Bonn zoological Bulletin 71 (1): 1-7

2022 -Bohme W. & Jablonski D.

https://do1.org/10.20363/BZB-2022.71.1.001

ISSN 2190-7307

http://www.zoologicalbulletin.de

Research article

urn:|sid:zoobank.org:pub:36B7D96A-85F5-4C 8E-B7A9-04128BAA0A 83

Making forgotten information available: An early study on the Afghanistan

Mountain Salamander Paradactylodon (Afghanodon) mustersi (Smith, 1940)

(Caudata: Hynobiidae)

Wolfgang Béhme!* & Daniel Jablonski?

'Stiftung Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany

?Department of Zoology, Comenius University in Bratislava, Ilkovicova 6, Mlynska dolina, SK-842 15 Bratislava, Slovakia

“Corresponding author: Email: w.boehme@leibniz-zfmk.de

'urn:lsid:zoobank.org:author: FFAC2972-9F52-404B-BA9C-489C7793FF8D

2 urn:Isid:zoobank.org:author:B624407A-7A F2-4871-9B03-C0A694959B8A

Dedication. We dedicate this paper to all enthusiastic students, teachers and researchers in Afghanistan to encourage them de-

voting themselves to the study of biodiversity of their country, as did the author of the study on the Afghan Mountain Salamander

already nearly six decades ago.

Abstract. The first study after the official species description of the endemic Afghanistan Mountain Salamander Para-

dactylodon (Afghanodon) mustersi (Smith, 1940) was published in Kabul, Afghanistan, and only in Dari Persian. We,

therefore, provide here an English translation of this paper, together with so far unpublished background information on

this rare and endangered amphibian, and on the former scientific German-Afghan cooperation project from the 1960s in

the framework of which this study had been performed.

Key words. Batrachuperus, Central Asia, Hindu Kush, ecology, conservation, history.

INTRODUCTION

The hynobiid Afghanistan Mountain Salamander Para-

dactylodon (Afghanodon) mustersi (Smith, 1940) is en-

demic in a relatively small part of northeastern Afghani-

stan (Provinces of Ghazni, Kabul, Parwan, and Wardak:

see BOhme 1982; Wagner et al. 2016; Ahmadzadeh et al.

2020). However, one dubious record of an urodelan am-

phibian (very likely a Paradacytlodon) is also known

from the Chitral valley of Pakistan (Wall 1911). This

means, the species is strictly endemic to the Hindu Kush

mountain range and represents thus one of the most en-

dangered amphibian species in the world (Stuart et al.

2008); according to the IUCN Red Data List of Threat-

ened Species (Papenfuss et al. 2004), this species falls

under the category “Critically endangered”. Its first de-

scription (Smith 1940) had considered it the westernmost

species of the otherwise East Asian (Chinese) genus Ba-

trachuperus Boulenger, 1878. Up to date, there are only

few mostly short, semi-popular or research papers on this

rare salamander, particularly on its life habits or biology

(Mertens 1970; Seufer 1974; Van Meeuwen 1977; Spar-

reboom 1977, 1978; Bohme 1982; Reilly 1983; Jablonski

et al. 2020).

However, it is not well known that an early study of its

ecology and biology after the official species description

Received: 16.06.2021

Accepted: 01.12.2021

(Smith 1940) was performed by Storai Nawabi when she

was a Student of Prof. Dr. Ernst Kullmann (1931-1996).

It was published in an Afghan journal named “Science”

which had appeared in Kabul (Nawabi 1965) and was

written in Dari Persian, a dialect of Farsi used in western,

central and northeastern Afghanistan.

The late Dr. Josef Eiselt (1912—2001), herpetologist at

the Vienna Natural History Museum, became interested

in this hardly accessible manuscript when he started to

work on this group of urodelans himself together with

a colleague (Eiselt & Steiner 1970) by describing a new

presumed congener from Iran which they named Ba-

trachuperus persicus Eiselt & Steiner, 1970 (see

Schmidtler & Schmidtler 1971). Thanks to his efforts,

a handwritten translation of the manuscript into Ger-

man was performed which made the information on this

rare salamander species available at least to the German

speaking scientific community so that for instance it could

be used for B6hme’s (1982) notes on this species. The de-

scription of another species from NE Iran, viz. B. gorgan-

ensis, by Clergue-Gazeau & Thorn (1978), forced more

interest in these westernmost hynobiids (e.g., Stock 1999)

and finally led to a generic partition, with the erection of

a new genus Paradactylodon Risch, 1984 for B. gorgan-

ensis. This concept was adopted by numerous authors to

accommodate the three western Batrachuperus taxa as

Corresponding editor: P. Wagner

Published: 13.01.2022

2 Wolfgang Bohme & Daniel Jablonski

opposed from their Chinese/Tibetan congeners for near-

ly three decades (e.g., Zhang et al. 2006; Raffaélli 2007;

Poyarkov 2010; Ahmadzadeh et al. 2011). Dubois & Raf-

faélli (2012) and Dubois et al. (2021), however, regarded

Risch’s (1984) name, although published under A. Du-

bois’ editorship, as incompatible with the regulations of

the Code (ICZN 1999) and replaced it by two new gen-

era, which were extremely shortly, only in a tabular form

diagnosed, viz. Jranodon Dubois & Raffaélli, 2012 for

the two Iranian taxa and the monotypic Afghanodon Du-

bois & Raffaélli, 2012 for the Afghan representative. Al-

though the calculated age of the split between these two

clades is 22.72 Mya (Ahmadzadeh et al. 2020) and thus

would justify generic separation, we follow these authors

in maintaining Paradactylodon as the genus name, the

more as Frost (2021) has convincingly argued for the

availability of Paradactylodon again. If it is warranted to

equip the Iranian (P. gorganensis is nested within persi-

cus!) and the Afghan species with an own monotypic ge-

nus-group name each, a subgeneric distinction would be

sufficient and beneficial for nomenclatural stability. Such

a distinction could also highlight the endemicity of these

divergent clades. Jranodon should then be endemic for

the Iranian Alborz and Talysh Mountains, and Afghano-

don for (according to our current knowledge) the Afghan

part of the Hindu Kush Mountains.

In this paper, we shall provide an English translation

of Nawabi’s (1965) paper, based on the handwritten Ger-

man version, in order to make the early, rare and local

information on this poorly known and critically endan-

gered salamander available to the scientific community,

the more as neither this work nor the note by Bohme

(1982) are cited in the recent checklist of the Afghan

herpetofauna by Wagner et al. (2016). A facsimile copy

of the original Farsi-written paper is also provided as an

appendix. We also provide some information on the Ger-

man-Afghan cooperation (partnership contract between

the universities of Cologne and Bonn on the one side,

and of Kabul on the other) in the 1970s, to illustrate the

intellectual climate in which the zoological research on

the fauna of Afghanistan flourished in those days. The

German projects in Afghanistan, mainly the foundation

of a zoological garden and a zoological museum nearby

were by-products, but not official parts of the mentioned

university partnership (Kullmann 1970; Naumann &

Nogge 1973; Nogge 2010). It was the zoo in Kabul

which brought Storai Nawabi in contact with Prof. Ernst

Kullmann who later became her academic supervisor in

Bonn. The first Dari-written study from Afghanistan car-

ried out and published by her (Nawabi 1965) remained

largely unknown, obviously due to linguistic reasons. We

here provide the first English translation of the handwrit-

ten German version initiated by J. Eiselt (some minor

additions for better understanding being in square brack-

ets).

Bonn zoological Bulletin 71 (1): 1-7

Fig. 1. (Fig. 2 in the original paper). Egg sacs of Paradacty-

lodon (Afghanodon) mustersi with developing larvae. Photo:

Storai Nawabi

English translation

A rare amphibian species in Afghanistan

Among the animals occurring in Afghanistan amphibi-

ans are hardly explored. Amphibians are vertebrates and

form a group between fishes and reptiles. Amphibian

larvae are aquatic and breathe with gills, but the adults

can also live on land because they get through their met-

amorphosis a new respiratory system. The gills are re-

duced and lungs are developed.

As far as currently known, there are more than 400 bird

species in Afghanistan, but only four species of amphibi-

ans. As a reason (for this poor amphibian fauna) it could

be assumed that Afghanistan is a dry country where large

parts are very warm in summer, but very cold in winter,

particularly above 1500 m [above sea level]. Moreover

several rivers are shallow and dry completely out in sum-

mer. Some rivers, however, have such a steep gradient

that they are, from this reason alone, unsuitable for such

animals.

Three of the amphibians belong to the group Anura

(tailless amphibians), and two of them are frogs, the sci-

entific names of which are (I) Rana ridibunda, and (2)

Rana sternosignata. The third one is a toad and is called

Bufo viridis. Jt is largely distributed and can be found

on creeks and in the mountains, as well as in cities and

is even common and present in large numbers in Kabul.

Only one species of the group of Urodela (tailed am-

Dhibians) which is called Batrachyperus [sic] mustersi,

has been found in 1935 and was described by Smith in

1940 as a new species (Contribution to the herpetology

of Afghanistan. Annals and Magazine of Natural History,

ser. IT. Vol 5, 382-384).

©ZFMK

An early study on the Afghanistan Mountain Salamander Paradactylodon (Afghanodon) mustersi )

Batrachyperus mustersi belongs to the Hynobiidae,

which are partitioned in five groups:

(1) Batrachyperus has two subgroups: B. pinchoni,

and B. tibetanus, which has only been found in the

mountains of South China.

(2) Hynobius with nine subgroups. One of them

is Hynobius keyserlingli [sic], occurring in

Kamtschatka, Mongolia, Mandshuria,

Europe at the Ural Mts.

(3) Onychodactylus has two subgroups: O. yaponicus

[sic] and O. fischeri, ranging from Korea to Japan.

(4) Pachypalaminus /iving in Japan.

(5) Ranodon one species of which, Ranodon sibiricus,

occurs in western China and Turkestan.

reaching

The family Hynobiidae has been found only in Asiatic

countries and represents the early urodelans. Represent-

atives of this family are characterised by their teeth, they

are called angle-toothed urodelans. Many of them have

four toes each at the fore- and hindlimbs.

B. mustersi has been found so far only in Paghman,

until now no other place has been recorded.

During all student excursions of our Institute numer-

ous water-breeding animals have been collected. So far,

their development, though potentially quite interesting,

has not yet been studied in detail.

We found B. mustersi in a tributary or side arm of the

Paghman River which runs through the Paghman valley.

The area is water-rich and partly forested. Because the

river itself is fast-running, its side arms provide better

conditions for the development of the eggs. The eggs are

forming long sac-like structures. One end is fixed at the

lower part of a stone and in direct contact with water,

while in Hynobius the eggs are deposited on plants at

the water surface. The shape of the egg sacs can be seen

in Fig. 2; remarkable is that one end of this structure

is pointed. Each egg consists of a gelatinous substance.

Interiorly is a germinal disc which grows in the course

of its development until the larva hatches. Such a round

egg measures 12 mm in diameter, the larva 13 mm. The

following table provides some measurements:

Values found on 17 June 1965 per egg sac

Lengt Width Number of eggs

I5cm Dies Ew

I3.cm Ze 29.

I3.cm 2.2 Di

10,5 cm is ee,

10cm ES, 30

10cm 2.4 PAL

Scm 1.6 25

Bonn zoological Bulletin 71 (1): 1-7

8S cm 2 2]

7 cm 2.4 19

6cm 2.4 13

Not all of the eggs within one sac are finishing their de-

velopment, some being unfertilized, some dying during

the embryonic development and some being viable. For

example: Out of 24 eggs, six remained unfertilized, five

remained in the embryonic stage, and 13 hatched. The

young larvae which leave the egg sac lie on the lower

part of the bottom.

We made the observations of this procedure under

artificial conditions. Whether the procedure would take

place in the same way also under natural conditions, is

not known. With our (limited) possibilities we were una-

ble to keep the larvae alive, because they need sufficient

amounts of oxygen and fresh water. Moreover, we are still

not sure which kind of food they need. Until now, we were

unable to collect larvae at exactly those places where we

had found the egg sacs. We assume that the hatched lar-

vae move to the Paghman River itself.

Young larvae have a fish-like appearance (Fig. 3) and

are 14-16 mm long immediately after hatching. The

whole body with tail, except of head and breast, is sur-

rounded by an uninterrupted median fin seam which is

reduced with the further development of the larva. Trac-

es of this seam are, however, still visible in older larvae

and in freshly metamorphosed animals. The larvae have

three pairs of juxtaposed gills. According to our studies,

the gills are still persisting in larvae up to 8.5 cm in size.

With the reduction of the gills the lungs start function-

ing, but the beginning of the metamorphosis stage itself

was not yet observed. In very young larvae, we observed

filiform appendices closely behind the eyes which are

subsequently lost again. These structures serve for main-

taining the larva’s equilibrium in the water (Fig. 3). At

the beginning of the larval stage, close to the gills, the

forelimbs start to develop, followed by the hind limbs lat-

er on, so that in larvae of 8.5 cm length as in Fig. 4 the

gills as well as the four extremities are visible.

The larvae have a whitish ground color with small

brown flecks scattered over the entire body including the

fin seams, which become scarcer towards the underside

and are invisible on breast and belly. During metamor-

phosis, dark green to brownish spots are increasing on

the dorsum. The developed larvae are silvery-brown at

their undersides with a greenish shine and have often

ventral silver-colored spots which are not well visible in

the metamorphosed animal but can persist on the low-

er jaw. The outer shape of the animal looks somewhat

flattened and is laterally segmented (Fig. 5). An adult

specimen may reach a length of 21.5 cm, the tail, with

10.5 to 11 cm, making approximately the half of the total

length. Our further investigations lead to the result that

after their egg deposition, the adults do not return to a

©ZFMK

4 Wolfgang Bohme & Daniel Jablonski

Fig. 2. (Figs 3—5 in the original paper). Young larva (above), larva ripe for metamorphosis (middle), and adult specimen (below).

Drawing and photos: Storai Nawabi

terrestrial habitat but stay permanently under water. We

emphasize this here because we found the salamanders

exclusively in the water, submersed under stones. This

was the result of our investigations as far as we could

perform them in the frame of our possibilities, and which

we plan to continue.

Fig. 1. Region where we found B. mustersi

Fig. 2. Eggs of these animals and their egg sacs which

contain already larvae

Fig. 3. Freshly hatched larva

Fig. 4. Developed larva with gills and extremities

Fig. 5. Adult B. mustersi

My cordial thanks for his support go to Dr. Kullmann

Bonn zoological Bulletin 71 (1): 1-7

DISCUSSION

The above text must be seen today against its historical

background: It was written 57 years ago by a young Af-

ghan female student in Kabul, Afghanistan, who carried

out a study including some fieldwork on the only uro-

delan species of her country, supervised by a German zo-

ologist. In fact, she was the first Afghan student who pub-

lished a herpetologically-related paper from Afghanistan.

The nomenclature she used is that of her time. The

subject of her study is now, after the discovery of a re-

lated species in Iran, and as stated already in the intro-

duction, assigned to the genus Paradactylodon Risch,

1984, which was partitioned again and even replaced by

©ZFMK

An early study on the Afghanistan Mountain Salamander Paradactylodon (Afghanodon) mustersi 5

hes

hh Oh

7 tea <t

‘ +

EEL pe, We

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z os > Sur er r)4 os) tye ty At

or “= =~ , ¥

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Fig. 3.

two names, by Dubois & Raffaélli (2012), the one for

our species here being Afghanodon. The genus concept

is to unite (monophyletically) related species, thus giving

information on phylogenetic relationship, while mono-

typic genera are merely a prefix of a single species name

(Richter 1943). Here, we maintain Paradactylodon and

group Afghanodon only as a subgenus of the Afghan en-

demic brook salamander which is sufficient to express

its long evolutionary isolation from the Iranian congener.

One character in the short and only tabular diagnosis of

Afghanodon and its Iranian fellow (sub)genus /ranodon

was an abbreviation for the “adaptability in terrarium*:

HAT (high) for the latter, and LAT (low) for the former,

certainly a weak diagnostic character for a new genus

name. Moreover, several reports have shown, that also

P. (A.) mustersi is well adaptable to captive conditions

(Mertens 1970; Seufer 1974; Sparreboom 1977, 1978:

Bohme 1982) and yielded several natural history data

(food intake by tongue protrusion, partial terrestriality,

reproductive behavior) obtained by these authors from

specimens kept in aqua-terraria.

The other amphibian species mentioned in Nawabi’s

(1965) paper, viz. Rana ridibunda, Rana sternosignata,

and Bufo viridis, are currently identified as Pelophylax

terentievi (Mezhzherin, 1992) Chrysopaa_ sternosigna-

ta (Murray, 1885) and a member of the Bufotes viridis

(Laurenti, 1768) complex (see Wagner et al. 2016), most

probably B. pseudoraddei (Mertens, 1971) (see Dufres-

nes et al. 2019). All these species are well known to be

present in the Paghman area (Wagner et al. 2016).

Due to the German partnership project between the

universities of Cologne and Bonn on the one side, and

the University of Kabul on the other, Storai Nawabi had

the chance to move to Bonn for the continuation of her

studies where she worked with an entomological-parasi-

Bonn zoological Bulletin 71 (1): 1-7

— - F e »,

? ots . - y F :

O0e sn PL FD = (- ‘ : bal Woe >. £, 2%

tological topic (Kullmann & Nawabi 1971) and also with

an arachnological-histological issue for her master thesis

(Nawab 1974).

The ZFMK received a valuable series of more than

fifty P. (A.) mustersi specimens including also larval

stages and egg sacs, collected by Clas M. Naumann

(1939-2004), lepidopterist and founder of the Zoological

Museum in Kabul (Naumann & Nogge 1973), from 1974

onwards husband of Storai Nawabi and later (from 1989)

ZFMK’s director (BOhme 2004; Hauser 2004; Schmitt

2005). Due to his efforts, a trial was made to collect

money for the reconstruction of the Zoological Museum

Kabul which was completely destroyed during the long

Afghan wartime, the initiative being called “Ein Stuhl fiir

Kabul” (= A chair for Kabul). However, this initiative

failed, because of the still ongoing military confronta-

tions which are likely to increase rather than to decrease

in the next time. As this causes always also a great loss

of biodiversity, the endemic, small-scaled distributed

and thus highly threatened Afghan mountain salamander

would thus make an ideal flagship species for the greatly

neglected nature conservation in Afghanistan (Jablonski

et al. 2021).

Acknowledgements. We thank Dr. Storait Naumann-Nawabi,

Bonn, for providing us with some information. Mrs. Hamideh

Fard, Museum Koenig Bonn, gave some linguistic advice. This

study was supported by the Slovak Research and Development

Agency under contract no. APVV-19-0076.

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6 Wolfgang Bohme & Daniel Jablonski

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bay Natural History Society 21: 121-145

Zhang P, Yue-Qin C, Hui Z, Yi-Fei L, Xiu-Lung W, Papenfuss

TJ, Wake D, Liang-Hu Q (2006) Phylogeny, evolution and

biogeography of Asiatic salamanders (Hynobiidae). Pro-

ceedings of the National Academy of Sciences of the United

States of America 103: 7360-7365

©ZFMK

An early study on the Afghanistan Mountain Salamander Paradactylodon (Afghanodon) mustersi

APPENDIX I

Facsimile copy of the title page of the original article in Farsi.

OCIENCE | KABUL, | SPECIAL SViToN | (A

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365) [p. 24-25)

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(Contribution to the herpetology of Afghanistan.

annals and magazin cf natural History.ser Il.vol 5

382 . 384)

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©ZFMK

BHL

Blank Page Digitally Inserted

Bonn zoological Bulletin 71 (1): 9-17

2022 Constant J. & Pham T.H.

https://do1.org/10.20363/BZB-2022.71.1.009

ISSN 2190-7307

http://www.zoologicalbulletin.de

Research article

urn:lsid:zoobank.org:pub:C4E72122-207A-4ED4-B71E-8BF6E35DC753

First record of the lanternfly genus Limois Stal, 1863 in Vietnam

with a new species, L. sonlaensis sp. nov.

(Hemiptera: Fulgoromorpha: Fulgoridae)

Jéréme Constant!" & Thai Hong Pham?

' Royal Belgian Institute of Natural Sciences, O.D. Phylogeny and Taxonomy, Entomology, Vautier street 29,

B-1000 Brussels, Belgium.

? Mientrung Institute for Scientific Research, Vietnam Academy of Science and Technology, 321 Huynh Thuc Khang, Hue, Vietnam

?Vietnam National Museum of Nature & Graduate School of Science and Technology, Vietnam Academy of Science and

Technology, 18 Hoang Quoc Viet, Hanoi, Vietnam

“Corresponding author: Email: jerome.constant@naturalsciences.be

'urn:lsid:zoobank.org:author:6E6072A 1-941 5-4C8D-8E60-2504444DB290

?urn:lsid:zoobank.org:author:E34CB863-7E3B-4E8F-8738-B41CO7D9F5F9

Abstract. The lanternfly genus Limois Stal, 1863 is recorded from Vietnam for the first time based on a new species,

L. sonlaensis sp. nov. from Son La Province in North Vietnam. The new species is described and illustrations of the holo-

type and male genitalia and a distribution map, are provided. An updated identification key to the nine species of Limois

is given.

Key words. Fulgoroidea, planthopper, Auchenorrhyncha, Tonkin, Indochina, China.

INTRODUCTION

The family Fulgoridae Latreille, 1807 contains about

774 species in 142 genera according to the FLOW da-

tabase (Fulgoromorpha Lists On the Web — Bourgoin

2021), representing slightly less than 6 % of the species

of Fulgoromorpha Evans, 1946 globally. In Vietnam, the

family includes 37 species and the country is currently the

most diverse in terms of lanternfly species as compared

to other Indochinese countries such as Cambodia (17 spe-

cies — Constant et al. 2016; Constant & Bartlett 2019),

Laos (6 species — Bourgoin 2021), Thailand (16 species —

Bourgoin 2021), and even China (32 species — Bourgoin

2021).

The genus Limois Stal, 1863 contains eight extant

species and is distributed in the Oriental Region and

the southeastern and far eastern parts of the Palaearctic

Region: China (northeast China, Shanxi, Shaanxi, He-

bei, Beijing, Gansu, Ningxia, Xizang, Hunan, Sichuan,

Guangxi, Fujian), Taiwan, Japan, Korea, Far Eastern

Russia, Bangladesh, northern India and Myanmar (Wang

et al. 2020). It also contains two extinct species from

China, and is the type-genus of the tribe Limoisini Lal-

lemand, 1963 in the subfamily Aphaeninae Blanchard,

1847 (Wang et al. 2020; Bourgoin 2021).

The recent publication of a revision of the genus Limois

by Wang and co-authors (Wang et al. 2020) attracted the

Received: 07.07.2021

Accepted: 21.01.2022

attention of the second author to a specimen in the collec-

tion of the Vietnam National Museum of Nature, which

was recognised as both an additional lanternfly genus for

the fauna of Vietnam and a species new to science.

The aim of this paper is to describe the new species of

Limois, L. sonlaensis sp. nov. as an addition to the biodi-

versity of Vietnam and to provide an updated identifica-

tion key for the nine extant species of Limois.

MATERIALS AND METHODS

The male genitalia were extracted after boiling the distal

portion of the abdomen several minutes in a 10% solu-

tion of potassium hydroxide (KOH) at about 100°C. The

pygofer was separated from the remains of the abdomen

and the aedeagus dissected with a needle blade for ex-

amination. The organs were then placed in glycerine for

preservation in a tube attached to the pin of the speci-

men. The external morphological terminology follows

O’Brien & Wilson (1985), the wing venation terminology

follows Bourgoin et al. (2015) and for the male genitalia,

Bourgoin & Huang (1990). The metatibiotarsal formula

gives the number of spines on (side of metatibia) apex

of metatibia/apex of first metatarsomere/apex of second

metatarsomere.

Corresponding editor: R. Peters

Published: 25.01.2022

10 Jér6me Constant & Thai Hong Pham

Abbreviations for measurements

The measurements were taken as in Constant (2004).

BF = maximum breadth of the frons

Brey T= maximum breadth of the tegmen

BV = maximum breadth of the vertex

LE = length of the frons at median line

LTg = maximum length of the tegmen

LT = total length (apex of head to apex of

tegmina)

LV = length of the vertex at median line

The photographs of the collection specimens were tak-

en with a Canon 700D camera equipped with a Sigma

50 mm Macro lens; those of the terminalia, with a Leica

EZ4W stereomicroscope with integrated camera, and in

both cases, stacked with CombineZ software and opti-

mized with Adobe Photoshop CS3. The distribution map

was produced with SimpleMappr (Shorthouse 2010).

Institutional abbreviations

VNMN = Vietnam National Museum of Nature, Hanoi,

Vietnam.

GTI = Global Taxonomy Initiative.

RESULTS

Order Hemiptera Linnaeus, 1758

Suborder Auchenorrhyncha Dumeril, 1806

Infraorder Fulgoromorpha Evans, 1946

Superfamily Fulgoroidea Latreille, 1807

Family Fulgoridae Latreille, 1807

Subfamily Aphaeninae Blanchard, 1847

Tribe Limoisini Lallemand, 1963

Genus Limois Stal, 1863

Limois Stal, 1863: 230.

Type species: L. westwoodii (Hope, 1843).

Limois — Metcalf 1947: 170. — Lallemand 1963: 54. —

Chou & Lu 1981: 221. — Chou et al. 1985a: 30. —

Chou et al. 1985b: 108. — Nagai & Porion 1996: 22.

— Wang et al. 2020: 36.

Species included [distribution]

Limois bifasciatus Ollenbach, 1928 [India, Uttarakhand

State (Ollenbach, 1928)]

Limois chagyabensis Chou & Lu, 1981 [China: Xizang,

Shaanxi, Sichuan (Wang et al., 2020)]

Limois emelianovi Oshanin, 1908 [China: Gansu, Dong-

bei (Wang et al. 2020); Russia (Oshanin 1908; Anu-

Bonn zoological Bulletin 71 (1): 9-17

friev 2009); Korea (Doi 1932a, 1932b; Kwon & Huh

2001)].

Limois guangxiensis Chou & Wang, 1985 [China:

Guangxi, Fujian (Wang et al. 2020)|

Limois hunanensis Chou & Wang, 1985 [China: Hunan

(Wang et al. 2020)]

Limois kikuchii Kato, 1932 [China: northern China (Kato

1932, 1933), Shaanxi, Beijing (Liang 2005), Korea

(Kato 1933; Metcalf 1947)]

Limois sonlaensis sp. nov. [ Vietnam: Son La Province]

Limois sordida Wang, Xu, Constant & Qin, 2020 [China:

Shanxi; Hebei, Beijing (Wang et al. 2020)]

Limois westwoodii (Hope, 1843) [Bangladesh (Hope

1843); China: Xizang; Myanmar (Wang et al. 2020)|

Identification key to the species of Limois Stal, 1863

1. Frons with three longitudinal carinae (Wang et al.

2020: figs 1E-F, 4C, 6E, 11D) «0.2000 2

— Frons with two longitudinal carinae (Wang et al.

2020: figs 2E, 3B, 8D, 9D-F) ........... ee 5

2. Pronotum with one dark irregular patch along the

posterior margin on each lateral area (Wang et al.,

2020: fig. 11B); base of hindwings yellow in male

and red in female (Wang et al. 2020: fig. 11A, C) ...

eR eee eter eee te L. emelianovi Oshanin, 1908

— Pronotum without one dark irregular patch along

the posterior margin on each lateral area (Wang

et al., 2020: figs 1D, 4B, 6D); base of hindwings

concolorous in both sexes (Wang et al., 2020:

figs L[A—B, 4A, D, 6A—B) 00... 3

3. Pronotum brown (Wang et al., 2020: fig. 4B); genital

styles distinctly elongate, subtriangular in lateral

view (Wang et al., 2020: fig. SA-B) 0.0...

hi hah cerashtdSeresuckstenaAtiaan hatter L. kikuchii Kato, 1932

— Pronotum purplish red or dark brown (Wang et al.

2020: figs 1D, 6D); genital styles short, almost

equilateral in lateral view (Wang et al. 2020: figs 1H,

Wg od aan ae Ne A ee ES ORR 4

4. Pronotum purplish red (Wang et al. 2020: fig. 1D);

thorax sparsely covered with few dark spots (Wang

et al., 2020: fig. 1D); irregular stripe in tegmina short

and straight, not extending to anal angle (Wang et al.,

2020: fig. 1A—C); apical half of endosomal processes

exposed (Wang et al. 2020: figs IK—L, 14C—-D) ......

Rene os, eee L. chagyabensis Chou & Lu, 1981

— Pronotum dark brown (Wang et al. 2020: fig. 6D);

thorax densely covered with numerous dark spots

(Wang et al., 2020: fig. 6D); irregular stripe in tegmina

long, sinuately extending to anal angle (Wang et al.,

2020: fig. 6A—-C); apical ‘4 of endosomal processes

exposed (Wang et al. 2020: figs 7D-E, 15C-—D) ......

aot, eae L. sordida Wang, Xu, Constant & Qin, 2020

©ZFMK

The new lanternfly Limois sonlaensis sp. nov. from Vietnam (Hemiptera: Fulgoridae) 1]

5. Thorax densely covered with numerous minute dark

spots, disc of mesonotum without black marking

(Fig. 1B; Wang et al. 2020: fig. 2D); tegmina

submedially without strongly contrasting broad

irregular band (Fig. 1A; Wang et al. 2020: fig. 2A)..

— Thorax with minute dark spots sparse or absent, disc

of mesonotum with black markings (Wang et al.,

2020: figs 3A, 8B, 9B); tegmina submedially with

contrasting broad irregular band (Wang et al. 2020:

ess SA Ce ROAR Reel ba Pe cog Ten ere if

6. Minute black spots density equal on disc of pro- and

mesonotum (Wang et al., 2020: fig. 2D); postclypeus

pale yellowish with minute darker spots (Wang et al.,

2020: fig. 2E); dorsal angle of gonostyli rounded in

lateral view (Wang et al. 2020: fig. 2G—H) ..............

NE ce es Sa L. guangxiensis Chou & Wang, 1985

— Minute black spots covering on disc of pronotum

distinctly less dense than that on disc of mesonotum

(Fig. 1B); postclypeus dark brown with irregular

pale yellowish markings (Fig. 1C); dorsal angle

of gonostyli forming a right angle in lateral view

(eR Ceo.) | ROI ee Re ee ee L. sonlaensis sp. nov.

7. Pronotum with 2 dark spots anteriorly, without one

longitudinal broken black band on each side of

median carina (Wang et al. 2020: fig. 3A) «0...

an MIs dots a te L. hunanensis Chou & Wang, 1985

— Pronotum without dark spots anteriorly, with one

longitudinal broken black band on each side of

median carina (Wang et al. 2020: figs 8B, 9B) ...... 8

8. Tegmina with an oblique narrow fascia from apex to

disc: (Wane ct al 2020) Te. 13). os, ects 8 iacs Boe 2

Sede Asa by Pena lobes Femaat L. bifasciatus Ollenbach, 1928

— Tegmina without an oblique narrow fascia from apex

to disc (Wang et al. 2020: figs 8A, C, E, 9A, G) ......

be ctckes| Be conc aM ays L. westwoodii (Hope, 1843)

Limois sonlaensis sp. nov.

urn: lsid:zoobank.org:act: BZA E4EF 3-DA99-4E9A-9A 79-25741F 7CO0EF

(Figs 1-4)

Material examined

Holotype

3 (Figs 1-3). Vietnam, Son La Province; Thuan

Chau District, Co Ma, alt. 1,400 m; 21°21711.5” N,

103°32’35.5” E; 7 Oct 2008; leg. Hoang Vu Tru; light

trap; VNMN.

Diagnosis

The species can be separated from the other Limois spe-

cies by the combination of the following characters:

1. Frons with two longitudinal carinae (Fig. 1C).

2. Pro- and mesonotum without large dark making on

disc, only with minute black spots; spots distinctly less

dense on pronotum than on mesonotum (Fig. 1B).

Bonn zoological Bulletin 71 (1): 9-17

3. Postclypeus dark brown with irregular pale yellowish

markings (Fig. 1C).

4. Tegmina with red suffusion on basal half of corium,

the reddish area not reaching base and not extending to

costal cell (Fig. 1A, D).

5. Posterior wigs with broad dark brown band along

posterior margin and broad dark brown band separat-

ing basal orange-red area from transparent distal third

(Fig. 1A, D).

6. Gonostyli with dorsal angle forming a right angle

(Fig. 2A).

Differential diagnosis

The most similar species is L. guangxiensis Chou &

Wang, 1985 from which L. sonlaensis sp. nov. can be sep-

arated, for example by the minute spots on disc of prono-

tum less densely distributed than on disc of mesonotum

(spotting density similar on disc of pro- and mesonotum

in L. guangxiensis) and by the pale yellowish ground co-

lour of the postclypeus (postclypeus dark brown with ir-

regular pale yellowish markings in L. sonlaensis).

Limois sonlaensis sp. nov. can be separated from the

remaining other species of the genus by the absence of

dark brown or black markings on the disc of the pro-

and mesonotum. The remaining other species, except

L. westwoodii, also lack the dark brown band separating

the orange-red basal area from the transparent distal area

of the posterior wings.

Etymology

The species epithet refers to Son La Province in North

Vietnam, where the type specimen was collected.

Description

Measurements and ratios. LT: ¢ (n= 1): 20.6 mm. LTg/

BTg = 2.6; LV/BV = 0.5; LF/BF = 1.03.

Head. Vertex excavate, with all margins carinate;

about two times broader than long; weakly transverse-

ly wrinkled; with a broadly U-shaped carina along me-

dian 2/, of posterior margin; pale yellow with irregular

brown markings in middle and at posterolateral angles

(Fig. 1B, E). Posterior face of head pale yellowish with

two longitudinal brown markings near middle (Fig. 1B,

E). Frons about as broad basally as long in mid-line

(excluding cephalic process); slightly convex in lateral

view; broadest with margins rounded near fronto-clypeal

suture, then moderately tapering dorsad with lateral

margins broadly incurved; portion above fronto-clypeal

suture slightly excavate, then median portion of disc el-

evated; elevation tapering dorsad and gradually turning

into two side-by-side carinae extending nearly to apex

of cephalic process; frons mostly dark brown mottled

with pale yellow and rather densely covered in minute

black points (Fig. 1B—C, H). Cephalic process directed

posterodorsally, tapering towards apex; in lateral view,

anterior margin rounded and posterior margin straight,

©ZFMK

12 Jér6me Constant & Thai Hong Pham

oblique; anterior face with lateral margins foliate lateral-

ly near base and two parallel median carinae stopping be-

fore a small lamina; small lamina followed by excavate

apex of process; posterior face of process flattened, elon-

gate with sides parallel and lateral margins carinate, and

dorsal margin angularly pointed dorsad; pale yellowish

slightly variagated with brown and with irregular minute

reddish brown points, and with carinae of anterior face

dark brown (Fig. 1B—C, E-F, H). Genae pale yellowish

with few brown markings (Fig. 1E, H). Eyes large, about

half as wide as vertex in dorsal view, strongly rounded

and protruding laterally (Fig. 1B—C). Ocelli well-devel-

oped, under compound eyes (Fig. 1H). Antennae rather

short, subcylindrical, black-brown (Fig. 1E, H). Clypeus

convex, elongate, subtriangular, carinate medially with

carina stronger on anteclypeus; lateral margins carinate

on basal % of postclypeus; base narrower than frons;

brown with anteclypeus darker and transverse irregular

pale yellowish markings; some minute black points on

postclypeus along lateral and median carinae (Fig. 1C—

D). Labium brown, very elongate and narrow, surpassing

apex of abdomen, with last segment much shorter than

penultimate (Fig. 1D).

Thorax. Pronotum wider than long with lateral angles

acutely pointed lateroventrally in dorsal view; anterior

margin strongly bisinuate, roundly emarginated behind

eyes, slightly emarginate behind vertex; posterior mar-

gin more or less straight, abruptly oblique near lateral

angle; weak median carina on anterior half of disc with a

deep impressed point at each side; disc transversely wrin-

kled posterior to impressed points; anterolateral margin

strongly carinate; paranotal lobes with strong carina

parallel to anterolateral margin on dorsal 4 and ventral

margin carinate; ventral 7%; of paranotal lobes coarsely

punctured; pale yellowish dorsally with some minute

dark brown dots and one bigger black point behind eye;

paranotal lobes with area between carinae dark brown

with large pale yellowish markings dorsally; ventral 7%

of paranotal lobes dark brown variegated with yellowish

(Fig. 1B—C, E-F, H). Mesonotum subtriangular with very

weakly marked median and peridiscal carinae; median

carina stopped before scutellum; peridiscal carinae in-

curved on anterior ‘4; scutellum slightly elevated, follow-

ing a shallow depression; ground colour pale yellowish;

disc densely covered in irregular small black dots; lateral

fields beyond peridiscal carinae smooth with some black-

brown markings including a curved marking anteriorly

(Fig. 1B, E, H). Tegulae pale yellowish with irregular

minute black points on ventral % (Fig. 1B, E, H).

Tegmina. (Fig. 1A, D, G) Elongate, slightly more than

2.5 x as long as broad, with a small round lobe posterior

to clavus (damaged on right wing of holotype); apical

angle rounded; apical margin strongly oblique; costal

cell pale yellowish brown with irregular rather large

black-brown markings; clavus pale yellowish with 1r-

regular dark brown markings, markings larger on distal

Bonn zoological Bulletin 71 (1): 9-17

¥3, basal 2 of corium opaque, pale yellowish with large

black-brown markings and a large reddish zone on disc,

not reaching base and more visible ventrally (Fig. 1D);

distal half transparent with large markings and veins,

dark brown (Fig. 1A, D, G). Venation dense and reticu-

late, with numerous secondary veins and cross-veinlets;

Pc+CP extending slightly beyond nodal line; ScP+R(+-

MA) very short; RP(+MA) merged with MP at basal 7/s

of clavus; PCu and Al fused at about % of clavus length

(Fig. 1A).

Hind wings. (Fig. 1A, D) Elongate, roundly pointed

apically, with posterior margin roundly indentate at cla-

val fold. Basal half bright orange turning to bright red

distally; distal half transparent with broad dark brown,

C-shaped band between bright red and transparent areas:

broad dark brown band along posterior margin; vein CuA

dark brown marked by an elongate dark brown marking

progressively widening towards the posterior and abrupt-

ly stopped at first fork of CuA; three dark brown makings

along vein Al in orange area.

Legs. (Fig. 1A, D) Elongate and slender. Dark brown;

femora with irregular pale yellow markings more or less

arranged in rings; tibiae with 2 well-defined pale yellow-

ish rings, the more distal one broader. Metatibiotarsal

formula: (5-6) 7 / 8-9/6.

Abdomen. Tergites orange with broad dark brown

band along basal margin (Fig. 1A, D); ventral face black-

brown (Fig. 1D).

Terminalia. Male. Pygofer (Py) with ventral margin

sinuate in lateral view; posterior margin in lateral view

slightly convex, emarginate on ventral 1/6 (Fig. 2A);

in dorsal view deeply, roundly emarginate posteriorly

(Fig. 2C, F); posterior margin slightly roundly projecting

posterad in ventral view (Fig. 2D). Gonostyli (G) subtri-

angular in lateral view, with dorsal angle right and poste-

rior margin rounded (Fig. 2A), in ventral view gonostyli

connected basally (Fig. 2D), lateral hook (/hg) slender,

directed posteroventrally, then curved ventrally towards

apex (Fig. 2A—B, D). Phallobase sclerotized in basal 1/6,

sheath cylindrical, phallobasal conjunctival processes

sclerotized over entire length, about five times as long

as sheath, apical 4 curved dorsad and terminally mem-

branous and inflated, phallus with a dorsal and a ven-

tral pair of elongate membranous processes (Fig. 3A—B,

D-G). Connective rod-like (Fig. 3A). Tectiductus rath-

er large, in lateral view elongate, in dorsal view broad,

subsquarish, distinctly concave ventrally (Fig. 3A—E, G).

Anal tube (An) massive, in lateral view reaching level of

apex of gonostyli, ventral margin nearly straight, apical

margin obliquely, roundly truncate (Fig. 2A), in dorsal

view, widening from base to apex, 1.09 x broader near

apex than long in median line, 1.34 x longer in maxi-

mum length than wide at maximum width, apical margin

roundly emarginate in dorsal view (Fig. 2C); anal column

(ac) elongate, surpassing posterior margin of anal tube

(Fig. 2A, C).

©ZFMK

The new lanternfly Limois sonlaensis sp. nov. from Vietnam (Hemiptera: Fulgoridae) 13

owe

i L i Ll

Fig. 1. Limois sonlaensis sp. nov., 6, holotype (VNMN). A. Habitus, dorsal view. B. Head and thorax, dorsal view. C. Head and

thorax, perpendicular view of frons. D. Habitus, ventral view. E. Head and thorax, laterodorsal view. F. Head and thorax, antero-

lateral view. G. Habitus, lateral view. H. Head and thorax, lateral view.

Bonn zoological Bulletin 71 (1): 9-17 ©ZFMK

14 Jér6me Constant & Thai Hong Pham

Fig. 2. Limois sonlaensis sp. nov., 4, holotype (VNMN), terminalia: anal tube, pygofer and gonostyli. A. Left lateral view. B. Cau-

dal view. C. Dorsal view. D. Ventral view. E. Posterolateral view. F. Laterodorsal view. Abbreviations: ac = anal column; An = anal

tube; G = gonostylus; /hg = lateral hook of gonostylus; Py = pygofer.

Bonn zoological Bulletin 71 (1): 9-17 ©ZFMK

The new lanternfly Limois sonlaensis sp. nov. from Vietnam (Hemiptera: Fulgoridae) 1K)

Fig. 3. Limois sonlaensis sp. nov., 3, holotype (VNMN), terminalia: aedeagus and connective. A. Lateral view. B. Laterodorsal

view. C. Dorsal view. D. Lateroventral view. E. Ventral view. F. Caudal view. G. Posteroventral view.

Bonn zoological Bulletin 71 (1): 9-17 ©ZFMK

16 Jér6me Constant & Thai Hong Pham

Note

Limois sonlaensis sp. nov. is the only species of the ge-

nus in which RP(+MA) is merging with MP on tegmina.

Both tegmina of the specimen show this character. How-

ever, as only one male is available, this character needs to

be confirmed from additional material in the future, and

might represent a good diagnostic character to recognize

this species.

Biology

The specimen was collected at a light trap in secondary

mountain tropical evergreen forest, at 1,400 m in altitude

(Fig. 4).

Distribution

Vietnam, Son La Province (Fig. 4).

DISCUSSION

The genus Limois now contains nine species, seven of

which being present in China. The genus 1s also found in

northern India, Myanmar, Bangladesh, Far Eastern Rus-

sia and South Korea (Wang et al. 2020), and it is here re-

corded from Vietnam for the first time. Despite the recent

collecting effort, especially in China, very little is known

about the natural history of these species, and host plants

remain completely undocumented so far. Specimens are

quite scarce in the collections, even if it happens that they

get collected from light traps. The fact that they seem to

prefer mountainous habitats leads to the hypothesis that

probably more species exist which will be discovered in

the future.

Acknowledgments. We thank Miss Mado Berthet (RBINS) for

improving the plates of habitus and genitalia; Dr Patrick Groo-

taert, Dr Frederik Hendrickx, Dr Marie-Lucie Susini Ondafe

and Dr Luc Janssens de Bisthoven (RBINS) for supporting our

Global Taxonomy Initiative project in Vietnam. This paper is a

result of the GTI project “A step further in the Entomodiversity

of Vietnam” supported through a grant issued by the capacity

building Programme of the Belgian Global Taxonomy Initiative

National Focal Point that runs under the CEBioS programme

with financial support from the Belgian Directorate-General for

Development Cooperation (DGD). The present study was also

= @ Limois sonlaensis sp. nov.

Fig. 4. Limois sonlaensis sp. nov., distribution map.

Bonn zoological Bulletin 71 (1): 9-17

©ZFMK

The new lanternfly Limois sonlaensis sp. nov. from Vietnam (Hemiptera: Fulgoridae) 17

supported by the Vietnam National Foundation for Science and

Technology Development (NAFOSTED) for the second author.

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©ZFMK

BHL

Blank Page Digitally Inserted

Bonn zoological Bulletin 71 (1): 19-22

2022 Takano H. & Laszlo G.M.

https://do1.org/10.20363/BZB-2022.71.1.019

ISSN 2190-7307

http://www.zoologicalbulletin.de

Research article

urn:|sid:zoobank.org:pub:07D84332-BBC3-48D1-9716-CB41D44DC574

A new species of Cryptopacha Prozorov & Zolotuhin, 2012

(Lasiocampidae: Lasiocampinae) from West Africa

Hitoshi Takano'* & Gyula M. Laszlo?

24 frican Natural History Research Trust, Street Court, Leominster, HR6 9OA, UK

*Corresponding author: Email: hitoshi.takano@anhrt.org.uk

‘urn:lsid:zoobank.org:author:22D08054-018D-44D5-8B32-1C0537E057F9

?urn:|sid:zoobank.org:author: 50CAA370-A89A-488E-A635-1B2D6FD71001

Abstract. A new species of Cryptopacha is described from West Africa: C. smithi sp. n. The habitus and genitalia are

illustrated and a comparison made with its closest congener, C. porphyria (Holland, 1893). These two taxa are separated

by the Dahomey Gap and their distribution is discussed in relation to this barrier to speciation.

Key words. Lappet moths, Dahomey Gap, biogeography, taxonomy.

INTRODUCTION

The lappet moth genus Cryptopacha Prozorov & Zolo-

tuhin, 2012 was erected by Prozorov & Zolotuhin (2012)

to include a single species, Metanastria porphyria Hol-

land, 1893, described from Gabon and thought to be

widely distributed in the lowland forests of Central and

West Africa. Cryptopacha porphyria (Holland, 1893) is

an unmistakable species with a characteristic forewing

pattern of white and purplish-brown bands on a red-

brown background. As part of recent DNA barcoding

efforts by the African Natural History Research Trust

(ANHRT), individuals from Central and West Africa

were sampled and the results suggested two, genetically

divergent populations existing either side of the Dahom-

ey Gap. Morphological studies of genitalia corroborated

the molecular results and a new species from West Africa

is described herein.

MATERIAL AND METHODS

Primary label data have been transcribed verbatim with

“//’ denoting a different label and “/’ denoting a line

break. Genitalia were dissected, stained with Eosin red

and mounted in Euparal on microscope slides applying

standard methods of preparation (Lafontaine & Mikkola

1987). Photos of adults were taken using a Nikon D90

SLR camera equipped with a Nikkor AF Micro 60 mm

lens. Genitalia were photographed using a Canon EOS

700D camera mounted on a Wild M3Z stereo micro-

scope. All images were edited in Adobe Photoshop. Gen-

italia terminology follows Prozorov & Zolotuhin (2012).

Received: 26.10.2021

Accepted: 18.02.2022

DNA barcodes were obtained by removing tarsal seg-

ments from adult specimens before being submitted to

the Canadian Centre for DNA Barcoding (CCDB, Bio-

diversity Institute of Ontario, University of Guelph).

Sequences were obtained using Single Molecule Re-

al-Time sequencing through the Sequel (PacBio) pipeline

at CCDB (Hebert et al. 2018). The resulting sequences

were aligned using MUSCLE in MEGA ver. X (Kumar

et al. 2018) and genetic distances were calculated using

Kimura’s two-parameter model (Kimura 1980).

RESULTS

Taxonomy

Family Lasiocampidae Harris, 1841

Subfamily Lasiocampinae Harris, 1841

Genus Cryptopacha Prozorov & Zolotuhin, 2012

Cryptopacha smithi sp. n.

urn:lsid:zoobank.org:act: FO6AA4E5-5 1 5A-412B-B33A-CC8B40B58618

Figs 1-5

Diagnosis

Cryptopacha smithi is very similar in appearance to

C. porphyria but in the latter, the forewing markings are

more heavily contrasting, the silvery-white bands (par-

ticularly the one beyond the submarginal band) are wid-

er and better defined and placed at slightly more of an

obtuse angle to the costal margin (Fig. 6). In the male

genitalia of C. porphyria, the tegumen 1s wider, the socil

Corresponding editor: M. Espeland

Published: 25.02.2022

20 Hitoshi Takano & Gyula M. Laszlo

are longer and the valves are more robust and shorter (the

valve-socii ratio is almost 1:1). The phallus is larger and

the pointed apical section longer in C. porphyria and the

digitiform projections of sternite VIII are more robust,

shorter and with greater serrations apically (Figs 7-9).

As these two species are distributed allopatrically, there

is unlikely to be any confusion in specimens with good

provenance.

Etymology

It is with great pleasure that this new species is dedicat-

ed to Richard Smith, the director of the African Natural

History Research Trust, for his continued support of and

dedication to African entomology and in particular Lepi-

doptera systematics.

Material examined

Holotype

WEST AFRICA -— Sierra Leone « @; “SIERRA LE-

ONE 120m / Tiwai Island, Moa River / N07°33'00",

W11°21'09" / 17—22.vi.2016 Light Trap / leg. Takano,

Miles & Goff / ANHRT:2017.18 // ANHRTUK /

00029947 // Gen. slide No. / LG 5618 m# / prep. by Gy.

M. Laszlo”; ANHRT.

Paratypes (38 d¢ 1 2)

WEST AFRICA — Guinea ¢ 5 3\'; Massadou campsite,

Forét Classée de Ziama; 08°20'36.25" N, 09°26'14.70" W;

alt. 541 m; 15—17.111.2019; Sz. Safian and S. Koivogui

leg.; Light Trap; ANHRT — Ivory Coast * 18 3: Tai

National Park; 05°50'00" N, 07°20'32.0" W; alt. 174 m;

25.i1I-17.iv.2017; A. Aristophanous, M. Aristopha-

nous, M. Geiser and P. Moretto leg.; MV Light Trap;

ANHRT ° 6 oo, 1 2: Tai National Park; 05°49'59.8" N,

07°20'32" W; alt. 174 m; 5—10.vii.2015; M. Aristopha-

nous, P. Moretto and E. Ruzzier leg.; Light Trap; ANHRT

— Liberia * 1 <; East Nimba Nature Reserve, Cellcom

Road; 07°31'2.18" N, 08°31'1.90" W; alt. 1300 m;

31 .11i-04.iv.2017; Sz. Safian and G. Simonics leg.;

Light Trap; ANHRT ¢ 3 33; Foya Proposed Protected

Area; 07°56'36" N, 10°16'36" W; alt. 530 m; 10-19.

x1.2017; M. Aristophanous, Sz. Safian, G. Simonics and

L. Smith leg.; MV Light Trap; ANHRT « 2 dc; Wele-

zu camp, Wonegizi Nature Reserve; 08°04'57.11" N,

09°34'47.86" W; alt. 561 m; 19-27.111.2019; Sz. Safian

and S. Koivogui leg.; Light Trap (Blended Bulb); ANHRT

° 1 &; Krahn-Bassa Reserve, Juboe River; 05°39'04" N,

08°39'04" W; alt. 140 m; 14—20.1.2018; M. Geiser, Sz.

Safian and G. Simonics leg.; Light Trap (Blended Bulb);

ANHRT — Sierra Leone « 1 <; same collection data as

for holotype; ANHRT + 1 3; Kambama village on the

banks of the Moa River; 07°33'29" N, 11°21'51" W;

alt. 110 m; 22.vi.2016; H. Takano, W. Miles and R. Goff

leg.; Light Trap; ANHRT.

Bonn zoological Bulletin 71 (1): 19-22

Description

External morphology of adults (Figs 1—2)

Forewing length

Holotype: 26 mm; range: 24—26 mm; female: 36 mm.

Upperside

Ground colour of head, thorax and forewings pur-

plish-brown mixed with densely scattered grey and white

scales. Head with a low longitudinal crest. Antennae

brown, bipectinate; rami lighter in colour. Patagia with

slightly greyer scales posteriorly. Tegulae with transverse

patch of white scales medially. Abdomen hirsute, cov-

ered in long red-brown hairs. Forewing rounded, outer

margin gently sinuate. Basal two-thirds (up to the sub-

marginal line) with silvery-mauve diffuse patches atop

the ground colour separated by dark brown bands and

silvery-white bands on their outer edge; basal band,

straight; double antemedial bands which kink inwards

towards dorsal margin at vein CuA2, posterior section

slightly sinuate; transverse band filling the space be-

tween veins Al+A2 and CuA2, bisecting the antemedial

bands just below CuA2 and extending to the outer mar-

gin. Discal spot white, well-developed. Postmedial and

submarginal bands slightly arcuate anteriorly kinking

sharply towards dorsal margin at vein M3, posterior sec-

tions beyond sinuate. Subterminal band zigzagged with

dark brown and white markings at corners, originating at

apical marking. Apical marking triangular, graphite-grey

with silvery markings on inner and outer edges. Hind-

wing outer margin slightly sinuate; same ground colour

as forewing, becoming paler towards the costa; anal fold

pale. Fringe greyish-brown, darkened at the veins.

Underside

Ground colour of head, thorax, legs purplish-brown; labi-

al palps brown. Forewing underside similar in colour to

upperside but slightly paler; basal and antemedian bands

absent; postmedial band well developed, submarginal

band less so, both bands converging towards dorsal mar-

gin. Broad diffuse greyish-white band beyond submargin-

al band, well-defined towards costal margin. Yellowish

scales along veins most clearly defined in the subtermi-

nal area. Hindwing ground colour similar to forewing but

overlaid with creamy scales except for tornal region and

postmedial, submarginal and subterminal bands. Termi-

nal band indicated by yellowish-white lunules. Yellowish

scales along veins most clearly defined in the subterminal

area reaching as far as postmedial line.

Female

Ground colour of body and wing pattern on both upper-

side and underside similar to male but slightly paler.

©ZFMK

A new species of Cryptopacha Prozorov & Zolotuhin, 2012 from West Africa 21

Figs 1-9. Cryptopacha smithi sp. n. and Cryptopacha porphyria (Holland, 1893), 4; Gabon, P.N. Ivindo [ANHRTUK00039000];

gen. slide No. LG 5616. 1. Cryptopacha smithi sp. n. holotype, 4, upperside. 2. Cryptopacha smithi sp. n., paratype, 2, upperside.

3. Cryptopacha smithi sp. n., holotype, 3, genital capsule; 4. idem., phallus. 5. idem., sternite VIII. 6. Cryptopacha porphyria

(Holland, 1893), upperside. 7. idem, genital capsule. 8. idem, phallus. 9. idem, sternite VIII. Scale bars: 1-2, 6 = 10 mm; 3-5,

7-9 = 1 mm.

Bonn zoological Bulletin 71 (1): 19-22 ©ZFMK

22 Hitoshi Takano & Gyula M. Laszlo

Male genitalia (Figs 3—5)

Uncus and gnathos absent; tegumen bell-shaped, apically

with a pair of robust conical soci, each basally as wide

as the width of tegumen. Tuba analis membranous. Valve

fully sclerotised, tubular, sickle-shaped, curved caudad,

two and a half times longer than the soci. Vinculum

sclerotised, curved dorsad. Saccus band-shaped. Anellus

membranous. Phallus short, strongly curved medially,

basally dilated, apically pointed. Sternite VIII with a pair

of heavily-sclerotised digitiform posterior projections

with serrations apically. Apodemes long, narrow and

rounded at extremities.

DNA divergences

The new species has been assigned the COI-5P cluster

number BOLD:AAZ9255. Interspecific pairwise dis-

tances between the new species and C. porphyria ranged

from 6.2-6.4%.

Distribution

Cryptopacha smithi is a species of the Upper Guinean

Forests distributed from Sierra Leone to Ivory Coast.

DISCUSSION

Cryptopacha smithi and its sister species C. porphyria

are distributed allopatrically on either side of the Daho-

mey Gap, a 200 km wide belt of open vegetation sepa-

rating the Upper and Lower Guinean Forest blocks. This

region was once covered by closed-canopy forest during

the mid-Holocene but rapid aridification following the

Holocene Climatic Pejoration resulted in the domination

of savannas in the Dahomey Gap (Demenot et al. 2018).

Numerous studies have shown that the Dahomey Gap

can be a significant barrier to gene flow for those species

restricted to forest habitats (e.g., Dongmo et al. 2019),

and further genetic and morphological studies of Lasio-

campidae, as well as other Lepidopteran groups will un-

doubtedly reveal many more sibling taxa separated by

this barrier.

Acknowledgments. We thank Edward M. Kargbo, Perma-

nent Secretary of the Ministry of Agriculture and Forestry,

for authorising entomological research in Sierra Leone and

to the Forestry Division for assistance in ground logistics. We

are most grateful to Aiah Lebbie and Richard Wadsworth of

the Department of Biological Sciences, Njala University for

authorising access to the research facilities on Tiwai Island.

Bonn zoological Bulletin 71 (1): 19-22

Research in Céte d’Ivoire was authorised by the Ministere de

l’ Enseignement Supérieur et de la Recherche Scientifique. The

Office Ivoirien des Parcs et Réserves (OIPR) and the Société

de Développement des Foréts (SODEFOR) are thanked for

authorising access to protected forests and providing export

permits. Darlington Tuaben, Mike C. Doryen and Kederick

F. Johnson of the Forestry Department Authority, Liberia are

thanked for issuing research permits and Annika Hiller (Wild

Chimpanzee Foundation, Liberia) and Jerry Garteh (Society for

the Conservation of Nature, Liberia) for their help in organising

ground logistics. Research in Guinea was authorised by Layaly

Camara, Directeur National, Ministere de |’ Environment et des

Eaux et Foréts. Mamdou Diawara (Directeur Exécutif, Guinée

Ecologie), Cece Papa Konde (Directeur General, Centre de

Gestion de |’Environnement du Nimba et du Simandou) and

Jamison Suter (Manager Responsibilité Environmentale et So-

cial de la Société des Mines de Fer de Guinée) are also thanked

for their assistance in planning and undertaking expeditions in

Guinea.

REFERENCES

Demenou BB, Doucet J-L, Hardy OJ (2018) History of the

fragmentation of the African rain forest in the Dahomey Gap:

insight from the demographic history of Zerminalia superba.

Heredity 120: 547-561

Dongmo J-B, DaCosta JM, Champlain D-L, Ngassam P, So-

renson MD (2019) Variable phylogeographic histories of five

forest birds with populations in Upper and Lower Guinea:

implications for taxonomy and evolutionary conservation.

Ostrich: Journal of African Ornithology 90 (3): 257-270

Hebert PDN, Braukmann TWA, Prosser SWJ, Ratnasingham S,

deWaard JR, Ivanova NV, Janzen DH, Hallwachs W, Naik S,

Sones JE & Zakharov EV (2018) A Sequel to Sanger: ampli-

con sequencing that scales. BMC Genomics 19: 219.

https://doi.org/10.1186/s12864-018-4611-3

Holland WJ (1893) Descriptions of new species and genera

of West African Lepidoptera. Psyche 6 (201): 373-376,

393-400, 411-418, 431-434, 451-454, 469-476, 487-490,

513-520, 531-538, 549-552, 565-568

Kimura M (1980) A simple method for estimating evolution-

ary rate of base substitutions through comparative studies of

nucleotide sequences. Journal of Molecular Evolution 16:

111-120

Kumar S, Stecher G, Li M, Knyaz C, Tamura K (2018) MEGA

X: Molecular Evolutionary Genetics Analysis across com-

puting platforms. Molecular Biology and Evolution 35:

1547-1549

Lafontaine JD, Mikkola K (1987) Las-och-nyckel systemen 1

de inre genitalierna av Noctuidae (Lepidoptera) som takson-

omiska kaennetecken. [Lock-and-key systems in the inner

genitalia of Noctuidae (Lepidoptera) as a taxonomic charac-

ter.] Entomologiske Meddelelser 55: 161—167

Prozorov AV, Zolotuhin VV (2012) Seven new monotypic gen-

era of African Lasiocampidae (Lepidoptera). Zoologicheski

Zhurnal 91 (8): 950-960

©ZFMK

Bonn zoological Bulletin 71 (1): 23-28

2022 Cortés-Fossati F.

https://do1.org/10.20363/BZB-2022.71.1.023

ISSN 2190-7307

http://www.zoologicalbulletin.de

Research article

urn:|sid:zoobank.org:pub:C9BAD342-B492-42FD-B7CD-E810D76A82AF

Assessing the distribution of the Andalusian endemic

Berberomeloe payoyo Sanchez-Vialas et al., 2020 (Coleoptera: Meloidae),

with comments on its ecology

Fernando Cortés-Fossati

EcoEvo Group, Area of Biodiversity and Conservation. Universidad Rey Juan Carlos, c/ Tulipan s/n,

E-28933, Mostoles, Madrid, Spain

Email: fernando.cfossati@urjc.es

urn:|sid:zoobank.org:author:F3 D8E2DD-58F0-41 16-80D4-A6A9248464B6

Abstract. Thanks to recent molecular studies, it was revealed that one of the biggest European beetles, the red-stripped

oil beetle Berberomeloe majalis (Linnaeus, 1758) (Coleoptera: Meloidae) comprised a complex of nine different species.

To improve the knowledge of this threatened group, chorology of the newly described microendemic B. payoyo Sanchez-

Vialas et al., 2020 is updated in pursuit of a better knowledge and future adequate conservation strategies. 252 field

sampling events were performed between 2012-2021. 668 specimens were studied. In the framework of the study, two citi-

zen science projects were carried out. The first detailed distribution map and notes about the species’ biology are provided.

Keywords. Biodiversity, blister beetles, entomology, Iberian endemism, insect conservation.

INTRODUCTION

Berberomeloe Bologna, 1989 is an endemic genus from

the Ibero-Maghrebian region and includes some of the

biggest European, which can reach a body length of up

to 7 cm (Bologna 1989; Garcia-Paris 1998; Cortés-Fos-

sati 2018a). This genus has remained unchanged for

thirty years including two species, the red-stripped oil

beetle Berberomeloe majalis (L., 1758) and B. insignis

(Charpentier, 1818). Recent molecular studies focused on

B. majalis revealed a complex of nine morphologically and

phylogenetically distinct species (Sanchez-Vialas et al.

2020). Among these nine species the new south-western

Iberian microendemic B. payoyo Sanchez-Vialas et al.,

2020 (Fig. 1) was described, being distributed in Cadiz

province, Malaga, and Granada provinces (Andalusia,

Spain) (Sanchez-Vialas et al. 2020). Currently, there are

hardly any data and only scarce studies available about

the natural history and biology of B. payoyo (Cortés-Fos-

sati 2018a, b; Cortés-Fossati & Cervera 2018; Sanchez-

Vialas et al. 2020). However, this information 1s crucial

to design appropriate and specific conservation strategies.

In this case — even though no Spanish oil beetle species 1s

protected under any legal framework — it was important

to consider that the only catalogued species within the

genus, Berberomeloe insignis, was classified as vulnera-

ble (VU) in the Andalusian and Spanish Red Book of In-

vertebrates (Barea-Azcon et al. 2008; Verdt et al. 2011).

The species is affected by several impacts, mainly related

with environment transformation (Garcia-Paris & Ruiz

Received: 08.03.2021

Accepted: 02.03.2022

2008, 2011). These threats are observed for B. payoyo

in some areas (Cortés-Fossati 2018a, b; Cortés-Fossati

and Cervera 2018, sub B. majalis) but the information

we have on the species is outdated and scarce , including

its distribution range. Up-to-date distribution information

and a georeferenced map are key tools in conservation

(D’Amen et al. 2013; Della Rocca et al. 2020), especially

in the cases of complexes of closely related species. This

could reveal potential cases of sympatry or allopatry and

clarify the geographic distribution ranges of each of the

species of the B. majalis complex, improving knowledge

and contributing to decrease the Wallacean shortfall, 1.e.,

the lack of knowledge about the geographical distribu-

tion of species (Lomolino 2004; Cardoso et al. 2011).

Therefore, the aim of this study is to provide a better un-

derstanding of the chorology and ecology of the Iberian

B. payoyo.

MATERIAL AND METHODS

The dataset has been generated by 1) own sampling cam-

paigns; 2) citizen science programs, and 3) available lit-

erature, from the oldest reported observation to the most

recent, covering the period 1921-2021.

1) Field sampling campaigns were carried out from

2012 to 2021, from January to August to safely cover the

time interval in which imagos of the species are present

in the field (from March to July in southern Spain: Bolo-

gna, 1989). Specimens were identified according to San-

Corresponding editor: D. Ahrens

Published: 08.03.2022

24 Fernando Cortés-Fossati

chez- Vialas et al. (2020). The species presents distinctive

morphological and biogeographical characteristics that

allowed all individuals to be identified (Sanchez- Vialas

et al. 2020). The first is that the taxonomic identity of the

species 1s very clear in most of the area it occupies, be-

ing restricted to the south-west of the Iberian Peninsula.

Secondly, B. payoyo presents among other characteristics

(for more details, see identification key and description in

Sanchez-Vialas et al. 2020) narrow, coloured post-tergal

bands (Fig. 1A), poorly impressed punctures on the head

(Fig. 1B), and anterior angles of the pronotum are not ex-

panded (Fig. 1C). These features differentiate B. payoyo

from the other species of the complex present in eastern

Andalusia (B. indalo Sanchez-Vialas et al., 2020 and

B. tenebrosus Sanchez-Vialas et al., 2020). Specimens

that could not be reliably determined based on their mor-

phology, as happened in four cases, were excluded from

the study. 252 samplings events were carried out in 86

different sampling points. 668 specimens were studied

in the field in 29 different locations. All populations /re-

cords were georeferenced. The number of individuals,

colouration of post-tergal bands, and biological/be-

havioural observations were also noted.

2) Additional observations were received by the au-

thor by mail, or were uploaded by citizens to two dif-

ferent Citizen Science Programs in which the author

was administrator (“Proyecto Biodiversidad de Anda-

lucia”, hosted in collaboration with iNaturalist.org and

the “Proyecto Meloidata”, hosted in collaboration with

Observation.org), and were also included in the database.

Likewise, the veteran Spanish nature conservation asso-

ciation Biodiversidad Virtual (biodiversidadvirtual.org)

provided their observations concerning the species from

its internal database. Only those observations were con-

sidered, in which the specimens could be reliably identi-

fied from photographs that had georeferencing and date.

3) The only seven publications that provided loca-

tion data were considered for this study (Bologna 1989;

Garcia-Paris 1998; Pérez-Moreno et al. 2003; Garcia-

Paris et al. 2003; Percino-Daniel et al. 2013; Cortés-Fos-

sati 2018, Sanchez-Vialas et al. 2020). In most cases, due

to the age of these works, there was not a very high spa-

tial precision in them, so the locations have subsequently

been referenced estimating their position with the data

provided in these works. On the other hand, in works be-

fore 2020, only locations have been considered in which,

according to Sanchez- Vialas et al. (2020), the taxonomic

identity of the species B. payoyo is clear, that is, the prov-

ince of Cadiz, and certain areas of Malaga and Granada.

The resulting distribution map was generated with

software QGIS ver. 3.16.1-Hannover (GIS Develop-

ment Team 2020) using a EPGS:4326-WGS 84 co-

ordinate system and the layers “Terrestrial 10x10 km

grid” from Ministerio para la Transicion Ecologica y el

Reto Demografico (miteco.gob.es) and “Lineas limite

Bonn zoological Bulletin 71 (1): 23-28

provinciales” from the Instituto Geografico Nacional

(centrodedescargas.cnig.es). In a complementary way,

notes about ecology and conservation of the populations

studied were registered.

RESULTS & DISCUSSION

An updated chorology of the species is provided based on

154 different locations (Appendix: Table S1), allowing

to establish a well-represented distribution map, since a

large part of its potential distribution area has been cov-

ered (Fig. 2). This total number has been formed by data

from fieldwork, citizen science and literature. During

field sampling, the species was present in 29 locations

of 86 sampled, belonging to 14 different municipal dis-

tricts. Citizen science programs functioned satisfactorily,

obtaining 84 valid observations from this source, belong-

ing to 84 different locations. It could be noted that, es-

pecially in rural areas, villagers generally have a good

knowledge about genus Berberomeloe, a phenomenon

already observed for the Cadiz region (Cortés-Fossati

2018a, Cortés-Fossati & Cervera 2018) and for the Ibe-

rian Peninsula (Garcia-Paris et al. 2016) probably due to

the ethnopharmacological uses that the hemolymph of

these animals has historically developed (Percino-Daniel

et al. 2012). This fact may be the main reason why this

program has given reliable results. Finally, the literature

review provided 41 locations, only eight of them georef-

erenced.

According to previous literature (Bologna 1989;

Garcia-Paris 1998; Cortés-Fossati 2018a, b) Berber-

omeloe payoyo is a diurnal xerophilous species, inhab-

iting diverse Mediterranean habitats (see Sanchez-Vialas

et al. 2020), being also distributed in suburban landscapes

with degraded vegetation (Cortés-Fossati 201 8a, b). Nev-

ertheless, the species seemed to have preference on open

areas, steppe lands and meadows with presence of bare

soil patches and not very dense, pioneer vegetation where

females oviposit (Cortés-Fossati 2018; Fig. 3). Here, its

presence can be massive (up to 200 individuals registered

in 0.27 ha ina single day). During evening time individu-

als have been observed on several occasions to leave the

foraging area to hide under leaf litter or grass in areas of

denser vegetation where they spend the night (pers. obs.).

Adults are present in the field from the end of February

to mid-July, being very rare outside this period (e.g., one

single adult record in January cited by Sanchez-Vialas

et al. 2020). Highest abundance occurred between end

of March and end of May (Fig. 4). Earliest oviposition

was observed at the end of March (03/21/2017 in Puerto

Real). Offspring hatched between 20 and 26 days later

(three replicates carried out in March 2014, March 2017,

and May 2017 in the laboratory in translucent 10L terrar-

iums in sand substrate with natural conditions of light,

humidity, and temperature). Characteristic of first instar

©ZFMK

Assessing distribution and ecology of B. payoyo 25

Fig. 1. Berberomeloe payoyo Sanchez-Vialas et al., 2020. A. Imago from Puerto Real, Cadiz. Dorsal view. B. Pronotum. C. Head

front view, with characteristic punctures. D. First instar larva in dorsal (1), ventral (II) and lateral view (III). Instars were damaged

during collection, lacking their terminal long chaetae, partially visible in III.

| ee

e © e

5 e

hy e

Aa e . ef =

OG e

e e t )

e 3 . q e

e e

\e e

)

e e

e? ee

e e o @ ¢@

aI e

e © 0 25 50 km

sd _—_k———_6—C—C—C~S~S

Fig. 2. Distribution of Berberomeloe payoyo composed with data from sampling campaigns, citizen science records and literature

records.

Bonn zoological Bulletin 71 (1): 23-28 ©ZFMK

26 Fernando Cortés-Fossati

Fig. 3. Typical environment where the species seems to have

preference, consisting of open areas with no dense vegetation

(Medina Sidonia, Cadiz).

larva (Fig. 1D) corresponds to the description made by

Bologna (1989) for the larvae from the B. majalis com-

plex, not observing appreciable differences.

Based on the findings, it 1s also possible to conclude

the absence of this species in the coastal strip and marshy

ecosystems, at least, in tidal influence areas and beach

sand substrate (after 113 samplings and studying all the

data provided by third parties, absent in the tidal zone

and marshes of Algeciras, Barbate, Cadiz, Chiclana de

la Frontera, Chipiona, Conil de la Frontera, Puerto Real,

Puerto de Santa Maria, Sanlucar de Barrameda, Tarifa).

J F M A

As expected, no sympatry cases with other Berberomeloe

species were detected in Cadiz, but a genetic study of the

possible contact areas is necessary, in this case, the limit

zone of distribution in the east, since there may be cases

of introgressive hybridization that do not allow the in-

dividuals to be correctly identified morphologically, and

even, hybrid populations may be present.

Intraspecific variation of the populations was detected,

in terms of the post-tergal band colouration, which can

range from light orange to reddish orange and even ver-

milion red, the orange-coloured form being probably the

most common in Cadiz province (Appendix: Table S2.,-

Fig. S2). No populations with mixed specimens of dif-

ferent band colours have been detected. No completely

black morphotype specimens were observed.

According to personal observation, imagos feed on

leaves of different pioneer and herbaceous plants species

as well as young shoots. However, they seem to show

preference on flower petals of certain plants species

such as those of the genus Echium L. (Boraginaceae).

Throughout this study, also repeated feeding on diverse

Asteraceae (highlighting Carduus L., Galactites Mo-

ench, and Pallenis Cass), and on Oxalis pes-caprae L.

(Oxalidaceae) has been observed. Berberomeloe payoyo

has also been observed feeding on Erodium L’ Her. ex

Aiton (Geraniaceae), Plantago L. (Plantaginaceae) and

Lysimachia (L.) U. Manns and Anderb. (Myrsinaceae).

150

100 +5

rot

M J J A

Fig. 4. Phenology of imago of Berberomeloe payoyo, built from the database generated for the study. Literature or citizen science

data that lacked the number of observed individuals by date have been plotted as n= 1.

Bonn zoological Bulletin 71 (1): 23-28

©ZFMK

Assessing distribution and ecology of B. payoyo pa:

No studies have been conducted on the conservation

status of the species. However, data collected during

this work suggest that populations near cities undergo

a negative anthropogenic pressure. The ecosystems of

the Cadiz and Malaga provinces are highly transformed

and fragmented, except for some well-preserved natural

cores such as Grazalema, Los Alcornocales or Sierra de

las Nieves Natural Parks. Agriculture and an aggressive

urbanistic process occupies most of the area, increasing-

ly taking ground in its development over the years. In

this way, in Cadiz it is very frequent to observe popula-

tions of B. payoyo located on croplands and peri-urban

regions with a clear degree of deterioration (Cortés-Fos-

sati 2018a). Totally or quasi-isolated populations due to

human infrastructures have been observed (e.g., diverse

populations in Chiclana de la Frontera, Puerto Real, Jer-

ez de la Frontera, Sanlucar de Barrameda) since the spe-

cies has a very low dispersal rate and a low capacity to

colonize new areas (Garcia-Paris 1998; Sanchez-Vialas

et al. 2020). Use of steppe lands or zones dominated by

colonizing pioneer r-strategist plant species as uncon-

trolled waste dumps, urban pressure, and the agricultural

fields as well as the use of chemicals are probably af-

fecting populations, threats already proposed as impacts

for other Meloidae (Garcia-Paris & Ruiz 2008a, b, 2011;

Ruiz & Garcia-Paris 2008a, b). In these zones, popula-

tions seem degraded and with lower population densities.

Also, road kills have been observed very frequently, a

phenomenon also observed for the Madrilenian Berber-

omeloe populations (Garcia-Paris et al. 2006; pers. obs.).

Even possible local extinction of some populations has

been observed during this work. In 2012, an expansion

to a mega commercial area was carried out in Jerez de

la Frontera (Area Norte and Area Sur, see Appendix: Ta-

ble S1). The species has no longer been observed there

since 2016, also not appearing again in the adjoining crop

fields. Moreover, a population located in a cropland in

Torre del Puerco (see Appendix: Table S1), completely

disappeared in 2017 after changes in land use in the area.

However, geographical situation of this population sug-

gests it could be derived from another nucleus, located on

a semi-natural shrubland in front of the sampling point,

found behind several linear structures that fragment the

area (two walkways and a road). So, if transformation

will not continue, there could be a possibility that the

beetle will return to the area. Finally, during 2017, in

Puerto Real: Parque Entrevias (see Appendix: Table S1),

the species apparently disappeared after diverse human

interventions, e. g., illegal deposit of debris and the in-

stallation of a pipeline that flooded the study area. As of

2018, a large decline in number of individuals was ob-

served. After that season, no specimen has been observed

in the area. However, there are no data on dynamic pop-

ulations that allow calculating possible regressions or the

real status of the species, so it would be necessary to car-

ry out studies focused on conservation ecology.

Bonn zoological Bulletin 71 (1): 23-28

CONCLUSIONS

Berberomeloe payoyo Sanchez-Vialas et al., 2020 is a

microendemic restricted to the south-west of the Iberian

Peninsula that is well represented throughout its area of

distribution. Information collected provides an update on

the distribution of B. payoyo and some ecological notes

that improve the knowledge about the species. The infor-

mation regarding its ecology and biological cycle seems

to fit very well to what was already stated by Bologna

(1989) when the genus was described (Bologna 1989;

Garcia-Paris 1998; Garcia-Paris et al. 2003; Cortés-Fos-

sati 2018). At least, peri-urban populations seem to suffer

from negative human impacts, e.g., habitat fragmentation

and change in land use. Future works on the conservation

ecology of the species are needed.

Acknowledgments. Thanks to the reviewers and journal ed-

itors for their constructive feedback. To Consejeria de Medio

Ambiente y Ordenacion Del Territorio of Junta de Andalucia

government for allowing this research providing their support

and required permissions. This research could not be possible

without field assistants Irene Martin-Rodriguez, Juan Manuel

Mufioz-Ocafia, Alberto Leandro Aranda Quirés and Fernando

Cortés Marquez. I would like to thank professors Juan Lucas

Cervera, Marco Bologna, Gonzalo Mufioz, and Marcos Mén-

dez for their invaluable suggestions. I also appreciate the help

of Julio Rabadan and Pablo Alvarez to manage, create, and host

the citizen science project “Meloidata” on Observation Spain. I

would like to express my gratitude to PhD Jazmin Deneb Orti-

gosa for co-creating and co-directing the citizen science project

“Biodiversity of Andalusia” hosted on iNaturalist California

Academy of Sciences. Also, to Biodiversidad Virtual team for

allowing to use their database. Thanks to PhD student Pablo

Escribano for arranging laboratory material for this research.

Finally, thanks to all entomologists, naturalists, villagers, citi-

zen scientists and students that kindly and decisively contribut-

ed to this work.

Ethical & legal aspects. This study was framed under the legal

regulations and laws for the collection and ethical treatment of

animals and has been endorsed by Andalusian government.

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APPENDIX I

(electronic supplement, available at www.zoologicalbulletin.de)

Table S1. Location dataset composed by data from field work

(F), citizen science programs (CS) and Literature (L: authors,

year). Dates indicated in intervals pertaining to the field cam-

paings represent a sustained work in the period between Janu-

ary and August of those years.

Table S2. List of specimens studied in the field in which the

color of the banding was noted. The observations are divided

between two classes, Orange (O) which is represented on the

map with orange dots and Red (Red) represented on the map

with red dots. Only those individuals from the sampling cam-

paigns were included in the study as the colors could not be

standardized from photographs provided by third parties.

Fig. S2. Mapping of specimens studied on field divided by co-

louration in the post-tergal banded pattern. The observations

are divided between with orange dots (populations with orange

bands and black dots (populations with red bands) represented

on the map with red dots. Only those individuals from the sam-

pling campaigns and Cortés-Fossati, 2018 were included in the

study as the colors could not be standardized from photographs

provided by third parties. According to citizen science data,

which have not been taken into account in the plot because it

cannot be corroborated — since despite having testimonies and

photos the colors have not been normalized with a scale — the

red individuals would be confined to the northern part of the

province and the southern part of the same Strait Zone), leav-

ing the central strip of the province dominated by populations

of orange coloration, thus being probably the most widespread

coloration.

©ZFMK

Table S1. Location dataset composed by data from field work (F), citizen science programs (CS) and Literature (L: authors, year). Dates indicated in intervals pertaining to the

field campaings represent a sustained work in the period between January and August of those years.

ey et EE AS Sg ee ee PS SPE.

Alcala de los Gazules

Algeciras

Arcos de la Frontera

Barbate

Chiclana de la Frontera

Conil de la Frontera

Jerez de la Frontera

Linea la Concepcién

Los Barrios

Medina Sidonia

Paterna de Rivera

Puerto Real

El Algibe

Sierra de la Plata

Arroyo del Salado

Sierra del Retin

East peri-urban area

South-West peri-urban area

Torre del Puerco

Fuente del Gallo

Area Sur Shopping centre

Arroyo Canaleja, South stretch

Arroyo Canaleja, East stretch

Arroyo de la Loba

Campifia del Area Norte

Campifia del Area Sur

Cartuja Santa Maria Defension

El Pedroso

Laguna de Medina

Zoological Park parkings

Sierra Carbonera

La Teja, Alcornocales Nat. Park

Arroyo del Azucar

Arroyo Golondrina/Croplands

Colada de la Espartera

Croplands near Saltillo

Croplands near Saltillo (2)

Arroyo Pozomedina/ croplands

Peri-urban area

36.44570,

36.11251,

36.74120,

36.17378,

36.41881,

36.41452,

36.33004,

36.29054,

36.69224,

36.67779,

36.67972,

36.69169,

36.68620,

36.69996,

36.65135,

36.52443,

36.61580,

36.69006,

36.19398,

36.28370,

36.46683,

36.42423,

36.43629,

36.45661,

36.46475,

36.52451,

36.53215,

-5.74695

-5.77617

-5.85514

-5.84883

-6.12198

-6.14204

-6.16010

-6.10900

-6.15621

-6.09269

-6.09339

-6.16478

-6.17414

-6.15733

-6.10955

-5.9869 1

-6.06087

-6.15332

-5.35895

-5.57462

-5.91008

-5.90806

-6.02801

-5.94742

-5.95086

-5.87369

-6.17407

106

2016-2017

2018-2019

22.05.16

2018-2019

09.04.17

2016-2017

01.05.13

2013-2019

2015-2017

2016-2017

2013-2019

09.05.14

2016-2017

17.03.17

2014-2019

2018-2019

07.04.17

01.06.17

2017

24.05.17

2016-2017

2016-2018

19.05.17

2017

CADIZ

Trebujena

Vejer de Frontera

Alcala de los Gazules

Algeciras

Barbate

Benalup-Casas Viejas

Castellar de la Frontera

Chiclana de la Frontera

Chipiona

Espera

Jerez de la Frontera

Jimena de la Frontera

Los Barrios

Manzanete

Medina Sidonia

Puerto Santa Maria

Puerto de Santa Maria

South croplands

Arroyo Montecote/ Croplands

Embalse de Barbate

Near Mirador de los Tallones

Urban zone

Near Arroyo Mondragon

Embalse del Celemin

Puerto Juan Alonso/Guadarranque

Sierra Montecoche

Cafiada Picapollos

El Marquesado

Pago de Humo

Pinar del Hierro

Urbanizacion Costa Ballena

Cafiada Jerez-Utrera

Near Arroyo de la Gallina

Laguna de Medina

Laguna del Tején

Las Canteras y el Tejon

Las Canteras el Tej6n Croplands

Arroyo de los Hoyones

Rio Hozgarganta

Embalse de Almodovar

Embalse Zanona

Near Garganta del Cabrero

Near Garaganta Curtidora

Sendero Palancar

Manzanete

Croplands

El Martillo Industrial area

Peri-urban croplands

Rancho Linares

36.85740, -6.19088

36.26264, -5.97657

36.39830, -5.70650

36.46497, -5.64809

36.14075, -5.45623

36.21585, -5.93205

36.30733, -5.72880

36.28233, -5.53112

36.45983, -6.11653

36.46005, -6.11715

36.40732 -6.08526

36.38523, -6.12780

36.70557, -6.41539

36.86150, -5.86470

36.59469, -5.57690

36.62210, -6.05190

36.57407, -6.07349

36.57845, -6.06554

36.58476, -6.06608

36.45684, -5.47904

36.45979, -5.47008

36.15347, -5.63273

36.24179, -5.67973

36.25150, -5.48990

36.25330, -5.47260

36.24939, -5.55560

36.43572, -5.90323

36.62448, -6.19817

2016-2017

01.06.17

14.06.18

19.04.19

03.04.17

29.04.17

26.05.15

22.04.18

12.04.21

18.04.21

23.05.18

19.05.18

26.05.18

20,0507

29.05.17

26.05.15

16.05.15

12.06.13

30.03.17

03.04.20

20.04.13

CADIZ

Puerto de Santa Maria

Puerto Real

San José del Valle

San Roque

Tarifa

Torre Alhaquime

Ubrique

Vejer de la Frontera

Sierra San Crist6bal

Arroyo del Castafio

Arroyo de la Zarza

Arroyo de las Yeguas

Camino Torrealta

Camino Los Barreros

East peri-urban forested area

Forested-shrubby area

La Chacona

Peri-urban cropland

Arroyo del Palmetin

El] Acebuchal

Peri-uban ring

Estaci6n Ambiental Madrevieja

Monte bajo

Peri-urban area

Pinar del Rey

Sierra Carbonera

Arroyo del Molino

Arroyo Cagancha

Camino de la Pista

Campifia Tarifefia

Cerro de Bartolo

Near Cortijo Aguilon

Embalse de Almodovar

Pico Luna

Rio de la Jara

Sierra la Plata

Abandoned cropland

Los Alcornocales Natural Park

South peri-uban area

Near Arroyo Donadillo

36.63250, -6.15120

36.54700,-6.08969

36.52619, -6.15571

36.53163, -6.16760

36.55845,-6.11097

36.55426,-6.08621

36.52646, -6.11869

36.53978,-6.10563

36.63580, -5.71410

36.59244, -5.76379

36.60721, -5.80027

36.20270, -5.40670

36.20974, -5.38457

36.20403, -5.69441

36.15347, -5.63273

36.06480, -5.64840

36.093 13, -5.72697

36.12480, -5.70490

36.15057, -5.65305

36.10292, -5.54157

36.05870, -5.63700

36.10799, -5.78828

36.89050, -5.25950

36.641672, -5.473934

36.27180, -5.93740

115

07.03.21

29.05.17

16.04.21

22.03.10

29.05.17

30.03.18

29.05.17

23.04.16

14.05.17

25.04.15

07.04.09

27.03.85

01.04.82

16.05.15

08.05.21

05.05.08

01.05.17

10.05.18

20.05.17

23.04.19

29.04.18

01.05.17

28.04.12

01.05.17

26.05.18

L: Bologna, 1989

L: Garcia-Paris, 1998

: Garcia-Paris et al., 2003

Garcia-Paris et al., 2003

: Garcia-Paris et al., 2003

L: Pérez-Moreno et al., 2003

L: Pérez-Moreno et al., 2004

L: Pérez-Moreno et al., 2003

L: Percino-Daniel et al., 2013

L: Cortés-Fossati, 2018

L: Sanchez-Vialas et al., 2020

et eee

CADIZ

Vejer de la Frontera

Villaluenga del Rosario

Algeciras

Alcala de los Gazules

Jerez de la Frontera

San Roque

Tarifa

Ronda

San Roque

Tarifa

Benalup-Casas Viejas

Chiclana de la Frontera

Vejer or Algodonales?

Medina Sidonia

Puerto Real

Sanlucar de Barrameda

Conil de la Frontera

Facinas (Barbate)

Near el Ciruelo

Near Rio Barbate

Peri-urban area

Bay of Cadiz Area, see Fig. 10

La Janda Area, see Fig. 10

Bay of Algeciras, see Fig. 10

N Bay of Cadiz Area, see Fig. 1

S Bay of Cadiz Area, see Fig. |

Bay of Algeciras, see Fig. 1

Laguna de Medina

Sierra de las Nieves

Benalup de Sidonia

La Muela

3km S Alcala de los Gazules

Arroyo del Saltillo

East croplands

Parque de Entrevias

Las Dunas industrial area

Fuente del Gallo

Facinas

36.235 10, -6.04980

36.23580, -5.93970

36.69740, -5.38767

36.45757, -5.94340

36.53127, -6.16051

36.53636, -6.19546

36.75019, -6.34113

36.29691, -6.11925

36.12633, -5.70572

870

13.05.18

23.05.12

05.05.08

21.04.21

23.04.21

24.04.21

16.03.85

27.03.83

00/05/1973

00/04/1971

10.04.87

00/05/1987

00/05/1971

20.03.89

2016-2018

2014-2021

04.06.17

13.04.13

19.06.16

: Sanchez-Vialas et al., 2020

L: Sanchez-Vialas et al., 2020

L: Garcia-Paris, 1998

Cotikeiien ke stem acne

CADIZ

MALAGA

Manzanete

Paterna de Rivera

San Roque

Alhaurin el Grande

Antequera

Campillos

Ciudad Jardin

Cortes de la Frontera

Fuente de Piedra

La Roda de Andalucia

Malaga

Ojén

Parauta

Palma-Palmilla

Puerto de la Torre

Sierra de Yeguas

Tolox

Yunquera

Montecorto

Sierra del Retin

2 km S Benalup Sidonia

7 km SW Benalup Sidonia

4km NE San José del Valle

3km S Alcala de los Gazules

Shrubby area

Acuartelamiento Bobadilla

Cropland

Sub-urban area

Laguna Redonda

Arroyo Hondo

Near Jardines de San Telmo

Loma de los Pinos

Sub-urban area

Mirador la Vicaria

Arroyo las Cafias

Sub-urban area

Near Area Recreativa Conejeras

Shrubby area

Camino Cupiana

Near Camino de Campillos

Mirador el Cuco

Mirador Luis Ceballos

Near Puerto Saucillo

Parque Natural Sierra las Nieves

Sierra de las Nieves

Near Malaga, see Fig. 1

36.20000,

-5.80000

36.62300,

37.03120,

37.06300,

37.04603,

37.03010,

36.77359,

36.76744,

36.56040,

37.13343,

37.13619,

36.72195,

36.722323,

36.56567,

36.663 16,

36.76606,

36.75087,

37.107585,

36.71058,

36.71734,

36.71040,

36.71142,

-4.70160

-4.74121

-4.71700

-4.85733

-4.84400

-4.38472

-4.41804

-5.39080

-4.72823

-4.75720

-4.48979

-4.485513

-4.85609

-5.09306

-4.44250

-4.49657

-4.829026

-4.98980

-4.96470

-4.96685

-4.96696

04.01.18

24.03.11

21.05.17

19.04.01

00/05/1973

29.04.07

14.04.15

25.05.19

05.04.12

03.04.07

04.04.15

04.03.15

15.07.19

06.04.08

13.05.11

22.07.17

19.03.21

14.03.16

25.05.13

03.05.14

26.03.16

25.04.91

L: Pérez-Moreno et al., 2004 MALAGA Sierra de Ronda NA NA NA 08.04.82

L: Sanchez-Vialas et al., 2020 GRANADA Santa Cruz Comercio NA NA NA NA

Table S2. List of specimens studied in the field in which the color of the banding was noted. The observations are divided between two classes,

Orange (O) which is represented on the map with orange dots and Red (Red) represented on the map with red dots. Only those individuals from the

sampling campaigns were included in the studv as the colors could not be standardized from photographs provided bv third parties.

oo gan eae eee ioe |r Bal

Fy, “Heer Fi. ar]

oy PE “a ee

mole aae he aes

Alcala de los Gazules

Algeciras

Arcos de la Frontera

Barbate

Chiclana de la Frontera

Conil de la Frontera

Jerez de la Frontera

Linea la Concepcion

Los Barrios

Medina Sidonia

El Algibe

Sierra de la Plata

Arroyo del Salado

Sierra del Retin

East peri-urban area

South-West peri-urban area

Torre del Puerco

Fuente del Gallo

Area Sur Shopping centre

Arroyo Canaleja, South

stretch

Arroyo Canaleja, East stretch

Arroyo de la Loba

Campifia del Area Norte

Campifia del Area Sur

Cartuja Santa Maria

Defension

El Pedroso

Laguna de Medina

Zoological Park parkings

Sierra Carbonera

La Teja, Alcornocales Nat.

Park

Arroyo del Azucar

Arroyo Golondrina/Croplands

36.44570, -5.74695

36.11251, -5.77617

36.74120, -5.85514

36.17378, -5.84883

36.41881, -6.12198

36.41452, -6.14204

36.33004, -6.16010

36.29054, -6.10900

36.69224, -6.15621

36.67779, -6.09269

36.67972, -6.09339

36.69169, -6.16478

36.68620, -6.17414

36.69996, -6.15733

36.65 135, -6.10955

36.52443, -5.98691

36.61580, -6.06087

36.69006, -6.15332

36.19398, -5.35895

36.28370, -5.57462

36.46683, -5.91008

36.42423, -5.90806

Pee OY FONG S. OC Oe, OOS oe: Con 2S

COR

L: Cortés-Fossati, 2018

CADIZ

Medina Sidonia

Paterna de Rivera

Puerto Real

Trebujena

Medina Sidonia

Puerto Real

Sanlucar de Barrameda

Colada de la Espartera

Croplands near Saltillo

Croplands near Saltillo (2)

Arroyo Pozomedina/

croplands

Peri-urban area

South croplands

Arroyo del Saltillo

East croplands

Parque de Entrevias

Las Dunas industrial area

36.43629, -6.02801

36.45661, -5.94742

36.46475, -5.95086

36.5245 1, -5.87369

36.53215, -6.17407

36.85740, -6.19088

36.45757, -5.94340

36.53127, -6.16051

36.53636, -6.19546

36.75019, -6.34113

BAO-0 OC OO 6°50

Assessing the distribution of the Andalusian endemic Berberomeloe payoyo Sanchez-Vialas

et al., 2020 (Coleoptera: Meloidae), with comments on its ecology

Fernando Cortés-Fossati

Appendix Fig. S2

Fig. S2. Mapping of specimens studied on field divided by colouration in the post-tergal banded pattern.

The observations are divided between with orange dots (populations with orange bands and black dots

(populations with red bands) represented on the map with red dots. Only those individuals from the

sampling campaigns and Cortés-Fossati, 2018 were included in the study as the colors could not be

standardized from photographs provided by third parties. According to citizen science data, which have not

been taken into account in the plot because it cannot be corroborated — since despite having testimonies and

photos the colors have not been normalized with a scale — the red individuals would be confined to the

northern part of the province and the southern part of the same Strait Zone), leaving the central strip of the

province dominated by populations of orange coloration, thus being probably the most widespread

coloration.

BHL

Blank Page Digitally Inserted

Bonn zoological Bulletin 71 (1): 29-39

2022 Schonfeld J. & Rohwedder D.

https://do1.org/10.20363/BZB-2022.71.1.029

ISSN 2190-7307

http://www.zoologicalbulletin.de

Research article

urn:|sid:zoobank.org:pub:0835A D8F-BFDE-407E-AC2F-05 DABEE4C9D4

Type specimen designation for

Onthophagus (Colobonthophagus) aenescens (Wiedemann, 1823) and

O. (Colobonthophagus) urellus Boucomont, 1920

with notes on the synonymy (Coleoptera: Scarabaeidae: Scarabaeinae)

Joachim Schonfeld!" & Dirk Rohwedder?

'Erlenweg 9, D-53489 Sinzig, Germany

?Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany

“Corresponding author: Email: Joachim_Schoenfeld@web.de

'urn:lsid:zoobank.org:author:D7A02B6F-7C87-4F47-929F-B | DOE2FFFBD3

2urn:Isid:zoobank.org:author:09BBOBES-E5F7-4654-BC85-C077D090A B95

Abstract. The taxonomic position of four species of Onthophagus (Colobonthophagus) Balthasar, 1963 was studied.

The neotype of O. (C.) aenescens (Wiedemann, 1823) is designated here, with O. (C.) volucer Balthasar, 1959 as a junior

subjective synonym of this species. Furthermore, O. (C.) usurpator Balthasar, 1960 was found to be a junior subjective

synonym of O. (C.) urellus Boucomont, 1920. The lectotype and two paralectotypes of O. (C.) urellus Boucomont, 1920

are designated. We show the first record of O. (C.) urellus of Nepal and the Palaearctic region.

Key words. Taxonomy, types, new synonymy, neotype designation, lectotype designation, Onthophagini, Oriental Regi-

on, Palaearctic Region.

INTRODUCTION

Onthophagus (Colobonthophagus) aenescens (Wie-

demann, 1823) was originally described as Copris ae-

nescens based on five male specimens from Bengal

(Wiedemann 1823: 15). During the 1943 bombardment

of Hamburg the original type series was lost (Abraham &

Husemann pers. comm.). Weidner (1976: 87) stated: “Die

Coleopterensammlung des Zoologischen Museums Ham-

burg wurde bis auf die in Alkohol aufbewahrten Larven

1943 beim Brand des Museums vollstandig vernichtet.”

— (translated from German: The collection of beetles at

the Zoological Museum of Hamburg was completely de-

stroyed in the fire at the Museum in 1943, except for the

larvae preserved in alcohol. ).

The original names Copris aenescens Wiedemann,

1823 (p. 177, Nr. 694/695) and Onthophagus aenescens

(p. 232, Nr. 1624/1625) are not recorded in the catalogue

*Verzeichnis der Typen und Typoide“ (Weidner 1976:

134).

According to Evenhuis (2008) and the rules of ICZN

(1999), a neotype should be designated if the original

type series is lost or damaged and designation of the neo-

type is necessary for clarification of taxonomic status of

the species. The position of O. aenescens in the subgenus

Colobonthophagus Balthasar, 1963 (Scheuern, 1995) and

the status of Onthophagus (Colobonthophagus) volucer

Received: 28.11.2020

Accepted: 11.03.2022

Balthasar, 1959 syn. nov. as a junior subjective synonym

of O. (C.) aenescens (Wiedemann, 1823) call for the des-

ignation of the neotype for the correct application of the

specific name.

After studying the holotype of Onthophagus volucer

Balthasar, 1959, and 155 specimens of Onthophagus (C.)

aenescens (Wiedemann, 1823), we are convinced that

this taxon is a junior subjective synonym of O. (Colo-

bonthophagus) aenescens (Wiedemann, 1823). Both taxa

show similar morphology of the head, punctation of the

pronotum, characteristics of elytra and surface of the

body (Fig. 1A—F). The aedeagus and the lamella copula-

trix also show the same structure (Fig. 2A, C). Further-

more, the type localities of both taxa are situated close to

each other (Fig. 6).

We also find O. (Colobonthophagus) usurpator

Balthasar, 1960 to be a junior subjective synonym of

O. (Colobonthophagus) urellus Boucomount, 1920; this

is based on examination of three syntypes of O. (C.) ur-

ellus, seven specimens from South India, one male from

Nepal and the holotype of O. (C.) usurpator. Both taxa

show similarities in all morphological characters of the

genitalia, pronotum and in the punctation. The armatures

of the head are different from other species of the subge-

nus (Figs 3A—F, 4A, C). In the present study we designate

the lectotype (4 major) of Onthophagus (Colobontho-

Corresponding editor: D. Ahrens

Published: 29.03.2022

30 Joachim Schénfeld & Dirk Rohwedder

phagus) urellus Boucomount, 1920 deposited at the

Muséum national d’ Histoire naturelle, Paris (MNHN).

This paper provides an overview of the distributions

of the two Onthophagus species in the Oriental and Pa-

laearctic regions (Middle Asian regions) (Fig. 6).

The revision of the subgenus Colobonthophagus

Balthasar, 1963 by Scheuern (1995, 1996) presents

27 species with keys separating O. (C.) aenescens and

O. (C.) urellus from other species.

MATERIAL AND METHODS

The following acronyms identify the collections housing

the examined material (curators in parentheses):

cAS = André Skale, Gera, Germany

cAW = Andreas Weigel, Wernburg, Germany

cAK = Andreas Kopetz, Eischleben, Germany

cAN = Alexander Napolov, Riga, Latvia

cDJ = Dieter Jungwirth, Ingolstadt, Germany

cJS = Joachim Schonfeld, Sinzig, Germany

cOH = Oliver Hillert, Schéneiche near Berlin,

Germany

cPS = Paul Schoolmeesters, Herent, Belgique

HNHM = Hungarian Natural History Museum,

Budapest, Hungary (Otto Merk)

MFNB = Museum fir Naturkunde Berlin, Germany

(Johannes Frisch)

MHNG = Muséum d’ Histoire Naturelle de Geneve,

Switzerland (Ivan Lobl)

MNHN = Muséum national d’ Histoire naturelle,

Paris, France (Olivier Montreuil)

NHMB = Naturhistorisches Museum, Basel,

Switzerland (Matthias Borer)

NHMD = Natural History MuseumofDenmark,Copen-

hagen, Denmark (Alexey Solodovnikov)

NMEG = Naturkundemuseum, Erfurt, Germany

(Matthias Hartmann)

NMPC = National Museum, Prague, Czech Republic

(Jiri Hajek)

PIME = Pontificio Istituto Missioni Estere

(P.I.M.E.), Entomological Museum,

Memphis, Detroit, Michigan (USA) Coll.

Dr Carlo Brivio

RBINS = Royal Belgian Institute of Natural Scien-

ces, Bruxelles, Belgium (Alain Drumont)

ZFMK = Zoologisches Forschungsmuseum Alexan-

der Koenig, Bonn, Germany (Dirk Ahrens)

ZMH = Zoologisches Museum Hamburg, Germany

(Martin Husemann)

Altogether, 156 specimens of O. (C.) aenescens and

12 specimens of O. (C.) urellus were studied (see ma-

terial listed below). Remarks of the authors and com-

ments are indicated in parentheses. The exact label data

Bonn zoological Bulletin 71 (1): 29-39

are cited for the type material only; individual labels are

indicated (only for types) by double slash (//), individual

lines on every label by single slash (/), [p] = preceding

data within quotation marks are printed, [hw] = preced-

ing data within quotation marks are handwritten.

Taxonomy and distribution

Onthophagus (Colobonthophagus) aenescens (Wiede-

mann, 1823)

Figs 1A—F, 2A—D, 5A-C, 6

Copris aenescens Wiedemann, 1823:13 (original de-

scription).

Onthophagus volucer Balthasar, 1959 [syn. nov.]: 190

(original description).

Onthophagus (s.str.) aenescens. Balthasar 1963: 264 (re-

description).

Onthophagus (s.str.) volucer Balthasar, 1963: 586 (rede-

scription); Bezdek & Hajek 2013:428 (catalogue, sta-

tus); Frey 1973: 102 (redescription, type material).

Onthophagus (Colobonthophagus) aenescens: Scheuern

1995: 416 (stat. nov., key); Mittal 2000: 263 (distribu-

tion); Kabakov 2006: 162 (status); Mittal & Jain 2015:

395 (distribution); Ziani & Bezdek, 2016: 175 (cata-

logue, distribution); Lobl et al. 2006: 163 (catalogue,

distribution).

Type locality of O. (Colobonthophagus) aenescens

(Wiedemann, 1823): Bengalia, Burma.

Type material examined (2 specs)

Onthophagus aenescens

Neotype

BENGALEN ° 3 (NHMD, here designated): “Bengal.

/ Septbr. 1810. [hw] // Zool. Museum DK Copenhagen

[p] / O. (Colobonthophagus) |p] / aenescens (WIED.)

[hw] / J SCHEUERN det.19 [p] 96 [hw] // Onthopha-

gus (Colobonthophagus) |p, fat] // aenescens (WIEDE-

MANN, 1823) [p] // NEOTYPUS © desig. 2022 [p, fat]

// J. SCHONFELD & D. ROHWEDDER Jp, red label]”.

Onthophagus volucer

Holotype

BURMA ° ¢@ (NMPC): “Burma [Myanmar] / Tenass[e-

rin] [hw] // Hoefer [hw] // Mus. Nat. Pragae [p] / 65 845

[hw] / Inv. [p, red label] // Onthophagus / volucer / 3

n.sp. / Balth.[hw] / Holotypus [p, pink label] // O. (Colo-

bonthophagus) |p| / aenescens WIED.) [hw] / J. SCHEU-

ERN det.19 [p] 96 [hw] // volucer BALTH. / Syn.nov. /

J. SCHEUERN det. [p] 96 [hw] // O. (Colobonthopha-

gus) / aenescens |(WIEDEMANN 1823) 4 / J. SCHON-

FELD det. 2017 [p]; // O. (Colobonthophagus) / volucer

/ BALTHASAR 1959/ syn. nov. / J. SCHONFELD det.

20.17 [pl

©ZFMK

Type specimen designation for Onthophagus (Colobonthophagus) aenescens and O. (Colobonthophagus) urellus 31

Fig. 1. A-C. Onthophagus (Colobonthophagus) aenescens (Wiedemann, 1823), neotype, 3. D-F. Onthophagus (Colobonthopha-

gus) volucer Balthasar, 1959 syn. nov., @. A, D. Habitus, dorsal view. B, E. Habitus, fronto-ventral view. C, F. Pronotal sculpture

in dorso-lateral view with close-up of the punctation (C = top; F = sloping). Scale = 5 mm.

Bonn zoological Bulletin 71 (1): 29-39 ©ZFMK

a2 Joachim Schonfeld & Dirk Rohwedder

0.(Colobonthophagus

ACh ESCEUS(GHE

J.SCHEVERN det.19

Tee cey

Lets

volucer BAUN,

{ nM .new —

J. SCHEUERN

Fig. 2. A-B. Onthophagus (Colobonthophagus) aenescens (Wiedemann, 1823), neotype, 4. C—D. Onthophagus (Colobonthop-

hagus) volucer Balthasar, 1959 syn. nov., 3. A, C. Left: Aedeagus, lateral view, raspula, lamella copulatrix, ventral view; right:

Parameres, dorsal view & apical view. B, D. Labels.

Other material examined (154 specs)

BANGLADESH ¢ EAST PAKISTAN [Bangladesh], Di-

najpur, X-1969, Barbe’, Paratype 2, Onthophagus voluc-

er Balt., G. FREY 1972; 12 (NHMB). EAST PAKISTAN

[Bangladesh], Dinajpur, IV-1970, Barbe’, Onthophagus

volucer Balt. 3, G. FREY 1969; 14 (NHMB). Benga-

la: Boldipukur, 6.1964 Grest leg., O. aenescens (WIED.)

det. PITTINO; 26), 19 (cRP).

INDIA ¢ [Nr.]26884, flavicornis meg. Germ., Zool. Mus.

Berlin; 1¢' (MFNB). Assam, Nr. 26884, Zool. Mus.

Berlin; 23, 32 (MFNB). Assam, Nr. 26884, aenescens

Bonn zoological Bulletin 71 (1): 29-39

Wied., West.[ermann], Zool. Mus. Berlin; 14 (MFNB).

Assam, Westermann, Zool. Museum DK Copenhagen;

14 (NHMD), 1 (cJS). Assam, Kaziranga, 10.X1I.1969,

leg. G. PILLERI; 200’, 349 (NMEG), 19 (ZFMK),

14, 19 (cAN), 24, 29 (cJS). [W Bengal] Barbaria; 19

(NHMD). Calcutta, 1938, Coll. Boucomont, Paris; 19

(MNHN). Calcutta, Ex Mus. V. LANSB.[ERGE], Coll.

Boucomont, Paris; 12 (MNHN). Darjeeling D., Sona-

pur, Mahahandra Riv., Bhakta B., 9.-15.1[X.1984; 19

(cJS). KHARI, [near] KASHINAGAR. W-BENGAL.,

leg. 26.1.1950. Rev.S. POLGAR 57, ex coll.: Ch.S.PAPP,

aenescens Wied., 13° (NHMB). Bengale (Mnisrech),

©ZFMK

Type specimen designation for Onthophagus (Colobonthophagus) aenescens and O. (Colobonthophagus) urellus 33

aenescens det. Gillet; 12 (RBINS). Sirampur, Zool.

Museum DK Copenhagen; 13° (NHMD), 19 (cJS).

Sunderbunds, Coll. Boucomont; 19 (MFNB). Ex Mus.

VAN LANSBERGE, ex coll. R. OBERTHUR 1952, det.

aenescens WIED.; 14 (MNHN).

Ex Museo E. HAROLD, aenescens WIED., G.J. ARROW

vidit. 1906, ex Coll R. OBERTHUR 1952; 14 (MNHN).

Coll. J. Thomson, Coll.R.I.Sc.N.B.; 14 (RBINS). Coll.

de Bonneuil, flavicornis Le Moult vendit,; 14 (RBINS).

Onthophagus (s.str.) volucer Balth. 3, Dr. V. Balthasar

det. 60; 14 (NHMB). Mus. S.S.T.L.; 14, 12 (NHMD).

NEPAL ¢ Bheri / D: Banke, Nepalganj, Hotel Sneha,

28°02’53”N, 81°36’56”E, 125 m, 05.VII.2009, leg. A.

Kopetz, LF #64; 24, 19 (cAK), 19 (cJS). Mahakali /

D: Kanchanpur, Mahendranagar, Hotel Sweet Dream,

N28°58135 E8111 O15; 21 0:m: 02;VII- 2009 leg tA:

Kopetz LF ,#60; 192 (cAK). Narayani Prov., Sauraha,

Royal Chitwan Nat.-P., Rapti River, Ufer, 180 m NN,

27°34’80"N, 84°29°49”E, 18.04.2000, LF [light trap],

A. Skale leg.; 23, 42 (cAS), 14, 19 (cJS).). Naraya-

ni Prov., Sauraha, Royal Chitwan Nat.-P., Rapti River,

Ufer, 180 m NN, 27°34’80"N, 84°29°49”E, 18.04.2000,

LF [light trap], A. Weigel leg.; 24, 29 (NMEG), 1¢

(cAW), 2 (cJS). Narayani Prov., Sauraha, Royal Chit-

wan Nat.-P., Rapti River, Ufer, 180 m NN, 27°34’80°N,

84°29°49”E, 18.04.2000, LF [light trap], F. Wolf leg.;

14, 12 (NMEG), 1° (cJS). Narayani Prov., Sauraha,

Royal Chitwan Nat.-P., Rapti River, Ufer, 180 m NN,

27°34’80"N, 84°29°49"E, 16.-18.04.2000, LF [Light

trap] A. Skale leg.; 12 (cAS), 19 (cJS), 146 (cOH).

Narayani Prov., Sauraha, Royal Chitwan Nat.-P., Rap-

ti River, Ufer, 180 m NN, 27°34’51”N, 84°29°30”E,

14.-15.V.2002, HF, leg. A. Kopetz; 12 (cAK). Chitwan

distr., Royal Chitwan Nat.P., Island Jungle Resort, 29-

30.X.1995, leg. L. Peregovits; 26, 22 (HNHM), 13

(ZFMK), 14, 22 (cJS). Royal Chitwan Nat.-P., 17. /

18.11.1998, SCHOOLMESTERS leg.; 192 (cJS). Royal

Chitwan N.P., Island Resort, 17.-18.11.1998, [anony-

mous], 2¢ (cJS). Narayani Zone, Chitwan Distr., Chit-

wan N.P., Sauraha — Thati Bagh Mara, 200 — 500 m, leg.

P. Cechovsky; 14, 49 (cDJ). Prov. Narayani, Chitwan:

Gunganagar, 200 m, 27°39’N, 84°19’E, V.2005, leg. D.

Ahrens; 12 (NMEG), 12 (cJS). Pokhara, 900 m, 2.-

3.1984, Bhakta B.; 146 (NHMB), 1), 19 (cJS).

TIBET (China, Xizang) * THIBET, 600/1/ Coll. Melly;

13 (MHNG). THIBET, 501/87/ Coll. Jurine, flavicornis

Germ.; 19 (MHNG).

Without exact data

Ex Mus. E. HAROLD, Coll. Boucomont; 54' (MNHN).

Bengala, Aenescens. ekke Type WIED., Zool. Museum

DK Copenhagen; 1 (cJS).

Bengalie, Ex Mus. Thorey, BATES, Coll. Boucomont,

Paris, 23, 22 (MNHN). Bengalia, Westermann; 19

Bonn zoological Bulletin 71 (1): 29-39

(NHMD). Bengal. /Septbr. 1810., Zool. Museum DK Co-

penhagen, O. (Colobonthophagus) aenescens (WIED.);

12° (NHMD).

Neotype redescription

Male specimen from Bengal (Figs 1A—C, 2A—B)

MEASUREMENTS (Fig. 1A—B). Length: 12.2 mm, width:

7.4 mm. Male major with two well developed horns,

and pronotum with an anteromedian round prominence;

moderately arched, mostly glabrous. Color blackish

brown, with very moderately green metallic sheen. In

dorsal view eyes broad oval; first elytra interval on base

without recess, but in the first quarter with a small longi-

tutinal elevation near suture. Antennal foot-stalks reddish

brown, club segments of antenna yellowish.

Heap. Clypeus broadly round, margin anteriorly dis-

tinctly reflexed, clypeofrontal carina moderately elevated

but distinct, circlar, backwards reaching to the bases of

two horns on vertex beneath well developed, oval eyes.

In frontal view horns slightly divergent on base, softly

incurved near apices, obliquely inclined backward. Ver-

tex between horns with a thin, low carena, situated back-

ward. Clypeus densely distinctly rugo-punctate, frons

densely rugo-punctate, vertex and genae more densely

rugo-punctate, building small, low ridges.

Pronotum (Fig. 1C) with a triangular, rounded an-

teromedian prominence, longitudinal midline missing;

on each side, behind the horns, with a deep depression;

anterior angles projected, rounded; lateral margins soft-

ly rounded, only slightly sinuate. Base rounded, only in

the middle bordered, marginal line reaching the second

striae of elytra. The middle of the disc with large, simple

rounded, deep punctures; punctures with different diam-

eters, mostly separated by one diameter, becoming com-

pact at a longitudinal area in the middle from anterome-

dian prominence to the base. Punctures of anterior angles

larger, more dense and more shallow, with some yellow,

minute setae; depressions with minute, oval punctures.

The sharp border of the base of the pronotum with a

dense line of large punctures, reaching the hind angles.

ELyTrA. Striae broad, punctures small, transverse,

weakly notching both margins of intervals; 7" striae dis-

tinctly curved in the middle, intervals slightly convex,

very weakly micro-reticulated, everywhere densely and

coarsely punctured, the distances between punctures are

shorter than their diameters. The inner intervals with

shallow, small punctures, each puncture bears a small

granule at its anterior margin. Toward outer intervals the

punctures becoming coarser and deeper, without gran-

ules, but with very short, fine, yellow-white setae. All

intervals apical with some short, yellow setae.

PyYGIDIUM carinate at base, weakly convex, micro-retic-

ulated, moderately sparsely punctated. Points moderately

©ZFMK

34 Joachim Schonfeld & Dirk Rohwedder

deep, with different diameters, and sub-regular in distri-

bution, simply rounded, with some short, yellow setae.

PROTIBIAE with four strong outer teeth, apical spur blunt

inner side straight, outer side curved to the obtuse tip,

slightly bent downward.

METASTERNUM. Anterior declination weakly convex, disc

plan, smooth, with a median longitudinal groove, apical

ending with an oval depression. Anterior declination and

ventral area with minute, shallow punctures.

AEDEAGUS and LAMELLA COPULATRIX (Fig. 2A).

Variation

Measurements (70 33’, Length 10.8-12.2 mm; 86 99,

Length 9.1-12.5 mm). In female and minor male ver-

tex with reduced horns, with a narrow straight lamina

between them. Pronotum with slight depressions and

reduced anteromedian prominence, notched in female.

Clypeus of female with very coarse, transverse rugose or

subrugose punctures. There is only a weak tendency to

variation in the punctures on the elytra.

Differential diagnosis

O. (C.) aenescens is very similar to O. (C.) dama (Fabri-

cius, 1798) and O. (C.) bengalensis Harold, 1886. O. (C.)

aenescens shows on dorsal surface large punctures sep-

arated from each other by approximately | diameter of

a puncture. O. (C.) bengalensis and O. (C.) dama show

very fine and sparse punctures separated further from

each other (1-3 puncture diameters). O. (C.) bengalensis

and O. (C.) aenescens have a completely bordered base

of the pronotum, while O. (C.) dama has only a short

remnant of a border in the middle of the base. The body

of O. (C.) aenescens and O. (C.) dama show a green or

brownish metallic sheen, the body of O. (C.) bengalensis

is black without metallic sheen.

Distribution

Bangladesh (East Pakistan), India, Myanmar, Nepal,

Tibet. (type locality of O. (C.) volucer Balthasar,

1959 syn. nov.: Burma [Myanmar], Tenass[erim])

(Fig. 6).

Taxonomic remarks

Based on four complete and one partial specimen Wie-

demann (1823) described Copris aenescens from “Ben-

galia” and deposited the type material at the Zoolo-

gisches Museum Hamburg, where it was destroyed by

fire in 1943. We have not found any types of this taxon

in the Zoologisches Museum Hamburg. Only in MFNB,

NHMB (Coll. FREY) and NHMD we have found some

determined specimens, but no valid types. As no original

material of Onthophagus (Colobonthophagus) aenescens

exists, we here designate a neotype from specimens col-

lected in the type locality Bengalia to clarify the taxo-

nomic status and for nomenclatural stability according to

Article 75.3 of the Code (ICZN 1999). The neotype is

Bonn zoological Bulletin 71 (1): 29-39

chosen from the collection of NHMD and deposited at

the Natural History Museum of Denmark, Copenhagen

(NHMD).

However, it was necessary to study the type material

of O. (C.) voluver Balthasar, 1959 syn. nov. and a lot of

well determined material of O. (C.) aenescens in order to

eliminate confusions in Balthasar’s key (Balthasar, 1963:

210, alt. 180 (193)) and with other closely related spe-

cies.

The present study is based on the examination of

156 specimens of O. (Colobonthophagus) aenescens

(Wiedemann, 1823) from Bangladesh, India, Myanmar,

Nepal and Tibet. The old historic region Bengalia had

no clearly defined borders, it apparently included several

regions today called Bangladesh, West Bengal and some

parts of Bihar, Jharkharnd, Tripuru and Odisha.

Some notes on the distribution have been published,

but we are not in the position to verify them: Mittal

(2000: 263): Himachal Pradesh; Mittal & Jain (2015:

395): Himachal Pradesh (Una), Uttarakhand (Dehra-

dun), Uttar Pradesh (Faizabad), Bihar (Chapra, Pusa), W.

Bengal (Purnea dist., Kolkata); Lobl et al. (2006: 163);

Ziani & Bezdék (2016: 175): The same Indian states and

Pakistan.

Balthasar described O. (C.) voluver syn. nov. based on

a single male specimen (holotype), the female was un-

known. Frey (1973: 102) studied 3 64 and 2 99 from

East Pakistan (Bangladesh, Dinajpur, X.1969), described

the female of O. (C.) voluver and incorrectly designated

an allotype and paratypes, which are not valid as such.

These specimens are deposited as follows: allotype in

collection Dr. Carlo Brivio (P.I.M.E.), paratypes in col-

lection G. Frey (NHMB).

The examined male holotype of O. (C.) voluver 1s con-

specific with the series of specimens and the neotype

of O. (C.) aenescens. They share the same morphology

of aedeagus, lamella copulatrix, structures of the head,

vertex armatures, pronotum punctation and body col-

oration. We cannot find any morphological character-

istics to separate O. (C.) aenescens Wiedemann, 1823

from O. (C.) volucer Balthasar, 1959. Consequently, we

consider O. (C.) volucer a junior subjective synonym of

O. (C.) aenescens.

The neotype of O. (C.) aenescens, here designated,

is chosen from the type locality Bengalia. Notable en-

tomologists, for example G.J. Arrow, V. Balthasar, A.

Boucomont, G. Frey, E. Harold and R. Pittino, have de-

termined specimens especially from Bengalia as O. (C.)

aenescens. They correspond with the diagnosis of the

subgenus Colobonthophagus of Scheuern (1995), the di-

agnostic characters used in Balthasar (1963), the above

redescribed taxon, and differs from all other species of

the genus Onthophagus.

Among other morphological characters, all species

of subgenus Colobonthophagus have large eyes with a

horn or a small bulge beneath, and a swelling or a longish

©ZFMK

Type specimen designation for Onthophagus (Colobonthophagus) aenescens and O. (Colobonthophagus) urellus 35

bulge on the base of the first elytral interval. The key of

Scheuern (1995) listed characters of O. (C.) aenescens:

Black or brownish, often with a metallic green sheen.

Clypeus clearly rounded, base of pronotum only in the

middle with clear marginal line. Dorsal surface every-

where densely and coarsely punctured, the distances of

punctures are shorter than their diameters.

Onthophagus (Colobonthophagus) urellus Boucomont,

1920

(Figs 3A—-F, 4A—D, 5D-F, 6)

Onthophagus urellus Boucomount, 1920: 310 (original

description).

Onthophagus urellus var. nilgirinus Arrow, 1931: 298

(description, var. ).

Onthophagus (s.str.) usurpator Balthasar, 1959: 190

(original description) [syn. nov.].

Onthophagus (Colobonthophagus) urellus: Scheuern

1995: 418 (listing, key, pictured); Scheuern 1996: 12

(key).

Onthophagus (Colobonthophagus) usurpator: Bezdék &

Hajek 2013: 427 (status, catalogue); Mittal & Jain

2015: 402 (distribution).

Type locality of O. (Colobonthophagus) urellus Bouco-

mont, 1920: BURMA [Myanmar]: Pegu.

Type material examined (4 specs)

Onthophagus (Colobonthophagus) urellus

Lectotype

MYANMAR «° 1 major (MNHN, here designated):

“Pegu [Myanmar] / Inde [hw] // Boucomont det. 19 [p]

19[hw] / Onthophagus / urellus n.sp. [hw] // TYPUS [p,

pink label] // MUSEUM PARIS / 1936 / COLL. A. BOU-

COMONT [p] // SYNTYPE [p, red label] | // SYNTYPE

/ Onthophagus / urellus Boucomont, 1920 // MNHN /

EC10031 // Onthophagus (Colobonthophagus) |p,fat] //

urellus A. BOUCOMONT, 1920 [p] // LECTOTYPUS

Sdesig. 2022 [p,fat] // J. SCHONFELD & D. ROHWED-

DER [p, red label]”.

Onthophagus (Colobonthophagus) urellus

Paralectotypes

MYANMAR * 13, 12 (MNHN, here designated): 1¢

minor (MNHN): “Pegu [Myanmar] / India [p] // Bou-

comont det. 19 [p] 20[hw] / O. urellus B. [hw] // 3 [p]

// TYPUS [p, pink label] // MUSEUM PARIS / 1936 /

COLL. A. BOUCOMONT [p]//SYNTYPE [p, red label]

/! SYNTYPE / Onthophagus / urellus Boucomont, 1920

// MNHN / EC10033 // Onthophagus (Colobonthopha-

gus) [p,fat] // urellus A. BOUCOMONT, 1920 [p] // LEC-

TOTYPUS © desig. 2022 [p, fat] // J. SCHONFELD &

D. ROHWEDDER [p, red label]”. 19 (MNHN): “Pegu

[Myanmar] / India_[p] // Boucomont det. 19 [p] 20 [hw]

Bonn zoological Bulletin 71 (1): 29-39

/ O. urellus [hw] // 2 [p] // TYPUS [p, pink label] //

MUSEUM PARIS / 1936 / COLL. A. BOUCOMONT

[p] // SYNTYPE [p, red label] // SYNTYPE / Onthop-

hagus / urellus Boucomont, 1920 // MNHN / EC10032

/! Onthophagus (Colobonthophagus) |p, fat] // urellus

A. BOUCOMONT, 1920 [p] // PARALECTOTYPUS 92

desig. 2022 [p, fat] // J. SCHONFELD & D. ROHWED-

DER [p, red label]”.

Onthophagus (Colobonthophagus) usurpator

Holotype

INDIA © 3 (NMPC): “INDIA / LIDIO CIPRIANI [p]

// COORG/ PONNAMPET / M. GHAT. XII-1934 [p]

// Onthophagus | s.str. 3 // usurpator / Balth.[hw] /

HOLOTYPUS [p] [handwritten corrections, pink la-

bel] // Mus. Nat. Pragae/ 65844 / Inv. [p, red label] //

O. (Colobonthophagus) [p] / urellus BOUC. 3 [hw] /

J. SCHEUERN det. 19 [p] 96 [hw] // usurpator Balth.

/ syn. nov. [hw] / J. SCHEUERN det. [p] 96 [hw] //

O. (Colobonthophagus) / urellus 3 | BOUCOMONT

1919 / J. SCHONFELD det.2017 [p] // O. (Colobonthop-

hagus) / usurpator / BALTHASAR 1960 / Syn. nov. / J.

SCHONFELD det. 2017 [p]”.

Other material examined (8 specs)

INDIA * Bombay [Mumbai], Onth. (Colob.) urellus

BOUC. var. nilgirinus ARROW, 2; 292 (cJS). S. Co-

org-Ammatti_[Ammathi], 3100 ft., XI-1952 P.S. Nathan,

Coll.R.I.Sc.N.B.; 14 (cJS). Ghates, R.P.F. Tabourel, VII-

IX 1898, MUSEUM PARIS 1952, Coll.: R. OBERTHUR,

Onthophagus urellus 3, det. G. Frey, 1955; 14 (NHMB).

NILGIRI HILLS, DEVALA, 3200 ft.,??.10.1960, (ex

PSCH); 192 (cJS). Nilgiri Hills, H.-L. Andrewes, An-

drewes Bequest., B-M. 1922-221, Coll.R.I.Sc.N.B.; 19

(RBINS). Tavargatti, Belgaum Div., ex Dehra Dun Coll.

B.M.1931-2., B.M. Bhatia., Onthophag. urellus Bouc.,

G. J. Arrow det.; 14 (NHMB).

NEPAL « P: Mahakali, D: Kanchanpur, vic. Mahen-

dranagar, Suklaphanta Nat. Res., Peters Area, Dsauda

river, 160 m, O1. VII. 2017, # 17-19. leg. A. Weigel,

second. forest riverside, 28°53’51”N 80°13’39”E, LFF

[light trap]; 14 (NMEG).

Distribution

South India: Western Ghats, Myanmar, Nepal (first re-

cord, see material examined), first record of Palaearctic

region (Fig. 6).

Remarks

The present study is based on the examination of eight

specimens from South India (Western Ghats), one male

from Nepal, and three MNHN syntypes (2 63, 1 29)

of Onthophagus (C.) urellus from Pegu [Myanmar] of

which we designate here a lectotype (3). Arrow (1931:

298) described the variety nilgirinus from South India,

Nilgiri Hills. There are only minor differences between

the types and the material from South India. In the Cata-

©ZFMK

36 Joachim Schénfeld & Dirk Rohwedder

Sway

Fig. 3. A-C. Onthophagus (Colobonthophagus) urellus Boucomont, 1920, neotype, 3. D-F. Onthophagus (Colobonthophagus)

usurpator Balthasar, 1959 syn. nov., 3. A, D. Habitus, dorsal view. B, E. Habitus, fronto-ventral view. C, F. Pronotal sculpture in

dorso-lateral view (top) with close-up of the punctation. Scale = 5 mm.

Bonn zoological Bulletin 71 (1): 29-39 ©ZFMK

Type specimen designation for Onthophagus (Colobonthophagus) aenescens and O. (Colobonthophagus) urellus 37

Boucomont det, 191

outhe

uefA-

SYNTYPE

Onthophagus

urelus Boucomont, 1920

. Ps Urelus

LIDIO GIPRIANL || 9. SCHEUERN ent

usurpator &

* PONNAMPET “|.

M.GHAT. X11-4934

0. (colabonthopheaus)

INDIA

COORG

| ECHEUERN. ask. :

Fig. 4. A-B. Onthophagus (Colobonthophagus) urellus Boucomont, 1920, neotype, 3. C-D. Onthophagus (Colobonthophagus)

usurpator Balthasar, 1959 syn. nov., 4. A, C. Left: Aedeagus, lateral view, lamella copulatrix, ventral view; right: Parameres,

dorsal view & apical view. B, D. Labels.

logue of Life (2018, Annual Checklist) the var. nilgirinus

is listed as synonym of the nominotypical species.

The taxon Onthophagus usurpator Balthasar, 1959

was described based on a single male (holotype) collect-

ed in South India, Ponnampet / M. Ghat. This examined

holotype is conspecific with the lectotype of Onthopha-

gus (C.) urellus. They share the shape of the aedeagus,

pronotum, and punctation of the body. There are, how-

ever, considerable differences in the armature of the

head, a phenomenon exhibited by most other species of

the subgenus Colobonthophagus (Scheuern, 1995). Mit-

tal (2015) has listed Onthophagus (Colobonthophagus)

urellus Boucomont, 1920 from Tamil Nadu (Nilgiri Hil

Bonn zoological Bulletin 71 (1): 29-39

s), Kerala (Thirunelly forest) and Onthophagus usurpa-

tor from Kerala, (Thirunelly forest). Karnataka (Coorg

Ponnampet) was the type locality of O. (C.) usurpator.

In Nepal, O. (C.) urellus was collected using a light

trap in young secondary forest with herds of cattle on

the right side of the river (A. Weigel pers. comm.). This

record provides the only ecological hint on this species.

We determined specimens from Western Ghats as O. (C.)

urellus, but we did not see any female of O. (C.) usurpa-

tor Balthasar, 1959.

The discontinuous distribution between southern India,

Nepal and Myanmar needs to be verified by more materi-

al, especially from Myanmar or adjacent countries.

©ZFMK

38 Joachim Schénfeld & Dirk Rohwedder

3 4A

of)

a |

“~,

Fig. 5. A-C. Onthophagus (Colobonthophagus) aenescens (Wiedemann, 1823) 9. D-F Onthophagus (Colobonthophagus) urellus

Boucomont, 1920, paralectotype, 2. A, D. Habitus, dorsal view. B, E. Habitus, fronto-ventral view. C, F. Vagina & spermathec.

Scale = 5 mm.

Bonn zoological Bulletin 71 (1): 29-39

Type specimen designation for Onthophagus (Colobonthophagus) aenescens and O. (Colobonthophagus) urellus 39

75 80 85 90 95 100 105 110

@ O. aenescens

“™ A OO. urellus

105 110

Fig. 6. Map. Blue square = Onthophagus (Colobonthopha-

gus) aenescens (Wiedemann, 1823); red triangle = Onthopha-

gus (Colobonthophagus) urellus Boucomont, 1920.

Acknowledgements. We would like to thank all colleagues

and institutions included in the Material and Methods section.

Thanks go to Konjev Desender & Marcel Cludts, RBINS,

Manfred Uhlig & Joachim Schulze, MFNB, Yves Cambefort,

MNOHN, Ole Martin & Sree Selvantharan, NHMD, Michel

Brancucchi (7) & Eva Sprecher, NHMB, for allowing us to

study the material in their care. We are also grateful to Ales

Bezdék, NMPC, Bernd Jaeger, MFNB, Dirk Ahrens, ZFMK

and Oliver Hillert, Schoneiche/Berlin for their helpful advice,

and to Martin Husemann (Hamburg, Germany) for searching

for material at the Zoologisches Museum Hamburg (ZMH). Fi-

nally, we are thankful to Katrin Heuser (Zurich, Switzerland)

and Jan Decher (ZFMK) for reviewing the manuscript and cor-

recting the English text, and to Antoine Mantilleri, MNHN, for

providing some photographs.

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Bengalen und dem Vorgebirge der guten Hoffnung; beschrie-

ben von E. R. W. Wiedemann. Zoologisches Magazin Band

II (1): 3-132

Ziani S & Bezdék A (2016) Scarabaeidae, subfamily Scarabaei-

nae, tribe Onthophagini. Pp. 180-204 in: Lobl I. & Lobl D.

(eds) Catalogue of Palaearctic Coleoptera. Volume 3: (Rev-

ised and Updated Edition). Scarabaeoidea — Scirtoidea — Das-

cilloidea — Buprestoidea — Byrrhoidea. Brill, Leiden/Boston

©ZFMK

BHL

Blank Page Digitally Inserted

FORSCHUNGS

AM 6ENIG Bonn zoological Bulletin 71 (1): 41-49 ISSN 2190-7307

2022 Franga R.C. et al. http://www.zoologicalbulletin.de

https://do1.org/10.20363/BZB-2022.71.1.041

Research article

urn:|sid:zoobank.org:pub: EOFEA3A D-B582-49B8-A720-1FCC44894FC5

Historical collection of snakes from Brazil by herpetologist and biogeographer

Paul Miiller (1940-2010), deposited at the Zoological Research Museum

Alexander Koenig, Germany

Colecaéo histérica de serpentes do Brasil do herpetélogo e biogeégrafo Paul Miller (1940-2010),

depositada no Zoological Research Museum Alexander Koenig, Alemanha

Rafaela C. Franca®"*, Frederico G.R. Franca ©?, Dennis Rédder ©? & Mirco Solé”!

' Programa de Pés-graduacdo em Ecologia e Conservagdo da Biodiversidade, Universidade Estadual de Santa Cruz, Rodovia

Jorge Amado, Km 16, CEP 45662-900 Ilhéus, Bahia, Brazil

? Departamento de Engenharia e Meio Ambiente, Centro de Ciéncias Aplicadas e Educagao, Universidade Federal da Paraiba —

UFPB, Av. Santa Elizabete, s/n — Centro. CEP 58297-0000, Rio Tinto, PB, Brazil

4 Herpetology Section, Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany

*Departamento de Ciéncias Bioldgicas, Universidade Estadual de Santa Cruz, Rodovia Jorge Amado, Km 16, 45662-900 Ilhéus,

Bahia, Brazil

“Corresponding author: Email: rcfranca@Quesc.br

'urn:Isid:zoobank.org:author: 8F44D056-3DB 1-4E7B-B5D6-28BA6BD9EC36

2urn:Isid:zoobank.org:author:D1554607-B161-41C2-905D-47DE00C728C6

3urn:Isid:zoobank.org:author:C7DOE2A F-2147-43FA-A89A-D770AAEA 1 50E

*urn:Isid:zoobank.org:author:CO62E3CB-D966-441 A-8B77-1F94DC85FA92

Abstract. Natural history collections are constituted of a wide variety of biological specimens preserved around the world.

They represent a continuous source of knowledge and play a fundamental role in the synthesis on the diversity, compo-

sition, distribution, and conservation of species. Paul Miller (1940-2010) was a German zoologist who collected amphi-

bians and reptiles in Brazil between 1964 and 1976, with the aim of increasing knowledge about the Brazilian fauna and

understanding the general patterns of Neotropical biogeography. We examined and re-determined all snakes found in Paul

Miuller’s collection, deposited at the Zoological Research Museum Alexander Koenig (ZFMK), and also reconstructed

the itinerary of his journeys through Brazil. We identified 556 snake specimens belonging to 80 species from six families

(Aniliidae, Boidae, Colubridae, Dipsadidae, Elapidae, and Viperidae). Muller collected snake specimens from all regions

of Brazil, although most are from the south (76% of the species) and southeast (14% of the species). This relevant material

can contribute to historical, biogeographic and conservation studies of the Brazilian snake fauna.

Key words. Biodiversity, Reptilia, Serpentes, Southeast Brazil, South Brazil, Scientific Collections.

Resumo. As colecoées de historia natural s4o constituidas de uma grande variedade de espécimes bioldgicos preservados

em todo mundo. Elas sao fontes continuas de novos conhecimentos e apresentam um papel fundamental na sintese sobre

a diversidade, composicao, distribuigao e conserva¢ao das espécies. Paul Muller (1940—2010) foi um zodlogo alemao que

coletou espécies de anfibios e répteis no Brasil entre os anos de 1964 a 1976, com o intuito de aumentar 0 conhecimento

sobre a fauna brasileira e entender os padr6es gerais da biogeografia neotropical. Nos examinamos e re-determinamos to-

dos os individuos encontrados na colecao de Paul Miller, depositada no Zoological Research Museum Alexander Koenig

(ZFMK) e também recriamos 0 roteiro de suas viagens pelo Brasil. Nos identificamos 556 espécimes de serpentes de 80

espécies pertencentes a seis familias (Aniliidae, Boidae, Colubridae, Dipsadidae, Elapidae and Viperidae). Miller coletou

exemplares de serpentes de todas as regides do Brasil, embora a maioria seja das regides sul (76% das espécies) e sudeste

(14 % das espécies). Este relevante material pode contribuir para estudos historicos, biogeograficos e conservacionistas

da fauna de serpentes brasileiras.

Palavras-chave. Biodiversidade, Reptilia, Serpentes, Sudeste do Brasil, Sul do Brasil, ColegGes cientificas.

INTRODUCTION their purpose is to document biodiversity and its distri-

bution and to serve as a resource for research and educa-

Natural history collections are made up of a wide variety tion (Winker 2004). This set of specimens comprise an

of biological specimens preserved around the world and invaluable record of the evolution of life and is the basis

Received: 06.04.2021 Corresponding editor: W. Bohme

Accepted: 29.03.2022 Published: 15.04.2022

42 Rafaela C. Franca et al.

for many biological researches such as taxonomy and

systematics, ecology, biogeography, mapping and mon-

itoring (Renner & Ricklefs 1994; O’Connell et al. 2004;

Winker 2004; Pyke & Ehrlich 2010; Ballard et al. 2017).

Paul Miller (1940-2010) was a German zoologist who

studied and collected amphibians and reptiles in Brazil

from 1964 to 1976. His main interests were related to

zoogeographical-ecological issues and problems of the

evolutionary genetics of amphibians and reptiles of the

Neotropics (Muller 1971). Muller visited different re-

gions of Brazil and gathered a huge collection of ver-

tebrates, including more than 6000 herpetological spec-

imens (Monzel 2016). His research on the Neotropical

herpetofauna focused on processes of differentiation of

amphibians and reptiles on islands on the east coast of

Brazil, such as the island of Sao Sebastiao in Sao Paulo

and the island of Santa Catarina (e.g., Muller 1968a, c,

1969b, c). Biogeographical studies on the island of Sao

Sebastiao resulted in his PhD thesis (see Muller 1968).

In addition, his research on islands has resulted in a large

number of new species distribution records (e.g., Muller

1968b, c, d, 1974, 1975, 1978). He also studied important

Brazilian herpetological collections, such as the Museu

Goeldi in Belém (Para), the Museu Nacional in Rio de

Janeiro, and the Instituto Butantan in Sao Paulo, where

he collected data to obtain an overview of the morpho-

logical variability of the species he collected in the field

(Monzel & Bohme 2010). Muller published in 1973 his

famous work “Dispersal centres of terrestrial vertebrates

in the Neotropic realm“ (see Muller 1973), which was the

result of his major field research from 1964 until 1970

and his investigations in Neotropical biogeography.

Throughout his research in Brazil, Paul Muller met

important Brazilian herpetologists such as Paulo Emilio

Vanzolini, Afranio do Amaral, Alphonse Richard Hoge,

Thales de Lema and Paulo Sawaya, the latter became

a friend and, to some extent, his “Brazilian supervisor”

(Monzel & Bohme 2010). On his return to Germany, Paul

Miller took his collection of amphibians and reptiles to

the University of Saarbriicken, where he was a profes-

sor and appointed head of the Institute of Biogeography

in 1971, and later, in 1999, moved his collection to the

University of Trier, where he accepted an offer to estab-

lish a new biogeographic institute (Monzel & Bohme

2010). After he retired in 2006, Paul Muller donated his

important herpetological collection to the Zoological Re-

search Museum Alexander Koenig, Germany (Monzel &

Bohme 2010).

In this study, we have examined and re-determined all

snake individuals found in Paul Muller’s collection and

also reconstructed Paul Muller’s journey through Brazil

over the years.

Bonn zoological Bulletin 71 (1): 41-49

MATERIAL AND METHODS

We identified all snake specimens following the current

nomenclature (e.g., Campbell & Lamar 2004; Grazziotin

et al. 2012; Pyron et al. 2013; Hoogmoed et al. 2019).

In addition, we recovered some of the material that was

poorly preserved, changed all the containers that were

damaged and renewed the alcohol. After having carefully

analyzed all the samples, we inserted the museum label

(ZFMK), but we also kept Paul Miller’s original field

labels. We then entered the field information of the spec-

imens into the museum’s database.

For the construction of the map with Paul Muller’s

travel itinerary through Brazil, we used the information

of the location, and the date on which the snake speci-

mens were collected. We georeferenced the points of the

locations where Paul Muller collected the species, that is,

in this process we considered only the snake specimens

that were collected by him. Then, we inserted all the in-

formation in ArcGIS 10.1 (ESRI 2004) and built a map.

RESULTS

We identified 80 snake species distributed in 556 spec-

imens and six families (Aniliidae, Boidae, Colubridae,

Dipsadidae, Elapidae, and Viperidae) in Paul Muller’s

collection (Table 1, Fig. 1). The family Dipsadidae is

the most represented in the collection, with the largest

Fig. 1. Paul Miuller‘s Snake Collection deposited at the Zoo-

logical Research Museum Alexander Koenig, Germany. Philo-

dryas olfersii Gunther, 1885 (ZFMK 102104), collected in Rio

Grande do Sul (A), snake specimens from the collection (B),

Bothrops alternatus Duméril, Bibron & Duméril, 1854 (ZFMK

102119) collected in Rio Grande do Sul (C).

©ZFMK

Collection of snakes by Paul Miller 43

Table 1. Number of specimens (N) of each snake species in Paul Miiller’s collection, the respective catalogue number of the

Zoological Research Museum Alexander Koenig (ZFMK) and the location where the species was collected. The abbreviations in

the localities column correspond to the following Brazilian states: AM (Amazonas), BA (Bahia), MG (Minas Gerais), MT (Mato

Grosso), PA (Para), RJ Rio de Janeiro, RS (Rio Grande do Sul), SC (Santa Catarina) and SP (Sao Paulo).

Family /Species N (ZFMKk) Localities

Aniliidae

Anilius scytale (Linnaeus, 1758) 1 102325 AM: Manaus

Boidae

EL aren Pn ei ae Ee 10 091896; 096323-28: 096357- AM: Manaus, Careiro; PA: Santarém,

; 59 Belém, Marituba, Ilha de Marajo

Epicrates crassus Cope, 1862 1 102405 MG: Pedro Leopoldo

Colubridae

Chironius bicarinatus (Wied-Neuwied, 1820) 7 — 102129-34; 102453 Lite ae

Chironius exoletus (Linnaeus, 1758) 6 Ae a A oe rae Se Ecopolde 8 Bloranopaks,

Chironius fuscus (Linnaeus, 1758) 1 102607 Brazil

Chironius quadricarinatus Boie, 1827 1 102608 MT: Cuiaba

Drymarchon corais (Bote, 1827) 1 102407 RS: Taquari

Leptophis ahaetulla (Linnaeus, 1758) 2 102380-81 AM: Uaupés, Manaus

Palusophis bifossatus (Raddi, 1820) 2 102170; 102624 RS: Sao Leopoldo

Spilotes pullatus (Linnaeus, 1758) 3 102110; 102602-03 pe ee an pepe

Spilotes sulphureus (Wagler, 1824) 2 102090-91 AM: Manaus

Dipsadidae

Apostolepis assimilis (Reinhardt, 1861) 1 102120 SC: Florianopolis

Atractus paraguayensis Werner, 1924 5 102191; 102439-43 RS: Taquara; SC: Florianopolis

Atractus reticulatus (Boulenger, 1885) 2 102140-102534 RS: Sao Leopoldo

Cercophis auratus (Schlegel, 1837) 2 102484-85 SC: Florianopolis

Dipsas albifrons (Sauvage, 1884) 2- 1O2510; LO2S1F SP: Ilha de Sao Sebastiao

Dipsas indica Laurenti, 1768 1 102201 RS: Sao Leopoldo

Dipsas mikanii (Schlegel, 1837) 1 102458 RS: Sao Leopoldo

Dipsas neuwiedi (Ihering, 1911) 11 102660-69; 102671 SC: Ilha de Santa Catarina

Disc ia Cope, 8 DE aaa eer te

Dipsas ventrimaculatus (Boulenger, 1885) 18 Toren hiss as A zi i BS RS: Santa Cruz do Sul; SAo0 Leopoldo

Fe ee ie DE DEMEROS to® leDAgtee SP: Ilha de Sao Sebastiao

Echinanthera cyanopleura (Cope, 1885) 5 ele re Bowe a Houinoehe Bao: Leopoldo Taquars,

Echinanthera melanostigma (Wagler, 1824) t* sLO2Z002 SP: Ilha de Sao Sebastiao

Erythrolamprus aesculapii (Linnaeus, 1758) 2 102178; 102383 MT: Cuiaba; SP: Sao Sebastiao

Erythrolamprus almadensis (Wagler, 1824) 9 oe ee Treen aaa os Pee cee ee ete

102513; 102605 ;

102126; 102206; 102496;

Erythrolamprus jaegeri (Gunther, 1858) 9 102542-43; 102600; 102648; RS: Sao Leopoldo, Taquara

Bonn zoological Bulletin 71 (1): 41-49

102673-74

©ZFMK

Table 1. continued.

Family /Species

Erythrolamprus miliaris (Linnaeus, 1758)

Erythrolamprus poecilogyrus (Wied-Neu-

wied, 1825)

Erythrolamprus semiaureus (Cope, 1862)

Erythrolamprus typhlus (Linnaeus, 1758)

Helicops carinicaudus (Wied-Neuwied, 1825)

Helicops infrataeniatus Jan, 1865

Lygophis anomalus (Gunther, 1858)

Lygophis flavifrenatus Cope, 1862

Lygophis lineatus (Linnaeus, 1758)

Oxyrhopus clathratus Duméril, Bibron &

Dumeéril, 1854

Oxyrhopus formosus (Wied-Neuwied, 1820)

Oxyrhopus guibei Hoge & Romano, 1977

Oxyrhopus petolarius (Linnaeus, 1758)

Oxyrhopus rhombifer Dumeéril, Bibron &

Dumeéril, 1854

Phalotris lemniscatus (Duméril, Bibron &

Dumeril, 1854)

Phalotris reticulatus (Peters, 1860)

Philodryas aestiva (Dumeéril, Bibron &

Dumeril, 1854)

Philodryas argentea (Daudin, 1803)

Philodryas olfersii (Lichtenstein, 1823)

Philodryas patagoniensis (Girard, 1858)

Pseudoboa haasi (Boettger, 1905)

Pseudoboa neuwiedii (Duméril, Bibron &

Dumeril, 1854)

Siphlophis pulcher (Raddi, 1820)

Bonn zoological Bulletin 71 (1): 41-49

—_ Oo

—

Rafaela C. Franga et al.

(ZFMK)

102128: 102136: 102137-39:

102165-69: 102438: 102456-

57: 102473-74: 102488-

90: 102535-39: 102580:

102594-99: 102626; 102675-

76: 102682-83

102187; 102189: 102199:

102200; 102205: 102350-51:

102379: 102436-37; 102447:

10245052: 10246467;

102477; 102482: 102486-87:

102514; 102524-29: 102544-

48: 102618; 102625-28:

102640; 102649-58: 102677-

80: 102684-87

102497-98; 102516

102320

102153-58: 102384: 102469:

102499: 102500: 102504-58:

102561

102159-64: 102501-09:

102515; 102559: 102612:

102616; 102630

102152; 102468

102481; 102672

102551-54

102478-83

102633

102173

102202

102143-51; 102195-97

102194

102639

102121; 102122: 102171:

102179: 102454: 102471:

102495: 102587-90; 102643

102406; 102619-20

102093-99: 102104: 102109:

102172; 102180: 102442:

102455; 102470: 102480:

102549: 102550: 102644:

102647

102135; 102446: 102472:

102476; 102547: 102611:

102631-32: 102670

102208

102531

102638

Localities

RS: Campo Bom, SAo Francisco de Paula,

Sao Leopoldo, Taquara; SC: Floriandépo-

lis; SP: Ilha de Sao Sebastiao

MT: Cuiaba, PA: Marajo; RS: Balneario

Pinhal, Campo Bom, Sao Leopoldo,

Novo Hamburgo, Panambi, Passo Fundo,

Portao, Porto Alegre, Santa Cruz do Sul,

Sao Francisco de Paula, Sao Leopoldo,

Taquara, Viamao, SC: Florianopolis

RS: Sao Leopoldo, Sao Sebastiao do Cai

RS: Taquara

RS: Sao Leopoldo

RS: Campo Bom, Porto Alegre, Sao Leo-

poldo

RS: Sao Leopoldo, SC: Ilha de Santa

Catarina

RS: Passo Fundo, Viamao

PA: Marajo

RS: Sao Leopoldo

AM: Manaus

RS: Sao Sebastiao do Cai

SP: Sao Sebastiao

RS: Morro Reuter, Novo Hamburgo, San-

ta Cruz do Sul, Sao Leopoldo

RS: Sao Leopoldo

RS: Sao Leopoldo, SC: Floriandpolis,

Ilha de Santa Catarina, SP: Ilha de Sao

Sebastiao

AM: Manaus; Ecuador

SC: Ilha de Santa Catarina; RS: General

Camara, Morro Reuter, Panambi, Portao,

Santa Cruz do Sul, SAo Leopoldo, Taquara

RS: Passo Fundo, Sao Leopoldo, SC:

Florianopolis; SP: Ilha de Sao Sebastiao,

Sao Sebastiao

RS: Sao Leopoldo

AM: Manaus

SC: Florianopolis

©ZFMK

Table 1. continued.

Family /Species

Taeniophallus bilineatus (Fischer, 1885)

Thamnodynastes sp.

Thamnodynastes nattereri (Mikan, 1828)

Thamnodynastes hypoconia (Cope, 1860)

Thamnodynastes lanei Bailey, Thomas &

Silva-Jr, 2005

Tomodon sp.

Tropidodryas serra (Schlegel, 1837)

Tropidodryas striaticeps (Cope, 1870)

Xenodon dorbignyi (Bibron, 1854)

Xenodon matogrossensis (Scrocchi & Cruz,

1993)

Xenodon merremii (Wagler, 1824)

Xenodon neuwiedii Ginther, 1863

Elapidae

Micrurus altirostris (Cope, 1860)

Micrurus averyi Schmidt, 1939

Micrurus corallinus (Merrem, 1820)

Micrurus frontalis Duméril, Bibron &

Dumeril, 1854

Micrurus spixii Wagler, 1824

Viperidae

Bothrops alternatus Duméril, Bibron &

Dumeril, 1854

Bothrops atrox (Linnaeus, 1758)

Bothrops bilineatus (Wied-Neuwied, 1821)

Bothrops cotiara (Gomes, 1913)

Bothrops diporus Cope, 1862

Bothrops insularis (Amaral, 1921)

Bothrops jararaca (Wied-Neuwied, 1824)

Bothrops jararacussu Lacerda, 1884

Bothrops leucurus Wagler, 1824

Bothrops moojeni Hoge, 1966

Bothrops pubescens (Cope, 1870)

Crotalus durissus Linnaeus, 1758

Collection of snakes by Paul Miller

Rea WY WR WN

(ZFMK)

102601; 102645

102681

102182-86; 102511-12;

102532; 102584

102192

102533

102659

102117; 102634-35

102636

102174-77; 102518-23:

102540: 102593: 102637

102604

102101-07; 102333-36;

102382; 102391-04; 102444

102123: 102127: 102181:

102646

096392-01; 102111-15;

102188; 102190; 102621

102629

102141; 102387-90; 102408-

102386; 102623; 102108

102119: 102319: 102329:

102338-39: 102341; 102346:

102355; 102372

102142

102378

102118; 102310-11

102124; 102125; 102344

93234

32540: 102100; 102321-24:

102337; 102342: 102345:

102347: 102352-54; 102356-

60: 102362-71: 102377:

102433-35: 102688-99

102326; 102327; 102373

102116

102328; 102343

102312-18; 102330-32;

102340; 102348-49; 102361

093229-33; 102375-76

Localities

SC: Ilha de Santa Catarina; SP: Ilha de

Sao Sebastiao

RS: Sao Leopoldo

SP: Ilha de Sao Sebastiao

PA: Ilha de Marajo

RS: Taquara

SC: Florianopolis

RS: Campo Bom, Praia da Pinhal, Sao

Leopoldo; SC: Florianopolis; SP: Ilha de

Sao Sebastiao

MT: Rondonopolis

RS: Santa Cruz do Sul, S40 Francisco de

Paula, Sao Leopoldo, Taquara

RS: Morro Reuter, S40 Leopoldo; SC:

Florianopolis, Ilha de Santa Catarina

RS: Campo Bom, Portéo, Santa Cruz do

Sul, S40 Leopoldo

AM: Manaus

SC: Florianopolis; SP: Ilha de Sao Se-

bastiao

MT: Cuiaba; SP: Sao Sebastido

AM: Manaus

RS: Portaéo, Sao Leopoldo; SP: Sao Se-

bastiao

AM: Manaus

Ecuador: Quito

RS: Sao Leopoldo

RS: Erval seco, Portéo

SP: Ilha de Queimada Grande

RJ: Rio de Janeiro; RS: SAo0 Leopoldo;

SC: Florianopolis; SP: Ilha de Sao Sebas-

tio, Sao Sebastiéo; Ecuador: Quito

SP: Ilha de Sao Sebastiao

BA: Costa Oeste da Bahia

MG: Pedro Leopoldo

RS: Panambi, Portéo, Sao Leopoldo

MG: Belo Horizonte; RS: Sao Leopoldo;

SP: Campinas

Bonn zoological Bulletin 71 (1): 41-49

©ZFMK

46 Rafaela C. Franga et al.

100%

200 ne

, 180 me 3

Sc 0 fo!)

® 60% &

& 120 ° a

A 50%

= 100 ° 9

5 80 40% §

FE 60 30% =

Z 40 20% 6

20 10%

0 0%

1964 1965 1966 1967 1968 1969 1970 1971 1972 1973 1974 1975 1976 1977

Year

Fig. 2. Number of specimens collected per year (dotted line) and the cumulative percentage of specimens from Paul Miller’s col-

lection between 1964 and 1976 (non-dotted line).

Legend iCuiaba.” *

sanoaee 1976 H Zs *

: e..

unnnnun 1975 ae, 3

PTTL 1974 :

1970

nooeune 1969

cnmanrs 1968

ucosmns 1OG/ . a Ee : awed f=)

waeeuna 1966 4 * ene - as ae N

* fi * x i w E 3

sestees 1965 pa Florianépolis

shterans 1964 ; es

Sao Leopoldo

[ Brazil hed

South America 0 330 660 1.320 Km | _

-9000000 -8000000 -7000000 -6000000 -5000000 -4000000 $

Fig. 3. Paul Miller‘s journey through Brazil between 1964 and 1976 based on records of dates and locations of snake specimens.

The color of the dotted lines represents the year of the journey and the arrow indicates the direction of movement throughout the

year.

Bonn zoological Bulletin 71 (1): 41-49 ©ZFMK

Collection of snakes by Paul Miller A7

Number of Specimens

300

250

200

100 i

50 al

0 mz ——— — ——— == ——

Rio Grande do Sao Paulo

Sul

Amazonas Minas Gerais Para

Brazilian states

Fig. 4. Number of snake specimens from Paul Miller’s collec-

tion collected in nine Brazilian states (Amazonas, Bahia, Minas

Gerais, Mato Grosso, Para, Rio de Janeiro, Rio Grande do Sul,

Santa Catarina and Sao Paulo).

number of species (50 spp.) and specimens (N = 357),

followed by the family Viperidae, with 12 species and

109 specimens. Bothrops jararaca (N = 65) and Eryth-

rolamprus poecilogyrus (N = 56) are the species with the

highest number of specimens in the collection.

Most of the specimens (86%) were collected in Brazil

between 1964 and 1976 (Fig. 2). During this period, Paul

Muller made several herpetological expeditions to differ-

ent states of the country (Fig. 3). However, Paul Muller

did not collect all specimens himself, 29% of the speci-

mens were collected by collaborators. In the collection,

there are five specimens of the genus Bothrops (B. jar-

araca, B. leucurus, B. insularis, B. cotiara, and B. pu-

bescens) collected in Brazil, between 1909 and 1963,

which were donations, as well as five other specimens

(Bothrops bilineatus, Bothrops jararaca, Chironius ex-

oletus, and Philodryas argentea (N = 2) which are from

Ecuador.

In Paul Muller’s collection there are snake specimens

from all regions of Brazil, although most are from the

south (N = 422; 76%) and southeast (N = 81; 14%). Most

of the specimens in the collection were captured in the

state of Rio Grande do Sul (53%), in the municipalities of

Sao Leopoldo, Campo Bom, Portaéo and Taquara, in the

state of Santa Catarina (22%), on the island of Santa Ca-

tarina, and in the state of SAo Paulo (13%), on the island

of Sao Sebastiao (Fig. 4). There is only one specimen

from the Northeast region, a Bothrops leucurus collected

in Bahia in 1912 which was donated to the collection.

DISCUSSION

Paul Miller’s first expeditions to Brazil were aimed at

obtaining as much information as possible about the

vertebrate fauna and butterflies from all over the coun-

try and thus increasing knowledge about the diversity of

the fauna and inferring patterns from Neotropical bio-

geography (Miller 1971; Monzel & Bohme 2010). Ac-

cording to Miller (1971), Brazil is of extreme zoogeo-

graphical interest, since it is the largest country in South

America, with a tropical climate and a much differenti-

ated vegetation zoning. It presents a transition between

Bonn zoological Bulletin 71 (1): 41-49

the humid forests of Ecuador, Colombia and Guyanas,

on the one hand, and the dry areas of the Argentinean

Pampa and Paraguay, on the other. Although Muller had

a vast knowledge of several groups of animals, most of

his studies were in the area of herpetology (Monzel &

Bohme 2010).

During his expeditions to Brazil, Miller visited several

states, but concentrated his collections on the Brazilian

coastal states, mainly Rio Grande do Sul, Sao Paulo and

Santa Catarina. Of these, Rio Grande do Sul was the state

with the largest number of snake specimens collected.

Although the focus of Paul Muller’s work in Brazil was

on island fauna (e.g., Muller 1968a, b, c, 1969a, b, c),

most of the collection sites in Rio Grande do Sul are not

islands. The number of specimens collected both in Rio

Grande do Sul and Santa Catarina is also due to the help

of Paul Muller’s friends, who supported him in collecting

the specimens and whom he thanked in his work, namely

Erno Bohler and Flavio Silva (both from Sao Leopoldo)

and Canisius Ritter (Florianopolis) (e.g., Muller 1968c,

1971).

The most numerous species in Paul Miuller’s collec-

tion were Bothrops jararaca and Erythrolamprus poecil-

ogyrus. B. jararaca is a species widely distributed in the

South and Southeast of Brazil (Sazima 1992; Grazziotin

et al. 2006; Monzel & Wister 2008) and common in the

Serra do Mar region (Centeno et al. 2008). EF. poecilogy-

rus 1s common and widely distributed throughout Brazil

(Dixon & Markezich 1992; Alencar & Nascimento 2014).

No species registered in Paul Miuller’s collection is in the

threatened species category of the Brazilian red list of

threatened species, except B. insularis, which 1s Critical-

ly Endangered (ICMBio 2018). Among the snakes in the

collection are two specimens of Uromacerina ricardinii,

used by Paull Miller to report the first record of the spe-

cies for the state of Santa Catarina (see Muller 1978). Re-

cently the species underwent a synonymization process

and today it is known as Cercophis auratus (Hoogmoed

et al. 2019).

In the collection, there are some species that provide

information from interesting collection locations. For

example, there are two specimens of Dipsas turgidus

collected in Florianopolis, Santa Catarina. In Brazil,

the distribution of D. turgidus covers the states of Mato

Grosso, Minas Gerais, Rio de Janeiro, Rio Grande do Sul

and Sao Paulo (Wallach et al. 2014). There are four indi-

viduals of Xenodon dorbignyi from Sao Sebastiao Island,

in Sao Paulo. This species has a known distribution in

the extreme south of Brazil, Uruguay, southern Paraguay

and central-northern Argentina (Orejas-Miranda 1966;

Giraudo 2001; Nenda & Cacivio 2007; Kunz et al. 2011;

Cacciali et al. 2016). In addition, other species with lo-

cation information for SAo0 Sebastiao and Sao Sebastiao

Island, in Sao Paulo, have not yet been registered in these

locations. For example, B. alternatus (N = 4), Micrurus

Jrontalis (N = 1) and Oxyrhopus petolarius (N = 1) col-

©ZFMK

48 Rafaela C. Franga et al.

lected in Sao Sebastiao, and the species Philodryas aes-

tiva (N = 1), Echinanthera melanostigma (N = 1) and

Erythrolamprus almadensis (N = 1) collected on Sao

Sebastiao Island (see Cicchi et al. 2007; Centeno et al.

2008). The absence of records of these species for these

locations can be explained by a variety of factors, such as

the rarity of some species in the region or the preference

of habitats of some species. Errors may have occurred in

the records of the collection locations of these species or

even some of these species may have disappeared from

these locations due to changes in land use in recent de-

cades.

The analysis of snake specimens from Paul Muller’s

collection can contribute to historical biogeographic

and conservationist studies of the Brazilian snake fauna.

In addition, given the current scenario of material loss

caused by recent incidents in large Brazilian collections,

such as the fires at the Instituto Butantan in 2010 and the

Museu Nacional in 2018 (see Mega 2019), these collec-

tions can provide valuable information and help to soften

the impact of material loss.

Acknowledgements. RCF thanks Fundagaéo de Amparo a

Pesquisa de Estado da Bahia (FAPESB) for a PhD scholarship

(BOL0353/2016) and Conselho Nacional de Desenvolvimento

Cientifico e Tecnol6gico (CNPq) for a Doutorado Sanduiche

no Exterior (SWE) scholarship (208442/2017-0). MS thanks

CNPq for a research scholarship (304999/2015-6) and Alex-

ander von Humboldt Foundation/CAPES for a grant (BEX

0585/16-5). FGRF thanks the financial support from CNPq

(Universal grant 404671/2016-0).

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BHL

Blank Page Digitally Inserted

Bonn zoological Bulletin 71 (1): 51-67

2022 Pietka J. et al.

https://do1.org/10.20363/BZB-2022.71.1.051

ISSN 2190-7307

http://www.zoologicalbulletin.de

Research article

urn:|sid:zoobank.org: pub:7A 17D328-4B7E-4D0D-80D4-87 1 F993DB695

Tree-fungus beetles collected on sawdust substrate with mycelium of selected

fungal species on trees at the Experimental Forest Station in Rogéw

(central Poland)

Jacek Pietka', Jerzy Borowski? & Adam Byk*"

'23 Department of Forest Protection, Institute of Forest Sciences, Warsaw University of Life Sciences — SGGW, Nowoursynowska

159/34, 02-776 Warsaw, Poland

“Corresponding author: Email: adam_byk@sggw.edu.pl

'urn:Isid:zoobank.org:author:60AE6C5 1-6D00-4196-8D95-F425501C4C14

7urn:|sid:zoobank.org:author:D763FED7-63 1 A-4673-B484-06CA 1078FC04

3urn:Isid:zoobank.org:author:27905F56-A 1 DC-444E-919D-4533899457A8

Abstract. This study of beetles attracted by a sawdust substrate overgrown with mycelium of rare species of wood-decay

fungi was aimed to assess (1) ecological groups of the beetles caught in traps with such a substrate; (2) species compositi-

on of beetle communities attracted to sawdust substrate with the selected wood-decay fungi. To determine which tree-fun-

gus beetles are associated with selected species of wood-decay fungi, we caught beetles in traps with sawdust substrate

overgrown by the mycelium of seven species of wood-decay fungi: Bondarzewia mesenterica (Schaeff.) Kreisel, Fistulina

hepatica (Schaeff.) With., Fomitopsis rosea (Alb. & Schwein.) P. Karst., Grifola frondosa (Dicks.) Gray, Hericium alpe-

stre Pers., H. coralloides (Scop.) Pers. and Meripilus giganteus (Pers.) P. Karst. The research was carried out in central

Poland, at the Forest Experimental Station in Rogow. One research season (April—October) yielded a total catch of 133

beetle species associated with wood-decay fungi, including 60 mycetobiontic species. The most abundant species of beet-

les were: Trixagus carinifrons (Bonvouloir, 1859), Cortinicara gibbosa (Herbst, 1793), Dienerella ruficollis (Marsham,

1802), Rhizophagus bipustulatus (Fabricius, 1792), and Aulonothroscus brevicollis (Bonvouloir, 1859). Two autonomous

clusters of tree-fungus beetles were distinguished in the collected material: beetles associated with brown rot and beetles

associated with white rot. The sawdust substrates with mycelium of wood-decay fungi, used in traps, attract mycetophilic

and mycetobiontic beetles.

Key words. Coleoptera, mycetophilic beetles, mycetobiontic beetles, wood-decay fungi, Poland.

INTRODUCTION

The largest taxonomic group of animals that live in and

on fungi are insects, mostly beetles. This group of organ-

isms plays a major role in decomposition of tree fungi

(Gilbertson 1984). Many fruiting bodies of wood-decay

fungi are very attractive food for beetles, which feed also

on the mycelium that penetrates wood (Benick 1952;

Hammond & Lawrence 1989; Butin 1995; Pecci-Madd-

alena & Lopes-Andrade 2017). Some beetles are highly

specific and associated only with a single fungal species.

For example, bark beetles (Curculionidae: Scolytinae)

live in symbioses with primarily ophiostomatoid Asco-

mycetes (Linnakoski et al. 2012; Six 2012).

However, little is known about beetles associated with

rare wood-decay fungi because of their limited area of

distribution (usually in natural or nearly natural forests),

short period of occurrence of fruiting bodies, and their

quick decomposition. Apart from the fruiting bodies of

fungi that are commonly found on trees, such as Fomes

fomentarius (L.) Fr., Fomitopsis pinicola (Sw.) P. Karst.

Received: 21.09.2021

Accepted: 22.01.2022

or 7rametes spp., also fruiting bodies of protected species

are visited by many beetle species. Poland was the first

country in Europe to introduce fungal species protection,

in 1983 (Ordinance 1983). Since 2014, 54 species of mac-

rofungi are strictly protected, while 63 are partly protect-

ed, so in total 117 fungal species are protected in Poland,

including 20 wood-decay fungi (Ordinance 2014). A ma-

jority of the protected wood-decay fungi are associated

mostly with primeval forests. Such habitats are located

primarily in national parks and nature reserves, i.e., in

areas where large amounts of dead wood lie on the for-

est floor. Creation of suitable conditions for development

of wood-decay fungi by leaving dead wood or stopping

timber harvesting in forests does not mean that valuable,

rare species will immediately appear there. The natural

process of wood colonization by fungi takes many years,

and production of fruiting bodies is only one of the final

stages of mycelium development in wood.

Among beetles of wood-decay and other tree fungi,

some are mycetobiontic (obligatorily associated with

fungi), while others are mycetophilic (facultatively as-

Corresponding editor: D. Ahrens

Published: 12.05.2022

52 Jacek Pietka et al.

sociated with fungi). According to published data, the

following mycetobiontic beetles have been recorded on

7 species of wood-decay fungi (six species protected in

Poland) selected for this study:

— on Fistulina hepatica (Schaeff.) With.: Cryptophagus

pubescens Sturm, 1845, C. scanicus (Linnaeus, 1758),

Micrambe abietis (Paykull, 1798), Dacne bipustulata

(Thunberg, 1781), Mycetophagus piceus (Fabricius,

1777), Psudotriphyllus suturalis (Fabricius, 1801),

Triphyllus bicolor (Fabricius, 1777), Abdera flexuo-

sa (Paykull, 1799), Orchesia micans (Panzer, 1793)

(Benick 1952; Burakowski et al. 1986, 1987; Nikitsky

1993; Nikitsky & Schigel 2004; Borowski 2006),

— on Fomitopsis rosea (Alb. & Schwein.) P. Karst.: Ag-

aricochara latissima (Stephens, 1832), Dolichocis

laricinus (Mellié, 1848), Cis dentatus Mellié, 1848,

C. glabratus Mellié, 1848, Ennearthron cornutum

(Gyllenhal, 1827) (Scheerpeltz & Hofler 1948; Kra-

sutsky 1995, 1997),

— on Grifola frondosa (Dicks.) Gray: Sepedophilus testa-

ceus (Fabricius, 1793) (Schigel 2007),

— on Hericium coralloides (Scop.) Pers.: Scaphisoma

agaricinum, Mycetophagus decempunctatus Fabricius

1801, M. multipunctatus Fabricius, 1792, M. quadri-

guttatus Muller, 1821 (Benick 1952; Nikitsky 1993),

— on Meripilus giganteus (Pers.) P. Karst.: Scaphisoma

agaricinum (Linnaeus, 1758), Cryptophagus dentatus

(Herbst, 1793), C. uncinatus Stephens, 1830, Carto-

dere nodifer (Westwood, 1839), Cortinicara gibbosa

(Herbst, 1793), Cis micans (Fabricius, 1792), Or-

chesia micans (Scheerpeltz & Hofler 1948; Benick

1952; Burakowski et al. 1987).

On the remaining two species of wood-decay fungi

used in this study, 1.e., Bondarzewia mesentrica (Schaeff. )

Kreisel and Hericium alpestre Pers., no mycetobiontic

beetles have been reported in the literature so far.

This study of beetles attracted by a sawdust substrate

overgrown with mycelium of rare species of wood-decay

fungi was aimed to assess (1) ecological groups of the

beetles caught in traps with such a substrate; (2) species

composition of beetle communities attracted to sawdust

substrate with the selected wood-decay fungi.

MATERIAL AND METHODS

Study area

The study was conducted in 2008 in the arboretum of the

Experimental Forest Station in Rogow (central Poland)

(Fig. 1). The arboretum was created in 1925 in a forest

and was associated since the very beginning with forest

research. The arboretum now covers 53.76 ha, including

41 ha being open to visitors (Pewniak et al. 2004), but

the research plot was established in the part that is closed

to visitors. The plot is located in a modified low-lying

oak-hornbeam forest Tilio-Carpinetum stachyetosum.

Bonn zoological Bulletin 71 (1): 51-67

It is dominated by Pinus sylvestris L. (aged about 130

years) and Quercus robur L. and Quercus petraea (Matt.)

Liebl. (aged 100—200 years), with admixture of Larix de-

cidua Mill., Carpinus betulus L., Abies alba Mill., Bet-

ula pendula Roth., and Picea abies (L.) H. Karst. The

well-developed herb layer includes Ste/laria holostea L.,

Anemone nemorosa L., Galeobdolon luteum Huds., Des-

champsia caespitosa (L.) P. Beauv., Poa trivialis L., and

Rubus spp.

The woodlands at Rogow are composed of small forest

patches, covering less than 10 km*. Moreover, the forest

patches are clearly separated from one another and form

a mosaic of small forest islands surrounded by farmland,

characteristic of central Poland. They are poor in typical

forest beetle species. However, the insect fauna of the

Experimental Forest Station in Rogow is well studied,

especially forest beetles (Borowski 2001).

Trapping

In the experiment, we used modified window traps

20 x 30 cm Plexiglas plates crossing each other, a funnel

and a preservative-filled bottle (ethylene glycol), and bait

(sawdust substrate overgrown with mycelium) (Fig. 2).

In the experiment we used mycelium of seven species of

wood-decay fungi belonging to the division Basidiomy-

cota, two of them causing brown rot: Fistulina hepatica

(Schaeff.) With. and Fomitopsis rosea; and five causing

white rot: Bondarzewia mesenterica, Grifola frondosa,

Hericium coralloides, H. alpestre, and Meripilus gigan-

teus (Table 1). Pure cultures of the fungi used to prepare

the bait substrate originated from the collection of the De-

partment of Forest Phytopathology and Mycology, War-

saw University of Life Sciences (SGGW). Identity of the

mycelium was confirmed using molecular techniques, on

the basis of the internal transcribed spacer (ITS) region

of nuclear ribosomal DNA, which is used most often for

research on genetic diversity of fungi (Seifert 2009). The

analyses were performed at the laboratory of the Institute

of Botany, Polish Academy of Sciences, Krakow.

The bait substrate was prepared by mixing and water-

ing of sawdust of an appropriate tree species with ground

wheat grain and inoculating it with mycelium. Beech

sawdust (Fagus sylvatica L.) was prepared for G. frondo-

sa, H. coralloides, and M. giganteus, fir sawdust (Abies

alba) for B. mesenterica and H. alpestre, spruce sawdust

(Picea abies) for F- rosea, and oak sawdust (Quercus ro-

bur) for F. hepatica. Each autoclavable polypropylene

biohazard bag (7” x12”, Cole-Parmer, US) was filled

with 180 g of the substrate (150 g of sawdust of the ap-

propriate tree species and 30 g of ground wheat grain).

The components were thoroughly mixed with 180 ml of

distilled water. The initial moisture content of the sub-

strate was about 100%. The bags were placed in an au-

toclave and sterilized at 121°C for 2 h, next moved to a

©ZFMK

Tree-fungus beetles collected on mycelium of selected fungal species in central Poland 53

Table 1. Characteristics of the fungi used to prepare the substrate attracting beetles (type of rot and major host plants according to:

Kotlaba (1984), Kreisel (1987), Ryvarden & Gilbertson (1993, 1994), Wojewoda (2003)).

Species Family Rot type

Bondarzewia mesenterica Bondarzewiaceae white

Fistulina hepatica Fistulinaceae brown

Fomitopsis rosea Fomitopsidaceae brown

Grifola frondosa Fomitopsidaceae white

Hericium alpestre Hericiaceae white

Hericium coralloides Hericiaceae white

Meripilus giganteus Meripilaceae white

Protection status in

Poland

partial protection

not protected

Major host plants

Abies, Picea

Quercus, Castanea

Picea, Abies

Quercus, Acer, Betula, Carpinus, Fagus

Abies, Picea

Fagus

Quercus, Fagus, Acer, Aesculus

laminar air flow cabinet and left for a few hours, until

they cooled down. The substrate was then inoculated

with mycelium of an appropriate species (from cultures

on Petri dishes), the bag was plugged with a cotton wool

ball, and the neck of the bag was tightly wrapped in a

colourless tape. The cotton wool was a filter that ensured

gas exchange and simultaneously protected the substrate

against microorganisms from the environment. Next the

bags were kept for 8-10 weeks at 22°C in a Q-Cell 700

incubator, until the substrate was overgrown by myceli-

um, and then used in an experiment.

Fig. 1. Location of the study area and a scheme of distribution of the traps with mycelium of wood-decay fungi on sawdust substrate

in the forest arboretum in Rogow. Symbols: @ = traps with Bondarzewia mesentrica, © = Grifola frondosa, = = Fomitopsis rosea,

Bonn zoological Bulletin 71 (1): 51-67

oO = Meripilus giganteus, A = Hericium coralloides, A= Hericium alpestre, = Fistulina hepatica.

©ZFMK

54 Jacek Pietka et al.

Fig. 2. Trap with mycelium of wood-decay fungi on sawdust

substrate, used to catch beetles (photo J. Pietka).

Then, small pieces of the polypropylene film were cut

off (at the top and at the bottom), and the substrate was

mounted above a window trap hung on a tree. The traps

were hanged at a height of 2 m, on young trees without

injuries, to make sure that the caught insects were attract-

ed by the substrate, rather than by a dying tree. From

April to October in the study area 70 such bait traps were

used, 10 for each of the studied species of wood-decay

fungi. Once a month, the caught material was collected,

transported to the laboratory, and next sorted and identi-

fied. The taxonomic classification and names of the iden-

tified insects follow the Catalogue of Palearctic Coleop-

tera (LObI & Smetana 2003—2013). When analysing their

dominance structure, Kasprzak & Niedbata’s (1981)

scale was used: superdominants (>30.00%), dominants

(5.01-—30.00%), subdominants (1.01—5.00%), and acci-

dentals (<1.00%). The faunistic similarity of tree beetle

communities caught in traps with mycelium of individual

species of wood-decay fungi on sawdust substrate was

analysed using numerical cluster analysis. This analysis

was based on the species composition (presence/absence

of species). To distinguish between groups, Ward’s algo-

rithm was used, and squared Euclidean distance was a

measure of similarity. Additionally, the faunistic similar-

ity of tree-fungus beetle communities was analysed using

Bonn zoological Bulletin 71 (1): 51-67

Nonmetric Multidimensional Scaling (NMDS). Data on

number of individuals were subjected to Hellinger trans-

formation and NMDS based on Bray-Curtis similarity

index. NMDS stress 0.099.

Results

From April to October 2008, we caught 2038 beetles of

133 species, representing 35 families (Table 2). In the

collected material, the families represented by the largest

numbers of species were the Latridiidae (11.3% of the

total number of species), Staphylinidae (9.0%), Ciidae

(6.0%), Cryptophagidae (6.0%), Curculionidae, Melan-

dryidae, and Mycetophagidae (5.3% each). The families

represented by the largest numbers of individuals were the

Throscidae (23.6%), Latridiidae (19.6%), Monotomidae

(7.6%), Cryptophagidae (7.0%), Salpingidae (6.6%), and

Leiodidae (5.5%). The most abundant species were: 7rix-

agus carinifrons (14.9%), Cortinicara gibbosa (7.1%),

Dienerella ruficollis (7.1%), Rhizophagus bipustulatus

(6.9%), and Aulonothroscus brevicollis (5,5%). These

species and 15 others (Acalles camelus, Agathidium ni-

gripenne, A. seminulum, Anaspis frontalis, A. rufilabris,

Cerylon histeroides, Cryptophagus dentatus, C. dorsalis,

Dacne bipustulata, Ennearthron palmi, Litargus con-

nexus, Mycetophagus multipunctatus, Salpingus plan-

irostris, S. ruficollis, and Trixagus dermestoides) were

caught on all types of bait. On the other hand, 35 species

were recorded only with one of the seven variants of the

bait, and only three of them were caught in the number of

more than one individual, 1.e., Hemicoelus canaliculatus

(two individuals in one trap with Grifola frondosa myce-

lium), Dendrophilus punctatus (two individuals in two

traps with Hericium alpestre mycelium) and Cis festivus

(three individuals in two traps with Meripilus giganteus

mycelium).

Among biotope groups of beetles, the major contribu-

tors to the total number of species were the organisms as-

sociated with mycelium and fruiting bodies of fungi (my-

cetocoles — 48.1%) (Table 3). The most abundant among

them were: 7rixagus carinifrons, Cortinicara gibbosa,

Dienerella ruficollis, and Aulonothroscus brevicollis.

The second most species-rich biotope group is associat-

ed with partly or completely dead wood (saproxylocoles

— 24.1%), represented most abundantly by Anaspis ru-

filabris and Melasis buprestoides. Corticoles were anoth-

er relatively large group (17.3%), and the most abundant

among them were Rhizophagus bipustulatus, Salpingus

ruficollis, and S. planirostris.

Among the trophic groups (Table 4), mycetophagous

(fungivorous) beetles were the most numerous (46.6%

of the total number of species), and the most abundant

species of this group were Dienerella ruficollis, Corti-

nicara gibbosa, and Trixagus carinifrons. Zoophages

ranked second (24.1%), and saproxylophages ranked

©ZFMK

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Bonn zoological Bulletin 71 (1): 51-67

Tree-fungus beetles collected on mycelium of selected fungal species in central Poland

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Bonn zoological Bulletin 71 (1): 51-67

Jacek Pietka et al.

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Bonn zoological Bulletin 71 (1): 51-67

Tree-fungus beetles collected on mycelium of selected fungal species in central Poland

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Bonn zoological Bulletin 71 (1): 51-67

Jacek Pietka et al.

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©ZFMK

Bonn zoological Bulletin 71 (1): 51-67

Tree-fungus beetles collected on mycelium of selected fungal species in central Poland 61

800

—@— individuals

700

—l-—species

300

Number of individuals or species

200

100

Jun

Jul Aug Oct

Sep

Fig. 3. Seasonal variation in numbers of individuals and species of beetles caught in traps with mycelium of wood-decay fungi on

sawdust substrate.

third (20.3%). Species feeding on slime moulds were the

least numerous (myxomycophages — 9.0%).

The largest number of individuals and species was

caught in spring months (Fig. 3). Among the dominant

species, Rhizophagus bipustulatus was caught primari-

ly in early spring (April-May), Cortinicara gibbosa in

spring (April-June) and autumn (October), whereas Di-

enerella ruficollis, in late summer and autumn (August—

October). The seasonal dynamics of species of the family

Throscidae shows that among Aulonothroscus brevicol-

lis, Trixagus carinifrons (dominant species), and 7. der-

mestoides, only 7: dermestoides has one peak of abun-

dance (in June), while the other two species have two

peaks of abundance. This suggests that the latter species

are bivoltine, 1.e., have two generations a year. The first

generation peaks in May, whereas the second generation

of A. brevicollis, in July, and that of 7? carinifrons, in

September (Fig. 4).

Cluster analysis of species composition of the catch for

the seven bait variants allowed us to distinguish two clus-

ters of species composition. The first cluster was formed

by the two beetle assemblages attracted to sawdust sub-

strate with mycelium of F) hepatica and F: rosea, 1.e., the

fungi causing brown rot (Fig. 5). The second cluster was

formed by all the other assemblages, including all spe-

cies being attracted by sawdust substrate with the fungi

causing white rot: B. mesenterica, G. frondosa, H. alpes-

tre, H. coralloides, and M. giganteus. The latter cluster

is composed again of two groups of species: the beetles

associated with fungi of the genus Hericium, and beetles

Table 3. Biotope groups of beetles caught in traps with mycelium of wood-decay fungi on sawdust substrate..

Number of species

Biotope group S %

Corticoles 23 173

Mycetocoles 64 48.1

Myrmetocoles 2 es)

Myxomycetocoles 12 9.0

Saproxylocoles 32 24.1

Bonn zoological Bulletin 71 (1): 51-67

Number of individuals

N %

39] 19.2

1303 63.9

5 0.2

170 8.3

169 8.3

©ZFMK

62 Jacek Pietka et al.

Table 4. Trophic groups of beetles caught in traps with mycelium of wood-decay fungi on sawdust substrate.

Number of species

Number of individuals

Trophic groups 5 % N %

Mycetophages 62 46.6 1297 63.6

Myxomycophages 12 9.0 170 8.3

Saproxylophages 27 20.3 146 Wee

Zoophages 32 24.1 425 209

associated with the other fungi causing white rot (Fig. 5).

A similar result was obtained using the Nonmetric Multi-

dimensional Scaling analysis (Fig. 6).

In the cluster composed of beetles attracted by saw-

dust substrate with brown rot fungi, the dominant species

were Cortinicara gibbosa (11.2%), Rhizophagus bipus-

tulatus (9.3%), Trixagus carinifrons (7.2%), and Cryp-

tophagus dentatus (6.7%) (Fig. 7). The most frequently

caught species specific to brown rot (absent from the sec-

ond cluster) was Corticarina similata (three individuals

in three traps).

In the cluster composed of beetles attracted by saw-

dust substrate with white rot fungi, the dominant spe-

cies were Trixagus carinifrons (17.8%), Dienerella

ruficollis (8.2%), Rhizophagus bipustulatus (6.0%), Au-

160

—s—Aulonothroscus brevicollis

140 —e— Trixagus carinifrons

—#— Trixagus dermestoides

lonothroscus brevicollis (5.9%), and Cortinicara gibbosa

(5.5%) (Fig. 8). The most frequent species in the white rot

were Ptilinus pectinicornis (15 individuals in 12 traps),

Scaphisoma agaricinum (nine individuals in nine traps),

and Clypastraea pusilla (eight individuals in six traps).

DISCUSSION

Trapping of 133 species of tree-fungus beetles with

the use of sawdust substrate with mycelium during one

growing season is a methodological success. As many as

90 of the species were recorded in the Experimental For-

est Station in Rogow for the first time, and two new spe-

cies for the fauna of Poland were found: Corticaria pineti

120

100

Number of individuals

Apr May Jun

Jul Aug sep Oct

Fig. 4. Seasonal variation in abundance of species of the family Throscidae caught in traps with mycelium of wood-decay fungi

on sawdust substrate.

Bonn zoological Bulletin 71 (1): 51-67

©ZFMK

Tree-fungus beetles collected on mycelium of selected fungal species in central Poland 63

Distance

Fr Fh He

Ha Mg Gf Bm

Fig. 5. Faunistic similarity of communities of beetles of wood-decay fungi (Cluster analysis), caught in traps with mycelium on

sawdust substrate. Abbreviations: Bm = Bondarzewia mesenterica; Fr = Fomitopsis rosea, Fh = Fistulina hepatica, Gf = Grifola

Jrondosa, Ha = Hericium alpestre,; Hc = Hericium coralloides; Mg = Meripilus giganteus.

and Euplectus infirmus. The former is a rarely reported

species living on moulds, known mostly from Central

Europe and Scandinavia, reaching south to Greece and

Italy. The latter species lives under the bark and has an

Atlantic distribution, extending to Italy, Croatia, and

Greece in the south and Germany in the east. It is known

also from Morocco and Canary Islands. Its record in cen-

tral Poland moves its eastern distribution limit more than

300 km eastward from records reported in the literature

(Borowski et al. 2010).

Beetle species recorded in this study have various mi-

crobiotope preferences (mycetocoles, saproxylocoles,

corticoles, myxomycetocoles, myrmetocoles). Some of

them develop exclusively on fruiting bodies of tree fungi

(e.g., Orchesia micans, Cis micans) or are spore-eating

species (e.g., Scaphisoma spp.). Some others develop on

fruiting bodies of tree fungi as well as mushrooms, e.g.,

Dacne bipustulata in fruiting bodies of tree fungi (Bura-

kowski et al. 1986) and in mushrooms of the genus Cor-

tinarius (Pers.) Gray (Borowski 2006). Zoophages were

also numerous in the samples (e.g., species of the genus

Rhizophagus Herbst). Johansson et al. (2006) confirmed

that fruiting bodies of tree fungi attract not only beetles

that feed on fungi but also many predatory species.

In total, the family Throscidae was represented most

numerously, by 480 individuals of three species: Au/ono-

Bonn zoological Bulletin 71 (1): 51-67

troscus brevicollis, Trixagus dermestoides, and T: carin-

ifrons. Burakowski (1991) reported that species of this

family are rarely found in the field, so their ecology is

poorly studied. Supposedly they live in the soil or litter,

near roots colonized by mycorrhizal fungi. Thanks to the

applied trapping method, we collected new information

on the ecology and phenology of emergence of the three

species mentioned above. The presented results indicate

that 7. carinifrons and A. brevicollis have two peaks of

abundance during the year, whereas 7? dermestoides has

only one. The existence of two peaks of abundance in

A. brevicollis and one peak in 7: dermestoides was re-

ported also earlier by Leseigneur (2004). According to

Hardersen et al. (2014), Throscidae were always pres-

ent with two or three species, from early April to early

November, mainly in Malaise traps at ground layer. Van

Meer (1998) supposes that larvae of A. brevicollis can

develop on dry rotten wood, very much like larvae of the

closely related A. Jaticollis Rybinsky, 1897, which were

observed by him in dry rotten oak wood. Mertlik & Le-

seigneur (2007) indicate that imagines of these species

overwinter in dead wood. The luring properties of the

substrate with mycelium and the presence of eggs and

larvae of insects could be the causes of the high abun-

dance of Rhizophagus bipustulatus, Salpingus ruficollis,

and S. planirostris. According to Nunberg (1967), R. bi-

©ZFMK

64 Jacek Pietka et al.

0.10 0.15

0.05

brown rot

NMDS2

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-0.2 0.0 0.2

NMDS1

Fig. 6. Faunistic similarity of communities of beetles of wood-decay fungi (Nonmetric Multidimensional Scaling analysis), caught

in traps with mycelium on sawdust substrate Abbreviations: Bm = Bondarzewia mesenterica, Fr = Fomitopsis rosea, Fh = Fistulina

hepatica, Gf = Grifola frondosa, Ha = Hericium alpestre,; Hc = Hericium coralloides, Mg = Meripilus giganteus.

pustulatus is predatory, living under the bark, or rarely — pae and excrements of various insects and fungi growing

in rotten wood, feeding mostly on eggs, larvae, free pu- _at their feeding sites. Mendel et al. (1990) list it among

Cortinicara gibbosa

11%

Rhizophagus

bipustulatus

Trixagus carinifrons

side 7%

remaining species

50%

Cryptophagus

dentatus

Aulonothroscus

brevicollis

\ ° 4

Agathidium ae, 5%

nigripenne Salpingus ruficollis Dienerella ruficollis

3% 4% 4%

Fig. 7. Contributions of species to the total number of beetles attracted by sawdust substrate with fungi causing brown rot.

Tree-fungus beetles collected on mycelium of selected fungal species in central Poland 65

remaining species

46%

)

Trixagus carinifrons

18%

Dienerella ruficollis

pae Rhizophagus

bipustulatus

Aulonothroscus

brevicollis

Cortinicara gibbosa

Trixagus Ge. ;

dermestoides Poe on e bipustu pe: Salpingus ruficollis 3%

3% A% 4%

Fig. 8. Contributions of species to the total number of beetles attracted by sawdust substrate with fungi causing white rot.

predators feeding on bark beetles of the genera TJomicus

Latreille, Orthotomicus Ferrari, and Pityogenes Bedel. It

is also known that larvae and imagines of S. planirostris

prey on bark beetles (Young 1991). The most abundant

Species among the trapped beetles included also Corti-

nicara gibbosa, Dienerella ruficollis, and Cryptophagus

dentatus. They are some of the most common mould-eat-

ing insects.

The applied sawdust substrate with mycelium of

wood-decay fungi attracted much more mycetobiontic

beetles than reported in earlier studies of fruiting bodies

of these fungal species (Scheerpeltz & Hofler 1948; Be-

nick 1952; Burakowski et al. 1986, 1987; Nikitsky 1993;

Krasutsky 1995, 1997; Nikitsky & Schigel 2004, 2007;

Borowski 2006). In traps with mycelium of G. frondosa,

we found 31 species of mycetobiontic beetles that were

not reported earlier from this fungus, with F) hepatica

25 species, with F’ rosea 24 species, with H. coralloides

21 species, and with M. giganteus 32 species. For the

other two species of fungi, there are no published data

on mycetobiontic beetles recorded on their fruiting bod-

ies, but we found 29 species in traps with mycelium of

H. alpestre and 27 species in those with B. mesenterica.

According to Borowski (2006, 2007), 259 mycetobion-

tic beetles are reported from Poland, whereas 60 spe-

cies were caught in our study. The findings of Pietka &

Borowski (2011a) in the forest reserve “Las Natolinsk1”

in Warsaw indicate that the traps with mycelium are high-

ly effective in catching mycetobiontic beetles. Numerous

Bonn zoological Bulletin 71 (1): 51-67

fruiting bodies of F: hepatica in that area resulted in a rel-

atively high abundance of the beetle 7riphyllus bicolor,

which was caught in traps with its mycelium. As many as

32 individuals of this rare mycetophagous species were

trapped there. Interestingly, in that reserve, between the

traps with mycelium of F) hepatica, traps with myceli-

um of another fungal species were located (G. frondosa,

found on oaks), but no individual 7: bicolor was caught

in them.

In the collected material, two autonomous faunistic

clusters of beetles of wood-decay fungi are noticeable:

one associated with brown rot fungi, and the other with

white rot fungi. The beetle communities attracted by saw-

dust substrate with brown rot fungi were dominated by

Cortinicara gibbosa, Rhizophagus bipustulatus, Cryp-

tophagus dentatus, and Trixagus carinifrons. The beetle

communities attracted by sawdust substrate with white

rot fungi were dominated by the same species except

C. dentatus, but with the addition of Dienerella ruficollis

and Aulonothroscus brevicollis. The three shared species

are beetles living on moulds or their predators. Three

beetle species were specific to the cluster composed of

beetles attracted by traps with white rot fungi: saprox-

ylophagous Ptilinus pectinicornis and mycetophagous

Scaphisoma agaricinum, and Clypastraea pusilla. These

beetles were caught exclusively in traps with mycelium

of white rot fungi. Pietka & Borowski (2011b) reported

that P. pectinicornis was very numerous (32.8% of the to-

tal catch) in beech stands in Swietokrzyski National Park.

©ZFMK

66 Jacek Pietka et al.

It was caught there in traps with sawdust substrate and

mycelium of a white rot fungus, 7rametes gibbosa (Pers. )

Fr. Leather et al. (2014) confirm that Prilinus pectinicor-

nis commonly associated with dry and exposed beech

wood was significantly associated with infested logs by

Hypholoma fasciculare (Huds.) P. Kumm., which also

causes white rot of wood.

The sawdust substrates with mycelium of wood-decay

fungi, prepared in laboratory conditions, constituted a

free and attractive ecological niche for many species.

CONCLUSIONS

The sawdust substrates with mycelium of wood-decay

fungi, used in traps, attract mycetophilic and mycetobi-

ontic beetles. In the study period (April-October 2008)

we caught 73 mycetophilic species and 60 mycetobion-

tic species of beetles. This bait type was effective in the

trapping of beetles associated with wood-decay fungi.

Two new species for the fauna of Poland were recorded

in this study: the mycetophilic Euplectus infirmus and the

mycetobiontic Corticarina pineti. In the tree-fungus bee-

tle community, mycetophages and zoophages feeding on

them were the major trophic groups, while mycetocoles

were the major microhabitat group. In the collected ma-

terial, two clusters of beetle species of wood-decay fun-

gi can be distinguished. The first one is associated with

brown rot fungi (Fistulina hepatica, Fomitopsis rosea),

and the other with white rot fungi (Bondarzewia mesen-

terica, Grifola frondosa, Hericium alpestre, H. coralloi-

des, Meripilus giganteus). It seems that this method can

be applied in studies on diversity of tree-fungus beetles,

valorisation of forests and monitoring of environmental

processes.

Acknowledgements. We thank Marek Stawski for his help in

preparing the NMDS analysis.

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BHL

Blank Page Digitally Inserted

Bonn zoological Bulletin 71 (1): 69-76

2022 Gorczyca J. et al.

https://do1.org/10.20363/BZB-2022.71.1.069

ISSN 2190-7307

http://www.zoologicalbulletin.de

Research article

urn:|sid:zoobank.org: pub:8A64E43A-B60B-45 15-BA2B-70115B67CBF8

The genus 7rynocoris Herring, 1976 (Heteroptera: Miridae: Cylapinae)

is no longer monotypic:

T. costaricaensis sp. nov. from Guanacaste, north-western Costa Rica

'2 Institute of Biology, Biotechnology and Environmental Protection, Faculty of Natural Sciences, University of Silesia in

Katowice, Bankowa 9, 40-007 Katowice, Poland

3Institute of Biology, University of Opole, Oleska 22, 45-052 Opole, Poland

4Service Public de Wallonie, DGO3, DEMNA, Av. Maréchal Juin, 23, BE-5030, Gembloux, Belgium, U.E.

“Corresponding author: Email: artur.taszakowski@us.edu.pl

'urn:lsid:zoobank.org:author:6E164D8D-B7D1-48E3-B771-C36E211AB7DC

2urn:|sid:zoobank.org:author: BB6E3148-D9BC-4DC1-A212-E99EC5F96E28

3urn:Isid:zoobank.org:author:845C8DAF-B265-4C4E-954E-6B8C44A EA27B

4urn:Isid:zoobank.org:author:308707E0-ED14-4AC2-AD3D-7A 1624020BDC

Abstract. A revised diagnosis of the genus 7rynocoris Herring, 1976 is presented and a new species, 7rynocoris costari-

caensis sp. nov., is described based on specimens collected in Costa Rica. Colour photographs of the adult and illustrations

of the male genitalia are provided for the new species. Scanning electron micrographs of selected structures of the new

species are also given.

Key words. Biodiversity, Fulviini, new species, Neotropics, plant bugs, taxonomy, true bugs.

INTRODUCTION

The genus 7rynocoris Herring, 1976 was described as

monotypic for 7. /Jawrencei Herring, 1976 based on spec-

imens collected in Panama (Herring 1976). The genus

was placed in the tribe Fulviini within Cylapinae (Her-

ring 1976). In addition to Fulviini, the subfamily Cylapi-

nae also includes representatives of the tribes Cylapini

and Vanniini in the Neotropical Region. Distribution and

biology of the New World Cylapinae still remain poorly

known, although several genera have been revised and

new taxa have been described recently (e.g., Chérot &

Carpintero 2016; Wolski 2017; Wolski et al. 2020).

Some representatives of Cylapinae (particularly in the

tribe Cylapini) are mycetophagous (Wheeler 2001), but

Trynocoris lawrencei probably feeds on larvae of beetles

that live in fungi (Herring 1976). Gorczyca et al. (2019)

redescribed 7rynocoris and T. lawrencei, and cited new

sampling localities in Panama, Mexico, Ecuador and Nic-

aragua.

While studying the material collected in Costa Rica in

the collection of the Division of Entomology, University

of Kansas Natural History Museum (Lawrence, Kansas,

USA; KUNHM), the authors found two specimens of a

new species of the genus 7rynocoris. In the present paper

we provide a revised diagnosis of the genus and describe

Received: 14.03.2021

Accepted: 18.02.2022

the new species. Furthermore, we compare the morphol-

ogy, including genitalia, of both species of 7rynocoris.

MATERIAL AND METHODS

The specimens were imaged using the following equip-

ment: Leica M205C stereo microscope with high diffuse

dome illumination Leica LED5000 HDI, Leica DFC495

digital camera and Leica application suite 4.12.0 soft-

ware; Leica DM 3000 upright light microscope with

Leica MC 190 HD digital camera and Leica Applica-

tion Suite ver. 4.13.0 software. SEM photographs were

obtained using Phenom XL field emission scanning

electron microscope at 10 and 15 kV accelerating volt-

age with a BackScatter Detector (BSD). Graphic editor

Adobe ®Photoshop ver. CS6 was used to edit the fig-

ures. Measurements were made with Leica application

suite ver. 4.12.0 software and are presented in millime-

ters (mm). The measured body parts were defined in

Wolski (2014). Dissections of male genitalia were done

according to Kerzhner and Konstantinov (1999). The

original description was used to identify the genus (Her-

ring 1976). Adult terminology used in the text follows

Schuh & Weirauch (2020).

Corresponding editor: X. Mengual

Published: 12.05.2022

70 Jacek Gorezyca et al.

The terminology of the male genital structures follows

Kerzhner & Konstantinov (1999), Konstantinov (2003),

Cassis (2008) and Schuh & Weirauch (2020).

RESULTS

Taxonomy

Trynocoris Herring, 1976

Trynocoris Herring 1976: 91 (as new genus) [type spe-

cies: Trynocoris lawrencei — original designation].

Trynocoris: Schuh, 1995: 38 (catalog), Schuh, R. T.

(2002—2013 On-line Systematic Catalog), Gorczy-

ca 2000: 49 (list of the genera of the tribe Fulviini),

Gorczyca 2006: 69 (catalog), Gorcezyca et al. 2019: 52

(redescription).

Diagnosis

Distinguished from the other genera of Cylapinae by the

following set of characters: body small (< 3.1 mm), oval

to elongate oval, dorsum densely and deeply punctured

(Figs 1A, 3A), covered with dense, scale-like setae, vary-

ing from linear to incrassate; second antennal segment

thickened, sometimes flattened laterally; the fourth an-

tennal segment subdivided (Fig. 1A—B; Gorczyca et al.

2019: fig. 1); cuneus short, apex of hemelytra at the level

of cuneal fracture bent downward; tarsi two-segmented

(Gorczyca et al. 2019: figs 10-12); male genitalia: pygo-

phore trapezoidal, its dorsal wall long, almost as long as

ventral wall, aperture of pygophore directed posteriorly;

aedeagus thin; ductus seminis long and thin; endosoma

membranous; left paramere with apical process broad in

dorsal view and with relatively long, thin process on ex-

treme apex; right paramere C-shaped its apical process

broad when viewed dorsally with more or less devel-

oped process in the middle of right lateral margin (Fig. 2;

Gorczyca et al. 2019: figs 17-20).

Remarks

This genus belongs to the tribe Fulviini and the morpho-

logical characters that warrant this placement as well as

the features indicating close similarity to some fulviines

were discussed by Gorczyca et al. (2019).

Herein we confirm the two-segmented tarsi with tar-

somere II undivided as one of the diagnostic features

of 7rynocoris. In Cylapinae, the number of the tarsal

segments is variable depending on the tribe. The repre-

sentatives of Bothriomirini, Cylapini, and Rhinomirini

possess three-segmented tarsi (Wolski 2017; Namyato-

va et al. 2019; Namyatova & Cassis 2016, 2019, 2021).

In contrast, in the tribe Fulviini, as circumscribed by

Gorczyca (2000), the number of tarsal segments is vari-

able depending on the genus. Genera Carvalhofulvius

Stonedahl & Kovac, 1995 (Stonedahl & Kovac 1995:

fig. 8), Cylapocoris Carvalho, 1954 (Wolski 2013),

Bonn zoological Bulletin 71 (1): 69-76

Howefulvius Schmitz & Stys, 1973 (Schmitz & Stys

1973: fig. 7), Peritropisca Carvalho & Lorenzato, 1978

(Wolski & Gorczyca 2014; Namyatova & Cassis 2019),

Sulawesifulvius Gorczyca, Chérot & Stys, 2004 (Gorczy-

ca et al. 2004: fig. 4; Wolski et al. 2017: fig. 10) and

others have two-segmented tarsi, as well as many spe-

cies of the genera Fulvius Stal, 1862 (Gorezyca 2000:

figs 23E, 27D; Yasunaga 2000: figs 10-11; Wolski et al.

2018: fig. 45) and Peritropis Uhler, 1891 (Wolski &

Henry 2012: fig. 81). Three-segmented tarsi are found

in Bironiella Poppius, 1909 (Namyatova et al. 2016:

fig. 19), Punctifulvius Schmitz, 1978 (Yasunaga 2000:

fig. 13; Namyatova & Cassis 2019: fig. 10n), Yamato-

fulvius Yasunaga, 2000 (Yasunaga 2000: figs 14-16) and

in some species of Peritropis (Namyatova et al. 2016:

191). Recently, Namyatova & Cassis (2019) pointed out

that most fulviines, which they included in their phyloge-

netic analysis, have three-segmented tarsi and suggested

that the two-segmented tarsi with the second tarsomere

divided observed by many previous authors are in fact

three-segmented tarsi. The question whether the tarsi ob-

served among others by Gorczyca & Eyles (1997: fig. 5),

Yasunaga (2000: fig. 12), Gorczyca (2002: 12) or Wol-

ski et al. (2018: fig. 53) have two or three segments re-

mains open. However, given the fact that many fulviines

possess two-segmented tarsi, with no subdivision of the

second segment, unlikely that all these observations are

incorrect. Taking into account the existing differences in

number of the tarsal segments among the fulviine gen-

era, we believe that the two-segmented tarsi are a helpful

character in defining the genus 7rynocoris in combina-

tion with other diagnostic features of this genus herein

presented.

Key to the species of the genus 7rynocoris Herring,

1976

1. Body covered with broadened, scale-like setae

(Fig. 3A-—B); second antennal segment weakly

flattened laterally, covered with thick, scale-like

ye] Fe Teh aaa ane Ae a T. lawrencei Herring, 1976

— Body covered with linear setae (Fig. 1); second

antennal segment not flattened, somewhat thickened

toward the apex, covered with simple setae (Figs 1A,

OS, Oe Ne ee eee eb i T. costaricaensis sp. nov.

Trynocoris costaricaensis sp. nov.

urn: lsid:zoobank.org:act: 17498A 48-3 133-4E24-98A9-FEQEDEES DD38

Figs 1-2, 3C, 4C—D

Diagnosis

The new species is similar to 7: Jawrencei (Figs 3A—B,

4A-B) but can be easily separated by the following char-

acter states: (1) Second antennal segment (Fig. 1) of fe-

male not foliaceous and laterally flattened, but basally

and medially cylindrical, apically slightly club-like, its

©ZFMK

Trynocoris costaricaensis, new species from Costa Rica 71

0.5mm

Fig. 1. Zrynocoris costaricaensis sp. nov., 2°, holotype (SM0210321 KUNHM-ENT). A. Dorsal view. B. Ventral view. C. Lateral

view.

72 Jacek Gorezyca et al.

Fig. 2. Trynocoris costaricaensis sp. nov., 3, paratype (SM0210194 KUNHM-ENT), genitalia. A-B. Left paramere. C—D. Right

paramere. E. Dorsal view of endosoma.

colour pattern more complex (basally brown, medially

orange-brown to orange, apically yellow versus evenly

red-brown except its dorsal and ventral black margins in

T: lawrencei). (2) Pilosity of second antennal segment

(Fig. 1) limited to long, suberect to erect setae, similar to

setae of third and fourth antennal segments (versus pilos-

ity of second antennal segment including scale-like se-

tae and thick, simple setae in 7: /awrencei). (3) Absence

of orange-red ‘ventral neck or ring’ easily visible under

the eyes in 7. Jawrencei. (4) Labium reaching ovipositor

(Fig. 1B—C) (versus in T: lawrencei reaching base of the

abdomen in the female, but the pygophore in the male).

(5) Body covered with setae curved, recumbent, bare-

ly widened apically, silvery bright under incident light

(Figs 1A, 3C) (versus scale-like, whitish, dull, obvious-

ly widened apically in T: lawrencei (Fig. 3A—B)). (6) In

males, the left paramere apical process in dorsal view

with apex curved toward the right side; right paramere

apical process in dorsal view with a moderately devel-

oped medial spine on right lateral margin (Fig. 2B, D).

Etymology

The specific epithet refers to the country where the type

locality is found, Costa Rica.

Type material

Deposited at the University of Kansas Natural History

Museum (KUNHM) (Lawrence, Kansas, USA).

Bonn zoological Bulletin 71 (1): 69-76

Holotype

COSTA RICA « 9; ’COSTA RICA Guanacaste / Cacao

Biological Station, 1050 m / 10°55'38" N, 85°27'7" W /

11 JUL 2000, leg. J.Ashe, R. Brooks, / Z.Falin

CRIABFOO 100 / ex. fogging fungus covering log //

SM0210321 / KUNHM-ENT’.

Paratype

COSTA RICA « 4, damaged and incomplete; COSTA

RICA: Guanacaste / Cacao Biological Station, 1050 m /

LO°S5°38"" N.t85927'7 Wel JUTE 2000) less J.Ashe;

R.Brooks, / Z.Falin CR1 ABFO0 100 / ex. fogging fungus

covering log // SM0210194 / KUNHM-ENT”’.

Description

Female

Measurements (in mm). Body (in dorsal view): Length:

3.1, width 1.58. Head: Interocular distance (vertex

width): 0.43, width of eye: 0.2, length of antennal seg-

ment I: 0.28, II: 0.63, III: 0.28, IV: 0.25. Pronotum: Me-

dial length: 0.6, posterior width of disk: 1.33, length of

lateral margin (between anterior and humeral angle): 0.6.

Scutellum: Length (mesoscutum excluded): 0.53, length

(mesoscutum included): 0.68, width: 0.73. Cuneus:

Length: 0.3, width at base: 0.2.

Colouration (Fig. 1A—C). Body black. Head. Vertex

(including narrow carina), frons and mandibular plates

black. Clypeus and maxillary plates reddish brown. Eyes

©ZFMK

Trynocoris costaricaensis, new species from Costa Rica 73

Fig. 3. A. Zrynocoris lawrencei Herring, 1976, dorsal view. B. 7: Jawrencei Herring, 1976, dorsal surface of the body (according

Gorczyca et al. 2019). C. T. costaricaensis sp. nov., 3, paratype (SM0210194 KUNHM-ENT), dorsal surface of the body.

grey. First antennal segment reddish with a yellowish ring

at basal third. Second antennal segment basally brown,

medially orange-brown to orange, apically yellow. Third

and base of fourth antennal segment brown, the apex

of fourth antennal segment black. Labium brown, the

first segment apically yellow. Thorax. Pronotum and

scutellum evenly black, pronotal collar reddish-brown.

Metafemur wide, reddish-brown with dark reddish spots.

Metatibia black with yellowish rings. Mesoscutum black,

the lateral part brown. Clavus black, the apex yellow to

orange. Endocorium black, its apex with a brownish me-

dian area and one yellowish spot at the inner corner. Exo-

corium brown with an elongate submedian longitudinal

black stripe and an apical yellow spot. Embolium brown.

Membrane grey. Cuneus orange-brown. Pro-, meso- and

metapleuron black to reddish black, shining. Abdomen.

Black.

Structure, texture and vestiture (Fig. 1A—C). Body

oval to elongate-oval. Head. Declivous. Frons and vertex

narrowly and shallowly punctate. Vertex posterior area

slightly raised, narrowly carinate. Eyes relatively small,

contiguous to the pronotal collar (partially hidden by the

eyes laterally). The first antennal segment with relatively

short, suberect setae, particularly apically. Second anten-

Bonn zoological Bulletin 71 (1): 69-76

nal segment bearing long, suberect to erect setae, longer

apically, similar to setae of third and fourth antennal seg-

ments. The fourth antennal segment subdivided. Thorax.

Pronotal collar very short. Pronotal lateral and posterior

margins carinate. Pronotal disk punctate, the punctation

wider and deeper than dorsal punctation of the head. Pro-

notal callosities barely visible. Mesoscutum wide, pos-

teriorly convex and laterally marginate. Punctation of

mesoscutum and scutellum similar to pronotal punctation

(Fig. 3C). Claval vein raised. Clavus and corium with

two different type of setae: one short, black, recumbent

the other slightly longer and curved, recumbent, barely

widened apically, silvery bright under incident light, in

rows on the basal part of the corium. Cuneus elongate

oval, oblique, with sparse, very short, stiff, black setae.

Abdomen. Thickened, not reaching the apex of mem-

brane. Female genitalia. (Fig. 4C—D). Bursa copulatrix

(= vagina, = genital chamber) totally membranous, de-

void of sclerite. Sclerotized rings not visible in available

specimens. Posterior wall not easily recognizable.

Male

Similar to female but smaller in size. The only male spec-

imen is seriously damaged, headless and without trochan-

©ZFMK

74 Jacek Gorezyca et al.

Fig. 4. Trynocoris spp., female genitalia in different views. A—-B. Trynocoris lawrencei Herring, 1976. C—D. 7: costaricaensis

Sp. NOV.

ters and femora. Thorax. Pronotum, mesoscutum and

scutellum entirely black. Length of pronotum 0.50 mm,

anterior margin 0.50 mm, lateral margins 0.65 mm, pos-

terior margin 1.30 mm. Hemelytra black, covered with

shining setae. Cuneus brown to dark brown tinged with

orange at the base and along the external margin. Body

ventrally chestnut to dark brown, coxae chestnut. Male

genitalia (Fig. 2). Left paramere hook-shaped; apical

process thin 1n lateral view, broadened basally, narrowed

toward the apex, apical process broad in dorsal view, its

apex curved toward the right side; paramere body with a

bundle of long, thick setae dorsally, paramere body when

viewed laterally with inner margin weakly convex basal-

ly and nearly straight and inner margin strongly convex

Bonn zoological Bulletin 71 (1): 69-76

medially. Right paramere C-shaped; apical process thin

and curved in lateral view, in dorsal view apical process

broad, nearly ellipsoid, with the moderately developed

process in the medial part of the right lateral margin. Ae-

deagus narrow, ductus seminis thin and long, endosoma

membranous.

Distribution

Costa Rica.

Biology

Unknown, but the specimens of this species were collect-

ed from a log covered with fungi. 7. /awrencei was also

collected in a similar habitat.

©ZFMK

Trynocoris costaricaensis, new species from Costa Rica WS

Acknowledgements. We would like to acknowledge Katrina L.

Menard (University of Connecticut) for finding the specimens

in the University of Kansas collection used in this study. We

also want to thank the reviewers and the editor, Ximo Mengual

(Zoologisches Forschungsmuseum Alexander Koenig, Bonn,

Germany) for all the comments that improved this manuscript.

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Bonn zoological Bulletin 71 (1): 77-85

2022 Gani A. et al.

https://do1.org/10.20363/BZB-2022.71.1.077

ISSN 2190-7307

http://www.zoologicalbulletin.de

Research article

urn:|sid:zoobank.org: pub:2EB89B65-13FE-4194-AA5F-06C172C78554

A new endemic species of pelvic-brooding ricefish

(Beloniformes: Adrianichthyidae: Oryzias)

from Lake Kalimpa’a, Sulawesi, Indonesia

Abdul Gani', Novian Suhendra’, Fabian Herder’, Julia Schwarzer’, Jan Moéhring*, Javier Montenegro®,

Muh. Herjayanto’ & Daniel F. Mokodongan**

'Faculty of Fisheries, Muhammadiyah Luwuk University, Banggai Regency, Central Sulawesi, Indonesia

?Fish Quarantine Station, Quality Control and Safety of Fishery Products, Palu, Central Sulawesi, Indonesia

3° Leibniz Institute for the Analysis of Biodiversity Change (LIB) — Museum Koenig, Section Ichthyology Adenauerallee 127,

D-53113 Bonn, Germany

*Leibniz Institute for the Analysis of Biodiversity Change (LIB) — Museum Koenig, Section Evolutionary Genomics,

Adenauerallee 127, D-53113 Bonn, Germany

°Japan Agency for Marine-Earth Science and Technology, Kanagawa, Yokozuka, Japan

’Department of Fisheries Science, Faculty of Agriculture, University of Sultan Ageng Tirtayasa, Serang,

Banten 42124, Indonesia

’Museum Zoologicum Bogoriense, Research Center for Biosystematics and Evolution,

the National Research and Innovation Agency, Cibinong, Indonesia

“Corresponding author: Email: daniel mokodongan@yahoo.com

'urn:Isid:zoobank.org:author: EE6892C6-F3FB-4794-BE 1 8-323078AEAE58

2urn:Isid:zoobank.org:author:8AA46AC3-1D92-4264-899 1-388F40217B6C

3urn:Isid:zoobank.org:author:A9906501-CD1C-4030-AA 13-CF80F98BA 106

4urn:Isid:zoobank.org:author: 70E7E37B-39A0-449E-9CF9-BBBS51E83C577

Surn:Isid:zoobank.org:author:8AB210AF-11F4-452C-9470-159BD187160D

Surn:Isid:zoobank.org:author:01CBD81E-C4DD-45D5-A19D-A3878DDOC9IBC

7urn:lsid:zoobank.org:author:4 1 A2A48D-1F92-4DB8-9ED8-D7B397691F5A

§urn:Isid:zoobank.org:author: DS B8COEF-47B1-4CE1-9850-36A6FDOSE1 88

Abstract. Oryzias kalimpaaensis sp. nov. is a new species of the genus Oryzias Jordan & Snyder, 1906, endemic to Lake

Kalimpa’a in Lore Lindu National Park, Central Sulawesi, Indonesia. The species is a pelvic-brooder, a non-monophyletic

group of ricefishes in which females carry an egg cluster until hatching. Pelvic-brooding is known from only a few taxa,

and the new species reported here is only the fifth pelvic-brooding species known so far. Oryzias kalimpaaensis sp. nov.

differs from all others Oryzias species by the following combination of characters: 61—67 scales in lateral line, 11-13

dorsal-fin rays, 11-13 pectoral-fin rays, body depth 16.0—22.2% SL, and length of head 30.1—33.7% SL. It is distinguished

from all species of the genus Adrianichthys Weber 1913, by its small size (max. 52.8 mm SL). A molecular phylogeny

based on mitochondrial ND2 sequences supports the distinctiveness of O. kalimpaaensis sp. nov. Oryzias kalimpaaensis

sp. nov. is closely related to pelvic-brooding O. eversi Herder, Hadiaty & Nolte 2012 endemic to Tilanga Pond in Tana

Toraja, and to the two species of Lake Lindu in Central Sulawesi, O. sarasinorum (Popta 1905) and O. bonneorum Parenti

2008. As Lake Kalimpa’a is a popular destination for nature tourism, anthropogenic pressure is high. The presence of

invasive fish species in the lake and parasites on collected specimens support thiss assumption.

Key words. Endemic, Oryzias, freshwater fish, Kalimpa’a, pelvic-brooder.

INTRODUCTION

Sulawesi is one of the five largest islands in Indonesia,

and harbors a rich fauna of freshwater fishes, including

endemic lake- and stream-dwelling species (Kottelat

1990a, 1990b; Miesen et al. 2016; Hadiaty 2018). The

island is a hotspot of ricefish diversity (Adrianichthyidae;

reviewed by Hilgers & Schwarzer 2019), and almost all

ricefish species known from it are endemic. Distribution

areas of most species are small, typically restricted to sin-

Received: 18.03.2022

Accepted: 24.06.2022

gle lakes like those of the Malili Lakes system, Lake Lin-

du, or Lake Poso; some ricefishes are even restricted to

tiny pools or ponds, or stretches of rivers (Parenti 2008;

Herder & Chapuis 2010; Parenti & Hadiaty 2010; Herder

et al. 2012; Parenti et al. 2013; Mokodongan et al. 2014).

The genus Oryzias Jordan & Snyder, 1906 is the most

diverse and consists of thirty-two species; four species

restricted to Lake Poso make up the endemic genus Adri-

anichthys Weber, 1913.

Corresponding editor: R. Peters

Published: 30.06.2022

78 Abdul Gani et al.

OT, fn s meth

Fig. 1. Lake Kalimpa’a: type locality of Oryzias kalimpaaensis sp.

Reproduction of Sulawesi ricefishes can be assigned

to two general strategies, transfer-brooders, 1.e., spe-

cies that deposit fertilized eggs on a substrate, as it is

the case with most adrianichthyids, and pelvic-brooders

(Parenti 2008; Spanke et al. 2021). In pelvic-brooding

ricefishes, females carry the fertilized eggs until the fry

hatches, protected in a ventral concavity, and covered by

elongated pelvic fins (Parenti 2008; Parenti & Hadiaty

2010; Parenti et al. 2013; Mokodongan et al. 2014; Man-

dagi et al. 2018; Spanke et al. 2021). Interestingly, pel-

vic-brooding occurs in species of both genera, Oryzias

and Adrianichthys. So far, a total of four pelvic-brood-

ing ricefishes has been reported: A. oophorus (Kottelat,

1990), A. poptae Weber & de Beaufort, 1922, O. sara-

sinorum (Popta, 1905) and O. eversi Herder, Hadiaty &

Nolte, 2012 (Kottelat 1990a; Parenti 2008; Herder et al.

2012; Mokodongan & Yamahira 2015).

Here we describe a new species of pelvic-brooding

Oryzias endemic to a small lake in Central Sulawesi,

Lake Kalimpa’a, which is located within Lore Lindu Na-

tional Park, close to Lake Lindu. This discovery brings

the number of recognized ricefish species on Sulawesi to

23, including at least five pelvic brooders (Parenti 2008;

Herder & Chapuis 2010; Parenti & Hadiaty 2010; Herder

et al. 2012; Parenti et al. 2013; Mokodongan et al. 2014:

Mandagi et al. 2018; Utama et al. 2022).

MATERIAL AND METHODS

Species description and specimens of the new Oryzias

were collected from Lake Kalimpa’a (Fig. 1) in Cen-

tral Sulawesi, at 1°19'36.1" S, 120°18'26.1" E, approx-

imately 23 km east of Lake Lindu (Fig. 2). The fish were

caught with a hand net 13 Sep. 2020. Immediately after

Bonn zoological Bulletin 71 (1): 77-85

nov., in Central Sulawesi, Indonesia.

capture, specimens of both sexes were photographed in

a small tank to document life coloration. For morpho-

logical investigations, 18 individuals (10 males and 8

females) were anesthetized using ice slurry, preserved

in 5% formalin for morphological analyses and later

transferred to 70% ethanol for storage. Specimens are

deposited in the fish collections of Museum Zoologicum

Bogoriense (MZB), BRIN in Cibinong; Museum Koe-

nig, Bonn (ZFMK); the Zoological Reference Collection

(ZRC) of the Lee Kong Chian Natural History Museum,

National University of Singapore.

Following Parenti (2008) we distinguished the speci-

mens as genus Oryzias by several characters including

maximum size in adult fish 52.8 mm SL. Specimens

were compared to all known Sulawesi Oryzias species,

with emphasis to those from Central Sulawesi and oth-

er pelvic-brooding species (Lake Lindu: O. sarasinorum

and O. bonneorum; Tilanga Pond: O. eversi; Lake Poso:

O. nebulosus Parenti & Soeroto 2004, O. nigrimas

Kottelat 1990, O. orthognathus Kottelat 1990, and Lake

Tiu: O. soeroto Mokodongan, Tanaka & Yamahira 2014).

Assessment of morphological traits follows Herder et al.

(2012). Number of fin rays and scales were counted di-

rectly from each individual using a stereo-microscope

(SWIFT- S306S-20-2L).

To investigate phylogenetic relationships, NADH

dehydrogenase subunit 2 (ND2) gene region was se-

quenced using two primers ND2L (5’-GGGCCCCAT-

ACCCCAAACATGTTGG-3’) and ND2H (5’-TTAAT-

TAAAGTGTCTGTTTTGC-3’) following Mokodongan

and Yamahira (2015). DNA was extracted from the right

pectoral fin of two different uncatalogued individuals of

O. kalimpaaensis sp. nov. using standard protocol Qia-

gen DNeasy Blood & Tissue Kit. PCR conditions were

as following: 94 °C for 3 min for denaturation, 35 cycles

©ZFMK

A new endemic species of pelvic-brooding ricefish from Lake Kalimpa’a, Sulawesi, Indonesia the)

of 94 °C for 30 s, 50 °C for 30 s, and 72 °C for 60 s, with

a final extension of 72 °C for 2 min. Positive amplifica-

tion was verified on a 1% agarose electrophoresis gel.

PCR products were then sent to PT. Genetika Science in

Jakarta (http://actagen.com) for sequencing. Sequence

information for the new species were aligned together

with sequences of 21 other species (two Adrianichthys

and 19 Oryzias) obtained from Mokodongan & Yama-

hira (2015); accession numbers LC051687-LC051739.

Alignments were done via ClustalW ver. 1.4 (Thompson

et al. 1994) and manually curated. The final alignment

consisted of 1046 bp. Sequence data of O. kalimpaaen-

sis sp. nov. and O. bonneorum were deposited in DNA

Data Bank of Japan (DDBJ) with accession number

LC669549-LC669550, and LC685422-LC685423, re-

spectively. Phylogenetic reconstructions using Maximum

Likelihood were performed in raxmlGUI 2.0.6 (Edler

et al. 2021), following the substitution model selected as

best fitting (GTR-I-G) by ModelTest-NG (Darriba et al.

2019) and with 1000 bootstrap replicates conducted.

Cololabis saira (Brevoort 1856) (Beloniformes: Scomb-

120°2'40"E 120°5'20"E 120°8'0"E 120°10'40"E 120°13'20"E

2355

Nokilalaki Mount

1°14'40"S

4°22'40"S

1°25'20"S

120°2'40"E 120°5'20"E 120°8'0"E 120°10'40"E 120°13'20°E

120°16'0"E

eresocidae) was used as outgroup with DDBJ accession

number AP002932.

Taxonomy

Oryzias kalimpaaensis sp. nov.

urn: Isid:zoobank. org: act:045555C9-BD02-445A-AFC0-1D2958A806EE

Lake Kalimpa’a Ricefish

Figs 3-4

Holotype

MZB. 26462 (Figs 3 (top), 4), 3, 41.9 mm SL, Indonesia,

Sulawesi Tengah, Regency of Lore Utara, District Poso,

Lake Kalimpa’a, 13 Sep. 2020, Abdul Gani and Novian

Suhendra.

Paratypes

MZB 26463-26466; 26528-26529, 4 33 (44.1-50.2 mm

SL), 2 99 (41.1-42.4 mm SL); ZFMK ICH-128486—

128492, 4 3 (43.7-49.1 mm SL), 3 9° (41.8-43.6 mm

SL); ZRC 62531, 2 3d (47.3-47.6 mm SL), 2 2° (46.9-

52.8 mm SL), collected with the holotype.

120°18'40"E 120°21'20"E 120°24'0"E 120°26'40"E 120°29'20"E

SUS 12F30S F260S O3T30N

1°14'40"S

Kalimpa'a Lake

Dp.

1°17'20"S

1°22'40"S

4°25'20"S

Scale 1: 100.000 |

120°18'40"E 120°21'20"E 120°24'0"E 120°26'40"E 120°29'20"E

Fig. 2. Map of streams and lakes in Lore Lindu national park (Central Sulawesi, Indonesia) including the type locality of Oryzias

kalimpaaensis sp. nov. (map by Moh. Arif Rahman). The location of the lake Kalimpa’a in Sulawesi is marked by a green dot on

the map in the upper right corner.

Bonn zoological Bulletin 71 (1): 77-85

©ZFMK

80 Abdul Gani et al.

Fig. 3. Living specimens of Oryzias kalimpaaensis sp. nov., & (top) and 2 (bottom) from Lake Kalimpa’a, Central Sulawesi,

Indonesia.

Diagnosis. Oryzias kalimpaaensis sp. nov. is a pel-

vic-brooding ricefish with pronounced sexual dimor-

phism. Females share with females of other pelvic-brood-

ing ricefishes (O. eversi, O. sarasinorum, A. oophorus)

morphological structures that enable the maternal fish to

carry bundles of fertilized eggs. The eggs remain con-

nected to the female by filaments, and are carried in a

ventral concavity present in females but not in males.

Oryzias kalimpaaensis sp. nov. differs from all other

Oryzias species from Sulawesi by unique lateral line

scale counts (60-67 in O. kalimpaaensis sp. nov. Vs.

70-75 in O. sarasinorum, vs. <58 in all remaining Sula-

wes! Oryzias). It has a deeper body than O. sarasinorum

(16.0—22.2% SL vs. 13-15% SL). Oryzias kalimpaaensis

sp. nov. has more dorsal-fin rays (11-13) than O. hadiat-

yae Herder & Chapuis 2010 (8-10), O. celebensis (Weber

1894) (8-10), O. woworae Parenti & Hadiaty 2010 (8),

O. asinua (7-9), O. wolasi Parenti, Hadiaty, Lumban-

tobing & Herder 2013 (7-9) and O. dopingdopingensis

Mandagi, Mokodongan, Tanaka & Yamahira 2018 (8-9),

and more pectoral-fin rays (11-13) than O. eversi (10).

In contrast to ricefishes of the O. woworae species

group (SE Sulawesi: O. woworae, O. asinua, O. wola-

si) O. kalimpaaensis sp. nov. does not exhibit red color

Bonn zoological Bulletin 71 (1): 77-85

in fins or bluish body coloration. Black blotches or lines

on the lateral side of body are characteristic for O. cel-

ebensis (Parenti, 2008), but absent in O. kalimpaaensis

sp. nov.. Oryzias kalimpaaensis sp. nov. has a relative-

ly long head (30.1-33.7% SL) compared to O. sara-

sinorum (29% SL), O. eversi (28.4—-30.7% SL), O. soer-

otoi (21.4—25.4% SL), O. orthognathus (22—26.1% SL),

O. nigrimas (21.5—25% SL), O. nebulosus (23-26% SL),

O. marmoratus (Aurich 1935) (24.0-27.2% SL),

O. matanensis (Aurich 1935) (25-29% SL), O. pro-

fundicola Kottelat 1990 (22—25.4% SL), O. celebensis

(24—26% SL), O. woworae (24-29% SL), O. asinua (25—

30% SL), O. wolasi (25-30% SL) and O. dopingdopin-

gensis (25.8—29.1% SL). Its anal-fin base 1s short (19.2—

24.0% SL) compared to O. soerotoi (24.1-30.5% SL),

O. orthognathus (26.2-31.6% SL), O. nigrimas (24.5-

29.9% SL), O. nebulosus (25-29% SL), O. marmoratus

(31.4-36.9% SL), O. matanensis (30.5—35.0% SL) and

O. profundicola (37.4—41.4% SL). The snout concavity

characterizing O. hadiatyae and O. orthognathus is ab-

sent in O. kalimpaaensis sp. nov..

Oryzias kalimpaaensis sp. nov. is distinguished from

Adrianichthys spp. by its smaller adult body size (max-

imum size 52.8 mm SL, vs. 200 mm) (Parenti, 2008),

©ZFMK

A new endemic species of pelvic-brooding ricefish from Lake Kalimpa’a, Sulawesi, Indonesia 81

Fig. 4. Preserved specimens of Oryzias kalimpaaensis sp. nov., holotype (MZB 26462), 4, 41.9mm SL (top); paratype (MZB 26466),

2, 42.4 mm SL (bottom).

and differs from all Adrianichthys except A. oophorus by

having less anal-fin rays (20—22 vs. >23). From A. oo-

phorus, it is distinguished by having more dorsal-fin rays

(11-13 vs. 10) and longer head (30.1—33.7% SL vs. 25—

27% SL (Parenti, 2008).

Etymology. The species epithet, ‘kalimpaaensis’, de-

notes the occurrence of this species in Lake Kalimpa’a,

Central Sulawesi, the type locality.

Description. See Fig. 3 and Fig. 4 for general appear-

ance in lateral view and table 1 for morphometric data.

Female with pronounced abdominal concavity between

pelvic fin and anal fin covered by adpressed pelvic fins

(11.7—19.0% SL). Body compressed laterally, body depth

16.0—22.2% SL. Caudal fin truncate; principal caudal-fin

rays 1,4/5,1; procurrent caudal-fin rays 6/6—6/8. Mouth

supra-terminal, lower and upper jaw equal or lower jaw

slightly longer than upper jaw. Length of caudal pedun-

cle 10.3-14.4% SL, depth of caudal peduncle 10.0-

12.1% SL. 11-13 dorsal-fin rays; 20-22 anal-fin rays;

5—6 pelvic-fin rays; 11-13 pectoral-fin rays.

Head length 30.1-33.7% SL and eye diameter 25.0—

29.6% HL. Dorsal head profile appears slightly concave

just above the orbit to the nape. Dorsal body profile rel-

atively straight without noticeable arch from nape to

dorsal-fin origin. Ventral body profile relatively straight,

with slight arching from head to anal. Genital papilla sin-

gle lobed in both sexes.

Live coloration. Body yellowish-brown with a

brown-greenish lateral line, more pronounced in females

than in males. Belly and throat light yellowish to white.

Bonn zoological Bulletin 71 (1): 77-85

7-10 faint blackish bars on lateral side of body. Dorsal

surface of head blackish, extending posteriorly as narrow

black dorsal stripe to dorsal-fin origin. Opercle with sil-

ver greenish sheen. Membranes of paired and unpaired

fins hyaline, rays light cream colored. Base of dorsal fin

yellowish. Anal-fin base with narrow blackish stripe.

Males with black submargin in dorsal and anal fin. Fe-

males with blackish submarginal band in anal fin, sub-

marginal marking in dorsal fin black, submargin lacking

in females. Caudal-fin base yellowish, followed posteri-

orly by a faint blackish bar. Dorsal and ventral caudal-fin

margins yellowish to orange, more pronounced in males

than in females. Courtship coloration unknown (Fig. 3).

Color in alcohol. Body of males and females light

yellowish-brown. 7—10 faint and irregular dark brown

to blackish bars on lateral body in both sexes. Males

and females with a faint blackish lateral stripe on later-

al midline, extending from uppermost posterior opercle

to caudal-fin base. Belly blackish grey in males, whitish

in females. Throat whitish in males and females. Dorsal

surface of head blackish in males and females, extending

posteriorly as narrow blackish dorsal stripe. Unpaired

fins whitish. Anal-fin base with narrow blackish stripe.

Faint blackish submargin in anal fin. Blackish submargin

present in dorsal fin in males, but not in females. Base of

caudal fin yellowish in both sexes (Fig. 4).

Sexual dimorphism. Females have a slightly wider

body compared to males (body width: 12.3-14.1% SL in

females vs. 11.7—13.4% SL in males). The single-lobed

genital papilla of males is tubular and slender compared

©ZFMK

82 Abdul Gani et al.

to the more rounded female papilla. Adult males have

elongated rays in dorsal and anal fin, a typical sexual di-

morphism in ricefishes. Length of dorsal fin extends be-

yond caudal-fin base in adult males (21.2-34.4% SL in

males vs. 14.4—24.9% SL in females). Elongated fin-rays

are absent in dorsal and anal fins of females. Females

have longer (13.7-19.0% SL vs. 10.3-12.7% SL) pel-

vic fins compared to males and a pronounced abdominal

concavity between anal fin and pelvic fin; egg clusters

in the concavity are supported by the pelvic fins (Fig. 3;

Fig. 4). Due to the less pronounced concavity, males

have a larger relative body depth at anal-fin origin than

females (19.3—22.2% SL vs. 16.0—-20.3% SL). See ‘Live

coloration’ for coloration of male and female specimens.

Reproduction. Oryzias kalimpaaensis sp. nov. is a

‘pelvic-brooder’. Pelvic-brooding is defined by females

carrying a cluster of eggs, connected by attaching fila-

ments to their gonoduct, until the fry hatches (Kottelat

1990a). All pelvic-brooding ricefish species described

to date (A. oophorus, A. poptae, O. eversi and O. sara-

sinorum) share certain external female-specific morpho-

logical characteristics like elongated pelvic fins (com-

pared to their respective males and to all transfer-brooding

species) and the presence of a ventral concavity (Herder

et al. 2012; Mokodongan & Yamahira 2015; Spanke et al.

2021). Female specimens of O. kalimpaaensis sp. nov.

carry their eggs on the abdominal concavity covered by

the pelvic fins. As described for the other pelvic-brood-

ing species (Kottelat 1990a; Iwamatsu et al. 2008; Herder

et al. 2012), the eggs do not adhere to each other, and are

suspended by attaching filaments to the female’s genital

pore. Approximately 24 eggs were counted from one fe-

male specimen depicted in Fig. 4. The eggs are clearly

developed and partially pigmented with embryos visible.

The size of each fertilized egg is about 2.19+0.10 mm in

diameter.

Distribution and habitat. Oryzias kalimpaaensis

sp. nov. is known only from Lake Kalimpa’a, about

22 km from Lake Lindu, Central Sulawesi, situated at

ca. 1,660 m above sea level. The lake is relatively small,

with the longest distance across the surface being ap-

proximately 300 meters from Southwest to Northeast.

The lake is used by local people as a place for nature

tourism. At time of sampling, most female O. kalim-

paaensis sp. nov. observed were carrying eggs. The

lake habitat is characterized by calm water with sandy

and muddy substrate with vegetation dominated by the

weed Phragmites karka (Retz.) (Cyperales: Poaceae).

The depth of Lake Kalimpa’a is about 11 meters (D. H.

Kristianto & Wantoko Staff of BBTNLL, pers. comm.)

and water temperature was 22 °C with pH 5-6.5 and DO

13.9 Mg/L, measured during daytime. There is a main

inlet of Basakura Stream near the collection site and four

more inlets of small streams around the lake. The Lake

outlet is connected to the Sopu River, a tributary of the

Palu River, which drains into the Makassar Strait. Other

Bonn zoological Bulletin 71 (1): 77-85

species present in the Lake Kalimpa’a were introduced

african tilapia Oreochromis sp. (Cichliformes: Cichlidae)

and snakeskin gourami 7richopodus pectoralis Regan

1910 (Anabantiformes: Osphronemidae), and native eel

Anguilla sp. (Anguilliformes: Anguillidae).

Phylogenetic relationships. The resulting ML phylog-

eny (Fig. 5) is largely congruent with the consensus tree

of Mokodongan & Yamahira (2015) in all well supported

branches, including the monophyly of all known Oryzias

species from Sulawesi (the “Oryzias celebensis species

group’). The two O. kalimpaaensis sp. nov. individuals

(DDBJ accession numbers: LC669549—LC669550) clus-

ter together and form the sister-group to the two other

pelvic-brooding Oryzias species (O. eversi and O. sar-

asinorum) and O. bonneorum. Oryzias sarasinorum

appears paraphyletic based on our phylogeny, as one

individual clusters with O. bonneorum. Sister-group to

this clade is the river-dwelling transfer-brooding spe-

cies O. dopingdopingensis. Interestingly, O. bonneorum

formed a clade together with the pelvic-brooding Oryzias

species, although it is still unclear whether this species is

a pelvic-brooder or not.

DISCUSSION

Ricefishes (Beloniformes: Adrianichthyidae) are a

strikingly diverse group of fishes, inhabiting riverine,

lacustrine, and brackish water habitats (Parenti 2008;

Hilgers & Schwarzer 2019). On Sulawesi, they show a

great diversity in coloration, shape, adaptations to diver-

gent habitat conditions and belong to two genera (Adri-

anichthys and Oryzias) with two main reproductive strat-

egies: transfer-brooding and pelvic-brooding. To date

pelvic-brooding is described in four species (A. oopho-

rus, A. poptae, O. sarasinorum and O. eversi), contrasted

by all other ricefish species that are either transfer-brood-

ing or have an unknown reproductive status (1.e., O. bon-

neorum, A. roseni Parenti & Soeroto 2004 and A. kruyti

Weber 1913). More undiscovered diversity appears like-

ly, as ricefishes tend to inhabit small isolated habitats (see

Parenti & Hadiaty 2010). Several species of Sulawesi

ricefishes are known only from water bodies with small

surface area, 1.e., O. soerotoi from Lake Tiu (Mokodon-

gan et al. 2014), O. hadiatyae from Lake Masapi, a small

satellite lake of Lake Towuti (Herder & Chapuis 2010),

O. eversi from Tilanga pond, a tiny karst pool (Herder

et al. 2012), as well as the new species described herein.

Oryzias kalimpaaensis sp. nov. is known only from the

Lake Kalimpa’a, a small upland lake at ~1660 m above

sea level. This is the highest altitude at which ricefish-

es have so far been recorded in Sulawesi. Distance be-

tween Lake Kalimpa’a and the closest habitat of other

pelvic-brooding ricefishes — Lake Lindu, the habitat of

O. sarasinorum and O. bonneorum — is only ~22 km

(Fig. 2). However, both are separated by a substantial

©ZFMK

A new endemic species of pelvic-brooding ricefish from Lake Kalimpa’a, Sulawesi, Indonesia 83

99)

Cololabis saira

O. marmoratus (Lake Lantoa A)

O. marmoratus (Laka Lantoa B)

O. marmoratus (Lake Mahalona A)

987! 0. marmoratus (Lake Towuti A)

93]! O. marmoratus (Lake Mahalona B)

O. hadiatyae (Lake Masapi A)

87| | O. hadiatyae (Lake Masapi B)

O. profundicola (Lake Towuti B)

O. profundicola (Lake Towuti A)

a O, marmoratus (Lake Towuti B)

100; O. matanensis (Lake Matano B)

O. matanensis (Lake Matano A)

‘0. nebulosus (Lake Poso B)

O. nebulosus (Lake Poso A)

O. nigrimas (Lake Poso A)

100) O. soerotoi (Lake Tiu A)

O. soerotoi (Lake Tiu B)

O. bonneorum (Lake Lindu 5)

O. oversi (Tilanga Fountain B)

1000 Sarasinorum (Lake Lindu A)

gg] L\y O. kalimpaaensis (Lake Kalimpa‘a m)

O. kalimpaaensis (Lake Kalimpa’‘a f)

Q. dopingdopingensis (Doping-doping River A)

O. dopingdopingensis (Doping-doping River B)

O. celebensis (Malino River B)

O. celebensis (Limbangang River B)

°8] 0. celebensis (Malino River A)

O. celebensis (Walanae River A)

O. celebensis (Walanae River B)

O. woworae (Fotuno Fountain B)

100] O. woworae (Fotuno Fountain A)

O. woworae (Waleale River B)

87/10. woworae (Waleale River A)

O. wolasi (Wolasi River A)

O. asinua (Asinua River A)

100} | O. asinua (Asinua River B)

O. wolasi (Moramo Waterfall A)

97'0. wolasi (Moroamo Waterfall B)

82 garA. oophorus (Lake Poso A)

99 A. oophorus (Lake Poso B)

agp A. poptae (Lake Poso A)

A. poptae (Lake Poso B)

O. javanicus (Java)

98+ O. javanicus (Muna)

100) ‘0. javanicus (Sulawesi)

O. haugiangansis (Vietnam)

0. latipes (Okinawa)

O. sakaizumii (Okinawa)

O. curvinotus (Hong Kong)

Fig. 5. Maximum Likelihood (ML) phylogenetic tree with the bootstrap values from 1000 bootstrap replicates. Oryzias kalim-

Paaensis sp. nov. is marked in red.

barrier, the Nokilalaki mountain (2357 m elevation).

Geographic distance to pelvic-brooding Adrianichthys

exceeds 56 km (Lake Poso: A. oophorus, A. poptae) and

to O. eversi (Tilanga Pond) the distance is 194 km.

Based on our phylogenetic analyses, O. kalimpaaen-

sis sp. nov. is closely related to both, the Lake Lindu

ricefishes, and O. eversi (Fig. 5). Its placement within

the clade of pelvic- brooding Oryzias appears plausible;

however, the reproductive biology of O. bonneorum re-

mains undocumented (Parenti 2008). The new species is

distinguished from other ricefish species by several mor-

phological characters, supporting that O. kalimpaaensis

Sp. nov. is a distinct species.

Although the reproductive ecology of O. kalimpaaesis

sp. nov. has not been studied in detail yet, records of fe-

male O. kalimpaaensis sp. nov. with developing eggs on

their belly (Fig. 3, Fig. 4 bottom) indicate — in line with

the phylogenetic and morphological evidence — that it is

a pelvic-brooding species. As in the other pelvic-brood-

ing species, females show a set of adaptations enabling

brooding, namely elongated pelvic fins, presence of a

ventral concavity and long filaments attaching the eggs

to the genital pore.

Lake Kalimpa’a is located in the Lore Lindu National

Park, and is a popular destination for nature tourism. Pro-

tection of the flora and fauna inhabiting the lake should

Bonn zoological Bulletin 71 (1): 77-85

be implemented, especially with regard to endemic spe-

cies, like O. kalimpaaensis sp. nov.. The most substantial

threats to Sulawesi’s endemic freshwater fishes are exotic

fish introduction, eutrophication, overfishing, and pollu-

tion (Parenti & Soeroto 2004, Herder et al. 2022). Lake

Kalimpa’a already contains non-native fishes such as

African tilapia (Oreochromis sp.) and snakeskin gourami

(Trichopodus pectoralis) (A. Gani & N. Suhendra, pers.

obs.), which might affect together with the direct anthro-

pogenic impact (e.g., through nature tourism) the lake

flora and fauna. We found anchor worms Lernaea sp. in

a female O. kalimpaaensis sp. nov., which indicates that

the population might already suffer stressful conditions;

parasites and disease are one of the causes of the decline

in other endemic lake fishes in Sulawesi (Kottelat 1990a;

Herder et al. 2022).

Comparative material examined

Oryzias asinua: MZB 21464, 2, Indonesia, Sulawesi

Tenggara, District Asinua; Oryzias bonneorum: MZB

15499, holotype, Indonesia, Sulawesi Tengah, Lake Lin-

du; Oryzias celebensis: MZB 2688, 4, MZB 5862, 3, In-

donesia, Sulawesi Selatan, Regency of Maros Indonesia:

Oryzias eversi. MZB 20780, holotype, MZB 20781, 1

paratype, Indonesia, Sulawesi Selatan, District Rante-

©ZFMK

Abdul Gani et al.

Table 1. Meristic and morphometric data of Oryzias kalimpaaensis sp. nov. from Lake Kalimpa’a. Data of the holotype are pre-

sented in a separate column.

females

paratypes (N=10)

paratypes (N=7)

max. mode min max. mode

22 20 20 22 22

13 11 1] 12 12

6 6 6 6 6

13 12 12 {2 12

1.4/5.1 1.4/5.1 14/51 14/54 1.4/5.1

6/8 6/7 6/6 6/7 6/7

67 66 61 66 64

max. mean (SD) min. max. mean (SD)

50.2 47.5 (42.2) Al.] 52.8 44 4 (44.1)

1229", “ROSE Bye D9 D9 TOTP Ee 10)

33.1 31.4 (41.0) 30.6 33.2 32:0-(41.0)

3.4 QO (ESD) 18.7 3279 26.5 (+6.6)

PAN 20.0 (+1.1) 18.5 22.4 20.0 (+1.6)

78.0 75.4 (41.1) La 9 74.9 (41.0)

51.8 50.5 (+0.8) 49,2 5229 51.19 (41.4)

64.6 62.7 (£1.0) 61.8 657 63.29 (41.4)

Do 21.0 (41.0) 16.0 20.3 18.49 (+1.5)

13.4 12.7 (40.3) (Bea) 14.1 13.49 (+0.6)

133 11.9 (+1.0) 10.7 14.4 12.09 (+1.3)

11.5 1124402) 10.0 124 11.09 (40.8)

2971 24.5 (42.4) 14.4 16.1 15.29 (40.8)

9 12.7 (40.7) 10.7 13.1 11.89 (40.8)

24.0 22.7 (+1.4) 19.4 22:5 21.99 (+1.1)

pale: 19-2 -GEIyS) 17.3 19.8 18.69 (40.9)

12.6 11.4 (40.7) 13.7 18.9 15.89 (+1.8)

36.5 33.0 (42.6) 30.6 34.2 32.49 (41-5)

28.4 26.7 (+1.1) 26.3 29.6 28.09 (+1.0)

39.2 35-9: (£2)3) 32:5 39.6 35.29*(42'3)

males

holotype

MZB26462 min.

anal-fin rays (total) 21 20

dorsal-fin rays (total) 12 1]

pelvic-fin rays 5 6

pectoral-fin rays 13 1]

principal caudal-fin rays 1.4/5.1 1.4/5.1

procurrent caudal-fin rays 6/7 6/6

scales in lateral row 66 62

min

standard length (mm) 41.9 44]

% standard length

total length 1223 193

head length 33:7 SOT

head depth ga 16:7

head width 195 18.0

predorsal length 75.4 74.4

prepelvic length 49.7 49.7

preanal length 62.6 (on By

body depth 21.0 19.3

body width HAC? 124

length of caudal peduncle 10.6 10.3

depth of caudal peduncle 10.1 10.9

length of dorsal fin 34.4 212

length of dorsal fin base 12.8 Mes

length of anal fin base 23:9 oD

length of pectoral fin 2237 | Pi

length of pelvic fin 10.6 10.3

% head length

interorbital width 29.4 27.8

eye diameter 26.9 25.0

34.7 31.5

snouth length

pao; Oryzias hadiatyae: MZB 18491, holotype, MZB

18503, 1 paratype, MZB 18504, 1 paratype, MZB 18505,

1 paratype, MZB 18506, 1 paratype, Indonesia, Sulawesi

Selatan, Lake Masapi; Oryzias marmoratus: MZB 2686,

7, MZB 2690, 1, Indonesia, Sulawesi Selatan, Lake

Wawontoa; MZB 2695, 5, MZB 2697, 5, Indonesia, Su-

lawesi Selatan, Lake Towuti; Oryzias matanensis: MZB

21379, 1, Indonesia, Sulawesi Selatan, Lake Matano;

Oryzias nebulosus: MZB 21381, 2, Indonesia, Sulawesi

Bonn zoological Bulletin 71 (1): 77-85

Tengah, Lake Poso; Oryzias nigrimas: MZB 5859, holo-

type, MZB 5872, 5 paratypes, Indonesia, Sulawesi Ten-

gah, Lake Poso; Oryzias orthognathus: MZB 5870, ho-

lotype, Indonesia, Sulawesi Tengah, Lake Poso; Oryzias

profundicola. MZB 5868, holotype, MZB 5861, 1 para-

type, MZB 5866, | paratype, MZB 5867, 2 paratypes, In-

donesia, Sulawesi Selatan, Lake Towuti; Oryzias wolasi:

MZB 21465, 2, Indonesia, Sulawesi Tenggara, District

©ZFMK

A new endemic species of pelvic-brooding ricefish from Lake Kalimpa’a, Sulawesi, Indonesia 85

Wolasi; Oryzias woworae: MZB 21380, 1, Indonesia,

Sulawesi Tenggara, Muna Island, District Parigi.

Acknowledgements. We would like to thank to Sutrisno K.

Dwaja, the President of Muhammadiyah Luwuk University,

Erwin, W., the Dean of Faculty of Fisheries, Muhammadiyah

Luwuk University, Risno Mina, the Head of LP3M Muham-

madiyah Luwuk University) and Hamzah, M.Si, the Head of

Fish Quarantine Station, Quality Control and Safety of Fishery

Products, Palu, Central Sulawesi with staff Irmawan Syafitri-

anto, Nur Halik Lande for supporting this research. Thanks to

Ir. Jusman, the Head of Lore Lindu National Park Hall for the

permit to conduct research in the Lake Kalimpa’a and park staff

Wantoko and Doni Kristianto for supporting us in conducting

fieldwork. We thank Fajri Ramadhan (MIPA Faculty, Tadulako

University), Suyanto, Zarlif, Novarman (Mapala Sagartmatha,

Agriculture Faculty, Tadulako University), Rocky Zakaria,

Ramlin, Fadly, Ekspedisi Riset Akuatika (ERA) Indonesia team

for the field help. Thanks also to Tan Heok Hui and one anon-

ymous reviewer for improving the manuscript. This study was

partially supported by Grant-in-Aid for Early-Career Scientists

number 19K 16203 to Javier Montenegro.

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APPENDIX I

(electronic supplement, available at www.zoologicalbulletin.de)

Table S1. All measurements and counts of examined

O. kalimpaaensis; separated in 33 in 29. * holotype.

©ZFMK

Table S1. All measurements and counts of examined O. kalimpaaensis; separated in 33 in 29. * holotype.

anal-fin rays (total)

dorsal-fin rays (total)

pelvic-fin rays

pectoral-fin rays

principal caudal-fin rays

procurrent caudal-fin rays

scales in lateral row

standard length (mm)

% standart length

total length

head length

head depth

head width

predorsal length

prepelvic length

preanal length

body depth

body width

length of caudal peduncle

depth of caudal peduncle

length of dorsal fin

length of dorsal fin base

length of anal fin base

length of pectoral fin

length of pelvic fin

% head length

interorbital width

eye diameter

snouth length

119.3

30.4

17.1

18.0

74.7

50.5

61.9

19.3

12.6

10.9

11.0

Oot

12.7

23.8

17.5

10.9

27.8

26.4

35:2

ind 12

ind 23

122.3

33.7

ie a

19.5

75.4

49.7

62.6

21.0

10.6

10.1

34.4

12.8

23.9

221

10.6

29.4

26.9

34.7

120.4

Shy

18.7

19.5

aor ba

61.9

19.0

12.9

10.8

10.5

15.8

Fal

22.4

LZ:

16.0

30.6

29.6

32:5.

females

ind 15

120.8

30.6

18.5

74.9

49.9

63.8

17.1

14.0

14.4

10.9

14.4

11.5

19.4

19.8

13.7

31.4

28.8

34.7

ind 17

122.0

32.9

19.3

O49

61.8

18.2

13.3

12.1

11.1

14.5

10.7

22:5

18.9

17.4

31.5

27.4

34.1

min. max. mode

20.0 22.0 21.0

11.0 13.0 12.0

5.0 6.0 6.0

11.0 13.0 12.0

14/51 1.4/5.1 1.4/5.1

6/6 6/8 6/7

61.0 67.0 64.0

min. max. _mean(SD)

41.1 52.8 45.8 (43.5)

119.0 122.9 120.6 (+1.2)

30.1 33.7 31.7 (41.1)

16.7 33.7 23.7 (£7.0)

18.0 224 20.0 (+1.3)

73.3 78.0 75.2 (+1.0)

49.2 53.0 50.7 (£1.1)

61.7 65.7 62.9 (+1.2)

16.0 22.2 20.0 (£1.7)

11.7 14.1 12.9 (40.6)

10.3 14.4 11.9 (41.1)

10.0 12.1 11.1 (40.6)

144 344 21.5 (+5.9)

10.7 13.9 12.4 (40.8)

19.2 24.0 22.4 (+1.4)

17.3 22.7 19.2 (£1.5)

10.3 19.0 13.1 (42.6)

27.8 36.5 32.7 (+2.3)

25.0 29.6 27 2: (+1.2)

31.5 39.7 35.3 (+2.2)