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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 We, therefore, provide here an English translation of this paper, together with so far unpublished background information on this rare and endangered amphibian, and on the former scientific German-Afghan cooperation project from the 1960s in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).
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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
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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
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An early study on the Afghanistan Mountain Salamander Paradactylodon (Afghanodon) mustersi 5
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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
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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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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.
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annals and magazin cf natural History.ser Il.vol 5 382 . 384)
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Bonn zoological Bulletin 71 (1): 1-7
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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-
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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
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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,
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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
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¥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).
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The new lanternfly Limois sonlaensis sp. nov. from Vietnam (Hemiptera: Fulgoridae) 13
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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.
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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.
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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.
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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
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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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Oshanin VT (1908) Beitrage zur Kenntnis der palearktischen Hemipteren. I. Neue Gattungen und Arten von Homopteren aus Russisch-Asien. Annuaire du Musée zoologique de l’Académie impériale des Sciences de St.-Pétersbourg 12: 463-477
Stal C (1863) Beitrag ziir Kenntnis der Fulgoriden. Entomol- ogische Zeitung. Herausgegeben von dem entomologischen Vereine zu Stettin 24: 230-251
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Wang W-Q, Xu S-L, Constant J, Qin D-Z (2020) Revision of the lanternfly genus Limois Stal (Hemiptera: Fulgoromor- pha: Fulgoridae) with description of a new species from Chi- na. European Journal of Taxonomy 720: 35-61. https://doi. org/10.5852/ejt.2020.720.1113
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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.
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A new species of Cryptopacha Prozorov & Zolotuhin, 2012 from West Africa 21
Oe
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
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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
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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 © e 5 e e hy e Aa e . ef = OG e e e t ) e 3 . q e e e \e e ) e e e e? ee e e o @ ¢@ e e Cd 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).
4
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 n c S
100 +5 ® rot D o =
50
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 Chiclana de la Frontera Chiclana de la Frontera Conil de la Frontera Jerez de la Frontera Jerez de la Frontera Jerez de la Frontera Jerez de la Frontera Jerez de la Frontera Jerez de la Frontera Jerez de la Frontera Jerez de la Frontera Jerez de la Frontera Jerez de la Frontera Linea la Concepcién Los Barrios Medina Sidonia Medina Sidonia Medina Sidonia Medina Sidonia 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 21
2016-2017 2018-2019 22.05.16 2018-2019 09.04.17 09.04.17 2016-2017 01.05.13 2013-2019 2015-2017 2015-2017 2016-2017 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 CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ
Trebujena Vejer de Frontera Alcala de los Gazules Alcala de los Gazules Algeciras Barbate Benalup-Casas Viejas Castellar de la Frontera Castellar de la Frontera Chiclana de la Frontera Chiclana de la Frontera Chiclana de la Frontera Chiclana de la Frontera Chipiona Espera Jerez de la Frontera Jerez de la Frontera Jerez de la Frontera Jerez de la Frontera Jerez de la Frontera Jimena de la Frontera Jimena de la Frontera Los Barrios Los Barrios Los Barrios Los Barrios Los Barrios Manzanete Medina Sidonia Puerto Santa Maria Puerto de 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 NA 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 NA 36.43572, -5.90323 36.62448, -6.19817 NA NA
2016-2017
01.06.17 14.06.18 19.04.19 03.04.17 29.04.17 NA NA NA 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 29.05.17 29.05.17 29.05.17 26.05.15 16.05.15 NA 12.06.13 30.03.17 NA NA 03.04.20 20.04.13 NA NA
CS CS CS CS CS CS CS CS CS CS CS CS CS CS CS CS CS CS CS CS CS CS CS CS CS CS CS CS CS CS CS CS
CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ
Puerto de Santa Maria
Puerto Real Puerto Real Puerto Real Puerto Real Puerto Real Puerto Real Puerto Real Puerto Real Puerto Real San José del Valle San José del Valle San José del Valle San Roque San Roque San Roque San Roque San Roque Tarifa Tarifa Tarifa Tarifa Tarifa Tarifa Tarifa Tarifa Tarifa Tarifa Torre Alhaquime Ubrique 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 NA NA 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 NA 36.20974, -5.38457 NA NA 36.20403, -5.69441 36.15347, -5.63273 36.06480, -5.64840 NA 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
NA 36.27180, -5.93740
115
07.03.21 29.05.17 NA NA 16.04.21 22.03.10 NA 29.05.17 30.03.18 29.05.17 23.04.16 NA 14.05.17 25.04.15 NA 07.04.09 27.03.85 01.04.82 NA 16.05.15 08.05.21 NA 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
CS CS CS L: Bologna, 1989 L: Bologna, 1989 L: Bologna, 1989 L: Garcia-Paris, 1998 L: Garcia-Paris, 1998 L: Garcia-Paris, 1998 : Garcia-Paris et al., 2003 : Garcia-Paris et al., 2003 : Garcia-Paris et al., 2003 Garcia-Paris et al., 2003 Garcia-Paris et al., 2003 : 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., 2003 L: Pérez-Moreno et al., 2004 L: Pérez-Moreno et al., 2003 L: Percino-Daniel et al., 2013 L: Percino-Daniel et al., 2013 L: Percino-Daniel et al., 2013 L: Percino-Daniel et al., 2013 L: Cortés-Fossati, 2018 L: Cortés-Fossati, 2018 L: Cortés-Fossati, 2018 L: Cortés-Fossati, 2018 L: Sanchez-Vialas et al., 2020 L: Sanchez-Vialas et al., 2020
et eee
CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ
Vejer de la Frontera Vejer de la Frontera Villaluenga del Rosario NA NA NA NA NA NA Algeciras Algeciras Algeciras Alcala de los Gazules Jerez de la Frontera Jerez de la Frontera San Roque Tarifa Ronda San Roque San Roque Tarifa Benalup-Casas Viejas Chiclana de la Frontera Vejer or Algodonales? NA Medina Sidonia Puerto Real 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 NA NA NA NA NA Laguna de Medina NA NA Sierra de las Nieves NA NA NA Benalup de Sidonia NA 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
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA
NA 36.45757, -5.94340 36.53127, -6.16051 36.53636, -6.19546 36.75019, -6.34113 36.29691, -6.11925 36.12633, -5.70572
30
870
13.05.18 23.05.12 05.05.08 NA NA NA NA NA NA 21.04.21 23.04.21 24.04.21 NA 16.03.85 27.03.83 00/05/1973 00/04/1971 10.04.87 00/05/1987 00/05/1971 20.03.89 NA NA NA NA 2016-2018 2016-2018 2014-2021 04.06.17 13.04.13 19.06.16
: Sanchez-Vialas et al., 2020 : Sanchez-Vialas et al., 2020 : Sanchez-Vialas et al., 2020 : Sanchez-Vialas et al., 2020 : Sanchez-Vialas et al., 2020 : Sanchez-Vialas et al., 2020 : Sanchez-Vialas et al., 2020 : Sanchez-Vialas et al., 2020
CS
CS
CS
CS
CS
CS
CS
CS
CS
CS
CS
CS
CS
CS
CS
CS
CS
CS
CS
CS
CS L: Sanchez-Vialas et al., 2020 L: Sanchez-Vialas et al., 2020 L: Garcia-Paris, 1998
Cotikeiien ke stem acne
CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ MALAGA MALAGA MALAGA MALAGA MALAGA MALAGA MALAGA MALAGA MALAGA MALAGA MALAGA MALAGA MALAGA MALAGA MALAGA MALAGA MALAGA MALAGA MALAGA MALAGA MALAGA MALAGA MALAGA MALAGA
Manzanete Manzanete Paterna de Rivera NA NA NA NA San Roque Alhaurin el Grande Antequera Antequera Campillos Campillos Ciudad Jardin Ciudad Jardin Cortes de la Frontera Fuente de Piedra La Roda de Andalucia Malaga Malaga Ojén Parauta Palma-Palmilla Puerto de la Torre Sierra de Yeguas Tolox Yunquera Yunquera Yunquera NA Montecorto NA
Sierra del Retin Sierra del Retin NA 2 km S Benalup Sidonia 7 km SW Benalup Sidonia 4km NE San José del Valle 3km S Alcala de los Gazules NA 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 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 NA Near Malaga, see Fig. 1
NA
36.20000,
-5.80000
NA NA NA NA NA NA
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
NA NA NA
04.01.18 24.03.11 21.05.17 19.04.01 19.04.01 19.04.01 NA 00/05/1973 29.04.07 NA 14.04.15 25.05.19 25.05.19 05.04.12 03.04.07 04.04.15 NA NA 04.03.15 15.07.19 06.04.08 13.05.11 22.07.17 19.03.21 NA 14.03.16 25.05.13 03.05.14 26.03.16 25.04.91 NA NA
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 Chiclana de la Frontera Chiclana de la Frontera
Conil de la Frontera Jerez de la Frontera
Jerez de la Frontera Jerez de la Frontera Jerez de la Frontera Jerez de la Frontera Jerez de la Frontera
Jerez de la Frontera Jerez de la Frontera Jerez de la Frontera Jerez de la Frontera Linea la Concepcion
Los Barrios Medina Sidonia 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
F
F L: Cortés-Fossati, 2018 L: Cortés-Fossati, 2018 L: Cortés-Fossati, 2018 L: Cortés-Fossati, 2018
CADIZ CADIZ CADIZ
CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ CADIZ
Medina Sidonia Medina Sidonia Medina Sidonia
Paterna de Rivera Puerto Real Trebujena Medina Sidonia Puerto Real 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 i
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
B
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
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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-
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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.
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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.
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38 Joachim Schénfeld & Dirk Rohwedder
q
? Vy 3 4A + (7 of) i]
= 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.
REFERENCES
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Balthasar V (1959) Neue Onthophagus-Arten (97. Beitrag zur Kenntnis der Scarabaeidae, Col.). Entomologische Blatter 55: 186-195
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tini, Onthophagini, Phanaeini, Scarabaeini and Sisyphini. Acta Entomologica Musei Nationalis Pragae 53: 387-442
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Harold E (1886) Coprophage Lamellicornien. Berliner Ento- mologische Zeitschrift 30 (2): 141-149
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Kabakov ON (2006) Beetles of the subfamily Scarabaeinae (In- secta: Coleoptera: Scarabaeidae) of the fauna of Russia and adjacent countries. Scientific Publications KMK, Moscow. [in Russian]
Lobl I, Krell, FT, Ziani S & Kral D (2006) Scarabaeidae, sub- family Scarabaeinae, tribe Onthophagini. Pp. 159-176 in: Lobl, I. & Smetana, A. (eds) Catalogue of Palaearctic Cole- optera, Volume 3: Scarabaeoidea — Scirtoidea — Dascilloidea — Buprestoidea — Byrrhoidea. Apollo Books, Stenstrup
Matsumura S (1938) Onthophagid-insects from Formosa, In- secta Matsumurana, Sapporo 12 (2-3): 53-63
Mittal IC (2000) Survey of scarabaeid (Coleoptera) fauna of Himachal Pradesh (India). Journal of Entomological Re- search 24: 259-269
Mittal IC & Jain R (2015) A checklist of Indian dung beetles (Coleoptera: Scarabaeidae). Indian Journal of Entomology 77. 383-404
Scheuern J (1995) Taxonomie, Sexualdimorphismus und Neu- beschreibungen orientalischer Onthophagus-Arten unter be- sonderer Berticksichtigung des Subgenus Colobonthophagus Balthasar, 1935 (Coleoptera, Scarabaeaidae). Entomologica Basiliensia 18: 413-453
Scheuern J (1996) Neue und verkannte Onthophagus-Arten aus der Orientalischen Region. (Coleoptera, Scarabaeaidae). Stuttgarter Beitrage zur Naturkunde A 542: 1-20
Weidner H (1976) Die Entomologischen Sammlungen des Zoologischen Instituts und des Zoologischen Museums der Universitat Hamburg IX. Teil 1) Insecta VI. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut 73: 87-264
Wiedemann, ERW (1823) Zweihundert neue Kafer von Java, 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
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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).
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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
44
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
56
—_ Oo
—
12
19
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
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
18
30
Rea WY WR WN
65
14
(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- 32
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
45
Localities
SC: Ilha de Santa Catarina; SP: Ilha de Sao Sebastiao
RS: Sao Leopoldo SP: Ilha de Sao Sebastiao SP: Ilha de Sao Sebastiao PA: Ilha de Marajo
RS: Taquara SC: Florianopolis 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).
3 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 | _ o -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 wv
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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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 designed by Okland (1996). A trap was made of two 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
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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 partial protection partial protection partial protection partial protection 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,
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oO = Meripilus giganteus, A = Hericium coralloides, A= Hericium alpestre, = Fistulina hepatica.
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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
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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 a oO oO
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
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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
80
60
Number of individuals
40
20
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
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Tree-fungus beetles collected on mycelium of selected fungal species in central Poland 63
70
65
60
55
Distance Ss
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-
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64 Jacek Pietka et al.
0.10 0.15
0.05
brown rot
NMDS2 -0.05 0.00
-0.10
-0.15
-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 9%
Trixagus carinifrons
side 7% remaining species
50%
Cryptophagus dentatus 7%
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.
Bonn zoological Bulletin 71 (1): 51-67 ©ZFMK
Tree-fungus beetles collected on mycelium of selected fungal species in central Poland 65
remaining species 46%
\
4 4 )
Trixagus carinifrons 18%
Dienerella ruficollis
pae Rhizophagus bipustulatus 6%
Aulonothroscus brevicollis 6%
|
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