata Jacoby, 1889: 229.

Type material. Jacoby mentioned at least two spec-

imens “Bhamoo, August 1885, also Pulo Penang (coll.

Jacoby)” in his original publication. One syntype is

available in MCGD “Bhamo, Birmania, Fea VHI.1885

/ flavofasciata Jac. / Monolepta flavofasciata Jac”. The

specimen from MCZH (Fig. 22) is not a type, since col-

©LIB

154 Izfa Riza Hazmi & Thomas Wagner

a an: a

M. flavofasciata

a 1

M. orientalis

iss

Sia

Fig. 21. Distribution of Monolepta orientalis Jacoby, 1889 and M. flavofasciata Jacoby, 1889.

leted later and from another locality, but 1s surely con-

specific.

Further material examined. — India. 2 ex., As-

sam, Palkai, 26°12’ N/92°56' E, Fry Coll. 1905.100,

Doherty (BMNH). — Malaysia. | ex., Island of Pen-

ang, 5°15' N/100°29' E, Baker (NHRS); 1 ex., Penang,

5°15' N/100°29' E, G. E. Bryant, XI.1913 (BMNH);

1 ex., Malay Penin. Kedah, Alor Star, Gunong Ker-

lang, 6°11’ N/100°19' E, 1.1V.1928 (UKM); 1 ex., Ma-

lay Penin., West Coast, Langkawi Is., 6°21' N/99°48' E

25.1V.1928 (BMNH); 1 ex., Perak, Tapah, Lata Iskandar,

4°19' N/101°19' E, 3-4.IV.1990, Zaidi, Ismail, Ruslan

(UKM); 1 ex., Pahang, Pulau Tioman, 2°47' N/104°08' E,

16-18. VIII.1994, Ismail & Jazmi (UKM); 1 ex., Kedah,

P. Langkawi, Lubuk Sembilang, 6°21' N/99°48' E,

20.V.1995, Ismail, Ruslan & Sham (UKM); 2 ex., Perlis,

Taman Negeri Wang Klian, 6°40' N/100°11' E, 29.1X—

4.X.1999, Zaidi, Ismail, Azman (UKM). — Thailand.

1 ex., E. Thailand, Nakhon Ratchasima Prov., Sakaer-

Bonn zoological Bulletin 71 (2): 139-176

Fig. 22. Material of Monolepta flavofasciata Jacoby, 1889, no

type, but collected nearby type locality at same time (MCZH).

at Biosphere Reserve, 14°30' N/101°55' E, VI.1995,

Ghazoul & Inward (BMNH).

Redescription

Total length. 4.30—5.80 mm (mean 5.05 mm; n= 0).

Head. Very finely punctuated and entirely pale yellow.

Labrum and mandible black. Antenna slender and extend-

ing almost to the middle of the elytra, pale yellow and

©LIB

Revision of Monolepta Chevrolat, 1836 from the Sundaland area E55

only terminal antennomere partly brownish (Fig. 22b,

23A). Third antennomere slightly longer than second;

ratio length of second to third antennomere 0.75—1.00

(mean 0.88); ratio length of third to fourth antennomere

0.44—0.50 (mean 0.47; Fig.23B).

Thorax. Pronotum very finely punctuated, entirely pale

yellow, parallel-sided and a bit widened posteriorly. Pro-

notal width 1.40—1.80 mm (mean 1.60 mm), ratio length

to width 0.56—0.58 (mean 0.57). Scutellum, meso- and

metathorax black. Elytron finely punctuated, black with

a broad yellowish transverse band beyond the middle and

brown-reddish band in the apical part (Figs 22b, 23A).

Elytral length 3.40-4.60 mm (mean 4.00 mm), maxi-

mal width of both elytra together 2.50—3.30 mm (mean

2.90 mm), ratio of maximal width of both elytra together

to length of elytron 0.72—0.74 (mean 0.73). Foreleg yel-

low, mid and hindlegs blackish (Fig. 22b).

Fig. 23. Monolepta flavofasciata Jacoby, 1889. A. Colour pat-

tern. B. Basal antennomeres. C. Median lobe, a. dorsal, b. later-

al, c. ventral, without endophallic structures. D. Spermathecae.

E. Bursa-sclerites, a. dorsal, b. ventral.

Abdomen. Pale brown to reddish, strongly contrasting

to the black metathorax.

Male genitalia. Median lobe of this species is compar-

atively small, parallel-sided in the basal two thirds and

narrowed in the apical third (Figs 23Ca, Cc). Tectum is

short, narrow in the middle, lanceolate (Fig. 23Ca). Ven-

tral spiculae most likely weakly sclerotised and hardly

visible lateral spiculae broad, simply curved structures

(Figs 23Ca, Cb), median spiculae short, slender bristles

(Figs 23Ca, Cb).

Bonn zoological Bulletin 71 (2): 139-176

Female genitalia. Spermatheca with small spherical

nodulus with broad ridge, long and curved cornu (Fig

23D). Bursa sclerites small and slender (Fig. 23E).

Diagnosis. Elytral colouration of M. flavofasciata is

very characteristic by its broad black base, a small trans-

verse yellow band around the middle, followed by a

broader transverse black band that is reaching the outer

elytral margin, and a reddish elytral apex. Most similar in

elytral colouration is M. orientalis, but with brown-red-

dish elytra with broad yellowish transverse band in the

middle, bounded above and below by a narrow black

band, and another narrow black band is placed across the

basal margin (Figs 19b, 20A, 22b, 23A). The genitalic

characters of both species are very different in both sexes

(Figs 20C-E, 23C-E).

Distribution. Known from East-India, Myanmar and

Thailand to the Malayan Peninsula (Fig. 21).

Monolepta zonula Weise, 1916

Figs 24—26

Monolepta zonula Weise, 1916: 40.

Replacement name for M. fasciatipennis Jacoby, 1892:

983, a junior homonym of M. fasciatipennis Blackburn,

1888: 180.

= Monolepta fasciatipennis Jacoby, 1892: 983 (Weise,

1916), replacement name for M. albofasciata Jacoby,

1889: 228, a junior homonym of M. albofasciata Jacoby,

1884b: 235.

Fig. 24. Type material of Monolepta zonula Weise, 1916. —

Monolepta albofasciata Jacoby, 1889, syntype (MCZH).

Type material. Syntypes: “Bhamo, Birmania, Fea

vill. 1886 / Monolepta albofasciata Jac. / 1 st Jacoby Coll.

/ Type 18443” (MCZH,; Fig. 24), each a further syntype

with the same data in MCGD and BMNH.

Further material examined. — Indonesia. | ex., Java

Occ., MG Piepers (RMNH); 2 ex., Sumatra, Manna,

4°27' S/103°01' E, 1902, M. Knappert (RMNH); 1 ex.,

Sarawak, Kuching, Bako NP, 1°42’ N/110°28' E, V.1999,

P. Votruba (CJB). — Malaysia. 19 ex., Malay Penin, Kuala

Lumpur, 3°06' N/101°39' E, Expd. Agrc. Dept. (BMNH);

1 ex., Sandakan, 5°50' N/118°03' E, C. V. Creagh, 1896

(BMNH); 1 ex., Kuching, 1°31' N/110°20' E, J. E. A.

Lewis, 1910 (BMNH); | ex., Penang, 5°25' N/100°29' E,

©LIB

156 Izfa Riza Hazmi & Thomas Wagner

Fig. 25. Monolepta zonula Weise, 1916. A. Colour pattern.

B. Basal antennomeres. C. Median lobe, a. dorsal, b. lateral,

c. ventral, without endophallic structures. D. Spermathecae.

E. Bursa-sclerites, a. dorsal, b. ventral.

J. E. A. Lewis, 1910-116 (BMNH); 7 ex., Malay Penin,

Kuala Selangor, 3°20' N/101°15' E, [X.1912, Expd.

Agrc. Dept. (BMNH); 4 ex., Malay Penin., Blackwa-

ter Est Klang, 3°02' N/101°26' E, VI.1916, Expd. Agrc.

Dept. (BMNH); 1 ex., Malay Penin, Selangor-Kua-

la Lumpur, 3°30' N/101°31' E, 1929, H. M. Pendle-

bury (BMNH); 1 ex., West Malaysia, Perak, Maxwell

Hills, 4°47' N/100°45' E, 3700-4500 ft., 13.11.1932, H.

M. Pendlebury (BMNH); 1 ex., Malay Penin, Negeri

Sembilan, Port Dickson, 2°32’ N/101°48' E, 21.11.1933

(BMNH); 2 ex., GAP Malaysia, 21.III.1974, Y. Kiy-

oyama (CTJ); 1 ex., Sarawak, Gunong Mulu Nat. Park,

3°55' N/114°46' E, R. G. S. Exped. 24.V1.1977/78, J. D.

Holloway et al. (BMNH); 1 ex., Kuala Lumpur, Jin Panta

Baru, 3°06’ N/101°39' E, 11.11.1982, R. Sulaiman (UKM);

1 ex., N. Sembilan, Hulu Bendul, 2°44' N/102°08' E,

27.11.1989, Kamaruzaman (UKM); 1 ex., Sabah, Pulau

Manukan, 5°58' N/116°E, 1.X.1991, Zaidi & S. Abin

(UKM); 1 ex., West Malaysia, Perak, Maxwell Hills,

4°47' N/100°45' E, 900-1000 m, 12.—16.1.1995, S. Be-

evar (CJB).

Redescription

Total length. 3.50-4.90 mm (mean 4.36 mm, n= 10).

Head. Finely punctuated, entirely blackish. Labrum

and mandibulae blackish. Antenna slender, extending

almost to middle of the elytra, yellowish, the extreme

apex of the last antennomere blackish (Fig. 25A). Sec-

ond and third antennomere roughly of the same length;

ratio length of second to third antennomere 0.87—1.00

(mean 0.97); ratio length of third to fourth antennomere

0.38—0.43 (mean 0.42; Fig. 25B).

Bonn zoological Bulletin 71 (2): 139-176

Thorax. Pronotum very finely punctuated, usually pale

yellow rarely up to pale brown, transversely convex, and

the sides slightly rounded. Pronotal width 1.05—1.55 mm

(mean 1.32 mm), ratio length to width 0.61—0.64 (mean

0.63). Scutellum, meso- and metathorax black. Elytron

finely punctuated, black with one yellow spots in the

middle on disc (Fig. 25A), often apical two thirds along

the suture reddish (Fig. 24b), rarely also with dark-red-

dish spot at humerus. Elytral length 2.75—3.85 mm (mean

3.34 mm), maximal width of both elytra together 2.00—

2.80 mm (mean 2.43 mm), ratio of maximal width of

both elytra together to length of elytron 0.71—0.75 (mean

0.73). Legs yellow to yellowish-brown.

a =M.ranuensissp.n. >

% M-hitam sp. n.

M. zonula

@ 1-2

@ 3-4

@ 5-7

@ s-19

Fig. 26. Distribution of Monolepta zonula Weise, 1916, M. ran-

uensis sp. nov. and M. hitam sp. nov.

Abdomen. Pale yellow to brown.

Male genitalia. Median lobe parallel-sided at base,

strongly narrowed in the apical quarter (Figs 25Ca, Cc).

Median spiculae few short and slender spines, lateral

spiculae broad, spiny structures, ventral spiculae very

fine (Fig. 25C).

Female genitalia. Spermatheca with spherical nodu-

lus having strong ridge, long and curved median part and

short al slender cornu (Fig. 25D). Bursa sclerites, small

and both pairs approximately of same size (Fig. 25E).

Diagnosis. Monolepta zonula looks most similar to

M. signata, M. empatbulat sp. nov. and M. mohamed-

Saidi sp. nov. These four species have black elytron with

circular yellowish spot at least on the basal half of ely-

tron (Figs 14A, 25A, 47A, 56A) in common, with api-

cal elytral quarter usually yellow. Head is usually black

contrasting to the yellow pronotum in M. zonula, M. em-

patbulat sp. nov. and M. mohamedsaidi sp. nov. while

pale brown to reddish-brown in M. signata. From M. em-

patbulat sp. nov. and M. mohamedsaidi sp. nov. it can

be differentiated partly by size (total length zonula

3.50-4.90 mm; M. empatbulat sp. nov. 3.25—3.80 mm,

M. mohammedsaidi sp. nov. 3.35—4.00 mm). In doubtful

cases the median lobe with strong conical shape and the

©LIB

Revision of Monolepta Chevrolat, 1836 from the Sundaland area 157

broad lateral spiculae (Figs 14C, 25C, 47C, 56C) allows

a Clear differentiation.

Distribution. This species is known from Peninsular

Malaysia, Borneo and Sumatra (Fig. 26).

Monolepta mentaweiensis (Jacoby, 1896)

Figs 27-30

Candezea mentaweiensis Jacoby, 1896: 143.

= Monolepta basalis Jacoby, 1884: 55 (Weise 1924:

78).

= Monolepta hageni Weise, 1916: 40; syn. nov.

Candezea mentaweiensis is the next available name

given priority over Monolepta basalis Jacoby, 1884 be-

cause of its junior homonymy over M. basalis Harold,

1880: 26. The senior homonym, described from East

Africa, is meanwhile transferred to Bicolorizea (Heu-

nemann et al. 2015), but the junior homonym cannot be

revalidated after article 57.2. IZCN-Code).

Monolepta mentawiensis

(Jacoby, 1896).

Hazmi & Wagner det. 12

Figs 27-28. Type material of Monolepta mentaweiensis

(Jacoby, 1896). — 27. Candezea mentaweiensis Jacoby, 1896,

syntype (BMNH). 28. Monolepta basalis Jacoby, 1884, lecto-

type, 9 (RMNH).

Type material. Monolepta basalis. Lectotype: 2, “Ma

Loe 1.77 / Monolepta basalis Jac.” (RMNH; (Fig. 27).

Paralectotypes: 4 ex., same data as lectotype (RMNH),

1 ex., “Ma Loe, 10.77” (RMNH), 1 ex., Spjg I.77

(RMNH), | ex., “Ma Loe II.77 / Sumatra Exped. / Ist Ja-

coby Coll. / Type 18461” (MCZAH). A lectotype is herein

designated to fix the name on a single specimen since

Jacoby gave no details on specimen numbers in his ori-

ginal publication.

Candezea mentaweiensis. Syntypes: “Mentawei Sipo-

ra Sereinu V-VI, 94 Modigliani / Museo Civ. Genova /

Candezea mentaweiensis Jac. / Jacoby Coll. 1909-28a /

Co-type” (BMNH; Fig. 28). A further syntype with the

same data is in MCGD. Type locality: 2°10' S/99°41' E.

Bonn zoological Bulletin 71 (2): 139-176

Further material examined. — Brunei. | ex., Labi,

Bukit Teraja, 4°18’ N/114°26' E, 60 m, 23.VIII.79, S. L.

Sutton (BMNH). — Indonesia. | ex., Sumatra Exped.,

Jacoby Coll. 1909-28a (BMNH); | ex., Sumatra, Manna,

4°27' S/102°59' E, M. Knappertt., Coll. Vett (RMNH):

1 ex., N. O. Sumatra, Tandjong Morawa, Serdang,

0°35' S/101°18' E, Dr. B. Hagen (BMNH); 7 ex., Suma-

tra, Moera Laboe, XI.1877 (RMNH); 4 ex., Borneo, Mt.

Tibang, Mjoberg (NHRS),; 1 ex., East Borneo, Sanga San-

ga, 0°38' S/117°12' E, 1907-203, H. D. Jansen (BMNH):

1 ex., Sumatra, Moera Laboe, X.1877 (RMNH); 1 ex., Sar-

awak, Kapit distr., Sebong, 1°33' N/114°17' E, HI.1994, J.

Horak (CJB). — Malaysia. 2 ex., Borneo (BMNH); 7 ex.,

Perak, 4°48' N/101°09' E, Doherty, Fry Coll. 1905.100,

Jacoby Coll. (BMNH); 1 ex., Borneo, Baly Coll.

(BMNH); 1 ex., Penang, 5°15' N/100°29' E, (Lamb.),

Pascoe Coll. (BMNH); 1 ex., Sarawak, Bidi, 411.1909,

C. J. Brooks (BMNH); 7 ex., West Sarawak, Mt. Matang,

1°34’ N/110°16' E, 16.—30.X1I.1913, 1.1914, G. E. Bryant

(BMNH); 3 ex., W. Sarawak, Lundu, 1°40’ N/109°48' E,

1.1914, G. E. Bryant (BMNH); 1 ex., Malay Penin, Se-

langor-Pahang, 2700 ft., 3°30' N/101°31' E, 1.1915, ex.

F. M. S. (BMNH); 3 ex., Malay Penin. Kedah nr Jitra,

catchment area, 6°15' N/100°25' E, 4.-7.1V.1928, H. M.

Pendlebury (BMNH); 1 ex., Malay Penin, Pahang, F. M.

S., Fraser Hills, 4200 ft., 3°42’ N/101°41' E, 11.TX.1931,

H. M. Pendlebury (BMNH); 1 ex., Sarawak, Matang,

1°34’ N/110°17' E, 10.V.1909, C. J. Brooks (BMNH);

1 ex., Kuala Lumpur, Gardens, 3°08' N/101°41' E,

18.X11.1938, H. M. Pendlebury (BMNH); 1 ex., Lang-

kawi Island, 6°21’ N/99°49' E, 30.IV.1928 (BMNH);

ZAR

Fig. 29. Monolepta mentaweiensis (Jacoby, 1896). A. Colour

pattern. B. Basal antennomeres. C. Median lobe, a. dorsal,

b. lateral, c. ventral, without endophallic structures. D. Sperma-

thecae. E. Bursa-sclerites, a. dorsal, b. ventral.

©LIB

158 Izfa Riza Hazmi & Thomas Wagner

1 ex., Selangor, Bukit Kutu, 3500 ft., 3°33' N/101°43' E,

18.11.1931, H. M. Pendlebury (BMNH); 2 ex., Perak, F.

M. S. Larut Hills, 3700 ft., 5°01’ E/100°53' E, 14.11.1932,

H. M. Pendlebury (BMNH); 1 ex., Pahang, Taman Nega-

ra, 3°58' N/102°26' E, 1.-13.]II.1984, L. Jessop (BMNH);

1 ex., Sabah, Poring, 6°02' N/116°42’ E, 6.XI.1986, F.

Abang (UKM); 1 ex., Sabah, nr Danum Valley Field,

5°25' N/118°23' E, 20.VI.—12.VII.1987, C. v. Achther-

berg & D. Kennedy (RMNH); 1 ex., Kedah, Langkawi,

Lubuk Sembilang, 6°21' N/99°48' E, 7.1V.1993, Ismail,

Sham, Yusof (UKM); 1 ex., Perak, Temenggor, Ekspedi-

si MNS-Belum, 5°33’ N/101°20' E, 29.X1.1993, Ismail,

Yusof, Bidi & Saiful (UKM); 1 ex., Perlis, Wang Kelian,

6°40' N/100°11' E, 19. VII.1994, Salleh, Ismail & Ruslan

(UKM); 3 ex., Terengganu, Setiu, H. Lipur Peladang,

5°27' N/102°45' E, V.1994, Ismail & Zabidi (UKM);

1 ex., Kedah, Pulau Langkawi, Gunung Raya, 933 m,

6°21' N/99°48' E, 21.V.1995, Ismail, Ruslan & Sham

(UKM); 6 ex., Sabah, Danum Valley, 5°25’ N/118°23' E,

6.—15.V.2007, 11.EX.2007, 11.[X.2007, B. H. Izfa(UKM);

5 ex., Sabah, Lembah Danum, 5°25' N/118°23' E, 16.-

19.V.1991, 27-31.VIIL.1991, 3.-5.XII.1991, 22-25.

VIII.1992, 6.V.2007, Ismail et al. (UKM); — Singapore.

2 ex., Singapore, 1°21' N/103°49' E, 97-109, H. N. Rid-

ley (BMNH); 1 ex., Singapore, H. N. Ridley (BMNH).

— Thailand. | ex., Siam, Renong, 13°45’ N/100°29' E,

Doherty, Fry Coll. (BMNH); 2 ex., Nakon Sri Tamarat,

Khao Ram, 1500-2900 ft., 17°01' N/99°49' E, 28.IT.1922,

H. M. Pendlebury (BMNH); 1 ex., Phang-nga Prov.,

5 km S Kao Lak, 8°36' N/98°15' E, VIII.2014, A. Skale

(NME). — Vietnam. 3 ex., Na Hang, 160 km, NNW Ha-

noi, 22°16' N/105°17' E, 200 m, VI.1996, A. Napolov

(NME).

Redescription

Total length. 4.90-6.25 mm (mean 5.91 mm, n= 10).

Head. Very finely punctuated, brown-reddish and frons

partly black. Labrum and mandible black. Antenna slen-

der, extending almost to half of the elytra, rarely either

entirely yellowish, and than often with brownish parts in

middle of antennomeres (Fig. 28b), or brownish or black

with two apical yellowish antennomeres (Fig. 29A).

Third antennomere slightly longer than second; ratio

length of second to third antennomere 0.75—1.00 (mean

0.80); ratio length of third to fourth antennomere 0.38—

0.50 (mean 0.42; Fig. 29B).

Thorax. Pronotum finely punctuated, entirely red-

disch-brown and posterior sides rounded and basally

widened. Pronotal width 1.60—2.00 mm (mean 1.84 mm),

ratio length to width 0.53—0.57 (mean 0.55). Scutellum,

meso- and metathorax reddish-brown. Elytron distinct-

ly widened behind, yellow to reddish-brown and in

most specimens darker brownish-red in the basal third

(Figs 27b, 29A). Elytral length 3.90-5.00 mm (mean

4.68 mm), maximal width of both elytra together 2.80—

3.70 mm (mean 3.49 mm), ratio of maximal width of

Bonn zoological Bulletin 71 (2): 139-176

mm |

alte |

= |

| a

Fig. 30. Distribution of Monolepta mentaweiensis (Jacoby,

1896) and M. putri Mohamedsaid, 2001.

both elytra together to length of elytron 0.70—0.78 (mean

0.74). Legs brown-reddish.

Abdomen. Yellow, contrasting paler to the more

brownish metathorax.

Male genitalia. Median lobe broad at base and more

Slender in the apical half (Fig. 29C). Median spiculae

comparatively short and slender, three lateral spiculae as

short broad spines or brush-like structures, ventral spic-

ulae very small and weakly sclerotised (Figs 29Ca, Cb).

Female genitalia. Spermatheca with large spher-

ical nodulus, middle part and cornu short and broad

(Fig. 29D). Bursa sclerites weakly sclerotised (Fig. 29E).

Diagnosis. In colouration, Monolepta mentaweiensis is

most similar to M. bruneiensis sp. nov., both species have

a slight contrast between the brown-reddish pronotum

the paler elytra. In most cases, M. mentaweiensis pos-

sesses a brown-reddish elytral base that does not occur

in M. bruneiensis sp. Monolepta mentaweiensis (4.90-

6.25 mm) is much larger than M. bruneiensis sp. nov.

(3.75—4.70 mm). The frons and mouthpart are partly

black in M. mentaweiensis while in M. bruneiensis, it is

brown-reddish as the head. The genitalic characters are

quite different between these two species (Figs 29C-—E,

44 C-E).

Distribution. Northern Vietnam to Thailand through

Malay Pensinsula towards Sumatra and Borneo (Fig. 30).

Monolepta militaris Jacoby, 1896

Figs 31-33

Monolepta militaris Jacoby, 1896: 484.

Type material. Syntypes: “Sumatra Si-Rambe, XII.90—

I1I.91, E. Modigliani / Monolepta militaris Jac. / Jacoby

Coll. 1909-28a / Type” (BMNH; Fig. 31). Further 5 syn-

types (3 ex. BMNH, at least 1 ex. MCGD, 1 ex. MCZH)

with the same data. Type locality: 5°24' S/105°17' E.

Further material examined. — India. | ex., India, occ.

Centr. GOA prov, 15°47»N/74°02)E, 12.—14. VIII.2002,

©LIB

Revision of Monolepta Chevrolat, 1836 from the Sundaland area 159

Fig. 31. Type material of Monolepta militaris Jacoby, 1896,

syntype (BMNH).

P. Sipek & M. Fikacek (CJB). — Indonesia. 4 ex., Su-

matra, Singalang, Sibajakvulkan, 0°25' S/100°20' E,

Myjoberg (NHRS); 4 ex., N. Sumatra, Bivouac two Mt.

Bandahara, 3°45' N/97°45' E, 5-10. VII.1972, J. Krikken

(RMNH). — Malaysia. 1 ex., N. Sembilan, Gemencheh,

2°35' N/102°24' E, 5.VHI.1990, Mahbob, Fog Perigen

(UKM).

Redescription

Total length. 3.70—5.40 mm (mean 4.55 mm; n= 10).

Head. Very finely punctuated, entirely yellow-

ish-brown to black. Labrum and mandible black. An-

tenna slender, extending to the middle of the elytra, yel-

lowish and usually two terminal antennomeres brownish

(Fig. 32A), sometimes also middle of antenna brownish.

Third antennomere slightly longer than second; ratio

length of second to third antennomere 0.75—1.00 (mean

0.88); ratio length of third to fourth antennomere 0.33—

0.50 (mean 0.41; Fig. 32B).

Thorax. Pronotum finely punctuated, entirely yel-

low. Pronotal width 1.20-1.60 mm (mean 1.34 mm),

P= ¢%

Fig. 32. Monolepta militaris Jacoby, 1896. A. Colour pattern.

B. Basal antennomeres. C. Median lobe, a. dorsal, b. lateral,

c. ventral, without endophallic structures. D. Spermathecae.

E. Bursa-sclerites, a. dorsal, b. ventral.

Bonn zoological Bulletin 71 (2): 139-176

ratio length to width 0.65—0.67 (mean 0.66). Scutellum,

meso- and metathorax black. Elytron pale brownish-red,

black transverse band in the basal third. Elytral length

3.00-4.10 mm (mean 3.56 mm), maximal width of both

elytra together 2.30—2.75 mm (mean 2.66 mm), ratio of

maximal width of both elytra together to length of ely-

tron 0.73-0.77 (mean 0.75; Figs 31b, 32A). Legs yellow,

often hindfemur, rarely additionally mid femur blackish

without the distal tip.

" Le

be Sy

. “ M. rufipennis Y (-

go P q

Mm 2-3 : y

ee! — } a

M. militaris

@ 1-3

@ 4-6

Fig. 33. Distribution of Monolepta militaris Jacoby, 1896, and

Monolepta rufipennis Jacoby, 1899.

Abdomen. Yellowish-brown, strongly contrasting to

the dark metathorax.

Male genitalia. Median lobe long and slender. Paral-

lel-sided and slightly conical towards apex, twisted be-

fore apex (Figs 32Ca, Cc). Median spiculae short and

brush-like structures at base and one pair of long, broad

spines, lateral spiculae long, strong with c-shaped api-

cal part, ventral spiculae jagged-liked with three spines

(Figs 32Ca, Cb).

Female genitalia. Spermatheca with comparatively

small nodulus and strong ridge, cornu long slender and

strongly curved (Fig. 32D). Two pairs of bursa sclerites

strongly serrate and roughly of same size (Fig. 32E).

Diagnosis. Monolepta militaris is a medium-sized

species of Monolepta that can be easily distinguished

by dorsal colouration. Elytra are brown-reddish, with

black transverse band at base, contrasting to a pale yel-

low pronotum (Fig. 32A). Monolepta rufipennis has also

reddish elytra, but can be differentiated from M. militaris

by a transverse band stretch across the basal elytral third

(Fig. 37). The median lobe of this species is very differ-

ent from M. rufipennis (Figs 32C, 37C).

Distribution. Known from southern India, Sumatra

and Java (Fig. 33).

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160 Izfa Riza Hazmi & Thomas Wagner

Monolepta marginicollis Jacoby, 1896

Figs 9, 34-35

Monolepta marginicollis Jacoby, 1896: 485.

Type material. Syntypes: “Sumatra Si-Rambe, XII.90—

111.91, E. Modigliani / Monolepta marginicollis Jac. /

Museo Civ. Genoa / Jacoby Coll. 1909-28a / Co-type”

(BMNH,; Fig. 34). One further sytpe with the same data

in MCGD. Type locality: 6°11' S/106°48' E.

Fig. 34. Type material of Monolepta marginicollis Jacoby,

1896, syntype (BMNH).

Further material examined. — Brunei. 4 ex., Labi,

Bukit Teraja 60 m, Mxt. Dipt. Forest, 4°18' N/114°26' E,

21.VIII.1979, B.M. 1983-39, Light trap, 1.75 m above

ground, S. L. Sutton (BMNH); 7 ex., Bukit Sulang nr

Lamunin, 4°39' N/114°44' E, 20. VIII. —-10.1X.1982, NE

Stork (BMNH); 1 ex., Kuala Belalong FSC, Diptero-

carp Forest, 4°29' N/115°11' E, 16.V1I.1991, N Mawdsley

(BMNH); 2 ex., Temburung District Ridge, NE of Kuala

Belalong, 4°37' N/115°8' E, 300 m, X.1992, Light Trap,

J. H. Martin (BMNH). — Indonesia. | ex., N Sumatra,

Bivouac Two, Mt Bandahara, 3°44’ N/97°43' E, 5-10.

VII.1972, J. Krikken (RMNH). — Malaysia. 3 ex., Bor-

neo, Mt. Tibang, 1700 m, Mjoberg (NHRS); 3 ex., Mt.

Matang, West Sarawak, XII.1913—I1.1916, G. E. Bryant

(BMNH),; 1 ex., Malay Penin, Kedah Peak, 16.11.1928

(BMNH); 1 ex., Sarawak, Engkelili, 1°8' N/111°39' E,

27.VIII.1967, Vincent Coll. (BMNH); 3 ex., Sarawak,

Gunong Mulu Nat. Park, 3°55' N/114°47' E, R. G. S.

Exped. 24.VI.1977, J. D. Holloway et al. (BMNH);

11 ex., Sabah, Tawau Plat. 1300 ft., 8 m S. Telupid,

5°35' N/117°07' E, 8.[X.1977, M. E. Bacchus (BMNH);

19 ex., Sarawak, 4th Division Gn. Mulu NP, nr. Base

Camp 50-100 m, 3°55’ N/114°46' E, V—VII.1978, P.

M. Hammond & J. E. Marshall (BMNH); 1 ex., Malay-

sia-SW, Sabah nr Long Pa Sia (west), 5°20' N/117°10' E,

1200 m, 2.—-14.1V.1987, C. v. Achterberg (RMNH);

2 ex., Malaysia, SE Sabah nr Danum Valley Field C,

5°25' N/118°23' E, 15. —25.II1.1987, C. v. Achterberg &

D. Kennedy (RMNH); 1 ex., N. Borneo, Sabah, 60 km of

Lahad Datu DVFC, EO sg. Segama, 4°58’ N/117°48' E,

18.X.1987, J. Huisman & R. de Jong (RMNH); | ex., Sa-

bah, Sandakan, Sepilok FR (obs tower), 5°51' N/118°0' E,

1.X1.1987, Krikken & Rombaut (RMNH); 1 ex., Pa-

hang, Kuala Lompat, 3°41' N/102°11' E, 27. VIII.1990,

Bonn zoological Bulletin 71 (2): 139-176

Fog Malathion, Mahbob (UKM); 1 ex., Sabah, Lembah

Danum, 5°25’ N/118°23' E, 3.-5.XII.1991, Zaidi et al.

(UKM); 1 ex., N. Borneo, Trus Madi, 5°43' N/116°25' E,

11.1V.1993, H. Karube (CTJ); 1 ex., Sarawak, Lundu,

T. Negara Gunung Gading, 1°44' N/109°50' E, 22-27.

IV.1994, Salleh dan Ismail (UKM); 1 ex., W. Per-

ak, 25 km NE Ipoh, Banjaran Tititwangsa Mts. Korbu,

4°41' N/101°18' E, 6.-12.V.2001, M. Riha (CJB).

Fig. 35. Monolepta marginicollis Jacoby, 1896. A. Colour pat-

tern. B. Basal antennomeres. C. Median lobe, a. dorsal, b. later-

al, c. ventral, without endophallic structures. D. Spermathecae.

Redescription

Total length. 3.25—4.00 mm (mean 3.55 mm, n= 10).

Head. Very finely punctuated, yellowish to red-

dish-brown, vertex often with short median black stripe.

Labrum and mandible black. Antenna slender, extending

almost to the middle of elytra, blackish, and three basal

antennomeres yellow-brownish (Fig. 35A). Third an-

tennomere significantly longer than second; ratio length

of second to third antennomere 0.67—0.75 (mean 0.73);

ratio length of third to fourth antennomere 0.43—0.67

(mean 0.62) (Fig. 35B).

Thorax. Pronotum finely punctuated, yellow to brown-

ish-yellow, lateral and basal margin black (Figs 34b,

35A). Pronotal width 1.00-1.20 mm (mean 1.06 mm),

ratio length to width 0.60—0.64 (mean 0.62). Scutellum,

meso- and metathorax blackish. Elytron finely punctuat-

ed, yellowish, all margins, including suture, black. Elytral

length 2.45—2.85 mm (mean 2.68 mm), maximal width of

both elytra together 1.80—2.10 mm (mean 1.96 mm), ra-

©LIB

Revision of Monolepta Chevrolat, 1836 from the Sundaland area 161

tio of maximal width of both elytra together to length of

elytron 0.71—0.74 (mean 0.73). Legs entirely yellowish.

Abdomen. Yellowish, strongly contrasting to the black

metathorax.

Male genitalia. Median lobe slender, parallel-sid-

ed, conidal in the apical third. Tectum pointed. Median

spiculae three broad spines, further brush-like structures

at sacculus (Figs 35Ca, Cb), lateral spiculae short spurs

close to apex, ventral spiculae as two fattened, spiny

structures (Figs 35Ca, Cb).

Female genitalia. Spermatheca with spherical com-

paratively big nodulus and small ridge. Cornu very long

and curved (Fig. 35D). Bursa sclerite very weakly sclero-

tised and hardly visible.

Diagnosis. Most similar to M. marginicolloides sp. nov.

on the first glance. Both species have yellow elytra with

a Slightly stripy appearance and narrow black outer ely-

tral margins and suture in common (Figs 34A, 52b). An-

tennomeres in both species are blackish and only three

basal antennomeres yellowish. Monolepta marginicollis

is on average smaller (3.25—4.00 mm) than M. margini-

colloides sp. nov. (3.70—5.10 mm). Other characters to

differentiate these two species are the yellow-brownish

head and vertex with short central black stripe in M. mar-

ginicollis, while head in M. marginicolloides sp. nov. is

completely black. The genitalic characters are very dif-

ferent, and allow a clear differentiation of both species in

any doubtful cases (Figs 34C, D, 53C-E).

Distribution. Known from Sumatra, Malayan Penin-

sula and Borneo (Fig. 9).

Monolepta rufipennis Jacoby, 1899

Figs 33, 36-37

Monolepta rufipennis Jacoby, 1899: 325.

Type material. Lectotype: “Perak / Monolepta rufipennis

Jac. / Jacoby Coll. 1909-28a / Type” (BMNH; Fig. 36).

Type locality: 4°48)N/101°09)E. Jacoby mentioned two

localities in his original publication and thus we desig-

nate a lectotype herein to fix the name on a single spec-

imen.

Further material examined. — Malaysia. 2 ex.,

Perak, Doherty, 4°48’ N/101°09' E, Fry Coll. 1905.100

(BMNH); 2 ex., Perak, F. M. S. Larut Hills, 3700 ft.,

5°N/100°53' E, VII.1908; 14.11.1932, H. M. Pendle-

Fig. 36. Type material of Monolepta rufipennis Jacoby, 1899,

lectotype (BMNH).

Bonn zoological Bulletin 71 (2): 139-176

bury (BMNH); 1 ex., Malay Penin, Selangor, Gombak

Valley, 3°25' N/101°47' E, 11.%.1921, H. M. Pendlebury

(BMNH); 3 ex., Perak, FM. S. Batang Padang Jor Camp,

3°54' N/101°26' E, 31.V.—5.VI.1923, H. M. Pendlebury

(BMNH); | ex., Pahang, F. M. S. Cameron’ s Highlands,

4000-4500 ft., 4°27' N/101°22’ E, 15.VI.1935, H. M.

Pendlebury (BMNH); 1 ex., Perak, Temenggor, Ekspe-

dist MNS-Belum, 5°35' N/101°21' E, 10.—-15.V.1994, Is-

mail & Sham (UKM).

Fig. 37. Monolepta rufipennis Jacoby, 1899. A. Colour pattern.

B. Basal antennomeres. C. Median lobe, a. dorsal, b. lateral,

c. ventral, without endophallic structures. D. Spermathecae.

E. Bursa-sclerites, a. dorsal, b. ventral.

Redescription

Total length. 4.75—6.35 mm (mean 5.61 mm; n= 10).

Head. Very finely punctuated, entirely black. Anten-

na slender, extending beyond the middle of elytra, pale

yellow yellowish-brown, two basal antennomeres black

(Fig. 37A). Third antennomere slightly longer than sec-

ond; ratio length of second to third antennomere 0.80—

0.83 (mean 0.81); ratio length of third to fourth antenno-

mere 0.46—0.50 (mean 0.48; Fig. 37B).

Thorax. Pronotum very finely punctuated, entire-

ly black, shiny, surface rather convex. Pronotal width

1.35-1.90 mm (mean 1.64 mm), ratio length to width

0.61—0.64 (mean 0.62). Scutellum brown reddish. Meso-

and metathorax black. Elytron entirely reddish-brown

(in living specimens most likely carmine red), dense

and finely punctuated, widened posteriorly. Elytron

length 4.30—5.05 mm (mean 4.57 mm), maximal width

of both elytra together 3.20—3.70 mm (mean 3.39 mm),

ratio of maximal width of both elytra together to length

of elytron 0.73—-0.75 (mean 0.74; Fig. 54). Legs entirely

brownish-red to blackish, distal part of femora paler.

Abdomen. Blackish.

©LIB

162 Izfa Riza Hazmi & Thomas Wagner

Male genitalia. Median lobe lanceolate, broad, be-

comes continuously narrow towards apex. Tectum point-

ed. Median spiculae long and slender, one pair shorter

and stronger (Figs 37Ca, Cb), lateral spiculae, short,

broad, lobe-like, ventral spiculae weakly sclerotised and

not clearly visibly.

Female genitalia. Nodulus reduced, middle part and

especially cornu long and curved (Fig. 37D). Bursa

sclerites large, of two different types, the dorsal one with

strong spines (Fig 37Ea).

Diagnosis. Monolepta rufipennis can be distinguished

from others Monolepta species by the black pronotum

and head contrasting to the reddish-brown elytra. Con-

cerning body size and dark dorsal colouration and brown-

ish-reddish elytra it is most similar to M. rubra. Both

species are presumably closely related and form — next

to some others — a distinctive clade within the Oriental

species of Monolepta characterized by a lanceolate me-

dian lobe with simple brush-like median spiculae, lobed

lateral spiculae and the lack of ventral spiculae (Figs 8C,

37C), and spermathecae possessing extremely reduced

nodulus (Figs 8D, 37D), and two large, and strongly dif-

ferent pairs of bursa sclerites (Figs 8E, 37E). Both spe-

cies can be in doubtful cases differentiated by this fine

but constant genitalic differences, furthermore, is their

distribution allopatric (Figs 9, 33).

Distribution. This species is only known from the Ma-

lay Peninsula (Fig. 33).

Monolepta tiomanensis Mohamedsaid, 1999

Figs 38-40

Monolepta tiomanensis Mohamedsaid, 1999: 247.

Type material. Holotype: “Pahang, Pulau Tioman, 6-9

Jun 1990, Zaidi, Ismail, Ruslan / Holotype Monolepta tio-

manensis n. sp. Mohammedsaid 1997” (UKM; Fig. 38).

— Paratypes. 2 ex., same data as holotype (UKM). Type

locality: 2°47' N/104°10! E.

Fig. 38. Type material of Monolepta tiomanensis Mohamed-

said, 1999, holotype (UKM).

Further material examined. — Brunei. 1 ex.,

Temburong District, ridge NE of Kuala Belalong,

Bonn zoological Bulletin 71 (2): 139-176

4°37' N/115°8' E, 300 m, X.1992 (BMNH). — Indonesia.

1 ex., Sibolangit, Sumatra, 3°18’ N/98°34' E, Myjéberg

(NHRS); 1 ex., Java, Gg. Moeria, Tjolo 700-1000 m,

7°36' S/110°42' E, 28.XII.1973, P. H. v. Doesburg Col-

lectie Van Doesburg (RMNH); 1 ex., Sumatra Utara,

Pasar Baru, 0°35' S/101°20' E, 26.II1.1996, K. Maruyama

(CTJ). — Malaysia. | ex., South China Sea, Pulau Tioman,

Sedagang at light, 2°47' N/104°10' E, V.1927, N.S.M.R.H

(BMNH); 2 ex., Perlis, Kaki Bukit, Wang Kelian,

6°40' N/100° E, 7.XII.1992, 19.VII.1994, Zabidi et al.

(UKM); 3 ex., Pahang, Pulau Tioman, 2°47' N/104°' E,

22 —28.IV.1993, Zaidi et al. (UKM);1 ex., N. Sembilan,

Serting Ulu, 2°59' N/102°E, 30.1II.1995, Ismail & Rus-

lan (UKM); 3 ex., N. Sembilan, Rembau, Gunong Datok,

2°34' N/102°3' E, 30. VII.1995, Ismail & Sham (UKM);

1 ex., Sarawak, Mt. Dulit, 4000 ft., 3°20' N/114°9' E,

21.X.1932, Moss Forest (BMNH); 2 ex., Perak, F.

M. S. Batang, Padang Jor Camp, 4°48’ N/100°48' E,

11.111.1924/1925, H. M. Pendlebury (BMNH). — Sin-

gapore. 2 ex., Singapore, 1°21' N/103°49' E, H. N. Rid-

ley, 1904/3 (BMNH).

Fig. 39. Monolepta tiomanensis Mohamedsaid, 1999. A. Co-

lour pattern. B. Basal antennomeres. C. Median lobe, a. dorsal,

b. lateral, c. ventral, without endophallic structures. D. Sperma-

thecae. E. Bursa-sclerites, a. dorsal, b. ventral.

Redescription

Total length. 4.75—6.15 mm (mean 5.56 mm; n= 10).

Head. Nearly impunctate, entirely black with very

large eyes. Antenna slender, long, extending towards

the apical third of elytra, first antennomeres brownish

to black at base, second to terminal antennomere yellow

(Fig. 39A). Third antennomere slightly longer than sec-

ond; ratio length of second to third antennomere 0.75—

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Revision of Monolepta Chevrolat, 1836 from the Sundaland area 163

1.00 (mean 0.88); ratio length of third to fourth antenno-

mere 0.27—0.45 (mean 0.36; Fig. 39B).

Thorax. Pronotum entirely pale yellow, transversely

convex, anterior side broadened. Pronotal width 1.40-—

1.80 mm (mean 1.64 mm), very broad, ratio length to

width 0.57-0.61 (mean 0.59). Scutellum, meso- and

metathorax yellow. Elytra punctuated like pronotum,

yellowish and elongated. Elytral length 3.60-4.50 mm

(mean 4.14 mm), maximal width of both elytra together

2.50-3.20 mm (mean 2.83 mm), ratio of maximal width

of both elytra together to length of elytron 0.66—0.70

(mean 0.68; Fig. 38). Legs entirely yellowish.

Abdomen. Yellow.

L— Vv

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M. tiomanensis ak \ I LL | ' f J +

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Fig. 40. Distribution of Monolepta tiomanensis Mohamedsaid,

1999, M. kuninghitam sp. nov., and M. sulawensis sp. nov.

Male genitalia. Median lobe short, broad, parallel-sid-

ed and rounded at apex. Tectum broad and nearly reach-

ing the apex of median lobe. Median spiculae are strong,

long, and horn-shaped, ventral spiculae comb-like, and

lateral spiculae spur-like, close to the apex of median

lobe (Fig. 39C).

Female genitalia. Spermatheca with large spherical

nodulus, cornu long and curved (Fig. 39D). Two pairs of

bursa sclerites of similar size, both carrying strong spines

(Fig. 39E).

Diagnosis. Monolepta tiomanensis can be distin-

guished by the entirely yellowish abdomen and thorax

to the contrasting black head. Entirely yellow dorsum is

found also in M. jacobyi, where head is not black, but the

narrow base of elytra. This species is also less slender

and has a less broad pronotum than M. tiomanensis (Fig-

S17A, 39A). There are several other species described in

Monolepta with throughout yellow colouration and large

eyes in the Oriental Region. This pattern is usually cor-

related with nocturnal activity. But all other species with

this general pattern belong not to “true” Monolepta; a

genital check is senseful in all doubtful cases.

Distribution. Known from Borneo (Malaysia, Brunei)

Singapore, Sumatra and Java (Fig. 40).

Bonn zoological Bulletin 71 (2): 139-176

Monolepta putri Mohamedsaid, 2001

Figs 30, 41-42

Monolepta putri Mohamedsaid, 2001: 137-169.

Type material. Holotype: 9 “Asah Panji, Lake Tamblin-

gan, C. Bali 6 V 1998, H. Takizawa / Holotype Mono-

lepta putri n. sp. des. Mohammedsaid 2000” (CTJ). —

Paratypes: 1 2, same data as holotype (UKM, Fig. 41);

1 2, same data, but 20.[X.1998 (UKM). Type locality:

8°15’ S/115°05' E.

Asah Panji pa

Lake Tamblingan

C. Bali 6 V .

1998 H.Takizawa

Fig. 41. Type material of Monolepta putri Mohamedsaid, 2001,

paratype (UKM).

Further materials examined. — Indonesia. | ex.,

C Bali, Candikuning, Kebun Raya, alt m 1400, 27.-—

31.X.1991, Krikken, Huijbregts, de Vries, multistr evergr

forest (degraded) (RMNH); | ex., same data as holotype,

but 25.V.2005 (UKM).

Redescription

Total length. 4.60—4.90 mm (mean 4.74 mm; n= 4).

Head. Very finely punctuated, yellowish, shiny. La-

brum and mandible blackish. Antenna slender, extended

to the middle of elytron, entirely yellowish (Fig. 42A).

Third antennomere slightly longer than second; ratio

Fig. 42. Monolepta putri Mohamedsaid, 2001. A. Colour pat-

tern. B. Basal antennomeres. C. Spermatheca. D. Bursa-scler-

ites, a. dorsal, b. ventral.

©LIB

164 Izfa Riza Hazmi & Thomas Wagner

length of second to third antennomere 0.75—1.00 (mean

0.88); ratio length of third to fourth antennomere 0.38—

0.44 (mean 0.41; Fig. 42B).

Thorax. Pronotum very finely punctuated, nearly

parallel-sided, shiny, entirely yellowish. Pronotal width

1.40-1.45 mm (mean 1.41 mm), ratio length to width

0.64—0.66 (mean 0.65). Scutellum, meso- and metatho-

rax blackish. Elytra entirely yellowish, basal margin, hu-

merus, basal one-fourth of lateral margin, a stripe beyond

humerus, and basal one fourth of suture black, given this

colouration a “saddle-like” structure (Figs 41b, 42A).

Elytral length 3.60-3.80 mm (mean 3.71 mm), maxi-

mal width of both elytra together 2.50—2.70 mm (mean

2.63 mm), ratio of maximal width of both elytra together

to length of elytron 0.70—0.72 (mean 0.71). Legs yellow.

Abdomen. Yellowish.

Male genitalia. Not known.

Female genitalia. Spermatheca with small spherical

nodulus. The middle part is long, cornu short (Fig. 42C).

The two pairs bursa sclerites of nearly equal size, both

with strong spines (Fig. 42D).

Diagnosis. Monolepta putri looks most similar to

M. kuninghitam sp. nov. Both species have yellowish

elytra with black basal and subhumeral margins. Head

and pronotum are yellowish, and mouthparts are black

in both species. The differences between M. putri and

M. kuninghitam sp. nov. are the yellow antenna of M. pu-

tri, while it is black in M. kuninghitam sp. nov. Pronotum

in M. putri is broader (ratio length to width 0.64—0.66)

than in M. kuninghitam sp. nov. (0.71), and the shape of

spermatheca are different. That of MZ putri is with spher-

ical nodulus, while in M. kuninghitam sp. nov., sperma-

theca is very small (42C), most similar to the presumably

closer related M. bifasciata, M. rubra, and M. rufipennis

(5D, 8D, 37D).

Distribution. Only known from Bali (Fig. 30).

Monolepta bruneiensis sp. nov.

urn. |sid:zoobank.org:act:AEQOCBE3-4EC8-4F 4D-A 7A0-08C866CC2279

Figs 43-45

Type material. Holotype: ¢ “Brunei, E 115 7'N4 34’ ,

Kuala Belalong FSC, Dipterocarp forest, Dryobalanops

beccariil / BM (NH) 1991-173 / Aerial Malaise 1A 260 m,

0 ‘

aaa ag] ri

8) F455

to. ol" az

(ESSee ge.

wee eNO.

520th GHG

weege SS

a ae 3 '

(22882 $85 ) \

(§288 282 /

2 eel Sed PIPrirrey|

Fig. 43. Type material of Monolepta bruneiensis sp. nov., ho-

lotype (BMNH).

Bonn zoological Bulletin 71 (2): 139-176

alt. 2.vi.91 / N. Mawdsley. NM 175” (BMNH; Fig. 43).

Type locality: 4°34' N/115°07' E. — Paratypes: 4 ex., same

data as holotype (BMNH).

Etymology. The name of the new species refers to the

state where it exclusively occurs.

Fig. 44. Monolepta bruneiensis sp. nov. A. Colour pattern.

B. Basal antennomeres. C. Median lobe, a. dorsal, b. lateral,

c. ventral, without endophallic structures. D. Spermathecae.

E Bursa-sclerites, a. dorsal, b. ventral.

Description

Total length. 3.75—4.70 mm (mean 4.26 mm; n = 6).

Head. Very finely punctuated, entirely brownish-red

and shining. Labrum and mandible as head, apically a

bit darker. Eyes very large. Antenna slender, extending

almost to the middle of elytra, blackish and only three

basal antennomeres yellow-brownish (Fig. 44A) or the

terminal antennomeres more brownish (Fig. 43b). Sec-

ond antennomere significantly longer than third; ratio

length of second to third antennomere 1.00—1.50 (mean

1.25); ratio length of third to fourth antennomere 0.25—

0.29 (mean 0.27; Fig. 44B).

Thorax. Pronotum finely punctuated, entirely brown-

ish-red, broad. Pronotal width 1.15—1.30 mm (mean

1.24 mm), ratio length to width 0.56—0.58 (mean 0.57).

Scutellum, meso- and metathorax brownish-red. Elytra

entirely yellow to yellowish-brown. Elytral length 2.90—

3.55 mm (mean 3.19 mm), maximal width of both elytra

together 2.20—2.70 mm (mean 2.46 mm), ratio of max-

imal width of both elytra together to length of elytron

0.75—0.79 (mean 0.77; Fig. 44A,). Legs yellowish-brown.

Abdomen. Yellowish-brown.

©LIB

Revision of Monolepta Chevrolat, 1836 from the Sundaland area 165

P Y

age M. bruneiensis sp. n.

— M.empatbulat sp. n.

Fig. 45. Distribution of Monolepta bruneiensis sp. nov., and

M. emphatbulat sp. nov.

e 1

@ 2-6

Male genitalia. Median lobe nearly parallel-sided and

slender. Tectum long and broad. Median spiculae broad,

flat with comb-like apex, overlapping in dorsal view

(Fig. 44Ca), lateral spiculae as simply slender spikes,

ventral spiculae slender curved hook (Fig. 44C).

Female genitalia. Spermatheca with spherical nodulus,

short middle part and long and slender cornu (Fig. 44D).

Bursa sclerites weakly sclerotized and hardly visible.

Diagnosis. Monolepta bruneiensis sp. nov. is most sim-

ilar in body colouration to M. mentaweiensis (Figs 29A,

44A). Both species possess a brownish-red pronotum and

yellowish elytra, but 1 mentaweiensis, additionally has

a brownish-red transverse basal band on the elytra (in

some old specimens, this band is a bit fainted) lacking

in M. bruneiensis sp. nov.. External differences are the

very short third antennomere in M. bruneiensis sp. nov.

(mean of ratio length second to third antennomere 1.25)

while it is longer in M. mentaweiensis (mean of same

ratio 0.80; Figs 29B, 44B), both species can be also dif-

ferentiated by size (total length M bruneiensis sp. nov.

3.75—4.70 mm; M. mentaweiensis 4.90—6.25 mm), and in

all doubtful cases, the genitalic characters allow an easy

identification.

Distribution. Only recorded from Brunei in North-

West Borneo (Fig. 45).

Monolepta empatbulat sp. nov.

urn: lsid:zoobank. org: act: 7640433 E-F BO8-4E32-956A-20EE24D45BDA

Figs 45-47

Type material. Holotype: ¢ “Brunei, Labi, Bukit Teraja

60 m, Mxt. Dipt. Forest, B.M. 1983-39, Light trap 1,75 m

above ground, 21.VIII.79, S. L. Sutton, 906, 2634”

(BMNH,; Fig. 46). Type locality: 4°18' N/114°26' E. —

Paratypes: Brunei. 2 ex., same data as holotype; 3 ex.,

same data but 24.VHI., 26. VIII. (BMNH). — Indonesia.

1 ex., Indonesia, Borneo, Kalimantan Tengah Busang/

Rekut conf., 0°03' S/113°59' E, VIII.2001, Brendell/Men-

del (BMNH). — Malaysia.; | ex., Tawai, 1300ft., 8 m S.

Bonn zoological Bulletin 71 (2): 139-176

, —

8, €8

So 4

gees 35

= 5 ESle o- ae Meee . 5 i ,

WES orn 1D:

Zev Pia ooo Et aie ©

ZERS FBR . rd

od aie nisi yp

Be, Pe oe

ARN 7 i

fa Pia eee b

Fig. 46. Type material of Monolepta empatbulat sp. nov., ho-

lotype (BMNH).

Telupoid, 5°30' N/117°24' E, [X.1977, M. E. Bacchus

(BMNH).

Etymology. The name refers on the elytral colouration

with four spots. Empatbulat means “four spots” in Malay.

Fig. 47. Monolepta empatbulat sp. nov. A. Colour pattern.

B. Basal antennomeres. C. Median lobe, a. dorsal, b. lateral,

c. ventral, without endophallic structures. D. Spermathecae.

E. Bursa-sclerites, a. dorsal, b. ventral.

Description

Total length. 3.25—3.80 mm (mean 3.55 mm; n= 10).

Head. Very finely punctuated, entirely blackish and

shining, also labrum and mandible blackish. Anten-

na long and slender, blackish and only three basal an-

tennomeres yellowish (Fig. 47A). Second antennomere

slightly shorter than third; ratio length of second to third

antennomere 0.67—1.00 (mean 0.93), ratio length of third

to fourth antennomere 0.29—0.43 (mean 0.34; Fig. 47B).

Thorax. Pronotum finely punctuated, broad, entirely

yellow to brownish-yellow. Pronotal width 0.95—1.20 mm

(mean 1.06 mm), ratio length to width 0.63—0.66 (mean

0.64). Scutellum, meso- and metathorax black. Elytra

elongated and broadened towards apex. Elytron black,

with four yellowish spots (Fig. 46b, 47A). Elytral length

2.50—2.75 mm (mean 2.67 mm), maximal width of both

©LIB

166 Izfa Riza Hazmi & Thomas Wagner

elytra together 1.80—2.00 mm (mean 1.92 mm), ratio of

maximal width of both elytra together to length of elytron

0.70—0.73 (mean 0.72). Legs yellow-brownish and distal

parts of femur and tibia brown to blackish (Fig. 46b).

Abdomen. Yellowish-brown, darker in the middle and

apical part of the abdomen.

Male genitalia. Median lobe broad and conical towards

apex. Tectum broad at base and narrowing towards apex.

Median spiculae of long, slender and flattened, ventral

spiculae shorter, club-shaped, jagged-like apically, later-

al spiculae club-shaped, basally widened (Fig. 47C).

Female genitalia. Spermatheca with small spheri-

cal nodulus, median part and cornu long and slender

(Fig. 47D). Bursa sclerites of similar size, strongly scle-

rotized and with strong spines (Fig. 47E).

Diagnosis. Monolepta empatbulat sp. nov. is most

similar to M. mohamedsaidi sp. nov. and M. zonula in

colouration and size. These species have a black head,

black elytra with yellow spots, and black meso- and

methothorax in common. M. zonula can be distinguished

by yellow antennae (Fig. 25A) from the other two spe-

cies with predominantly black antenna (Figs. 47A, 56A).

The legs of M. empatbulat sp. nov. are partly brownish to

black, while the other two species have entirely yellow

legs. In any doubtful cases the strong differences in the

genitalic morphology of both species allow a clear allo-

cation (Figs 25C—E, 47C—E, 56C-—E).

Distribution. Only known from Borneo (Brunei, Kali-

mantan and Sabah; Fig. 45).

Monolepta hitam sp. nov.

urn: |sid:zoobank.org:act:5BC28167-B9D7-4C97-9B9A-E24B7153D593

Figs 26, 48-49

Type material. Hototype: <, “Indonesia, Sulawe-

si Utara, Dumoga-Bone N. P., July 1985 / R. Ent. Soc.

London Project Wallace B. M. 195-10 / Tray 60 / Fog

13 230 m, 11. VII.85, BMNH Plot A” (BMNH,; Fig. 48).

Type locality: 0°32' N/123°58' E. — Paratypes: Indone-

sia. 9 ex., Sulawesi Tengah, Nr. Morowali, Ranu River

Area, 27.1. —20.1V.1980, S. L. Sutton & C. J. Rees; M. J.

D. Brendell, B. M. 1980-281 (BMNH); 24 ex., same data

as holotype, II. —XII.1985 (BMNH); 6 ex., Dumoga Bone

N. P. Toraut, 233 m, Sulawesi Utara, 21.V.1985, multistr.

Evergreen forest, Rothamstead strap (RMNH); 2 ex., Ed-

Fig. 48. Type material of Monolepta hitam sp. nov., holotype

(BMNH).

Bonn zoological Bulletin 71 (2): 139-176

wards subcamp, 664 m, at light, 2—5.V.1985, multistr.

Evergreen forest, Rothamstead strap (RMNH).

Etymology. The name refers to the elytral colouration;

hitam means “black” in Malay language.

Description

Total length. 4.60—6.00 mm (mean 5.16 mm; n= 10).

Head. Very finely punctuated, reddish-brown to dark-

brown, labrum and mandible of same colour. Antenna

long and slender, extending to the middle of the elytra,

yellowish, seventh, eighth and terminal antennomere are

partly dark-brown to blackish (Figs 48b, 49A). Second

and third antennomere approximately of the same length:

ratio length of second to third antennomere 0.78—1.00

(mean 0.95); ratio length of third to fourth antennomere

0.33—0.44 (mean 0.38; Fig. 49B).

Thorax. Pronotum finely punctuated, reddish-brown

to dark-brown as head. Pronotal width 1.35—1.60 mm

(mean 1.48 mm), ratio length to width 0.63—0.66 (mean

0.64). Scutellum brownish, meso- and metathorax black.

Elytron entirely blackish or in some specimens with nar-

row brownish-red apical margin (Figs 48b, 49A). Elytral

length 3.65-4.45 mm (mean 3.98 mm), maximal width

of both elytra together 2.50—-3.00 mm (mean 2.65 mm),

ratio of maximal width of both elytra together to length

of elytron 0.64—0.68 (mean 0.67). Legs entirely red-

dish-brown.

Abdomen. Reddish-brown.

Male genitalia. Median lobe parallel-sided basally and

becomes significantly narrow towards apex, tectum also

apically narrowed. Median spiculae long and slender,

Fig. 49. Monolepta hitam sp. nov. A. Colour pattern. B. Bas-

al antennomeres. C. Median lobe, a. dorsal, b. lateral, c. ven-

tral, without endophallic structures. D. Spermathecae. E. Bur-

sa-sclerites, a. dorsal, b. ventral.

©LIB

Revision of Monolepta Chevrolat, 1836 from the Sundaland area 167

few very long, lateral spiculae club-shaped, ventral spic-

ulae club-shaped with spurs at apex (Fig. 49C).

Female genitalia. Spermatheca with spherical nod-

ulus, comparatively large. Cornu long and curved

(Fig. 49D). Bursa sclerites of similar size, slender, finely

serrate (Fig. 49E).

Diagnosis. Monolepta hitam sp. nov. can be easily

distinguished from other Sundaland species of Monolep-

ta by colouration of the elytra. This is the only species

that has almost entirely black elytra, in some specimens,

brown reddish at apical margin. Somewhat similar is the

equally sized Monolepta rufipennis (total length 4.75—

6.35 mm; M. hitam sp. nov. 4.60-6.00 mm) with black

head and pronotum and reddish elytra (Figs 36A). The

black seventh and eight antennomere (Figs 48b, 49A) is

also a peculiar character of this new species.

Distribution. Only recorded from two sites on Su-

lawesi (Fig. 26).

Monolepta kuninghitam sp. nov.

urn: |sid:zoobank. org: act: AD6 96406-0BA0-4E3 1-88 D9-44C F7386C344

Figs 40, 50-51

Type material. Holotype: 3, “Sarawak, 4th Division,

Gn. Mulu NP, nr. Base Camp 50-100 m, Malaise trap”

(BMNH,; Fig. 50). Type locality: 3°55' N/114°46' E. — Pa-

ratypes: Indonesia. 2 ex., Borneo, Sarawei u. Leb. Hara,

1.1925, Sammelreise Prof. Dr. H. Winkler ded. 1924-

1925 (MNHU). — Malaysia. 1 ex., N. Borneo, Betto-

tan, nr. Sandakan, 5°51' N/118°3' E, VII.1927 (BMNH);

18 ex., Sarawak, Mt. Dulit, 4.000 ft., 3°20’ N/114°9' E,

VIII.—XI1.1932, Hobby & A. W. Moore (BMNH),; 28 ex.,

Sarawak, 4th Division, Gn. Mulu, 3°55' N/114°46' E, V—

VIII.1978, P. M. Hammond & J. E. Marshall (BMNH).

Etymology. The name of the new species refers to the

dorsal colouration, yellow and black - kuninghitam in

Malay language.

lotype (BMNH).

Description

Total length. 3.70—4.35 mm (mean 3.96 mm; n= 10).

Head. Very finely punctuated, yellow to brownish-yel-

low, labrum and mandible blackish. Antenna slender, ex-

tending almost to the middle of the elytra, blackish and

Bonn zoological Bulletin 71 (2): 139-176

<<a ®

as :

B s

Fig. 51. Monolepta kuninghitam sp. nov. A. Colour pattern.

B. Basal antennomeres. C. Median lobe, a. dorsal, b. lateral,

c. ventral, without endophallic structures. D. Spermathecae.

E. Bursa-sclerites, a. dorsal, b. ventral.

Ea Eb

only three basal antennomeres yellowish (Figs 50b, 51A).

Second antennomere shorter than third; ratio length of

second to third antennomere 0.67—0.75 (mean 0.69), ratio

length of third to fourth antennomere 0.50—0.57 (mean

0:52; Fig. 51B).

Thorax. Pronotum finely punctuated, narrow, yellow.

Pronotal width 1.00-1.25 mm (mean 1.09 mm), ratio

length to width 0.70—0.73 (mean 0.71). Scutellum, meso-

and metathorax blackish. Elytra yellowish, basal margin

black, in some specimens also apical margin blackish

(Figs 50b, 51A). Elytral length 2.65—3.30 mm (mean

2.94 mm), maximal width of both elytra together 1.90-—

2.40 mm (mean 2.10 mm), ratio of maximal width of

both elytra together to length of elytron 0.70—0.73 (mean

0.72). Femora yellowish, tibiae (with exception of the

proximal yellow third), and tarsi dark-brown (Fig. 50b).

Abdomen. Yellowish.

Male genitalia. Median lobe slender, narrow, contin-

uously conical towards apex (Fig. 51C), tectum corre-

sponding slender and pointed. Median spiculae are slen-

der and fine spines, lateral spiculae broad and v-shaped,

ventral spiculae are weakly sclerotised (Figs 51Ca, Cb).

Female genitalia. Spermatheca with very small

nodulus, broad middle part and slender, curved cornu

(Fig. 51D). Bursa sclerites large, of similar size, margins

strongly serrate (Fig. 51E).

Diagnosis. The colouration of Monolepta kuninghitam

Sp. nov. is somewhat similar to M. jacobyi in some spec-

imens with very narrow black margined elytral base, but

antennal colour, very small size and the narrow prono-

tum of M. kuninghitam sp. nov. allow a clear differenti-

ation. Genital structures patterns and size are similar to

M. bifasciata (Figs SC—E, 51C—D) and both species are

probably closely phylogenetically related. Median lobe,

©LIB

168 Izfa Riza Hazmi & Thomas Wagner

and pronotum is much slenderer in M. kuninghitam (ra-

tio length to width 0.70—0.73), broader in M. bifasciata

(0.61—0.65).

Distribution. Only known from northern Borneo (Sa-

bah and Sarawak; Fig. 40).

Monolepta marginicolloides sp. nov.

urn: lsid:zoobank.org:act:55D980FA-A34D-40B8-9F 12-4DFE28E3 147D

Figs 52-54

Type material. Holotype: 2 “Johor, Gunung Ledang,

19-23. V1.93, Yusof, Saiful, Meor’ (UKM; Fig. 52).

Type locality: 2°22' N/102°36' E. — Paratypes: Bru-

nei. | ex., Labi, Bukit Teraja, 60 m Mxt. Dipt. Forest,

4°18' N/114°26' E, 26. VIII.1979, S. L Shutton (BMNH).

— Indonesia. 3 ex., N Sumatra, Bivouac one Mt Ban-

dahara, 3°43’ N/97°41' E, 25.VI.—5S.VUI.1972, J. Krik-

ken (RMNH); 1 ex., Kalimantan, Timur Apokayan,

Long Sungei Barang, 900 m, 1°15’ S/116°49' E, 15-23.

11.1997, CuP Zorn (CJB); 1 ex., Borneo, Kalimantan

Tengah Busang/Rekut conf., 0°3' S/113°59' E, VIII.2001,

Brendell/Mendel (BMNH). — Malaysia. 1 ex., Penang,

5°15' N/100°29' E (BMNH); | ex., Sarawak, 4th Divi-

sion, Gn. Mulu NP., 3°55' N/114°46' E, P- M. Hammond

et al. (BMNH); | ex., Perak, Cameron Highlands, Rin-

glet, 4°25N/101°23E, V.2001, M. Riha (CJB).

Etymology. The name refers to the similarity in co-

louration pattern to M. marginicollis, suffix -oides

(grec.): similar as.

panne

Fig. 52. Type material of Monolepta marginicolloides sp. nov.,

holotype (BMNH).

Description

Total length. 3.70—5.10 mm (mean 4.38 mm; n = 6).

Head. Very finely punctuated, blackish and shiny, as

labrum and mandible. Antenna long and slender, extend-

ing to the middle of the elytra, blackish, only three basal

antennomeres yellowish (Figs 52b, 53A). Third antenno-

mere longer than second; ratio length of second to third

antennomere 0..67—0.75 (mean 0.69); ratio length of third

to fourth antennomere 0.38—0.43 (mean 0.41; Fig. 53B).

Thorax. Pronotum very finely punctuated, broad, en-

tirely yellowish. Pronotal width 1.20-1.55 mm (mean

1.37 mm), ratio length to width 0.58—0.63 (mean 0.60).

Scutellum, meso- and metathorax black. Elytron yellow-

Bonn zoological Bulletin 71 (2): 139-176

Fig. 53. Monolepta marginicolloides sp. nov. A. Colour pattern.

B. Basal antennomeres. C. Median lobe, a. dorsal, b. lateral,

c. ventral, without endophallic structures. D. Spermathecae.

E.Bursa-sclerites, a. dorsal, b. ventral.

ish with completely black margins (Fig. 52b), or only

most parts of the suture, base, and part of margins beyond

humerus black (Fig. 53A). Elytral length 2.90-3.95 mm

(mean 3.48 mm), maximal width of both elytra together

2.00—2.70 mm (mean 2.38 mm), ratio of maximal width

of both elytra together to length of elytron 0.68—0.70

(mean 0.69). Legs entirely yellowish.

Abdomen. Yellowish, darker in the middle of abdo-

men.

Male genitalia. Median lobe long and strongly nar-

rowed in the apical quarter (Fig. 53C), tectum short,

broad at base. Median spiculae fine short bristles, later-

al spiculae flattened, v-shaped, ventral spiculae weakly

sclerotised as group of small spines (Figs 53Ca, Cb).

Female genitalia. Spermatheca with small spheri-

cal nodulus, long and curved median part and cornu

(Fig. 53D). Dorsal pair of bursa sclerites with strong

spines, ventral part more elongated, with serrate margins

(Fig. 53E).

Diagnosis. Colouration pattern very similar to M. mar-

ginicollis, but M. marginicolloides sp. nov. 1s on aver-

age larger (total length 3.70—-5.10 mm; M. marginicollis

3,254.00 mm), and elytra are a bit slenderer (ratio of

maximal width of both elytra together to length of elytron

0.68-0.70; M. marginicollis 0.71—0.74). Other characters

to differentiate these species are the yellow-brownish

head and vertex with short central black stripe in M. mar-

ginicollis, while head in M. marginicolloides sp. nov. is

completely black. The genitalic characters are very dif-

ferent and allow a clear differentiation of both species in

any doubtful cases (Figs 34C, D, 53C-E).

©LIB

Revision of Monolepta Chevrolat, 1836 from the Sundaland area 169

M. mohamedsaidi sp.n. >

ei (|

@ 3-2 ~ A

oer ——

a Ls

* M. marginicolloides sp. n.

| PLIES 4

Fig. 54. Distribution of Monolepta marginicolloides sp. nov.,

and M. mohamedsaidi sp. nov.

Distribution. Known from Peninsular Malaysia and

Borneo (Sarawak, Kalimantan, Brunei) (Fig. 54).

Monolepta mohamedsaidi sp. nov.

urn: lsid:zoobank. org: act: 2485 1 E45-4856-4726-854B-71386D5F'4794

Figs 54—56

Type material. Holotype: 3 “Sarawak, 4th. Division

Gn. Mulu NP/ nr. Base Camp 50-100 m, V.—VHI.1978, P.

M. Hammond & J. E. Marshall, B. M.1978-49” (BMNH;:

Fig. 54). Type locality: 3°55' N/114°46' E. — Paratypes:

Brunei. 46 ex., Labi, Bukit Teraja 60 m, Mxt. Dipt.

Forest, 4°18' N/114°26' E, 21.VIII.79, B.M. 1983-39,

Light trap 1, 75 m above ground, S. L. Sutton (BMNH);:

24 ex., Temburung District Ridge, NE of Kuala Bela-

long, 300 m, 4°37' N/115°08' E, X.1992, 125W MV Light

Trap, J. H. Martin coll. (BMNH) ; 1 ex., Darussalam, Ulu

Temburong NP, 4°32' N/115°9' E, II.2015, P. Koéarek

(CJB). — Indonesia. 3 ex., N Sumatra, Bivouac one

Mt Bandahara, 3°43' N/97°41' E, 25.VI—5.VUHI.1972,

J. Krikken (RMNH); 1 ex., N-Sumatra, Hutapa-

dang, 1°25' N/99,28E, VHI.1991, Lichtfang, D. Erber

(ZFMK). — Malaysia. | ex., Penang, 5°24' N/100°29' E,

Lam. 93.60 (BMNH); 3 ex., Mt. Matang, W. Sarawak,

1°34’ N/110°16' E, [-II.1914, Bryant coll. (BMNH);

1 ex., W. Sarawak, Quop, 1°37' N/110°24' E, IV.1914,

G. E. Bryant (BMNH); 1 ex., Sarawak, foot of Mt. Du-

lit, Junction of rivers Tinjar & Lejok, 3°20’ N/114°09' E,

29.VIII.1932 (BMNH); 1 ex., Malay Penin., Selangor,

Bukit Kutu, 3500ft., 3°33' N/101°43' E, 11.1X.1929,

| eae |

; a aig

‘nr. Base |

ee corp | decumented

{ . 0 mi.

Ho i. Pbbaaa

Fig. 55. Type material of Monolepta mohamedsaidi sp. nov.,

holotype (BMNBH).

Bonn zoological Bulletin 71 (2): 139-176

H. M. Pendlebury (BMNH); 2 ex., Sabah, Tawau Plat.

1300 ft., 8 m S. Telupid, 5°35' N/117°07' E, 8.1X.1977,

M. E. Bacchus (BMNH),; 4 ex., Sarawak, Gunong Mulu

Nat. Park, R. G. S. Exped. 1977/78, 3°55' N/114°46' E,

24.V1.1978, J. D. Holloway et al. (BMNH); 24 ex.,

Sarawak, 4th. Division Gn. Mulu NP, nr. Base Camp

50-100 m, 3°55' N/114°46' E, V.—VIII.1978, P. M. Ham-

mond & J. E. Marshall (BMNH); 2 ex., Sabah, Sanda-

kan, Sepilok FR (obs tower), 50 m, 5°52' N/117°56' E,

1.X1.1987, Krikken & Rombaut (RMNH); 1 ex., Sabah,

Interior zone, road Keningau-Sepulut, km 39 Sg. Sook,

350 m, 5.02' N/116.27' E, 17.X1.1987, J. Huisman & R.

de Jong (RMNH); 1 ex., Sabah, West coast zone, 12 km

NNE of Ranau, Poring hot Springs, 6.04’ N/116.42' E,

9.X1.1987, J. Huisman & R. de Jong (RMNH); | ex.,

Sabah, Keningau Area, Tenom, 4°59' N/115°55' E,

23.X1.1987, Krikken & Rombaut (RMNH); 1 ex., Ma-

laysia, Fraser’ s Hill, 3°42’ N/101°44' E, 22.11.1991,

RM Exped. (NHRS); 1 ex., Johor, Gunung Ledang,

2°22' N/102°36' E, 19-23.V1.1993, Yusof, Saiful, Meor

(UKM); 1 ex., Pahang, Cameron Highlands, Tanah

Rata, 4°28' N/101°22' E, 1400 m, II.1997, Schuh &

Lang (ZFMK); 1 ex., Pahang, Endau Rompin NP,

2°31N/103°24! E, IV.2008, P. Cechovsky.

Etymology. Dedicated in honour to Mohammed Salleh

Mohamed Said, taxonomist from Malaysia, who did a

tremendous work on the South-East Asian Chrysomel-

idae.

Description

Total length. 3.25—4.00 mm (mean 3.60 mm; n= 10).

Head. Very finely punctuated, varied from yellowish

to blackish, labrum and mandible black. Antenna slender,

short, extending almost to the middle of elytra, black-

ish and only three basal antennomeres yellow (Figs 55b,

56A). Second antennomere shorter than third; ratio

length of second to third antennomere 0.67—1.00 (mean

0.72); ratio length of third to fourth antennomere 0.50—

0.60 (mean 0.53; Fig. 56B).

Thorax. Pronotum finely punctuated, entirely yel-

low-brownish and shiny. Pronotal width 1.00—1.25 mm

(mean 1.12 mm), ratio length to width 0.58—0.62 (mean

0.60). Scutellum, meso- and metathorax black. Elytra

shows considerable variation. Almost 80 % has black el-

ytra with longitudinal yellow spot in the middle of the

disc (Fig. 55b, 56Ab), others two yellow spot on the first

basal halfof elytra, and another two spots on the apical

part that stretch to elytral margin (Fig. 56Aa). Elytral

length 2.50—-2.90 mm (mean 2.71 mm), maximal width

of both elytra together 1.90—2.20 mm (mean 2.08 mm),

ratio of maximal width of both elytra together to length

of elytron 0.75—0.79 (mean 0.77; Fig. 56A). Legs entire-

ly yellowish.

Abdomen. Yellowish.

Male genitalia. Median lobe is slender and narrowed

towards apex. Tectum is long, lanceolate with marked

©LIB

170 Izfa Riza Hazmi & Thomas Wagner

Fig. 56. Monolepta mohamedsaidi sp. nov. A. Colour pattern.

B. Basal antennomeres. C. Median lobe, a. dorsal, b. lateral,

c. ventral, without endophallic structures. D. Spermathecae.

E.Bursa-sclerites, a. dorsal, b. ventral.

tip. Median spiculae long and slender structure, lateral

spiculae fine comb-like, ventral spiculae weakly sclero-

tised and hardly visible (Fig. 56C).

Female genitalia. Spermatheca with small spheri-

cal nodulus, middle part short, cornu long, slender and

curved (Fig. 56D). Bursa sclerites of same size, both with

very strong spines (Fig. 56E).

Diagnosis. Monolepta mohamedsaidi sp. nov. shows

considerable variation on elytra. Small body size with

yellow elytra and broad black outer and sutural margins

(Figs 55b, 56Ab) 1s very characteristic for most speci-

mens. There is some similarity with M. marginicollis

that is of same size, but has a black spot on vertex and

also black pronotal lateral and basal margins (Figs 34A,

56Ab). Specimens with yellowish spot on elytra look sim-

ilar to M. zonula and M. empatbulat sp. nov. (Fig. 56Aa).

M. zonula, is on average larger (total length 3.50—4.90,

M. mohamedsaidi sp. nov. 3.25—4.00 mm); M. empatbu-

lat sp. nov. is of same size as M. mohamedsaidi sp. nov.

but has partly black legs. The genitalic characters, partic-

ularly median lobe, allow also here a clear identification.

Highest similarity with M. zonula ,in external and genital

morphology suggest a close phylogenetic relationship.

Distribution. Known from Peninsular Malaysia and

Borneo (Sabah, Sarawak and Brunet; Fig. 54).

Monolepta ranuensis sp. nov.

urn: lsid:zoobank.org:act:

Figs 26, 57-58

Type material. Holotype: “Sulawesi Tengah, Nr. Mo-

rowali, Ranu River Area, 27.1.—20.1V.1980, S. L. Sut-

ton & C. J. Rees; M. J. D. Brendell, B. M. 1980-281”

(BMNH; Fig. 57). — Paratypes. 108 ex., same data as ho-

lotype (BMNH). Type locality: 6°14' S/106°49' E.

Bonn zoological Bulletin 71 (2): 139-176

Fig. 57. Type material of Monolepta ranuensis sp. nov., holo-

type (BMNH).

Etymology. The name refers to the type locality, Ranu

River in Sulawesi.

Description

Total length. 3.25—3.70 mm (mean 3.44 mm; n= 10).

Head. Finely punctuated, yellowish, with deep suture

between antennal sockets, labrum and mandible yellow-

ish-brown. Antenna very slender, extending to the middle

of the elytra, yellow, last antennomere darker (Figs 57b,

58A). Second antennomere on average shorter than third;

ratio length of second to third antennomere 0.67—1.00

(mean 0.85); ratio length of third to fourth antennomere

0.43—0.50 (mean 0.42; Fig. 58B).

Thorax. Pronotum yellow, broad, shining and with

shallow depression in the middle, comparatively coarse-

ly punctuated. Pronotal width 0.95-1.00 mm (mean

0.97 mm), ratio length to width 0.63—0.65 (mean 0.64).

Scutellum, meso- and metathorax yellow, as elytron that

is coarsely punctuated and shining. Elytral length 2.35-

2.60 mm (mean 2.43 mm), maximal width of both elytra

Fig. 58. Monolepta ranuensis sp. nov. A. Colour pattern.

B. Basal antennomeres. C. Median lobe, a. dorsal, b. lateral,

c. ventral, without endophallic structures. D. Spermathecae.

E. Bursa-sclerites, a. dorsal, b. ventral.

©LIB

Revision of Monolepta Chevrolat, 1836 from the Sundaland area 171

together 1.70—1.90 mm (mean 1.74 mm), ratio of max-

imal width of both elytra together to length of elytron

0.69-0.73 (mean 0.71; Fig. 58A). Legs yellow.

Abdomen. Yellowish.

Male genitalia. Median lobe lanceolate, becoming

slightly towards apex. Median spiculae as few stron-

ger spines, often slightly curved in dorsal view, on base

with two horn-shaped structures (Fig. 58Cb), lateral

spiculae slender with two small spines apically, ventral

spiculae weakly sclerotized and hardly visible or absent

(Fig. 58C).

Female genitalia. Spermatheca with small spheri-

cal nodulus and long and slender middle part and cor-

nu (Fig. 58D). Bursa sclerites broad, nearly of same size

with spiny margins (Fig. 58E).

Diagnosis. Monolepta ranuensis sp. nov. is with a to-

tal length of 3.25-3.70 mm the smallest “true” species

of Monolepta from Sundaland area. Somewhat similar

is the also unicolorous, but reddish-brown M. sulawen-

sis sp. nov. that is of similar size and occurs syntopic at

the same place on Sulawesi. In doubtful cases, the gen-

ital characters of both sexes allow a clear identification

(Figs 58C—E, 60C-—E). Also, the shallow transverse pro-

notal depression, and the elytral punctuation is charac-

teristic for M. sulawensis sp. nov. There a several further

small completely yellow species described in Monolepta

(and probably even not yet described) in the region, and

in many cases the genitalic dissection only allows an al-

location to genus and species.

Distribution. Only recorded from the type locality in

Central-Eastern Sulawesi (Fig. 26).

Monolepta sulawensis sp. nov.

urn: lsid:zoobank. org: act: A42DE2FD-5 B03-4665-930 1-426212644D89

Figs 40, 59-60

Type material. Holotype: 6 ’Sulawesi Tengah, Nr. Mo-

rowali, Ranu River Area, 27.1—20.1V.1980, S. L. Sut-

ton & C. J. Rees; M. J. D. Brendell, B.M. 1980-281”

(BMNH,; Fig. 138). — Paratypes. 13 ex., same data as ho-

lotype, mainly captured by light (BMNH).

Etymology. Patronym.

Fig. 59. Type material of Monolepta sulawensis sp. nov., holo-

type (BMNH).

Bonn zoological Bulletin 71 (2): 139-176

Fig. 60. Monolepta sulawensis sp. nov. A. Colour pattern.

B. Basal antennomeres. C. Median lobe, a. dorsal, b. lateral,

c. ventral, without endophallic structures. D. Spermathecae.

E. Bursa-sclerites, a. dorsal, b. ventral.

Description

Total length. 3.70—4.25 mm (mean 3.90 mm; n= 10).

Head. Very finely punctuated, entirely brownish, as

labrum and mandible. Antenna slender, extending to the

middle of elytra, brownish, and the last antennomere

often a bit darker (Figs 59b, 60A). Third antennomere

shorter than second; ratio length of second to third an-

tennomere 1.00—1.50 (mean 1.25); ratio length of third

to fourth antennomere 0.37—0.42 (mean 0.38; Fig. 60B).

Thorax. Pronotum very finely punctured, entirely yel-

lowish to reddish-brown, shining and without depres-

sion. Pronotal width 1.20—1.25 mm (mean 1.22 mm),

ratio length to width 0.60—0.64 (mean 0.62). Scutellum,

meso- and metathorax brownish. Elytra entirely yellow-

ish-brown to brownish. Elytral length 2.80-3.40 mm

(mean 3.04 mm), maximal width of both elytra together

2.10—2.40 mm (mean 2.20 mm), ratio of maximal width

of both elytra together to length of elytron 0.70-0.74

(mean 0.72; Fig. 60A). Legs yellowish-brownish.

Abdomen. Yellowish-brownish.

Male genitalia. Median lobe broad, short, parallel-sid-

ed. Group of median spiculae at base and one pair of flat-

tened structures with brushy apex in the middle, lateral

spiculae as slightly curved plates, ventral spiculae club-

shaped with brushy apex (Fig. 60C).

Female genitalia. Spermatheca with large spherical

nodulus, broad middle part, slender cornu (Fig. 60D).

Two pairs of differently structures bursa sclerites, both

with fine saw-like margins (Fig. 60E).

Diagnosis. Monolepta sulawensis look somewhat

similar to the also uniformly colored, but pale yellowish

M. ranuensis sp. nov., but is larger (total length 3.70—

4.25 mm, M. ranuensis 3.25—3.70 mm). Short third anten-

©LIB

172 Izfa Riza Hazmi & Thomas Wagner

nomere is also characteristic for MZ. sulawensis sp. nov.,

as is the genitalic morphology (Figs 58C—E, 60C-—E).

Distribution. Only recorded from the type locality in

brown-reddish; median lobe broad, with three types

of spiculae (Fig. 20C); widely distributed from India

towards Java (Fig. 21)... M. orientalis Jacoby, 1889

Central-Eastern Sulawesi (Fig. 40).

Narrow yellowish transverse-band in the middle,

broad black at base and narrow black behind the

Identification key yellow band, brown-reddish in the apical third

(Fig. 22b, 23A); scutellum black; median lobe

1 Elytron bi- or tricolorous, with transverse bands comparatively small, ventral spiculae hardly visible

(Figs 5A, 20A, 23A, 29A, 32A), circular yellow

spots (Figs 25A, 47A, 56A), or spot with undulate

black and yellow colouration (Fig. 14A) ................ Zz

(Fig. 23C); widely distributed from India towards

Javathig. DVS. Bald M. flavofasciata Jacoby, 1889

Fivtron!predbiminantly-uiiedloroustyellow., reddish 7. Head and pronotum of same colouration, reddish-

gies Sa i da gaa ag Figs 10b-13b, 14A i

or black (Figs 8A, 17A, 39A, 44A, 58A, 60A), See eS ee eee yale Genes oan

. spot at elytral base and a second in the apical third,

yellow with black suture and/or basal and lateral Ani

, but usually weakly defined, resulting in somewhat

elytral margins (Figs 34A, 42A, 51A, 53A), with dulati fovea Ivtra (Fis 10b-12b

reddish base (Fig. 3a), or brownish with red elytral We Scale al ee a (Figs ‘

HDRCEIS AOA) sel ceccecsy fe alas Rosle Bec bec abe 10 ee etal anal ag Clearly BCE NEL Bey Lap ols

2 Elytron with transverse bands (Figs 5A, 20A, 23A, (Fig. 13); total length 3.50-5.25 mm, widely

DS Ps SON eels hese east ey Iwas cute ad rae) Uva 3 distributed and abundant from Northwest-India and

— Elytron with circular or longitudinal yellow spot Central China throughout the entire Oriental Region

(Figs 25A, 47A, 56A), or spot with undulate black to Luzon, Palawan, Borneo and Java, not known

and yellow colouration (Fig. 14A)....0..00.0ccce. 7 from Wallacea (Fig. 15) ..M. signata (Olivier, 1808)

3 Elytron only with one brown or black transverse — Pronotum yellow, head usually black (rarely yellow),

band at base, extension varies from very narrow up elytron yellow and black only, yellow spots not close

to one third of elytral length (Figs 29A, 32A)........ 4 to humerus and well defined (Figs 25A, 47A, 56A).

= Sel ytronwatheat least-nyo-clearly denned transverse -2ygisk. doe veh escvesat gah ohn Rae gd ces yehore ee a 8

bands (Figs 1b, 4a, 5A, 20A, 23A), rarely subterminal 8 Elytron black with one circular yellow spot in the

batidireduced'tospots (Piss 2D) o.28 et sccen seine: eo. 5 middle (Fig 25A), apical third of elytron or at least

4 Elytron with dark brown to black base, remaining broad apical margin yellowish, suture sometimes

parts of elytron pale reddisch-brown, head and reddish (Fig. 24b), antenna with exception of the

pronotum contrasting yellow (Figs 31b, 32A); terminal antennomere yellow; on average larger,

smaller, total length 3.70—5.40 mm, wide distribution total length 3.50-4.90 mm; Sundaland distribution

from India to Southeast Asia up to Borneo (Fig. 33) without Wallacea (Fig. 26)...M. zonula Weise, 1916

Te ee ay ga M. militaris Jacoby, 1896 a Elytron black with two circular yellow spots

— Elytron with paler brownish base, other parts more (Figs 47A) where the hind one can extend to the

yellow, head and pronotum brownish-red, net or less apical margin (Fig. 56Aa), or longitudinal band

contrasting to the elytral base (Figs 27b, 29A); larger, (Fig. 56Ab), antenna with exception of the yellow

a eaeu cere rae ce AR a three basal antennomeres black; on average smaller,

atta PE eaan Soe rae aS gue ey Singapore total fensth: 3: 25-4 OOM oi hoc ee ee ik other aca 9

CRAG ASO) A Mns as. cc: M. mentaweiensis (Jacoby, 1896) ;

; 9 Pronotum broad, pronotal width to length 0.58—0.62,

5 Subapical transverse band ends far from elytral ; .; ;

eo, elytral colouration quite variable (Fig. 56A); median

margin (Figs 1b, 4a, 5A), sometimes reduced to small : ;

; lobe slender (Fig. 56C), bursas sclerites with very

spots (Fig. 2b); smaller, total length 3.70—5.10 mm; ; Fis. RELY Bartneulne Malan rr

pronotum less broad, pronotal width to length ath spines (Fig. Pies ae d aoa oe

0.61-0.65; spermatheca without spherical nodulus ORC ON fais Be sed neste . mohamedsaidi sp. nov.

(Fig. 5D); widely distributed from India through SE-__ ~ _ Pronotum less broad, pronotal width to length 0.63—

Asia towards New Guinea (Fig. 6).........ccccccccceeee 0.66; elytron with completely black margins and two

AL guy el Wry M. bifasciata (Hornstedt, 1788) circular spots (Fig. 47A), median lobe very broad

— Subapical transverse band extending to elytral (Fig. 47C), bursa sclerites less spiny (Fig. 47E),

margin (Figs 20A, 23A); larger 4.10-5.80 mm; restricted to Borneo.............. M. empatbulat sp. nov.

pronotum broader, pronotal length to width 0.55— 10 Elytron yellow with at least parts of the suture black,

0.59 (mean 0.58); spermatheca with spherical usually often also parts of outer margins black

nodulus (Figs 20D, 23D).....0....0.cccccccccecce cence 6 (Pigs 34A0 AQAA SGA) oko. t ihc nda 11

6 Broad yellowish transverse band bounded above and -—_ Elytron entirely yellow (Figs 39A, 44A, 58A),

below by a narrow black band, and another narrow

black band on basal margin (19b, 20A), scutellum

Bonn zoological Bulletin 71 (2): 139-176

yellow with black base (Figs 17A, 51A), or reddish

(3) ]6) (TC) Cah ada se Urea shakin Dahesh Ra her iid aera Raa 13

Revision of Monolepta Chevrolat, 1836 from the Sundaland area

Only basal third of suture black, antenna yellow

(Fig. 42A); total length 4.60-4.90 mm; endemic in

Johan: See ee RED M. putri Mohamedsaid, 2001

Suture mainly black, antenna brown to black

(Figs 34A, 53A); Malay Peninsula, Borneo or

SUMMA rr Reena rer ch tad heaasiat erat ante as 12

Head yellow to brownish-yellow, vertex with

longitudinal median black spot, outer and basal

pronotal margins black (Fig. 34A); smaller, total

length 3.25—4.00 mm; median lobe conical (Fig. 34C)

spermatheca with large nodulus (Fig. 34D); Malay

Peninsula, Borneo and Sumatra (Fig. 9) .............00....

Fase Oe ee eee M. marginicollis Jacoby, 1896

Head black, pronotum completely yellow (Fig. 53A);

larger, total length 3.70—5.10 mm; median with very

Slender apical third (Fig. 53C), spermatheca with

small nodulus (Fig. 53D); Malay Peninsula and

Borneo (Fig. 54).......... M. marginicolloides sp. nov.

Head black or yellow, pronotum completely yellow,

very broad sutural and outer margins (Fig. 56Ab);

smaller, total length 3.25—4.00 mm; median lobe like

ig. SOCs COLOUrAONs Ly Pe OL. mer... wee, oo emee fe we

keg cerns vse fee M. mohamedsaidi sp. nov. (see 9)

Elytron entirely yellow (Figs 39, 58) or yellow with

black (Figs 17, 51) or reddish (Fig. 3a) base......... 14

Elytron unicolorous reddish, brownish or black

(Figs 8A, 37A, 60A), or brownish to black with red

elytral tip: (Pie AG As cn). Sei 8, A. Boreas. Songs 19

Elytron entirely yellow (Figs 39A, 44A, 58A); total

Lene th. 3°25 GSM e 5, Be ents iij aig etek oe 15

Base of yellow elytra black (Fig. 51A), sometimes

black colour very narrow (Fig. 17A), or reddish

CEES Dili teak es Ec eg Oe ee 17

Head and pronotum contrasting brownish-red, to

the yellow elytra; basal antennomeres yellowish

to brown, others brown to black (Figs 43a, 44A);

second antennomere significantly longer than third,

ratio length of second to third antennomere 1.00-

1.50 (Fig. 44B); total length 3.75—-4.70 mm; endemic

LOSS UMCIME 7, 355, cade a are M. bruneiensis sp. nov.

Pronotum yellow, same color as elytra; antenna

entirely yellow (with exception of terminal

antennomere, Figs 39A, 58A); second antennomeres

not shorter than third (Figs 39B, 58B).......ssc0ssee0 16

Head black, strong contrast to yellow pronotum

(Fig. 39A); larger, total length 4.75-6.15 mm;

median lobe very short and broad (Fig. 39C); Malay

Peninsula, Borneo, Sumatra and Java (Fig. 40)........

hr a Re M. tiomanensis Mohamedsaid, 1999

Head and pronotum of same colour (Fig. 58A); much

smaller, total length 3.25—-3.70 mm; median lobe

slender, conical (Fig. 58C); endemic to Sulawesi

(EHO ON etc rgemnantles Beeesioe M. ranuensis sp. nov.

Very large, total length 6.40—-7.70 mm; pronotum

broad, pronotal width to length 0.59-0.61 (Fig. 17A);

black basal elytral margin very fine; antenna yellow;

Bonn zoological Bulletin 71 (2): 139-176

173

widely distributed in Sundaland (Fig. 18).................

pS er US ores Delle ots Mee M. jacobyi Weise, 1908

Much smaller, total length 3.70-4.35 mm. ............ 18

Pronotum less broad, pronotal width to length 0.70-

0.72 (Fig. 51A); broad black basal elytral margin

and also elytral tip black; outer antennomeres black;

endemic to Borneo ............ M. kuninghitam sp. nov.

Pronotum broader, pronotal width to length 0.61-

0.65; elytra reddish to brownish at base (Fig. 3b); ;

rare colour pattern of................ M. bifasciata (see 5)

Elytron dark brown to black with reddish apex

(Fig. 49A)); total length 4.60-6.00 mm; endemic to

SULAWESI eRe Nee linn oe ee M. hitam sp. nov.

Elytron unicolorous reddish, brownish or black ... 20

Small, total length 3.70-4.40 mm; median lobe broad

(Fig. 60C), spermatheca with spherical nodulus

(Fig. 60E); endemic to Sulawesi.........0000....00ccee

er es Seen Keer ee M. sulawensis sp. nov.

Larger, total length 4.50-6.35 mm, median lobe

slender (Figs 8C, 37C), spermatheca without

spherical nodulus (Figs 8E, 37E); unknown from

SUGGS, 2oPe een nnmraes” lame ener weeny over tony PUer. ) 22 lappa neahtS VaK 21

Head, pronotum and underside black, elytron

reddish-brown (Figs 36b, 37A); median lobe broad

(Pigea7€ je. MalayvePeminsulavOnbyey.. cee okies ian

Pi Ree RR her teas ha Fa M. rufipennis Jacoby, 1899

Upper- and underside uniformly brownish-red, outer

elytral margins gradually darker (Figs 7b, 8A);

median lobe slender (Fig. 8C); Java, Borneo and

ddjacentislandssCras.. Ge A a0) kM Co laa ve,

Checklist of Monolepta Chevrolat, 1836 from Sunda-

land

The following list comprises all known species of Mono-

lepta after revision including synonymies:

Monolepta bifasciata (Hornstedt, 1788)

Chrysomela bifasciata Hornstedt, 1788

= Cryptocephalus multicolor Gmelin, 1790

= Crioceris quadrinotata Fabricius, 1801

= Luperodes latefascia Motschulsky, 1858

= Monolepta parvonotata Jacoby, 1886; syn. nov.

= Monolepta mustaphai Mohamedsaid, 1997;

syn. nov.

= Monolepta entimauensis Mohamedsaid, 1998;

syn. nov.

Monolepta bruneiensis sp. nov.

Monolepta empatbulat sp. nov.

Monolepta flavofasciata Jacoby, 1889

Monolepta hitam sp. nov.

Monolepta jacobyi Weise, 1908

Monolepta kuninghitam sp. nov.

Monolepta marginicollis Jacoby, 1896

Monolepta marginicolloides sp. nov.

©LIB

174 Izfa Riza Hazmi & Thomas Wagner

Monolepta mentaweiensis (Jacoby, 1896)

Candezea mentaweiensis Jacoby, 1896

= Monolepta basalis Jacoby, 1884

= M. hageni Weise, 1916; syn. nov.

Monolepta militaris Jacoby, 1896

Monolepta mohamedsaidi sp. nov.

Monolepta orientalis Jacoby, 1889

= Monolepta konbiriensis Duvivier, 1891

Monolepta putri Mohamedsaid, 2001

Monolepta ranuensis sp. nov.

Monolepta rubra (Gyllenhal, 1808)

= Luperodes javanensis Jacoby, 1887

Monolepta rufipennis Jacoby, 1899

Monolepta signata (Olivier, 1808)

Galeruca signata Olivier, 1808

= Crioceris neglecta Sahlberg, 1829: 29

= Luperodes quadripustulatus Motschulsky, 1858

= Luperodes hieroglyphicus Motschulsky, 1858

= Monolepta elegantula Boheman, 1859: 183

= Luperodes dorsalis Motschulsky, 1866: 415

= Luperodes quadriguttata Fairmaire, 1887: 333

= Monolepta picturata Jacoby, 1896; syn. nov.

= Monolepta simplex Weise, 1913

Monolepta sulawensis sp. nov.

Monolepta tiomanensis Mohamedsaid, 1999

Monolepta zonula Weise, 1916

Species of Monolepta transferred to other genera

The following list comprises species originally described

in Monolepta known from the core of Sundaland with-

out the Philippines (except of Palawan), Thailand and

Indo-China, or, as a positive list, comprises species

known from the states of Malaysia, Singapore, Brunei

and Indonesia without parts East of the Lydekker line,

i.e., including the fauna of Sulawesi, Timor and Seram

up to the East (Lohmann et al. 2011). Some of these spe-

cies have been synonymized or transferred by us to other

genera, others need to be transferred to other genera in

subsequent studies, since they are not con-generic with

Monolepta s. str.

Our decision to not include the following taxa in Mono-

lepta is based on a check of most of the type material.

Similar to the taxonomic revision on base of monophy-

letic groups of the Afrotropical galerucine fauna, with

nearly 50 papers published up to now, also for the orien-

tal taxa tremendous revisionary work needs to be done.

We started this with revisions on Ochralea (Hazmi &

Wagner 2010a), Arcastes (Hazmi & Wagner 2010b,

2010c), Neolepta (Hazmi & Wagner 2013) and the rede-

scription of the generotype of Paleosepharia (Rizki et al.

2016). On base of those results, several of the following

species belong to Paleosepharia, but up to now only one

transferral has been published (Rizki et al. 2014). Species

of Nadrana Baly, 1865 from Sundland have been also

revised recently (Zulfadli et al. 2015), and there might be

Bonn zoological Bulletin 71 (2): 139-176

also few species from the following list that need to be

transferred to this group.

Monolepta aemula Weise, 1922

Synonym of Paraneolepta imitans (Jacoby, 1894)

(Hazmi & Wagner 2013)

Monolepta affinis Jacoby, 1886

Monolepta albofasciata Jacoby, 1884

Monolepta approximans Jacoby, 1896

Monolepta azlani Mohamedsaid, 1998

Transferred to Paleosepharia (Rizki et al. 2014)

Monolepta bimaculata (Hornstedt, 1788)

Monolepta borneensis Mohamedsaid, 1994

Monolepta c-album (Jacoby, 1899)

Monolepta cantik Mohamedsaid, 2000

Monolepta castanea Allard, 1888

Monolepta castanoptera Weise, 1924

Monolepta danumica Mohamedsaid, 1993

Monolepta erythromelas Weise, 1922

Synonym of Ochralea nigripes (Olivier, 1808)

(Hazmi & Wagner 2010a)

Monolepta foveicollis Baly, 1888

Monolepta haemorrhoidalis (Fabricius, 1801)

Monolepta impressicollis (Jacoby, 1896)

Monolepta inornata (Jacoby, 1894)

Monolepta irpa Mohamedsaid, 2000

Monolepta kedenburgi Weise, 1922

Transferred to Nadrana (Zulfadli et al. 2015)

Monolepta kenit Mohamedsaid, 2000

Monolepta kerangas Mohamedsaid, 1998

Monolepta kraepelini Weise, 1922

Monolepta latefasciata Jacoby, 1896

Monolepta laticornis (Jacoby, 1899)

Monolepta longitarsis Jacoby, 1896

Monolepta malaysiana Mohamedsaid, 1993

Monolepta melancholica Jacoby, 1886

Monolepta merah Mohamedsaid, 1993

Monolepta modigliani Jacoby, 1896

Monolepta moluquensis Allard, 1888

Monolepta murphyi Mohamedsaid, 2002

Monolepta nigriceps Weise, 1915

Monolepta nigrilabrum (Jacoby, 1899)

Monolepta nigripes (Olivier, 1808)

Revalidated as Ochralea nigripes (Olivier, 1808)

(Hazmi & Wagner 2010a)

Monolepta nigromarginata Jacoby, 1896

Monolepta obtusa Jacoby, 1896

Monolepta pagi Mohamedsaid, 2001

Monolepta parva Mohamedsaid, 2001

Monolepta pectoralis Boheman, 1859

Monolepta piceola Weise, 1915

Monolepta rubricollis Jacoby, 1905

Monolepta rugosa Mohamedsaid, 1998

Monolepta sangirensis Jacoby, 1894

Monolepta semifovea Mohamedsaid, 1993

Monolepta sexmaculata Jacoby, 1886

©LIB

Revision of Monolepta Chevrolat, 1836 from the Sundaland area 175

Monolepta subsulcata Boisduval, 1835

Monolepta thoracica (Jacoby, 1896)

Monolepta terminata (Guérin-Méneville, 1830)

Monolepta tibowensis Mohamedsaid, 2000

Monolepta timorensis Jacoby, 1894

Monolepta unicolor Jacoby, 1886

Monolepta wallacei Baly, 1888

Monolepta wangkliana Mohamedsaid, 2005

Transferred to Ochralea (Hazmi & Wagner 2010a)

Monolepta wilsoni Kimoto, 1989

Acknowledgements. We thank all colleagues who made mate-

rial available to us. Michael Kuhlmann photographed the type

of Crioceris quadrinotata, Ron Been and Jan Bezdék made

many valuable comments on the manuscript. Jan additionally

offered us generously his photos of type material from NHRS

and MCGD.

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= of Biodiversity Change

AL KGENIG

Bonn zoological Bulletin 71 (2): 177-183

2022 Saigusa T & Sinclair B.J.

https://do1.org/10.20363/BZB-2022.71.2.177

ISSN 2190-7307

http://www.zoologicalbulletin.de

Research article

urn:|sid:zoobank.org:pub:68F9C 120-14D4-4562-B3FB-289 17927E795

A new genus of Clinocerinae (Diptera: Empididae)

from Taiwan and Yunnan (China)

Toyohei Saigusa Oig Bradley J. Sinclair

©>.

'7-1-402 Baikoen 2-chome, Chuo-ku, Fukuoka-shi 810-0035, Japan

?Canadian National Collection of Insects & Canadian Food Inspection Agency, OPL-Entomology, K.W. Neatby Bldg., C.E.F.,

960 Carling Ave., Ottawa, ON, Canada KIA 0C6

“Corresponding author: Email: bradley.sinclair@inspection.gc.ca

'urn:lsid:zoobank.org:author:390F7B16-1B9D-419F-BOCD-F3279B89DF 1D

? urn:lsid:zoobank.org:author:45 16327F-B73E-456C-927F-18EFBOB9E08B

Abstract. A new genus of Clinocerinae (Diptera: Empididae), Asioclinocera gen. nov. and two new species, A. formosana

gen. et sp. nov. and A. yunnanica gen. et sp. nov., are described from Taiwan and Yunnan (China), respectively. An updated

key to Palaearctic genera of Clinocerinae 1s provided and the phylogenetic affinities of the new genus are discussed.

Key words. Aquatic dance flies, new species, description, generic key.

INTRODUCTION

The Clinocerinae Melander, 1928 is a subfamily of aquat-

ic dance flies (Diptera: Empididae), with adults frequent-

ly encountered in seeps, streams and rivers worldwide

(Sinclair 1995). The genera are distinguished by: narrow

wings; non-raptorial fore legs; broad, sucker-like labella;

stout, erect costal setae; and upcurved male terminalia

(Sinclair 1995). In the Northern Hemisphere there are 14

recorded genera, with Hypenella Collin, 1941 and Rhy-

acodomia Saigusa, 1986 restricted to eastern Asia (Saigu-

sa 1986; Sinclair 1995). In recent years, odd new species

of clinocerines were identified from Yunnan (China) and

Taiwan. These species are distinct from other clinocer-

ines, characterized by long slender arista-like stylus (usu-

ally strong, shorter than head), convex labrum (usually

flattened), elongate radial fork (branching of vein R,,.

usually short), unmodified female terminalia (some gen-

era with acanthophorites) and distinctive male terminalia

(phallus usually elongate and separated).

In this study, we describe these specimens as a new ge-

nus, recognizing two new species. In addition, the phy-

logenetic affinities are discussed and an updated identi-

fication key to the Palaearctic genera of Clinocerinae is

provided.

MATERIAL AND METHODS

This study is based on material deposited in the following

institutions: Biosystematics Laboratory, Kyushu Univer-

Received: 11.07.2022

Accepted: 17.10.2022

sity, Fukuoka, Japan (KUMF); and Canadian National

Collection of Insects, Ottawa, Canada (CNC). Label data

for primary types are cited from the top downward, with

the data from each label in quotation marks. Labels are

cited in full, with original spelling, punctuation, and date,

and label lines are delimited by a slash (/). The repository

of each type is given in parentheses. Secondary type data

are abridged and listed alphabetically.

Methods used in specimen preparation follow that of

Sinclair (1994, 1995) and Saigusa & Sinclair (2016).

Photographs were taken with a Leica camera model

DFC5400 using Leica Application Suite X. The distri-

bution map was created with SimpleMappr (Shorthouse

2010). Terms used for adult structures follow those of

Cumming & Wood (2017).

RESULTS

Taxonomy

Asioclinocera gen. nov.

urn: lsid:zoobank. org: act: DDB824 13-42C5-4792-A5EF-26F9AD8AF9BF

Figs 1-10

Type-species

A. yunnanica gen. et sp. nov.

Diagnosis. This new genus is distinguished from all other

genera of Clinocerinae by the following combination of

characters: elongate, filamentous arista-like stylus, face

Corresponding editor: X. Mengual

Published: 24.10.2022

178 Toyohei Saigusa & Bradley J. Sinclair

with deep cleft and ridge, labrum convex, radial fork

elongate with vein R,,. branching near apex of cell dm

and cell r, about one-third length of wing, apical fifth of

fore femur with spur-like anteroventral seta, male tergum

8 greatly narrowed medially and broadly plate-like lat-

erally, and phallus broadly attached to hypandrium with

large ejaculatory apodeme.

Etymology. The generic name is derived from Asia,

referring to its distribution, and Clinocera Meigen, 1803,

a related genus. The gender is feminine.

Description. Eye with dense ommatrichia. Arista-like

stylus very slender and long, about as long as head height

(Fig. 2). Face with median ridge on lower half. Gena di-

vided by frontoclypeal suture extending from palpus to

lower margin of eye. Mouthparts with labrum convex

(Fig. 2); labellum sucker-like, without pseudotracheae.

Fore femur with preapical anteroventral spine-like seta,

slightly shorter than width of femur (Fig. 2); without sub-

apical anterior comb, only cluster of short stiff setae; an-

teroventral and posteroventral rows of slender short setae

(Fig. 5). Fore tibia with anteroventral row of short stout

setae (Fig. 5). Tarsomere 5 without apical prolongation.

Hind tibia with distinct anteroventral seta on apical third

(Fig. 1). Scutum with long dorsocentral setae (Fig. 5);

without acrostichal setae; notopleuron with weak ante-

rior seta and strong posterior seta; scutellar setae very

closely approximated at apex (width of anterior ocellus);

laterotergite with setae; dorsal mesepimeral pocket pres-

ent. Wing with basal costal seta; without pterostigma

(Fig. 3); Sc complete; costa circumambient with strong,

erect setae beyond R,; R, ends before middle of wing;

elongate cell dm; radial fork very long, arising near apex

of cell dm; R, slightly arched to straight and divergent

from R.; cell r, about one-third length of wing; cell cua

narrow, with truncate apical margin. Female abdomen

telescopic, apparently unmodified, with cerci of A. yun-

nanica (unknown for formosana) short, shiny, sharply

pointed (not dissected). Male terminalia (Figs 6-7) with

setae of cercal plate not confined to small region (Fig. 8);

phallus without distiphallus; base of phallus and hypan-

drium expanded; male tergum 8 narrow with inflated lat-

eral margin bearing many setae (Fig. 9).

Distribution. This new genus 1s currently known from

Taiwan and the Chinese province of Yunnan (Fig. 10).

Remarks. This new genus of Clinocerinae appears to

be most similar to Rhyacodromia on the basis of the con-

vex-shaped labrum and spine-like setae on the fore femur.

The convex labrum is a rather unique character within

Clinocerinae and previously considered a synapomorphy

of the clade Rhyacodromia + Trichoclinocera Collin,

1941 (Sinclair 1995, character 25). However, the male

terminalia distinctly differ between Rhyacodromia and

Asioclinocera gen. nov., specifically the connection of

phallobase and hypandrium is unique in the former genus

(anterolateral corner of hypandrium has unique one-point

articulation with the anteroventral corner of epandrium

Bonn zoological Bulletin 71 (2): 177-183

and posterolateral corner of sternum 7 (Saigusa 1986:

fig. 9; Sinclair 1995: fig. 15)), whereas the connection

in Asioclinocera gen. nov. is similar to Clinocera and

Wiedemannia Zetterstedt, 1838; the surstylus is widely

connected with the distal margin of the epandrium in

Rhyacodromia compared to the narrow connection with

the epandrium in Asioclinocera gen. nov.; and tergum 7

bears a denticulate posterior swelling in Rhyacodromia

and is unmodified in Asioclinocera gen. nov. The new

genus is further distinguished in the following identifi-

cation key.

Key to Palaearctic genera of Clinocerinae

1. Sc evanescent apically (Sinclair 1995: fig. 64) ........

Bh ee er eh A ok Proclinopyga Melander

— Sc complete, reaching costal vein ........00....000006. 2

2. R, with macrotrichia arising from dorsal surface

(Saigusa & Sinclair 2021: fig. 40) oo...

eo: ete eee Ee al Trichoclinocera Collin

aa) ARS WALNOUUIMACTOUMEIN AT 5 ota tnnmit coin Peadevignbi B)

3. Proboscis as long as height of head; labrum long and

blade-like, extending length of proboscis; labellum

narrow, not sucker-like (Sinclair 1995: fig. 58) .......

Pes sree a AL ent Ses Pe oe Roederiodes Coquillett

— Proboscis shorter than height of head; labrum

short, subtriangular, encircled by large sucker-like

IF er AU NUIN heres mae a eee Wl OL. Me Mr es AL, po ete 4

4. Gena divided by frontoclypeal suture extending

from palpus to lower margin of eye (Sinclair 1995:

jOF SNe 618) Nite A IAMN Ry Bey ALE A Ee Rg, oR ae ei 5)

— Gena divided by frontoclypeal suture extending from

palpus to facial ridge (Sinclair 1995: fig. 59) ....... 1]

5. Neck arising high on occiput; head usually narrow,

extending obliquely forward (Sinclair 1995: fig. 60);

wing usually with circular white spots (Sinclair

OGD s WSOP) 0dr Dolichocephala Macquart

— Neck arising near centre of occiput; head

hypognathous; wing without circular white spots .. 6

6. R,,, forked distal to apex of R,,, (Smith 1965:

IGF ny 8 Pam eee kn re Rhyacodromia Saigusa

— R,,, forked proximal to apex of R,,, ...........00c0.

7. Arista-like stylus very slender, about as long as

head height (Fig. 2); apical scutellar setae closely

approximated; radial fork arising near apex of cell

dm, with cell r, nearly one-third length of wing

Jad oon eter wis tind ee Asioclinocera gen. nov.

— Arista-like stylus usually thickened, shorter than

height of head (Sinclair 1995: fig. 57); apical

scutellar setae widely separated; radial fork arising

well distant of cell dm, with cell r, nearly one-fifth

length of wing (Sinclair 1995: fig. 67)... 8

8. Lower margin of face entire, without median notch

and ridge; phallus with non-articulated apical

filament or distiphallus (Sinclair 1995: fig. 21);

female abdomen attenuated ......... Clinocera Meigen

©LIB

A new genus of Clinocerinae (Diptera: Empididae) from Taiwan and Yunnan (China) 179

— Lower margin of face with notch; if notch lacking, 10. Pulvilli vestigial and empodia reduced, length less

phallus with biarticulated distiphallus (Sinclair than one-third length of tarsal claws ......0....00...c.06.

1[99s* e849 tandsicmalesabdomensimuncatce(G Oa, Adar Parneerae hens Seeman gtd ald Bergenstammia Mik

Bergenstammia nudipes (Loew, 1858)) .............04. 9 — Pulvilli and empodia well developed and pad-like, at

9. Face with setulae along inner margin of eye ............ least as long as tarsal claws ............ Phaeobalia Mik

rane tuners! 2 remy orsewssseadl weer os wa re tk Kowarzia Mik — 11. Clypeus prominent and clearly separated from face

— Face lacking setulae along inner margin of eye ... 10 CSinclaital 995211075 9)). 12a cece | Hypenella Collin

— Clypeus not prominent or separated from face .... 12

Figs 1-3. Asioclinocera formosana gen. et sp. nov., lateral view. 1. Habitus. 2. Head, antennae and fore leg, cleared specimen.

3. Wing. Scale bars: 1 = 1.0 mm; 2 = 0.25 mm; 3 = 0.3 mm. Abbreviations: dm = discal medial cell; lbr = labrum; R, = anterior

branch of radius; R, = upper branch of third branch of radius; R, = lower branch of third branch of radius.

180 Toyohei Saigusa & Bradley J. Sinclair

Figs 4-5. Asioclinocera yunnanica gen. et sp. nov., lateral view. 4. Habitus. 5. Head, fore legs and thorax. Scale bars: 4 = 1.0 mm;

5=0.5 mm.

12. Gena strongly tapered with similar sized proboscis;

posterior margin of eye straight 2.0.0.0...

Fee net ea tpg eRe Se ak Whe Ao Clinocerella Engel

— Gena gradually tapered with broad _ proboscis;

posterior margin of eye rounded (Sinclair 1995:

IDES HOM WS) Erma cereus. We rel Wiedemannia Zetterstedt

Asioclinocera formosana gen. et sp. nov.

urn: lsid:zoobank.org:act:04456EED-§ 177-4B40-BAD3-539C9D10BA56

(Figs 1-3, 6, 8, 10)

Diagnosis. This species is distinguished from Asio-

clinocera yunnanica gen. et sp. nov. by the dark scutum,

lacking dark vittae; anteroventral and posteroventral

rows of setae less than half width of fore femur; and

shape of the clasping cercus.

Etymology. This species is named after the country of

origin, ‘Formosa’, a common name used by Europeans

until the 20" century.

Type material. Holotype. <, labelled: “[Taiwan: Tai-

chung-hs.] / Suchilanchi [24.32073° N, 121.28708° E]

1600m / Huangshan nr Lishan / Nov. 24, 1997 /

T. SAIGUSA col.”; “HOLOTYPE / Asioclinocera / for-

mosana / Saigusa & Sinclair [red label]” (KUMEF).

Bonn zoological Bulletin 71 (2): 177-183

Paratype. | 3, ,same collection data as for holotype,

except K. Masunaga leg. (CNC) [entire body dissected/

cleared ].

Description (wing length 3.5—3.8 mm). Male. Head

(Fig. 2): Oval, tapered ventrally, dark brown. Face with

thin pruinescence, narrower than width of antennal bases.

Ocellar triangle with pair of straight, anteriorly-directed

ocellar setae, longer and stronger than vertical setae; pair

of vertical setae. Postocular row of setae on upper half

strong; lower half slender; occipital setae scattered, se-

tae posterior to mouth slender. Antenna brown (Fig. 2);

scape shorter than pedicel, with several setulae; pedicel

with circlet of apical setae; postpedicel short-ovate; basal

segment of arista-like stylus wider than long; arista-like

stylus as long as head height. Thorax: Scutum dark

brown, with thin pruinescence; prescutellar depression

with whitish pruinescence in anterior view. Pleura dark

brown, concolourous with scutum (Fig. 1). Scutal setae

long, slender; acrostichals absent; 5 dorsocentral setae; 1

postpronotal seta; 1 presutural supra-alar seta; notopleu-

ron with several anterior setulae, 1 upper posterior no-

topleural seta; 2 postsutural supra-alar setae; 1 postalar

seta; pair of scutellar setae, closely approximated, longer

than prescutellar dorsocentral. Laterotergite with sever-

al long, slender setae. Legs: Coxae and femora brown,

©LIB

A new genus of Clinocerinae (Diptera: Empididae) from Taiwan and Yunnan (China) 181

paler than pleura, remaining segments dark brown. Fore

femur with rows of fine anteroventral and posteroven-

tral setae, length less than half width of femur; preapical

anteroventral spine-like seta longer than width of tibia

(Fig. 2). Fore tibia with row of strong, short anteroven-

tral setae, less than half length of tibia. Mid tibia with

basal posterodorsal seta, twice width of tibia. Hind tibia

with strong anteroventral seta on distal third, nearly twice

width of tibia; row of strong, erect posterodorsal setae

on distal third, subequal to width of tibia. Wing (Fig. 3):

Infuscate, without cloudy pattern; R, nearly straight,

ending in costa slightly beyond middle between R,,,

and R,. Halter brown. Abdomen: Sclerites dark brown,

darker than thorax, with short, scattered setae and thin

pruinescence. Tergum 7 membranous medially; sternum

7 with anterolateral corner prolonged into distinct projec-

tion. Tergum 8 roundly expanded laterally with numer-

ous long setae; sternum 8 as long as sternum 7, posteri-

or margin truncate, not prolonged beneath hypandrium.

Terminalia (Figs 6, 8): Hypandrium triangular, similar

in length to epandrium, with 6 apical setae. Phallus with

broad triangular base arising from hypandrium, articu-

lated with straight, stout apical section; apex expanded

with narrow, membranous crest. Ejaculatory apodeme

broad, rounded, narrowly extended into base of phallus.

cl cerc

ej apod ~

Epandrium rounded, with long setae along dorsal mar-

gin. Cercal plate with long setae on apical half. Clasping

cercus broad, flattened, spatulate; inner face clothed with

peg-like setae over most of surface. Surstylus narrow, ta-

pered to pointed apex; shorter than half length of clasp-

ing cercus.

Female. Unknown.

Asioclinocera yunnanica gen. et sp. nov.

urn: |sid:zoobank. org: act:6474F'D44-4EC 2-44 7F-SEBF-AA8AB92C754F

(Figs 4-5, 7, 9-10)

Diagnosis. This species is distinguished from Asio-

clinocera formosana gen. et sp. nov. by the pale brown

scutum, with dark median vittae and dark border around

pruinescence of prescutellar depression; anteroventral

and posteroventral rows of setae longer than half width

of fore femur; and shape of the clasping cercus.

Etymology. The species is named after the Chinese

province of Yunnan.

Type material. Holotype. <, labelled: “[Yunnan:

Lushui] / Gaolikungshan Nat. / Res.(2550-2700m) nr. /

Lushui [25.968736° N, 98.711538° E], 13. Aug. 1995 /

T. Saigusa col.”; “HOLOTYPE / Asioclinocera / yunnan-

ica / Saigusa & Sinclair [red label]” (KUMF).

Figs 6—7. Male terminalia of Asioclinocera gen. nov., lateral view. 6. A. formosana gen. et sp. nov. 7. A. yunnanica gen. et sp. nov.

Scale bars: 6 = 1.0 mm; 7 = 0.1 mm. Abbreviations: cl cerc = clasping cercus; ej apod = ejaculatory apodeme; hypd = hypandrium;

ph = phallus; sur = surstylus.

Bonn zoological Bulletin 71 (2): 177-183

©LIB

182 Toyohei Saigusa & Bradley J. Sinclair

clcerc. @ es

Sl Ph

Figs 8—9. Male terminalia of Asioclinocera gen. nov., lateral view. 8. A. formosana gen. et sp. nov. 9. A. yunnanica gen. et sp. nov.

Scale bars: 8 = 1.0 mm; 9 =0.1 mm. Abbreviations: cerc pl = cercal plate; cl cerc = clasping cercus; ej apod = ejaculatory apodeme;

epand = epandrium; hypd = hypandrium; ph = phallus; S = sternum; sur = surstylus; T = tergum.

Paratypes. Same collection data as for holotype (1 9,

KUMF); Yunlong-x. [25.79032° N, 99.10223° E], 3 km

NNW Zhichangm, 2500-2600 m, 7.v1.1996, T. Saigusa,

col. (1 4, CNC) [dissected].

Description (wing length 3.3-3.8 mm). Male. Head

(Fig. 5): Oval, tapered ventrally, dark brown. Face with

thin pruinescence, narrower than width of antennal bas-

es. Ocellar triangle with pair of straight, anteriorly-di-

rected ocellar setae, longer and stronger than vertical

setae; pair of vertical setae. Postocular row of setae on

upper half strong; lower half slender; occipital setae scat-

tered, setae posterior to mouth slender. Antenna brown

(Fig. 5); scape shorter than pedicel, with several setulae;

pedicel with circlet of apical setae; postpedicel short-

ovate; basal segment of arista-like stylus wider than

long; arista-like stylus as long as head height. Thorax:

Scutum pale brown, with thin pruinescence; medially

with broad, dark brown vitta extending to prescutellar

depression; prescutellar depression with whitish pru-

inescence with dark border in anterior view. Pleura pale

brown, concolourous with scutum (Fig. 5). Scutal setae

long, slender; acrostichals absent; 5 dorsocentral setae;

1 postpronotal seta; 1 presutural supra-alar seta; noto-

pleuron with several anterior setulae, 1 upper posterior

notopleural seta; 2 postsutural supra-alar setae; 1 postalar

seta; pair of scutellar setae, closely approximated, longer

than prescutellar dorsocentral. Laterotergite with several

long, slender setae. Legs: Pale brown, concolourous with

pleura, segments gradually darker apically. Fore femur

with rows of long, fine anteroventral and posteroventral

setae, longer than half width of femur, similar to length

of spine-like seta (Fig. 5); preapical anteroventral spine-

like seta longer than width of tibia. Fore tibia with row

Bonn zoological Bulletin 71 (2): 177-183

of strong, short anteroventral setae, less than half width

of tibia (Fig. 5). Mid tibia with basal posterodorsal seta,

twice width of tibia. Hind tibia with thin anteroventral

seta on distal third, less than twice width of tibia; row

of strong, erect posterodorsal setae on distal third, lon-

ger than width of tibia. Wing (Fig. 4): Infuscate, without

cloudy pattern; R, distinctly arched, ending in costa near

middle between R,,, and R.. Halter brown. Abdomen:

Sclerites dark brown, darker than thorax, with short, scat-

tered setae and thin pruinescence. Tergum 7 membranous

medially; sternum 7 with anterolateral corner prolonged

into distinct projection. Tergum 8 roundly expanded lat-

erally with numerous long setae; sternum 8 as long as

sternum 7, posterior margin prolonged beneath hypan-

drium. Terminalia (Figs 7, 9): Hypandrium rectangular,

similar in length to epandrium, apex slightly tapered with

6 apical setae. Phallus with broad triangular base aris-

ing from hypandrium, stout apical section sinuous; apex

expanded with narrow, membranous crest. Ejaculatory

apodeme broad, rounded, narrowly extended into base

of phallus. Epandrium rounded, with long setae mostly

along dorsal margin. Cercal plate with long setae on ap1-

cal half. Clasping cercus digitiform, tapered apically with

rounded apex; strong, thickened setae along inner poste-

rior margin and inner apex; apex crowned with 3 spine-

like setae; without peg-like setae on inner surface. Sur-

stylus digitiform, strongly arched, tapered to rounded

apex; longer than length of clasping cercus.

Female. Similar to male except face slightly broader

than width of antennal sockets.

Acknowledgements. The following collector kindly provid-

ed specimens to the first author: K. Masunaga. Jeff Cumming

©LIB

A new genus of Clinocerinae (Diptera: Empididae) from Taiwan and Yunnan (China) 183

Fig. 10. Distribution of Asioclinocera gen. nov.

(Canada) and Maria Ivkovi¢ (Croatia) kindly commented on

the manuscript.

REFERENCES

Collin JE (1941) Some Pipunculidae and Empid-

idae from the Ussuri region on the far eastern bor-

der of the U.S.S.R. (Diptera). Proceedings of the En-

tomological Society of London (B) 10: 218-248.

https://doi.org/10.1111/).1365-3113.1941 tb00684.x

Cumming JM, Wood DM (2017) [Chapter] 3. Adult morphol-

ogy and terminology. Pp. 89-133 in: Kirk-Spriggs AH &

Sinclair BJ (eds) Manual of Afrotropical Diptera. Volume 1.

Introductory chapters and keys to Diptera families. Suricata

4, South African National Biodiversity Institute, Pretoria

Loew H (1858) Noch eine neue Clinocera. Wiener Entomolo-

gische Monatschrift 2: 386-387

Meigen JW (1803) Versuch einer neuen Gattungs-Eintheilung

der europaischen zweifligligen Insekten. Magazin fur Insek-

tenkunde 2: 259-281

Saigusa T (1986) New genera of Empididae (Diptera) from

eastern Asia. Sieboldia 5: 97-118

Saigusa T, Sinclair BJ (2016) Revision of the Japa-

nese species of Trichoclinocera Collin (Diptera: Em-

Bonn zoological Bulletin 71 (2): 177-183

110 115

125

(A. formosana ®)

(A. yunnanica )

le (e) on

= oe

120 7 7's BA WwaR

pididae: Clinocerinae). Zootaxa 4103: 201-229.

https://doi.org/10.11646/zootaxa.4103.3.1

Saigusa T, Sinclair BJ (2021) Revision of the Trichoclinocera

yixianensis species-group from eastern Asia (Diptera: Emp-

ididae: Clinocerinae). Bonn zoological Bulletin 70 (1): 51-

65. https://doi.org/10.20363/BZB-2021.70.1.051

Sinclair BJ (1994) Revision of the Nearctic species of 7ri-

choclinocera Collin (Diptera: Empididae; Clinocer-

inae). The Canadian Entomologist 126: 1007-1059.

https://doi.org/10.4039/Ent1261007-4

Sinclair BJ (1995) Generic revision of the Clinocerinae

(Empididae), and description and phylogenetic relation-

ships of the Trichopezinae, new status (Diptera: Em-

pidoidea). The Canadian Entomologist 127: 665-752.

https://doi.org/10.4039/Ent127665-5

Shorthouse DP (2010) SimpleMappr, an online tool to

produce publication-quality point maps. Online at

http://www.simplemappr.net [last accessed 27 Jun. 2022]

Smith KGV (1965) Diptera from Nepal Empididae. Bulletin of

the British Museum (Natural History), Entomology 17 (2):

63-112. https://doi.org/10.5962/bh1.part. 14808

Zetterstedt JW (1838) Sectio tertia. Diptera. Dipterologia Scan-

dinaviae amicis et popularibus carissimus. Pp. 477-868 in:

Insecta Lapponica L. Voss (ed.) Lipsiae, vi + p. 1140

©LIB

BHL

Blank Page Digitally Inserted

LIB Leibniz Institute for the Analysis

== of Biodiversity Change

Bonn zoological Bulletin 71 (2): 185-191

2022 Szawaryn K.

https://do1.org/10.20363/BZB-2022.71.2.185

museum

: KOENIG

ISSN 2190-7307

http://www.zoologicalbulletin.de

Research article

urn:|sid:zoobank.org:pub:61C4C8E6-3867-4E0 1-8456-079C3797F4B4

New species of Monocoryna Gorham, 1885 (Coccinellidae: Monocoryninae)

from India

Karol Szawaryn

Museum and Institute of Zoology, Polish Academy of Sciences, Wilcza 64, 00-679 Warszawa, Poland

Email: k.szawaryn@gmail.com

urn:|sid:zoobank.org:author:D741C759-6CDD-4B61-BE93-3ECC2918A73F

Abstract. The peculiar ladybird genus Monocoryna Gorham, 1885 is distributed in Eastern part of the Oriental Region,

and it inhabits mainly the Malay Archipelago, with only nine species known. Recent molecular studies revealed that

Monocoryna forms a separate branch on the phylogenetic tree of Coccinellidae, the subfamily Monocoryninae. In this

study, a new species M. indica sp. nov. is described from Andippatti Hills in India. The discovery extends the range of the

genus to the Western part of the Oriental Region. Additionally, male genitalia of M. yamashinai Sasaji from Thailand are

described and illustrated for the first time. A distribution map and an updated key to all species of Monocoryna are also

provided.

Key words. Coleoptera, Coccinelloidea, taxonomy, Tamil Nandu, ladybird beetles, new species.

INTRODUCTION

The genus Monocoryna Gorham, 1885 was described by

Gorham (1885) to accommodate a single species, M. de-

cempunctata Gorham, 1885 from Sumatra, Indonesia.

Due to its peculiar structure of antennae, extraordinary

large and rounded terminal antennomere, and lobbed

four-tarsomere tarsi it was classified by Gorham (1885)

as Endomychidae Leach, 1815. That placement was also

followed by subsequent authors (Arrow 1920; Strohecker

1953). After detailed examination of the morphology of

specimens of Monocoryna Miyatake (1988) realized that

it belongs to the ladybird beetles (Coccinellidae Latreille,

1807). He also proposed a new tribe Monocorynini for

this peculiar genus.

Gorham (1885) suggested a close relationship of Mono-

coryna with the ladybird genus Jetrabrachys Kapur, 1948

(= Litophilus Frolich, 1799). Later Firsch (1996) pointed

out that Monocoryna also resembles in some morpholog-

ical features the South African genus Mimilitophilus Ar-

row, 1920, which was placed either in Litophilini (Crow-

son, 1952) or Coccidulini (lablokoff-Khnzorian, 1974).

Studies of Seago et al. (2011) and Robertson et al. (2015)

based on molecular data showed, however, that Mono-

coryna does not form a monophyletic clade with Tetrab-

rachys but is grouped with Mimolitophilus. A recent

molecular analysis (Che et al. 2021) based on 96 molec-

ular markers showed that Monocoryna forms a separate

branch within Coccinellidae between subfamilies Mi-

croweseinae and Coccinellinae, thus authors proposed a

new subfamily Monocoryninae for this genus.

Received: 02.01.2022

Accepted: 17.10.2022

Monocoryna is endemic to the Oriental Region and

has been known from its eastern part so far. Slipinski

and Jadwiszczak (2000) listed nine currently recognized

species of Monocoryna, of which four are known from

Borneo (Malaysia), two from Mindanao (Philippines),

one from mainland Malaysia, one form Chiang May

(Thailand), and one occurring in Sumatra and Java (In-

donesia). Specimens of Monocoryna are rarely found in

collections, and each species is known from a single or

just a few specimens. No information about its behavior

or food preferences is known except scarce information

about general habitat that can be concluded from the label

data as a mainly forest species occurring at high altitudes

(1250-3500 m as.l.) (Miyatake 1988; Slipinski & Jad-

wiszczak 2000).

During research in the collection of the Museum and

Institute of Zoology in Warsaw, a single specimen of

Monocoryna from India was found which appeared to be

a new species. The new species here described extends

the range of the genus to the Western part of the Oriental

Realm.

MATERIAL AND METHODS

The holotype is deposited in the Museum and Institute of

Zoology, Polish Academy of Sciences in Warsaw (MIZ).

Material from the Muséum d’ Histoire Naturelle, Geneva

(MHNG) and the Staatliches Museum fiir Naturkunde,

Stuttgart (SMNS) were also studied.

Corresponding editor: D. Ahrens

Published: 24.10.2022

186 Karol Szawaryn

Genitalia were dissected, cleared in a 10% solution of

KOH, rinsed with distilled water, transferred to glycerol,

and examined on slides. All colour images were taken

using a stereo microscope Leica MZ 16 with a digital

camera IC 3D; final images were produced using Helicon

Focus 5.0X64 and AdobePhotoshop CS6 software. Mor-

phological terminology follows Slipinski (2007).

RESULTS

Taxonomy

Family Coccinellidae Latreille, 1807

Subfamily Monocoryninae Miyatake, 1988

Tribe Monocorynini Miyatake, 1988

Genus Monocoryna Gorham, 1885

Monocoryna Gorham, 1885: 527.

Type species: Monocoryna decempunctata Gorham,

1885 (by monotypy).

Walteria Sicard, 1913: 511.

Type species: Walteria antennalis Sicard, 1913 (by

monotypy). Synonymized by Miyatake 1988: 28.

A | B

“—

Sicara Strand, 1942: 392. New name for Walteria Si-

card, 1913, not Schultze, 1885. Synonymized by Fursch

1996: 195.

Monocoryna indica sp. nov.

urn: |sid:zoobank.org:act: DC3 EA 19F-8 EDB-4B74-A9DC-6D42841A 2427

(Figs 1—2, 5)

Diagnosis. M. indica sp. nov. is most similar to

M. yamashinai Sasaji, 1989 described from Thailand

(Chiang Mai). Both species are dark brown to black,

however, in M. yamashinai each elytron is bordered with

a reddish-orange band with central disc dark brown,

whereas M. indica sp. nov. has entirely dark brown el-

ytra. It can be also separated by the shape of male geni-

talia, in M. indica parameres are slender, broadest in the

middle, with apices narrowly rounded, penis guide mod-

erately well developed and sub-triangular, and penis with

single needle-like spine apically, while in M. yamashinai

parameres are broadened apically with broadly rounded

apices, penis guide reduced to a narrow projection, and

penis with two apical spines. From the other species,

M. indica sp. nov. can be separated by uniformly dark

brown colouration, while other have bright yellowish to

red-orange maculae on elytra, sometimes fused forming

C Rrdigpalki Wills

ek ee

Besuchet Lob! Mussard

R.D. Pope det. 1957

Fig. 1. Monocoryna indica sp. nov. A. Habitus. B. Pronotum. C. Holotype labels.

Bonn zoological Bulletin 71 (2): 185-191

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New species of Monocoryna Gorham, 1885 from India 187

aS :

Fig. 2. Monocoryna indica sp. nov. A. Abdomen. B. Tegminal strut. C. Tegmen. D. Male abdominal segments IX and X. E. Tip

of penis. F. Penis, lateral. G. Tegmen, inner. H. Tegmen, lateral. I. Ventrite 6. J. Tergite VII. B—D in the same scale; F—H in the

same scale.

188 Karol Szawaryn

bands. It also has apically broadened penis apex with dis-

tinct needle like sclerite while other known species have

simple, pointed apex.

Etymology. The newly described species is named af-

ter the country of its origin, India.

Type material

Holotype. o', “INDIA, Madras, Andippatti Hills,

3.X1.72, Besuchet Lob] Mussard/ Monocoryna sp. R.D.

Pope det. 1987” (MIZ); Fig. 1C.

Description. Male. Length = 5.9 mm, width = 3.9 mm.

Body and colour. Broadly oval (Fig. 1A), moderate-

ly convex dorsally, covered with moderately long, or-

ange-brown vestiture. Pronotum dark brown with paler

lateral sides (Fig. 1B), hypomeron reddish-brown, elytra

entirely dark brown, epipleura dark brown with inner

margin reddish-brown, meso and metaventrite brown-

ish-black, mouthparts and tarsi brown, antennomeres 1—5

brown, 6—8 black. Head and pronotum densely covered

with punctures of uniform size, elytra covered with inter-

mixed smaller and larger punctures arranged irregularly.

Wingless.

Head. Dorsally with long supraorbital sulci; with deep,

long ventral antennal grooves. Eyes coarsely faceted,

small, distance between eyes more than half width of the

head. Terminal maxillary palpomere slightly broadened

apically, longer than its width. Antenna with scape dis-

tinctly swollen with anterior surface roundly expanded

WwW |

forwards; pedicel small, transverse; antennomere 3 elon-

gate, longer than antennomeres 4—6 combined; antenno-

meres 4—7 transverse.

Pronotum. With complete anterior, lateral margins

bordered, lateral borders anteriorly not touching anterior

pronotal margin, posteriorly extending to lateral parts of

hind margin of pronotum. Base of pronotum with two

small, subtriangular sulci. Hypomeron with broad con-

cave area in anterior part. Anterior prosternal margin pro-

duced forwards forming a chinpiece, partially covering

mouthparts. Prosternal process broad with sinusoidal,

subparallel, separate and complete carinae.

Scutellar shield small, subtriangular, bordered. Elytral

margin narrow, not visible from above. Epipleuron in-

complete apically. Mesoventral intercoxal process broad-

er than mesocoxal diameter. Metaventral postcoxal lines

joined medially forming a straight line on mesoventral

process, descending laterally. Metanepimeron with cari-

na near anterior margin.

Legs. Slender with trochanters sub-quadrate, tibiae

without apical spurs, tarsi with four tarsomeres, tarsal

claw simple without basal angulation.

Abdomen. With six ventrites (Fig. 2A); ventrite | long,

more than 1.5 x as long as ventrite 2 at the level of post-

coxal lines, postcoxal lines complete, recurved, deep,

angulate, reaching posterior margin of the ventrite and

sub-parallel to its margin; ventrite 5 broadly truncate api-

Fig. 3. Monocoryna yamashinai Sasaji, 1989. A. Habitus, dorsal. B. Habitus, lateral. C. Habitus, frontal.

Bonn zoological Bulletin 71 (2): 185-191

©LIB

New species of Monocoryna Gorham, 1885 from India 189

Fig. 4. Monocoryna yamashinai Sasaji, 1989. A. Tip of penis. B. Penis, lateral. C-D. Tegmen, inner. E. Tegmen, lateral.

cally; ventrite 6 narrow, broadly rounded (Fig. 21); tergite

VIII large, arcuate (Fig. 2J). Apodeme of sternite IX long

(Fig. 2B, D), rod like, distinctly broadened basally with

median part translucent. Tergite X transverse.

Genitalia. Tegmen symmetrical (Fig. 2C, G—H), phal-

lobase with long additional projection on outer surface:

penis guide in inner view small, sub-triangular, moder-

ately broad, with distinct median carina on outer surface;

parameres in inner view large, long oval, broadest in the

middle, densely setose on lateral and apical margins, in

lateral view narrow, with margins straight; tegminal strut

simple rod-like, very long. Penis long (Fig. 2E—F), slen-

der, without capsule, apex slightly broadened with dis-

tinct needle-like projection.

Female. Unknown.

Distribution. India, Tamil Nandu, Andippatti Hills.

Bonn zoological Bulletin 71 (2): 185-191

Monocoryna yamashinai Sasaji, 1989

Monocoryna yamashinai Sasaji, 1989: 117.

Figs 3—5

Material examined. | 0’, Thailand, Chiang Mai Prov.,

Doi Sanyao 1000 m, Schwendinger 7.9.91/ Monocoryna

yamashinai det. A. S. Slipinski/ COL 3831 (MHNG) —

1 2, N. Thailand, NE Chiang Mai 6 km NE Doi Saket,

27.11.1998, R. Grimm/ Museum Stuttg (SMNS).

Description. Genitalia. Tegmen symmetrical (Fig. 4C—

E), phallobase with additional short projection on outer

surface; penis guide in inner view very small, sub-trian-

gular, narrow; parameres in inner view large, long oval,

distinctly broadened apically, sparsely setose on apical

margin, in lateral view more broad, with margins more

rounded; tegminal strut simple rod-like, very long. Penis

long (Fig. 4A—B), slender, without capsule, apex distinct-

ly narrowed in apical part with two terminal projections,

one short and sinuate, the second longer and needle-like.

Remarks. Originally 1 yamashinai was described by

Sasaji in 1989 based on a single female specimen from

©LIB

190 Karol Szawaryn

indica sp. nov.

decempunctata

Fig. 5. Distribution map of species of Monocoryna Gorham, 1885.

Thailand (Chiang Mai). Since that time no new records

were published. Monocoryna yamashinai can be eas-

ily separated from the most similar M. indica but also

from the other congeners by its colour pattern of elytra,

with disc dark brown and lateral margin with broad red-

dish-orange band (Fig. 3A—C). Examination of the addi-

tional material allowed description of the male genitalia

of this species.

Distribution. Thailand, Chiang Mai.

Key to species of Monocoryna Gorham, 1885

1. Elytral disc without spots or transverse bands,

uniformly dark brown to blackish |... 2

— Elytral disc with distinct paler spots, sometimes

arranged in transverse bands .............00...ccceeeeee 3

2. Elytra uniformly dark brown to black, lateral margin

of elytra without paler margin. India .........000....000....

Reena) be hry a emia ae M. indica sp. nov.

— Elytra with lateral margin with paler yellow-orange

border. Thailand .......... M. yamashinai Sasaji, 1989

3. Hind wings well developed; humeral callus distinct

— Hind wings atrophied; humeral callus indistinct .... 7

4. Elytra brown to blackish brown with 10—12 isolated

— Elytra blackish brown with 3 reddish brown bands.

Malay Peninsula ................ M. fasciata Arrow, 1920

5. Elytra with more than 10 spots. Malaysia (Borneo:

= SELVA WET O! SOLS sa2s ils: tlw ts cet ld cae erashice ces 6

6. Elytral spots small and roundish; body shortly oval;

smaller (4-5 mm in length). Malaysia (Borneo:

OTC: | rea M. moultoni (Sicard, 1913)

Bonn zoological Bulletin 71 (2): 185-191

philippinensis

nicoleberthiae

antennalis

— Elytral spots larger and elongate; body oblong-

oval, larger (7-8 mm in length). Malaysia (Borneo:

hs) Yc LUN Soampas decks M. magnopunctata Miyatake, 1988

7. Elytra piceous brown with 13 paler spots, of which

postscutellar spot is situated on the elytral suture, two

subapical spots are connected with each other at their

tips, forming a V-shaped marking; scutellar shield

moderate in size. Malaysia (Borneo: Sarawak) ........

ee RT Mk x Ace WE M. antennalis (Sicard, 1913)

— Elytra blackish brown to black with 10 paler spots,

none of them situated on the elytral suture; scutellar

Shilel Chavet: Siti Lee aes tee, 09h. Rete eee choi easy 8

8. Body length less than 6.5 mm. Indonesia (Java,

Sumatra) ............. M. decempunctata Gorham, 1885

— Body length more than 7.5 mm. Philippines .......... 9

9. Antennal club black, lateral pronotal sulcus joined

to anterior margin of pronotum, body larger, around

9mm in length .....M. nicolebertiae Jadwiszczak &

Slipinski, 2000

— Antennal club orange, lateral pronotal sulcus not

reaching anterior margin of pronotum, body shorter,

AROUMIG Sei Mia CM OE Ree aay be 8 Nea ae

...M. philippinensis Jadwiszczak & Slipinski, 2000

Acknowledgments. Przemek Szymroszczyk (MIZ), Giulio

Cuccodoro (MHNG), Wolfgang Schawaller (SMNS) and Adam

Slipinski (CSIRO Canberra, Australia) are thanked for a loan

of the materials used in this study. Maria Kosciesza (MIZ) is

thanked for providing necessary literature. Hermes Escalona

and a second anonymous reviewer are thanked for valuable

comments on the early version of the manuscript.

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New species of Monocoryna Gorham, 1885 from India 191

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©LIB

LIB. Leibniz Institute for the Analysis

=— of Biodiversity Change

ISSN 2190-7307

http://www.zoologicalbulletin.de

Bonn zoological Bulletin 71 (2): 192—203

2022 Barjadze S. et al.

https://do1.org/10.20363/BZB-2022.71.2.192

museum

: KOENIG

Research article

urn:|sid:zoobank.org:pub:5F5C593E-BB21-4A99-A36A-9E7F92DB3957

Descriptions of a new aphid species of Lipaphis Mordvilko, 1928

(Hemiptera: Aphididae) from South Korea and the hitherto unknown

oviparae of Lipaphis ruderalis Borner, 1939 from the Czech Republic

‘Institute of Zoology, Ilia State University, Giorgi Tsereteli 3, 0162, Tbilisi, Georgia

? Institute of Biology, Biotechnology and Environmental Protection, Faculty of Natural Sciences,

University of Silesia in Katowice, Bankowa 9, 40-007 Katowice, Poland

?3 Institute of Zoology, Ministry of Education and Sciences, Republic of Kazakhstan, Academgorodok, Al-Farabi avenue, 93,

Almaty, 050060, Kazakhstan

“Corresponding author: Email: shalva.barjadze@yahoo.com

'urn:lsid:zoobank.org:author:63 A BE1 B2-8A56-42C8-BC34-1119D3A2ECBC

2urn:Isid:zoobank.org:author:78C290A3-D07B-4A F9-9358-ED8C05A702BF

3urn:|sid:zoobank.org:author:8AE088A0-72C 1-4282-A A4E-EB16CCD9FC24

Abstract. Apterous viviparous 9° 9 of Lipaphis (Lipaphidiella) holmani sp. nov., living on Lepidium virginicum L. (Bras-

sicaceae) in South Korea, are described and differences with all known species belonging to the subgenus Lipaphidiella

Doncaster, 1954 and Lepidium-feeding Lipaphis spp. are presented. In addition, the poorly known Lipaphis sisymbrii Bo-

zhko, 1976 is redescribed based on apterous and alate viviparous 2 9 from Kazakhstan and its subgeneric position based

on the morphological features is revised. Finally, the hitherto unknown oviparous 9 9 of Lipaphis ruderalis Borner, 1939

are described from the Czech Republic. An identification key for apterous viviparous 9 9 of all species currently included

in the subgenus Lipaphidiella is given.

Key words. Aphids, Asia, Macrosiphini, Lipaphidiella, taxonomy.

INTRODUCTION

The genus Lipaphis Mordvilko, 1928 (Aphididae: Aphi-

dinae: Macrosiphini) was erected for Aphis erysimi

Kaltenbach, 1843 (type species). The genus comprises

12 species within two subgenera: Lipaphidiella Don-

caster, 1954 (3 species) and Lipaphis Mordvilko, 1928

(9 species) which are associated without host alternation

with Brassicaceae mostly in the Western Palaearctic re-

gion (Blackman 2010; Blackman & Eastop 2022). The

mustard aphid Lipaphis pseudobrassicae (Davis, 1914)

is an important pest of Brassica crops worldwide (Black-

man & Eastop 2000). Lipaphis is related to Brevicoryne

Das, 1915, and is characterized 1n apterae by weakly de-

veloped antennal tubercles, short antennae, the absence of

secondary rhinaria on antennal segment II, a wrinkled or

reticulate sclerotic dorsum, cylindrical or distally slight-

ly swollen siphunculi, broadly based, tongue-shaped, or

elongate triangular cauda with 4-6 setae, and 3:3:2 or

3:3:3 first tarsal chaetotaxy (Heie 1992; Blackman &

Eastop 2006; Blackman 2010).

During an examination of Jaroslav Holman’s aphid

collection deposited at Biology Centre CAS, Institute of

Entomology, Ceské Budéjovice, Czech Republic (IECA),

fourteen slides with aphids determined by J. Holman as

“Lipaphis” or “Lipaphis aff. lepidii”, sampled on Lepidi-

um virginicum L. (Brassicaceae) in South Korea and two

slides with aphids determined as “Lipaphis ruderalis”’ by

J. Holman, sampled on Lepidium ruderale L. (Brassica-

ceae) in the Czech Republic were found.

The following 12 aphid species are recorded on Lepid-

ium virginicum worldwide: Aphis craccivora (Koch,

1854), A. fabae Scopoli, 1763, A. gossypii Glover, 1877,

Brevicoryne brassicae (Linnaeus, 1758), Lipaphis pseu-

dobrassicae (Davis, 1914), Macrosiphum euphorbiae

(Thomas, 1878), Myzus ascalonicus Doncaster, 1946,

M. ornatus Laing, 1932, M. persicae (Sulzer, 1776), Ne-

omyzus circumflexus (Buckton, 1876), Pemphigus pop-

ulitransversus Riley, 1879, and Protaphis middletonii

(Thomas, 1879) (Holman 2009; Villalobos Muller et al.

2010; Blackman & Eastop 2022), while 11 aphid species

(Aphis craccivora Scopoli, 1763, A. gossypii Glover,

1877, Aulacorthum solani (Kaltenbach, 1843), Brevic-

oryne brassicae (Linnaeus, 1758), Lepidaphis deformans

(Nevsky, 1929), Lipaphis erysimi (Kaltenbach, 1843),

L. lepidii (Nevsky, 1929), L. ruderalis Borner, 1939,

Macrosiphum euphorbiae (Thomas, 1878), Myzus cerasi

(Fabricius, 1775), M. persicae (Sulzer, 1776)) are record-

Received: 03.04.2022

Accepted: 18.10.2022

Corresponding editor: D. Ahrens

Published: 24.10.2022

New species of Lipaphis Mordvilko, 1928 193

ed on Lepidium ruderale (Holman 2009; Blackman &

Eastop 2022).

Among the examined material an undescribed Li-

paphis species from South Korea feeding on Lepidium

virginicum, and hitherto unknown oviparae of Lipa-

phis ruderalis Borner, 1939 were recognized. The un-

described species and Lipaphis ruderalis belong to the

subgenus Lipaphidiella because they have a medial su-

pracaudal process on abdominal tergite VII (Doncaster

1954; Heie 1992; Blackman & Eastop 2006). Lipaphis

sisymbrii Bozhko, 1976 was described extremely briefly

on Sisymbrium polymorphum (Murray) Roth (Brassica-

ceae) from Ukraine (Bozhko 1976). Therefore, it became

necessary to redescribe it in more detail.

Here we describe the new Lipaphis species from South

Korea and the hitherto unknown oviparae of Lipaphis

ruderalis; in addition, we redescribe the poorly known

L. sisymbrii and revise its subgeneric position.

MATERIAL AND METHODS

The measurements of the morphological characters of

aphid specimens mounted on slides were done according

to Blackman & Eastop (2006) using an Accu-Scope—

Exc—350 compound microscope, and each character was

measured using CaptaVision Software ver. 3.9. and pho-

tographed with a Excelis HDS digital camera. Measure-

ments are given in millimeters (mm). Actual host plant

names are given according to The Plant List (2013).

Abbreviations for morphological terms

ABD TERG = abdominal tergite

ANT = antenna

ANT I-VI, Vb, VIb = antennal segments I-VI, Vb , VIb

respectively

ANT III BD = basal diameter of antennal

segment III

BL = length of body along midline,

excluding cauda

BW = maximum width of body

FTC = first tarsal chaetotaxy

HFEM = hind femur

HTIB = hind tibia

HT II = second segment of hind tarsus

HW = head width across compound eyes

L = length

MSL = maximal setal length

MTu = marginal tubercle(s)

PT = processus terminalis

SIPH = siphunculus

URS = ultimate rostral segments IV and

V together

W = width

Bonn zoological Bulletin 71 (2): 192—203

Institutional abbreviations

BMNH = Natural History Museum, London, UK

IECA = Biology Centre CAS, Institute of Ento-

mology, Ceské Budéjovice, Czech Republic

IZAK = Institute of Zoology, Ministry of Education

and Sciences, Almaty, Kazakhstan

IZISU_ = Institute of Zoology, Ilia State University,

Tbilisi, Georgia

RESULTS

Taxonomy

Lipaphis holmani sp. nov.

urn:Isid:zoobank.org:act: 1A3EC6B8-592F-41FC-BC 10-82730DC8F712

Figs 1, 3, 5, Table 1

Type material

Holotype. N 99HO39A, apterous viviparous ° onslide,

on Lepidium virginicum L., SOUTH KOREA, Gyeongg1

Province, Suwon, Yeogisan (as “Yogi-san, Kvonggi-do

prov.” On slide label), 37°16’49” N, 126°59’0” E, 105 m

a.s.1., 09.v.2003, J. Holman leg. (IECA).

Paratypes. 13 apterous viviparous 9@ on slides: N

99HO39A, 10 apterous viviparous 99 on slides, the

same data as for holotype, TECA); N TA—T-—00054-55,

2 apterous viviparous 9 on slides, the same data as

for holotype, (IZISU); N 99HO39A, 1 apterous vivipa-

rous 9 on slide, the same data as for holotype, (BMNH).

Etymology. The authors have chosen the specific name

in honor of Dr Jaroslav Holman, who was collector of

this species and outstanding aphid taxonomist.

Description

Apterous viviparous 9 (n= 14)

Color in life: Unknown. Pigmentation on slide: ANT I-

II brown, concolorous to head dorsum; ANT III-IV pale:

ANT V pale with brown apex; ANT VI, clypeus, rostral

I—V and tarsi dark brown; cluster of round—shaped dark

brown spots always developed in pleural position on the

thorax; coxae and trochanters pale brown; femora with

pale bases and gradually darker remaining parts; tibiae

with pale middle section and pale brown bases and dark

brown apices; dark intersegmental sclerites on abdomen

always absent; SIPH pale brown with darker apical 4 or

Ys, brown transverse bands present on ABD TERG VII-—

VIII; subgenital and anal plates and cauda pale brown;

stigmal plates brown (Fig. 1).

Morphological characters: Body oval. Frons w-shaped

(Figs 1, 3). ANT 6-segmented (Figs 1, 3). ANT tubercles

lower than rounded medial frontal tubercle. ANT cuticle

imbricated. Primary rhinaria ciliated. Secondary rhiniaria

always absent. Rostrum short, reaching to middle cox-

ae. URS sub-parallel sided with pointed apex and fine

©LIB

194 Shalva Barjadze et al.

Figs 1-9. 1, 3, 5. Lipaphis holmani sp. nov. apterous viviparous @. 2, 4, 6-9. L. ruderalis Borner, 1939 oviparous 2. 1. Apterous

viviparous 9, general view. 2. Oviparous @, general view. 3. Head and antenna. 4. Head. 5. Posterior end of abdomen. 6. Antenna.

7. Hind tibia with scent plaques. 8. Dorsal abdominal cuticle. 9. Posterior part of abdomen.

New species of Lipaphis Mordvilko, 1928

195

Table 1 (continued on next page). Metric and meristic characteristics of apterous viviparous 9 9 of Lipaphis (Lipaphidiella) holm-

ani sp. nov. and oviparous °° of L. (Lipaphidiella) ruderalis Borner, 1939. Explanations about abbreviations used in the table are

given in the Material and methods section.

Apterae, L. holmani sp. nov.

Oviparae, L. ruderalis Borner, 1939

n=14 n=6

Characters

BL 1.482-1.854 1.457-1.795

BW 0.853-—1.026 0.856—1.062

ANT 0.576—0.676 0.565—0.710

ANT II 0.168—0.212 0.185—0.237

MSL ANT III 0.003—0.007 0.005—0.006

ANT IV 0.069—0.087 0.050—0.074

ANT V 0.071—0.095 0.070-0.089 (if 6 segmented)

ANT V bor VI b 0.063—0.079 0.069—0.076

PT 0.094—0.122 0.112—0.149

HW 0.387-0.410 0.419-0.445

MSL on cephalic dorsum 0.009-0.012 0.008—0.010

URS 0.098—0.107 0.085—0.090

Posterior setae on hind trochanter 0.014—0.026 0.013-0.018

HFEM 0.329-0.404 0.314-0.358

Dorsal MSL on femora 0.006—0.011 0.007—0.010

HTIB 0.594—0.700 0.521—0.606

Dorsal MSL in the middle of HTIB 0.009-0.013 0.010-0.018

HT IIL 0.111-0.124 0.107—0.123

MSL ABD TERG III 0.006-0.011 0.007—0.010

MSL ABD TERG VIII 0.007—0.010 0.009-0.011

SIPH-L 0.226-0.271 0.199-0.273

Cauda L 0.143—0.184 0.191-0.213

Ratios

ANT/BL 0.35—0.40 0.34-0.44

PT/ANT V b or VI b 1.33-1.61 1.62—2.06

PT/ANT V (if 6 segmented) 1.15-1.48 1.62—1.94

PT/ANT IV 1.18-1.58 1.94-2.98

PT/ANT HI 0.49-0.63 0.47-0.73

PT/HW 0.24—0.28 0.27-0.35

PT/URS L 0.94-1.16 1.32-1.67

PT/SIPH 0.37—-0.48 0.45—0.68

ANT IV/ANT V (if 6 segmented) 0.81—1.04 0.59-0.93

ANT III/ANT IV+ANT V (if 6 segmented) 1.08-1.27 1.27-1.75

ANT III+PT/SIPH 1.04—1.40 1.29-1.61

ANT ITI/URS L 1.67—2.12 2.05—2.79

ANT III/SIPH L 0.64—0.94 0.76-0.95

MSL ANT III/ANT III BD 0.15—0.33 0.20-0.28

MSL on cephalic dorsum/ANT III BD 0.43—0.60 0.36—0.45

URS L/URS W 1.72-2.26 1.27-1.51

URS L/ANT VI b 1.26-1.60 1.17-1.28

URS L/HW 0.24—0.26 0.20-0.21

URS L/HT II L 0.81-0.92 0.72-0.79

URS L/SIPH L 0.38-0.44 0.33-0.45

Posterior setae on hind trochanter/suture 0.31—0.56 0.19-0.30

HFEM/BL 0.20-0.25 0.19-0.23

HFEM/HW 0.83—1.00 0.75—0.83

Dorsal MSL on femora/ANT HI BD 0.27—0.55 0.32-0.43

Dorsal MSL on femora/suture 0.15—0.24 0.11-0.14

Bonn zoological Bulletin 71 (2): 192—203

©LIB

196 Shalva Barjadze et al.

Table 1. continued.

Apterae, L. holmani sp. nov. Oviparae, L. ruderalis Borner, 1939

n=14 n=6

Ratios

HTIB/BL 0.36-0.44 0.31-0.38

HTIB/HW 1.46-1.71 1.24-1.39

Dorsal MSL in the middle of HTIB/ANT III BD 0.43-0.65 0.43-0.82

Dorsal MSL in the middle of HTIB/diameter of 0.25-0.41 0.16—0.34

HTIB at the middle section

HT I L/HW 0.28-0.31 0.25-0.29

MSL ABD TERG III/ANT HI BD 0.26-0.52 0.30-0.48

MSL ABD TERG VIII/ANT HI BD 0.30-0.48 0.39-0.52

SIPH L/ SIPH BW 3.23-4.59 3.16—-4.06

SIPH L/BL 0.13-0.17 0.14-0.16

SIPH L/HW 0.55—-0.65 0.47-0.65

SIPH L/HT II L 1.85-2.31 1.63-2.32

Cauda L/Cauda W 1.17-1.57 1.50-1.95

Cauda L/ANT III 0.79-0.99 0.81-1.03

Cauda L/HW 0.36-0.43 0.44—0.50

Cauda L/SIPH L 0.57-0.78 0.77-0.96

Cauda L/URS L 1.43-1.77 2.12-2.39

Chaetotaxy

SN ANT III 3-4 4-5

SN (accessory) on URS 3-5 2

First tarsal formula hie ALG So Bat bs) 5°32 OF 32373

SN on ABD TERG VUHI 4-6 6-10

SN on discal part of subgenital plate 2-4 2-5

SN on hind margin of subgenital plate 7-9 11-16

SN on Cauda (3)4 7-11

and pointed accessory setae. Setae on body very pale;

setae on dorsum and ANT rarely blunt, mostly capitate

or fan-shaped. Ventral setae on abdomen pointed and

longer than dorsal ones located there. Dorsum of head,

thorax and abdomen sclerotized, wrinkled and reticu-

lated. Reticulation pattern comprising small cells with

irregular edges. SIPH rough, imbricated with spinulose

imbrication on apical halves, tapering or slightly swollen

in the apical third and with marked flange, its swollen

part 1.03—1.10 x the immediate proximal narrow part

(Figs 1, 5). ABD TERG VIII with low, rugose medial

supracaudal swelling with 2 setae on it. Subgenital plate

oval, sclerotic, with spinulose imbrications. 0-3 small

MTu present on ABD TERG II-V on one side (O44 in

total). Cauda elongate triangular or tongue—shaped with

laterally situated long, curved and pointed setae and nu-

merous strong dark spinules (Figs 1, 5). Measurements,

ratios and chaetotaxy are provided in Table 1.

Differential diagnosis

Lipaphis holmani sp. nov. belongs to the subgenus Li-

paphidiella Doncaster, 1954 because of (1) the weakly

developed antennal tubercles, (2) the presence of the me-

dial supracaudal process on ABD TERG VIII, and (3) the

Bonn zoological Bulletin 71 (2): 192—203

ratio of the ANT/BL which is less than 0.5 (Doncaster

1954; Heie 1992; Blackman & Eastop 2006). At pres-

ent, four species are known in the subgenus: L. jungarica

Kadyrbekov, Renxin & Shao, 2002, L. lepidii (Nevsky,

1929), L. ruderalis Borner, 1939 and L. sisymbrii Bozh-

ko, 1976 (Favret 2022).

Apterous viviparous 29 of Lipaphis holmani sp. nov.

are distinguished from the same morph of L. jungarica

by (1) the absence of secondary rhinaria on ANT III,

which are present in L. jungarica; (2) by the number of

accessory setae on URS: 4 (rarely 3 or 5) setae in the new

species, but only 2 in L. jungarica; (3) by the presence

of reticulation on dorsum of thorax and abdomen being

absent in L. jungarica; and (4) by the number of setae on

the posterior margin of the subgenital plate: 7-9 in the

new species, but only 2 setae in L. jungarica (see also

Kadyrbekov et al. 2002).

Apterous viviparous 29 of Lipaphis holmani sp. nov.

can be distinguished from the same morph of L. /epidii

based on Doncaster 1954; Blackman & Eastop 2022, in-

cluding investigated type material of L. /epidii, by (1) the

number of accessory setae on URS: 4 (rarely 3 or 5) setae

in the new species, but only 2 in L. /epidii; (2) the ratio

of the CAUDA L/ANT III: 0.79-0.99 in the new species,

©LIB

New species of Lipaphis Mordvilko, 1928 197

but 0.45-0.75 in both subspecies of L. lepidii (L. lepidii

lepidii (Nevsky, 1929) and L. lepidii cardariae Knech-

tel & Manolache, 1944).

Apterous viviparous 29 of Lipaphis holmani sp. nov.

are distinguished from the same morph of L. ruderalis

and L. sisymbrii by (1) the number of accessory setae on

URS: 4 (rarely 3 or 5) setae in the new species, while 2,

rarely 3 in L. ruderalis and L. sisymbrii; (2) the ratio of

the URS L/HT II L: 0.81—0.92 in the new species, but

0.60-0.70 in L. ruderalis and 0.62—0.82 in L. sisymbrii;

(3) the presence/absence of intersegmental sclerites on

abdomen: absent in the new species, but always present

in L. ruderalis and L. sisymbrii; (4) the shape of the me-

dial supracaudal process on ABD TERG VIII: a small

rugose medial swelling in the new species, but a large,

medial, conical-shaped supracaudal process in L. ruder-

alis and L. sisymbrii (Doncaster 1954; Heie 1992; Black-

man & Eastop 2022).

Apart from above mentioned Lipaphis species, two

species — L. erysimi and L. pseudobrassicae — utilize

Lepidium spp. as host plants (Holman 2009; Blackman &

Eastop 2022), from which the new species should be dis-

tinguished easily by the presence of the medial supra-

caudal process on ABD TERG VIII, which is absent in

L. erysimi and L. pseudobrassicae (Blackman & Eastop

2022). Only Lipaphis erysimi has been recorded previ-

ously from the Korean Peninsula (Lee et al. 2002).

Biology. The new species lives on Lepidium virgin-

icum L. (Brassicaceae). Other morphs and life cycle of

the new species are unknown.

Distribution. It is known only from the type locali-

ty—Yeogisan hill, Suwon, Gyeonggi Province, South

Korea, Korean peninsula. The new aphid species should

be widely distributed because it lives on a widely distrib-

uted invasive host plant, Virginia pepperweed (Lepidium

virginicum) (Stoyanov & Vladimirov 2015). This plant

species is native to North America. It was introduced to

Africa, Asia, Australia, Europe and South America and is

naturalized widely elsewhere (Zhou et al. 2001; Stoyan-

ov & Vladimirov 2015).

Lipaphis ruderalis Borner, 1939

Lipaphis ruderalis Borner, 1939: 79.

Figs 2, 4, 6, 7-9, Table 1

Material examined

N 4395, 3 oviparous 29 on | slide, on Lepidium rud-

erale L., CZECH REPUBLIC, Central Bohemian Re-

gion, Nymburk District, Lysa nad Labem, 50°12'5" N,

14°49'58" E, 183 ma.s.l., 09.v.2003, J. Holman leg. and

det. TECA) — N 4395, 3 oviparous 9° on 1 slide, same

collection data as for preceding (IZISU).

Bonn zoological Bulletin 71 (2): 192—203

Description

Oviparous &° (n= 6)

Color in life: Unknown. Pigmentation on slide: Head,

ANT I-II, IV—VI, clypeus, rostral IlI—V, tarsi, large mar-

ginal sclerites on thoracic I-III, intersegmental sclerites

on abdomen, SIPH, postsiphuncular sclerites, subgeni-

tal and anal plates, spinopleural bar on ABD TERG VI,

transverse bands on ABD TERG VII-VIII dark brown;

ANT III with paler basal and darker apical halves; cauda

pale brown or dark brown; coxae, trochanters and basal

part of femora pale brown with dark brown remaining

part; tibiae with dark brown bases and apices and pale

brown middle section; stigmal plates pale brown (Fig. 2).

Morphological characters: Body oval. Frons slightly

convex (Figs 2, 4). ANT 5 or 6-segmented (Figs 2, 4,

6). ANT tubercles weakly developed. ANT cuticle im-

bricated. Primary rhinaria ciliated. Secondary rhiniaria

always absent. Rostrum short, reaching to middle coxae.

URS sub-parallel sided with pointed apex and fine and

pointed accessory setae (Fig. 2). HTIB swollen (Fig. 7),

diameter of its swollen part 1.27—1.67 x its narrowest di-

ameter at base with 51—81 round pseudosensoria, mostly

single or in clusters of two or three. Setae on body dusky

on dusky sockets; setae on head, thorax and abdominal

dorsum and ANT capitate or fan—shaped. Ventral setae on

abdomen are pointed and longer than dorsal ones located

there. Dorsum of head, thorax and abdomen is sclerotic,

wrinkled and reticulated. Reticulation pattern comprising

small cells with irregular edges (Fig. 8). SIPH rough, im-

bricated with spinulose imbrication on apical 1/3, slight-

ly swollen in the apical third and with marked flange, its

swollen part 1.06—-1.20 x the immediate proximal nar-

row part (Figs 2, 9). ABD TERG VIII with high, medial

conical process with 2 setae on it. Subgenital plate oval,

sclerotic, with spinulose imbrications. 14 small MTu

present on ABD TERG IJ—V on one side (2-8 in total).

Cauda elongate triangular, not or slightly constricted in

the middle, with laterally situated long, curved and point-

ed and dorsally situated short, straight and point or slight-

ly curved and capitated setae and numerous strong dark

spinules (Figs 2, 9). Measurements, ratios and chaetotaxy

are provided in Table 1.

Differences between apterous viviparous and ovip-

arous °°. Differences were found in (1) the number of

setae on ABD TERG VIII: maximum 6 in apterae, while

6—10 in oviparae; (2) the number of setae on cauda: max-

imum 7 in apterae, while 7—11 in oviparae; (3) shape of

HTIB and (4) presence/absence of pseudosensoria: it is

none-swollen without pseudosensoria in apterae, while it

is swollen with pseudosensoria in oviparae.

Biology. The species lives on undersides of leaves,

stems and flower-stalks of Berteroa incana (L.) DC.,

Cardaria draba (L.) Desv., Lepidium spp. (L. apetalum

Willd., L. densiflorum Schrad., L. ruderale L., L. sativum

L., Lepidium sp.) and Sisymbrium altissimum L. (Brassi-

©LIB

198 Shalva Barjadze et al.

caceae) (Holman 2009; Blackman & Eastop 2022). It is

clear that this species is a monoecious holocyclic.

Distribution. It is distributed from Eastern Europe

to China: Germany, Poland, Czech Republic, Slovakia,

Hungary, Romania, Ukraine, Russian Federation, China

(Tao 1999; Holman 2009).

Lipaphis sisymbrii Bozhko, 1976

Lipaphis sisymbrii Bozhko 1976: 102.

Figs 10-16, Table 2

Material examined

N 1927, 1 apterous viviparous 9, on Sisymbrium poly-

morphum Murray) Roth, KAZAKHSTAN, West Kazakh-

stan region, Aksay town environments, 51° 9°52.28” N,

52°58’23.08” E, 57 ma.s.l., 27.v1.1990, R.Kh. Kadyrbe-

kov leg. IZAK)-—N 2772, 5 apterous viviparous 9 9 and

1 alate viviparous 2, on Sisymbrium polymorphum Mur-

ray) Roth, KAZAKHSTAN, East Kazakhstan region,

env. st. Kajnar, 23.v1.1978, N.E. Smailova leg. (ZAK)

—N 230a, 4 apterous viviparous 9 and 3 alate vivip-

arous 29, on Sisymbrium loeselii L., KAZAKHSTAN,

Almaty region, Karatal river, 40 km to North-West Ush-

tobe town, 19.v.2005, M.O. Aitzhanova leg. IZAK) —N

1456, 6 apterous viviparous 9°, on Sisymbrium loesel-

ii L., KAZAKHSTAN, Almaty region, Zhungar Alatau

mounting system, Zhabyk ridge, 30 km to east Koktuma

village, 6.vili.1989, R.Kh. Kadyrbekov leg. IZAK) — N

2077, 5 alate viviparous 9°, on Sisymbrium loeselii L.,

KAZAKHSTAN, Almaty region, Zhungar Alatau mount-

ing system, Kojandytau ridge, 25.v1.1991, R.Kh. Kadyr-

bekov leg. IZAK) — N 2295, 7 apterous viviparous 9 9

and 1 alate viviparous 2, on Sisymbrium polymorphum

(Murray) Roth, KAZAKHSTAN, Almaty region, Tentek

river, 4 km to West from Usharal town, 46°10’3.28” N,

Figs 10-13. Lipaphis (Lipaphidiella) sisymbrii Bozhko, 1976. Apterous viviparous 9. 10. General view. 11. Head, thorax, and

antenna. 12. Abdominal dorsum with reticulation pattern. 13. Posterior part of abdomen.

Bonn zoological Bulletin 71 (2): 192-203

©LIB

New species of Lipaphis Mordvilko, 1928 199

80°54’7.03” E, 384 ma.s.l., 29.v.2005, M.O. Aitzhanova

leg. (IZAK).

Redescription

Apterous viviparous 9 (n= 23)

Color in life: Yellowish or light green, with a slight

light bloom, on the dorsal side of the body there are par-

allel segmental light brown small spots, SIPH, cauda,

coxae and trochanters are light, sometimes dusky; legs,

except for tops of tibiae and tarsi, light. Apices of tibi-

ae and tarsi brown. Pigmentation on slide: Head pale or

brown. ANT I-II pale or dusky in some specimens; ANT

IH-IV pale, apex of ANT V and ANT VI brown. Clypeus,

rostral III—V, apices of tibiae and tarsi pale brown; sub-

genital and anal plates dusky; marginal spots on thorac-

ic I-III and small intersegmental sclerites - light brown;

SIPH and cauda are light, dusky in some specimens; stig-

mal plates light brown (Figs 10-13).

414)

Morphological characters: Body oval (Fig. 10). Frons

uniformly convex without ANT tubercles (Figs 10-11).

ANT 5 or 6-segmented (Fig. 11). ANT tubercles not

developed (Figs 10-11). ANT cuticle wrinkled. Prima-

ry rhinaria ciliated. Secondary rhinaria always absent.

Rostrum short and not reach the middle coxae; URS

sub-parallel-sided with pointed apical and accessory se-

tae (Fig. 10). Dorsum setae pale, short, obtuse. Cuticle of

head and genital plate sclerotized, squamous (Figs 11—

13). Thoracic I-III with small, sometimes larger, squa-

mous marginal sclerites (Fig. 11). The rest of body are

not sclerotized, with delicate cellularity. Reticulation

pattern comprising small cells with irregular edges. SIPH

imbricated, with slightly swollen in the apical third and

with marked flange, its swollen part 1.10—1.20 x the im-

mediate proximal narrow part (Figs 10, 13). ABD TERG

VIII with high medial conical-shaped supracaudal pro-

cess with 2 setae on it (Figs 10, 13). Subgenital plate

oval, sclerotized, squamous. 0-3 small MTu present on

Figs 14-16. Lipaphis (Lipaphidiella) sisymbrii Bozhko, 1976. Alate viviparous °. 14. General view. 15. Head, thorax, and anten-

nae. 16. Posterior part of abdomen.

Bonn zoological Bulletin 71 (2): 192—203

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200

Shalva Barjadze et al.

Table 2 (continued on next page). Metric and meristic characteristics of apterous and alate viviparous 9° 9 of Lipaphis (Lipaphi-

diella) sisymbrii Bozhko, 1976. Explanations about abbreviations used in the table are given in the Material and methods section.

Apterae

n=23

Alatae

n=10

Characters

ANT

ANT III

MSL ANT III

ANT IV

ANT V

ANT V borANT VI b

MSL on cephalic dorsum

URS L

Posterior setae on hind trochanter

HFEM

Dorsal MSL on femora

HTIB

Dorsal MSL in the middle of HTIB

12 ii

MSL ABD TERG II

MSL ABD TERG VIII

SIPH L

Cauda L

1.290-2.180

0.576—1.037

0.498-0.911

0.149-0.311

0.007

0.058-0.115

0.069-0.104

0.048—0.081

0.092—0.150

0.323—0.420

0.008—0.009

0.075—0.104

0.008—0.009

0.265—0.426

0.008—0.009

0.438—0.783

0.008—0.009

0.104—0.138

0.008—0.009

0.011—-0.012

0.186—0.334

0.161—0.230

1.580-1.958

0.576—0.898

1.012—1.602

0.334—0.495

0.007

0.144—0.265

0.115-0.219

0.092-0.127

0.255—0.380

0.369-0.403

0.008

0.081—0.115

0.008—0.009

0.436—0.622

0.008—0.009

0.818—1.140

0.008—0.009

0.127-0.161

0.008—0.009

0.011-0.012

0.149-0.230

0.127—-0.184

Ratios

ANT/BL

PT/ANT V b or VI b

PT/Ant V or VI

PT/ANT IV

PT/ANT II

PT/HW

PT/URS L

PT/Siph

ANT IV/ANT V or VI

ANT III/ANT V or VI

ANT III+PT/Siph

ANT III/URS L

ANT III/Siph L

MSL ANT III/ANT III BD

MSL on cephalic dorsum/ANT III BD

URS L/URS W

URS L/HW

URS L/H ILL

URS L/Siph L

Posterior setae of hind trochanter/suture

HFEM/BL

HFEM/HW

Dorsal MSL on femora/ANT HI BD

Dorsal MSL on femora/suture

HTIB/BL

HTIB/HW

Bonn zoological Bulletin 71 (2): 192—203

0.33—-0.46

1.40-2.51

0.58-0.72

1.25-2.46

0.40—0.85

0.21-0.44

1.23-2.13

0.33—-0.54

0.28-0.72

0.71—-1.56

0.84—2.07

1.84—3.55

0.54—0.96

0.18-0.19

0.33—-0.37

2.27—2.31

0.18—0.26

0.62—-0.82

0.24—0.33

0.36—0.37

0.16—0.23

0.71—1.03

0.35—0.38

0.20-0.21

0.28-0.41

1.18-1.86

0.67—0.83

2.61—-3.68

0.72-0.78

1.39-2.40

0.65—0.90

0.68—-0.94

2.61-4.12

1.29-2.09

0.32-0.52

0.80—1.23

3.46-4.48

3.10—5.37

1.95-2.67

0.18-0.19

0.33-0.37

2.19-2.25

0.21—-0.30

0.58—0.83

0.41-0.77

0.36—-0.37

0.24—-0.32

1.15-1.54

0.35—0.38

0.20-0.21

0.46-0.59

2.15—2.83

©LIB

New species of Lipaphis Mordvilko, 1928

Table 2. continued.

Dorsal MSL in the middle of HTIB/ANT III BD

Dorsal MSL in the middle of HTIB/Diameter of

HTIB at the middle section

H II L/HW

MSL ABD TERG III/ANT II BD

MSL ABD TERG VIII/ANT HI BD

Siph L/ Siph BW

Siph L/BL

Siph L/HW

Siph L/H II L

Siph L/Cauda L

Cauda L/ Cauda W

Cauda L/ANT III

Cauda L/HW

Cauda L/Siph L

Cauda L/URS L

Apterae

n= 23

0.35-0.37

0.21-0.22

0.27-0.39

0.37—0.38

0.45—0.50

4.17-5.97

0.15-0.19

0.66—0.85

2.11-—2.90

1.34-1.93

1.36—-1.77

0.69-1.11

0.44—0.59

0.59-0.74

2.10—2.84

0.35-0.37

0.21-0.22

0.37—0.38

0.45—0.50

2.56—3.96

0.09-0.12

0.39-0.57

1.08-1.54

1.00-1.54

1.18-1.45

0.37—-0.46

0.33—-0.48

0.80—1.00

1.10—-2.14

Chaetotaxy:

SN ANT III

SN (accessory) on URS

First tarsal formula

SN on ABD TERG VIII

SN on discal part of subgenital plate

SN on hind margin of subgenital plate

SN on Cauda

ABD TERG H-V on one side (2-6 total). Cauda elongat-

ed triangular or tongue-shaped with curved and pointed

setae, with numerous strong spinules (Figs 10, 13). Mea-

surements, ratios and chaetotaxy are given in Table 2.

Alate viviparous ° (n= 10)

Color in life: head, ANT, legs, SIPH, cauda, thorax,

marginal spots ABD TERG II-IV are brown. Abdomen

light green with a slight light bloom. Pigmentation on

Slide: head, thorax, ANT, except the paler base of ANT

III, clypeus, rostral III—V, legs, except pale bases of fem-

ora and middle of tibiae, SIPH, cauda, subgenital and

anal plates, marginal spots on ABD TERG II-V brown

(Figs 14-16).

Morphological characters. Body elongated-oval

(Fig. 14). Frons W-shaped with a median tubercle, on

which the ocellus is located, and only just marked ANT

tubercles (Figs 14-15). ANT six-segmented (Fig. 15).

ANT cuticle wrinkled; primary rhinaria ciliated. Second-

ary rhinaria are present on ANT III (9-36), IV (1-14),

and V (0-5) (Figs 14-15). Rostrum short, not reach the

middle coxae; URS sub-parallel-sided with pointed api-

cal and accessory setae. Dorsum setae pale, short, obtuse.

Cuticle of head, thorax, and genital plate sclerotized,

squamous. Abdomen not sclerotized, finely reticulated.

Reticulation pattern comprising small cells with irregular

Bonn zoological Bulletin 71 (2): 192—203

3-7

2-3

RB SO Sse

2-3

2-5

6-8

edges. SIPH imbricated, with slightly swollen in the api-

cal third and with marked flange, its swollen part 1.15—

1.20 x the immediate proximal narrow part (Fig. 16).

ABD TERG VIII bears high medial conical-shaped

subracaudal process, with 2 setae. Subgenital plate oval,

sclerotized, scabrous. 0-5 small MTu present, on ABD

TERG II-V on both sides (2-8 total). Cauda elongated

triangular with curved and pointed setae, with numerous

spinules (Fig. 16). Measurements, ratios and chaetotaxy

are given in Table 2.

Differential diagnosis. Lipaphis sisymbrii is morpho-

logically similar to L. ruderalis in shape of the high me-

dial conical-shaped supracaudal process on ABD TERG

VIII, the presence of intersegmental sclerites on the

abdomen, and the chaetotaxy of the URS (2 accessory

setae). It differs from L. ruderalis in shape of a faintly

convex frons without ANT tubercles, the fine and less

distinct reticulation of the light dorsal parts of the thorax,

and ABD, light SIPH and cauda, chaetotaxy of the ABD

TERG VIII (2 setae versus 4—6) and cauda (4 setae ver-

sus 5—7), ratios of the cauda to SIPH (0.59-0.74 versus

0.77-0.96). Besides, they use different host plants.

Biology. The species lives on Sisymbrium loeselii L.

and S. polymorphum (Murray) Roth (Brassicaceae).

Distribution. Ukraine and Kazakhstan (west, east,

south-east) (Bozhko 1976; Kadyrbekov 2009, 2017).

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202 Shalva Barjadze et al.

Taxonomic position of Lipaphis sisymbrii Bozhko,

1976

Lipaphis sisymbrii Bozhko, 1976 was described extreme-

ly briefly based on apterous and alate viviparous 9° on

Sisymbrium polymorphum (Murray) Roth (Brassicaceae)

from Ukraine (Bozhko 1976). In spite of the fact that in

the original description presence of the medial supra-

caudal process on ABD TERG VIII was mentioned (p.

102), it was considered as a member of the subgenus Li-

paphis in Blackman & Eastop (2006, 2022) and in Favret

(2022). Based on the presence of the medial supracaudal

process, mentioned in the original description and shown

in specimens from Kazakhstan (Figs 10, 13), this species

is transferred in the subgenus Lipaphidiella Doncaster,

1954. At present, five species are united in the subgenus

Lipaphidiella: L. jungarica Kadyrbekov, Renxin & Shao,

2002, L. lepidii (Nevsky, 1929), L. ruderalis Borner,

1939, Lipaphis sisymbrii Bozhko, 1976 and L. holmani

sp. nov. The identification key for these species is given

below.

Identification key to the species of the subgenus

Lipaphidiella Doncaster, 1954

1. Reticulation on dorsum of thorax and abdomen

absent. Secondary rhinaria on ANT III present.

Posterior margin of the subgenital plate with only

Ane L. jungarica Kadyrbekov, Renxin & Shao, 2002

Dorsum of thorax and abdomen reticulated.

Secondary rhinaria on ANT III absent. Posterior

margin of the subgenital plate with more than

CSAC E Forza ber Mrtetd dd Pea ena con Ae TE a0) 2

1. ABD TERG VIII with high medial conical-

shaped supracaudal process (Figs 2, 9, 10, 13).

Intersegmental sclerites on abdomen present (Figs 2,

TC BL) ore cars eres MP, 0 aN ER se s 3

ABD TERG VIII with medial supracaudal process as

low rugose swelling (Fig. 5). Intersegmental sclerites

on. abdomenabsent.Chige Wen eo 6. ce eee 4

3. Thoracic and ABD reticulation less distinct (Fig. 12).

Cauda L 0.59-0.74 x SIPH L with 4 setae .......0.......

bade cca Meas Ave cased L. sisymbrii Bozhko, 1976

— Thoracic and ABD reticulation more distinct (Fig. 8).

Cauda L 0.77-0.96 x SIPH L with 5-7 setae ...........

eae Teele nee Aes a L. ruderalis Borner, 1939

4. Cauda L 0.45-0.75 x ANT HI. URS with 2 accessory

Sela G tut arenas es L. lepidii (Nevsky, 1929)

— Cauda L 0.80-1.10 x ANT III. URS with 3-5

ACCESSORVESCIde. vy pen snews lies es.4! 5, L. holmani sp. nov.

Acknowledgments. We would like to thank AleS Bezdék (In-

stitute of Entomology, Ceské Budéjovice, Czech Republic) for

the loan of specimens, Andrey Stekolshchikov (Institute of Zo-

ology, St. Petersburg, Russia) for providing some data of the

type material of Liphapis lepidii and Colin Favret (Université

Bonn zoological Bulletin 71 (2): 192—203

de Montréal, Montréal, Canada) for providing PDF versions

of the publications on the genus Lipaphis. We are grateful to

Susan Halbert (Florida Department of Agriculture and Con-

sumer Services, Division of Plant Industry, Gainesville, USA)

and anonymous referees for invaluable information and critical

review of the manuscript. The first author’s visit in Czech Re-

public was supported by the Ministry of Education, Science,

Culture and Sport of Georgia as a part of the grant: “Animal

Diversity in Natural and Agroecosystems of Georgia, Monitor-

ing and Biocontrol”. First author’s research was supported by

the project: “Investigation of the Georgian aphids (Hemiptera:

Aphidoidea) using modern approaches” (SRSFG — R-21-1509)

funded by the Shota Rustaveli National Science Foundation of

Georgia. Third author did his part of the work thanks to the

grant project of the Science Committee of the Ministry of Edu-

cation and Science of the Republic of Kazakhstan (subject No.

OR11465437-OT-21).

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Blackman RL, Eastop VF (2000) Aphids on the World’s Crops:

An Identification and Information Guide. Second Edition.

John Wiley & Sons, Chichester

Blackman RL, Eastop VF (2006) Aphids on the world’s herba-

ceous plants and shrubs. John Wiley & Sons, London

Blackman RL (2010) Aphids — Aphidinae (Macrosiphini).

Handbook for identification of British insects. Vol. 2. Part 7.

Royal Entomological Society, London

Blackman RL, Eastop VF (2022) Aphids of the World’s Plants:

An Online Identification and Information Guide. Online

at http://www.aphidsonworldsplants.info (accessed 2 Jan.

2022)

Borner C (1939) Neue Gattungen und Arten der mitteleu-

ropadischen Aphidenfauna. Arbeiten tiber physiologische und

angewandte Entomologie aus Berlin-Dahlem 6 (1): 75—83

Bozhko MP (1976) Aphids of forage plants. Vishcha shkola,

Kharkiv [in Russian]

Das BC (1915) Beitrage zur Kenntnis der Hollandischen

Blattlause. Eine Morphologisch-Systematische Studie.

Friedlander, Berlin

Davis JJ (1914) New or little known species of Aphididae. The

Canadian Entomologist 46 (7): 226-236

Doncaster JP (1954) Notes on the genus Lipaphis Mordvilko,

1928 (Homoptera: Aphididae) and description of a new spe-

cies. Proceedings of the Royal Entomological Society of

London, Series B, Taxonomy 23 (5-6): 83-88

Favret C (2022) Aphid Species File. Version 5.0/5.0. Online at

http://Aphid. SpeciesFile.org (accessed 2 Jan. 2022)

Heie OE (1992) The Aphidoidea (Hemiptera) of Fennoscandia

and Denmark. IV. Family Aphididae: Part I of tribe Macro-

siphini and subfamily Aphidinae. Fauna Entomologica Scan-

dinavica 25: 1-188

Holman J (2009) Host plant catalog of aphids. Pa-

laearctic region. Springer, Branisovska.

https://doi.org/10.1007/978-1-4020-8286-3

Kadyrbekov RKh, Renxin H, Shao H (2002) To aphid fauna

(Homoptera, Aphididae) of Xinjiang-Uygur region of Chi-

na. Tethys Entomological Research 6: 13-32

Kadyrbekov RH (2009) To the fauna of aphids (Homoptera,

Aphidinea) of the Saur ridge. Trudy Instituta zoologii MES

RK 50: 79-91 [in Russian]

Kadyrbekov RH (2017) Aphids (Homoptera: Aphidoidea, Phyl-

loxeroidea) of Kazakhstan (Check list). LLP 378, Almaty [in

Russian]

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lause (Phytophthires). In Commission der Roschiitz’schen

Buchandlung, Aachen

Knechtel WK, Manolache CI (1944) Neue Blattlause fir

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(Cruciferae). Pp. 1-193 in: Wu ZG & Raven PH (eds) Flora

of China (Brassicaceae through Saxifragaceae). Vol. 8. Sci-

ence Press, Beijing

©LIB

LIB. Leibniz Institute for the Analysis

== of Biodiversity Change

Bonn zoological Bulletin 71 (2): 204-208

2022 Mabrouk1 Y. et al.

museum

: KOENIG

https://do1.org/10.20363/BZB-2022.71.2.204

ISSN 2190-7307

http://www.zoologicalbulletin.de

Research article

urn:|sid:zoobank.org:pub: AC9E7EF2-8443-4A57-9A 7D-FCS5SBDBAO0B97

Mahrazia benlemithi gen. et sp. nov., a new subterranean snail

(Gastropoda: Hydrobiidae) from Morocco

'Université Sidi Mohamed Ben Abdellah, Faculté des Sciences de Dhar El Mahraz, Laboratoire de Biotechnologie,

Conservation et Valorisation des Ressources Naturelles, Fes, Morocco

2 Université Mohammed Premier, Faculté Pluridisciplinaire de Nador, Equipe de Recherche en Biologie et

Biotechnologie Appliquées, Morocco

3 Schulstr, 3, D-25491 Hetlingen, Germany

“Corresponding author: Email: younes_mab@hotmail.fr

'urn:|lsid:zoobank. org:author:9FE7D 1 12-2607-4855-A BF7-9BFOBA38B8C8

2urn:Isid:zoobank.org:author:358C248 1-FAD9-4524-AEFC-E1 8008EE04AC

3urn:Isid:zoobank.org:author: 8CB6BA7C-D04E-4586-BA I D-72FAFF54C4C9

Abstract. The species richness of the Hydrobiidae in Morocco is one of the highest in North Africa. Mahrazia benlemlihi

gen. et sp. nov. is a new stygobiont hydrobiid gastropod from the Saiss aquifer, north-central Morocco. It can be distin-

guished by the morphology of the shell and anatomical details.

Key words. Minute snails; Saiss aquifer; stygobiont snail; new genus; groundwater.

INTRODUCTION

Freshwater gastropods often dominate superficial aquat-

ic ecosystems in both biomass and numbers, where they

graze on periphytic or epiphytic algae and biofilms. They

constitute an important dietary component for several

predatory animals. Freshwater snails can also be found

in water bodies fed by karstic aquifers; some species are

adapted to live in caves and subterranean habitats (Ke-

bap¢i 2013; Gloer 2019); these are called stygobionts.

The aquatic gastropod family Hydrobiidae Stimpson,

1865 occurs mainly in the western Palaearctic and the

Nearctic (Lydeard & Cummings 2019). The family sen-

su lato comprises more than 400 extant genera (Kabat &

Hershler 1993), many of which are stygobionts.

In Morocco, only four stygobiont species of hydrobiids

are known to have depigmented bodies and to lack eye

spots, these are: Atebbania bernasconi Ghamizi, Bodon,

Boulal & Giusti, 1999, Heideella andreae Backhyus &

Boeters, 1974, Heideella makhfamensis Bodon, Ghami-

zi & Giusti, 1999 and Rifia yacoubii Ghamizi, 2020. In

addition, there are two stygobiont species of Moitessierii-

dae: Iglica seyadi Backhyus & Boeters, 1974 and Iglica

soussensis Ghamizi & Boulal, 2017 (Ghamizi 2020).

New research conducted recently in north-central Mo-

rocco revealed a new stygobiont genus of the family Hy-

drobiidae.

Received: 06.09.2022

Accepted: 02.11.2022

MATERIAL AND METHODS

Field surveys were conducted since 2014 (still ongoing),

in which several localities were prospected in the north-

ern part of Morocco. Stygobiont molluscs were collected

from wells by means of a phreatobiological net, through

filtering subterranean water by passing it through a ny-

lon net and sieving mud and sediment. The samples were

fixed in 75% ethanol. The dissections and measurements

of the genital organs and the shells were carried out using

a stereo microscope (Leica M205C) with a digital camera

(Leica DMC5400). The type material is stored partly in

the Zoological Museum Hamburg (ZMH), partly in the

collection of the first author.

RESULTS

Phylum Mollusca Cuvier, 1795

Class Gastropoda Cuvier, 1795

Superorder Caenogastropoda Cox, 1960

Superfamily Truncatelloidea Gray, 1840

Family Hydrobiidae Stimpson, 1865

Mahrazia gen. nov.

urn:lsid:zoobank.org:act:38D2A566-9C D0-4D71-B4F 8-648 762506E0E

Type species

Mahrazia benlemiihi gen. et sp. nov.

Corresponding editor: B. Huber

Published: 04.11.2022

Mahrazia benlemiihi gen. et sp. nov., a new subterranean snail from Morocco 205

Table 1. Shell measurements of Mahrazia benlemiihi gen. et

sp. nov. (in mm; N=9; measurement accuracy: 0.05).

Shell measures Min Max Mean — sd

Shell height x15 3.35 325 0.10

Aperture height 1.00 1.10 1.05 0.05

Spire height 2.10 2.30 2.20 0.10

Shell width liege 1.75 1.65 0.10

Aperture width 0.90 1.00 0.95 0.05

Type material

Holotype

ZMH 141428; from a well located at the Faculty of

Sciences Dhar El Mahraz, University Sidi Mohamed

Ben Abdellah, Fez-Meknes region; 34°01'59.16" N,

4°58'37.86" W; collected 9 Apr. 2022.

Paratypes

3 spec.; ZMH 141429; same collection data as for holo-

type. — 6 spec.; Y. Mabrouki leg.; collected 10 Oct. 2022;

same locality as for holotype.

; : we Ae Figs 1-5. Mahrazia benlemilihi gen. et sp. nov. 1. Holotype

Diagnosis. The new genus can be distinguished from (ZMH 141428). 2. Holotype, penis. 3-5. Paratypes (3 dried).

the other Moroccan stygobiont hydrobiids by its elon-

Fig. 6. Habitus of live specimens of Mahrazia benlemilihi gen. et sp. nov.

206 Youness Mabrouki et al.

6°0'0"W

36°0'0"N

33°0'0"N

3°0'0"W

editerranean

[rm

* Faculty of Sciences Dhar El Mahraz

Groundwater of Fez-Meknes

~ €3 Sebou River basin

Fig. 7. Location of the type locality of Mahrazia benlemilihi gen. et sp. nov.

gated ovate shell with less than 5 whorls, its long pe-

nis, thickened at the basis with a bulbous part near the

pointed penis tip. Heideella Backhuys & Boeters, 1974

has a slim, elongated conical shell with more than 5 flat

whorls, a conical penis that is more or less elongated,

wrinkled near its base, with one lobe, more or less devel-

oped and bulging. Atebbania Ghamizi, Bodon, Boulal &

Giusti 1999 has an elongated conical shell with more

than 5 whorls, with the penultimate whorl slightly broad-

er than the body whorl, a penis with an apical stylet, one

lobe bent downwards on its left side, and the penial duct

running near the right side. Rifia Ghamizi, 2020 has a

valvatoid shell and a bifurcated penis. The other genera

are surface water molluscs and have eyespots and pig-

mented bodies.

Description. See description of single known species

below.

Etymology. The genus name ‘Mahrazia’ refers to its

type locality: the Faculty of Sciences Dhar El Mahraz

FSDM, located on the university campus Dhar El Mah-

raz and covering an area of 6 ha. It mainly serves the

region of Fez-Boulemane located in the northern center

of Morocco.

Mahrazia benlemilihi gen. et sp. nov.

urn. lsid:zoobank.org: act: 73B222F F-B0D7-4143-BC10-6B3518DBEFB

Figs 1-6

Bonn zoological Bulletin 71 (2): 204—208

Etymology. The species was named after Professor Mo-

hammed Benlemlih (the Dean of the Faculty of Sciences,

Dhar El Mahraz), in recognition of his encouragement

and support for our research work and for his help in

finding the new species.

Description

Shell. Minute, thin-walled, elongated ovate, glossy

and transparent when fresh, spire of 4.5 whorls growing

regularly, and separated by a deep suture. Shell surface

smooth, with growth lines that are hardly visible. Apex

broad and flat. Aperture ovate with a sharp peristome.

Umbilicus closed. Outer lip slightly sinuated in later-

al view. The shell measurements (N=9) are presented

in Table 1. Since the number of individuals studied is

quite small, these morphometric characteristics need to

be completed by further catches of the new subterranean

snail.

Operculum. The corneous operculum is smooth and

thin, and brown in colour.

Soft parts morphology and anatomy. Body translu-

cent when alive, white when preserved, pigment-less;

tentacles very long and thin, without eyes. Penis long,

thickened at basis with a bulbous part near pointed penis

tip, four times longer than wide with folds at the basis.

Distribution. This new genus and species 1s known

only from the type locality, which is a well located in the

Faculty of Sciences Dhar El Mahraz (Fig. 7). It belongs

©LIB

Mahrazia benlemiihi gen. et sp. nov., a new subterranean snail from Morocco 207

to the Saiss aquifer system, one of the main aquifer sys-

tems of Morocco, limited to the North by the Pre-Rif, to

the East by the valley of the Oued Sebou, to the West by

the tributaries of the Oued Beht and to the South by the

Middle Atlas. It is formed by the deep Lias aquifer and

the Quaternary groundwater. The water level is on aver-

age at a depth of 50 m below the ground level.

DISCUSSION

The Moroccan freshwater ecosystem is among the most

diverse within the northern African region, where a di-

versified molluscan fauna appears to live in the aquifers,

in the karstic underground (Van Damme et al. 2010).

In addition to the historical records, many new species

and genera have been recently discovered in this coun-

try (Gloér et al. 2020a, 2020b; Boulaassafeer et al. 2021;

Mabrouki et al. 2020, 2021a, 2021b, 2022a, 2022b; Taybi

et al. 2021; 2022a, 2022b, 2022c). While there has been

a significant advancement of knowledge on the surface

water malacofauna of Morocco, gaps remain and knowl-

edge on the occurrence of freshwater species from sever-

al aquifers is still poor, with some entirely ‘blank areas’.

Five species of stygobiont Hydrobiidae sensu stricto

are currently recognized, including Mahrazia benlemli-

hi gen. et sp. nov. For the moment, the new genus and

species is only known from the Sais aquifer. The stygobi-

ont valvatoid Rifia yacoubii is widespread in the phreatic

waters in the southern border of the Rif region, in the

upstream of the Moulouya, Sebou and Loukkos basins.

Atebbania bernasconii is restricted to the subterranean

waters of the Tiznit plain (southern Morocco). Heideella

andreae is distributed in southern-central Morocco and

the Marrakech vicinity, and H. makhfamanensis is en-

demic to the hyporheic waters of Oued Makhfamane in

the centre of the Haouz plain, in central Morocco (Gham-

izi 2020; Gloer 2022).

Pollution, habitat loss, and an ever-drier climate seem

to be the main factors responsible for the decline of

freshwater molluscs in North Africa (Taybi et al. 2017).

Numerous freshwater wells are presently at risk of sali-

nization (Van Damme et al. 2010; Khalloufi et al. 2017),

threatening the stygobiont fauna which they shelter,

including molluscs. Further intensified studies and col-

lecting expeditions, and description of new taxa are an

urgent need.

Acknowledgements. We are grateful to the editor and three

anonymous reviewers for valuable corrections and comments.

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Boulaassafer K, Ghamizi M, Machordom A, Albrecht C, Delica-

do D (2021) Hidden species diversity of Corrosella Boeters,

1970 (Caenogastropoda: Truncatelloidea) in the Moroccan

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420. https://doi.org/10.1007/s13127-021-00490-3152.

Ghamizi M (2020) New stygobiont genus and new species

(Gastropoda, Hydrobiidae) from the Rif (Morocco). Ecolog-

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Gloer P (2019) The freshwater gastropods of the West-Pa-

laearctis, Volume I Fresh- and brackish waters except spring

and subterranean snails. Identification key, anatomy, ecology,

distribution. Hetlingen, Germany

Gloer P (2022) The freshwater gastropods of the West-Pa-

laearctis, Volume 3 Hydrobiidae, Identification key, anatomy,

ecology, distribution. Hetlingen, Germany

Gloer P, Mabrouki, Y Taybi AF (2020a) A new ge-

nus and two new = species (Gastropoda, Hydrobii-

dae) from Morocco. Ecologica Montenegrina 28: 1-6.

https://do.org/10.37828/em.2020.28.1

Gloer P, Mabrouki Y, Taybi AF (2020b) Two new val-

vatoid genera (Gastropoda, NHydrobiidae) from Mo-

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https://doi.org/10.37828/em.2020.30.12

Kabat AR, Hersler R (1993) The Prosobranch family Hydrobu-

dae (Gastropoda: Rissooidea): Review of classification and

supraspecific taxa. Smithsonian Contributions to Zoology

547: 1-94

Kebapci U (2013) Cave and subterranean snails of Turkey.

Muzeul Olteniei Craiova, Oltenia, Studii $i comunicari, Sti-

intele Naturii 2 9 (2): 127-130

Khalloufi N, Béjaoul M, Delicado D (2017) A new genus and

Species of uncertain phylogenetic position within the family

Hydrobiidae (Caenogastropoda, Truncatelloidea) discovered

in Tunisian springs. European Journal of Taxonomy 328:

1-15

Lydeard C, Cummings KS (2019) Freshwater Mollusks of the

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gastropod fauna (Gastropoda: Hydrobiidae, Lymnaei-

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Mabrouki Y, Taybi AF, Gléer P (2021a) Two new species of

the genera Js/amia and Mercuria (Gastropoda, Hydrobii-

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Mabrouki Y, Taybi AF, Gléer P (2021b) Further records of

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Van Damme D, Ghamizi M, Soliman G, MclIvor A, Seddon

M.B (2010) The status and distribution of freshwater mol-

luscs. Pp. 30—50 in: Garcia N, Cuttelod A & Abdul Malak D

(eds) The status and distribution of freswater biodiversity

in Northern Africa. IUCN, Gland, Switzerland; Cambridge,

UK; Malaga, Spain

©LIB

LIB. Leibniz Institute for the Analysis

=— of Biodiversity Change

Bonn zoological Bulletin 71 (2): 209-214

2022 Casiraghi A. et al.

museum

: KOENIG

ISSN 2190-7307

http://www.zoologicalbulletin.de

https://do1.org/10.20363/BZB-2022.71.2.209

Research article

urn:|sid:zoobank.org: pub: BBBC7AFF-7D47-4047-A BFE-5S5FD2ECB8DA3

The presence of Prociphilus (Prociphilus) bumeliae (Schrank, 1801)

(Aphididae: Eriosomatinae: Pemphigini) in the Iberian Peninsula

‘Instituto Valenciano de Investigaciones Agrarias. (IVIA). Unidad de Entomologia, Centro de Proteccién Vegetal y

Biotecnologia. Ctra. Moncada-Naquera Km. 4,5. E-46113 Moncada, Valencia, Spain

? Asociacion “Fotografia y Biodiversidad”, Torrejon de Ardoz. 28850 Madrid, Spain

> Departamento de Biodiversidad y Gestién Ambiental, area de Zoologia. Universidad de Leon. 24071 Leon Spain

* Departamento de Artroépodos, Museo de Ciencias Naturales de Barcelona, 08003, Barcelona, Spain

“Corresponding author: Email: casiraghi_alia@gva.es

'urn:lsid:zoobank.org:author:905837A0-9C77-4D57-A 895-B19DD9FF790B

2urn:Isid:zoobank.org:author:5 173B589-EBF0-44B4-A774-94049F905F49

3urn:Isid:zoobank.org:author: DOFA98CF-CA4D-4927-8FE5-EC4EA C9BFA30

*urn:Isid:zoobank.org:author:C134E648-51C1-48A2-8676-2E9517657FAE

Abstract. Prociphilus (Prociphilus) bumeliae (Schrank, 1801) was recorded for the first time in the Iberian Peninsula,

while developing colonies on the branches of Fraxinus angustifolia Vahl, attended by Lasius niger (Linnaeus, 1758) and

a Tapinoma ant of nigerrimum group (Nylander, 1856). Relationships with these two ant species are new records. The

identification was carried out using morphological, biological, and molecular characteristics.

Key words. Aphids, ash, Pemphigini, Barcoding, ant-aphid relationships, COL

INTRODUCTION

The genus Prociphilus Koch, 1857 groups over 50 spe-

cies (Blackman & Eastop 2022; Favret 2022), which are

characterized by being large aphids, usually without cor-

nicles, covered in abundant filamentous wax produced

by well-developed waxy glands also present in the fun-

datrices (Blackman & Eastop 2022).

In this genus, we can find species that have apparent-

ly recently moved their sexual phase from Populus spe-

cies to other hosts such as Rosaceae, Caprifoliaceae and

Oleaceae, as it happens in most Pemphigini, but they still

maintain different conifers as secondary hosts, in which

they can develop parthenogenetic populations (Zak 1965;

Purkart et al. 2019).

Most species of genus Prociphilus are characterized by

dioecious and holocyclic cycles, with a primary (or win-

ter) host on which the fundatrix and winged fundatrige-

niae develop, and a secondary (or summer) host in which

winged virginogeniae develop and from which the sex-

uparae will leave to give birth on their primary hosts to

the sexual morphs that, after mating, will produce over-

wintering eggs (Blackman & Eastop 2022). However,

many species, like Prociphilus tessellatus (Fitch, 1851),

have the capacity to develop anholocyclic populations

despite living in areas where their primary hosts are pres-

ent (Zak 1965; Purkart et al. 2019). The genus’ complex

Received: 24.02.2022

Accepted: 09.11.2022

biological cycles and the polyphenism of its morphs have

caused many species to be little known or their descrip-

tions to be poor. Currently, their cycles (and all their

morphs) are only fully known in three species (Blackman

& Eastop 2022) and at a molecular level there are only

COI sequences of ten species, making it a genus in need

of a deep revision.

Prociphilus species are mainly distributed in the North-

ern Hemisphere (20 in North America, 7 in Europe, 3 in

Central Asia and 15 in East Asia), basically restricted to

the areas where their primary hosts develop, with the ex-

ception of the Asiatic Prociphilus (P.) osmanthae Essig

& Kuwana, 1918 that has recently been introduced in

Hungary (Ripka et al. 2020) and Prociphilus (Meliarrhiz-

ophagus) fraxinifolii (Riley, 1879), a species of Nearctic

origin (Smith & Parron 1978; Maw et al. 2000), which

has been expanding its distribution range due to the or-

namental use of its hosts, and is now present practical-

ly throughout the Palearctic region, from Great Britain

and Iberian Peninsula to China (Remaudiere & Ripka

2003; Bienkowski & Orlova-Bienkowskaja 2018), Mex-

ico (Torres-Acosta & Sanchez-Pefia 2015), South Amer-

ica (Chile) (Carrillo 1977) and South Africa (Muller &

Scholl 1958).

Four species were known to date in the Iberian Penin-

sula: Prociphilus (Prociphilus) fraxini (Fabricius, 1777),

Prociphilus (Prociphilus) oleae (Leach, 1826) and Pro-

Corresponding editor: R. Peters

Published: 16.11.2022

210

ciphilus (Stagona) pini (Burmeister 1835) from the re-

view carried out for the Iberian Fauna project by Pérez

Hidalgo & Nieto Nafria (2003) and Prociphilus (Meliar-

hizophagus) fraxinifolii that was detected in 2011 (Pérez

Hidalgo & Mier Durante 2012).

A colony of winged viviparous females collected on

branches of Fraxinus angustifolia Vahl in Madrid and

other colonies found on the same plant in the cities of

Burgos and Leon allows us to mention, for the first time

for the Peninsula, a fifth species, Prociphilus bumeliae

(Schrank 1801).

MATERIAL AND METHODS

Initially we studied several dozens of winged viviparous

females and nymphs recorded on 10" June 2018 from the

locality of San Agustin de Guadalix in the Province of

Madrid, Spain (40.788270 N, 3.705692 W). The colony

developed on branches of F? angustifolia (reference sam-

ple 2137) and was clearly attended by ants.

At first, the colony was identified using the identifica-

tion keys of Blackman & Eastop (2022) and Heie (1980).

The identification was confirmed by COI molecular anal-

ysis with the methodology exposed in Casiraghi et al.

(2019).

During the spring of 2021, several Prociphilus col-

onies were also recorded on Fraxinus angustifolia, in

several green areas of the cities of Leon (42.591725 N,

5.559641 W, 8" May 2021, reference sample 4017) and

Burgos (42.358805 N, 3.681891 W, 15 May 2021, refer-

ence sample 4066), attended in both locations by Lasius

niger (Linnaeus, 1758).

All associated ants were identified by Xavier Espadal-

er (University of Barcelona, Spain). Both aphid and ant

specimens were deposited in the aphidological collection

of the Institute for Integrative Systems Biology (Univer-

sity of Valencia-CSIC).

Futhermore, to investigate phylogenetic relationships

of Prociphilus species, COI sequences identified as Pro-

ciphilus and Pachypappa warshavensis (outgroup) were

downloaded from the NCBI nucleotide database (Nu-

cleotide 2021) on November 14", 2021 [queries: proc-

iphilus AND “aphids” [porgn] AND (COI OR CO] OR

COX1) NOT COIL; “Pachypappa warshavensis” AND

“aphids” [porgn] AND (COI OR COI OR COX1) NOT

COI].

After manually reviewing the fasta files, 106 sequenc-

es were obtained [1 problem sequence, | outgroup se-

quence, 103 template sequences]. Multiple sequence

alignment was conducted with muscle ver. 3.8 (Edgar

2004).

Phylogeny was inferred using a Neighbour-Joining al-

gorithm (Saitou & Nei 1987) with the F81 model (Fel-

senstein 1981) using the packages adegenet ver. 2.1.4

(Jombart 2008), ape ver. 5.5 (Paradis & Schliep 2018)

Bonn zoological Bulletin 71 (2): 209-214

Alice Casiraghi et al.

and phangorn ver. 2.7.1 (Schliep 2011) under R ver. 4.1.2

(R Core Team 2021). Pachypappa warshavensis (acces-

sion number MK936318.1) was used as outgroup, and

node support was computed with 1000 bootstrap rep-

licates. The tree was represented with ggtree package

ver. 1.16.6 (Yu et al. 2018).

According to the obtained phylogeny, the problem se-

quence was identified as Prociphilus bumeliae (Schrank,

1801) (Fig. 2).

RESULTS

Morphological characters and COI sequence (Fig. 2)

confirm without doubt that the aphid colony on F: angus-

tifolia was composed of P. bumeliae. Identification keys

from Blackman & Eastop (2022) and Heie (1980) allow

an easy distinction between P. bumeliae and P. fraxini,

basically due to the presence of depigmented wax plates

on the head of P. bumeliae (Fig. 1G). In P. fraxini these

plates are badly defined (Fig. 1H).

Morphology

Heie (1980) gives a good description of the viviparous

females of this species (fundatrix, winged fundatrige-

niae, apterous virginogeniae and sexuparae), which we

complement here with data on the only forms currently

known in the Iberian Peninsula.

Winged fundatrigeniae (based on measurements of

10 specimens) measure 3.8—5.5 mm. In life, they have a

reddish-brown coloration and are covered with abundant

filamentous wax (Fig. 1B, IF) that varies depending on

the body distribution of their wax plates and their stage

of development (Fig. 2C). In microscopic preparation,

head and thorax are blackish brown, with lighter areas

that correspond to the plates of the wax glands. The head

presents a well-developed posterior pair of waxy gland

plates (Fig. 1G) and, in very rare cases, two other small

ones between the antennae in the anterior part of the

head. Antennae measure 1.2—2.2 mm; they are formed by

6 segments and are 0.40—0.60 x the body. The antennal

segment HI measures 6.0—7.3 x the antennal segment II,

and from 1.07—1.23 the sum of the antennal segments IV

and V; the IV"—VI" antennal segments measure 0.31—

0.38, 0.29-0.37 and 0.36—0.42 mm, respectively, and the

terminal process 0.15—0.20 = the base of the VI" antennal

segment. The antennal segments III-IV present 16—24

and 2-9, respectively, secondary sensoria, with a trans-

verse oval shape, while the VI" lacks them. The rostrum

measures 0.20—0.32 x the body and its apical segment

carries about 6 complementary setae, and it is 3.25—4.25

x as long as its basal width and (0.6) 0.9-1.04 x the sec-

ond segment of the hind tarsi. This morph lacks siphun-

culi and the cauda is rounded and carries 7—10 setae and

in the first tarsal segment they usually have 4 setae (ex-

ceptionally 3).

©LIB

The presence of Prociphilus (Prociphilus) bumeliae (Schrank, 1801) in the Iberian Peninsula 211

— ll

Fig. 1. Prociphilus (P.) bumeliae (Schrank, 1801). A. leaves nest produced by the fundatrix and the winged fundatrigeniae in

Fraxinus angustifolia Vah|. B—D. Colonies on branches with different degrees of detail. E. Same colony attended by an ant of the

Tapinoma nigerrimum group. F. Aspect of the winged fundatrigeniae. G. Detail of its head (arrows indicate waxy plates). H. Pro-

ciphilus (Prociphilus) fraxini (Fabricius, 1777). Detail of the head of the winged fundatrigeniae (arrows indicate waxy plates).

212 Alice Casiraghi et al.

0 P. xylostei

P. kuwanai

; P. fraxinifolii

P. erigeronenis

P. americanus

1° P. bumeliae

P. tesselatus

P. caryae 9.01

— Pachypappa warshavensis

Fig. 2. Phylogenetic tree with studied samples, based on mitochondrial marker COI, obtained with NJ method (K2P), bootstraps:

1000. Sample indicated in bold red corresponds to the record of the present study. Abbreviation: P. = genus of Prociphilus Koch

1857.

The presence of Prociphilus (Prociphilus) bumeliae (Schrank, 1801) in the Iberian Peninsula

Biology

Prociphilus (Prociphilus) bumeliae presents a dioecious

holocyclic cycle, with species of Fraxinus (usually Frax-

inus excelsior L., and exceptionally Fraxinus ornus

L., Fraxinus angustifolia subsp. oxycarpa (Willd.) and

Fraxinus pallisae Wilmott), Ligustrum vulgare L. and

Syringa vulgaris L. as primary hosts, and roots of Abies

(usually Abies alba Mill. and, optionally, Abies balsamea

(L.) Mill. and Abies veitchii Lindley) as secondary hosts

(Holman 2009). Fundatrices make leaf curls resembling

bird nests in early spring (Fig. 1A), but as colonies de-

velop, they can move to trunk bark and thin branches

(Fig. 1B). Winged females migrate to the roots of their

secondary hosts where they form colonies covered with

abundant waxy secretion and from which the winged

sexuparae will emerge in late August-September to re-

turn to their primary host (Mordvilko 1935; Blackman &

Eastop 2022).

Associations between P. bumeliae and the ants 7° cf.

nigerrimum and L. niger are new aphid-ant relationship

for this species.

Distribution

Prociphilus bumeliae is a widely recorded species

throughout Europe (Holman 2009; Nieto Nafria et al.

2013), but there are some citations from East Asia that

need confirmation because they could be other species

(see Blackman & Eastop 2022). As for now, these re-

cords from the province of Madrid and Castilla y Léon

are the first of P. bumeliae in the Iberian Peninsula, but

it is likely to be distributed in other parts, where its host

plants are present both natural and ornamentally.

DISCUSSION

Taxonomic-bionomic problems

Prociphilus is a genus of aphids that presents complex

taxonomic and biological problems (Smith & Stroyan

1972), specifically due to the complexity of their life cy-

cles and their ability to develop paracycles in their sec-

ondary hosts, as it happens in other groups of Pemphigini

and Eriosomatini. However, thanks to the use of molec-

ular tools combined with traditional taxonomy, some of

the abovementioned problems can be solved (Pike et al.

2012).

As it happens with various Eriosomatinae, the identi-

fication of P. bumeliae presents some problems: P. (Pro-

ciphilus) fraxini is also widely distributed in Europe and

very similar in aspect and size to P. bumeliae (MatoSevi¢

2004; Blackman & Eastop 2022). This species also al-

ternates between F: excelsior and A. alba and its apter-

ous viviparous females (exules) are practically indistin-

guishable from those of P. bumeliae (Blackman & Eastop

2022). The two species can only be well differentiated

thanks to the winged stage corresponding to winged fun-

Bonn zoological Bulletin 71 (2): 209-214

213

datrigeniae developing inside the leaf nest, which we

used to determine the new record (Dransfield & Bright-

well 2021).

Faunistic studies

With our record, there are five Prociphilus species re-

corded in the Iberian Peninsula (P. fraxini, P. bumeliae,

P. oleae, P. (S.) pini and P. (M.) fraxinifolii (Pérez. Hidal-

go & Nieto Nafria 2003; Pérez Hidalgo & Mier Durante

2012). With the exception of P. fraxinifolii, which is be-

ginning to be widely distributed, due to the ornamental

use of Fraxinus pennsylvanica Marshall, the rest of sam-

ples are due to few specimens captured on their hosts or

with traps.

The distribution of P. fraxini in the Iberian Peninsu-

la corresponds to the natural area of its secondary host

A. alba (Pérez Hidalgo & Nieto Nafria 2003), but the

presence of P. bumeliae on F: angustifolia requires fur-

ther studies to confirm whether the ranges of both species

are limited to that of their hosts or may be broader.

It is important to consider that F’ excelsior is scarce in

the Sierra de Guadarrama, however F) angustifolia is a

very common species in this area, so it is to be supposed

that when it develops also on this species it is more wide-

spread, being able to carry out its root cycle in the firs

scattered throughout the garden areas.

On the other hand, the discovery of P. bumeliae in

the green spaces of the cities, where its secondary hosts

(Abies and Ligustrum) are also present, may suggest that

it may be much more widespread in the peninsula than

one might initially think.

Normally, in contrast to the invasive P. fraxinifolii

(Hataj & Osiadacz 2017), European Prociphilus spe-

cies are not considered harmful for their Fraxinus hosts,

despite the impressive leaf malformations and the great

amount of honeydew produced (Pérez Hidalgo & Mier

Durante 2012). However, different authors sustain the

hypothesis of an indirect damage from P. bumeliae and

P. fraxini sucking activity, which could lead to facilitate

ash pathogens infections, like those of the fungus Hyme-

noscyphus fraxineus (Chalara fraxinea) Queloz et al.,

2011 (Jankovsky & Holdenrieder 2009; Longauerova

et al. 2013).

Acknowledgements. The authors are very grateful to Doc.

Francesca Maria Gatti (Université de Strasbourg, Master in

Translation, specialized in English and Russian) for reviewing

the article in English. The authors want to thank the reviewers

and editor for their precious help and comments to improve this

paper.

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