SUGGESTED CITATION FOR THIS VOLUME:

Telnov D., Barclay M. V. L. & Pauwels QO. S. G. (eds) 2021. Biodiversity, biogeography and nature

conservation in Wallacea and New Guinea. Volume IV. The Entomological Society of Latvia, Riga, 443 pp.

SUGGESTED CITATION FOR SEPARATE PAPERS:

Greke K. 2021. New species and records of the Papuan Diplommatinidae (Caenogastropoda:

Cyclophoroidea): 111-134. In: Telnov D., Barclay M. V. L. & Pauwels O. S. G. (eds) Biodiversity, biogeography

and nature conservation in Wallacea and New Guinea. Volume IV. The Entomological Society of Latvia,

Riga, 443 pp.

ISBN of the volume IV: 978-9984-9768-9-1

ISSN of the series: 2255-9728

http://zoobank.org/24CE43G6E-338A-40CC-9F4E-C30F10379CD0

PUBLISHER:

The Entomological Society of Latvia, Riga http://leb.daba.|v 7-

es NABU

NABU, Naturschutzbund Deutschland, Germany https://en.nabu.de

The Natural History Museum, London, United Kingdom nhm.ac.uk

EDITORIAL BOARD:

DMITRY TELNOV (executive editor, the Natural History Museum, London & the Entomological Society

of Latvia, Riga), MAXWELL V. L. BARCLAY (the Natural History Museum, London) and OLIVIER S. G.

PAUWELS (Royal Belgian Institute of Natural Sciences, Brussels)

ORDERS & INQUIRIES:

The Entomological Society of Latvia, c/o Institute of Biology,

O. Vaciesa iela 4, LV-1004, Riga, Latvia NATU RAL

Executive editor: anthicus@gmail.com H ISTO RY

MUSEUM

ETHICS, TERMS & CONDITIONS: http://leb.daba.lv/book/terms.html

Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea is published by the

No parts of this book may be reproduced in any form by digital or mechanical means (including

photocopying, scanning, recording, or information storage and retrieval) prior to obtaining permission in

writing from the publisher.

OPEN ACCESS ARCHIVE OF PREVIOUS VOLUMES: https://www. biodiversitylibrary.org

LAYOUT: Dmitry Telnov

PRINTED BY: Jelgavas tipografija, Jelgava, Latvia June 2021)

DISCLAIMER:

The views and opinions expressed by the authors in this volume do not necessarily represent or reflect

those of the editors or copyright owners. While each article or chapter is believed to contain accurate

information, neither the editors nor copyright owners offer any warranty, expressed or implied, or assume

any legal liability or responsibility for the accuracy, completeness, or usefulness of any information,

product, or process disclosed, or represent that its use would not infringe privately owned rights.

Title page photo: Papuan house surrounded by primary rainforest, Star Mountains, New Guinea (image

by Dmitry Telnov).

Book4.indd 2 02-Jun-21 21:46:29

Drawn by Dmitry Paramonov (Riga, Latvia, March 2021)

Dedicated to Otto Merkl (1957-2021)

This volume is respectfully dedicated to our friend and colleague Ott6 MERKL (1957-2021) who unexpectedly

died on the 19" February during the late stages of its production. Otto was the Head of Coleoptera and Head

of Zoological Collections at the Hungarian Museum of Natural History and the Budapest Zoo, and was a world-

renowned specialist in the Coleoptera family Tenebrionidae, especially the subfamily Lagriinae, as well as a museum

curator of the highest standard and of international repute.

Appointed soon after graduation in 1981, Otto served 40 years as an employee of the Hungarian Museum, where

he followed in the footsteps of his mentor and predecessor, the tenebrionid specialist Zoltan Kaszab (1915-1986).

During his career Otto did much to raise the reputation and scope of the Hungarian Museum’s globally significant

collections. He formed a bridge with the world scientific community, and was known and admired by taxonomic and

faunistic entomologists, professional and amateur, as an example of a scientist with a deep specialisation, but at

the same time, a broad overview of entomology and the management of Museums and Collections. He published

over 220 scientific papers, several books and book chapters, has almost 100 species named in his honour, and was

deeply involved in the Hungarian Entomological Society all his life. In 2019 the Society awarded him their highest

honour, the “Frivaldszky Gold Memorial Plaque”, of which, in his 62, he was the youngest ever recipient.

It is to be regretted that during Ottd’s last years, the status and future of the Museum to which he dedicated his

life was plagued by uncertainties outside his control, which must have increased his stress; he died on his way into

work, not long after taking on the role of Head of the whole Zoological Collections.

Otto will be fondly remembered all over the world, for many years to come, as a model of a scientist-curator

and as the man who helped build the remarkable collections of the Hungarian Museum, and put them firmly on the

world map.

Words and opinions by M. Barclay and D. Telnov, with data taken from “In memoriam Ott6 Merk! (1957-2021)” by G. Szél.

Book4.indd 3 02-Jun-21 21:46:35

Book4.indd 4 02-Jun-21 21:46:35

Contents

FECL GOI A oe sae eee cce er alesia te haan ea te te ae we enc hes nO ad eras Mi tech, Se ete nae i aicins caer tests Ore asec eee tere eS a ve Ue ee et a ee 6

Foreword, by O'SHEA, Mark: Wallacea —- A hotspot Of Snake GIVEPSItY ..........ccccessssessssseseeeeeeeeeeeeeeeeeesssesssssscsaneeseeeseeeeeeeeeess £

AGKMOWICU BE IMGIESs aseceusasb teense cuca seen, Mess. od teel ted sean hat chalice hate ea ted cans As ant urs anal oon aah 0 Ooi lf aR ee 18

SECTION ONE: VERTEBRATE ZOOLOGY AND ECOLOGY

RICHARDS, Stephen J., JATURADI, Burhan, KREY, Keliopas & DONNELLAN, Stephen C.: A new stream-dwelling frog

of the genus Litoria Tschudi, 1838 (Anura: Pelodryadidae) from Salawati Island, INGONESIA ............:ccccceeeeeeeeeeeeeees 19

ROSLER, Herbert & GAULKE, Maren: On the biology of Ernstkeller’s cave gecko, Gekko ernstkelleri Rosler, Siler,

Brown, Demegillo et Gaulke, 2006 (Squamata: Gekkonidae), with emphasis on reproductive biology ...........:cee 35

SHEA, Glenn M. & ALLISON, Allen: A new species of Sohenomorphus (Squamata: Scincidae) from Mount Kaindi,

Viorobe Province. -Raptiat NGWIGUINMGay. ..crec:ccectessosttestecacsctatecstbatea srsuvecsaltteneentrncundeearesnttecessucscsstteavadihssifadtrsatetteesnadaretecets 49

SECTION TWO: INVERTEBRATE ZOOLOGY AND ECOLOGY

BALLERIO, Alberto: First contribution to the knowledge of the Ceratocanthinae (Coleoptera: Scarabaeoidea:

Hybosoridae) from the Solomon Islands, with descriptions Of five NEW SPECIES. ..........::sscccccceceeecseeeeesssssssssssssseeeeeeeees 61

BORDONI, Arnaldo: New data on the Oriental Xantholinini, 49. A new genus, species and records from the

Philippines:and: Indonesia, (Goleopterary Stapnylinid ae) tec. Ti% danneseee te Recncnestus iovtensda see? Vestas deaneuvtetene Daas Qvtvtandsll a RA 5

DRUMONT, Alain, KOMIYA, Ziro & WEIGEL, Andreas: Catypnes marazziorum sp. nov. (Coleoptera: Cerambycidae:

Preninae) TremnurapliarNew GuUInea-.2. ese a. den acct nce: wt cece edn toe cee alien et xm apis acioatedla ten deceennnnnrseeteh arent orbnsune eadaadoceey 83

GOLOVATCH, Sergei I|., AKKARI, Nesrine, GOUD, Jeroen & TELNOV, Dmitry: Review of the Papuan millipede

genus Acanthiulus Gervais, 1844 (Diplopoda: Spirobolida: PaChybolidae) ...........ccccccccccceceeccsssssssssssseseceeeeeeeeeeeeeeeeseess 91

GOLOVATCH, Sergei I.: A new species of the genus Acanthiulus Gervais, 1844 (Diplopoda: Spirobolida: Pachybolidae)

EOD PRA es I INT chaos 2 ee ace 2 A ts ane sc eas stan aatace ads na doh age Sha et ohaaueta oat tee Ae ee Sa tsuay abe ree eens no hedaanct 107

GREKE, Kristine: New species and records of the Papuan Diplommatinidae (Caenogastropoda: Cyclophoroidea) .. 111

GREKE, Kristine & SLAPCINSKY, John: New Taheitia H. et A. Adams, 1863, with revisional notes on the Papuan

Truncatellidae (Caenogastropoda: TruncatellOid@a) .............:::ccccccccccsecesesesesesssssssscceececeeeeeeeeeseseeeeesenssssneceeceeeeeeeeeeeeeess 135

GREKE, Kristine, TELNOV, Dmitry, KAGAINIS, Ugis & TELNOV, Edwin: The possible functional role of periostracal

processes in Japonia saetigera (van Benthem Jutting, 1958) (Caenogastropoda: Cyclophoridae) based on SEM

micrographanalysis-andinumenical CaleulatiGnS +A... 8.2. sede ee lean e- secer tenes oe candice venue cade sokee-cwneeeekenn oa enstaaemeett pe wecetaeatne 185

HAVA, Jiti: Orphinus (Orphinus) terminalis (Sharp in Blackburn & Sharp, 1885) (Coleoptera: Dermestidae) from

THe SOLOMON AS AMSA WaT IAL Rea es cei eee ae ee ee re ec are Pacer tenet ees teats ante saan yeaa eee teerantetee nectar eteaes 199

HAVA, Jit: A new synonym in the genus Orphinus Motschulsky, 1858 (Coleoptera: Dermestidae) from Papua

NEW LIN a eeshatees, esheets ce Mee cr gm 502 MCR Nt ds obtain aed ehac ith aed musa Met emeL ews, Mute ah bad eecePns, Macc ini nant 201

INGRISCH, Sigfrid: New species and records of Tettigoniidae Krauss, 1902 from the Papuan Region (Orthoptera).. 203

KALASHIAN, Mark Y.: A new species of Endelus Deyrolle, 1864 (Coleoptera: Buprestidae) from Sulawesi,

Indonesia, with description of Endelus (HExAZONOCEIUS) SUDZEN. NOV. .....csesseseeeeeeeeeeseseeeeeeeeeeeeeeeeeeeseseeseneeeeseessseanees 215

KALNINS, Martins: Contribution to the knowledge of dragonflies (Odonata) from the Raja Ampat (Indonesia),

WATHPAOLES: ON AIEIMEC OOS Vie me esha teil USey, tel Syd LA Be SR oh Os Ps ee ee core PT ae Pails

KALNINS, Martins: Contribution to the knowledge of aquatic bugs (Hemiptera: Heteroptera) from Wallacea and

New Guineas With CCOIOBIGAl NOLES 625.556 a) Cad wedecl ewes ub acs tocea th ducer taal aceecdeta acs cubuee Sele arsautes icine nndeoscncwcse nee 239

KAZANTSEV, Sergey V.: New Plateros Bourgeois, 1879 from New Guinea and the Moluccas, with a checklist of

SHECIES Olathe TEBION (COlEG Plea AV CIA ACY we toh trate seal eds octal ntl ed tee inane kneech net enkueecedirenglaaaus stllslbeuecdrendteiadeedeene 245

KAZANTSEV, Sergey V.: New metriorrhynchine species (Coleoptera: Lycidae) from SUIAWES I ........cceeessseeeeeeeseneeeeeeees 25S

LOBL, Ivan: Contribution to the knowledge of the Scaphisomatini (Coleoptera: Staphylinidae: Scaphidiinae) of

MindanaosRniliPDINeS. «wn ese eee ee Seats ete ne ee, en erin ne See tA ba Sel ee ft Os Di na es 265

OWEN, Ifor L. & BORAY, Joseph C.: Distribution of Orientogalba viridis (Quoy & Gaimard, 1832) (Gastropoda:

Lymnaeidae), the snail intermediate host of Fasciola hepatica Linnaeus, 1758, in Papua New Guinea ............... 273

OWEN, Ifor L.: A brief review of animal parasites recorded in Western Province, Papua New Guinea ............:::0000000 281

TELNOV, Dmitry & POLHEMUS, Dan A.: A new Tanycricos La Rivers, 1971 (Hemiptera: Heteroptera: Naucoridae)

from the Cyclops Mountains, New Guinea, with a key tO SP@CIES ..............ccccesseseeeessssececceceeeeeeeeeesnsaaeasensesccccceesesseeess 329

WEIGEL, Andreas & SKALE, André: On the taxonomy, synonymy and faunistics of the Apomecynini of the Asian-

Australian region (Coleoptera: Cerambycidae: Lamiinae). Part 8 ..............cccsssssesesesssececcceeseceeeeceeenssseesesessscececeeseseseeees 345

YOKOI, Yaheita: Taxonomic notes on the tribe Callidiopini Lacordaire, 1869 (Coleoptera: Cerambycidae) and some

other taxa of the Papuan Region, mainly from the Solomon Islands, with descriptions of five new species .......... 363

ZIDEK, Jiri: Revision of Philacelota Heller, 1900 and Engertia Dalla Torre, 1912, with description of a new genus

(Coleoptera: Scarabaeidae: MelOIONthiNnae) .............:::ccccccccccccececesseseseessssccccceeccesseeeeesensneseesesesscececeeeseeeeeeeeeessaeeseessessnees 397

ZILLI, Alberto, de VOS, Rob & EDWARDS, Edward D.: The torsilinea-species group (subg. Mocrendes Nye, 1975)

of the genus Platyja Hubner, [1823] (Lepidoptera: Erebidae), with descriptions of four NeW SPECIES ..........:::eee 413

SECTION THREE: NATURE CONSERVATION

BIENKOWSKI, Lara S., STEPHAN, Robin S., KIRSCHEY, Tom & SARYANTHI, Ria: Population density of Tarsius

supriatnai along a forest degradation gradient in Popayato-Paguat landscape (Gorontalo Province, Sulawesi) .... 434

Index to NEW taxa CGESCHIDEM IN CNIS VOLUME 2... ccceeeccecceeecceeeceeceeeeceeeeeeneeaeecuueceeeeeuueeeueeaeaeesauaeeeesaueeeaeesaueaeneeaeeesueeaneeseeeeees 443

Book4.indd 5 02-Jun-21 21:46:36

Editorial

The period during which this new volume was edited has been somber. Humanity as a whole

has dramatically suffered, mainly as a result of our galloping demography, exacerbate mobility and

urbanization, and of our uncontrolled and increasing contacts with wildlife. This gloomy episode, which

possibly originated in an East Asian food market selling exotic bush meat, has generated millions of losses

of human lives and economic and social disasters. It has stressed how much medicine, human welfare

and biological sciences are strongly interdependent, and how deeply we are connected to the other animal

species sharing the Earth. As a light in the darkness, this period has also revealed extraordinary examples

of selfless solidarity and of rapid collaborative progresses in Science.

Scientific knowledge is indeed key to better manage and protect both the wildlife and ourselves. The

present, fourth volume of the series “Biodiversity, Biogeography and Nature Conservation in Wallacea

and New Guinea” is an additional brick to the edification of this knowledge.

With four chapters on mollusks, combined to the malacological contributions in the previous volumes,

the series already counts among the major contributions to the knowledge of the terrestrial snails of

the Wallacean Region. Most chapters of the present new volume unsurprisingly deal with arthropods,

which represent the vast majority and the most poorly known animals. Newly discovered invertebrates

described here include, among other little natural treasures, a predatory aquatic bug from the Cyclops

Mountains, a non-spiky species in an otherwise spiky millipede genus from Wau, and a new snail from

Waigeo Island, a giant of its kind. Several chapters are dedicated to vertebrates, including one on the

conservation of a recently described tarsier, and the diagnoses of a new frog and of a new skink species.

In total three dozens of new taxa are here brought to light and enrich the already highly diverse, still vastly

understudied, Wallacean biodiversity.

Par vents et marées, in spite of the challenges of this period, my friend zoologist Dmitry Telnov has

stayed on course and managed to involve the passionate authors who contributed to this volume, for the

sake of the knowledge, the recognition and the conservation of the fascinating and unique fauna and

flora of the Papuan Region and Wallacea.

OuivieR S. G. PAUWELS

Curator (Recent Vertebrates Collections)

Royal Belgian Institute of Natural Sciences

Brussels, Belgium (opauwels@naturalsciences.be)

Book4.indd 6 02-Jun-21 21:46:36

O'SHEA, M.: (FOREWORD) Wallacea - A hotspot of snake diversity

Foreword

Wallacea - A Hotspot of Snake Diversity

Mark O’SHEA

Mark O’Shea MBE is Professor of Herpetology at the University of Wolverhampton, United Kingdom. Although he has

conducted herpetological fieldwork throughout the world, his specialist area remains the snakes of New Guinea and

Wallacea.

Wallacea comprises five main archipelagos, including Sulawesi in the west, the Lesser Sunda Islands

in the southwest, and the northern, central and southern Moluccas to the east. Alfred Russel Wallace

included this area in his Australo-Malayan Sub-region (Wallace 1876), along with New Guinea and the

Solomon Islands, one of four sub-regions in his Australian Region, a definition followed by other authors

(Beddard 1895; George 1962; Lincoln et al. 1982), although Udvardy (1975) placed Wallacea into his

Indomalayan Realm, and New Guinea into Oceania. Darlington (1957) separated the area between the

Wallace Line (Wallace 1859; 1860), modified by Huxley (1868), and the Lydekker Line (Lydekker 1896)

as distinct from both larger regions or realms. Huxley’s modified Wallace Line places the Philippines

within Wallacea but here | intend Wallacea to include only those archipelagos listed earlier.

The non-oceanic snake fauna* of Wallacea comprises 46 genera, of which five (11%) are endemic

(Cyclotyphlops, Sundatyphlops, Pseudorabdion, Rabdion and Brachyorrhos), and 115 species, of which

67 (58%) are endemic, with an additional 11 endemic subspecies. From the world’s 41 extant snake

families, 16 (39%) are represented across Wallacea’s 347000 km? land area. This representation, for

a relatively small land area, is impressive when compared with the family counts for seven much larger

biogeographical realms?: Nearctic - 8; Neotropical - 14; Palaearctic - 11; Afrotropical - 19; Indomalaya

- 22; Australasia (excluding Wallacea) - 9, and Oceania - 5 families.

The Indo-Australian Archipelago has long been an area of intense interest for herpetologists (Barbour

1912; de Haas 1950; de Rooij 1915; 1917), and it remains so today (Kaiser et a/. 2011 2013; Koch

2012; de Lang 2011'; 2013; de Lang & Vogel 2005; O'Shea et al. 2012; 2015; Sanchez et a/. 2012).

Those species that also occur extralimitally to Wallacea demonstrate affinities with the snake faunas

of mainland Asia, the Philippines, and the Sundaland islands, New Guinea, the Solomon Islands, the

southwest Pacific, and mainland Australia (See Table 1).

Typhlopidae: Asiatyphlopidae. The typhlopid blindsnakes are represented in Wallacea by one

subfamily, five genera and ten species. The most widely distributed species is the famous “flowerpot

snake”, Indotyphlops braminus, so named because, as the world’s only obligate parthenogenetic snake

species, it has been transported circumglobally in the root balls of soil surrounding horticultural and crop

plants and become a super-coloniser, establishing beachhead populations far from its origin in South

Asia. Even though this tiny snake is the most widely distributed terrestrial snake, most of its 23 congeners

are confined to mainland Asia, except |. schmutzi, which is endemic to Flores and the Komodo Islands in

the Lesser Sundas.

The genus Malayotyphlops is centred on the Philippines with 10 of its 12 species endemic to that

archipelago, but two species occur in Indonesia, M. koekkoeki on Bunyu Island, North Kalimantan, and

M. kraali in the Kei Islands and Seram in the Moluccas. The latter taxon is quite possibly two separate

species. Half of all blindsnakes in Wallacea belong to the genus Ramphotyphlops, with five species on

1 Sea kraits (Laticauda) and true seasnakes (Hydrophis) are omitted.

2 Antarctica is omitted.

Book4.indd 7 02-Jun-21 21:46:38

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Sulawesi and in the Moluccas, and 16 species distributed from the Malay Peninsula to the Caroline Islands

and New Caledonia.

The final two blindsnake genera are both monotypic (Hedges et al. 2014), and endemic to Wallacea.

Sundatyphlops polygrammicus is represented by five subspecies distributed from Lombok to Timor. It is

most closely related to Anilios, an Australo-Papuan genus that contains two elevated former subspecies

of polygrammicus. Sulawesi is home to Cyclotyphlops deharvengi, a tiny snake with a distinctive circular

frontal scale on top of its head.

Gerrhopilidae. This blindsnake family was elevated from within the Typhlopidae, based on molecular

data (Vidal et a/. 2010). It contains two genera, the monotypic Cathetorhinus, believed to come from

Mauritius, and Gerrhopilus, a genus of 20 species distributed from India to the Philippines and Papua

New Guinea. A single widespread Indonesian species, G. ater, is recorded in Wallacea, from Sulawesi, and

Halmahera and Ternate in the northern Moluccas.

Cylindrophiidae. Asian pipesnakes, genus Cylindrophis, include 15 species, mostly endemic to

Southeast Asia, excluding the Philippines, but with a single Sri Lankan species. Wallacea is clearly a

hotspot for Cylindrophis with eight species recorded from the Lesser Sundas, Sulawesi and the Moluccas.

These include the widely distributed Southeast Asian C. ruffus and the Wallacean endemics: C. aruensis,

C. boulengeri, C. isolepis, C. melanotus, C. opisthorhodus, and C. yamdena. The most recently described

Wallacean species was C. osheai from Boano Island to the northwest of Seram and named for this author

(Kieckbusch et a/. 2018).

Xenopeltidae. This monotypic South and Southeast Asian family contains only two species of

iridescent-scaled sunbeam snakes. One species is endemic to Hainan Island, China, but the other

species, Xenopeltis unicolor, is widely distributed, from India to China, and enters Wallacea on Sulawesi,

and neighbouring Buton Island.

Pythonidae. The pythons are very well represented in Wallacea. The most iconic and widely distributed

species, and also the longest snake in the world, is the reticulated python, Malayopython reticulatus,

which occurs from the Southeast Asian mainland to the Philippines, as far north as Itbayat Island, and

south to Sulawesi, the Lesser Sundas, and the Moluccas, with two endemic, insular Subspecies on Selayar

and Tanahjampea Islands, south of Sulawesi (Auliya et al. 2002). Its sister taxon is M. timoriensis, which

inhabits Flores and other Inner Banda Arc islands, but, despite its name, not Timor in the Outer Banda Arc.

The correct common name for this species should therefore be Lesser Sunda or Flores python, rather than

Timor python. Recent molecular research (Reynolds et a/. 2014) has demonstrated that Malayopython is

more closely related to the Australo-Papuan pythons of the genera Morelia and Simalia, than to the Afro-

Asian pythons, genus Python.

Simalia is represented in the Moluccas by four species. Simalia tracyae in the northern Moluccas,

S. clastolepis in the central Moluccas, and S. nauta in the southern Moluccas (Harvey et a/. 2000), while

a widespread Australo-Papuan species, S. amethistina, inhabits a few small islands at the eastern edge

of Wallacea. The genus Liasis is also Australo-Papuan, but one species is endemic to the Lesser Sundas.

Liasis mackloti is a water python commonly encountered on Timor and Alor, with subspecies on Wetar,

L. m. dunni, and Sawu, L. m. savuensis. Some authorities elevate these two island endemics to specific

status.

The only truly Asian python in Wallacea is the Burmese python, Python bivittatus. Primarily distributed

across northern India and Nepal to China, and south to Thailand and Vietnam, but absent from the Malay

Peninsula, Borneo and Sumatra, isolated populations are known from east Java, Nusa Barang, and Bali,

and within Wallacea from Sumbawa and south Sulawesi, this last population being allocated subspecific

status as P. b. progschai (Jacobs et al. 2009).

Candoiidae. The five species of Pacific boas are primarily distributed from New Guinea, north to Palau,

—_—_

tl i

Book4.indd 8 02-Jun-21 21:46:38

O'SHEA, M.: (FOREWORD) Wallacea - A hotspot of snake diversity

south to the Loyalty Islands, and as far east as Fiji, Tonga, Samoa and Tokelau (Smith et a/. 2001). Two

New Guinean species extend their ranges eastwards to the Talaud and Sangihe Islands north of Sulawesi.

Candoia paulsoni tasmai is an endemic Moluccan subspecies of the Solomon Islands ground boa, C.

paulsoni, otherwise known from Papua New Guinea and the Solomon Islands, but not from intervening

western New Guinea. The other taxon present is the widespread and gracile New Guinea tree boa, C.

carinata carinata.

Acrochordidae. The three species of highly aquatic Asian filesnake genus Acrochordus are so named

because of their strange loose-skinned, baggy bodies covered with tuberculate scales (McDowell 1979),

which distinguish them apart from all other living snakes. The two largest species, A. javanicus from

mainland Southeast Asia and Sundaland, and A. arafurae from northern Australia and southern New

Guinea, are freshwater inhabitants of slow-moving watercourses, but their smaller, widely distributed,

saltwater-dwelling relative, A. granulatus, has been recorded throughout Wallacea, excluding the southern

Moluccas.

Calamariidae. This family was until recently a subfamily of the Colubridae (See below) and its largest

genus, with 64 species, is Calamaria, the reedsnakes. Sulawesi is home to eleven endemic Calamaria,

and one widespread Indo-Philippine species, while a further endemic, C. ceramensis, inhabits Seram and

Ambon. The dwarf reedsnake genus Pseudorabdion contains 15 species, mostly in the Philippines and

Southeast Asia, but two species, P. sarasinorum and P. torquatum, are endemic to Sulawesi.

Two further genera, Calamorhabdium and Rabdion, are endemic to Wallacea. Calamorhabdium

acuticeps, R. forsteni, and R. grovesi are endemic to Sulawesi while C. kuekenthali is endemic to Bacan

Island in the northern Moluccas.

Colubridae: Ahaetullinae. This subfamily contains five genera of Asian and Australasian treesnakes,

three of which enter Wallacea. Genus Ahaetulla contains 17 species of vinesnakes, one of the most

widely distributed being A. prasina, which occurs as four subspecies, from India to the Philippines. The

nominate subspecies is present on Lombok, Sumbawa, Sulawesi and in the northern Moluccas. Of the

five flying snakes, genus Chrysopelea, two species are recorded within Wallacea (Mertens 1968). The

eastern subspecies of C. paradisi, C. p. celebensis, is present on Sulawesi, while C. rhodop/euron inhabits

eastern Wallacea: the nominate form from Bacan to Tanimbar, with a subspecies, C. r. viridis endemic to

the Sangihe Islands, north of Sulawesi.

The largest genus in the Ahaetullinae is Dendrelaphis with 47 species, whose members are known

as bronzebacks in Asia and as treesnakes in Australasia. Eight species occur in Wallacea, including the

Asian D. pictus, the Philippine D. marenae, and the Australo-Papuan D. calligastra. The other five species

are Wallacean endemics, including D. inornatus in the Lesser Sundas, with two subspecies, D. terrificus

on Sulawesi, and D. modestus, D. grismeri and D. keiensis in the northern, central and southern Moluccas

respectively (Vogel & van Rooijen 2008; van Rooijen & Vogel 2012).

Colubridae: Colubrinae. This large colubrid subfamily is represented in Wallacea by seven genera,

mostly of Asian origin. The nocturnal and highly arboreal catsnake genus Boiga contains 35 Asian and

two Australasian species. The mostly widely distributed Asian species is B. dendrophila, which comprises

nine subspecies, only one of which, B. d. gemmicincta, occurs east of the Wallace Line on Sulawesi

(Brongersma 1934). An equally widely distributed Australo-Papuan species is the brown treesnake, Boiga

irregularis, which inhabits northern Australia, New Guinea, the Solomon Islands, which was accidentally

introduced to Guam following the Second World War and where it has had a catastrophic effect on the

native avifauna (Rodda et al. 1999). It also occurs in the Moluccas and on Sulawesi. The two remaining

Boiga species are Wallacean endemics, B. hoeseli from the Lesser Sundas and B. tanahjampeana from

Tanahjampea Island, south of Sulawesi.

Six species of ratsnakes inhabit Wallacea. The genus Coelognathus is represented by one species

in the Lesser Sundas, C. subradiatus, and two species on Sulawesi, C. flavolineatus and C. erythrurus,

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

the latter represented by an endemic subspecies, C. e. celebensis. Sulawesi is also home to two endemic

ratsnake species, Gonyosoma jansenii and Ptyas dipsas, while the widely distributed Asian ratsnake, G.

oxycephalum, is one of several Asian snake species that are recorded no farther east than Lombok.

The Asian wolfsnake genus Lycodon contains 67 species, three of which occur in Wallacea. Lycodon

Capucinus is a perianthropic Southeast Asian species that has been distributed as far as New Guinea

(O’Shea et al. 2018), and remote Pacific and Indian Ocean islands. Lycodon subcinctus occurs throughout

the Lesser Sundas and it is probably the snake behind the stories of the “Timor krait” due to its black-

and-white banded patterning. There is a single Wallacean endemic, L. stormi/ on Sulawesi. The Asian kukri

snakes, Oligodon, are named for their sharp, kukri-like teeth which are used to slice into reptile eggs - or

for its effective defence against unware herpetologists. With 83 species this is the world’s second most

diverse snake genus. Four species occur in Wallacea, one Javanese species, O. bitorquatus, on Sumbawa,

and three Wallacean endemics: O. forbesi and O. unicolor, in the southern Moluccas, and O. waandersi

on Sulawesi and in the central Moluccas.

All these colubrine genera are primarily Asian in their origin and diversity, but one important Australo-

Papuan colubrine genus is also present in Wallacea. The groundsnake genus Stegonotus is the Australo-

Papuan equivalent of the Asian wolfsnakes Lycodon, and the genus is currently the subject of considerable

taxonomic revision. At the time of writing the genus contains 25 species (Kaiser et a/. 2018; 2019; 2020;

O’Shea & Richards 2021). Fourteen species inhabit New Guinea, two occur on Borneo, and one each

are known from the Philippines and Australia. The remaining seven species are Wallacean endemics,

including the northern Moluccas, S. batjanensis; central Moluccas, S. modestus; Kei Islands, S. keiensis;

Aru Islands, S. aruensis; Flores, S. florensis; Sumba; S. sutteri, and Semau, S. lividus. The description of

a new species from Timor-Leste is in preparation.

Pseudaspididae. This is a small family containing three genera and four species from Africa and Asia.

The Asian representatives belong to the genus Psammodynastes and are known as “mock vipers” because

of their viperine appearance and behaviour. The most widely distributed species, P. pulverulentus, occurs

from India and China, through the Sundaland and Philippine archipelagos, and into Wallacea where it is

recorded from Sulawesi, the Sula and Togian Islands, and Sumba and Sumbawa in the Lesser Sundas.

Sibynophiidae. The Sibynophiidae is still treated as a subfamily of the Colubridae by some authors.

It is a small family, containing three genera, two Asian and one American. The largest is Sibynophis with

nine species distributed from India to China, and south into Sri Lanka, Indonesia, and the Philippines.

Only S. Seminatus enters Wallacea, on Lombok.

Pareidae. There are slug- and sSnail-eaters throughout the world's tropics, and herpetologists

colloquially refer to the guild as the “goo-eaters”. The Pareidae contains four genera of Asian slug-eaters,

the largest being Pareas, with twenty species, but only P. carinatus has been recorded in Wallacea, from

Lombok.

Natricidae. Another former subfamily of the Colubridae, the Natricidae contains the freshwater-

dwelling keelbacks and watersnakes, including familiar genera like Natrix in Europe and Thamnophis

and Nerodia in North America. Not all species are benign, however, and Rhabdophis tigrinus has caused

fatalities in Japan, with serious bites also attributed to R. subminiatus from mainland Southeast Asia.

Three of the four genera present in Wallacea are of Asian origin, and one is more Australo-Papuan.

Hebius is a large Asian genus with 44 species, of which two occur on Sulawesi and neighbouring islands,

including H. sarasinorum and H. celebicum. Genus Rhabdophis contains 27 species, with three entering

Wallacea. Rhabdophis chrysargoides is widespread on Sulawesi and its satellite islands, while R. callistus

is endemic to north Sulawesi. A single specimen of R. chrysargos has been collected from Flores, but the

species is widespread west of the Wallace Line. Xenochrophis trianguligerus is also widespread west of

the Wallace Line, but in Wallacea it is limited to Sulawesi and its satellite islands.

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O'SHEA, M.: (FOREWORD) Wallacea - A hotspot of snake diversity

In the Moluccas the influence of the Australo-Papuan keelback genus Tropidonophis is strongly in

evidence. The genus contains 19 species, 12 of which are endemic to New Guinea or New Britain, one

is Australo-Papuan, two are Philippine endemics and the final four inhabit the Moluccas, including T.

elongatus, on Seram and Ambon, and T. truncatus, T. halmahericus, and T. punctiventris on Halmahera

and its neighbours (Malnate & Underwood 1988). The first two of these species occur extralimitally in

western New Guinea, while the latter two are Wallacean endemics.

Homalopsidae. Another unique Indo-Australian family, the Homalopsidae contains 29 genera and

56 species (Murphy & Voris 2014). Most members are rear-fanged and mildly venomous, aquatic,

marine, brackish, or freshwater inhabitants, although the family also contains three genera of fangless

terrestrial snakes from the Moluccas, western New Guinea, and Sumatra, which were for a long time

considered incertae sedis. The four Moluccan species, known as short-tailed snakes, belong to the genus

Brachyorrhos, the fifth endemic Wallacean genus. Brachyorrhos albus inhabits Seram, Ambon, Nusa

Laut, and Bisa; B. gastrotaenius occurs on Buru, B. raffrayi on Ternate, and B. wallacei on Halmahera

(Murphy et al. 2012), although perhaps these last two should have been named the other way around

since Wallace had his famous ‘eureka moment’ on Ternate. The genus Hypsiscopus contains two species

of freshwater aquatic watersnakes, and both are found on Sulawesi. Hypsiscopus plumbea is widespread

west of the Wallace’s Line, while H. matannensis is a Wallacean endemic.

Two of the other Wallacean homalopsids are widespread Indo-Australian species, Cerberus

schneiderii and Fordonia leucobalia, the former preying on mudskippers and frogs, the latter feeding on

freshly moulted crabs or mud lobsters. Cerberus will enter rice paddies but Fordonia is a purely mangrove

and estuarine dweller. Another mangrove swamp inhabitant occurs in the extreme southeast of Wallacea,

Myron karnsi, a small piscivorous species endemic to the Aru Islands.

Elapidae: Elapinae. The Elapinae is the subfamily containing the cobras, kraits, and coralsnakes

of Asia, cobras and mambas of Africa, and coralsnakes of the Americas. Only two species extend their

ranges into Wallacea. The Indonesian spitting cobra, Naja sputatrix, occurs in the Lesser Sundas, in the

Inner Banda Arc islands as far east as Alor. The king cobra, Ophiophagus hannah, the longest venomous

snake in the world, is the only terrestrial elapid recorded from Sulawesi and its satellite islands, Buton,

and Peleng in the Banggai Islands.

Elapidae: Hydrophiinae. The second elapid subfamily is the Hydrophiinae. It contains all terrestrial

venomous snakes from Australia, New Guinea, the Solomon Islands, and Fiji, and the marine seasnakes

and sea kraits (these latter groups are omitted from this discussion). Death adders, genus Acanthophis,

are short, squat, nocturnal, sit-and-wait ambushers that occupy the vacant “viper niche” across Australia

and New Guinea, and both New Guinean species also occur in Wallacea. Acanthophis laevis is a smooth-

scaled species that occurs throughout New Guinea and extends its range into the Moluccas, where it is

found from the Obi Islands, through Seram and its satellites, to the Kei and Aru Islands. The rough-scaled

A. rugosus has a limited range in southern New Guinea, but it too has been recorded from the Moluccas,

on the Tanimbar Islands. Only one other New Guinea elapid is represented in Wallacea. Muller’s crowned

Snake Aspidomorphus muelleri, is widely distributed throughout New Guinea, and it has also been

collected on Seram in the central Moluccas.

Viperidae: Crotalinae. The pitvipers inhabit the Americas and Asia, and they are well presented in

the Philippines and Sundaland, and three endemic species also inhabit Wallacea. The Island pitviper,

Trimeresurus insularis, inhabits virtually every island in the Lesser Sundas, as far east as Romang and

Kisar Islands, which, along with Timor, represent the closest viper populations to Australia. A second

species, T. fasciatus, is endemic to Tanahjampea Island and is the fourth endemic snake species on

that small (< 200 km?) island. The famous temple pitvipers of Malaysia and the Philippines are also

represented in Wallacea, by the endemic Sulawesi temple pitviper, Tropidolaemus laticinctus, which is

distributed island wide, and also found on Buton Island in the south and the Sangihe Islands in the north.

llacea is the eastern Russell’s

€ =

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

viper, Daboia siamensis. The Viperinae is a group primarily associated with Europe, the Middle East and

Africa, but two species occur further east. The western Russell’s viper, D. russelii, inhabits most of South

Asia, including Sri Lanka, while the eastern Russell’s viper demonstrates a much patchier distribution. It

has large but disjunct ranges in Myanmar and Thailand, and smaller ranges in southern China, Taiwan,

and eastern Java. Populations of this highly venomous snake, probably the most dangerous snake in the

region, are also located on Sumbawa and the Komodo Islands (Thorpe et a/. 2007).

From the above it is clear that Wallacea is home to an eclectic assortment of snake families, genera,

and species, and that it illustrates elements of the much larger regions that surround it, plus a few

endemic Wallacean taxa for good measure.

Table 1. Snake genera of Wallacea.

Legends: T/W/E - Total species in genus / Species in Wallacea/Endemic species in Wallacea.

Extralimital distribution: AF - Africa; AR - Arabia; AU - Australia; EA - East Asia; EM - Eastern Melanesia (Samoa,

Fiji); IC - Indochina; MI - Micronesia (Palau, Caroline Is.); NC - New Caledonia; NG - New Guinea; PH - Philippines;

SA - South Asia; SL - Sundaland (Borneo, Sumatra, Java, Bali); SO - Solomons; WA - Western Asia; Introduced

populations of Indotyphlops braminus worldwide, Python bivittatus in Florida, Boiga irregularis on Guam, and

Lycodon capucinus on Indian & Pacific Ocean islands are not listed.

Wallacean archipelagos: LS - Lesser Sundas; SU - Sulawesi; NM - northern Moluccas (Halmahera and satellites);

CM - Central Moluccas (Seram and satellites); SM - southern Moluccas (Kei, Aru, Tanimbar islands and satellites).

Numbers in the columns indicate the number of species present.

Taxon —Cis|s«CT/W/E | ~——___sé#Exttrallimitaldistribution S| LS | SU | NM | CM | SM |

TYPHLOPIDAE: ASIATYPHLOPINAE

[Gyeiouphions | _4/i/a_ | ___EndemiotoSuawesi | [a] | |

[Malayotyohiops | ia/it SPSL] STS tft

[Ramphotyphlops__|_16/5/2_| IC, SL, PH, NG, SO, NC, MI (Palau, Carolines) | | 2 | 2 | 2 | 2 |

|Sundatyphlops | 4/4/1_ | ———EndemictolesserSundas | 1 |) | CCU]

GERRHOPILIDAE

SA, IC, SL (Java), PH, NG RTS es

[CYLINDROPHIIDAE

CYLINDROPHIIDAE

XENOPELTIDAE

Xenopeltis | 2/4/0_ | SA IGEASL | ATT

(a O20 2 0 (OO

Python | 9/40 | CAR SAIGSL dE A ATT

Simaia | 6/4/38 | ANG

NG, SO, NC (Loyalty Is.), EM, Mi(Palau)_ | | 2 | 4 | 4 | 1

ACROCHORDIDAE

Acrochordus | 3/4/0_ | ST SAIC,SLNG AUT AE AT A Tt |

Calamaria (93/42/11) SAIC PHASE | at Tt

[Calamorhabdium | _2/2/2 | __EndemictoSulawesi&Moluccas | | 4 | 4 | |

[Pseudorabdion | 15/2/2 | IG PHL | | TT

Endemic to Sulawesi ae

Ahaetulla

Chrysopelea

Dendrelaphis 47/8/4

Book4.indd 12 02-Jun-21 21:46:40

O'SHEA, M.: (FOREWORD) Wallacea - A hotspot of snake diversity

Table 1 (continuation)

| Taxon | T/W/E | __—Extralimitaldistribution | -LS_| SU | NM | CM | SM_

COLUBRIDAE: COLUBRINAE

[Coelognathus | 7/3/0 | SAIGSLPH | A] 2 TT

[Gonyosoma | 6/2/4 | SAICSLPH | A] ATT

tycoon | 67/3/4 | CWASAICGEASLPH | 2 | 2 || 2 |

Oligodon | 83/4/38 | WASAICEASLPH | 2 | 2 | a | 2

Ptyas | 13/4/4 | WASAICEASLPH | | A |

Stegonotus | 24/7/7 | NG AUUSLPH | S| A | 2 | 2

[Psammodynastes | _2/4/0 | SANIGEASLPH | A] AT A] |

Sibynophis | 9/4/0 | SAICEASL,PH | A ||

Pareas | 20/4/0 | SAG EASL CT A ||

Hebius | 47/2/2 | SAG EASL | CE

[Rhabdophis | 27/3/41 | SAICEASLPH | 2 | 2 ||

Tropidonophis____—|_-49/4/2 | NG AU PH | | A

Xenochrophis ee ee

HOMALOPSIDAE

5/1/0

2/2/14

AF, AR, WA, SA, IC, EA, SL, PH

pC SAICEASLPH Tt

|Aspidomorphus | 3/4/0 | NGC

VIPERIDAE: CROTALINAE

Trimeresurus | 54/2/2_ | SAICEASL.PH | 2 | A ||

Tropidolaemus_— | 5/4/1_ | SA(South India) IC,SL.PH | | A ||

AF, WA, SA, IC, EA (China, Taiwan),SL(Java) | 1 | | | | |

po Total species per archipelago! 31 | 57 | 29 | 23 | 26 |

Acknowledgements

The author would like to thank Hinrich Kaiser (Zoologisches Forschungsmuseum Alexander Koenig,

Bonn, Germany) for reading and commenting on a draft of this Foreword.

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TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figures 1-8. Snakes of Wallacea. 1 - Brahminy blindsnake, Indotyphlops braminus, aka the parthenogenetic

“flowerpot snake” distributed worldwide; 2 - Timor blindsnake, Sundatyphlops polygrammicus, a species with five

subspecies in the Lesser Sundas, this being the Timorese form; 3 - Timor pipesnake, Cylindrophis boulengeri, one

of seven Wallacean endemic pipesnakes; 4 - New Guinea ground boa, Candoia aspera, a species that also occurs in

the northern Moluccas; 5 - Reticulated python, Malayopython reticulatus, the longest snake in the world reaches at

its southeastern extreme on Timor; 6 - Little filesnake, Acrochordus granulatus, a coastal species widely distributed

across Wallacea; 7 - Lesser Sunda bronzeback, Dendrelaphis inornatus, with Inner and Outer Banda species, this

being the Timorese subspecies; 8 - Lesser Sunda ratsnake, Coelognathus subradiatus, which inhabits the Lesser

Sundas and also Enggano Island off Sumatra.

Book4.indd 16 02-Jun-21 21:46:42

O'SHEA, M.: (FOREWORD) Wallacea - A hotspot of snake diversity

ce eee

GE «.

Figures 9-16. Snakes of Wallacea. 9 - Island wolfsnake, Lycodon capucinus, is a widely distributed species that is

introduced to many islands; 10 - Timor groundsnake, Stegonotus sp. nov., the species description is in preparation;

11 - Dog-faced watersnake, Cerberus schneideri, a common mangrove and coastal rice-paddy species in Timor;

12 - Crab-eating mangrove snake, Fordonia leucobalia, a widespread and highly variable mangrove swamp species;

13 - King cobra, Ophiophagus hannah, the longest venomous snake in the world, this specimen from India; 14 -

Smooth-scaled death adder, Acanthophis laevis, from New Guinea but also present on Seram and neighbouring

islands; 15 - Island pitviper, Trimeresurus insularis, green specimens inhabit Timor-Leste, yellow specimens also

occur in eastern Timor-Leste, and on Wetar, while Komodo specimens are green or cyan in colour; 16 - Eastern

Russel’s viper, Daboia siamensis, from the Komodo Islands, the most dangerous snake in Wallacea.

€ -

Book4.indd 17 02-Jun-21 21:46:43

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Acknowledgements

Unlike the previous volumes, this new work took more than three years to compile and produce, and was

brought to fruition through the support of numerous generous people - our family members, friends

and colleagues. We would express our deep gratitude to them as well as their institutions!

Board of the Entomological Society of Latvia, Riga (https://facebook.com/LatvijasEntomologijasBiedriba)

Board of NABU (Naturschutzbund Deutschland, Berlin (https://en.nabu.de)

Matthias HARTMANN, M.Sc. and Director (Naturkundemuseum Erfurt, Germany)

Christoph NEUMANN, Dr. (Crop Life International, Brussels, Belgium)

Andrey SHKARUPIN, M.Sc. (Riga, Latvia), an altruistic Supporter of biodiversity research and conservation

in Papuan Region

Laszlo WAGNER (Budapest, Hungary) (http://east-indonesia.info)

Kristine GREKE, M.Sc. (Latvian National Museum of Natural History, Riga), Edwin TELNOV and Alisa

TELNOVA (both Dzidrinas, Latvia) for their support and understanding

All the friendly (and not friendly) and welcoming people of the Moluccas, Lesser Sunda Islands, Raja

Ampat, New Guinea, the Philippines, Solomon Islands and Sulawesi, who shared their amazing

islands, cultures, even homes with us, and provided all other assistance during our sometimes too

frequent and not necessarily wanted visits

Responsible institutions, local counterparts and their staff in charge for arranging required permits

All the other persons, including all the authors, who friendly contributed to this volume

All papers are peer reviewed, so we take this opportunity to thank all our competent anonymous

referees and consultants!

Those referees who decided not to remain anonymous are acknowledged in person: Luca BARTOLOZZI

(Museo di Storia Naturale “La Specola”, Florence, Italy), AleS BEZDEK (Institute of Entomology, Ceské

Budejovice, Czech Republic), Leonidas-Romanos DAVRANOGLOU (Oxford, United Kingdom), Rainer

GUNTHER (Museum fiir Naturkunde, Berlin, Germnay), Klaus-Gerhard HELLER (Friedrich-Alexander-

University of Erlangen-Nuremberg, Erlangen, Germany), Andreas HERRMANN (Stade, Germany),

Martin HONEY (Natural History Museum, London, United Kingdom), Denis KEITH (Museum des

Sciences naturelles et de Préhistoire, Chartres, France), Bernard LANDRY (Muséum d'histoire

naturelle, Geneva, Switzerland), Jackson MEANS (Virginia Museum of Natural History, Martinsville,

U.S.A.), Alexandr I. MIROSHNIKOV (Russian Entomological Society, Krasnodar, Russia), Paul OLIVIER

(Griffith University, Brisbane, Australia), Dan A. POLHEMUS (Bernice P. Bishop Museum, Honolulu,

U.S.A.), Robert W. SITES (Columbia, U.S.A.), Gunther THEISCHINGER (Australian Museum, Sydney),

Tomas TICHY (Technical University of Ostrava, Czech Republic), and Mark G. VOLKOVITSH (Zoological

Institute of the Russian Academy of Sciences, Saint Petersburg, Russia).

——

if |

Book4.indd 18 02-Jun-21 21:46:43

RICHARDS, S. J., TuATURADI, B., Krey, K. & Donnellan, S. C.: A new stream-dwelling frog of the genus Litoria ...

pp. 19—33

A new stream-dwelling frog of the genus Litoria

Tschudi, 1838 (Anura: Pelodryadidae) from Salawati

Island, Indonesia

urn:lsid:zoobank.org: pub: /B/91247-399A-4E15-9DB2-C9535/7A/7EF28

STEPHEN J. RICHARDS *, BURHAN TJATURADI *, KELIOPAS Krey ° & STEPHEN C.

DONNELLAN “

1 - Corresponding author: Herpetology Department, South Australian Museum, North Terrace,

Adelaide, South Australia 5000, Australia; steve.richards@samuseum.sa.gov.au

2 - Center for Environmental Studies, Sanata Dharma University (CESSDU), Yogyakarta,

Indonesia; btjaturadi@gmail.com

3 - Department of Biology, Faculty of Mathematics and Natural Sciences, University of Papua,

Manokwari, Indonesia; keliopaskrey@ymail.com

4 - Herpetology Department, South Australian Museum, North Terrace, Adelaide, South Australia

5000, Australia; steve.donnellan@samuseum.sa.gov.au

Abstract: We describe a moderately small (10 males 23.3-26.9 mm, one female 31.5 mm SVL) new species of the

speciose pelodryadid genus Litoria Tschudi, 1838 from Salawati Island in the Raja Ampat Archipelago, a group of

islands off western New Guinea. The new species is mottled green and brown, has extremely long limbs, and was

found in lowland tropical rainforest where males called from low vegetation along a rocky, clear-flowing stream. The

advertisement call is a series of sharp ‘chip’ notes produced in variable combinations, and sometimes preceded by

a longer ‘buzz’ note. Genetically, the new species is related to a clade of predominantly montane, torrent-breeding

frogs that is most diverse in the mountains of mainland New Guinea. It appears to be unique among members of

this clade in having pigmented eggs. This description brings to three the number of frog species recently described

from Salawati that are to date known only from that small land-bridge island.

Key words: Amphibia, treefrog, new species, Raja Ampat Islands, New Guinea.

Introduction terrestrial herpetofauna of Salawati remained

poorly documented. However, recent studies there

Book4.indd 19

The Raja Ampat islands lie immediately to

the west of the Vogelkop Peninsula in West Papua

Province, Indonesian New Guinea. The archipelago

includes four major islands: Misool, Salawati,

Batanta and Waigeo. Unlike Waigeo and Batanta

which are oceanic islands with notable bird

(Beehler 2007), mammal (Helgen 2007), reptile

(Jacobs 2003; Oliver et al. 2008) and invertebrate

(Polhemus & Allen 2007) endemism, Misool and

Salawati are land-bridge islands that share a

common shallow water platform with the adjacent

mainland (Beehler 2007) and they are generally

considered to have a depauperate subset of the

nearby mainland fauna (e.g. Helgen 2007). Although

the Raja Ampat islands are part of the famous coral

triangle that is well known for its significant marine

biodiversity (Asaad et al. 2018), until recently the

have resulted in the descriptions of two new species

of geckos (Cyrtodactylus irianjayaensis Rosler,

2001 and Lepidodactylus pollostus Karkkainen,

Richards, Kraus, Tjaturadi, Krey et Oliver, 2020)

(Oliver et al. 2008; Karkkainen et al. 2020), and

two new species of microhylid frogs (Cophixalus

salawatiensis Gunther, Richards, Tjaturadi et

Krey, 2015, and Xenorhina salawati Gunther,

Richards, Tjaturadi et Krey, 2020) (Gunther et al.

2015; 2020). Here we describe a new species of

the pelodryadid frog genus Litoria from Salawati,

bringing to three the number of frogs currently

known only from this small land-bridge island. More

survey effort on the nearby mainland and on all of

the Raja Ampat islands is required before levels of

amphibian endemism on Salawati can be assessed

with confidence.

02-Jun-21 21:46:43

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TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Material and methods

Frogs were located using head torches and by

tracking advertisement calls. Voucher specimens

were fixed in 10% formalin and stored in 70%

ethanol. Liver samples for genetic analysis were

stored in 90% ethanol prior to specimen fixation.

All preserved animals and most genetic samples

of the new species are lodged in the Museum

Zoologicum Bogoriense, Cibinong, Indonesia

(MZB), and several genetic sub-samples are stored

in the Australian Biological Tissues Collection

(ABTC) at the South Australian Museum, Adelaide,

Australia (SAMA). Measurements, terminology,

and abbreviations follow Tyler (1968) and Oliver

et al. (2019). Measurements were made to the

nearest 0.1 mm with callipers (SVL - body length

from snout to vent, TL - tibia length from heel to

outer surface of flexed knee, HL - head length, from

tip of snout to posterior margin of tympanum, HW

- head width at level of tympana) or a dissecting

microscope fitted with an optical micrometer (all

other measurements): EN - distance from anterior

corner of eye to posterior margin of naris; IN -

internarial distance, between medial margins of

external nares; EYE - horizontal diameter of eye;

TYM - horizontal diameter of tympanum including

tympanic annulus; 3FD - transverse diameter

of disc of finger 3; 3FP - transverse diameter of

penultimate phalanx of finger 3; 4TD - transverse

diameter of disc of toe 4 and 4TP - transverse

diameter of penultimate phalanx of toe 4. Sex was

determined by examination of vocal slits, nuptial

pads, the presence of eggs or maturity of oviducts,

and by observation of calling. Calls were recorded

using a Sennheiser ME66 microphone, K6

powering module, and a Marantz PMD-660 digital

recorder. Calls were analysed using Avisoft-SASLab

Pro (v4.34, available from Avisoft Bioacoustics

http://www.avisoft.com/) using procedures and

terminology recommended by Kohler et a/. (2017).

Comparative material was examined at

the American Museum of Natural History, New

York (AMNH), Natural History Museum, London

(BMNH), Museum of Comparative Zoology,

Harvard (MCZ), Naturalis, Leiden (RMNH), South

Australian Museum, Australia (SAMA), Museum

fur Naturkunde Berlin (ZMB), the Natural Sciences

Resource Centre of the University of Papua New

Guinea, Port Moresby (UPNG), Museum Zoologicum

Bogoriense, Cibinong, Indonesia (MZB) and the

Queensland Museum, Brisbane (QM) (Appendix 1).

Additional information for comparisons was taken

from published papers on the relevant groups (e.g.

Tyler 1968; Menzies 1993). SJR refers to original

field collection numbers of S.J. Richards. Tissues

used in the genetic analysis are associated with

vouchers in the following additional institutions:

Australian Museum, Sydney (AMS), Museum and Art

Gallery of the Northern Territory, Darwin (MAGNT),

and the Western Australian Museum (WAM).

Frozen or alcohol preserved tissues were

available from 36 Litoria and Nyctimystes

specimens representing 23 species, and five

outgroup species (Appendix 2). Selection of ‘torrent

associated’ taxa was based on the mitochondrial

DNA sequence phylogenetic analysis of Rosauer et

al. (2009) anda broader unpublished mitochondrial

barcode survey of Melanesian pelodryadid frogs

(Donnellan, Richards & Mahony unpublished). DNA

was extracted using a Puregene DNA isolation kit

(Gentra Systems, Minneapolis, MN, U.S.A.) following

the manufacturer’s protocol for DNA purification

from solid tissue. A fragment of the mitochondrial

NADH dehydrogenase subunit 4 (ND4) gene

was amplified and sequenced using the forward

primers 5’-TGACTACCAAAAGCTCATGTAGAAGC-3’

with the reverse primer

5’-CATTACTTTTTACTTGGATTTGCACCA-3’. Each PCR

was carried out in a volume of 25 ul with a final

concentration of 1X GeneAmp PCR Gold buffer, 2-4

mM MgCl, 200 M of each dNTP, 0.2 mM of each

primer and 0.5 U of AmpliTaq Gold DNA polymerase

(Applied Biosystems, Foster City, CA, U.S.A.).

Amplifications consisted of an initial denaturation

step of 94°C for 9 min, followed by 34 cycles of PCR

with the following temperature profile: denaturation

at 94°C for 45 s, annealing at 55°C for 45 s, and

extension at 72°C for 1 min, with an additional final

extension at 72°C for 6 min. The double-stranded

amplification products were visualised on 1.5%

agarose gels and purified using an UltraClean PCR

clean-up DNA purification kit (Mo Bio Laboratories

Inc., CA) before cycle-sequencing using the BigDye

Terminator v3.1 cycle-sequencing kit (Applied

Biosystems). The cycling protocol consisted of 25

cycles of denaturation at 96°C for 30 s, annealing

at 50°C for 15 s, and extension at 60°C for 4

min. All samples were sequenced on an Applied

Biosystems 3/700 DNA sequencer. Sequences

were aligned with Muscle v6.814b (Edgar 2004)

implemented in Geneious Pro v8.1.4 (Kearse et al.

2012) and are deposited in GenBank (Appendix 2).

We used the maximum likelihood approach

to reconstruct phylogenetic relationships among

the mitochondrial DNA sequences, implemented

in 1Q-tree (Nguyen et al. 2014) on the IQ-TREE

webserver (Trifinopoulos et a/. 2016). We estimated

02-Jun-21 21:46:44

Book4.indd 21

RICHARDS, S. J., TuATURADI, B., Krey, K. & Donnellan, S. C.: A new stream-dwelling frog of the genus Litoria ...

the best substitution model with ModelFinder

(Kalyaanamoorthy et al. 2017) following the BIC

criterion. We assessed branch support with 1000

ultrafast bootstrap pseudo-replicates (Hoang et al.

2017).

Net average sequence divergence between

lineages (dA) was calculated in MEGA v7 (Kumar et

al. 2016) as: dA = AXY - (dX + dY)/2, where AXY is

the average distance between groups X and Y, and

dX and GY are the within-group mean (Table 2).

Results

Description of a new species

The species described herein is assigned

to Litoria sensu Tyler and Davies (1978) based

on its molecular genetic relationships, pending

a phylogenetic based resolution of generic

boundaries within Pelodryadidae. Our genetic,

morphological and acoustic data support the

distinctiveness of the new species and indicate

that its relationships lie with a clade of Litoria from

mainland New Guinea that is characterised by its

torrent-breeding ecology.

Litoria amnicola sp. nov. (Figs 1—6, 8)

http://zoobank.org/9E2Z0CE59-18E3-4DDD-8642-

5BE/7E12/76B62

Suggested English common name:

Ampat torrent treefrog.

Raja

Holotype adult 6’ MZB Amph.12116 (FN = SJR7728):

Weybya Camp, Salawati Island, Raja Ampat Archipelago,

West Papua Province, Indonesia (00°57.383’S,

130°47.060’E; 50 m a.s.l.), collected on 24 June 2005

by S. Richards, B. Tjaturadi and K. Krey.

Paratypes 11 specimens: MZB Amph.12099-12104

(FN SJR7738, 7734, 7737, 7758, 7729, 7754); MZB

Amph.12106 (SJR7736) MZB Amph.12113-12115,

12117 (SJR 7739, 7759, 7753, 7740), same details as

holotype, except MZB Amph.12114 adult female and

MZB Amph.12099 subadult female, collected 24-25

June 2005.

Derivatio nominis: From the Latin ‘amnis’ = a

stream of water, and ‘cola’ = dweller, referring to

the preferred habitat of this species.

Diagnosis: A species of Litoria that can be

distinguished from all congeners by the following

combination of characters: 1) size moderately

small (10 adult males 23.3-26.9 mm SVL, one adult

female 31.5 mm SVL), 2) dorsum finely granular,

pp. 19—33

mottled green and brown, 3) vomerine teeth barely

detectable, 4) ear small, not exceeding 40% of eye

diameter, 5) vocal slits present, 6) limbs extremely

long (TL/SVL 0.59-0.64), 7) heel without prominent

conical tubercle(s), 8) finger webbing not extending

beyond tubercle at base of penultimate phalanx

on Finger 4, and 9) advertisement call a series

of distinctly pulsed, unmusical chirps and buzzes

uttered singly or in groups of up to 12 notes.

Description of holotype: Adult male with vocal

slits and pale brown nuptial pads. Morphometric

data are summarised in Table 1. Body moderately

slender, limbs long (TL/SVL 0.61), head moderately

wide (HW/SVL 0.34), slightly less than length

(HL/SV 0.35, HL/HW 1.05). Vomerine teeth tiny,

between and close to edge of choanae; scarcely

visible but detectable by scraping with tip of forceps.

Tongue broadly oval with distinct, broad posterior

notch. Vocal slits long, extending from slightly

anterior of angle of jaws halfway to front of mouth.

Snout broadly rounded with tip slightly truncated

in dorsal view (Fig. 1), truncate (nearly vertical) in

lateral view. Canthus rostralis well defined, curved;

loreal region steep, concave; nostrils closer to tip

of snout than to eyes; internarial distance greater

than distance from external naris to eye (EN/IN

0.85, IN/SVL 0.11, EN/SVL 0.09); eyes large (EYE/

SVL 0.14), prominent, protruding in dorsal and

ventral views; pupil horizontal, without pigment

flecks on nictitating membrane. Tympanum small

(TYM/SVL 0.05), slightly less than one third

diameter of eye (TYM/EYE = 0.32), annulus distinct

except dorsal third obscured by thick, slightly

curved supratympanic ridge; series of small,

pale tubercles located below tympanum. Skin of

dorsum finely granular, less so (nearly smooth) on

dorsal surfaces of limbs; ventrally throat and chest

smooth with numerous scattered low tubercles;

abdomen coarsely granular (Figs 1, 3). Ventral

surfaces of limbs smooth. A patch of large, white

tubercles below vent extends distally approximately

4 mm along posterior margins of each thigh. A

series of low, pale tubercles along outer edge of

foot, extending from heel to midpoint of T5. Some

scattered pale tubercles on outer edge of forearm,

not forming linear ridge. Fingers moderately short

with expanded terminal discs (3FD/SVL 0.05)

with distinct marginal grooves; relative lengths of

fingers: 3>4>2>1; webbing limited, on inside of

F4 reaching to just below subarticular tubercle at

base of penultimate phalanx, continuing as broad

flange along edge of finger to base of disc; on

outside of F3 reaching only halfway to tubercle at

base of penultimate phalanx then continuing to

Ree

PAA

02-Jun-21 21:46:44

Book4.indd 22

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

disc as broad flange (Fig. 2), webbing only a basal

trace between 2 and 3, absent between 1 and 2.

Hand with poorly defined narrow inner, and oval

outer, metacarpal tubercles, and distinct narrow

dermal ridge along outer edge of hand. Nuptial

pad on first finger pale brown, elongate (1.6 mm

long), broader distally (0.5 mm) than proximally

(0.25 mm). Toes with expanded terminal discs

with terminal grooves, 4TD same as 3FD (3FD/4TD

1.00). Webbing between toes reaches to base of

disc on inside of T5, and on outside of T3 and T2;

and to about halfway to subarticular tubercle at

base of penultimate phalanx on both sides of T4,

then extending to disc as narrow flange; relative

lengths of toes: 4>5>3>2>1, but T5 only marginally

longer than T3. Foot with narrow, elongate inner,

and small, low, round outer metatarsal tubercle.

Colouration in life (Fig. 5): body and limbs rufous

brown dorsally, extensively mottled with bright

green; brown areas most intense on anterior of mid-

dorsum, on snout including loreal region, and in

broad band across dorsal surfaces of both thighs,

these brown patches enclosing more intense

green blotches. Green patch between eyes forming

poorly defined inter-orbital bar. Posterolateral

surfaces of body translucent, with scattered low,

pale-cream tubercles; prominent pale-cream

tubercles extend in short row from posterior edge

of eye below tympanum to posterior edge of curved

Supratympanic ridge. Ventrally abdomen white,

throat yellowy cream without dark stippling around

angle of jaws; ventrolateral surfaces of body and

ventral surfaces of thighs and tibiae translucent

pinkish-grey, tibiae bones visible through skin (Fig.

3); posterior surfaces of thighs uniform grey and

hidden surfaces of thighs peppered with brown

spots. Peppering of fine dark-brown specks present

on dorsal surfaces of limbs, extending onto dorsal

surfaces of fingers, toes and webbing. A row of

low, pale-cream tubercles along outer edge of foot

and several small pale-cream tubercles on outer

edge of forearm. Iris pale yellow with rufous brown

reticulations.

Colouration in preservative: green markings on

dorsum and limbs have become blue, background

brown and fine brown peppering have become

purplish brown; fine brown peppering on limbs more

conspicuous in preservative than in life. Ventral

Surfaces uniformly cream, except for extensive fine

brown peppering on plantar surfaces; peppering

restricted to few scattered spots on palmar

Surfaces.

Variation in the type series: Measurements

and body proportions of the type series show

limited variation (Table 1). Male body size 23.3-

26.9 mm SVL; one adult female (MZB Amph.12114)

with SVL of 31.5 mm, one immature female (MZB

Amph.12099) with SVL of 27.3 mm. Snout normally

broadly rounded with truncate tip in dorsal view, but

MZB Amph.12102 with more pointed tip. Vomerine

teeth normally detectable only by scraping with

forceps, but visible (though very small) in several

specimens including MZB Amph.12114 (adult

female). 3FD is larger than 4TD in all but three

of the types, in which 3FD = 4TD. In preservative

dorsal colour variable, ranges from pale (e.g. MZB

Amph.12115) to dark (MZB Amph.12113) brown;

extent of blue markings is highly variable, from few,

small, scattered spots (e.g. MZB Amph.12104) to

extensive blue patches across much of dorsum

(e.g. MZB Amph.12113). Fine dark brown stippling

on body and limbs more conspicuous on paler

Specimens and on lower jaw. Most specimens with

narrow blue (green in life) bar between eyes or

immediately anterior of eyes; this bar complete or

broken into series of spots (e.g. MZB Amph.12114),.

Row of pale tubercles below tympanum invariably

present but varies from scarcely detectable to

prominent (see e.g. fig. 6).

Advertisement call: Our analysis of the call

structure of Litoria amnicola sp. nov. is based on

detailed examination of 11 calls produced by the

holotype (MZB Amph.12116) at an air temperature

of 26°C. Calls of L. amnicola sp. nov. are series of

2-13 short, distinctly pulsed notes sounding to the

ear like a harsh ‘chip’, produced singly ‘single’, in

couplets ‘double’ notes, or (rarely) in triplets ‘triple’

notes, occasionally preceded by one or two longer

‘buzz’ notes (Fig. 8). Because the sequence and

rate at which ‘single’, ‘double’, and ‘triple’ notes

are produced Is variable we defined groups of notes

as separate calls if the first note in one group was

separated from the last note in the previous group

by at least two seconds, irrespective of the note

type. Eleven calls of the holotype were produced

at a rate of 0.15 calls/s, with intervals of 2.95-

7.02 s between calls (mean = 4.49; SD = 1.38).

Dominant frequency is 2858-3081 Hz (mean =

3004, SD = 85.80). Of the eleven calls analysed,

four contain one ‘buzz’ note and one contains

two consecutive ‘buzz’ notes; ‘buzz’ notes always

precede ‘chip’ notes. ‘Buzz’ notes are 0.13-0.19

Ss long (mean = 0.16, SD = 0.02) and contain 18-

25 pulses (mean = 21.7, SD = 2.5), produced at

a rate of 126-133 pulses/s (mean = 129.7, SD =

2.29, n = 6). In contrast, 81 shorter ‘chip’ notes

are just 0.007-0.046 s long (mean = 0.021, SD =

0.008) and contain 2-7 pulses (mean = 3.6, SD

02-Jun-21 21:46:44

Book4.indd 23

RICHARDS, S. J., TuaTuRADI, B., Krey, K. & Donnellan, S. C.: A new stream-dwelling frog of the genus Litoria ...

Figures 1-4. Litoria amnicola sp. nov. 1 - Dorsal view of preserved holotype (MZB Amph.12116) [scale bar = 5 mm];

pp. 19—33

== >> . “ee 7

ks Se : 4 De

a P of Sag eee

7 ~ * ae

2 - Left hand of holotype (MZB Amph.12116) in palmar view [scale bar = 2 mm]; 3 - Ventral view of holotype (MZB

Amph.12116) in life - note visibility of bones through skin; 4 - Three pigmented eggs extracted from adult female

paratype (MZB Amph.12114) [scale bar = 2 mm].

= 0.94), produced at a rate of 130-214 pulses/s

(mean = 154.7, SD = 19.01, n = 81). Twenty-one

of the ‘chip’ notes analysed (26%) are produced as

‘single’ notes, 54 (67%) are produced as ‘double

notes’ (Fig. 8), and six (7%) are produced as triple

notes. The substantial differences in both length

and pulse number between ‘buzz’ and ‘chip’ notes

Suggest that they do not represent ends of a

spectrum but are discrete note types.

Differential diagnosis: Litoria amnicola_ sp.

nov. is compared here with small (male SVL ~20-

30 mm), green or green and brown New Guinean

Litoria that do not breed in lotic waterbodies or

for which breeding biology is unknown, as follows:

Litoria amnicola sp. nov. differs from L. albolabris

(Wandolleck, 1911) and L. mystax (Van Kampen,

1906) in having dorsum brown and green (vs

uniform green), dorsum granular (vs smooth), in

having a series of pale cream tubercles beneath the

tympanum (vs a single white bar), and limbs longer

(TL/SVL 0.59-0.64 vs 0.50-~0.55 in L. albolabris

and 0.54 in L. mystax; Tyler 1968); from members

of the L. bicolor (Gray, 1842) group (L. bibonius

Kraus et Allison, 2004, L. chloristona Menzies,

Richards et Tyler, 2008, L. contrastens (Tyler, 1968),

L. eurynastes Menzies, Richards et Tyler, 2008,

L. lodesdema Menzies, Richards et Tyler, 2008,

and L. viranula Menzies, Richards et Tyler, 2008),

02-Jun-21 21:46:44

Book4.indd 24

TELNov, D.

et al. (eds) 2021: Biodiversity,

Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figures 5-/. Litoria amnicola sp. nov. 5 - Hele (MZB ae 12116) in life: 6 - Banas (MZB aah 12115) in

life; 7 - Habitat of Litoria amnicola sp. nov. at the type locality. Males called from low vegetation adjacent to this

stream.

1000

500

fs 0

-500

-1000

S 6

oN 4 itil

om 2 nN bi |

a= 0 —_

bail lig ' . mere i heii ail ' " Wii whe

Time sec

Figure 8. Waveform (top) and spectrogram (bottom) of an advertisement call of Litoria amnicola sp. nov. (holotype:

MZB Amph.12116) showing variation in note structure including an introductory ‘buzz’ note and nine ‘chip’ notes

produced individually and in couplets.

02-Jun-21

21:46:46

Book4.indd 25

RICHARDS, S. J., TsATURADI, B., Krey, K. & Donnellan, S. C.: A new stream-dwelling frog of the genus Litoria ...

in having dorsum brown mottled with green (vs

predominantly green or occasionally brown), tarsi

with row of tubercles on outer margins (vs absent),

and snout truncate in lateral view and at tip in dorsal

view (vs rounded in lateral view and obtusely pointed

or rounded in lateral view; Kraus & Allison (2004a);

Menzies et al. (2008)); from L. christianbergmanni

Gunther, 2008 in its smaller size (male SVL 26.9-

31.2 mm in L. christianbergmanni), dorsum brown

and green (vs uniform green with white, yellow or

pale green spots), and white bar below eye lacking

(vs present in L. christianbergmanni); from L.

chloronota (Boulenger, 1911) in having body size

smaller (males 27-32 mm in L. chloronota), dorsum

brown and green (vs mottled pale and darker green

with or without yellow spots), snout broader (EN/

IN 0.79-0.97 vs 0.63-0.71 in L. chloronota), and

limbs longer (TL/SVL 0.59-0.64 vs 0.52-0.58 in

L. chloronota; Menzies 1993); from members of

the L. gracilenta (Peters, 1869) group (L. aruensis

(Horst, 1883), L. auae Menzies et Tyler, 2004, L.

callista Kraus, 2013, L. elkeae GUnther et Richards,

2000, L. eschata Kraus et Allison, 2009, L. kumae

Menzies et Tyler, 2004, and L. robinsonae Oliver,

Stuart-Fox et Richards, 2008) in having dorsum

brown and green (vs plain green with or without

pale or dark spots), and pale canthal and postocular

Stripes absent (vs present: Menzies and Tyler

2004; Kraus 2013); from L. havina Menzies, 1993

in having dorsum brown and green (vs uniformly

green), and fleshy rostral spike in males lacking

(vs present); from L. rubrops Kraus et Allison,

2004 in having dorsum brown and green (vs green,

usually speckled with black or darker green),

vomerine teeth small but detectable (vs absent

in L. rubrops), limbs longer (TL/SVL 0.59-0.64 vs

0.48-0.59 in L. rubrops), and iris pale yellow with

rufous brown reticulations (vs iris red in L. rubrops;

Kraus & Allison 2004b); from L. nigropunctata

(Meyer, 1875), L. iris (Tyler, 1962), L. majikthise

Johnston et Richards, 1994, L. ollauro Menzies,

1993, L. richardsi Dennis et Cunningham, 2006, L.

singadanae Richards, 2005, and L. verae Gunther,

2004, in having fingers with limited webbing (not

extending beyond subarticular tubercle at base of

penultimate phalanx, vs finger webbing extensive,

extending beyond subarticular tubercle at base

of penultimate phalanx), and further from L. iris

in lacking (vs having) violet patch in axilla and

having posterior surfaces of thighs uniform grey (vs

posterior of thighs blue, red, or yellow, frequently

blotched with white or purple, in L. iris), from L.

ollauro in having dorsum variably green and brown

(vS uniform green or green with yellow spots in

pp. 19—33

L. ollauro) (Menzies 1993), from L. majikthise by

having posterior surfaces of thighs uniform grey in

life (vs red in L. majikthise), and by pear|-white post-

ocular bar lacking (present in L. majikthise), from

L. richardsi in having dorsum mottled brown and

green (vs dorsum with irregular black lines), throat

and finger and toe webbing without extensive black

markings (vs present in L. richardsi), and periphery

of tympanic membrane not transparent (vs

transparent in L. richardsi), from L. singadanae in

having posterolateral surfaces of belly and posterior

surfaces of thighs uniform translucent pinkish grey

in life (vS posterior of venter and hidden surfaces of

legs uniform orange in L. singadanae), tympanum

much smaller (TYM/EYE 0.25-0.40 vs 0.69-0.87 in

L. singadanae), tympanic membrane pigmented (vs

transparent in L. singadanae), and from L. verae in

prominent crenulated skin fold along outer margins

of tarsi lacking (vs present), and dorsum without

small brown spots aligned transversely (vs present

in L. verae).

Litoria amnicola sp. nov. is compared here

with small (male SVL ~20-30 mm) green, green

and brown or predominantly brown New Guinean

Litoria that breed in lotic waterbodies or that are

presumed to do so, as follows: Litoria amnicola

sp. nov. differs from L. bulmeri (Tyler, 1968) by

size smaller (males 23.3-26.9 mm vs 29-34 mm

SVL), dorsum brown and green (vs uniform green)

and black lateral stripe absent (vs present); from

L. brongersmai by having legs longer (TL/SVL 0.59-

0.64 vs 0.52-0.54) snout truncate (vs rounded)

and dorsum brown and green (vs dark brown) (Tyler

1968); from L. dorsivena (Tyler, 1968) by canthus

rostralis curved (vs straight), vomerine teeth poorly

developed (vs prominent) and dorsum brown and

green (vs brown) (Tyler 1968); from L. fuscula Oliver

et Richards, 2007, by size smaller (males 23.3-

26.9 mm vs 29-30 mm SVL), legs longer (TL/SVL

0.59-0.64 vs 0.52-0.54) and having dorsum brown

and green (vs uniform brown) (Oliver & Richards

2007); from L. longicrus (Boulenger, 1911) in

having dorsum brown and green (vs green), dorsum

granular (vs smooth), and in having a series of pale

cream tubercles beneath the tympanum (vs a single

white bar); from L. modica (Tyler, 1968) in having

snout tip truncate at tip (vs rounded in both dorsal

and lateral views), skin of dorsum granular (vs

smooth with scattered low tubercles), tongue with

posterior notch (vs absent), and eggs unpigmented

(vS pigmented); from L. napaea (Tyler, 1968) by

size larger (males 23.3-26.9 mm vs 18.8-22.7 mm

SVL), skin granular (vs smooth), snout truncate

(vs obtusely pointed) and eggs unpigmented (vs

wee

02-Jun-21 21:46:46

Book4.indd 26

TeLNov, D. et al. (eds) 2021:

Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Callimedusa_tomopterna_64894

Pseudacris_species_ 55395

Hyla_japonica_64965

100 100 , Hyla_crucifer_32559

Hyla_crucifer_32560

Litoria_luteagrp_SJR1443

100 Litoria_eucnemis_32215

Litoria_eucnemis_90052

100 , Litoria_sauroni_76032

99 Litoria_sauroni_98775

Litoria_infrafrenata_15302

Litoria_infrafrenata_48722

Nyctimystes_humeralis_44909

Nyctimystes_humeralis_44903

Litoria_brevipalmata_24910

100 Litoria_prora_136053

Litoria_prora_98252

400| Litoria_amnicola_90195

Litoria_amnicola_90101

Litoria_amnicola_90197

Litoria_rivicola_100683

Litoria_rivicola_100685

Litoria_spartacus_98336

100; Litoria_arfakiana_100659

Litoria_arfakiana_100689

Litoria_oenicolen_118073

Litoria_oenicolen_118074

Litoria_pratti_142224

Litoria_dorsivena_112514

Litoria_modica_45060

Litoria_modica_46364

100, Litoria_becki_116940

Litoria_becki_116987

Litoria_bulmeri_133760

Litoria_angiana_48210

Litoria_angiana_48337

Litoria_micromembrana_98290

Litoria_micromembrana_48643

Litoria_leucova_101474

Litoria_bicolor_28366

Litoria_bicolor_100072

100

40 99

66 100

7 fe

21 0.1

50 73 100

100

Figure 9. Maximum likelihood (ML) phylogram of relationships between mitochondrial ND4 nucleotide sequences of

Litoria species. Numbers at nodes are ML bootstrap proportions.

pigmented) (Tyler 1968);

1911) by size smaller (males 23.3-26.9 mm vs

29.5-31.5 mm SVL) (Tyler 1968), canthus rostralis

curved (vs straight) and vomerine teeth poorly

developed (vs well developed); from L. rara Gunther

et Richards, 2005 by size larger (males 23.3-26.9

mm vs 20.4 mm SVL), skin dorsally finely granular

(vs smooth), and dorsum brown mottled with

green (vs dark grey); and from L. scabra Gunther

et Richards, 2005 in having skin of dorsum finely

granular (vs strongly tubercular), crenulated dermal

ridge on foot and forearm absent (vs present), and

snout truncate in lateral view (vs acute, projecting

beyond lower jaw). From the new species’ closest

known relative, Litoria rivicola Gunther et Richards,

2005 (see Molecular genetic comparisons below),

Litoria amnicola sp. nov. can be distinguished by

size larger (males 23.3-26.9 mm vs 20.4-22.1 mm

SVL), dorsum finely granular (vs coarsely tubercular),

and snout truncate (vs acutely projecting) (GUnther

& Richards 2005).

Distribution and habitat:

sp. nov. iS a scansorial species that is currently

known only from a single location at an altitude of

about 50 m a.s.|. near the north coast of Salawati

Island (Map 1), where males called from heights of

between 50 cm and 3 mon low foliage along a clear-

flowing, rocky stream in lowland rainforest (Fig. 7).

None were observed adjacent to forest pools or

other lentic habitats. It is not known whether Litoria

amnicola sp. nov. is more widespread on Salawati.

The type locality is at the northern edge of the only

substantial mountain range on the island, and it

is possible that the species’ distribution there is

constrained by the availability of cool, fast-flowing

streams draining the mountains. Images of a small

treefrog taken recently along a similar clear-flowing

stream on Batanta Island by Dr. Tibor Kovacs

may represent this species, but voucher material

is required to confirm that record. On Salawati,

calling males climbed onto shrubs and started

calling before dark, and calling activity continued

until after midnight each night. An adult female

(MZB Amph.12114) contained heavily pigmented

from L. pratti (Boulenger,

Litoria amnicola

02-Jun-21 21:46:48

Book4.indd 27

RICHARDS, S. J., TuATURADI, B., Krey, K. & Donnellan, S. C.: A new stream-dwelling frog of the genus Litoria ...

eggs; three eggs extracted for examination (Fig.

4) measured 1.3-1.5 mm diameter (mean = 1.4

mm). Although the deposition of pigmented eggs

in forest streams is common among some groups

of Melanesian pelodryadid frogs (e.g. the Litoria

genimaculata (Horst, 1883) group), this is the first

reported case of a species within the large clade of

montane, torrent-breeding Litoria from New Guinea

(Fig. 9) that is Known to have pigmented ova (e.g.

Tyler 1968; Tyler & Davies 1978; Menzies 2006;

S.J. Richards, unpublished data). The eggs of the

closest known relative of L. amincola sp. nov,, L.

rivicola, have not been reported but tadpoles have

a large, ventrally located oral disc with numerous

papillae, as is typical of other torrent-breeding

species (Gunther & Richards 2005). The tadpole of

L. amnicola sp. nov. has yet to be discovered. The

eggs and tadpoles of many New Guinean torrent-

breeding Litoria have not been documented, and

it is possible that additional species within this

clade have pigmented ova. Integrated reproductive,

ecological and molecular studies are required to

provide a better understanding of the evolution of

reproductive traits in this group of frogs.

Molecular genetic comparisons: The

final alignment for the mitochondrial ND4 gene

comprised 845 bp. In a phylogram of relationships

among mitochondrial ND4 sequences, the

sequences from L. amnicola sp. nov. are the sister

to the two L. rivicola sequences with strong support

and both sit within the “torrent- breeding frog” group

(Fig. 9). The net uncorrected sequence divergence

(dA) for ND4 between L. amnicola sp. nov. and its

sister taxon, L. rivicola, was 0.14, which is at the

upper end of the range of values among other sister

taxon pair comparisons within the torrent-breeding

frog group (0.11-0.15, Table 2).

Acknowledgements

Fieldwork in the Raja Ampat Islands was

Supported by Conservation International and the

South Australian Museum. We are most grateful

to Yance de Fretes, Muhamad Farid and Jatna

Supriatna of Conservation International for their

organisational skills and Support, and to Herlina

Kafiar, Rizana Kurniati, Elias Kore, Sofia Roni, Arthur

Tipawael and Adelina Werimon for assistance

in the field. We are also extremely grateful to the

Indonesian Institute of Sciences (LIPI) for their

Support and approval of specimen export, the

Forestry Department, especially Kantor Sub BKSDA

2, Sorong and Directorate Jenderal PHKA, and to

pp. 19—33

Lisa Capon who produced Figs 1—8 and the map,

and Tibor Kovacs who kindly provided images of a

similar Litoria species from Batanta Island while this

manuscript was in review. Linda Ford (then AMN#),

Barry Clarke (BMNH), José Rosado (MCZ), Hellen

Kurniati and Mumpuni (MZB), Pim Arntzen (RMNH#),

Carolyn Kovach and Mark Hutchinson (SAMA), and

Rainer Gunther (ZMB) kindly provided access to

specimens in their care, and tissue samples were

provided by Paul Doughty, Rainer Gunther and

Michael Mahony. We also thank Luke Price, Steven

Myers and Tessa Bradford for performing the DNA

sequencing, and Paul Oliver and Rainer Gunther

for their careful and constructive reviews of the

manuscript.

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Table 1. Measurements (mm) and ratios of the type series of Litoria amnicola sp. nov.

All vouchers have MZB Amph registration numbers. *MZB Amph.12116 is the holotype. All soecimens are adult

males except MZB Amph.12114 which is an adult female and MZB Amph.12099 which is a subadult female.

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Book4.indd 30

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Table 2. Net average sequence divergence (dA) for the mitochondrial ND4 gene between sister species pairs

Litoria rivicola - L. amnicola sp. nov.

Sister species pair dA sd

L. angiana — L. bulmeri

[L. arfakiana-L.oenicolon ——C—C—C‘~dS‘ WL

[L.becki- L.modica ss —“‘(CSC‘«dCS‘ OCW‘:

Litoria albolabris (Wandolleck, 1911)

Papua New Guinea: Sandaun Province, Aitape,

SAMA R4947 (syntype).

Litoria auae Menzies et Tyler, 2004

Papua New Guinea: Gulf Province, Purari River, near

McDowell Is, UP2490 (holotype), SAMA R57262-63

(paratypes).

Litoria bulmeri (Tyler, 1968)

Papua New Guinea: Madang Province, Upper

Aunjung Valley, Schrader Mtns, SAMA R5625

(holotype).

Litoria chloristona Menzies, Richards et Tyler,

2008

Papua New Guinea: National Capital District,

Waigani, SAMA R9122a; Central Province, Brown

River, SAMA R13251a-c; Gulf Province, Kopi, SAMA

R63502-9 (all paratypes).

Litoria chloronota (Boulenger, 1911)

Indonesia: West Papua Province, Arfak Mountains,

BM1947.2.31.20 (syntype), UP8380-8.

Litoria contrastens (Tyler, 1968)

Papua New Guinea: Eastern Highlands Province,

Barabuna, SAMA R5845 (holotype), SAMA R6450

(paratype); Western Highlands Province, Noreikova,

SAMA R5847 (5 specimens).

Litoria dorsivena (Tyler, 1968)

Papua New Guinea: West Sepik Province, Telefomin,

SAMA R7901 (holotype).

Litoria elkeae Gunther et Richards, 2000

Indonesia: Papua Province, Siewa_ River,

Amph.3866-9, QMJ70490-2 (all paratypes).

MZB

Litoria eurynastes Menzies, Richards et Tyler,

2008

Indonesia: Papua Province, Siewa River, MZB

Amph.14651-5; Papua New Guinea: Manus

of Litoria from the “torrent-breeding” frog group.

Appendix 1. Specimens examined.

Province, SAMA R63497-501

paratypes).

Lorengau, (all

Litoria fuscula Oliver et Richards, 2007

Indonesia: Papua Province, Derewo River Basin,

MZB Amph.11822 (holotype), SAMA R60724

(paratype).

Litoria havina Menzies, 1993

Papua New Guinea: Western Province, Ok Kam,

UP7281 (holotype); Western Province, Ok Ma, UP

8406-7 (paratypes); Western Province, Ok Kam,

SAMA R38596-7; Southern Highlands Province,

Agogo Range, SAMA R60173-7.

Litoria iris (Tyler, 1963)

Papua New Guinea: Chimbu Province, Bamna,

BM 1961.1226 (holotype); Southern Highlands

Province, Tari, UP3115-35; Eastern Highlands

Province, Ubaigubi, UP8289-90; Enga Province,

Porgera, UP 7148-67, Lake Tawa, SAMA R7/1599;

Sandaun Province, Telefomin, SAMA R5423,

R58/4; Hela Province, Gigira Ridge, SAMA R71598.

Litoria kumae Menzies et Tyler, 2004

Papua New Guinea: Southern Highlands Province,

Tari, UP3108 (holotype), SAMA R52760-61

(paratypes).

Litoria leucova (Tyler, 1968)

Papua New Guinea: Sandaun Province, Mt Stolle,

SAMA R44091-2, UP8604-6.

Litoria lodesdema Menzies, Richards et Tyler,

2008

Papua New Guinea: East New Britain Province,

Kerevat, SAMAR7/046-47, R/049, R/055-56,

R7058-59, R8439a,b.

Litoria longicrus (Boulenger, 1911)

Indonesia: Papua _ Province, Wendessi, BM

1947.2.22.60-61 (syntypes).

02-Jun-21 21:46:49

Book4.indd 31

RICHARDS, S. J., TuATURADI, B., Krey, K. & Donnellan, S. C.: A new stream-dwelling frog of the genus Litoria ...

Litoria majikthise Johnston et Richards, 1994

Papua New Guinea: Western Province, Tabubil,

SAMA R44093 (holotype), UP6734, 7305-9, 8501-

8, 8602-3, SAMA R44094-44101 (all paratypes).

Litoria modica (Tyler, 1968)

Papua New Guinea: Eastern Highlands Province,

Oruge, MCZ 52856 (holotype), MCZ 52857-52861,

SAMA R8108 (paratypes).

Litoria mucro Menzies, 1993

Papua New Guinea: East Sepik Province, Near Rauit

Village, UP2741-3, 2745-56 (paratypes).

Litoria mystax (van Kampen, 1906)

Indonesia: Papua Province, Moaif, RMNH 4632

(holotype).

Litoria napeae (Tyler, 1968)

Indonesia: Papua Province, Snow Mountains,

Bernhard Camp, AMNH 49573, 49575, 4965/7,

(1022-24 (paratypes).

Litoria nigropunctata (Meyer, 1875)

Indonesia: Papua Province, Yapen Island, Mount

Waira, ZMB 63977, Yapen Island near Konti, SAMA

R61799.

Litoria ollauro Menzies, 1993

Papua New Guinea: Milne Bay Province, Agaun,

UP4644 (holotype), UP4634-43, 4645-51, 5070-

89 (paratypes).

Litoria pratti (Boulenger, 1911)

Papua Province, Wendessi,

Mountains,

BMNH

Indonesia:

1947.2.23.54-56, Arfak

1947.2.23.57-58 (syntypes).

pp. 19—33

Litoria richardsi Dennis et Cunningham, 2006

Papua New Guinea: Western Province, Tabubil,

SAMA R60283 (holotype); Upper Fly River, SAMA

R71602-5; Indonesia: Papua _ Province, Tir

River, Mamberamo Drainage, MZB Amph.11823

(paratype).

Litoria robinsonae Oliver, Stuart-Fox et Richards,

2008

Papua New Guinea: Gulf Province, Ivimka,

Lakekamu Basin, SAMA R55527 (holotype),

R55528-9 (paratypes).

Litoria scabra Giinther et Richards, 2005

Indonesia: Papua Province, Wapoga River Drainage,

MZB Amph.11335 (holotype), Amph.11336-11340,

SAMA R6070/7-9, ZMB6735/7-9 (paratypes).

Litoria singadanae Richards, 2005

Papua New Guinea: Morobe Province, Ridge above

Surim Camp, eastern Finisterre Mountains, SAMA

R60172 (holotype), UP 9968, SAMA R60171

(paratypes).

Litoria verae Giinther, 2004

Indonesia: Border of Papua and West Papua

Provinces, Wondiwoi Mountains, ZMB 62384.

Litoria viranula Menzies, Richards et Tyler, 2008

Papua New Guinea: Western Province, Bensbach

River, SAMA R63487 (holotype), Wegamu, SAMA

R63486-92 (paratypes); Indonesia: Papua

Province, Merauke, SAMA R13666a-c, R1366/7a-d.

Litoria wapogaensis Richards et Iskandar, 2001

Indonesia: Papua Province, Wapoga River, MZB

Amph.3873 (holotype), MZB Amph.3874-76, SAMA

R54595-98 (paratypes).

02-Jun-21 21:46:49

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Appendix 2. Tissue vouchers used.

Jere ecseorah aman Jimre et eee lee)

voucher ince Voucher Accession

ar Tay

> wenssae0 [7953578

a CC

iia ——apoies [60065 ra raat | roar pan ro5358

amnioola SP. nov

titoria [angiana AMIS RA24118

Litoria angiana Morobe AMS R124246

Litorila[arfakiana ZMB 62116

Utoria | arfakiana

Lito Westem [PNG [SAMAR65I67

Litera ven

100072 Mitchell Plateau (Queensland, [Australia WAM |WAM R167822 | |MT953590 |

Territor

Wales

SsaT6O_ |W Wnt [weston | NG 8 MTO5S59D

129812 —[Wavatipe —|Enga [ps — Ra0660— ron

Ilwraith Rang ges

Ciera [euenemis [80052 sineize ____[mwas76as

Litoria infrafrenata 15302 Australia |SAMAR34972 MT953596

8722 [Karkar land AWS RE24631__[MTO53507

ioia7a | MtStole SAMAR4I092__|MT953598

Litoria lutea group SJR1443 Bougainville Autonomous | PNG SJR1443 MT953599

Region of

Bougainville

Litoria | micromembrana 48643 Bundi = {Madang PNG [AMS R124552 MW357881

SECU. MW357880

Highlands

fe —}rez Yuk Creek near eo SJR14541 MT953605

Tabubil

Highlands

prora 136053 yeaa Su ae Western PNG | SJR14205 MT953606

ivicola | 100683 oo ZMB 67361 [MT953608

700685 __[30km SE Nabire ZBGALI7___ [MTO5360S

76032 [Dark End Lumber te SIRIGED_—___[MT9536I0

Highlands

Highlands MW357885

Highlands MW357884

Count

55805 ———— MTOSSOI6

Book4.indd 32 02-Jun-21 21:46:50

RICHARDS, S. J., TuATURADI, B., Krey, K. & Donnellan, S. C.: A new stream-dwelling frog of the genus Litoria ...

pp. 19—33

Batanta

ag a Amn

i. ! Se -<,

y - " oe a Soeyayapura

a . Po =

[Ca Papua New Guinea

- ™ ~~

SMadangy -_ . f

‘Vis . tee;

fo. ea on

~~" Merauke™,

Kilometers

Map 1. The type (and only known) locality for Litoria amnicola sp. nov. on Salawati Island.

Book4.indd 33 02-Jun-21 21:46:50

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Book4.indd 34 02-Jun-21 21:46:50

Book4.indd 35

Roster, H. & GauLke, M.: On the biology of Ernstkeller’s cave gecko, Gekko ernstkelleri (Gquamata: Gekkonidae) ...

pp. 35-48

On the biology of Ernstkeller’s cave gecko, Gekko

ernstkelleri Rosier, Siler, Brown, Demegillo

et Gaulke, 2006 (Squamata: Gekkonidae),

with emphasis on reproductive biology

urn:lsid:zoobank.org:pub:133AEEF3-CB26-4EAC-AC1E-EBCOAEF39991

HERBERT ROSLER * & MAREN GAULKE 2

1 - Corresponding author: Senckenberg Naturhistorische Sammlungen Dresden, Museum fur

Tierkunde, Sektion Herpetologie, Konigsbrucker Landstrasse 159, D-01109, Dresden, Germany:

herbertroesler@aol.com

2 - GeoBio-Center Ludwig-Maximilians-Universitat,

Munchen, Germany; mgaulke@web.de

Richard-Wagner-Strasse 10, 80333,

Abstract: Gekko ernstkelleri Rosler, Siler, Brown, Demegillo et Gaulke, 2006 is a medium sized member of the

genus Gekko Laurenti, 1768, endemic to the central Philippine Island of Panay. This rupicolous species inhabits

limestone outcrops, especially entrance areas of limestone caves. It is mainly nocturnal. Four pairs of G. ernstkelleri

were kept in husbandry for eight years, and their species-specific behaviour and breeding biology recorded. No

agonistic behaviour was observed among couples living permanently together, while males display territorial induced

aggressive behaviour towards each other. G. ernstkelleri is oviparous. Egg clutches contain one or two hard-shelled

eggs. Fertile eggs are attached to inorganic substrates such as glass or plastic tubes. In captivity, juveniles hatched

after 64-179 days. Snout-vent length (SVL) of hatchlings is 29.5-39.5 mm, tail length (TaL) 31.0-47.0 mm, and

weight 0.6382-1.1632 g. The relative clutch weight varies between 9.87% and 21.08%.

Key words: Sauria, Gekko ernstkelleri, description, distribution, ecology, behaviour, reproduction.

Introduction

With a land area of around 12 000 km’,

Panay is the sixth-largest island of the Philippine

Archipelago. The herpetology of Panay is fairly well

explored (e.g. Ferner et al. 2001; Gaulke 2011).

Thirteen gecko species (family Gekkonidae

Gray, 1825) are recorded from the island. Seven

of them are endemic to the Philippines, but only

Gekko ernstkelleri is endemic to Panay (Rosler et al.

2006). The present chapter deals with behavioural

observations made during eight years of husbandry

of this species, as well as with observations in

nature.

Material and methods

Occasional field observations on Gekko

ernstkelleri are recorded by one of the authors

(M. Gaulke) since 2004. Observations and

investigations on four pairs of G. ernstkelleri kept

in husbandry by the first author (HR) are recorded

since 2013. Both data sets are compared and /

or supplemented, to obtain a profound knowledge

on the biology of G. ernstkelleri, with emphasis on

reproductive biology.

The geckos were kept in four equivalent glass-

terraria with similar equipment. Air temperature

and relative humidity were recorded in a 24 hour

cycle for one year with a hygrothermograph. Climatic

terraria conditions were not equivalent to conditions

in nature, especially winter temperatures inside the

terraria were significantly lower than on Panay. The

geckos were fed with different insects and insect

larvae, vegetarian food was not ingested.

Morphometric parameters (size of living

adults and juveniles, egg-length and egg-width)

were measured with a digital caliper, thickness

of eggshells with a micrometer. The weight of

eggs, eggshells, and juveniles was taken with a

pharmaceutical micro scale, adult weight with a

laboratory scale.

The reproductive effort (= relative clutch mass,

02-Jun-21 21:46:51

Book4.indd 36

TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

RCM) was determined by using formulas from

Pianka & Pianka (1976) and Hielen (1993), egg-

volume with the formula 4/3nab2.

Abbreviations used in the text:

SVL - snout-vent length;

TaL — tail length;

BW - body weight;

EL - egg length;

EW - egg width.

Results

Short description and systematics

With a total length of around 200 mm (max.

SVL 92 mm), G. ernstkelleri Rosler, Siler, Brown,

Demegillo et Gaulke, 2006 is a medium-sized

member of the genus Gekko. SVL of mature males

82-92 mm (N = 8), of mature females 78-85 mm

(N = 10), TaL/SVL 1.18-1.36 (data recorded from

free ranging individuals on Panay, Gaulke 2011).

Gekko ernstkelleri is specific due to the

following features: 11-15 dorsal tubercle rows;

14-18 subdigital scansors on first, and 16-19 on

fourth finger; 15-17 subdigital scansors on first,

and 17-19 on fourth toe; dorsal tail tubercles

only up to fifth caudal whorl; subcaudals enlarged.

Males can be determined by enlarged cloacal

tubercles and 36 - 42 preanofemoral pores, which

are absent in females (Rosler et a/. 2006).

The life coloration of G. ernstkelleri is somewhat

variable. The dorsal side is olive to brown-olive,

with distinctive whitish and dark-brown spots.

The different-sized whitish spots are arranged

in irregular transverse rows across the dorsum,

being more prominent on head and nuchal region,

and fading caudad in intensity and size. Paired

dark-brown spots are present on the vertebral

region. While they are arranged in transverse rows

posteriorly, they sometimes merge in the nuchal

region and on the head into darkish skin patches.

The dorsal side of the tail is dark and light banded,

with the light bands being about half as wide as the

dark bands. Around the posterior part of the tail,

the light bands can build confluent rings. The dark

interspaces have blackish-brown margins, and are

mottled brown in between. Especially in males they

can be blackish-brown throughout. Gular region

and venter are light yellowish to light greyish-brown,

with fine brown dotting. The ventral side of the tail is

light yellowish-brownish with large brown blotches,

which become denser towards the tail-tip (Fig. 1).

Males are occasionally more distinctly colored than

females (Fig. 2).

Within the genus Gekko the species G.

ernstkelleri belongs to the Gekko monarchus group

(Rosler et al. 2011).

Wood et al. (2020) published a phylogeny

and systematic concept of the Gekko clade, and

suggested an arrangement in seven subgenera.

For the Philippine species-complex they introduced

the subgenus Archipelagekko Wood, Guoa,

Travers, Su, Olson, Bauer, Grismer, Siler, Moyle,

Andersen et Brown, 2020. This phylogenetic

lineage contains, with exception of G. gecko, all

other Philippine species of the genus Gekko (G.

athymus Brown et Alcala, 1962, G. carusadensis

Linkem, Siler, Diesmos, Sy et Brown, 2010, G. co/

Brown, Siler, Oliveros, Diesmos et Alcala, 2011, G.

crombota Brown, Oliveros, Siler et Diesmos, 2008,

G. ernstkelleri Rosler, Siler, Brown, Demegillo et

Gaulke, 2006, G. gigante Brown et Alcala, 1978, G.

mindorensis Taylor, 1919, G. monarchus [Schlegel,

1836], G. palawanensis Taylor, 1925, G. porosus

Taylor, 1922, G. romblon Brown et Alcala, 1978,

G. rossi Brown, Oliveros, Siler et Diesmos, 2009),

as well as G. kikuchii Oshima, 1912, which is

distributed up to Lanyu Island, Taiwan. All species

of the subgenus Archipelagekko resemble each

other widely regarding their phenotype (head-,

body- and tail-shape; SVL < TaL; similar coloration

and dorsal color pattern) (Brown & Alcala 1962;

1978; Brown et al. 2008; 2009; 2011; Linkem et

al. 2010; Rosler et a/. 2011).

Distribution

To present knowledge, G. ernstkelleri is

endemic to Panay Island, and on Panay only known

from different locations of the Northwest Panay

Peninsula, from the Provinces of Antique and Aklan.

It occurs from sea level up to around 300 m asl

(Gaulke 2011).

Ecology

Gekko ernstkelleri is an inhabitant of limestone

karst habitats. Most specimens were sighted in the

entrance areas of limestone caves within or close

to secondary or primary lowland limestone forests

(Fig. 3). Tunnel-like caves with wide entrance areas

on both sides are occupied throughout (Fig. 4). A

few individuals were sighted on trees surrounding

cave entrances, and on deeply corroded rocky

limestone outcrops close to the coastline.

02-Jun-21 21:46:51

Rdster, H. & GauLke, M.: On the biology of Ernstkeller’s cave gecko, Gekko ernstkelleri (Gquamata: Gekkonidae) ...

pp. 35-48

Figure 1. Adult male of Gekko ernstkelleri Rosler, Siler, Brown, Demegillo et Gaulke, 2006

(photo courtesy T. Hofmann).

‘N

Figure 2. Adult couple of Gekko ernstkelleri, male on the left and female on the right.

The female has an almost completely regenerated tail.

The diurnal and annual range of temperature Behaviour

and humidity in caves and deep limestone crevices

is more consistent than the outside air temperature Gekko ernstkelleri is a mainly nocturnal

and humidity (Alcala & Alcala 2005), besides, karst species. In nature, first active individuals were

habitats offer a wealth of places for egg-deposition. observed at around 4:30 pm, before darkness,

but their main activity time starts during sunset,

qQq ee 37

aan

Book4.indd 37 02-Jun-21 21:46:52

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figure 3. Gekko ernstkelleri often inhabits deeply karstified limestone walls and outcrops (NW Panay Peninsula).

Book4.indd 38 02-Jun-21 21:46:54

Book4.indd 39

Roster, H. & GauLke, M.: On the biology of Ernstkeller’s cave gecko, Gekko ernstkelleri (Gquamata: Gekkonidae) ...

between six and seven pm (Rosler et al., 2006;

Gaulke 2011). In their terraria, active specimens

were frequently observed outside their retreats

during daytime, especially following cool winter-

nights, to occupy sites close to light- and heat

sources. Even intensive movements outside their

terraria usually did not induce flight behaviour

during their diurnal thermoregulation periods,

while during almost complete darkness, the same

optical stimuli (in a distance of 0.5 to 2 m from the

terraria) led to flight behaviour. Acoustic stimull

without accompanying movements, such as human

voices, usually did not induce flight behaviour.

The G. ernstkelleri were kept in pairs in their

terraria throughout the investigation period of six

years, however, the partners were exchanged a few

times during this time. During this time, no intra-

sexual aggressive behaviour was observed, and

even an occasional exchange of the females did

not lead to noteworthy problems.

lt was not possible to keep G. gigante, another

endemic gecko from the Philippines, for a long

period of time under similar conditions as G.

ernstkelleri. Even the first socialization of a mature

male and a mature female of G. gigante (in March)

= i

pp. 35-48

resulted in fierce biting’s. This aggressive behaviour

was probably induced by a_non-compatible

hormonal state of the pair. The obviously sexually

motivated male attacked the disinterested female

of G. gigante fiercely, inflicting Severe bite injuries

on head and body, and biting off most toes of

the female’s hind limbs. Those conflicts can be

interpreted as intraspecific territorial behaviour.

Sexual unmotivated females of G. gigante provoke

the same behaviour in males, as they would show

towards conspecifics of the same sex. Continued

aggressive behaviour of males towards sexual

unmotivated females of G. gigante within artificial,

limited habitats is known from other members of

the genus Gekko as well (Kreuzer & Grossmann

2003).

At the start of the investigation period,

threatening behaviour among captive males of

G. ernstkelleri was regularly observed, because

they had constant visual contact with each other

(later, visual covers were provided). The threatening

behaviour started with staring at each other,

followed by wriggling movements of the slightly

raised tails. A trial to socialize two males had to be

cancelled. A direct confrontation involuntarily led to

02-Jun-21 21:46:56

Book4.indd 40

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

the mentioned threatening behaviour, followed by

fast biting attacks of the dominant individual (the

resident of the respective terrarium). The attacked

male tried to flee, but was chased and bitten by

the dominant male until it was removed from the

terrarium.

Unfortunately, the mating behaviour never

could be observed in G. ernstkelleri. Typical external

indications such as bite marks in the nuchal region

of the female could never be detected. Mating calls

of males, as known from different Gekko-species

(Mertens 1946; Bohme et al. 1985; Tang et al.

2001; Jono & Inui 2012; Chen et al. 2016) could

not be detected. Gekko ernstkelleri is not voiceless.

The geckos can emit a slight fear scream or a

defense call.

Many Squamata have developed species

behavioural models for the protection of their

offspring. Protective behaviour starts with the

search for adequate egg-laying sites, but can

also involve active defense of eggs and offspring

(Deegener 1918; Somma 1990).

In the case of G. ernstkelleri, observations in

natural habitats proved egg-guarding behaviour

(according to Shine 1988). Egg-clutches (often

attached to crevices on cave ceilings) usually are

guarded by one or two adults, which attack intruders

such as herpetologists, if getting close to the eggs.

Even during daytime, when at the first moment no

gecko can be seen, adults will immediately appear

at the clutch site if they notice disturbances (Gaulke

POTLAZ O12).

Egs-guarding behaviour also occurred in the

terraria. It could be directly observed if eggs were

not attached inside hiding places (plastic tubes)

but to open, easily accessible areas. Directly after

egg-deposition, adults remained in the direct

vicinity of the eggs. Sometimes, both partners

were seen close to the eggs, but more frequently

the female. Unlike observations in their natural

habitat, the adults in the terraria took refuge in

hiding places if disturbed, for example by handlings

inside the terrarium, and did not try to defend their

eggs. A while after egg-deposition, the females

only sporadically visited their clutch-sites. A more

or less constant egg-guarding behaviour was only

observed, if the eggs were attached inside plastic

tubes. Inside the tubes, almost always one or both

adults were seen during controls throughout the

entire incubation period. However, they were easily

dispelled from their guarding-post, without showing

any signs of threatening behaviour or trying to

attack.

All behavioural modes in relation to eggs and

hatchlings are summarized as “parental care”

(Petzold 1984; Shine 1988). Within the geckos,

three different stages of parental care can be

clearly differentiated: precaution (Search for, and

eventually preparing of a suitable clutch-site), care

(egg-suarding and egg-defence), and aftercare

(defence of hatchlings). Only few gecko species

show all three stages of parental care (Rosler

2005).

Stage one, searching for a suitable egg-

deposition place, is strongly developed in G.

ernstkelleri. In several caves with a high G.

ernstkelleri population, even intensive searches

did not reveal a single egg-clutch, obviously the

eggs were hidden very carefully in deep crevices.

On the other hand, sites considered as optimal by

the geckos, were used for extended periods of time,

often by several females. This was easily discernible

a5 OB

30 96

2 —

™ 30 2 =

2 =

15 a =

o or

= x

e 10 BR

5 86

Oo a4

month

BS Mean Maximum monthly tem perature

mean minimum monthly temperature

anmme Mean monthly ar humidity

Diagram 1. Annual climate (1990-2005) of Capiz

(N11° 23.33, E122° 37.67), Panay, the Philippines.

It should be noted that the microclimate within the cave

habitats of Gekko ernstkelleri is more balanced and, in

particular, the humidity is less subject to fluctuations.

35 95

20 8 =

Z =

Mas E

E i

fel

F 19

month

[8 mea Maximum monthly temperature

mean minimum monthly temperature

a= ea monthly ar humidity

Diagram 2. Climatic conditions in the Gekko ernstkelleri

terrariums in 2013 (deviations in the remaining years

are almost non-existent).

The geckos were kept in cooler and drier conditions than

within their habitat in Panay.

02-Jun-21 21:46:56

Book4.indd 41

Roster, H. & GauLke, M.: On the biology of Ernstkeller’s cave gecko, Gekko ernstkelleri (Gquamata: Gekkonidae) ...

by egg-clutches of different developmental stages

glued close to each other, often on the shell

remains of already hatched eggs. As mentioned

above, these places are regularly guarded by adults

(Gaulke 2012).

Under natural conditions the parental care

of G. ernstkelleri includes two of the three stages:

precaution and care, while no indications of

aftercare were observed. Observations under

captive conditions revealed a somewhat different

behaviour. In one case, the highly developed eggs

were devoured by one parent (it is not known

whether the male or the female, and it is not known

whether the embryos were alive or deceased

directly prior to the time of egg-eating). The

devouring of empty eggshells is a normal part of the

reproductive behaviour of females, this way they fill

up their depleted calcium resources effectively. If

ages

2,5

Jan Feb Mar Apr May Jun Jul Aue Se Oct Now Dec

month

Diagram 3. Monthly egg production (total n = 77) in

terrariums of four females of Gekko ernstkelleri from

2013-2019.

the embryos were dead prior to the egg eating, the

adult must have detected the condition olfactorily,

and used the eggs as a natural resource. But if the

embryos were still alive, it means that no bonding

towards the own eggs existed during the time of

egg eating. This would be a sign of a behavioural

disorder.

Two times, the own offspring (each time two

hatchlings) was devoured by the parents, both

times by different pairs. It is unknown, whether

both parents or only one adult was involved in this

cannibalistic behaviour. In both cases, the juveniles

hatched during night-time from eggs glued inside

plastic tubes, where they were guarded by the

parents.

In several species of the genus Gekko, e.g. G.

gecko (Linnaeus, 1758), G. grossmanni Gunther,

1994, G. petricolus Taylor, 1962, G. siamensis

pp. 35-48

and G. vittatus Houttuyn, 1782, observations in

captivity revealed a complex parental care (Koch

1968; Grossmann & Mudrack 1987; Grossmann

& Stein 1988; Treu 2001; Kreuzer & Grossmann

2003; Grossmann 2004). In other species, e.g. G.

badenii Szczerobak & Nekrasova, 1994, the own

hatchlings fall into the predator-prey system of

the adults (Hofmann 2007). Cannibalism could

also be observed in G. monarchus (Rosler 1981;

1982), a species related to G. ernstkelleri. As

phyletic criterion for both species, a low-threshold

of parental care could be primordial, or else, was

newly acquired because of the discontinuation of

predation pressure in the terrarium.

Among the Gekkonidae, parental care is

highest developed within the genus Gekko.

According to present knowledge the complexity of

parental care gives no group-specific indications

on infrasubgeneric level (Table 1). However, in

many Gekko species parental care has not yet

been investigated thoroughly. Further studies are

necessary to ascertain which behavioural elements

constitute parental care in other groups of affinity

(respectively subgenera sensu Wood et al. 2020)

within the genus Gekko, and whether a group-

specific distinction will be indicated.

100

eee mass (Yo)

0 10 20 30 4 30 6 7

mcubation time (days)

Diagram 4. Weight loss during the incubation period of

an egg from Gekko ernstkelleri. The egg was hatched

in an incubator, incubation temperature fluctuated

between 25-32 °C, relative air humidity > 90 %.

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02-Jun-21 21:46:56

Book4.indd 42

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Reproduction

Females used for breeding were photographed,

to enable recognition based on individual colour

patterns. They were numbered 1 to 4, and their

SVL and BW determined in 2014 and 2018. In

numerical sequence, mean values are: female 1

SVL 78 mm, BW 16.27 g, female 2 SVL 79 mm, BW

12.63 g, female 3 SVL 84 mm, BW 15.80 g, female

4 SVL 84 mm, BW 16.00 g.

All four female G. ernstkelleri were breeding,

captive conditions for the four pairs (terrarium

size, equipment, climatic conditions, diet) were

identical. Females 1 and 4 deposited fertile eggs

before completing their second year. Female 2 had

reached an age of two years and three months

before depositing its first fertile eggs, and female

3 four years and seven months. The late start of

egg laying in female 3 is not necessarily due to a

delay in reaching sexual maturity. Possible reasons

might be an acyclic hormonal process of the sexual

partners, individual sexual Uunacceptance, or

physiological problems.

Table 1. Parental care of species from three related groups of the genus Gekko.

Legends: The taxonomic structures follow the concept of Wood et al. (2020). Data on Gekko kuhli (Stejneger,

1902) and Gekko lionotum (Annandale, 1905) from Grossmann (2008), for further information see text.

The species-specific behavior of parental care is divided into three sections and assessed as non-existent (0)

or existing (1).

| Care __| Aftercare _

| Provision _

Subgenus Gekko a

IG.gecko. ——“—‘“‘ié;C

G. gecko

|Subgenus Archipelagekko | |

erties a a ae ae aaa

IGamonarchUsae a ween eee ei ee

Subgenus Lomatodactylus Se

[G.badenii-

G. badenii a as a es (ae

[Subgenus Ptychozoon— —(i|

[G.kuhi

[G.lionottum —s—“(‘irPSC Ud

Between July 5, 2013 and 10, 2019, the four

females produced 44 egg-clutches all in all. Most

clutches were deposited by females 1 (20) and

4 (14). Female 2 produced six clutches between

2015 and 2018, female 3 four clutches between

2016 and 2019. It could not be clarified, whether

the strongly differing number of egg-clutches

produced by the four females had physiological

reasons or has to be seen in context with captive

conditions (e.g. no free choice of mates).

On Panay, G. ernstkelleri lives in groups.

Knowledge on intraspecific patterns of relations

within these social structures does not exist.

Individual relationships (empathy) may influence

the egg-laying activities of a female as well as

physiological circumstances.

The duration of the breeding season as well as

the number of clutches within the breeding season

of the kept G. ernstkelleri differed among the four

individuals. The longest recorded breeding season

lasted around nine and a half months (October 16,

2017 to July 27, 2018). Within this timespan, F,

produced four egg-clutches in different time gaps.

During another breeding season, F, deposited the

same number of clutches within only about five

months (November 20, 2015 to April 30, 2016).

None of the females laid more than four egg-

clutches within one egg-laying cycle.

Similarly, quantitative graduated frequencies

are known from other gecko species. Osadnik

(1987) for example observed an_ individually

differing contribution to egg production in a group

of eight female Phelsuma dubia (Boettger, 1881)

throughout a two-year field study. Within six months,

the females produced between one and nine

clutches. The big difference in clutch quantity was

partly age- and size dependent. Nevertheless, two

same-sized females produced three versus nine

clutches within an egg-laying season of six months.

Differing egg-deposition frequencies influence

the population dynamic. A regularly spaced egg-

deposition frequency among all reproductive active

females leads to an exponential population growth.

Of the 44 egg-clutches produced by the four

02-Jun-21 21:46:57

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Rdster, H. & GauLke, M.: On the biology of Ernstkeller’s cave gecko, Gekko ernstkelleri (Gquamata: Gekkonidae) ...

females, 11 contained a single egg, and 33 two

eggs each (mean clutch size 1.78+0.42). Eleven of

the 77 eggs (14%) were infertile.

Clutches containing only one egg were

produced throughout the breeding season, but

much less so during the beginning and during

the midst of the breeding season (9% and 18%,

respectively). Most single-egg clutches were laid

towards the end of the breeding season (73%), a

strong indication for a state of exhaustion of the

reproductive active females towards the end of the

breeding season.

Different places for egg-deposition, partly

consisting of natural materials, partly of artificial

materials, were available in the respective terraria.

These were vertical and sloping crevices between

stone slabs (oil shale), large and flat pieces of

different types of bark, and plastic tubes with a

diameter of 35 mm and 50 mm, respectively.

The egg-laying behaviour of G. ernstkelleri is

fixed genetically. Within its natural habitat, females

glue their eggs in small crevices at cave walls or,

more often, on the ceiling of a cave (Fig. 5). In the

terraria fertile eggs were also glued to surfaces,

whereby hard inorganic materials such as glass or

hard plastic were preferred over organic matters

such as oak bark or cork. Egg-clutches with two

eggs were always glued in pairs together (Fig. 6).

Infertile eggs were sometimes found on the ground,

or aS a Smeared mass on the glass pane of the

terraria. Horizontal surfaces (the terrarium floor),

as well as unstructured clay soil or heaps of foliage

are rather untypical egg-deposition sites. Only once

pp. 35-48

a clutch was glued to a dry leaf on the ground (pers.

com. T. Hofmann).

At the start, the females preferred the thinner

plastic tubes (diameter 35 mm) for egg-deposition,

while no clutches were found in the plastic tubes

with a diameter of 50 mm.

To determine egg-weight, the preferred egg-

laying sites were modified later on. Therefore, the

interior of the plastic tubes was lined with thin

paper. This was meant to enable the observer to

remove the eggs from the surface after deposition

easily. However, it turned out that this manipulation

of the hitherto preferred egg-laying sites resulted

in the females looking for alternative egg-

deposition sites. Obviously they regarded paper

as an unsuitable substrate to glue the eggs on.

While choosing suitable sites for egg-deposition,

G. ernstkelleri’s primary selective criterion is

the quality of the substrate, while locality is of

secondary importance. The preference of substrate

quality over protecting shelters such as crevices

means that it is considered of higher importance

ethologically.

Searching for new egg-laying sites, the eggs

were now glued to freely accessible, mostly

moderately warm and darkish places along the

lower part of the sides of the terrarium glass. The

females did not use the available hiding places in

crevices. Lighter positions at the upper part of the

glass panes, which were exposed to more radiation

heating, were rarely used. Part of the eggs glued

outside the plastic tubes were protected by a

transparent, air-permeable cover by the observer.

Figure 5. A clutch of Gekko ernstkelleri stuck to the cave ceiling in a natural habitat, the Philippines, Antique

Province, Pandan Municipality, Barangay Sto. Rosario.

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02-Jun-21 21:46:57

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

The complete natural reproductive cycle of

G. ernstkelleri in nature is unknown. Developing

eggs were observed from October to end of June,

however, no field observations were conducted

during the months of July to September. Most newly

laid eggs were found in March. The tropical climate

of its natural distribution makes it likely that they

are reproducing throughout the year (compare

tropical taxa in Fitch 1970).

The climatic conditions provided for G.

ernstkelleri in captivity throughout the observation

period differed from the climate within their natural

habitat. Humidity as well as temperature provided

for the geckos in their terraria was lower (Diagrams

1 & 2). It is known that geckos kept in captivity can

tolerate differences in temperature and humidity

up to a certain degree. In some species it could be

determined already that these differences lead to

more or less pronounced deviation from the natural

species-specific reproductive cycle (Rdsler 2005).

Within the terraria G. ernstkelleri laid eggs

from October to July of the following year. During

August and September, no egg-laying activities

were noted (Diagram 3). It is noteworthy that part

of the egg-laying activities occurred during the

cool phase of husbandry. The observations show

that females of G. ernstkelleri were able to carry

eggs full term beside the decreasing, relatively low

ambient temperatures starting in October. Actually

a high number of egg-clutches were recorded in

January 2015, 2016, and 2019. A total of 10 eggs

were laid during these months. Only the relatively

low day- and night temperatures of the two following

months obviously influenced egg-development,

resulting in a low number of eggs. In February the

egg-production declined drastically to only four

eggs, followed in March by a slight increase to

seven eggs. In April and May egg-laying activities

reached their peak (14 eggs each). During the

winter months February/March neither a prolonged

gestation period nor a resorption of eggs were

observed (Rosler 2005).

To present knowledge, all members of the

genus Gekko glue their eggs to a substrate (Okada

et al. 2002; Rosler 2005; Kreuzer et al. 2013). In

about two-thirds of the eggs of G. ernstkelleri they

were glued more or less at one of the egg-poles

lying in their longitudinal axis, in the rest either flat

parallel, or slightly tilted towards the longitudinal

axis. This has to be considered for the assessment

of egg shape, and the computation of egg volume,

respectively. In eggs, which were glued at a pole,

the plane adhesive surface amounts to at least

two-thirds of egg-width or egg-length, respectively,

compared to being glued parallel to their longitudinal

axis. In the first instance this results in an objective

decrease of the egg-length, in the second case of

the egg-width. Accordingly, data on the proportion

EL/EW, as well as on the volume, must be regarded

as approximate values.

Only 26% of all eggs could be measured,

eggs glued to unfavourable sites could not be

measured. They had a length of 14.85-18.85 mm

(16.40+1.08, n=20), and a width of 11.40-13.90

mm (12.48+0.81, N = 20). Eggs laid in captivity

differ much less in size than those laid in nature.

The determined size of eggs laid in nature is: length

14.4-16.0 mm (N = 10) and width 10.9-12.8 mm (N

= 10) (Gaulke 2012). These slight deviations have

to be considered as normal intraspecific variations.

The form of the eggs of G. ernstkelleri is

roundish to oval. The value EL/EW is 1.07-1.42

(1.31+0.07, N = 20). Comparing the value of both

egs-axes, egg-length has a higher range (4 mm)

and variation (V 1.15) than egg-width (2.5 mm, V

0.6). The higher variation determined in egg length

is not a species specific character, as it is typical for

most species of the Gekkota (Kratochvil & Frynta

2005).

Merely a single egg was glued to a plastic tube

lined with paper, and therefore could be recovered.

lts weight immediately following deposition was

1.3967 g. During the incubation period of 76 days,

an egg mass decreased 10.5% (Diagram 4). A egg

mass reduction during embryogenesis is typical

for all hard-shelled eggs of the Sphaerodactylidae,

Phyllodactylidae, and Gekkonidae (Rosler 2005).

The determined egg mass reduction in the G.

ernstkelleri egg does not differ significantly from

those of other species in these three families

(Rosler in prep.).

The incubation period of geckonid eggs

shows species-specific differences. Besides,

the embryonic development is influenced by the

ambient temperature (Rdosler 1998). The prevalent,

relatively low temperatures measured at the egg-

laying sites of G. ernstkelleri in the terraria resulted

in a significantly prolonged incubation time. At the

egg-laying sites with the lowest temperatures (10-

15 cm above ground, distance towards the heat-

source 25-30 cm), the day/night temperatures

ranged between 22°C and 15.5°C during the winter

months, and between 25.5°C and 17.5°C during the

summer months. During the cool period, mean day

temperatures taken directly at the egg-deposition

sites ranged below 20°C (February, minimal

daily mean 18.8°C). Under these conditions, the

juveniles of G. ernstkelleri hatched after 82-179

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Roster, H. & GauLke, M.: On the biology of Ernstkeller’s cave gecko, Gekko ernstkelleri (Gquamata: Gekkonidae) ...

days (117424, n=33). Only two clutches were

deposited close to the light source (fluorescent

lamp), at a distance of six cm. The fluorescent

lamp operated 12 hours daily. The temperature at

the clutch site reached 29-33°C during daytime,

and 19-26°C during nighttime. One of the clutches

deposited close to the light source hatched after

64 days, the second one after 76 days.

Juveniles of G. ernstkelleri hatched during

daytime as well as during the night (Fig. 7). If they

hatched within the terraria, they were taken out on

the day of hatching, and measured and weighed

immediately. The SVL of newly hatched juveniles

ranged from 29.5 to 39.5 mm (33.8442.35, n=45),

the TaL from 31.0 to 47.0 mm (40.4343,52, n=45).

The TaL/SVL ratio is 1.00-1.37 (1.20+40.06, n=45).

In some clutches embryos underwent normal

development until reaching hatching time (egg

teeth in upper jaw completely developed (Rosler

2001)), but still deceased in the egg. Eggs

successfully hatching and those containing fully

developed but dead embryos were exposed to the

same incubation conditions. Therefore, reasons for

the death of some of the embryos are unknown. A

connection with the time of oviposition as well as

obvious malformations of the embryos as reason

for death can be excluded.

In part of the hatchlings, the TaL corresponded

with the SVL or was just slightly longer. This was the

case mainly in juveniles hatching too early. During

embryonic development of G. ernstkelleri, as in all

oviparous geckos, an early negative allometric tail

growth occurs (Noro et a/. 2009). After reaching a

certain embryonic SVL state this relation reverses,

resulting in a proportionate faster growth of the

TaL. A balanced ratio between SVL and TaL during

hatching is untypical for G. ernstkelleri. What causes

a premature hatching of juveniles could not be

determined. Characteristics for premature hatching

were, beside of the untypical body proportions,

remains of egg yolk within the egg shells, or egg

yolk clinging to the umbilical cord of the hatchlings.

No size differences between the juveniles hatched

after fully completed embryonic development in

the terraria, and those hatched in nature, were

recorded. Recorded hatchling sizes within the type

locality of G. ernstkelleri were: SVL 34.7-35.5 mm

and TaL 40.7-41.4 mm (N = 4 (Gaulke 2012)). The

smaller size range of the measurements compared

to those of the hatchlings bred in captivity is a

result of the lower number of individuals measured.

In the juveniles hatching with completely

absorbed egg yolk, the SVL values 38-45% (4143, N

= 30) of the SVL of the mother. Similar proportions

pp. 35-48

are known from G. gigante (42%), G. palmatus

Boulenger, 1907 (40%), and G. petricolus (47%),

while juveniles of the larger species G. siamensis

(SVL of adult 136.5 mm) are comparatively smaller

(35% of the maternal SVL).

The weight of juvenile G. ernstkelleri hatched

in captivity with completely absorbed egg yolk is

0.8917-1.1632 g (0.9975+0.0/764, N = 25), and

in nature 0.90-1.04 g (Gaulke 2012). Usually, the

juveniles hatched with completely absorbed yolk

sack. Those having not yet completely absorbed

the egg yolk, showed a significantly lower weight.

Hatchlings with torn off, only partly absorbed, or

yolk sack clinging to the umbilical cord weighed

0.6382-0.8018 g (0.7048+0.0553, N = 12). The

survival capability of these juveniles was low.

The weight of fully developed juveniles of G.

ernstkelleri amounts to 5.6-7.8% (6.3+0.6, N =

25) of the weight of the mother. Similar ratio data

are displayed by the Philippine species G. gigante

(6.0%), and G. mindorensis (4.9-5.2%), as well as

by the Vietnamese species G. palmatus (5.2%).

After hatching, we attempted to retrieve the

eggshells as completely as possible. Sometimes this

was impossible because the females discovered the

empty eggshells within short time, and devoured

them. The saved eggshells were dried for one day

by 60 °C, afterwards the thickness and weight of

the shells were determined. The eggshells of G.

ernstkelleri are 82-188 um thick (136428, n=32),

which amounts to 0.44-0.99% (0.80+40.13, n=18)

of the egg-length. Their weight is 0.1646-0.2552

g (0.2009+0.0247, n=18). This is equivalent to

about 15.8% of the weight of freshly deposited

eggs (n=1).

The reproductive effort conducted by a female

for the production of an egg-clutch, based on the

calculation of relative clutch mass (RCM), depends

mainly on clutch size. The energy expense to

produce a clutch with only one egg compared to the

normal sized clutch containing two eggs is about

half only. For the calculation of the relative clutch

mass two formulas were used:

RCM=cv/fbw x 100 [%]

(cv = clutch volume, fow = female body weight,

after Pianka & Pianka (1976))

RCM=fwem/(fwem+fbw) x 100 [%]

(fwem = fresh wet clutch mass, fow = female body

weight, after Hielen (1993))

02-Jun-21 21:46:57

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figure 6. Two clutches of Gekko ernstkelleri glued on Figure 7. Hatching of Gekko ernstkelleri in the terrarium

a glass panel. The upper clutch is older than the lower

one. To the right of the lower clutch is an egg mark of a

previously placed single egg.

~. 7 + +S!

& a

: 3 - ee oat =<

Figure 8. Nibble traces on the newly hatched eggs from Gekko ernstkelleri. Eggshells are valuable calcium

resources and are regularly eaten by the females.

—.

Book4.indd 46 02-Jun-21 21:47:00

Book4.indd 47

Roser, H. & GauLke, M.: On the biology of Ernstkeller’s cave gecko, Gekko ernstkelleri (Gquamata: Gekkonidae) ...

One quarter of the G. ernstkelleri egg-clutches

contained one egg. Those eggs had a slightly bigger

volume (1.30-1.61 cm’, mean 1.41+0.17, n=3)

compared to those from clutches containing two

eggs (1.03-1.87 cm’, mean 1.324+0.25, n=17).

Possibly, a female invests more from her own limited

body resources in the production of a single egg.

However, statistics do not support this hypothesis,

since differences in egg-volume between both

samples were insignificant (Mann-Whitney U Test,

p 0.8181). However, only few single-egg clutches

two eggs 19.95 ING7

18.03-21.08 ‘|

pp. 35-48

(n=3) were available.

Results regarding the reproductive effort of G.

ernstkelleri are presented in Table 2.

Calculated using the formula by Pianka &

Pianka (1976), G. ernstkelleri soends less effort in

egg production (RCM for two eggs 19.95%, one egg

10.20%) than G. gigante (RCM of two eggs 28.33%,

one egg 12.15%), but more than G. monarchus

(RCM of two eggs 13.50%, one egg 5.90%).

Table 2. Relative clutch mass of Gekko ernstkelleri.

| =n __| Formula according to

Pianka & Pianka (1976)

10.20 9.87-10.54 Pianka & Pianka (1976

ee eee

Conclusions

This paper gives comprehensive data on the

reproductive biology of an endemic Philippine gecko

species, resulting from extensive investigations

and observation in captivity during a time span of

several years, Supplemented by field investigations.

Analytical comparisons with other species of the

genus Gekko, as well as with other squamates of

the herpetofauna of Panay are planned.

Acknowledgements

We are grateful to Thomas Hofmann (Zittau,

Germany) for an animated exchange of information,

and the contribution of photos. Maren Gaulke

highly appreciates the support of the Department

of Environment and Natural Resources and the

Protected Areas and Wildlife Bureau (DENR and

PAWB, respectively, Quezon City, the Philippines),

given to the projects conducted by her as member

of the Philippine Endemic Species Conservation

Project (PESC). We thank Olivier S. G. Pauwels (Royal

Belgian Institute of Natural Sciences, Brussels) for

reviewing the manuscript and providing instructive

information.

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und Nachzucht von Gekko smithii (Gray, 1842). -

Sauria 9, No 3: 25-30.

Grossmann W. & Stein, V. 1988. Gekko petricolus Taylor,

1962 - Haltung und Zucht im Terrarium. - Sauria

10, No 3: 27-30.

Hielen B. 1993. Unterschiedliche

Fortpflanzungsstrategien bei Geckos der Gattung

Tarentola, Gray, 1825. - Salamandra 28, No 3/4:

179-194.

Hofmann T. 2007. Der Vietnam-Goldgecko. Vivaria

Verlag, Meckenheim: 63 pp.

Jono T., Inui Y. 2012. Secret calls from under the eaves:

acoustic behaviour of the Japanese house gecko,

Gekko japonicus. - Copeia 2012, No 1: 145-149.

Koch H. 1968. 3 Jahre Tokehzucht. - Aquarien Terrarien

15, No. 2: 67.

Kratochvil L., Frynta D. 2005. Egg shape and size

allometry in geckos (Squamata: Gekkota), lizards

with contrasting eggshell structure: why lay

spherical eggs? - Journal of Zoological Systematics

and Evolutionary Research 44: 217-222.

Kreuzer M., Grossmann W. 2003. Beobachtungen an

Gekko ulikovskii Darevski & Orlow, 1994 und Gekko

grossmanni Gunther, 1994 im Terrarium. — Sauria

25, No 3: 3-11.

Kreuzer M., Grossmann W., Hofmann T. 2013.

Terrarienbeobachtungen an Gekko badenii (Sauria:

Gekkonidae). - Sauria 35, No 3: 41-47.

Linkem C. W., Siler C. D. Diesmos A. C., Sy E., Brown

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reference to limb morphogenesis. - Develoomental

Dynamics 238: 100-109.

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Archipelago, Japan. - Journal of Herpetology 36,

No 3: 473-479.

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Taggeckos Phelsuma dubia (Boettger, 1881). PhD

thesis, Universitat Bochum, 311 pp. [unpublished]

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Erforschung der LebensauffSerungen der Niederen

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Received: 23.11.2020.

Accepted: 25.ix.2020.

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Book4.indd 49

SHEA, G. M. & ALLison, A.: A new species of Sohenomorphus (Squamata: Scincidae) from Mount Kaindi ...

pp. 49-60

A new species of Sphenomorphus (Squamata:

Scincidae) from Mount Kaindi, Morobe Province,

Papua New Guinea

urn:lsid:zoobank.org:pub:16286B38-O030A-4601-BB4B-6AED8E54C22E

GLenn M. Suea * 2? & ALLEN ALLISON °

1 - Corresponding author: Sydney School of Veterinary Science, BO1, University of Sydney, NSW

2006, Australia; glenn.shea@sydney.edu.au

2 — Australian Museum Research Institute, Australian Museum, 1 William Street, NSW 2010,

Sydney, Australia

3 - Bishop Museum, 1525 Bernice Street, Honolulu, 96817, Hawaii, U.S.A.

Abstract: Sohenomorphus wau sp. nov. is described from a single specimen from near Wau in Morobe Province,

Papua New Guinea. The new species is a member of the S. solomonis species group, but its affinities within that

group are unclear.

Key words: Reptilia, New Guinea, taxonomy, description.

Introduction

New Guinea has a diverse skink fauna, with

166 currently recognised described species,

distributed over 19 genera. The most species-rich

genus is Sohenomorphus Fitzinger, 1843, with 46

currently recognised species, over a quarter of all

New Guinean skinks. However, the species diversity

of Sohenomorphus is an underestimate, with many

new species to be described, and a number of

current synonyms to be resurrected.

Many of the New Guinea Sphenomorphus

are short-limbed semifossorial or leaf litter

inhabitants, constituting a single major lineage,

the Sphenomorphus solomonis species group,

characterised by the presence of a postsupraocular

scale (Greer & Shea 2004). Most members of

this group show a pattern of fragmentation of

the temporal and posterior supralabial scalation

that involves at a minimum the division of the

last Supralabial into an upper and a lower scale,

with many species showing addition division of

the penultimate supralabial, the primary temporal

and/or the lower secondary temporal. One species,

S. schultzei (Vogt, 19114), falls outside this pattern

in having division of the primary temporal into an

upper and lower scale, without any division of the

last two supralabials.

In this paper, we describe a second species of

Sphenomorphus with this peculiar pattern.

The new species is known from only a single

specimen from the upper slopes of Mt. Kaindi

near Wau in Morobe Province, collected in 1973.

While we would normally like to examine additional

specimens to confirm the diagnostic characters

prior to naming the species, the site is now

inaccessible due to closure of the Wau Ecology

Institute and landslides blocking the track to the

site, and despite large numbers of Sohenomorphus

specimens having been collected from this

and nearby sites in the period 1961-1988, no

additional specimens of the species have been

found. It is readily distinguished from all other

Sphenomorphus known from Morobe Province

by multiple character states, and hence we are

confident that the species is not just an aberrant

individual of an already described species.

Material and methods

Definitions of head scales largely follow Taylor

(1936), although we use the spelling supraciliaries

rather than superciliaries, Supralabials rather than

upper labials, and infralabials rather than lower

labials, in accordance with general usage.

In particular, we provide the _ following

definitions of the scales around the orbit.

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Supraciliaries are the small scales bordering the

lateral margin of the supraocular scales. The first

Supraciliary contacts both the first Supraocular

and the prefrontal. The last supraciliary extends

medially along the posterior border of the last

Supraocular, and overlaps the anterior margin of

the parietal and the uppermost postsubocular. In

the Sohenomorphus solomonis species group, the

last Supraciliary seems to have become enlarged,

extending medially to reach the frontoparietals, and

has become divided into an upper and lower scale,

the lower overlapping the upper. The upper scale

of the two has been termed the postsupraocular

by Greer & Shea (2004), as it is aligned with the

Supraocular series, but partially separated from

them by the lower element of the posterior most

supraciliary, and lies with its long axis at about

90° to the last supraocular. Presuboculars are the

series of angular scales contacting the supralabials,

from the first scale posterior to the posterior loreal

to the last scale contacting both the supralabial

below the centre of the eye and the one anterior

to that. Preoculars are the scales (usually singular)

that lie dorsal to the first presubocular, posterior

to the posterior loreal, and align with the anterior

end of both the upper and lower palpebral rows.

Some authors adopt a different nomenclature,

referring to two preoculars, and including the

first presubocular as the lower preocular, thereby

reducing the number of presuboculars by one.

Postsuboculars are the line of scales following the

anterior edge of the underlying jugal bone, from the

first scale contacting both the subocular supralabial

and the penultimate supralabial, posterodorsally to

the last scale reaching and overlapping the corner

of the parietal. Some authors, such as Greer &

Shea (2004) restrict the postsuboculars to that

part of the series contacting the penultimate

supralabial. The upper and lower postoculars are

the scales wedged between the last supraciliary,

the penultimate supraciliary, and the upper

postsuboculars, and are aligned with the posterior

extremity of the upper and lower palpebral rows

respectively. The evolutionary history and homology

of the upper postocular are complex. In many

lygosomines, and in the non-attenuate scincines

(Taylor’s 1936 “Eumeces”, now spread over several

genera), the upper postocular is a clearly distinct

scale partially wedging between the last two

supraciliaries, with the penultimate supraciliary

lying lateral to the last supraocular, and the last

supraciliary lying posterior to the last supraocular.

In some genera, such as Eugongylus Fitzinger,

1843, a basal member of the tribe Eugongylini, the

upper postocular variably wedges further between

the last two supraciliaries, often fully separating

them, and becoming a de facto penultimate

supraciliary as it pushes dorsally, though its lower

part is still aligned with the upper palpebral row.

In most species in the tribe Sphenomorphini

Welch, 1982, the upper postocular continues to

extend dorsally, such that it is indistinguishable

from the supraciliary series, and, like the last

sSupraciliary, extends medially along the posterior

margin of the last supraocular. However, in some

Sphenomorphins, such as the S. solomonis species

group, this enlarged scale seems to have divided

into a larger upper scale and a lower scale aligned

with the upper palpebral row, returning to the

former configuration of a penultimate supraciliary

dorsally and upper postocular ventrally. However,

the new penultimate supraciliary, extending along

the posterior margin of the last supraocular rather

than being lateral to it, is derived from the upper

postocular rather than being a true member of

the supraciliary row. The space occupied by the

penultimate supraciliary plus the upper postocular

in the S. solomonis species group corresponds

to the space occupied by the upper postocular in

other sohenomorphins, and the space occupied by

the postsupraocular plus last supraciliary, and their

relationship to other scales in the S. solomonis

group, iS in agreement with the relationships of

the last supraciliary of other sohenomorphins. As a

further complexity, Greer (1983) began to use the

new term pretemporals for the combination of the

last Supraciliary and uppermost postsubocular of

Taylor (1936). In this paper, we maintain Taylor’s

nomenclature for these scales.

Museum collection abbreviations follow Sabaj

(2016).

Results

Sphenomorphus wau sp. nov. (Figs 1-2)

http://zoobank.org/1C114F31-9EB5-4F86-

B9OF-9D99582A297B

Holotype 9: BPBM 14182, Mt. Kaindi, 1900m,

5km WNW Wau (7.337259°S 146.666090°E, AGD 66

datum), Morobe Province, Papua New Guinea, collected

A. Allison, 141 May 1973 (field tag AA 140).

Derivatio nominis: Toponymic. The species

name is a noun in apposition alluding to the type

locality (both the nearest town and the location of

the former Wau Ecology Institute), and suggestive

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SHEA, G. M. & ALLison, A.: A new species of Sohenomorphus (Squamata: Scincidae) from Mount Kaindi ...

of surprise, the first author’s initial thought when

he found the specimen among the unidentified

Sphenomorphus material at the Bishop Museum.

Diagnosis: Identifiable as a member of the tribe

Sphenomorphini by the medial pair of preanals

being enlarged and laterally overlapping the

adjacent preanals, upper secondary temporal

separated from first pair of nuchals by an upper

tertiary temporal, and with most of the supradigital

scales divided (Greer 1974; 1979; 1990). Assigned

to the Sohenomorphus solomonis species group of

Greer and Shea (2004) by lacking a windowed lower

eyelid, and possessing a postsupraocular scale.

Differs from all other members of the S. solomonis

species group in having the last two supralabials

undivided, the primary temporal divided, and the

first supralabial not fused to the nasal.

Description: Measurements: The mature female

holotype (follicles moderate-sized, oviducts

obliquely pleated) has snout-vent length (SVL)

61 mm; axilla-groin length, from posterior side of

forelimb to anterior side of hindlimb, with limbs

extended at right angles to the body wall, 33 mm

(54.1% of SVL); tail regenerated, original portion

63.5 mm, with regenerated portion included, 72.5

mm; forelimb length, from tip of claw of longest digit

to medial end of post-brachial crease, with limb

extended at a right angle to body wall, 12.5 mm

(20.5% of SVL); hindlimb length, from tip of longest

digit to medial end of postfemoral crease, with limb

extended at a right angle to body wall, 15.5 mm,

but missing terminal end of fourth (longest) digit

bilaterally; estimated complete hindlimb length

pp. 49-60

(based on relative proportions of toes being similar

to other Sohenomorphus) 17 mm (27.9% of SVL);

head length, from tip of snout to anterior margin of

ear, 11.7 mm (19.2% of SVL); head width at widest

point of head 8.9 mm (76.1% of head length); head

depth at deepest point over parietal table, and

including lower jaw, 6.5 mm (55.6% of head length).

Coloration in preservative (Fig. 1): Dorsal ground

colour yellow brown, head and most of body with a

few sparse dark brown small flecks. Posterior part

of body, beginning at about the anterior extent of

adpressed hind limbs, densely flecked with dark

brown, progressing on the tail dorsum to a dense

but fine dark reticulum, leaving only small spots of

the lighter ground colour visible. Laterally yellow

brown with a grey tinge, face with the ground colour

largely obscured by a fine dark brown reticulum;

upper and lower lips with dark margins to the

labial scales giving a coarsely barred pattern; body

dorsolaterally with a coarse dark brown reticulum

about two scale rows wide dorsolaterally, marking

a demarcation between dorsal and lateral patterns,

becoming a finer reticulum along the flanks. Tail

pattern laterally like tail dorsum. Limbs above with

a coarse dark brown reticulum, bolder on the hind

limbs than on the front limbs. Venter yellow-brown,

throat with dark brown streaks and spots, ceasing

at the level of the front limbs to leave a largely

immaculate body venter; just posterior to the vent,

dark spotting and streaking begins to redevelop,

rapidly becoming the major feature of most of

the tail venter. Parietal and visceral peritoneum

unpigmented.

Figure 1. Holotype of Sohenomorphus wau Sp. nov.

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Scalation: Rostral with a moderate medial lobe

extending posteriorly between nasals, and lateral

lobes extending posteriorly to a point below

the nostrils, in narrow contact with frontonasal.

Frontonasal laterally overlapped by nasals and

anterior loreal, posteriorly overlapping prefrontals;

prefrontals large, in broad contact; frontal

large, kite-shaped, laterally overlapped by first

Supraciliary, overlapping first two sSupraoculars;

a pair of obliquely oriented frontoparietals, in

moderate median contact, overlapped laterally

by last three supraoculars and postsupraocular.

Interparietal large, of similar size to frontal,

overlapped by frontoparietals and overlapping

parietals. Parietal eye spot absent. Parietals large,

overlapped by postsupraocular, lastsupraciliary, and

upper postsubocular, in median contact posterior

to interparietal. Supraoculars four, of similar

anteroposterior width, second tallest. Nasal with a

round central nostril, supranasals and postnasals

absent. Two loreals, each taller than wide, anterior

loreal overlapping frontonasal and_ prefrontal,

posterior loreal overlapping prefrontal, first

Supraciliary, preocular and anterior presubocular.

Supraciliaries ten left, nine right, in largely linear

contact with supraoculars, first extending medially

to contact frontal, separating prefrontal and first

supraocular; first three contacting first Supraocular,

penultimate large, oblique, extending medially to be

overlapped by fourth supraocular, last supraciliary

large, divided into upper (postsupraocular) and

lower scale. Postsupraocular medially overlaps

frontoparietal. Preocular single, separated from

prefrontal by contact of posterior loreal with

first Supraciliary, and followed by one oblique

scale wedged between anterior supraciliaries

and upper palpebrals. Presuboculars five (left)/

four (right), first two above second supralabial,

remainder interdigitating with supralabials two

to five. Suboculars one. Postsuboculars four

(left)/five (right), first interdigitating between

Supralabials five and six, uppermost overlapped

by last supraciliary and overlapping parietal, upper

secondary temporal and upper part of primary

temporal. Upper postocular single, elongate, below

penultimate supraciliary. Lower postocular divided

into an anterior and posterior scale, both overlapped

by upper postocular, and posterior postocular

interdigitating with upper postocular and lower part

of last supraciliary. Upper palpebrals in contact with

Supraciliaries 2-9 (L) and 2-8 (R). Lower eyelid

scaly; lower palpebral row of small squared scales

only slightly interdigitating with subpalpebrals.

Supralabials seven, first rectangular, longer than

second, second to fourth square, fifth below

subocular, sixth and seventh more pentagonal.

Postlabials two. Primary temporals two, lower

overlapping upper, and wedged between last two

Supralabials. Secondary temporals two, upper

elongate, bordering parietal, anteriorly separating

upper primary temporal from parietal, and

overlapping lower secondary temporal. Nuchals two

left/one right, anterior pair separated from upper

secondary temporals by one scale. Ear vertically

ovoid, lacking lobules along anterior margin. Mental

in broad contact with postmental. Infralabials five.

Postmental wider than long, in contact with only

first infralabial on each side. First pair of chin

Shields in broad median contact. Second pair of

chin shields separated and overlapped by a single

median scale. Third pair of chin shields divided

into a medial and lateral scale, medially separated

by three scales that overlap the more medial chin

shield. Midbody scales 34; paravertebral scales

barely broader than adjacent dorsal scales, 83 from

first scale posterior to parietal to last scale anterior

to level of anterior margin of hindlimbs; subdigital

lamellae paired, with lamellae of preaxial series

larger than postaxial series; subdigital lamellae

below fourth toe unknown, as fourth toe incomplete

on both limbs, but assuming a similar proportion of

digit lengths to other members of the S. solomonis

species group, an estimate of 11 lamellae below

this digit.

Osteology: Presacral vertebrae 26. Postsacral

vertebral series incomplete, but 30 to the point

of regeneration. Phalangeal formula of manus

2.3.4.4.3; phalangeal formula of pes 2.3.4.?.4.

Differential diagnosis: From S. schultzei (Vogt,

19114), the only other species in the S. solomonis

species group lacking division of the last supralabial

but with divided primary temporal scales, S. wau

sp. nov. differs in lacking the characteristic fusion

of first supralabial to nasal (Greer 1973, Greer &

Parker 1974). Sohenomorphus schultzei is also

a much smaller species, with maximum SVL only

50 mm, and very rarely more than 42.5 mm, and

has fewer midbody scales (22-28), paravertebral

scales (43-70), supralabials (six, with fourth

below centre of eye), the lower secondary temporal

overlapping the upper secondary temporal, and the

third pair of chin shields not divided.

In the key to the S. solomonis species group

provided by Greer & Shea (2004), S. wau sp. nov.

would key to S. nigriventris (de Rooij, 1915) in

having prefrontals in contact, supralabials seven

with the fifth below the eye, three supraciliaries

contacting the first Supraocular, presuboculars

02-Jun-21 21:47:02

from Mount Kaindi ...

)

Squamata: Scincidae

A.: Anew species of Sohenomorphus (

SHEA, G. M. & ALLISON,

pp. 49-60

le bar = 2 mm].

[sca

Figure 2. Head shields of Sohenomorphus wau Sp. nov.

Book4.indd 54

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

four (note that the definition of presuboculars

used in that paper is different to the definition

used here, so that our counts are one greater

than used by Greer), and infralabials contacting

the postmental one. Sohenomorphus nigriventris

iS a species complex, although the populations in

Papua New Guinea represent a single taxon. These

geographically nearest populations to S. wau sp.

nov. (though still separated by 485 km) differ from

it in having 28-32 midbody scales, last supralabial

divided, lower secondary temporal overlapping the

upper secondary temporal, and two suboculars.

They also have a distinctive colour pattern of

small white spots partially edged with dark brown,

giving a series of partial-ocelli that are the major

component of lateral and dorsal pattern.

Four other Sohenomorphus species are known

from the Wau area: S. darlingtoni (Loveridge,

1945), S. jobiensis (Meyer, 1874), S. neuhaussi

(Vogt, 1911b) and S. solomonis (Boulenger, 1887).

Of these, only S. darlingtoni and S. jobiensis have

been recorded from the type locality of S. wau sp.

nov. The local population of S. darlingtoni differs

from S. wau sp. nov. in a number of characters,

including smaller maximum SVL (52 mm), fewer

paravertebral scales (59-77), greater number of

Supraoculars (uSually six, with the first three in

contact with the frontal, occasionally the first two

only in contact with the frontal, or seven, with the

first three contacting the frontal) and supraciliaries

(10-14, modally 12), anterior loreal either excluded

from the supralabials by contact between nasal

and posterior loreal, or double (divided into a dorsal

and ventral scale), at least the last supralabial,

and often the last two, divided into upper and

lower scales, and the lower secondary temporal

overlapping the upper.

Sphenomorphus jobiensis is a much larger

species than S. wau sp. nov., with the minimum

mature size for local populations being SVL = 76 mm

for males and 79 mm for females. It also has many

more midbody scales (36-44), subdigital lamellae

(21-29), supralabials (8-10), and postsuboculars

(usually 6-8, rarely 5), as well as having a double

anterior loreal, the last two supralabials divided

and the lower secondary temporal overlapping

the upper secondary temporal, and the first two

pairs of chin shields in median contact. It is not a

member of the S. solomonis species group to which

S. wau sp. nov. belongs, lacking the diagnostic

postsupraocular scale of that species group.

Sphenomorphus neuhaussi is a member

of the S. pratti group, and like other members of

that complex is an elongate, short-limbed, small-

headed semifossorial species with a very pointed

snout. Its distinctive colour pattern of narrow dark

grey stripes ona light grey to pink/tan ground colour

is very different to that of S. wau sp. nov., and it

is a much larger species (adult SVL of the local

population 90 mm or more), with shorter forelimbs

(13.3-17.1% of SVL) and head (14.2-17.4% of SVL),

and greater number of midbody scales (36-38) and

paravertebral scales (90-111). It further differs in

lacking a postsupraocular, having prefrontal scales

separated (at nearest approach, barely touching in

a few individuals), anterior loreal long and low, only

2-3 presuboculars, the last supralabial divided,

and the second and third chin shields separated

from the infralabials by one (second chin shields)

or two (third chin shields) rows of oblique scales.

Sphenomorphus solomonis, like S. wau sp.

nov. a member of the S. solomonis group and

possessing a postsupraocular scale, and of similar

size to S. wau sp. nov. (local populations with adult

SVL 53.5-71.5 mm), differs in having a more

coarsely spotted lateral pattern, often with dark

dorsal spotting, fewer midbody scales (26-30)

and paravertebral scales (59-72), separated

prefrontals, 2-6 nuchals on each side, more

square loreals; 3 presuboculars, 2-4, usually 3,

Suboculars; usually single primary temporal but

last supralabial divided, and usually two pairs of

chin shields in median contact.

Distribution and habitat: Known only from a

single site on Mt. Kaindi on the Bulolo Watut Divide

at 1900 m (Figs 3-6). This site is along the old road

to Edie Creek from Wau. Landslips (Fig. 7) have

resulted in the old road to Edie Creek becoming

inaccessible, and the new road to Edie Creek,

constructed in the early 1970s, bypasses the site.

On the 1:100 000 Papua New Guinea topographic

map series (Sheet 8283 Wau, Series T6012 Edition

1-AAS, 1977), the site begins at DM638891 and

follows the old road along the side of the valley for

about one kilometre. The area immediately along

the road was mostly a mix of anthropogenous

grassland with scattered patches of second growth

(dominated by Saurauia sp.) and remnant mid-

montane Elaeocarpus forest. The area is just above

the Castanopsis zone (Gressitt & Nadkarni 1978).

Annual rainfall, based on limited measurements

from a nearby weather station, is around 2500

mm and the mean monthly temperature is around

16:5°C:

The only other similarly-sized Sohenomorphus

known from the type locality is S. darlingtoni. lt was

common under logs, pieces of wood and other

debris along the abandoned roadway and occurs in

02-Jun-21 21:47:02

SHEA, G. M. & ALLison, A.: A new species of Sohenomorphus (Squamata: Scincidae) from Mount Kaindi ...

pp. 49-60

4 Mount Hagen oY 7

Men ® lire A

PAPUA NEW 2.

GUINEA ge

PortMoresby

Figure 3. Distribution of Sohenomorphus wau sp. nov. (star).

Figure 4. Habitat at eastern end of the general collecting site for Sohenomorphus wau sp. nov., looking west,

at the junction of the old (to right) and new (to left) roads to Edie Creek. The site consists of both grasslands

and forested areas and extends along the old road for about 1 km.

Book4.indd 55 02-Jun-21 21:47:03

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figures 5-7. Sohenomorphus wau sp. nov. habitat.

5 - Habitat slightly further along the old road to Edie

Creek, showing forest typical of the area; 6 - Details of

dense understory at forested area in the general site for

Sphenomorphus wau sp. nov.; 7 - Landslip adjacent to

the site for Sohenomorphus wau sp. nov.

Book4.indd 56 02-Jun-21 21:47:05

Book4.indd 57

SHEA, G. M. & ALLison, A.: A new species of Sohenomorphus (Squamata: Scincidae) from Mount Kaindi ...

the area from around 1100 m to nearly 2200 m in

both primary and secondary forest.

Sphenomorphus wau sp. nov. appears to be

a mid-montane species and is likely a primary

forest inhabitant. The second author made

extensive collections from 1200-1800 m along

the Edie Creek Road and adjacent second growth

forests and on the south summit of Mt. Kaindi at

2362 m but obtained only the holotype. Although

he conducted relatively little field work in mid-

montane primary forests on Mt. Kaindi, he collected

fairly extensively in these forests on Mt. Missim,

across the Wau Valley to the east. Mt. Missim

shares many species of amphibians and reptiles

with Mt. Kaindi but Sohenomorphus wau sp. nov.

is either absent from there or exceedingly rare.

The Bishop Museum holds 284 Sohenomorphus

specimens from the vicinity of Wau, Mt. Kaindi

and Mt. Missim collected between 1961 and 1988

from a number of altitudes and habitats, but only

the single specimen of S. wau sp. nov. is among

them. Until additional records are obtained, we are

unable to determine whether the primary forest at

the type locality is core habitat for the species, and

it naturally occurs in low densities in that habitat,

or whether the low densities reflect the type locality

being on the periphery of the distribution for the

species.

Acknowledgements

We thank Bozena Jantulik (University of

Sydney, Australia) for preparing the final version of

figure 2, and Dane Trembath (Australian Museum,

Sydney, Australia) for figure 3. The first author also

thanks past and present collection managers at

the Bishop Museum, Honolulu (Carla Kishinami,

Pumehana Imada and Molly Hagemann) for the loan

of this specimen and others and assistance during

visits to that collection over many years, staff at

the Australian Museum, Sydney (Jodi Rowley, Allen

Greer, Ross Sadlier, Cecilie Beatson and Stephen

Mahony) for facilitating his work at that institution,

and researchers and collection managers past and

present at the following institutions for facilitating

examination of specimens: Museum of Comparative

Zoology, Harvard University, Cambridge (James

Hanken, José Rosado), Naturalis Biodiversity Centre,

Leiden (Marinus Hoogmoed, Esther Dondorp),

National Museum of Natural History, Washington

DC (George Zug, Addison Wynn), American Museum

of Natural History, New York (Darrel Frost, Linda

Ford, David Dickey, Lauren Vonnahme), Papua New

pp. 49-60

Guinea National Museum, Port Moresby (llaiah

Bigilale), University of Papua New Guinea Natural

History Collection, Port Moresby (Rose Singadan)

and South Australian Museum, Adelaide (Mark

Hutchinson, Carolyn Kovach). The second author

also thanks Reni and Thomaz of Wau (Papua New

Guinea) for their assistance with fieldwork at the

Wau Ecology Institute.

References

Boulenger G. A. 1887. Catalogue of the Lizards in

the British Museum (Natural History). Volume

lll. Lacertidae, Gerrhosauridae, Scincidae,

Anelytropidae, Dibamidae, Chamaeleontidae.

Trustees of the British Museum, London: xii + 575

pp, xl pls.

de Rooij N. 1915. The Reptiles of the Indo-Australian

Archipelago. |. Lacertilia, Chelonia, Emydosauria.

E. J. Brill, Leiden: xiv + 384 pp.

Greer A. E. 1973. Two new lygosomine skinks from New

Guinea with comments on the loss of the external

ear in lygosomines and observations on previously

described species. - Breviora 406: 1-25.

Greer A. E. 1974. The generic relationships of the

scincid lizard genus Leiolopisma and its relatives.

- Australian Journal of Zoology Supplementary

Series 31: 1-67.

Greer A. E. 1979. A phylogenetic subdivision of Australian

skinks. - Records of the Australian Museum 32 No

8: 339-371.

Greer A. E. 1983. A new species of Lerista from Groote

Eylandt and the Sir Edward Pellew Group in

northern Australia. - Journal of Herpetology 17, No

1: 48-53.

Greer A. E. 1990. Overlap pattern in the preanal scale

row: an important systematic character in skinks. -

Journal of Herpetology 24 No 3: 328-330.

Greer A. E., Parker F. 1974. The fasciatus group of

Sphenomorphus (Lacertilia: Scincidae): notes on

eight previously described species and description

of three new species. - Papua New Guinea

Scientific Society Proceedings [1973] 25: 31-61.

Greer A. E., Shea G. 2004. A new character within the

taxonomically difficult Sohenomorphus group of

lygosomine skinks, with a description of a new

species from New Guinea. —- Journal of Herpetology

38, No 1: 79-87.

Gressitt J. L., Nadkarni N. 1978. Guide to Mt. Kaindi.

Wau Ecology Institute Handbook No 5: 135 pp.

Loveridge A. 1945. New scincid lizards of the genera

Tropidophorus and Lygosoma from New Guinea. -

Proceedings of the Biological Society of Washington

58: 47-52.

Meyer A. B. 1874. Ubersicht Uber die von mir auf

Neu-Guinea und den Inseln Jobi, Mysore und

Mafoor im Jahre 1873 gesammelten Amphibien.

02-Jun-21 21:47:05

Book4.indd 58

TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

- Monatsbericht der Koniglich Akademie der

Wissenschaften zu Berlin 1874: 128-140.

Sabaj M. H. 2016. Standard symbolic codes for

institutional resource collections in herpetology

and _ ichthyology: an Online Reference. Version

6.5. American Society of Ichthyologists and

Herpetologists, Washington, D.C. Electronically

accessible at http://www.asih.org [accessed 1

October 2020].

Taylor E. H. 1936. A taxonomic study of the cosmopolitan

scincoid lizards of the genus Eumeces with an

account of the distribution and relationships of its

species. - University of Kansas Science Bulletin

Material studied:

Sphenomorphus darlingtoni (Loveridge, 1945)

n = 45) all from Morobe Province, Papua New

Guinea:

BPBM 2887, 3853, Wau 1200 m, 7.343297°S

146.712862°E; BPBM 3967, Wau, 1100 m,

7.343297°S 146.712862°E; BPBM 2890, Mt.

Missim, vic Wau, 7.218297°S 146.809899°E;

BPBM 4141, Mt. Missim, vic Wau, 1650 m,

7.275815°S 146.779891°E; BPBM 6182, Mt.

Kaindi, 2350 m, 7.355634°S 146.678053°E;

BPBM 8356-60, 19074-79, 29792-93, 4.5

km WNW Wau, Mt. Kaindi, 1750 m, 7.330616°S

146.682971°E; BPBM 8741, 8950, 8958, 9018,

9243, 19073, 28568, 30172, 5 km WNW Wau,

1800 m, 7.337259°S 146.666090°E; BPBM

8935, Edie Creek, vic Wau, 2150 m, 7.365242°S

146.664411°E; BPBM 19081, 19092-93,

23673-74, 24059, 24062, Mt. Missim, 1290

m, 7.282609°S 146.769022°E; BPBM 19082-

83, Mt. Missim, Camp 2, 1630 m, 7.274457°S

146.777174°E; BPBM 19084-85, Mt. Missim,

Camp 2, 1640 m, 7.274457°S 146.777174°E;

BPBM 19086-8/7, Mt. Missim, Camp 2, 1650

m, 7.274457°S 146.777174°E; BPBM 19088-

89, Mt. Missim, Camp 2, 1670 m, 7.274457°S

146.777174°E; BPBM 19090, Mt. Missim, Camp

2, 1680 m, 7.274457°S 146.777174°E; BPBM

19091, Mt. Missim, Camp 2, 1700 m, 7.274457°S

146.777174°E; BPBM 30141, Mt. Missim, vic Wau,

1350 m, 7.282609°S 146.769022°E.

Sphenomorphus jobiensis (Meyer, 1874)

n = 108, all from Morobe Province, Papua New

Guinea:

BPBM 8320, 8322, 8361, 8363, 10812, 14590,

14592-93, 14596-99, 14601-06, 14608-10,

14613-17, 14619-21, 14624, 19185, 19187-

—_—_

23, No 1: 1-643.

Vogt T. 1911a. Reptilien und Amphibien aus Neu-

Guinea. - Sitzungsberichte der Gesellschaft

Naturforschender Freunde zu Berlin 1911, No 9:

410-420.

Vogt T. 1911b. Reptilien und Amphibien aus Kaiser-

Wilhelmsland. - Sitzungsberichte der Gesellschaft

Naturforschender Freunde zu Berlin 1911, No 9:

420-432.

Received: 01.x.2020.

Accepted: 12.xi.2020.

Appendix 1. List of examined specimens.

88, 19191, 19193-94, 21476-77, 24737-43,

24745-59, 32644, 2.5 km NW Wau (Wau Ecology

Inst), 1230 m, 7.341486 °S 146.705616°E; BPBM

14591, 14594-95, 14600, 14622-23, 3 km

NW Wau, 1300 m, 7.324120°S 146.693650 °E;

BPBM 14612, 1.8 km W Wau (WEI House #1),

1360 m, 7.341466°S 146.705616°E; 19179, 5

km WNW Wau (Mt. Kaindi), 1800 m, 7.337259°S

146.666090°E; BPBM 19180-84, 24736,

32645-46, 1 km SW Wau, 1200 m, 7.349690°S

146.706460°E; 19189-90, Mt. Missim, vic Wau,

ca. 1350 m, 7.282609°S 146.769022°E; BPBM

19192, along Bulolo-Wau rd, nr Kalli Bridge, ca. 4

km NNW Wau aerodrome, ca. 950 m, 7.309783°S

146.719203 °E; BPBM 19195, Mt. Missim, 1470 m,

7.282609 °S 146.769022°E; BPBM 19196-20141, 2

km W Wau, 1150 m, 7.343300°S 146.694750 °E;

BPBM 19202, 21499, Wau Ecology Institute, 1200

m, 7.341486°S 146.705616°E; BPBM 20970,

20974, 20979, Mt. Missim, Lantana Creek, 1100-

1160 m, 7.278776°S146.749771° E; BPBM 20971,

20977, Mt. Missim, Lantana Creek, 1100-1130

m, 7.278776°S 146.749771°E; BPBM 20972, Mt.

Missim, Misery Creek, 1040-1060 m, 7.279952°S

146.741261°E; BPBM 20973, Mt. Missim,

Cold Water Creek, 1100-1150 m, 7.262681°S

146.743650°E; BPBM 20975, Mt. Missim, Lantana

Creek, 1100-1150 m, 7.278776°S 146.749771°E;

BPBM 20976, Mt. Missim, Misery Creek, 1025-

10305: mi; 4#.279952°S. -146:741261° E> “BPBM

20978, Mt. Missim, Cold Water Creek, 1150-1195

m, 7.262681°S 146.743650°E; BPBM 20980, Mt.

Missim, Misery Creek, 1040-1090 m, 7.279952°S

146.741261°E; BPBM 20981, Mt. Missim, Trap

32, 7.218297°S 146.809899°E; BPBM 21493-

94, Wau, 1200 m, 7.343297°S 146.712862°E;

BPBM 24721, Mt. Missim, Misery Creek, Trap 84,

1040-1060 m, /7.279952°S 146.741261°E;

BPBM 24732, 1km SW Wau, 1150 m, 7.349690°S

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Book4.indd 59

SHEA, G. M. & ALLison, A.: A new species of Sohenomorphus (Squamata: Scincidae) from Mount Kaindi ...

146.706460°E; BPBM 24733-35, 1 km SW Wau,

1250 m, 7.349690°S 146.706460°E.

Sphenomorphus neuhaussi (Vogt, 1911b)

n = 30, all from Morobe Province, Papua New

Guinea:

BPBM 8319, 2.5 km NW Wau, 1340 m,

7.341486°S 146.705616°E; BPBM 14649-52,

14654, 14656-58, 19095, 23668, 24728-29,

25549, 2.5 km NW Wau (Wau Ecology Inst), 1230

m, 7.341486°S 146.705616°E; BPBM 14653,

14655, 1.8 km W Wau (Wau #1 house), 1360 m,

7.341486°S 146.705616°E; BPBM 18977-81,

Wau, 1200 m, 7.343297 °, 146.712862°E; BPBM

19094, 1 km SW Wau, 1200 m, 7.349690°S

146.706460°E; BPBM 19096-1002, 2 km W

Wau, 1150m, 7.343300°S 146.694750°E; BPBM

24727, 1 km SW Wau, 1300 m, 7.349690°S

146.706460°E.

Sphenomorphus nigriventris (de Rooij, 1915)

n = 200:

AMS R30731-32, R30747, AMNH 107235-37,

MCZ R132568-83, PNGNM 24195-201, Imigabip,

A200 ft [1250 m], Western Province, PNG,

5.28°S 141.50°E; AMS R30755, R127837-66,

MCZ R132584-86, Tifalmin [1530 m], Sandaun

Province, PNG, 5.08°S 141.45°E; AMS R30758,

AMNH 107230, 107233, MCZ R132587-608,

PNGNM 24188-91, SAMA R11638, Wangbin,

4800 ft [1400 m], Western Province, PNG, 5.23°S

141.25°E; AMS R40788, AMNH 111742-43,

MCZ R139276-82, R142471-74, R142487,

PNGNM 22352A-B, USNM 204022, Gigabip,

5000 ft [1520 m], Western Province, PNG, 5.12°S

141.05°E; AMS R115515-16, R115518, Yapsiei,

200 m, Sandaun Province, PNG, 4.62°S 141.08 ° E;

AMS R119542-43, Miptigin, Telefomin Mtns

1700-2300 m, Sandaun Province, PNG, 5.17°S

141.58°E; AMS R122866, R122879, Sol River,

Telefomin area, 1460 m, Sandaun Province, PNG,

5.12°S 141.65°E; BPBM 34308, Kunida, Subutu

River area, 1810 m, Southern Highlands Province,

PNG, 5.650453°S 142.642773°E; BPBM

34309, Top House, 1910 m, Southern Highlands

Province, PNG, 5.65915°S 142.63538°E; BPBM

34315-36, 34318, unnamed locality, 1855 m,

Southern Highlands Province, PNG, 5.652061°S

142.643303°E; BPBM 34310-12, 34319-39,

34345-46, Semba, 1790 m, Southern Highlands

Province, PNG, 5.633963 °S 142.649765° E; BPBM

34313-14, 34340-44, Dickson’s House, 1777 m,

Southern Highlands Province, PNG, 5.64545°S

142.63904°E; PNGNM 22763, UPNG 4622,

Sh a -o —F. st

pp. 49-60

Tabubil, Western Province, PNG, 5.25°S 141.22°E;

PNGNM 24183-87, 24192-94, Olsobip patrol post,

Western Province, PNG, 5.38°S 141.50°E; RMNH.

RENA 30018-22, SAMA R6240-41, R6232A-C,

R6255-56, Busilmin, 1520 m, Sandaun Province,

PNG, 4.92°S 141.13°E; SAMA R6249, R6261,

Kawolabip, Western Province, PNG.

Sphenomorphus schultzei (Vogt, 1911a)

n= 133:

AMS R30763, AMNH 107226-27, 111744,

124041-43, MCZ R142339-40, PNGNM 24249-

50, Wangbin, 4800 ft [1400 m], Western Province,

PNG, 5.23°S 141.25°E; AMS R40786-87, AMNH

111733-35, MCZ R140675-80, R140682-95,

PNGNM 22362A-B, USNM 204024-25, Gigabip,

5000 ft [1520 m], Western Province, PNG, 5.12°S

141.05°E; AMS R118816-17, Namosado, 900

m, Southern Highlands Province, PNG, 6.25°S

142.78°E; AMS R127836, AMNH 107229, MCZ

120137-40, PNGNM 24248, Tifalmin [1530

m], Sandaun Province, PNG, 5.08°S 141.45°E;

AMNH 62375, 18 km [23 km] SW Bernhard Camp,

Idenberg River, 2150 m, Central Mamberamo

Regency, Indonesia, 3.52°S 139.08°E; AMNH

95594, Mt. Hunstein, 4000 ft [1200 m], East

Sepik Province, PNG, 4.50°S 142.65°E; AMNH

107228, MCZ R124044, PNGNM 24247, Imigabip,

4200 ft [1250 m], Western Province, PNG, 5.28°S

141.50°E; BPBM 2494, 2501, 2504, 2506, 2508,

2510, 2520, 2524-25, 3190, 3193, 3206, 3210,

3212-13, 3221-23, 3225-26, 3231-32, 3750,

Sibil Valley, Star Mountains, Pegunungan Bintang

Regency, Indonesia, 4.89°S 140.64°E; MCZ

R140696, Mt. Fubilan, 6000 ft [1850 m], Western

Province, PNG, 5.20°S 141.13°E; MCZ R141632-

34, Mt. Fubilan, 5000-6000 ft [1500-1850 ml],

Western Province, PNG, 5.23°S 141.12°E; PNGNM

23195, Telefomin [2050 m], Sanduan Province,

PNG, 5.13°S 141.63°E; PNGNM 24246, Olsobip

patrol post, Western Province, PNG, 5.38°S

141.50°E; RMNH.RENA 18198, Calap village, Bime

Valley, 1890 m, Pegunungan Bintang Regency,

Indonesia; RMNH.RENA 18199, Munggona, 1800

m, Eipomek Valley, Pegunungan Bintang Regency,

Indonesia, 4.45° 140.02°E; RMNH.RENA 29843-

49, 29853-54, 29858-73, Basiskamp [Base

Camp], Sibil Valley, Pegunungan Bintang Regency,

Indonesia, 1260 m, 4.90°S 140.62°E; RMNH.

RENA 29850-51, Sibil, Pegunungan Bintang

Regency, Indonesia, 4.90°S 140.62°E; RMNH.

RENA 29855-5/7, Ok Bon, Bivak 39 [Camp 39],

Pegunungan Bintang Regency, Indonesia, 1300 m,

4.88°S 140.8°; SAMA R11829, Gayah Lydnum,

oes

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Taman, New Guinea; SAMA R12710, Kowol River

[2 Kauwol River, Western Province, PNG]; UPNG

8640-41, Mt. Robinson, Western Province, PNG,

5.23°S 141.18°E; UPNG 5684-86, Tabubil,

Western Province, PNG, 5.25°S 141.22°E; ZMA

12078, Parana Valley, upper Swart Valley, between

Peak Doorman and Peak Trikora, Lanny Jaya

Regency, Indonesia, 3.92°S 138.50°E

Sphenomorphus solomonis (Boulenger, 1887)

n = 99, all from Morobe Province, Papua New

Guinea:

BPBM 2880, 3745, Wau, 1200 m, 7.343297°S

146.712862°E; BPBM 8858, Vickery Ck, 8 km N

Wau, 1100 m, 7.281701°S 146.710666°E; BPBM

9076, 9078, 9147, 9149, 9205, 9221, 9227, 9267,

9288-89, 9331, 14659, 14661-62, 14665-74,

17399-415, 19128, 32639, 32641, 2.5 km NW

Wau (Wau Ecology Inst), 1230 m, 7.341486°S

146.705616°E; BPBM 9326-27, 14112,

19130-69, 2 km W Wau, 1150 m, 7.343300°S

146.694750°E; BPBM 14660, 14663-64, 19129,

3kmW Wau, 1600 m, 7.343300 °S 146.685690 ° E;

BPBM 19125, 1 km SW Wau, 1400 m, 7.349690°S

146.706460°E; BPBM 19126, 1 km SW Wau,

1250 m, 7.349690°S 146.706460°E; BPBM

19127, 1 km SW Wau, 1200 m, 7.349690°S

146.706460°E; BPBM 19129, ca. 3 km NW Wau,

ca. 1400 m, 7.342911°S 146.694448°E; BPBM

20969, Mt. Missim, Lantana Creek, 1100-1150

m, 7.278776°S 146.7497 71° E; BPBM 25367, 3.5

km NW Wau, Mt. Kaindi, ca. 1450 m, 7.342911°S

146.694448°E.

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21:47:06

Book4.indd 61

BALLERIO, A.: First contribution to the knowledge of the Ceratocanthinae from the Solomon Islands ...

pp. 61-73

First contribution to the knowledge of the

Ceratocanthinae (Coleoptera: Scarabaeoidea:

Hybosoridae) from the Solomon Islands,

with descriptions of five new species

urn:lsid:zoobank.org: pub: /BCBFA28-A/7C 7-406 /-AB21-4140D 7/09BA10

ALBERTO BALLERIO

Viale Venezia 45, |-25123, Brescia, Italy; alberto.ballerio.bs@aballerio.it

Abstract: The subfamily Ceratocanthinae Martinez, 1968 (Coleoptera Scarabaeoidea Ceratocanthinae) is recorded

for the first time from the Solomon Islands, with the description of five new species of Perignamptus Harold,

1877: Perignamptus kerleyi sp. nov., P. capitatus sp. nov., P. longepilosus sp. nov., P. mundaensis sp. nov. and P.

rendovaensis sp. nov. Each species is described and illustrated and an identification key to all discussed taxa is

provided.

Key words: Taxonomy, New Georgia, Guadalcanal, Perignamptus.

Introduction

The fauna of Scarabaeoidea Latreille, 1802

(Coleoptera) of the Solomon Islands still remains

largely unknown. It is not surprising therefore that

no Ceratocanthinae Martinez, 1968 (Coleoptera

Scarabaeoidea Hybosoridae) have been recorded

from there until now (Ocampo & Ballerio 2006).

Since 1995 the author has gathered several

unpublished data on the Ceratocanthinae fauna

of the Solomon Islands, mainly coming from the

expeditions held in the years ’60 of last century by

Philip and Penelope Greenslade (e.g. Greenslade

& Greenslade (1970)), by the Bernice Bishop

Museum (Honolulu), again in the years ’60, and,

more recently, by the National Museum of Prague

and other Czech entomologists. The aim of this

first note is the description of five new species of

the genus Perignamptus Harold, 1877. Additional

species of Ceratocanthinae, belonging to other

genera, will be described or recorded in future

papers (in preparation).

Material and methods

Methods and terminological conventions follow

Ballerio and Grebennikov (2016) and references

therein quoted. The holotypes of the new species

here described are deposited in the collection

Kingdom). Label data are provided verbatim, with

a slash to separate labels. Author’s comments

are in square brackets, while depository collection

acronyms are in parenthesis unless otherwise

stated. Photographs of the habitus were taken with

a Canon EOS D5 MII with a macro objective MP 65

mm, while photographs of the male genitalia were

taken with a Canon EOS D5 MII connected by a

phototube to a Leica MZ 12.5 stereoscope. Multi-

layer images were then assembled using Helicon

Focus software and cleaned and unmasked using

a photo processing software.

Acronyms of scientific collections used in the text:

ABCB - Collection Alberto Ballerio, Brescia, Italy;

BMNH - The Natural History Museum (formerly British

Museum, Natural History), London, United

Kingdom;

NMPC - National Museum, Prague, Czech Republic.

Other abbreviations:

EL - maximum elytral length;

EW - maximum total elytral width;

HL - maximum head length;

HW - maximum head width;

L - length;

PL - maximum pronotal length at middle;

PW - maximum pronotal width at middle;

W - width.

02-Jun-21 21:47:06

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TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Results

Hybosoridae Erichson, 1847

Ceratocanthinae Martinez, 1968

Ceratocanthini Martinez, 1968

Perignamptus Harold, 1877

Perignamptus kerleyi sp. nov. (Figs 1 a, b, c, d, e,

f, g, h, i & 6)

Holotypus 4 deposited in the BMNH: Solomon IS.,

Guadalcanal, Mt. Austen, carrion trap / 24/8 1965,

19360, P. Greenslade/ Solomon lIs., Pres. P. J. M.

Greenslade, B. M. 1966.477/ Perignamptus kerleyi n.

sp. det. A. Ballerio 1999, Holotypus 3 / [dissected and

glued on a card. Male genitalia glued in DMHF resin on a

separate card, same pin].

Paratypes 5specimens: Solomon IS., Guadalcanal, Mt.

Austen / 7/3 1966, 22304, P. Greenslade/ Solomon Is.,

Pres. P. J. M. Greenslade, B. M. 1966.47 7/ Perignamptus

kerleyi n. sp. det. A. Ballerio 1999, Paratypus 2 /

(BMNH); / carrion trap / Solomon Islands: Guadalcanal,

Mt. Austen, 1936 / 24. VIIIL65 P Greenslade/ B. M.

1966.477/ Perignamptus kerleyi n. sp. det. A. Ballerio

1999, Paratypus 3 / (ABCB); / Solomon IS., Guadalcanal,

Mt. Austen / Feb. 1966, 1246 P. Greenslade /

Acanthocerus gen. + sp. incog. R. Madge det. 1967 /

Solomon Is., Pres. P. J. M. Greenslade, B. M. 1966.47 7/

Perignamptus kerleyi n. sp. det. A. Ballerio 1999,

Paratypus 2 / (BMNH); / Solomon IS., Guadalcanal, Mt.

Austen, 19/1I/1963/9954 P. Greenslade/ Perignamptus

kerleyi n. sp. det. A. Ballerio 1999, Paratypus 2 /

(BMNH); Solomon Islands, Guadalcanal ca. 4,5 km S

of Barana vill, forest nr., “Japanese camp & Moka river

09°30.3’S, 159°58.9’E, 275 m Jiri Hajek leg., 5.-6-

-XI1.2013 / Perignamptus kerleyi n. sp. det. A. Ballerio

2020, Paratypus <4 / (NMPC).

Derivatio nominis: Noun in the genitive case of

Latin. Patronymic, dedicated to Malcolm D. Kerley,

former collection manager at BMNH.

Diagnosis: Perignamptus kerleyi sp. nov. can

be easily differentiated from all other species

of Perignamptus by the following combination

of characters: a) metallic dark green color, b)

elytra impunctate (50x), c) pronotal bead on fore

margin complete and narrow, d) dorsal ocular area

present, e) genal canthus present but not fused

with occipital area of head, f) head surface with a

longitudinal shallow line bordering internally dorsal

ocular area, g) pronotum mostly impunctate with

only few shallow horseshoe-shaped punctures

basally and along sides.

Description of holotypus: HL= 0.80 mm; HW =

1.46 mm; PL = 1.30 mm; PW = 2.20 mm; EL = 2.16

mm; EW = 2.13 mm. Large-sized Perignamptus,

flightless, shiny, metallic green (under artificial

lighting the color looks metallic bronze), antennae,

tarsi and sternum reddish-brown. Dorsum

glabrous (50x). Whole dorsum covered by sparse

micropunctation (visible only with at least 30x

magnification). Head subpentagonal, wider than

long (W/L ratio= 1.37), clypeus subtriangular, with

sides smooth and gently curved and clypeal apex

obtuse with a very short apical acute projection;

clypeopleuron short, genae acute and slightly

protruding outwards, genal canthus present and

fused with occipital area of head, dorsal ocular

area large, interocular distance about seven times

the maximum width of dorsal ocular area. Ventral

ocular area medium sized, elongate, visible only

laterally, head dorsal surface smooth, apart from

micropunctation and apart from a_ longitudinal

Shallow line bordering internally dorsal ocular area,

genal canthus with some longitudinal irregular

Shallow lines, and a few transversal shallow lines

on fore margin. Antennae 10-segmented, scape

slightly clavate distad, pedicellus rounded and

abruptly bent forwards distally, flagellum made of

short articles distinctly wider than long, antennal

club three-segmented, uniformly hairy. Mouthparts.

Distal epipharynx bisinuate, longitudinally divided

by a strong anterior median process; median

brush and corypha absent; apical fringe made of

long, fine setae, absent in the middle. Mentum

ventrally flat, widely emarginated in the middle,

emargination regularly wide-U-shaped; labial palpi

(including palpiger) four-segmented, first segment

short and transverse, segment two short, segment

three longer and plumper than preceding two

together, segment four Ssubconical, about as long

as segment three, apically bearing some short

sensilla, all segments, apart from the last one,

fringed with long setae. Mandibles short, regularly

curved, apicalis with pointed apical tooth very short

and blunt, not protruding over mesal brush, lateral

sclerite of apicalis bearing a distinct large pore,

mesal brush wide and well developed. Pronotum

wider than long (W/L ratio = 1.62), base widely

rounded, fore angles not truncate, fore margin

bisinuate with a thin smooth bead, pronotal sides

and base with a thin bead; pronotal sculpturing as

follows: disc smooth apart from micropunctation,

sides and base with some shallow comma-shaped

punctures, sometimes fused together. Scutellar

Shield wider than long, sides distinctly notched

by elytral articular process, converging to form a

triangle with elongate acute apex and sides slightly

curved inward. Surface slightly depressed in the

middle, covered by irregular impressed comma-

Shaped punctures. Elytra convex, rounded (in

02-Jun-21 21:47:06

BALLeRIO, A.: First contribution to the knowledge of the Ceratocanthinae from the Solomon Islands ...

pp. 61-73

Figure 1. Perignamptus kerleyi sp. nov. paratypus from 4,5 km S of Barana vill. A - Extended, dorsal view; B -

Enrolled, ventral view; C - Enrolled, dorsal view; D - Enrolled, lateral view. Paratypus from Mt. Austen; E - Head

(white arrows on the right indicate inner longitudinal line, white arrow on the left indicates the genal canthus not

fused with the occipital area of head); F - Metatibia (black arrows indicate the apical corbel); G - Aedeagus; H -

other side of parameres; | - Ninth abdominal segment.

Des

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

dorsal view), slightly longer than wide (W/L ratio =

0.97), sutural stria absent, humeral callus absent,

whole surface glabrous and smooth (50x), apart

from micropunctation and the presence at sides

of two longitudinal shallow wide furrows, parallel to

inferior Sutural stria, the dorsal one being shorter

than the ventral one and occupying only distal third,

while ventral one occupying median and distal third.

Pseudoepipleure absent, marginal area narrow and

smooth, inferior sutural stria present and reaching

humeral area, striated articular area not visible,

elytral articular process developed, smooth and

shiny, separated from remaining elytral surface

by a deep narrow furrow. Protibiae ending with

two outer teeth, outer margin finely and irregularly

serrate. Meso- and metatibiae ending with two

apical spurs. Metatibiae with outer apex truncate

forming a small, punctured “apical corbel” (sensu

Ballerio & Grebennikov 2016). Male genitalia:

aedeagus with parameres feebly asymmetrical,

Short, about 1/3 the length of basal piece.

Sexual dimorphism and variability: Males

have the inner apical spur of mesotibiae bent

inwards at a right angle, while female mesotibiae

have both apical spurs straight. The type series

does not show strong variability, apart from the

development of larger pronotal punctation and the

lateral elytral longitudinal furrows, which in the two

female paratypes are very weak.

Remarks: Perignamptus kerleyi sp. nov. is a

typical Perignamptus, with a morphology similar

to P. loriae Gestro, 1898 and P. sharpi (Harold,

1877) (the latter being the type species of the

genus), both from New Guinea, and the only other

Perignamptus species known to have elytra smooth

or finely punctate. It differs from P. loriae by: a) the

presence of a longitudinal shallow line bordering

internally dorsal ocular area, absent in P. loriae, b)

the bead of pronotal base complete and narrow,

whereas in P. loriae it is wide and interrupted in

the middle, c) the punctation of pronotum, which,

besides micropunctation, is larger and limited

to base and sides, whereas in P. loriae is smaller

and uniformly distributed. It differs from P. sharpi

by: a) the presence of a longitudinal shallow line

bordering internally dorsal ocular area, whereas in

P. sharpi the line is replaced by an irregular row of

fine simple punctures, b) genal canthus not fused

with occipital area of head, whereas in P. sharpi the

genal canthus is fused with the occipital area of

head, c) the bead of pronotal base complete and

narrow, whereas in P. sharpi is wide and interrupted

in the middle, d) the punctation of pronotum, which,

besides micropunctation, is larger and present on

———

ty y

base and sides, whereas in P. sharpi is completely

absent. Perignamptus kerleyi sp. nov. also shows

a large metatibial apical corbel, which in all other

known species of Perignamptus is smaller or

absent.

Distribution and habitat. Known only from

Guadalcanal Island (Mt. Austen and surroundings

of Barana village to the west of Mount Austen). The

area is covered by lowland rainforest. The material

collected by the Greenslades was found through

bait trapping using carrion (although, based on our

knowledge of Ceratocanthinae biology, it is unlikely

the specimens were attracted by the carrion; it is

more likely they ended up in the trap by random

walking).

Perignamptus capitatus sp. nov. (Figs 2 a, b, c, d

& 6)

Holotypus < deposited in the BMNH: Solomon IS., New

Georgia, Wagina isl., 3/7 1966 (3218) P. Greenslade /

Solomon Is., Pres. P. J. M. Greenslade, B. M. 1966.477/

Perignamptus capitatus n. sp. det. A. Ballerio 1999,

Holotypus <@ / [dissected and glued on a card. Male

genitalia glued in DMHF resin on a separate card, same

pin].

Paratypus: Allotypus 19 deposited in the BMNH: same

data as holotypus (apart from identification label).

Derivatio nominis: Latin adjective meaning

“headed”, due to the unusual shape of the head

with the clypeal fore margin abruptly depressed.

Diagnosis. Perignamptus capitatus sp. nov. can

be easily differentiated from all other species

of Perignamptus by the following combination

of characters: a) clypeal fore margin abruptly

depressed with respect to remaining head surface,

b) dorsal ocular area absent, c) pronotal discal

punctation made of large deep simple punctures, d)

elytral punctation made of shallow large horseshoe-

Shaped punctures, e) inferior sutural stria of elytra

limited to distal third.

Description of holotypus: HL = 0.46 mm;

HW = 0.68 mm; PL = 0.73 mm; PW = 1.13 mm;

EL = 113 mm; EW = 1.06 mm. Small-sized

Perignamptus, flightless, shiny, brown, antennae,

tarsi and sternum reddish-brown. Dorsum setate

(45x). Head subpentagonal, wider than long (W/L

ratio = 1.41). Head: Clypeus subtriangular, with

sides smooth and gently curved and clypeal apex

broadly rounded. Fore margin of clypeus abruptly

depressed with respect to remaining head surface

all over its extension; clypeopleuron short, genae

rounded and slightly protruding outwards, genal

canthus indistinct, dorsal ocular area absent.

Head dorsal surface with vertex covered by sparse

02-Jun-21 21:47:07

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BALLERIO, A.: First contribution to the knowledge of the Ceratocanthinae from the Solomon Islands ...

pp. 61-73

Figure 2. Perignamptus capitatus sp. nov. A - Dorsal view of holotypus; B - Aedeagus; C - Spermatheca of allotypus;

D - Latero-dorsal view of holotypus.

large simple impressed punctures, frons almost

impunctate with only few simple large punctures,

clypeus with dense impressed horseshoe-shaped

punctures with opening directed inwards, fore

clypeal depression with some irregular simple

longitudinal lines, interpunctural distance shorter

or equal to punctural diameter; eyes small,

longitudinally elongate. Antennae 10-segmented,

scape slightly clavate distad, pedicel rounded and

abruptly bent forwards distally, flagellum made of

short articles distinctly wider than long, antennal

club three-segmented, uniformly hairy. Mouthparts.

Distal epipharynx bisinuate, longitudinally divided

by a strong anterior median process; median brush

and corypha absent; apical fringe made of long, fine

setae, absent in the middle. Mentum ventrally flat,

widely emarginated in the middle, emargination

regularly wide-U-shaped; labial palpi (includin

palpiger) four-segmented, first segment short and

transverse, segment two short, segment three

longer and plumper than preceding two together,

segment four Subconical, about as long as segment

three, apically bearing some short sensilla, all

segments, apart from the last one, fringed with

long setae. Pronotum wider than long (W/L

ratio = 1.85), roughly wide U-shaped, fore angles

truncate and slightly sinuate externally, fore margin

with a thick smooth bead, pronotal sides with a

thin bead, pronotal base without any visible bead

(dorsal view); pronotal sculpturing as follows: disc

slightly raised with respect to remaining surface,

extending backwards basally, whole surface, apart

from sides, uniformly covered by large impressed

simple punctures, with interpunctural distance

mostly equal to punctural diameter; sides covered

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

by shallower large horseshoe-shaped punctation

with interpunctural distance mostly shorter than

punctural diameter. Scutellar shield wider than

long, forming a triangle with elongate acute apex

and sides slightly curved inward. Surface slightly

depressed in the middle, covered by irregular

impressed punctures. Elytra_ strongly convex,

subovoidal (in dorsal view), slightly longer than

wide (W/L ratio = 0.92), sutural interstria absent,

humeral callus absent, whole surface covered by

dense relatively impressed punctation, punctures

near base small transverse comma-shaped with

a setigerous pore internally, remaining surface

covered by large horseshoe-shaped punctures

with opening directed backwards, and a setigerous

pore inside, interpunctural distance shorter than

punctural diameter (punctures almost touching

each other). Pseudoepipleure absent, marginal

area broad, expanded outwards, coarsely wrinkled,

inferior Sutural stria limited to distal third, striated

articular area not visible, elytral articular process

developed, smooth and shiny, separated from

remaining elytral surface by a deep narrow furrow.

Protibiae ending with two outer teeth, outer margin

almost smooth. Meso- and metatibiae ending with

two apical spurs. Metatibiae with outer apex not

truncate (apical corbel absent). Male genitalia:

aedeagus with parameres feebly asymmetrical,

Short, about 1/8 the length of basal piece.

Sexual dimorphism and variability: Males

have the inner apical spur of mesotibiae gently

bent inwards, while female mesotibiae have both

apical spurs straight. The allotypus has the head

vertex almost impunctate.

Remarks: The large sparse impressed pronotal

punctation and the abruptly depressed clypeal fore

margin make this species unmistakable among all

other known species of Perignamptus. The overall

morphology of P. capitatus sp. nov., in particular the

subsphaerical elytra and the lack of dorsal ocular

area, place this species in a group formed by P.

longepilosus sp. nov., P. rendovaensis sp. nov. and

P. mundaensis sp. nov. See under P. longepilosus

for a more detailed discussion.

Distribution and habitat: Known only from the

type locality (Wagina Island, also known as Wiggina

or Vaghena), a small island north of the New

Georgia Islands and next to Choiseul Island in the

Solomon Islands. No information is available about

the collecting circumstances. The type locality is

covered by rainforest.

Perignamptus longepilosus sp. nov. (Figs 3 a, b,

c & 6)

Holotypus deposited in the BMNH: Solomon

IS., Kolombangara. S. Kuzi 4,000’, 29.VIII.1965, P.

Greenslade, Roy. Soc. Exped. Brit. Mus., 1966-I / / litter

c& d / Perignamptus longepilosus n. sp. det. A. Ballerio

2000, Holotypus 3 / [dissected and glued on a card.

Male genitalia and mouthparts glued in DMHF resin on

two separate cards, same pin].

Derivatio nominis: Latin adjective meaning

“with long setation”, due to the presence of rows

and patches of setae longer than the other dorsal

setation and longer than the setation found in the

other species.

Diagnosis: Perignamptus longepilosus sp. nov.

can be easily differentiated from all other species

of Perignamptus by the following combination of

characters: a) dorsal ocular area absent, b) clypeal

disc with fine sparse punctation, c) head with a

short weak smooth tubercle on vertex, d) presence

of two patches of denser and longer setae in

correspondence with the two paradiscal pronotal

gibbosities, e) inferior sutural stria of elytra limited

to distal third, f) pronotal and elytral punctation

made of shallow dense punctures, g) elytra strongly

convex (Subsphaerical).

Description of holotypus: HL = 0.55 mm;

HW = 0.97 mm; PL = 0.98 mm; PW = 1.50 mm;

EL = 1.50 mm; EW = 1.45 mm. Small-sized

Perignamptus, flightless, shiny, brown, antennae,

tarsi and sternum reddish-brown. Dorsum setate

(45x). Head subpentagonal, wider than long (W/L

ratio= 1.41), clypeus subtriangular, with sides

smooth and gently curved and clypeal apex broadly

rounded; clypeopleuron short, genae rounded

and slightly protruding outwards, genal canthus

indistinct, dorsal ocular area absent. Vertex in the

middle with a smooth weak tubercle. Head dorsal

surface setate, setae short, vertex covered by some

Shallow transverse lines, frons slightly and gently

raised, frons and clypeal disc covered by sparse

simple punctures (interpunctural distance being

at least twice the punctural diameter, but often

more than twice), fore margin and sides with some

transverse shallow lines. Antennae 10-segmented,

scape slightly clavate distad, pedicellus rounded

and abruptly bent forwards distally, flagellum made

of short articles distinctly wider than long, antennal

club three-segmented, uniformly hairy. Mouthparts.

Distal epipharynx bisinuate, longitudinally divided

by a strong anterior median process; median

brush and corypha absent; apical fringe made of

long, fine setae, absent in the middle. Mentum

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BALLERIO, A.: First contribution to the knowledge of the Ceratocanthinae from the Solomon Islands ...

ventrally flat, widely emarginated in the middle,

emargination regularly wide-U-shaped; labial

palpi (including palpiger) four-segmented, first

segment short and transverse, segment two

short, segment three longer and plumper than

preceding two together, segment four subconical,

about as long as segment three, apically bearing

some short sensilla, all segments, apart from

the last one, fringed with long setae. Mandibles

short, regularly curved, apicalis with pointed apical

tooth very short and blunt, not protruding over

mesal brush, lateral sclerite of apicalis bearing

a distinct large pore, mesal brush wide and well

developed. Pronotum wider than long (W/L ratio

= 1.58), roughly wide U-shaped, fore angles gently

subtruncate, fore margin bisinuate with a thin

smooth bead, pronotal sides and base with a thin

bead; pronotal sculpturing as follows: disc slightly

raised with respect to remaining surface, slightly

“ Vy

4 "4

A Nee

Mei! ath a

lea

ih Pon

pp. 61-73

extending backwards basally, two gibbosities at

each side of disc (paradiscal gibbosities), whole

Surface covered by setae longer than the ones on

head, setae on paradiscal gibbosities, fore angles

and base denser and longer than the others, disc

covered by fine simple shallow punctures mixed to

irregular horseshoe-shaped shallow punctures with

opening directed outwards, sides of disc uniformly

covered by dense shallow large horseshoe-

shaped punctures, sometimes becoming ocellate,

horseshoe-shaped punctation with interpunctural

distance shorter than punctural diameter. Scutellar

shield wider than long, forming a triangle with

elongate acute apex and sides slightly curved

inwards. Surface slightly depressed in the middle,

covered by irregular impressed punctures. Elytra

strongly convex, almost rounded (in dorsal view),

about as long as wide (W/L ratio= 1.00), sutural

interstria absent, humeral callus absent, whole

Figure 3. Perignamptus longepilosus sp. nov. holotypus. A - Dorsal view; B - Latero-dorsal view; C - Aedeagus.

02-Jun-21 21:47:09

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TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Surface covered by dense shallow punctation,

some transverse lines near base then remaining

surface covered by dense shallow large horseshoe-

shaped punctures with opening mostly directed

backwards, each one bearing a simple puncture

in the middle, interpunctural distance shorter than

punctural diameter (punctures almost touching

each other), punctures as large as pronotal ones,

setation about as long as pronotal one, with two

irregular longitudinal rows of longer setae parallel

to elytral suture. Pseudoepipleure absent, marginal

area broad, expanded outwards, coarsely wrinkled,

inferior Sutural stria limited to distal third, striated

articular area not visible, elytral articular process

developed, smooth and shiny. Protibiae ending with

three outer teeth, outer margin almost smooth.

Metatibiae with outer apex not truncate (apical

corbel absent). Male genitalia: aedeagus with

parameres feebly asymmetrical, short, about 1/5

the length of basal piece. Aodominal ninth segment

with apodeme slightly longer than basal triangle.

Remarks: Perignamptus longepilosus sp. nov.

together with P. capitatus sp. nov., P- mundaensis

sp. nov., and P rendovaensis sp. nov. form a

distinctive group within the genus Perignamptus.

These four species are characterized by the following

combination of characters: a) dorsal ocular area

absent, b) strongly convex (Subsphaerical) elytra,

c) inferior sutural stria limited to distal third. This

group of species shares with the other species

of Perignamptus the presence of a basal pore on

mandibles, the expanded third labial palpomere,

the shape of epipharynx, the subclavate antennal

scape, and the feebly asymmetrical short

parameres. Despite the clear differences with

respect to the other known Perignamptus species

| prefer not to assign the above mentioned four

new species to a separate new genus, due to the

strong morphological variability present within

the Perignamptus group of genera, as defined by

Ballerio (2009, 2018). The genus Perignamptus,

apart from the five new species here described,

includes four other species: P. carinipennis Gestro,

1898, P. loriae Gestro, 1898, P. rossi Paulian, 1978

and P. sharpi (Harold, 1877), the type species of the

genus. The author is aware of several undescribed

species, mainly from New Guinea and, based on

the author’s unpublished data, it is also quite clear

that Macrophilharmostes major (Paulian, 1975)

is actually a true Perignamptus and that the two

species of Madrasostes described by Paulian from

localities east of the Weber Line, i. e., M. granulatum

(Paulian, 1975) and M. loebli Paulian, 1981, are

true Perignamptus too. It must be stressed that

the differentiation between the genus Madrasostes

Paulian, 1975 and the genus Perignamptus is quite

debatable at the state of the art and a careful re-

assessment of the two genera is needed.

Distribution and habitat. Known only from

the type locality. Kolombangara Island belongs to

the New Georgia Islands in the Western Province

of the Solomon Islands. No information is available

about the collecting circumstances apart from the

indication “litter”. The type locality is covered by

rainforest.

Perignamptus mundaensis sp. nov. (Fig 4 a, b, c,

d,e&6)

Holotypus 4 deposited in the BMNH: Solomon Islands,

New Georgia, Munda 22384, 5-ll-1966, P. Greenslade,

B. 1966 - | / Perignamptus mundaensis n. sp. det. A.

Ballerio 2000, Holotypus < /. [dissected and glued on a

card. Male genitalia glued in DMHF resin on a separate

card, same pin].

Paratypes 29: same data as holotypus (BMNH).

Derivatio nominis: Latin adjective meaning

“from Munda”, after the type locality.

Diagnosis: Perignamptus mundaensis sp. nov.

can be easily differentiated from all other species

of Perignamptus by the following combination of

characters: a) clypeal disc with irregular comma-

shaped shallow punctures and some horseshoe-

Shaped punctures mixed to few simple fine

punctures, b) head with a short weak smooth

tubercle on vertex, c) dorsal ocular area absent,

d) pronotal and elytral punctation made of shallow

dense horseshoe-shaped punctures mixed to

ocellate punctures (sides of pronotum with ocellate

punctures only), e) inferior sutural stria of elytra

limited to distal third, f) elytra strongly convex

(Subsphaerical).

Description of holotypus: HL = 0.64 mm; HW =

0.93 mm; PL = 0.97 mm; PW = 1.46 mm; EL= 1.55

mm; EW = 1.49 mm. Small-sized Perignamptus,

flightless, shiny, brown, antennae, tarsi and

sternum reddish-brown. Dorsum_ setate (45x).

Head subpentagonal, wider than long (W/L ratio=

1.53), clyoeus subtriangular, with sides smooth and

gently curved and clypeal apex broadly rounded;

clypeopleuron short, genae rounded and slightly

protruding outwards, genal canthus indistinct, dorsal

ocular area absent. Vertex with in the middle a short

longitudinal smooth weak tubercle; head dorsal

surface setate, setae short, vertex covered by some

Shallow transverse lines, frons slightly and gently

raised, frons and clypeal disc covered by a mixture

of shallow large horseshoe-shaped punctures and

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BALLERIO, A.: First contribution to the knowledge of the Ceratocanthinae from the Solomon Islands ...

comma-shaped punctures (interpunctural distance

being shorter than punctural diameter punctures

almost touching each other) mixed to few fine

simple punctures, fore margin and sides with some

transverse shallow lines. Antennae 10-segmented,

scape slightly clavate distad, pedicel rounded and

abruptly bent forwards distally, flagellum made of

short articles distinctly wider than long, antennal

club three-segmented, uniformly hairy. Pronotum

wider than long (W/L ratio= 1.75), roughly wide

U-shaped, fore angles strongly truncate, fore margin

bisinuate with a thin smooth bead, pronotal sides

and base with a thin bead; pronotal sculpturing

as follows: disc slightly raised with respect to

remaining surface, slightly extending backwards

basally, two gibbosities at each side of disc

(paradiscal gibbosities), whole surface covered by

setae longer than the ones on head, whole pronotal

Surface uniformly covered by dense shallow large

horseshoe-shaped punctures, often becoming

ocellate, sides of pronotum with dense large

pp. 61-73

ocellate punctures only, interpunctural distance

shorter than punctural diameter (punctures almost

touching each other). Scutellar shield wider than

long, forming a triangle with elongate acute apex

and sides slightly curved inward. Surface slightly

depressed in the middle, covered by irregular

impressed punctures. Elytra strongly convex,

subsphaerical (dorsal view), slightly longer than

wide (W/L ratio = 0.95), sutural interstria absent,

humeral callus absent, whole surface covered by

dense shallow punctation, elytral suture marked by

a dense fine simple punctation, remaining surface

covered by dense shallow large horseshoe-shaped

punctures with opening mostly directed backwards,

each one bearing a simple puncture in the middle,

interpunctural distance shorter than punctural

diameter (punctures almost touching each other),

punctures as large as pronotal ones, setation about

as long as pronotal one. Pseudoepipleure absent,

marginal area broad, expanded outwards, coarsely

wrinkled, inferior sutural stria limited to distal third,

Figure 4. Perignamptus mundaensis sp. nov. holotypus. A - Dorsal view; B - Latero-dorsal view; C - Detail of

pronotal sides; D - Spermatheca of allotypus; E - Aedeagus.

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Striated articular area not visible, elytral articular

process developed, smooth and shiny. Protibiae

ending with two outer teeth (a third one very small,

hardly visible), outer margin almost smooth. Meso-

and metatibiae ending with two apical spurs.

Metatibiae with outer apex not truncate (apical

corbel absent). Male genitalia: aedeagus with

parameres feebly asymmetrical, short, about 1/3

the length of basal piece.

Sexual dimorphism and variability: Males

have the inner apical spur of mesotibiae gently

bent inwards, while female mesotibiae have both

apical spurs straight.

Remarks: Very similar to P rendovaensis sp.

nov. from which differs by the punctation pattern

of pronotum and elytra (see Diagnosis). See

also under P. longepilosus sp. nov. for a detailed

discussion.

Distribution and habitat: Known only from the

type locality. Munda is a settlement located in the

island of New Georgia in the Western Province of

the Solomon Islands. No information is available

about the collecting circumstances. The type

locality is covered by rainforest.

Perignamptus rendovaensis sp. nov. (Figs 5 a, b, c,

d,e,f,g,h & 6)

Holotypus <& deposited in the BMNH: Solomon Islands,

New Georgia, Rendova Is., 15.16:IV:66, P. Greenslade,

Roy. Soc. Exped. B. M. 1966 - | / Perignamptus

rendovaensis n. sp. det. A. Ballerio 1999, Holotypus 3 /

[dissected and glued on a card. Male genitalia glued in

DMHF resin on a separate card, same pin].

Paratypes 7 specimens (23, 42, 1 sex unstated):

same data as holotypus (6 BMNH & 1 ABCB).

Derivatio nominis: Latin adjective meaning

“from Rendova”, after the type locality.

Diagnosis: Perignamptus rendovaensis sp. nov.

can be easily differentiated from all other species

of Perignamptus by the following combination

of characters: a) clypeal disc with dense fine

punctation mixed to irregular comma-shaped

shallow punctures and some horseshoe-shaped

punctures, b) dorsal ocular area absent, c) pronotum

with disc covered by simple fine punctures and

few long shallow irregular lines mixed to comma-

Shaped punctures, d) elytral punctation made

of shallow dense horseshoe-shaped punctures

becoming ocellate only on distal third, e) inferior

sutural stria limited to distal third, f) elytra strongly

convex (Subsphaerical).

Description of holotypus. HL=0.53 mm; HW =

0.82 mm; PL = 0.88 mm; PW = 1.30 mm; EL = 1.40

mm; EW = 1.33 mm. Small-sized Perignamptus,

——_

= ae ‘ j

“= a “ta

flightless, shiny, brown, antennae, tarsi and

sternum reddish-brown. Dorsum setate (45x).

Head subpentagonal, wider than long (W/L ratio=

1.50), clyoeus subtriangular, with sides smooth and

gently curved and clypeal apex broadly rounded;

clypeopleuron short, genae rounded and slightly

protruding outwards, genal canthus_ indistinct,

dorsal ocular area absent. Vertex with in the middle

a short longitudinal smooth weak tubercle; head

dorsal surface setate, setae short, vertex covered

by some shallow transverse lines, frons slightly and

gently raised, frons and clypeal disc covered by a

mixture of shallow small comma-shaped punctures,

oriented centrifugally and several shallow fine

simple punctures (interpunctural distance of

comma-shaped punctures being shorter than

punctural diameter), fore margin and sides with

some transverse shallow long lines. Antennae

10-segmented, scape slightly clavate distad, pedicel

rounded and bent forwards distally, flagellum made

of short articles distinctly wider than long, antennal

club three-segmented, uniformly hairy. Mouthparts.

Distal epipharynx bisinuate, longitudinally divided

by a strong anterior median process; median

brush and corypha absent; apical fringe made of

long, fine setae, absent in the middle. Mentum

ventrally flat, widely emarginated in the middle,

emargination regularly wide-U-shaped; labial palpi

(including palpiger) four-segmented, first segment

short and transverse, segment two short, segment

three longer and plumper than preceding two

together, segment four Ssubconical, about as long

as segment three, apically bearing some short

sensilla, all segments, apart from the last one,

fringed with long setae. Mandibles short, regularly

curved, apicalis with pointed apical tooth very short

and blunt, not protruding over mesal brush, lateral

sclerite of apicalis bearing a distinct large pore,

mesal brush wide and well developed. Pronotum

wider than long (W/L ratio = 1.69), roughly wide

U-shaped, fore angles strongly truncate, fore

margin feebly bisinuate with a thin smooth bead,

pronotal sides and base with a thin bead; pronotal

sculpturing as follows: disc slightly raised with

respect to remaining surface, slightly extending

backwards basally, two very weak gibbosities at

each side of disc (paradiscal gibbosities), pronotal

disc covered by simple fine punctures and few

long shallow irregular lines mixed to comma-

Shaped punctures, pronotal sides covered by large

horseshoe-shaped punctures mixed to few ocellate

punctures, interpunctural distance shorter than

punctural diameter (punctures almost touching

each other). Scutellar shield wider than long, with

02-Jun-21 21:47:10

BALLERIO, A.: First contribution to the knowledge of the Ceratocanthinae from the Solomon Islands ...

pp. 61-73

Figure 5. Perignamptus rendovaensis sp. nov., paratypus. A - Enrolled, ventral view; B - Enrolled, dorsal view;

C - Enrolled, lateral view; D - Detail of pronotal sides; E - Spermatheca; F - Aedeagus; G - Aedeagus; H - Ninth

abdominal segment.

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TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Perignamptus longepilosus sp. nov.

Perignamptus mundaensis sp. nov.

Perignamptus rendovaensis sp. nov.

Perignamptus capitatus sp. nov.

Perignamptus kerleyi sp. nov. ,

Figure 6. Distribution map showing the type localities of Perignamptus within the Solomon Islands archipelago.

elongate acute apex and sides slightly curved

inward. Surface slightly depressed in the middle,

covered by irregular impressed simple punctures.

Elytra strongly convex, subsphaerical (dorsal

view), almost as long as wide (W/L ratio= 1.04),

sutural interstria absent, humeral callus absent,

whole surface covered by dense punctation made

of large horseshoe-shaped punctures with opening

mostly directed backwards, each one bearing

a simple puncture in the middle, interpunctural

distance shorter than punctural diameter

(punctures almost touching each other), punctures

as large as pronotal ones, setation about as long

as pronotal one. Pseudoepipleure absent, marginal

area broad, expanded outwards, coarsely wrinkled,

inferior Sutural stria limited to distal third, striated

articular area not visible, elytral articular process

developed, smooth and shiny. Protibiae ending

with two outer teeth, outer margin almost smooth.

Meso- and metatibiae ending with two apical spurs.

Metatibiae with outer apex not truncate (apical

corbel absent). Male genitalia: aedeagus with

parameres almost symmetrical, short, about 1/3

the length of basal piece.

Sexual dimorphism and variability: Males

have the inner apical spur of mesotibiae gently

bent inwards, while female mesotibiae have both

apical spurs straight. Some paratypes have some

horseshoe-shaped punctures even on_ pronotal

disc.

Remarks: See above under P. mundaensis sp.

nov. and under P. longepilosus sp. nov. for a detailed

discussion.

Distribution and habitat: Known only from the

type locality. Rendova Island belongs to the New

Georgia islands in the Western Province of the

Solomon Islands. No information is available about

the collecting circumstances. The type locality is

covered by rainforest.

Key to the Solomon Islands Ceratocanthinae

An identification key is provided in order

to identify the species here described. The key

includes both the genus Pterorthochaetes Gestro,

1898, which occurs in the Solomons based on

the author's unpublished data, and the genus

Cyphopisthes Gestro, 1898 for which no data are

known but whose presence in the Solomon Islands

is possible, since the genus occurs in New Guinea

and in Australia (Queensland), with an old record

for New Caledonia too (Ballerio 2013).

02-Jun-21 21:47:11

Book4.indd 73

BALLERIO, A.: First contribution to the knowledge of the Ceratocanthinae from the Solomon Islands ...

1 Antennae 9-segmented, labrum not truncate,

somewhat depressed distally, elytra without a distinct

DSCUCEPIPICULON .......eseseerseeees Pterorthochaetes Gestro

- Antennae 10-segmented, labrum truncate or not

truncate, pseudoepipleure absent or present ............ 2

2 Labrum_ subtruncate, elytra with a_ distinct

pseudepipleure Cyphopisthes Gestro

- Labrum not truncate, somewhat depressed distally,

elytra regularly rounded (sagittal view) 3 Perignamptus

3 Larger species (>4 mm), elytra impunctate (50x), color

metallic green, dorsal ocular area PreSent ...........:222020e

CROP A bommmmnr sinies Simein LD, MARL cabin bit oh P. kerleyi sp. nov.

- Smaller species (<3 mm), elytra uniformly punctate

(50x), color dark brown to reddish-brown, dorsal ocular

area absent

4 Vertex of head without a smooth weak tubercle,

fore margin of clypeus abruptly depressed, pronotal

punctation large, impressed and Sparse .........ccccsssseeeeeees

P. capitatus sp. nov.

- Vertex of head with a weak smooth tubercle, fore

margin of clypeus on the same plane of remaining head

surface, pronotum with dense small shallow punctation

5 Head with clypeal disc with fine sparse punctation ....

d SadtniiahiaReeetobhs tee eaiuhe taut eR satay est P. longepilosus sp. nov.

— Head with clypeal disc densely punctate ................. 6

6 Pronotal sides with ocellate punctures, remaining

pronotal and elytral punctation made of shallow

dense horseshoe-shaped punctures mixed to ocellate

PUIG HITS Sys sete wena Paes ambeaetaae P. mundaensis sp. nov.

— Pronotum with disc covered by simple fine punctures

and few long shallow irregular lines mixed to comma-

shaped punctures, sides with horseshoe-shaped

punctures, elytral punctation made of shallow dense

horseshoe-shaped punctures becoming ocellate only

on distal third P. rendovaensis sp. nov.

Acknowledgements

The author wishes to thank Malcolm D.

Kerley, Maxwell V. L. Barclay (both BMNH) and Jiri

Hajek (NMPC) for the loan of relevant material.

Dmitry Telnov (BMNH) and Luca Bartolozzi (Museo

Zoologico “La Specola”, Florence, Italy) are thanked

for editorial assistance.

References

Ballerio A. 2009. Unusual morphology in a new genus

and species of Ceratocanthinae from New Guinea

(Coleoptera: Scarabaeoidea: Hybosoridae). - The

Coleopterists Bulletin 63, No 1: 44-53.

Ballerio A. 2013. Revision of the Australian

Ceratocanthinae (Coleoptera, Scarabaeoidea,

Hybosoridae). - ZooKeys 339: 67-91.

Ballerio A. 2018. Description of four new species of

pp. 61-73

Malaysian Madrasostes Paulian, 1975 forming

a distinctive group of species (Coleoptera,

Hybosoridae, Ceratocanthinae). - Entomologische

Blatter und Coleoptera 114: 89-100.

Ballerio A., Grebennikov V. V. 2016. Rolling into a ball:

phylogeny of the Ceratocanthinae (Coleoptera:

Hybosoridae) inferred from adult morphology

and single origin of an unique body enrollment

coaptation in terrestrial arthropods. - Arthropod

Systematics and Phylogeny 74, No 1: 23-52.

Gestro R. 1898. Sopra alcune forme di Acanthocerini.

- Annali del Museo Civico di Storia Naturale di

Genova 39: 450-498.

Greenslade P. J. M., Greenslade P. 1970 Studies on

the fauna of soil, litter and allied habitats in the

Solomon Islands: 209-212. In: Phillipson J. (ed.).

Methods of study in soil ecology. UNIPUB, New

York: 303 pp.

Harold E. von 1877. Enumération des lamellicornes

coprophages rapportés de |’Archipel Malais, de la

Nouvelle Guinée et de |l’Australie boréale par M.

M. J. Doria, O. Beccari et L. M. d’Albertis. - Annali

del Museo Civico di Storia Naturale di Genova 10:

38-110.

Ocampo F. C., Ballerio A. 2006. Phylogenetic analysis of

the scarab family Hybosoridae and monographic

revision of the New World subfamily Anaidinae

(Coleoptera: Scarabaeoidea). 4. Catalog of

the subfamilies Anaidinae, Ceratocanthinae,

Hybosorinae, Liparochrinae, and Pachyplectrinae

(Scarabaeoidea: Hybosoridae). - Bulletin of the

University of Nebraska State Museum 19: 178-

209.

Paulian R. 1975. Sur quelques Acanthoceridae

(Coleoptera) indo-australiens. - Annales

Historico-Naturales Musei Nationalis Hungarici

67: 151-154.

Paulian R. 1978. Révision des Ceratocanthidae [Col.

Scarabaeoidea] Il - Les espéces orientales

et australiennes. - Annales de la _ Société

Entomologique de France (N.S.) 14: 479-514.

Paulian R. 1981. Un nouveau Madrasostes des Illes

Bismarck (Coleoptera, Ceratocanthidae). - Revue

Suisse de Zoologie 88: 343-344.

Received: 31.viii.2020.

Accepted: 14.xii.2020.

(es:

02-Jun-21 21:47:12

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

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Book4.indd 75

Borponi, A.: New data on the Oriental Xantholinini. 49. A new genus, species and records from the Philippines ...

pp. 75-82

New data on the Oriental Xantholinini, 49.

A new genus, species and records from the Philippines

and Indonesia (Coleoptera: Staphylinidae)

316° contribution to the knowledge of the Staphylinidae

urn:lsid:zoobank.org:pub:95DOA41F-78C/7-4ADF-857E-EB9A9E 7D9AF5

ARNALDO BorDONI

Museo di Storia Naturale dell’Universita di Firenze, sezione di Zoologia “La Specola”, via Romana

17, |-50125, Florence, Italy; arnaldo.bordoni@fastwebnet.it

Abstract: Lalahonia gen. nov., with the type species Lalahonia pronotata sp. nov., are described and illustrated from

Mindanao, Philippines. The following new Lalahonia species are described and illustrated from the Philippines: L.

speciosa sp. nov. (Luzon), L. camarinensis sp. nov. (Luzon). Further Indonesian and Philippine staphylinid species

new to science described and illustrated, namely Erymus philippinus sp. nov. (Mindanao), Manilla camarinensis sp.

nov. (Luzon), M. candalagaensis sp. nov. (Mindanao) and M. leuseriana sp. nov. (Sumatra).

Key words: Lalahonia gen. nov., Erymus, Manilla, Luzon, Mindanao, Sumatra.

Introduction

Apart from the species described in the

revision of Xantholinini Erichson, 1839 staphylinids

(Bordoni, 2002) of the Oriental Region, the author

has dedicated two contributions to the Philippine

fauna (Bordoni, 2017, 2020) with descriptions of

new species. In the present paper, a particularly

interesting new genus Lalahonia gen. nov. is

described, with three new species referring to it.

Additionally, descriptions of new species of Erymus

Bordoni, 2002 and Manilla Bordoni, 1990 are

presented.

Material and methods

Specimens were studied using a Wild M5A

stereomicroscope and an Optika B-290 trinocular

microscope. The genitalia have been included in

Euparal to made permanent mounts. All taxa are

listed systematically. All taxa listed systematically.

Acronyms used in the text:

CAB - Collection Arnaldo Bordoni, Florence, Italy;

CJM —- Collection Jan Matéji¢ek, Hradec Kralové, Czech

Republic;

CAS - Collection Alexey Shavrin, Daugavpils University

and __ Technology,

Institute of Life Science

Coleopterological Research Center,

Latvia;

KUNM - Kansas University Natural History Museum,

Lawrence, U.S.A.;

SMNS - Staatliches Museum fur Naturkunde, Stuttgart,

Germany.

Daugavpils,

Results

Thyreocephalus rufus Cameron, 1941

Examined material: Mindanao, Gumitan, Davao del

Sur, loc. coll. 11.2019 (1 specimen CAB); Mindanao, 39

km E Malaybalay, Busdi, 1000 m, leg. Bolm 5-9.1V.1996

(6 specimens SMNS & 1 CAB); Mindanao 30 km NW

Maramag, Bagongsilang, 1700 m, leg. Bolm 13-17.

IV.1996 (2 specimens SMNS & 1 CAB).

Distribution: Endemic to the Philippines (Bordoni,

2002).

Thyreocephalus dustucheus Bordoni, 2002

Examined material: N Luzon, Cagayan, Sta Ana,

loc. coll. 1.2019 (1 specimen CAB); N Luzon, Cagayan,

Claveria, loc. coll. 11.2019 (1 specimen CAB); Mindanao,

Davao Prov., Mt Talomo (Mt Apo), 1300 m, 7°04’40.9’’N,

125°20'08.3’E, leg. A. Anichtchenko 22-28.1V.2019 (1

specimen CAS).

Distribution: Endemic to the Philippines (Bordoni,

2002).

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Thyreocephalus omaleus Bordoni, 2002

Examined material: Mindanao, Mt Apo, 1400 m, leg.

Bolm 18-19.IV.1996 (1 specimen SMNS).

Distribution: Endemic to the Philippines.

Thyreocephalus lumawigi Bordoni, 2020

Examined material: Mindanao, Mt Apo, Mt Apo N. P.,

6°59'15"N, 125°16'15"E, leg. |. Lumawig IV.2014 (1

specimen CJM).

Distribution: This species was described recently

(Bordoni, 2020) from Mindanao, Wao, Lanao del

Sur.

Lalahonia gen. nov. (Figs 1-20)

Type species: Lalahonia pronotata sp. nov.

Derivatio nominis: The generic epithet is the

name of the goddess of the volcanoes of the

Philippine mythology.

Description: The new genus is characterized

by the simultaneous presence of the following

characters: body of medium-large size (Figs 1-2);

maxillary palos with small first article, 2"? longer

than 3%, the last longer than the previous (Fig.

6); labial palos with small first article, the 2"

shorter than the 3™ (Fig. 7); labrum similar to that

of Thyreocephalus (Fig. 9 ); 3% antennomere very

longer than 2" (Fig. 8); frontal and ocular grooves

absent, at most indicated by very short carinae;

eyes small and moderately protruding; epistoma

protruding; mandibles with short and broad distal

groove, only one tooth and very evident prostheca

(Fig. 10); gular sutures juxtaposed for their entire

length (Fig. 11); pronotum with broad, areolate,

dense, lateral punctation; antesternal plate entire,

with a feeble trace of suture; upper epipleural line

of pronotum not joint with the lower line; sternum

short, with sub-rectilinear posterior margin and

short, acute median apophysis; epipleural portion of

elytra thickly dotted; anterior tarsi not dilated, with

1°, and 2™ articles visibly longer than the following

two. Sixth urite as in Figs 3-4. Male genital segment

composed of pleurae, tergite and sternite, without

peculiar characters; aedeagus large (Fig. 5), ovoid

elongate with basal bulb composed dorsally by two

membranes; median lobe long and narrow, laterally

sometime dilated in a denticulation; parameres

very long and narrow; inner sac ribbon-shaped,

folded on itself, covered with fine scales.

Differential diagnosis: The new genus is

similar in the general appearance to Arnaldolinus

Janak, 2018 from Madagascar from which it differs

in the shape of the maxillary and labial palps,

mandibles not falciform, the absence of frontal and

ocular grooves, structure of the antesternal plate

without sutures, different upper epipleural line, and

different aedeagus.

Distribution: Philippines.

Lalahonia pronotata sp. nov. (Figs 1-2, 6-15)

Holotype 6 CAB: Philippines, Mindanao, Bukidnon,

Dominorog, loc. coll. 1I.2019.

Derivatio nominis: The specific epithet is the

Latin “pronotatus” (punctate pronotum).

Description: Length of body 18 mm from anterior

margin of head to posterior margin of elytra: 9.5

mm. Body (Figs 1-2) black with bluish elytra;

antenna brown, legs brown with the external

surface yellowish. Head sub-rectangular, with

sub-rectilinear and sub-parallel sides and widely

rounded posterior angles. Surface with very dense,

deep, sometimes a little elongate punctures,

provided with long, yellowish setae on the sides.

Labrum as in Fig. 8. Pronotum longer and narrower

than head, with moderately oblique anterior

margins, rounded anterior angles, slightly sinuate

sides. Surface with numerous, not dense, deep,

lateral punctures, particularly on the anterior and

posterior portion of pronotum; a very large median

part of the surface without punctures and micro-

sculpture; Some micro-punctures distinct. Elytra

sub-rectangular, moderately dilated posteriad,

significantly longer and wider than pronotum, with

marked humeral angles. Surface with dense, deep

punctation, apart a lateral band; the folded portion

of elytra covered with finer punctures. Scutellum

with some punctures. Abdomen shiny, without

micro-sculpture, with deep punctation, arranged

in 5-6 series on each segment. Sixth urite as in

Figs 3-4. Aedeagus (Fig. 5) 3.7 mm long, very

voluminous, ovoid elongate, with narrow median

lobe; parameres short, inner sac in form of a ribbon,

folded several times on itself and covered with fine

scales.

Differential diagnosis: The larger size and

dense punctation on the lateral surfaces of the

pronotum and the structure of the aedeagus

differentiate this species from the two congeners

described below.

Distribution: The species is known only from the

type locality.

02-Jun-21 21:47:12

Borponi, A.: New data on the Oriental Xantholinini. 49. A new genus, species and records from the Philippines ...

pp. 75-82

al 2

Figures 1-2. Lalahonia pronotata sp. nov., holotype 3, habitus. 1 - Dorsal view; 2 - Ventral view

(images courtesy Silvio Cuoco).

Figures 3-5. Lalahonia pronotata sp. nov., holotype @. 3 - 6" tergite; 4 - 6" sternite;

5 - Aedeagus [not to scale] (images courtesy Silvio Cuoco).

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Lalahonia speciosa sp. nov. (Figs 1, 13-16)

Holotype 4 KUNM: Philippines, Camarines Sur, Naga

City, BGY, Panicuason, leg. D. General 11.1992.

Paratype 1° CAB: same data as holotype.

Derivatio nominis: The specific epithet is the

Latin “speciosus” (showy).

Description: Length of body 12.5 mm from

anterior margin of head to posterior margin of

elytra: 9.5 mm. Body (Fig. 16) black with reddish

brown dark elytra and abdomen; antennae

brown with the apical sixth segments covered

with pale pubescence; legs light brown. Head

subquadrangular, with evidently rounded sides and

widely rounded posterior angles. Eyes medium-

sized and almost flat. Surface of head totally

covered with deep, areolate, regular punctures,

alsmost arranged in series. Lateral and ventral

surface with similar punctation. Labrum as in Fig.

12. Pronotum massive, longer and, moderately

narrower than head, with slightly oblique anterior

margins, narrowly rounded anterior angles and

Sinuate sides. Surface with micro-punctation

and deep, very evident punctation, forward and

posteriad, apart a broad median portion. Elytra sub-

rectangular, slightly dilated posteriad, longer and

barely wider than pronotum, with slightly rounded

humeral angles. Surface with very fine, dense

punctation, arranged in numerous series, provided

with long setae; scutellum small, with some

superficial punctures. Abdomen similar to that of L.

camerinensis sp. nov. Tergite and sternite of male

genital segment as in Figs 13-14. Aedeagus (Fig.

15) 1.22 mm long, ovoid elongate, with rounded

median lobe; parameres narrow and long; inner sac

ribbon-like, folded one time on itself, covered with

fine scales.

Differential diagnosis: This species, like the

next one, differs from the type species of the genus

in its considerably smaller dimensions and the

Shape and structure of the aedeagus.

Distribution: The species is known only from the

type locality.

Lalahonia camarinensis sp. nov. (Figs 17-21)

Holotype 3 KUNM: Philippines, Camarines Sur, Naga

City, BGY, Panicuason, leg. D. General 1.1992

Paratype 1 CAB: same data as holotype.

Derivatio nominis: The specific epithet refers to

the Camarines region of Luzon Island.

Description: Length of body 11.5 mm from

anterior margin of head to posterior margin of

elytra: 6 mm. Body (Fig. 17) black with reddish

brown dark elytra; antennae and legs brown. Head

sub-quadrangular, with almost rectilinear sides and

widely rounded posterior angles. Eyes medium-

sized and slightly protruding. Surface of head with

dense, deep punctures, partially elongate between

thin carinae, sparse in the median portion and

absent on clypeus. Lateral portion with similar

punctation. Labrum as in fig. 18. Pronotum convex,

dilated anteriad, longer and narrower than head,

with moderately oblique anterior margins, rounded

anterior angles and slightly sinuate sides. Surface

with three large, deep punctures near the anterior

angles, a series of finer punctures along the lateral

margins. Elytra sub-rectangular, with sub-rectilinear

and sub-parallel sides, longer than pronotum, as

wide as pronotum in widest part, with rounded

humeral angles. Surface with deep, very dense

punctation, arranged in numerous series; scutellum

large with some punctures. Abdomen with traces of

transverse micro-striation and fine, evident, dense

punctation, arranged in numerous series on each

segment, except froma median stripe. Tergite and

sternite of male genital segment as in Figs 19-20.

Aedeagus (Fig. 21) 2.4 mm long, ovoid elongate,

with peculiar median lobe; parameres of peculiar

Shape; inner sac ribbon-like, folded two time on

itself, covered with fine scales.

Differential diagnosis: This species differs

from the previous one in the punctuation of the

pronotum, noticeably sparser, and in the shape and

structure of the aedeagus.

Distribution: The species is known only from the

type locality.

Erymus philippinus sp. nov. (Figs 22-24)

Holotype @ CAB: Philippines, SE Mindanao, Araibo,

Pantukan, Compostela Valley, Candalaga Mts, 900 m,

7°16'35.3''N, 126° 10'12.8’S, leg. A. Shavrin 4.V.2019.

Derivatio nominis: The specific epithet refers to

the Philippines.

Description: Length of body 4.4 mm from anterior

margin of head to posterior margin of elytra: 2.2

mm. Body shiny, reddish brown, with darker head.

Characterized by ovoidal head, dilated forward,

with very large and protruding eyes. Surface of

head with traces of transverse micro-striation

and few, scattered punctures. Pronotum dilated

anteriad, longer and narrower than head, with

oblique anterior margins, rounded anterior angles

and slightly sinuate sides. Surface with dorsal

series of five very fine and widely spaced punctures

and lateral series of two anterior punctures. Elytra

large, dilated posteriad, as long as, and significantly

02-Jun-21 21:47:12

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Borponi, A.: New data on the Oriental Xantholinini. 49. A new genus, species and records from the Philippines ...

Figures 6-16. Morphology of Lalahonia species. 6 - L. pronotata

sp. nov., maxillary palpus; 7 - ditto, labial palpus; 8 - ditto,

antennomeres 1-5; 9 - ditto, labrum; 10 - ditto, left mandible;

11 - ditto, gular suture; 12 - L. speciosa sp. nov., labrum; 13 -

ditto, 6" visible tergite; 14 - ditto, 6" visible sternite; 15 - ditto,

aedeagus [not to scale]; 16 - L. speciosa sp. nov., holotype 4

(image courtesy Silvio Cuoco).

wider than pronotum, with marked humeral

angles. Surface with fine punctation, arranged in

three series, one juxtasutural, one median and

one lateral; median series composed by more

numerous punctures. Abdomen with traces of

transverse micro-striation and a few, scattered

punctures, arranged in one series with long setae,

almost absent in the median pronotum portion of

the segments. Tergite and sternite of male genital

segment as in Figs 22-23. Aedeagus (Fig. 24) 0.48

mm long, sub-quadrangular, with narrow and long

parameres; inner sac apparently with few distal,

curved spinulae.

Differential diagnosis: The species differs from

the congeners in the ovoid head, dilated forward,

with very large and protruding eyes and in the

structure of the aedeagus with inner sacs provided

pp. 75-82

with spinules.

Distribution: The species is known only from the

type locality. This is the first Erymus Bordoni, 2002

representative in the Philippines.

Manilla camarinensis sp. nov. (Figs 25-30)

Holotype 3 KUNM: Philippines, Camarines Sur, Naga

City, BGY, Panicuason, D. General III.1992.

Derivatio nominis: Toponymic, referring to the

type locality.

Description: Length of body 5 mm from anterior

margin of head to posterior margin of elytra: 2.6

mm. Body shiny, reddish brown, with yellowish

fae,

02-Jun-21 21:47:13

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figures 18-24. Morphology of the Philippine Xantholininae.

18 - Lalahonia camarinensis sp. nov., labrum; 19 - ditto,

tergite of the male genital segment; 20 - ditto, sternite of

the same; 21 - ditto, aedeagus; 22 - Erymus philippinus

sp. nov., tergite of male genital segment; 23 - ditto, sternite

of the same; 24 - ditto, aedeagus [not to scale].

Legends: m - membranous part.

Figures 25-30. Morphology of the _ Philippine wm,

Xantholininae. 25 - Manilla camariniensis sp. nov., iy

tergite of male genital segment; 26 - ditto, sternite

of the same; 27 - ditto, aedeagus; 28 - Manilla

candalagaensis sp. nov., tergite of male genital

segment; 29 - ditto, sternite of the same; 30 - ditto, 28 29

aedeagus [not to scale].

Figure 17. Lalahonia camarinensis sp. nov.,

holotype 3 (image courtesy Silvio Cuoco).

Legends: m - membranous part.

Book4.indd 80 02-Jun-21 21:47:13

Book4.indd 81

Borponi, A.: New data on the Oriental Xantholinini. 49. A new genus, species and records from the Philippines ...

anterior margin of elytra; antennae and legs light

brown. Head ovoid with rounded lateral sides

from posterior margin of eyes to the neck. Eyes

large but almost flat. Surface of head with four

punctures on clypeus and few, fine punctures,

especially on sides. Pronotum narrow, longer and

narrower than head, with oblique anterior margins,

almost obsolete anterior angles and moderately

Sinuate sides. Surface with dorsal series of five

punctures and lateral series of three anterior

punctures. Elytra longer and wider than pronotum,

with almost obsolete humeral angles. Surface with

3-4 fine punctures, arranged in three series of 3-4

punctures, one juxtasutural, one median and one

lateral. Aodomen with two series of fine punctures on

each segment. Tergite and sternite of male genital

segment as in Figs 25-26; aedeagus (Fig. 27) 1.22

mm long, ovoid, very membranous, diaphanous,

with very small sub-triangular parameres; inner sac

apparently not visible.

Differential diagnosis: The species differs

from M. candalagaensis sp. nov. in the smaller

dimensions, the yellow anterior margin of the

elytra, the large but almost flat compound eyes, the

structure of the aedeagus with inner sac apparently

not visible.

Distribution: The species is known only from the

type locality.

Remark: The condition of the holotype is poor.

Manilla candalagaensis sp. nov. (Figs 28-30)

Holotype 3 CAS: Philippines, cS Mindanao, Araibo,

Pantukan, Compostela Valley, Candalaga Mts, 900 m,

7°16'35.3’N, 126°10'12.8’S, A. Shavrin 4.V.2019

Paratypes 84 CAS & 34 CAB: same data as holotype.

Derivatio nominis: The specific epithet refers to

the type locality.

Description: Length of body 6 mm from anterior

margin of head to posterior margin of elytra:

2.9 mm. Body shiny, reddish brown, with lighter

abdomen; antennae light brown; legs yellowish

with paler femora. Very similar to M. misamisae

Bordoni, 2002 from N and SW Mindanao, from

which differs for the following characters: body

stronger, wider and longer than head, with

significantly larger and more protruding eyes;

surface of head without micro-sculpture; pronotum

more massive; elytra subrectangular with red and

not yellowish humeral angles; surface with very

numerous punctures; different tergite and sternite

of male genital segment and different inner sac

of the aedeagus. Tergite and sternite of male

genital segment as in Figs 28-29; peculiar shape

pp. 75-82

of the sternite. Aedeagus (Fig. 30) 1.4 mm long,

ovoid, very membranous, diaphanous, with very

small sub-triangular parameres; inner sac barely

discernible, vaguely tubiform, partially covered with

Sparse scales.

Differential diagnosis: The new species is

closely related with M. misamisae Bordoni, 2002

from Mindanao from which differs in the large

and more protruding eyes, the head devoid of

microsculpture, the red humeral angles and the

inner sac covered of the aedeagus with sparse

scales.

Distribution: The species is known only from the

type locality.

Manilla leuseriana sp. nov. (Figs 31-33)

Holotype @ KUNMN: Sumatra, Aceh Gunung, Leuser

Nat. Park, Ketambe Res. Sta., 400 m, 3°41’0’N,

97° 39’0’E, leg. D. Darling 1-30.X!.1989.

Derivatio nominis: Toponymic, referring to the

type locality.

Description: Length of body 4.8 mm from anterior

margin of head to posterior margin of elytra: 2.6

mm. Body shiny, reddish brown, with darker head

and reddish elytra; antennae brown; legs yellowish.

Head ovoid, narrow posteriad, with rounded sides

from the posterior margin of eyes to the neck, Eyes

large, significantly protruding. Surface of head with

the usual four punctures on clypeus and few other

very fine punctures on sides. Pronotum narrow,

longer and narrower than head, with very oblique

anterior margins, obsolete anterior angles and

moderately emarginated sides. Surface with dorsal

series of five punctures and lateral series of three

anterior punctures. Elytra broad, sub-rectangular,

longer and wider than pronotum, with marked

humeral angles. Surface with three series of fine

and scattered punctures, one juxtasutural, one

median and one lateral. Aodomen with few, fine

punctures on the lateral portion of the segments.

Tergite and sternite of male genital segment as in

Figs 31-32. Aedeagus (Fig. 33) 0.7 mm long, ovoid,

very membranous, diaphanous, with very small

ovoid parameres; inner sac everted, tube-shaped,

with scattered, very small scales.

Differential diagnosis: The species differs

from the congeners of Malaysia above all in the

conformation of the aedeagus.

Distribution: The species is known only from the

type locality.

Remark: In Sumatra occur other two species of

Manilla Bordoni, 1990: M. bukittinggiana Bordoni,

2002 (westerncentral part of Sumatra: Bukittinggi)

02-Jun-21 21:47:13

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

and M. rougemonti Bordoni, 2002 (northern part of

Sumatra: Berastagi). The new species differs from

these taxa in shorter and paler body, and different

genitalia. The collecting locality of the new species

is the most northern record of Manilla in the islands.

Acknowledgements

| thank warmly the colleagues for providing

the discussed specimens for study, namely Arnaud

Faille (SMNS), Zack Falin (KUNM), Jan Matéjicek

(SJM) and Alexey Shavrin (CAS). | am indebted to

Silvio Cuoco (Livorno, Italy) for the colour images

used in the present manuscript.

Figures 31-33. Morphology of Manilla

lauseriana sp. nov. 31 - Tergite of male

genital segment; 32 - Sternite of the

same; 33 - Aedeagus [not to scale].

Legends: m - membranous part.

References

Bordoni A. 1990. Studi sugli Xantholinini della Regione

Orientale: il genere Manilla gen. n. (Coleoptera,

Staphylinidae). In: Berti N. (ed.) Miscellanées sur

les Staphylins. - Mémoires du Muséum national

d’Histoire naturelle, Paris (A): 171-180.

Bordoni A. 2002. Xantholinini della Regione Orientale

(Coleoptera: Staphylinidae). Classificazione,

filogenesi e revisione tassonomica. - Memorie del

Museo regionale di Scienze naturali (Torino) 33:

998 pp.

Bordoni A. 2017. New data on the Oriental Xantholinini.

39. New species and new records from the

Philippines (Coleoptera, Staphylinidae). - Bollettino

della Societa entomologica italiana 149, No 1: 33-

44.

Bordoni A. 2020. New data on the Oriental Xantholinini.

47. New species and new records from Mindanao,

Philippines (Coleoptera, Staphylinidae). - Bollettino

della Societa entomologica italiana 152, No 1: 3-8.

Cameron M. 1941. New species of Staphylinidae

(Coleoptera) from the Philippines. - Annals and

Magazine of Natural History 11, No 7: 430-447.

Received: 10.xi.2020.

Accepted: 10.xii.2020.

02-Jun-21 21:47:13

Book4.indd 83

Drumont, A., Komiya, Z. & WEIGEL, A.: Catyones marazziorum sp. nov. (Coleoptera: Cerambycidae) from Papua New Guinea

pp. 83-90

Catypnes marazziorum sp. nov. (Coleoptera:

Cerambycidae: Prioninae) from Papua New Guinea

urn:lsid:zoobank.org:pub:OEF88/24-4E91-4056-BFDE-99/7 DFE43D5A7

Axain Drumont +, ZiRo Komiya * & ANDREAS WEIGEL °

1 - O. D. Taxonomy and Phylogeny - Entomology, Royal Belgian Institute of Natural Sciences, Rue

Vautier 29, B-1000, Brussels, Belgium; alain.drumont@naturalsciences.be (ORCID: O0O00-OO000-

0001-9357-473xX)

2 - 3-2-12, Shimouma, Setagaya-ku, 154-0002, Tokyo, Japan

3 -Am Schlo8garten 6, D-O7381, Wernburg, Germany; rosalia-aw@gmx.de

Abstract: A new species of Catypnes Pascoe, 1864 is described and illustrated: Catyones marazzioruM sp. nov.

(type locality: Papua New Guinea, Morobe Province, Kapiro village, deposited in Royal Belgian Institute of Natural

Sciences, Brussels). The new species is compared with the single Catyones species previously known from Papua

New Guinea and with C. pirkli (Drumont, Komiya et Weigel, 2019), its closest congeneric taxon from Wallacea

(Sulawesi). A distribution map for all known species of Catypnes is given.

Key words: Aegosomatini, Catypnes, taxonomy, new species, Papua New Guinea.

Introduction

Quite recently the former subgenus Catypnes

Pascoe, 1864 of the genus Joxeutes Newman,

1840 was raised to genus level (Jin et al. 2020).

Catypnes currently comprises six species after the

recent descriptions of two species: one from New

Caledonia and one from Sulawesi in Indonesia

(Delahaye et a/. 2016; Drumont et a/. 2019).

Only one species of Catypnes, namely

C. dentifrons (Aurivillius, 1925), was hitherto

known from Papua New Guinea. It was described

on the basis of a unique male collected on the Mt.

Bolan located in south-eastern part of the country.

In continuation of our investigation on this peculiar

genus, we came across a series of specimens

collected from Papua New Guinea. After comparing

these specimens from Papua New Guinea with the

types of all taxa constituting the genus Catypones

(namely dentifrons - NHRS, negrosianus - ZSM,

macleayi - BMNH, pascoei - RBINS, pirkli - RBINS,

punctatissimus [= macleayi] - MNHN and salesnei

- NDC), we have come to the conclusion that these

belong to a new species that is described and

illustrated here.

Material and methods

All specimens examined in this study are

mounted and were observed under Leica stereo

microscope MZ-6. The dissected male genitalia

were fixed onto a white card together with the

specimen. Photographs were taken with a Leica

stereo microscope Z6 APOA, a Planapo-lens 1.0 was

used, and subsequently processed and measured

with the Leica Applications-software 4.0. The final

processing was made with Photoshop 7.0. Before

making photographs all parts of male genitalia

were immersed for a minimum of 24 hours in 80%

lactic acid and subsequently photographed in

99.5% glycerol.

Total body length is measured from the anterior

border of the clypeus to the apex of elytra and does

not include partially exposed abdominal segments.

Data from all specimen labels are reproduced

verbatim, without additions. If not stated, all labels

are printed. Authors’ comments are given in Square

brackets. If there are multiple labels on a specimen

these are separated by a slash. Each studied

specimen of the new species is provided with a

black-framed label on red paper as ‘HOLOTYPE’ or

‘PARATYPE’.

Pa.

wee

02-Jun-21 21:47:13

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TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

The material discussed below and types studied are

housed in the following collections:

ADC - Collection Alain Drumont, Brussels, Belgium;

AWC - Collection Andreas Weigel, Wernburg, Germany;

BMNH - Natural History Museum (formerly British

Museum, Natural History), London, United

Kingdom;

GVMC - Collection Giuseppe & Valentino Marazzi, Arese,

Italy;

JDC - Collection Jiti Dvoraéek, Krenov, Cesky Krumlov,

Czech Republic;

MNHN - Muséum national d’Histoire naturelle, Paris,

France;

NDC - Collection Norbert Delahaye, Plaisir, France;

NHRS - Swedish Museum of Natural History, Stockholm,

Sweden;

RBINS - Royal Belgian Institute of Natural Sciences,

Brussels, Belgium;

ZKC - Collection Ziro Komiya, Tokyo, Japan;

ZSM - Zoologische Staatssammlung Munchen, Munich,

Germany.

Other abbreviations:

HT - Holotype;

PT - Paratype.

Results

Catypnes marazziorum sp. nov. (Figs 1-9 & 13-

19)

Holotype ex ADC, will be deposited in RBINS: |.G.:

34.222) (Figs 1-3): Kapiro village, WAU, MOROBE,

Papouasie Nvelle Guinée, 21.X1Il.1998 / COLLECTION /

A. DRUMONT.

Paratypes 10¢ & 149: 19 (Allotype) AWC (Figs 4-6):

PAPUA NEUGUINEA, Bulolo, Morobe Prov., September

1999 / local collector; 14 AWC (Figs 7-9): Kapiro

village, Wau District, Morobe Prov., P. N. G. 12.IV.1997

/ I. dentifrons [underside of the label, handwritten] / ex

coll. G. et V. MARAZZI [handwritten] / COLLECTION / A.

DRUMONT; 1¢ ADC: Wau, P. N. G., VII.1990; 19 ADC:

Papua New Guinea, WAU, M.P., IIl-91 / Toxeutes dentifrons

9 Aurivillius, 1926 [handwritten], Det. A. DRUMONT,

2002; 14' ZKC: Papua New Guinea, Morobe Prov., Wau

dist., Kapiro vill., 5.1.1995, local collector leg.; 22 ZKC:

same locality, 14.1I1.1995, local collector leg.; 19 ex ZKC,

will be deposited in RBINS: I.G.: 34.223), same locality

and date, local collector leg.; 14 ZKC: same locality but

1.XII.1997, local collector leg.; 12 ZKC: same locality but

12.XI|.1997, local collector leg.; 29 ZKC: same locality

but III.1998, local collector leg.; 14 & 19 ADC: same

locality but 26.III.2004, local collector leg.; 19 ZKC:

same locality but 6.VIII.2007, local collector leg.; 13)

ZKC: same locality but 13.IV.2008, local collector leg.;

19 ADC: Wau Valley, 1200 M., Morobe prov., P.N.G.,

23.X1.1995; 24 ADC: Wau Valley, Morobe prov., P.N.G.,

15.11.1990; 14 & 12 GVMC: P.N.G., Morobe prov., Aseki,

1.1998; 12 GVMC: P.N.G., Morobe prov., Wau, Mt.

Kandi, 2.1I1.2005; 19 JDC: Papua New Guinea, Morobe

Prov., Wau dist., Kapiro vill., 13.11.2006, leg. Hudson;

13 ZKC: Papua New Guinea, Oro Prov., Timbek vill., 18.

Il.1996, local collector leg.

Derivatio nominis: Patronymic. We dedicate this

species jointly to Giuseppe and Valentino Marazzi,

two entomologists passionate about the worldwide

Cerambycidae, especially Prioninae and who

participate in our researches since more than 20

years.

Measurements: Holotype 3: 31.0 mm; allotype

2: 29.0 mm; PT 3: 27.5 (min.) - 33.5 (max.) mm, @:

31.0 (min.) - 37.5 (max.) mm.

Description: Habitus, male. Slightly elongated

and subparallel; body dark-reddish brown except

on the elytra which are lighter (especially apical

third of the elytra); underside reddish-brown. Head

slightly broader than long, upper side with dense

and strong, deep punctures, diameter of punctures

mostly much larger than their distances, punctures

also partly touching; wrinkled on the sides up to

the underside, partly with longitudinal furrows;

underside in the middle broadly smooth, not

punctured, underside between the eye lobes cross

furrowed and with erect yellowish hairs; genae

angularly protruding, almost rectangular; clypeus

wrinkled, in front with long, errect yellowish hairs;

mandibles dark brown, with blackish borders,

outer margin strongly curved, tip bidentate,

lower tooth shorter, outside covered with coarse

deep punctures of different sizes and sparsely

protruding yellowish hairs, inner margin behind

tip almost parallel sided, then truncated before

base; clypeus broadly deepened and wrinkled

just inside anterior margin; eyes kidney-shaped,

not edged, about twice as long as broad; distance

between eyes on front about 2.5 times the width of

the upper lobes in male. Antennae 11-segmented,

light brown, 1-3 segments dark brown; scape short,

length/apex width: 1.8; 3% antennal segment

longest, about 1.3 times as long as scape and

reaching anterior tooth of pronotum; 1-8 segments

covered with coarse punctures, these become

finer towards end and distances between become

larger towards apex; 9-11 segments chagrined, not

punctured and dull, apex of the 7-10 segments

angularly widened at the outside; apex of 11"

segment rounded; underside of segments 5-10 flat

and broadly furrowed, 3 and 4" furrowed only in

about apical half, 5-9 almost completely furrowed,

02-Jun-21 21:47:13

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Drumont, A., Komiya, Z. & WEIGEL, A.: Catyones marazziorum sp. nov. (Coleoptera: Cerambycidae) from Papua New Guinea

broad and flat, 6" segment weakly furrowed near

apical part only. Pronotum rectangular in shape,

median-length/base width = 0.62; anterior border

Straight with dense yellowish hairs, posterior

border slightly wavy, middle part (nearly 3/5) also

with short yellowish hairs, otherwise without hairs;

pronotal hind corners obtuse, at middle tooth as

wide as humeral angles; front tooth immediately

after anterior border of pronotum, acute-angled;

pronotal surface smooth shiny, punctures irregular,

deep, diameters of punctures mostly smaller than

their distances, small areas also without punctures;

prosternum somewhat wrinkled; borders of coxae

dark brown; prosternal process shiny, parallel-sided

between coxae, widened backward, and protruding

by about one third, broadly truncated at posterior

border, almost at same height as coxae. Venter:

Mesosternal process lower than coxa, triangular,

truncate at end; mesoventrite shiny, very finely

punctured; long protruding yellowish hairs on

meso- and metaventrite, epimeron and episternum,

the latter finely wrinkled; metepisternum about

four times longer than wide, less tapering distally;

abdomen shiny, smooth, glabrous, with very fine,

hardly visible punctures; last abdominal ventrite

slightly concave at the posterior border and with

yellowish, erect hairs, which become _ longer

towards tip and at sides. Elytra: about twice as

long as wide, length/width = 2.1; almost parallel

sided, shiny, the fine engraved punctures become

finer towards apex; finely wrinkled or rugulose near

apex; distances between the punctures variable,

from almost touching each other to larger than

their diameter; subscutellar area with rather dense

and coarse punctures; elytra also with three very

feeble costae; scutellum large, almost square,

slightly rounded apically, smooth and glossy, with

coarse punctures; elytral apex broadly rounded,

sutural edge extended into a pointed tooth; behind

the humeral region with a narrow lateral margin

which disappears before apex; epipleura narrow,

only under the shoulders very broad, chagrined,

disappearing before apex. Legs: Femora smooth

and shiny, very finely punctured, only slightly

thickened in middle; tibiae almost parallel sided,

widened towards apex, with coarse punctures, the

distances between them much larger than their

diameter, inner sides with oblique yellowish hairs;

tarsi with claw article the longest, 4 very small,

hardly visible, all joints with yellowish hairs above,

37 joint very deeply lobed, ratio of tarsal joints

1-5: 0.6 / 0.4 / 0.4 / 0.08 / 1.0. Male genitalia

(Figs 13-18): median lobe with non-inverted

endophallus (Fig. 13), 6.2 mm long, 1.0 mm wide,

pp. 83-90

nearly straight in lateral view, median struts about

2/5 length of median lobe, median lobe curved in

lateral view; dorsal plate almost parallel-sided with

apex broadly rounded triangularly, shorter than

ventral plate; ventral plate almost parallel-sided,

strongly narrowed before apex and extended into

parallel-sided spine; endophallus very long, not

distinctly subdivided into phallomeres (according

to Yamasako & Ohbayashi 2011), inside median

lobe (basal phallomere) with a pair of characteristic

crescent-shaped sclerites (Fig. 14), a band-shaped

folded structure inside first part of endophallus

(more than 6 mm long); tegmen (Figs 16-17) 5

mm long, about 1 mm wide at base of parameres,

parameres almost parallel-sided, slightly narrowed

to apex, and tight together at the base, closer

than the width of a paramere, width of paramere

before apex 0.25 mm; apex rounded and with long

yellowish hairs at anterior sixth, obtuse angled in

lateral view; base of the parameres with a small

wide tooth inside (Fig. 17); 8 tergite (Fig. 18)

2.5 mm long, about 2.8 mm wide, sides rounded

and slightly concave before apex; anterior margin

concave; apical 3/5 with long yellowish hairs,

longer and more dense around the anterior edges.

Sexual dimorphism: Female (Figs 4-6) with

antennae shorter (only reaching to the beginning

of the abdomen) and finer; scape less strongly

thickened; shorter mandibles; wider eyes, thus the

distance between eyes at the frons smaller, about

1.5 of the eye width; cheeks less strongly angularly

protruding; femora less thickened; elytral length-

width ratio: ¢ 2,1-2,2, 9 2,3-2,4 (4 specimens

measured). In a female (PT from Wau) about 123

eggs were found (see fig. 19), size of eggs about

2.5 mm long and 1.2 mm wide, in average.

Differential diagnosis: We have examined

holotype (male) of Catyones dentifrons (Figs 10-

12), the only other species present in Papua New

Guinea, preserved in NHRS, from which males of

C. marazziorum sp. nov. (Figs 1-3 & 7-9) differ

by the following morphological characters: width

of the head smaller than pronotum (as wide as

pronotum in C. dentifrons), median lateral tooth of

pronotum triangular (stretched in a tapering spine

in C. dentifrons), 1%t antennal segment elongated

(short and robust in C. dentifrons), 2" elytral costa

weak (distinct in C. dentifrons), apical tooth of the

sutural angle acute and distinct (weak and short in

C. dentifrons), femora thin and elongated (large and

robust in C. dentifrons). C. marazziorumM sp. nov.

is externally similar to C. pirkli (Drumont, Komiya

& Weigel, 2019) from Sulawesi in Indonesia, from

which the new species can be mainly distinguished

Bee

02-Jun-21 21:47:13

TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

5mm

Figures 1-3. Catypnes marazziorum sp. nov. 1-3 - Holotype @ (RBINS), body length 31 mm. 1 - Habitus, dorsal

view; 2 - ditto, ventral view; 3 - ditto, lateral view; 4-6 - Allotype 2 (AWC), body length 29 mm. 4 - Habitus, dorsal

Book4.indd 86

view; 5 - ditto, ventral view; 6 — ditto, lateral view.

02-Jun-21 21:47:14

Drumont, A., Komiya, Z. & WEIGEL, A.: Catyones marazziorum sp. nov. (Coleoptera: Cerambycidae) from Papua New Guinea

pp. 83-90

Figures 7-12. Papuan Catypnes species. 7-9 - C. marazziorum sp. nov., paratype 4 (AWC) from Kapiro village,

body length 29 mm. 7 - Habitus, dorsal view; 8 - ditto, ventral view; 9 - ditto, lateral view; 10-12 - C. dentifrons

(Aurivillius, 1925), holotype 3 (NHRS). 10 - Habitus, dorsal view; 11 - ditto, ventral view; 12 - ditto, lateral view.

(images 10-12 courtesy Jiff Pirkl).

FE a> LaDy,

Book4.indd 87 02-Jun-21 21:47:15

TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figures 13-19. Catypnes marazziorum sp. nov. 13-18 - Holotype <, genitalia. 13 - Median lobe with non inverted

endophallus, lateral view; 14 - ditto, dorsal view; 1.5 - ditto, ventral view; 16 - Tegmen, lateral view; 17 - Parameres,

ventral view; 18 - tergite 8, ventral view; 19 - Paratype 2 from Wau, eggs [scale bars 2 mm].

Book4.indd 88 02-Jun-21 21:47:19

Book4.indd 89

Drumont, A., Komiya, Z. & WEIGEL, A.: Catyones marazziorum sp. nov. (Coleoptera: Cerambycidae) from Papua New Guinea

MADE TG) Raley

pp. 83-90

" Philippines)

Palawan Negros a Catypnes dentifrons

af Mindanao

* ds 4 @ Catypnes macieayi

ae % Basilanilsland)

ts 2% & Catypnes marazziorum sp. n.

- * s EB Catypnes negrosianus

et “a H i

, oF Geer .

f ee wr _ Catypnes pascoei

‘Sulawes| ah ier.

EH Z> aS. ee oak gM aa ne he Catypnes pirkli

4 * fs Ser wey Bg ae 3 = Ss, 7 Bismarck Sea “*s ;

BB rconesis Se : 4 © Catypnes salesnei

; 1% Renda Ge ee =, = B (Pees Th t ee = Me

3 gia Papua New il ane i

bh : ar ryt Ze ‘ a Kratula sea by pe : ry aes _ Solomon Sea is = i

a) ate ; ore ad

1 be Si

NORTHERN :

TERRITORY

QUEENSLAND Lo New Caledonia

Australia

WEBRER A 0 2 ERRIMipeaiIe SIRT RMIEMe Mineg Sa aRR MAS POLL ORME mnt ay

AUSTRALIA Brisbane

oO.

SOUTH 4 Norfolk Island

AUSTRALIA yi

Perth '

‘=

NEW SOUTH

WALES & ie

Sydney

Adelaide 7

—— er o » &

ust in L, ere Oj

Big

2 oat

Melbourne a

Figure 20. Distribution map for all known species of Catypones Pascoe, 1864 (source: Google Maps).

by the following points: maximum width (including

medium teeth) of pronotum equal to the width at

shoulders while it is shorter in C. pirkli; surface of

elytra glossy instead of being mat in C. pirkli, and

by the elytral puncture which are deep and close

(especially visible on the elytral disc and the area

between the sutural line and the first costa) while

those are light and composed of small punctures

in C. pirkli.

Variability in the paratype series: none, except

the one related to the sex and the size of the

Specimens.

Distribution: The new species is distributed in

Papua New Guinea, presently known from the

Morobe and Oro provinces only (see fig. 20).

Checklist of species of Catyones Pascoe, 1864

Catypnes dentifrons (Aurivillius, 1925)

Distribution: Papua New Guinea: Bolan Mts.

Catypnes macleayi Pascoe, 1864

= Toxeutes punctatissimus Thomson, 1877

[female]

Distribution: Australia: New South Wales, Norfolk.

Catypnes marazziorum sp. nov.

Distribution: Papua New Guinea: Morobe and

Oro provinces.

Catypnes negrosianus (Hudepohl, 1987)

Distribution: Philippines: Mindanao, Negros.

02-Jun-21 21:47:19

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Catypnes pascoei (Lameere, 1904)

Distribution: Australia: Queensland.

Catypnes pirkli (Drumont, Komiya et Weigel,

2019)

Distribution:

Sulawesi.

Indonesia: South and _ Central

Catypnes salesnei (Delahaye, Drumont et Komiya,

2016)

Distribution: New Caledonia.

Acknowledgements

We sincerely thank the curators of the

various institutions who allowed us to examine

the ‘type material’ of the various Catypnes species

necessary to this long-term study of the genus:

Michael Balke (ZSM), Maxwell Barclay (BMNH),

Thierry Deuve (MNHN) and Bert Viklund (NHRS).

We are indebted to Jiri Pirkl (Czech Republic) who

manages the website www.prioninae.eu devoted

to the types of Prioninae of the world, providing

important and evident help for species identification

to the researchers all over the world in this subfamily

of Cerambycidae, and who kindly provided us

the pictures of the ‘type’ of T. dentifrons used in

this paper. We are grateful to Norbert Delahaye

(France), Jiri Dvoracek (Czech Republic) as well as

Giuseppe and Valentino Marazzi (Italy) for providing

us pictures and data from their collections.

Finally we also sincerely thank Hemant V. Ghate

(Department of Zoology, Modern College, Pune,

Maharasntra, India) for improving the language and

for suggestions made to the manuscript.

References

Aurivillilus C. 1925. Neue oder wenig bekannte

Coleoptera Longicornia. 21. - Arkiv for Zoologi 18A

(9): 1-22 [= 503-524], figs 141-163 (according to

Lin & Ge (2020)).

Delahaye N., Drumont A., Komiya, Z. 2016. Description

d’une nouvelle espéce de Toxeutes de Nouvelle-

Calédonie (Coleoptera, Cerambycidae, Prioninae,

Aegosomatini). - Les Cahiers Magellanes, New

Series, 23: 129-138.

DrumontA., Komiya Z., Weigel A. 2019. New contribution

to the knowledge of the genus Toxeutes Newman,

1840 with the description of a new species

from Sulawesi Island in Indonesia (Coleoptera,

Cerambycidae, Prioninae). - Besoiro 29: 3-13.

Jin M., Zwick A., Slipinski S. A., de Keyzer R., Pang

H. 2020. Museomics reveals extensive cryptic

diversity of Australian prionine longhorn beetles

with implications for their classification and

conservation. - Systematic Entomology 45: 1-26,

9 figs.

Lin M.-Y., Ge, S.-Q. 2020. Aurivillius’s ,Neue oder wenig

bekannte Coleoptera Longicornia” (1886-1927),

the correct years and page numbers. - ZookKeys

911: 113-137.

Yamasako J., Ohbayashi N. 2011. Review of the genus

Paragolsinda Breuning, 1956 (Coleoptera,

Cerambycidae, Lamiinae, Mesosini), — with

reconsideration of the endophallic terminology. -

Zootaxa 2882: 35-50.

Received: 18.xi.2020.

Accepted: 26.xii.2020.

02-Jun-21 21:47:19

Book4.indd 91

GOLovatcH, S. |., AKKARI, N., Goup, J. & TELNov, D.: Review of the Papuan genus Acanthiulus Gervais, 1844 (Diplopoda) ...

pp. 91-106

Review of the Papuan millipede genus Acanthiulus

Gervais, 1844 (Diplopoda: Spirobolida: Pachybolidae)

urn:lsid:zoobank.org:pub:BE62AA /C-13C0-4256-9AD2-AD8098ED53B4

SerGe! I. Gotovatcu *, NESRINE AKKARI *, JEROEN Goup ° & Dmitry TELNov * °

1 - Institute for Problems of Ecology and Evolution, Russian Academy of Sciences, Leninsky

prospekt 33, 119071, Moscow, Russia; sgolovatch@yandex.ru

2 - Naturhistorisches Museum Wien, Burgring 7, A-1010, Vienna, Austria; nes.akkari@gmail.com

3 - Naturalis Biodiversity Center, Darwinweg 2, 2333, Postbus 9517, 2300, RA Leiden,

The Netherlands;_jeroen.goud@naturalis.nl

4 - Natural History Museum, Cromwell Road, SW7 5BD, London, United Kingdom;

anthicus@gmail.com (ORCID: OO00-0003-3412-0089)

5 - Institute of Biology, University of Latvia, O. VacieSa iela 4, LV-1004, Riga, Latvia

Abstract: The genus Acanthiulus, which has hitherto been known to comprise three accepted species or Subspecies

endemic to the Papuan region, is revised, rediagnosed and shown to include only a single, quite variable species,

A. blainvillei (Leguillou, 1841), with A. blainvillei seotemtrionalis Attems, 1914 and A. wollastoni Hirst, 1914 both

considered as its new subjective junior synonyms, syn. nov. Pronounced morphological variations, all clearly

illustrated here, concern only peripheral characters, but the gonopodal structure remains very stable. Three morphs

are distinguished: A, B and C. The distribution of A. blainvillei is mapped, the genus and species apparently being

restricted to the Aru Archipelago, East Indonesia and much of New Guinea, both Indonesian and Papua New Guinea.

Certain clinal variation patterns and an evolutionary scenario can be suggested in the distribution and polymorphism

of the widespread species A. blainvillei at the northern periphery and in the centre of its distribution area.

Keywords: Millipede, taxonomy, new synonymy, iconography, polymorphism, microevolution, Papuan Region.

Introduction

According to the latest catalogue (Jeekel

2001), the Papuan genus Acanthiulus Gervais,

1844 is currently known to comprise only the

following three accepted species or subspecies.

These are A. blainvillei (Leguillou, 1841) (the type

species of the genus, with several synonyms and

a variety), from Indonesian, former Dutch New

Guinea (Alkmaar, “Kajo Bay”, “between Njad and

Sekopo” at Tami River, presently Papua Province of

Indonesia), northern New Guinea, former Kaiser-

Wilhelmsland, presently Papua New Guinea (Sepik

River and Astrolabe Bay), and Aru Archipelago, East

Indonesia (Dobo, Manoemai, Wokam, and Womar,

East Indonesia); A. blainvillei septemtrionalis

Attems, 1914, from western New Guinea (Tanah

Merah Bay, Hollandia, coast south of Humboldt

Bay, Bewani River, and Zoutbron near Begowri

River, Papua, Indonesia); and A. wollastoni Hirst,

1914, from southwestern New Guinea (Mimika

River, Papua, Indonesia).

Acanthiulus forms are very characteristic

due to most of the body rings/segments having a

number of conspicuous cones or spines (Figs 1A,

1B). This is in stark contrast to the more commonly

encountered, often large-sized and sympatric

Spirobolida, which typically lack an ornamented

integument. The generic etymology is based on this

prominent ornamentation with “acantha” meaning

“spine” in Greek.

Even though Acanthiulus is a small, oligotypic

genus, its taxonomic history is quite convoluted

(Jeekel 1971; 2001). It was first proposed as a

subgenus of /Julus Linnaeus, 1758 (recte: Julus)

by Gervais (1844) to encompass only J. Blainvillii

(recte: blainvillei) Leguillou, 1841, from = an

unspecified locality in New Guinea. The species was

briefly redescribed (Gervais 1844) and, a little later,

schematically illustrated (Gervais 1847), apparently

using the type material in the Paris Museum. The

species was thereby erroneously transferred to the

order Spirostreptida (Gervais 1847). Latzel (1884)

suggested, but did not formalize the synonymy of

Acanthiulus with Trachyiulus Peters, 1864 (recte:

Trachyjulus), again a member of Spirostreptida,

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while Bollman (1893) agreed to their close affinities,

but rejected the synonymy.

Daday (1893) very imperfectly described

and schematically illustrated a second species,

Spirobolus dentatus Daday, 1893, from Kaiser-

Wilhelmsland, New Guinea, which Silvestri (1895)

synonymized with Trigoniulus Blainvillii [sic!], and

provided its brief redescription. Acanthiulus was

thus sunk under Trigoniulus Pocock, 1894, not vice

versa.

Pocock (1893) was apparently the first to

consider Acanthiulus as a genus of full rank in

Spirobolida, close to Spirobolus Brandt, 1833,

but easily distinguished by the presence of eight

transverse rows of spines on most body rings. He

also described a new species, A. Murrayi (recte:

murrayl) Pocock, 1893, from “Wokan Dobbo”,

Aru Islands, nicely illustrated the body sculpture,

and corrected the spelling of the type species to

blainvillei, aloeit referred to it as Blainvillei.

Porat (1894) described another new species of

Acanthiulus, this time from Cameroon, west-central

Africa: A. tuberculosus Porat, 1894. In addition,

he erected a new genus, Thrinciulus Porat, 1894,

to accommodate two different new species from

Cameroon, but designated Acanthiulus murrayi as

its type species (cf. Carl (1913) and Jeekel (1971;

2001)). Acanthiulus tuberculosus has since Pocock

(1903) been known to have nothing to do with the

true Acanthiulus. Moreover, it has long become

(even twice!) the type species of two formal genera

of Spirostreptida, the valid one being Tropiiulus

Silvestri, 1896 (Jeekel 1971). Paradoxically,

Acanthiulus tuberculosus is still listed as such on

the web (https://www.gbif.org/species/1026912).

Bouvier (1903) described and _ beautifully

illustrated yet another new species, Acanthiulus

maindroni Bouvier, 1903, from southern India.

However, like A. tuberculosus, it has nothing to do

with the true Acanthiulus, albeit it does belong to

the same family Pachybolidae, order Spirobolida.

Bouvier (1903) also stated that he had examined

the holotype of Acanthiulus blainvillei in the Paris

Museum, and compared it to his A. maindroni.

Brolemann (1903) provided a comparative study of

the gonopods of Acanthiulus vis-a-vis several other

genera of Spirobolida, for which, however, he used

the gonopods of maindroni... At present, maindroni

is considered as the sole accepted species of

Eucentrobolus Pocock, 1903, a genus endemic to

southern India (Golovatch & Wesener 2016).

Pocock (1903) must have overlooked Silvestri’s

(1895) proposed synonymy of Daday’s dentatus

with blainvillei, as well as Porat’s (1894) earlier

typification of Thrinciulus. He was also so badly

misled by Daday’s (1893) erroneous statement

that the ozopores in dentatus were located behind,

not before the stricture on the body rings, that

he created a new genus, Polybunobolus Pocock,

1903, for murrayi alone, aS opposed to Dlainvillei

and dentatus which remained in Acanthiulus. From

the very start, Polybunobolus thus became an

objective junior synonym of Thrinciulus.

Carl (1913) not only extensively discussed the

taxonomy of Acanthiulus, but he also redescribed,

nicely illustrated and recorded A. murrayi from

three places in the Aru Islands: Teranhan: Ngaiguli;

Wammer: Dobo; and Wokam: Samang. In addition,

Carl (1913) formally synonymized Polybunobolus

with Acanthiulus, but Brolemann (1913) questioned

that synonymy and corrected the scope of

Acanthiulus compared to his own earlier treatment

(Brolemann 1903). He also provided a most detailed

redescription and a number of line drawings of A.

blainvillei coming from an unspecified locality in

New Guinea; more importantly, he was the first to

properly illustrate the gonopodal structure of A.

blainvillei (reproduced here in Figs 29-32).

Attems (1914a) very concisely described

a new subspecies and a variety of A. blainvillei

from mainland New Guinea, i.e. A. blainvillei ssp.

septemtrionalis Attems, 1914 and A. blainvillei

var. intermedius Attems, 1914, also presenting

a key to both subspecies and the variety of A.

blainvillei. Chamberlin (1920) slightly misspelled

the name septemtrionalis as septentrionalis [sic!]

and, more importantly, once referred to the variety

intermedius as A. blainvillei intermedius. However,

as the latter name was clearly listed as a variety

earlier in the same paper, it seems improper to

formally treat intermedius as a subspecies and to

allot it a taxonomic rank.

Finally, Hirst (1914) described a new species,

A. wollastoni Hirst, 1914, from Dutch New Guinea

(Mimika River), thus providing the latest contribution

to the scope of Acanthiulus. As a result, Jeekel

(2001) considered valid only the following three

species or subspecies: A. blainvillei (Leguillou,

1841), A. blainvillei septemtrionalis Attems, 1914,

and A. wollastoni Hirst, 1914.

Jeekel’s (2001) catalogue of Indo-Pacific

Spirobolida is fairly complete and relevant, but it

makes sense to reiterate it in full at least as regards

Acanthiulus, because it omitted a reference and

some original spelling details of all species-level

names.

The main objective of our contribution is to

identify the Acanthiulus material housed in the

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Zoological Museum of the Moscow University

(ZMUM), Russia, aS well as to present the first

relevant iconography of Acanthiulus.

Material and methods

Most of the fresh samples were collected

recently and donated to the ZMUM. All Acanthiulus

material, both type and non-type, identified by

Carl Attems and currently kept at the Natural

History Museum of Vienna, Austria (NHMW) and

the Naturalis Biodiversity Center, Leiden, The

Netherlands (NBCL) has also been considered

and properly illustrated. The type series of another

congener housed in the Natural History Museum in

London (BMNH) has also been accessed, revised

and illustrated. On-spot field observations and

images were made by one of us (DT) and Martins

Kalnins (Sigulda, Latvia). In addition, a specimen

of Acanthiulus filmed at Bosavi Volcano, Papua

New Guinea for BBC’s “Lost land of the volcano”

(https://www.bbc.co.uk/programmes/b00Mm82h7)

has been identified from video and mapped as well.

Pictures were taken with a Canon EOS 5D

digital camera and stacked using Zerene Stacker

software (ZMUM), with a Nikon DS-Ri-2 camera

mounted on a Nikon SMZ25 stereo microscope

using NIS-Elements Microscope Imaging Software

with an Extended Depth of Focus (EDF) patch

(NHMW), a Nikon D800 with a Nikkor AF-S 105

mm Macro lens (RMNH), or a Zeiss Axio Zoom V.16

(BMNH).

Because numerous locality and even

regional names in New Guinea have been

notoriously changed or repeatedly misspelled,

while some have simply vanished (e.g., Bronbeek,

Modderlust or Tanah Merah Bay around Jayapura

alone) over the island’s historical record, an

updated list of relevant toponyms Is provided in the

legend to Map. The modern name always follows

earlier Synonyms, if any.

In the catalogue sections per species

below, D stands for a description or descriptive

notes (sometimes also including a key, discussion,

new status, Synonymy or combination), R for new or

old records, and M for mere mention.

Results

Family Pachybolidae

Genus Acanthiulus Gervais, 1844

pp. 91-106

by subsequent designation of Gervais (1844).

= Thrinciulus Porat, 1894

Type species: Acanthiulus murrayi Pocock,

1893, by subsequent designation of Porat (1894).

= Polybunobolus Pocock, 1903

Type species: Acanthiulus § murrayi

1893, by original designation.

Pocock,

A full generic synonymy list is available in Jeekel

(2001). Because the genus appears to contain only

one species, below we combine its diagnosis and

description.

Diagnosis and description: A genus of

particularly large-sized Pachybolidae (adults 100-

210 mm long and 9-14.5 mm in diameter, and

the collum being smooth and bearing unusually

prominent, relatively narrowly rounded, flap-

Shaped and sinuous paraterga drawn clearly

below the venter, slightly more narrow in the <,

distinctly bordered: narrowly along the anterior and

lateral margins and especially broadly along the

caudal margin (Figs 3, 9-10, 16, 19, 42, 62-65,

68). Most body rings are heavily sculptured and

show a transverse row of 1+1, 3+3 or 4(5)+4(5)

conspicuously enlarged cones or spines on each

metazona, often with small intermediate crests,

ribs, striae or bosses (Figs 1, 3-5, 9-12, 16-23,

33-36, 42-46, 52-56, 62-68, 71-72); the

telson has a very small epiproct not overhanging

the paraprocts and lacking a caudal process, each

paraproct being distinctly impressed before the

caudal marginal lip (Figs 5, 12, 22-23, 36, 46,

56, 63, 67, 72). The hypoproct is very wide and

short, with an almost straight caudal margin. The

adult body is with 50-55p+T segments/rings. The

colouration is usually dark brown to blackish, rarely

dark olivaceous, only the antennae and ozopores

are sometimes contrasting red or yellowish (Figs 1,

3-5, 9-12, 16-23, 33-36, 42-46, 52-56, 62-68,

71-72), and the legs red-brown, occasionally with

yellow tarsi. Eye patches round, ca 50 ommatidia

arranged in eight vertical rows. Supralabral setae

2+2, the labrum has three distinct medial teeth

inside a deep notch at, and a short, but evident

axial suture/line above, the anterior margin (Figs

17, 18, 33, 43, 53, 65, 69). The antennae are

short, C-shaped, a little longer and reaching the

caudal margin of the collum in the 4; antennomere

2 is the longest, antennomere 8 has four apical

sensory cones (Figs 3, 9, 10, 16-19, 33, 42-43,

52-53, 62-65, 68-69). Striations on postcollum

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ringS are as uSual: very fine, dense, vertical and

often confused striolations on prozonae; arcuate,

somewhat more distinct, often similarly confused

striations on mesozonae; and usually rough striae,

also increasingly dense and distinct ventrad, on

metazonae (Figs 3-5, 9-12, 16-23, 33-36, 42-

46, 52-56, 62-68, 71-72). A pro-mesozonal line

is absent. The tegument is mostly shining, delicately

reticulate, in places micropunctate, only metazonae

(but not their ornamentations!) are often rough and

dull, sometimes coated with dirt. Scobinae absent,

the limbus is unmodified. Ozopores starting with

ring 6 and lying in the mesozona, well before the

meso-metazonal suture/line (Figs 9, 10, 16, 19,

42, 52, 62-64, 68, 71-72). 4 coxae are swollen

ventrally, with one spine located above and 2-3

below the claw; all podomeres are more or less

bulged ventrally, each of those lying proximal

to the tarsus normally supports one seta. The

penes are basally a single, but quickly and widely

diverging structure, each branch consisting of

5-6 constricted rings and apically surmounted by

a small papilla (Figs 17, 53). 4 ring 7 is swollen

ventrally, a complete ring bearing a clear ventral

bridge in the caudal half (Figs 33, 44, 54, 70). ¢

legs are a little longer than @ ones, all 4 coxae are

rounded and somewhat swollen, prefemora and

femora compressed laterally and slightly ridged

ventrally, tarsi without sole pads (Figs 53-54).

Both gonopod pairs mostly very heavily sclerotized.

The anterior gonopods (Figs 6, 7, 13-15, 25-27,

29, 30, 37-38, 47-48, 57-58, 73-74) are witha

broad sternite (Ss) attached laterobasally to tracheal

apodemes (a) and extended medially into a high,

distally bilobed, apically diverging and rounded

central process (sp) which divides both coxae (cx)

and consists of two medially fused halves still

visibly separated by a deep suture. The coxae (cx)

are Set laterally on, and are only a little higher than,

the sternal process (Sp); each coxa (cx) is roundly

subtriangular to squarish in shape, hollow on the

caudal face and contains a smaller and similarly

Shaped telopodite (t) sunken inside the hollow; t is

slightly bilobed apically, the more narrow, rounded

and finger-shaped apical lobe lying mesally. The

posterior gonopods (Figs 8, 28, 31-32, 39-41,

49-51, 59-61, 75-77) are 2-segmented, the

sternite (S) being strongly chitinized, elongate,

Subcylindrical and transverse, divided, but fused

centrally, either side with a small ampulla/vesicle

and borne on a large tracheal apodeme (a). The

coxa (cx) is rather simple, boat shaped, regularly

and clearly curved mesad, mostly hollow on the

mesal side; in the basal half it carries a seminal/

prostatic groove that originates from the ampulla

and ends up at about cx midheight near a small,

squarish, serrate and chitinized lobule (lo) at the

bottom of the cx hollow. The apical part of cx is trifid,

showing a larger, subapical and subtriangular lobe

(Ib), rounded to subacute, at the anterior margin,

and two particularly small, apical denticles (d), both

subacute to acute.

Acanthiulus blainvillei (Leguillou, 1841) (Figs

1-77, map)

Julus Blainvillei [sic!] Leguillou, 1841: 279 (D).

lulus Blainvillii [sic!] - Gervais 1844: 70 (D); 1847: 173

(D, pl.44, figs 8-8b).

Acanthiulus Blainvillii [sic!] - Preudhomme de Borre

1884: 52 (M).

Acanthiulus Blainvillei [sic!] - Preudhomme de Borre

1884: 63 (M); Pocock 1893: 137 (M); 1903: 531

(D); Bouvier 1903: 264 (M); Attems 1914b: 352

(M); 1915: 1, 6, 31 (D, figs 6-9); 1927: 62 (M).

Acanthiulus blainvillii [sic!] - Bollman 1893: 139 (M);

Hoffman, Keeton 1960: 6 (M).

Spirobolus dentatus Daday, 1893: 101 (D, pl.3, figs

1-7), Wilhelmsland, New Guinea; synonymized

by Silvestri (1895: 654); Carl 1913: 275, 276 (D);

Jeekel 2001: 56 (M, R).

Acanthiulus Murrayi [sic!] Pocock 1893: 136 (D, pl.9, figs

7-7b), Wokan Dobbo, Aru Islands; synonymized by

Attems (1914b: 351).

Acanthiulus dentatus - Pocock 1903: 531 (D); Carl

191352075; 2 76D):

Acanthiulus Murrayi [sic!] - Bouvier 1903: 264 (M);

Attems 1914b: 352 (M).

Trigoniulus Blainvillii [sic!] - Silvestri 1895: 654 (D).

Polybunobolus Murrayi [sic!] - Pocock 1903: 531 (D).

Acanthiulus murrayi - Carl 1913: 276 (D, pl.11, figs 14-

17), Aru Islands (Carl, 1913); Chamberlin 1920:

216 (M); Jeekel 2001: 56 (M, R).

Acanthiulus blainvillei - Brolemann 1913: 105, 109 (D,

pl.15, figs 23-26), New Guinea, no precise locality;

Attems 1914a: 383 (D); 1932: 4 (R, Manoembai,

Aru Islands); Chamberlin 1920: 215 (M); Jeekel

1971: 191 (M); 2001: 55 (M, R).

Acanthiulus blainvillei var. intermedius Attems 1914a:

381, 382, 384 (D). Northern Dutch New Guinea:

near Kajo Bay, between Njad and Sekopo at Tami

River, and Astrolabe Bay (Attems, 1914a); Jeekel

2001: 56 (M, R).

Acanthiulus blainvillei var. intermedius - Chamberlin

1920: 215 (M); Moritz & Fischer 1975: 245 (M).

Acanthiulus blainvillei ssp. septemtrionalis Attems,

1914a: 383 (D), syn. nov. Northern Dutch New

Guinea (Attems, 1914a); Jeekel 2001: 56 (M, R).

Acanthiulus Blainvillei [sic!] var. intermedius - Attems

1914b: 41, 84, 352 (M); 1915: 34 (M).

Acanthiulus Blainvillei [sic!] ssp. septemtrionalis Attems,

1914b: 41, 84, 352 (M); 1915: 32 (M).

Acanthiulus wollastoni Hirst, 1914: 330, fig. 17 (D), syn.

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nov. Southern Dutch New Guinea (Hirst, 1914);

Jeekel 2001: 56 (M, R).

Acanthiulus Blainvillei [sic!] - Attems 1915: 6, 7, 34 (D,

figs 6-9). Southwestern (Alkmaar) and northern

New Guinea (Kaiserin Augusta River), Aru Islands

(Attems, 1915).

Acanthiulus blainvillei intermedius [sic!] - Chamberlin

1920: 267 (M).

Acanthiulus wollastoni - Chamberlin 1920: 216, 267

(M).

material: Julus blainvillei: &

Type or original

holotype (Paris Museum, MNHN HB 008, J.-P. Mauriés &

J.-J. Geoffroy in litt.; not examined); Acanthiulus murray:

9 holotype (BMNH, not examined); Spirobolus dentatus:

~ « _—

pp. 91-106

3 lectotype, 4 paralectotype (Budapest Museum, HNHM

974.a.2/2, Korsos (1983) and in litt.; not examined);

Acanthiulus blainvillei var. intermedius syntypes: 1

3 (NHMW 8433, Umgebung von Kajo Bai, zwischen

Njad und Sekopo, P.N. van Kampen & K. Gjellerup leg.

[Papua Province, Jayapura Regency, Muaratami District,

between Nyao and Skouw seaside area], examined,

Figs 3-8), 2 299 (NBCL RMNH DIP 562 and DIP 563,

examined), 14,3 99 (Berlin Museum, ZMB 5220-5222

(Moritz, Fischer, 1975), not examined); Acanthiulus

blainvillei ssp. septemtrionalis syntypes: 2 do, 5 9°

(NBCL RMNH DIP 571-577, Indonesia, NE Papua,

Tanah Merah Bay; Indonesia, NE Papua, Hollandia (=

Jayapura, = Port Numbay), S2.538554, E140.705194;

Indonesia, NE Papua, between biwak Kasawari and

(s 2 * : 2 ee ¥ t : . ;

Atk a Sa ee stots ig Te Sia i 2

a SS Cw B-

Figures 1-2. Live 4 (1A-B) and eggs (2) of Acanthiulus blainvillei (Leguillou, 1841), morph C, in the field

from near Lake Kamakawalar, West Papua [not to scale] (image 2 courtesy Martins Kalnins).

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biwak Modderlust [S2.69, E140.85], = [Kustengebiet

sudlich von der Humboldtbai; Indonesia, NE Papua,

Bewani River [am Bewani-Fluss]; Indonesia, NE Papua,

between biwak Bronbeek and _ biwak Modderlust;

Indonesia, NE Papua, biwak Zoutbron, 1910-1911,

P.N. van Kampen & K. Gjellerup leg.; examined, Figs

9-15); Acanthiulus wollastoni syntypes: 1 ¢, 1 Q

(BMNH, British Ornithologists’ Union Expedition and

the Wollaston Expedition in Dutch New Guinea, Mimika

River; examined, Figs 16-28); Acanthiulus blainvillei: 4

O92 (NHMW 8432, Indonesia, Moluccas, Aru Islands,

Kobroor Island, Manumbai, Trip Prince Leopold Belgium

1928/29, don. 1931, det. C. Attems; examined).

New material examined: Acanthiulus — blainvillei

(forma typica): 2 64, 3 99 (ZMUM), E Indonesia, West

Papua Prov., S Bird’s Neck Isthmus, 47 km E of Kaimana,

Triton Bay, environs of Kamaka (former Warika) village,

surroundings of Lake Kamakawalar, S3°46’22”,

E134°12’02”, 60-310 ma.s.l., primary lowland tropical

rainforest on limestone, 8.IX.2010, D. Telnov leg.; 1 4, 4

9 (ZMUM), same locality, S3°45’33”, E134°12’05”, 90

ma.s.l., primary lowland tropical rainforest on limestone,

8.IX.2010, M. Kalnins leg.; 2 specimens (Sex unknown),

E Indonesia, West Papua Prov., S Bird’s Neck Isthmus,

2-4 km NE of Kaimana, S3°39’, E133° 46’, 150-200

m, 19-20.1X.2010, edge of disturbed lowland rainforest

on limestone, leg. M. Kalnins.

Acanthiulus blainvillei (forma septemtrionalis): 2 33, 1

9 (ZMUM), E Indonesia, West Papua Prov., 32 km SSE of

Jayapura, near Indonesia - Papua New Guinea border

cross point, S2°37’21”, E140°58’42”, 170-200 m

a.s.l., primary lowland tropical rainforest on limestone,

17.11.2018; 1 4 (ZMUM), E Indonesia, Papua Prov.,

Star Mountains, 13.5 km SSE of Oksibil, S4°59’33”,

E140°42’02”, 780 m a.s.l., primary lowland tropical

rainforest, 14.111.2018.

Morphological variations and taxonomic

implications

Numerous morphological characters of

Acanthiulus blainvillei have been shown to vary,

sometimes considerably so. The above diagnosis

and description reflect most of the variable traits.

By far the most conspicuous variations

concern tergal structures, in particular, the

number and shape of metatergal teeth, cones or

spines. Attems (1914a) was the first to recognise

intraspecific categories within A. — blainvillei,

allocating them either to the rank of a subspecies

(septemtrionalis) or a variety (intermedius) and

paying special attention to that particular feature.

His key to all three varieties can be repeated as

follows (translated from German):

“1a. Each metazonite has 8 or 6 large and sometimes

further smaller teeth, excluding metazonites 2 to 5

or 6, where rows gradually begin. All rows reach the

penultimate segment. 3’, 9 are with 50-52, mostly 51,

body segments (New Guinea, Aru ISIands) .......cceeeeeee 2

2a. Large teeth on metazonites are long and sharp, and

present in 8 rows. 3 9.6-11 mm wide

BON, stesso ee a aR ee ee eer blainvillei Le Guillou.

2b. Large teeth on metazonites are much shorter and

blunt, and present in 6 rows, the other rows are much

smaller, sometimes totally absent. 3 13.5 mm, @ up to

DAL TN WIGS saccriiatanneconantetdanrereemncenst var. intermedius Att.

1b. Each metazonite has only 2 large teeth, the rows

begin from segment 6 or 7 and end on segment (3) 4

or 5 from behind. @, 2 are with 53-56 body segments.

Width 13.5-14.5 mm (northern Dutch New Guinea) .....

We raer te castrate oons cet subsp. septemtrionalis n. subsp.”

In that key, Attems (1914a) unwillingly omitted

A. wollastoni Hirst, 1914, because its description had

not yet been published. Hirst (1914) characterized

A. wollastoni as being blackish in colouration, with

50 body rings, and 103 (4) or 115 mm () in length

and 9.1 (4) or 10.2 mm () in width. He compared

A. wollastoni (two syntypes, 3 and @) to A. murrayi

(2 holotype), stating that the antennomeres in A.

wollastoni were stouter and enlarged distad (vs

elongate and not so much enlarged in A. murrayi);

the metaterga supported eight rows of “thorns”, of

which four dorsal rows were much better developed

than in A. murrayi, being obsolete or absent from

rings 2 and 3, bosses on the 4", spiniform from

the 5" on, and all eight fully developed from the

7", The caudal parts of the rings in A. wollastoni

carried small ridges, and the paraprocts were less

coarsely punctured posteriorly than in A. murrayi.

Only the gonopods were illustrated (Figs 27-28).

Having revised both syntypes of A. wollastoni, we

take the opportunity to illustrate here most of its

peripheral and gonopodal characters (Figs 16-23),

with the exception of the posterior gonopod which

was found either lost or misplaced.

Given that A. murrayi has since Attems (1914b)

been treated as a junior synonym of A. blainvillei,

and that all of the characters of A. wollastoni fail to

reflect more than individual variations observed In

A. blainvillei, sometimes even sex-linked ones (e.g.

the length of the antennae), we do not hesitate

to formally consider A. wollastoni as another

Subjective junior synonym of A. blainvillei, syn.

nov. In this connection, no lectotype designation is

necessary for the type series of A. wollastoni.

The situation concerning the status of the

subspeciesseptemtrionalisisnotsostraightforward,

as the external differences between the typical

form of A. blainvillei and the subspecies (= taxon)

septemtrionalis are indeed very apparent (see

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pp. 91-106

Figures 3-8. Acanthiulus blainvillei (Leguillou, 1841) 2,

morph B, from between Nyao and Skouw, NHMW 8433.

3 - Anterior part of body, lateral view; 4 - Midbody rings,

dorsal view; 5 - Posterior part of body, lateral view; 6 -

Left anterior gonopod, posterior view; 7 - Sternum and

right anterior gonopod, anterior view; 8 - Left posterior

gonopod, anterior view [not to scale] (images courtesy

Oliver Macek).

Legends: a - Tracheal apodeme; cx - Coxa; s - Sternum;

sp - Sternal process; t - Telopodite.

Figures 9-15. Acanthiulus blainvillei (Leguillou, 1841)

3, morph A (3 syntypes of Acanthiulus blainvillei

ssp. septemtrionalis Attems, 1914), from Hollandia (= ,

Jayapura, = Port Numbay), and between biwak Kasawari a

and biwak Modderlust, NBCL RMNH DIP 572 and DIP j

573, respectively. 9-10 - Anterior parts of body, lateral °

views; 11 - Midbody rings, lateral view; 12 - Posterior

part of body, lateral view; 13-15 - Anterior gonopods,

anterior, lateral and posteroventral views, respectively

[not to scale].

Legends: a - Tracheal apodeme; cx - Coxa; s - Sternum;

sp - Sternal process; t - Telopodite.

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TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

the key above). However, given the features of the

variety intermedius bridging those of blainvillei

and septemtrionalis, we are inclined to follow

Attems (1914a; 1914b) in treating A. blainvillei as

a single, widespread and polymorphous species

containing three morphological forms. Only the

status of septemtrionalis is to be downgraded,

and the following new formal synonymy advanced:

A. blainvillei septemtrionalis is another subjective

Junior synonym of A. blainvillei, syn. nov. To avoid

the use of Latin names for the infrasubspecific

categories, and considering the least ornamented

morph as_ evolutionarily basal, we designate

septemtrionalis as morph A, intermedius as morph

B, and the main form of A. blainvillei as morph C.

No lectotype designation is likewise needed for the

type series of A. blainvillei septemtrionalis. The

same obviously concerns the series of syntypes of

A. intermedius.

Variations in metatergal sculpture, primarily the

size and shape of spines/teeth on the metazonae,

are the best pronounced in morph C, apparently

in relation to its particularly vast distribution and

much more abundant material available for a

comparative analysis. Cones start with ring 2 and

on rings 6 or 7 they turn into /Jarger spines/teeth

that can be sharp and strongly inclined caudad,

usually produced past the rear tergal margin, but

sometimes they remain clearly shorter, blunt or

rounded cones lying within the ring’s hind contour.

Each metazona has 8-10 strong spines, teeth or

cones, these being less regular in the anterior half

of the body and with some smaller intermediate

ridges or denticles which are more conspicuous on

rings 2-5(6). Spines invariably end on the last ring,

just before the telson (Figs 22-23, 36, 46).

In morph B, obviously much more rare and

localized, there are 3+3 strong, stout and mostly

blunt or rounded cones, additionally also an axial

and 2-4 much smaller, intermediate, longitudinal

crests/ribs located between the larger, but still

moderately strong teeth. Metatergal cones are

evident and also start already with ring 2, quickly

grow and turn into larger cones on all following rings

(Figs 3-5). Attems (1914a) considered this variety

as being more similar to morph C than to morph A.

Finally, morph A also seems to be rare and

quite restricted in distribution. It shows only 1+1

strong cones/teeth on midbody metazonae, both

located just below the ozopore level, with the

bases of the teeth taking up most of the metazona

length, and the tips acute or rounded and either

drawn past or remaining within the caudal margin.

Cones are traceable starting with ring 2, being

represented by rounded tubercles and gradually

growing towards rings 6 to 9, to abruptly get fully

or almost fully reduced only on the last few (usually

three) rings before the telson. The dorsum between

both lateral rows of teeth is roughly, longitudinally

and irregularly rugose, with additional and usually

barely traceable crests/ribs and/or denticles (Figs

9-12, 52-56, 62-68, 71-72).

It seems that only because morph A is

apparently |= more disjunct morphologically

compared to both B and C that Attems (1914a),

following the morphological subspecies concept,

allotted it the rank of a subspecies. Instead we

stick to the modern biological subspecies theory

and practice (https://simple.wikipedia.org/wiki

Biological species concept), regarding A, B and C

as morphs of a single polymorphous species.

As regards the number of body rings,

it only seems to vary very little, 50-55p+T,

since only adults seem to have hitherto been

captured. Juvenile instars have still escaped from

examination. This, against the background of very

considerable size variations (LO-21 cm long and

9-14.5 mm in diameter), clearly suggests several

mature stadia in the development of A. blainville.

As the entire order Spirobolida seems to grow with

hemianamorphosis alone (Enghoff et al. 1993),

it would be very interesting to trace the evolution

of polymorphism in that remarkable species.

Molecular studies seem very promising in this

respect as well.

Among the main features that definitely remain

very stable and prove to be most characteristic

of the species as a whole, are the shape of a

dorsally smooth collum, albeit with certain sex-

linked variations, and the gonopodal structure

(Figs 6-8, 13-15, 25-32, 37-41, 47-51, 57-61,

73-77). Morphological variation in the gonopodal

conformation, the basic character accepted in

modern diplopod systematics, appears to be

so minor and virtually negligible that, following

Attems (1914a), we tend to treat A. blainvillei as

a remarkable, but typical example of infraspecific

polymorphism, despite the often conspicuous

peripheral differences between populations

(morphs A, B and C). Another remarkable feature of

A. blainvillei seems to be strict allopatry observed

between the morphs (Map). The only exception

superficially concerns A. blainvillei and A. blainvillei

intermedius, both recorded from Kaiserin Augusta

(= Sepik) River by Attems (1915), but no exact

locality was presented for either form.

Moreover, thezoneofpronounced polymorphism

and microevolution appears to be relatively limited,

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pp. 91-106

o | B é

e 3 Figures 25-28. Gonopods of Acanthiulus

i iS blainvillei (Leguillou, 1841), morph C (¢ and @

|." syntypes of Acanthiulus wollastoni Hirst, 1914),

le a 4 from Mimika River. 25-27 - Sternum and right

+ = : ; : ¥

k ¥ anterior gonopod, anterior, posterior and anterior

| &

views, respectively; 28 - Left posterior gonopod,

anterior view.

Legends: a - Tracheal apodeme; cx - Coxa; s

- Sternum; sp - Sternal process; t - Telopodite.

Drawings (27-28) reproduced after Hirst (1914)

22 [not to scale].

PEO yl

Figures 16-24. Original label (24), as well as 3

(16-17 & 22) and 2 (18-21 & 23) of Acanthiulus

blainvillei (Leguillou, 18441), morph C (¢ and @

syntypes of Acanthiulus wollastoni Hirst, 1914),

from Mimika River. 16-19 - Anterior parts of

body, lateral, ventral, ventral and lateral views,

respectively; 20-21 - Middle part of body,

lateral and dorsal views, respectively; 22-23

- Posterior parts of body, lateral views [not to

scale].

Figures 29-32. Gonopods of Acanthiulus

blainvillei (Leguillou, 1841), morph C, 4 from

an unspecified locality in New Guinea. 29-30 -

Both anterior gonopods, posterior and anterior

views, respectively; 31-32 - Right posterior

gonopod, anterior and mesal views, respectively.

Legends: a - Tracheal apodeme; cx - Coxa;

d - Two apical denticles; Ib - Lobule at bottom

of coxal hollow; lo - Subapical mesal lobe; s -

Sternum; sp - Sternal process; t - Telopodite.

Drawings reproduced after Brodlemann (1913)

[not to scale].

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TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figures 33-36. Habitus of Acanthiulus _ blainvillei

(Leguillou, 1841), morph C, 4 from near Lake

Kamakawalar, West Papua. 33 - Anterior part of body,

ventral view; 34-35 - Middle part of body, dorsal and

lateral views, respectively; 36 - Posterior part of body,

lateral view [not to scale] (images courtesy Kirill V.

Makarov).

Figures 37-41. Gonopods of Acanthiulus _ blainvillei

(Leguillou, 1841), morph C, 4 from near Lake

Kamakawalar. 37-38 - Both anterior gonopods,

anterior and posterior views, respectively; 39-41 - Coxa

of right posterior gonopod, anterior, mesal and posterior

views, respectively [not to scale] (images courtesy Kirill

V. Makarov).

100

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Figures 42-51. Acanthiulus _ blainvillei

(Leguillou, 1841), morph C, a different

3S from near Lake Kamakawalar,

West Papua. 42-43 - Anterior part of

body, lateral and ventrocaudal views,

respectively; 44 - Ring 7 with intact

gonopods, ventral view; 45-46 - Middle

and posterior parts of body, respectively,

lateral views; 47-48 - Both anterior

gonopods, anterior and posterior views,

respectively; 49-51 - Coxa of right

posterior gonopod, anterior, mesal and

posterior views, respectively [not to scale]

(images courtesy Kirill V. Makarov).

Figures 52-61. Acanthiulus blainvillei

(Leguillou, 1841), morph A, @ from near

Jayapura. 52-53 - Anterior part of body,

lateral and ventral views, respectively (NB:

left paratergum on collum broken off); 54

- Ring 7 with intact gonopods, ventral

view; 55-56 - Middle and posterior parts

of body, respectively, lateral views; 57-

58 - Both anterior gonopods, posterior

and anterior views, respectively; 59-61 -

coxa of right posterior gonopod, anterior,

mesal and posterior views, respectively.

[not to scale] (images courtesy Kirill V. 46

Makarov).

pp. 91-106

101

02-Jun-21 21:47:53

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figures 62-67. Acanthiulus _ blainvillei

(Leguillou, 1841), morphA, 34 from near

Jayapura, Papua. 62 - Anterior part of

body, lateral view (to show an intact right

paratergum of collum); 63 - Habitus,

lateral view; 64-65 - Anterior part of body,

lateral and subventral views, respectively;

66-67 - Middle and posterior parts of

body, respectively, lateral views [not to

scale] (images courtesy Kirill V. Makarov).

Figures 68-77. Acanthiulus _ blainvillei

(Leguillou, 1841), morph A, 4 from near

Oksibil. 68-69 - Anterior part of body,

lateral and ventral views, respectively;

70 - Ring 7 with intact gonopods, ventral

view; 71-72 - Middle and posterior parts

of body, respectively, lateral views; 73-74

- Both anterior gonopods, anterior and

posterior views, respectively; 75-77 -

Coxa of right posterior gonopod, anterior,

mesal and posterior views, respectively.

[not to scale] (images courtesy Kirill V.

Makarov).

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pp. 91-106

"(¥SI| AW}]EOO] PS}USWWOD JO} aged x9U

99S) yYdiow J3ad puke Ajjesouas JSed 0] ISOM LOI PAJBQUUNU Je SAI{}edO} || “UOIeD0| |eX1YdeusOEs JOeXS JeajOUN Jay} UedW ‘| J, ‘Sjeyoeuq asenbs ul Saiyedo] Sy

(OT SIDUV YUM pesedaid) ajsuel -— 9 Ydiow ‘puoWeIp — g Yduow ‘ajould - YW Ydsouw sayjiAU/e/g SNjniyzUeoY JO UOIINGIISIP au “de,

103

02-Jun-21 21:48:01

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Book4.indd 104

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

being mainly restricted to northern and, to a

lesser extent, central New Guinea (Map). If the

presumably basalmost morph A indeed originated

somewhere in that area, then the evolutionarily

more advanced morph B and, especially, morph C

could be viewed as later offshoots and expansion

waves that have attained a still similarly restricted

and a much wider distribution, respectively. The

particularly successful and common morph C

presently occurs over much of New Guinea and in

the Aru Archipelago, but neither at the presumed

origin centre near Jayapura nor in more or less

central New Guinea (Map).

As an alternative viewpoint, superficially all

or some of the morphs could also be regarded as

independent species. For example, the particularly

rare A. intermedius might be considered as a

still somewhat underdeveloped A. blainville/, i.e.

its younger, but already fully mature ontogenetic

stadium or stadia (Enghoff et al. 1993). However,

this is hardly likely, because the 3 and Q of A.

intermedius attain the same, typically very large

size (13.5 or 14 mm in diameter, respectively, and

51-52 body rings), while all three morphs, and this

being especially important, share exactly the same

gonopodal structure.

Notes on Acanthiulus blainvillei bionomy

The largest observed specimens of Acanthiulus

blainvillei were the two females from northern

New Guinea, from near the Indonesian - Papua

New Guinean border crossing point (See above for

locality details). Measured in situ (live specimens)

by the fourth author: one female amounted to a

total body length of 210 mm, while the second one

to 190 mm. Unfortunately, that material was not

preserved for subsequent laboratory study.

At all sites of the recent observations made in

Indonesian New Guinea (the vicinities of Jayapura

and Kaimana, Star Mountains, and Triton Bay, see

above for details), Acanthiulus specimens were

found in limestone-rich areas or immediately at the

base of limestone rocks or hills. The presence of

limestone in the habitat seems to be ecologically

important if not crucial for this very large species

which definitely requires a serious calcium supply

to support its massive and hard exoskeleton.

The animals were observed feeding on fallen

leaf litter, only partially decomposed (already

brown) leaves being consumed by the millipedes.

The population density of adults is very low, usually

ca 2-3 specimens per 100-200 m? of primeval or

secondary rainforest, as their active and purposeful

104

search on open ground and in various potential

shelters like under logs and larger fallen leaves or in

tree stumps shows. Juveniles are especially rarely

encountered, since only a single young instar of ca

6 cm long was observed walking on the ground over

the whole survey time (a few weeks in 2010 and

2018).

Oviposition in A. blainvillei was observed on

the spot near the village of Warika, Bird’s Neck

Isthmus of New Guinea, by Martins Kalnins and

Dmitry Telnov. The female laid eggs separately, one

by one, encapsulating each egg inside an ovoid

“cocoon” ca 1-1.5 cm in diameter (Fig. 2). Each

cocoon, or egg capsule was produced of excrement

from an everted hind gut and, upon oviposition, it

was flattened dorsoventrally on sides through tightly

Spiralling the body. The female left egg capsules

directly on the rainforest ground.

Adult specimens taken in September 2010

survived for almost two years in captivity with

neither breeding nor an oviposition Success.

Acknowledgements

We are most grateful to Janet Beccaloni

(BMNH), Oliver Macek (NHMW) and Kirill V. Makarov

(Pedagogical State University, Moscow, Russia)

for their help in arranging access to type material

and taking some of the images for the present

paper. Martins Kalnins (Institute of Life Sciences

and Technology, Daugavpils University, Latvia) is

thanked for sharing specimens, observations, and

images. Both Jean-Jacques Geoffroy and Jean-

Paul Mauriés kindly provided information on the

holotype of Julus blainvillei in the Paris Natural

History Museum, while Zoltan Korsds on the

types of Spirobolus dentatus Daday, 1893 in the

Budapest Natural History Museum.

Special thanks go to Jackson Means (Virginia

Natural History Museum, Martinsville, U.S.A.) for

his critical pre-review of an advanced draft.

The first author was supported by the

Presidium of the Russian Academy of Sciences,

Programme No 41 “Biodiversity of natural systems

and biological resources of Russia”.

References

Attems C. 1914a. Zur Fauna von Nord-Neuguinea.

Nach den Sammlungen von Dr. P. N. van Kampen

und K. Gjellerup aus den Jahren 1910 und 1911.

Myriopoden. - Zoologische Jahrbucher, Abteilung

02-Jun-21 21:48:01

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GoLovatcH, S. |., AKKARI, N., Goup, J. & TELNov, D.: Review of the Papuan genus Acanthiulus Gervais, 1844 (Diplopoda) ...

fur Systematik, Geographie und Biologie der Tiere

37: 380-384.

Attems C. 1914b. Die indo-australischen Myriopoden. -

Archiv fur Naturgeschichte 80A, No 4: 1-398.

Attems C. 1915. Myriopoden von Neu-Guinea Il.

Gesammelt wahrend der Expeditionen 1904-

1909. - Nova Guinea 13 (Zoologie 1): 1-38.

Attems C. 1927. Myriopoden von Ambon- und

anderen Inseln der Banda-See. - Zoologische

Mededeelingen, Rijks Museum van Natuurlijke

Historie 10: 61-70.

Attems C. 1932. Myriopoden. - In: Van Straelen V.

(Ed.), Résultats scientifiques du voyage aux Indes

Orientales Néerlandaises de LL. AA. RR. le Prince

et la Princesse Léopold de Belgique 3, No 12:

3-34.

Bollman C. H. 1893. The Myriapoda of North America.

- Bulletin of the United States National Museum

46: 1-210.

Bouvier E.-L. 1903. Acanthiulus Maindroni, Myriapode

nouveau de la famille de Spirobolidés. - Bulletin du

Muséum d’histoire naturelle 6: 263-297.

Brolemann H. W. 1903. Le genre Acanthiulus

(Myriapodes). - Annales de _ la _ Société

entomologique de France 72: 469-477.

Brodlemann H. W. 1913. The Myriapoda in the Australian

Museum. Part ii - Diplopoda. - Records of the

Australian Museum 10: 77-158.

Carl J. 1913. Diplopoden der Aru- und Kei-Inseln.

Ergebnisse einer zoologischen Forschungsreise in

densudostlichen Molukken (Aru- und Kai-Inseln)im

Auftrag der Senckenbergischen Naturforschenden

Gesellschaft, Band 2. - Abhandlungen

herausgegeben von der Senckenbergischen

Naturforschenden Gesellschaft 34: 269-279.

Chamberlin R.V. 1920. The Myriopoda of the Australian

Region. - Bulletin of the Museum of Comparative

Zoology at Harvard College 64, No 1: 1-269.

Daday J. 1893. Uj vagy kevéssé ismert idegenfdldi

myriapodak a magyar nemzeti Muzeum

gyujteményében. Myriopoda extranea nova vel

minus cognita in collectione Musaei nationalis

hungarici. — Természetrajzi Fuzetek 16: 98-113.

Enghoff H., Dohle W., Blower J. G. 1993. Anamorphosis

in millipedes (Diplopoda) - the present state

of knowledge with some developmental and

phylogenetic considerations. - Biological Journal

of the Linnean Society 109: 103-234.

Gervais P. 1844. Etudes sur les Myriapodes. - Annales

des Sciences naturelles 2 (Série 3: Zoologie): 51-

80.

Gervais P. 1847. Histoire naturelle des Insectes. Aptéres

4: i-xvi, 1-623.

Golovatch S. |., Wesener T. 2016. A species checklist of

the millipedes (Myriapoda, Diplopoda) of India. -

pp. 91-106

Zootaxa 4129, No 1: 001-075.

Hirst S. 1914. Report on the Arachnida and Myriopoda

collected by the British Ornithologists’ Union

Expedition and the Wollaston Expedition in Dutch

New Guinea. - Transactions of the Zoological

Society of London 20: 325-334.

Hoffman R. L., Keeton W. T. 1960. List of the generic

names proposed in the diplopod order Spirobolida,

with their type species. - Transactions of the

American Entomological Society 86: 1-26.

Jeekel C. A. W. 1971 (for 1970). Nomenclator generum

et familiarum Diplopodorum. - Monografieén van

de Nederlandse Entomologische Vereniging No 5,

412 pp.

Jeekel C. A. W. 2001. A bibliographic catalogue of the

Spirobolida of the Oriental and Australian regions.

- Myriapod Memoranda 4: 5-104.

Korsos Z. 1983. Diplopod types in the Hungarian Natural

History Museum, |. - Annales historico-naturales

Musei Nationales Hungarici 75: 117-120.

Latzel R. 1884. Die Myriopoden der Osterreichisch-

Ungarischen Monarchie. Zweite Halfte. Wien, 414

Leguillou E. J. F. 1841. Catalogue raisonné des Insectes

recueillis pendant le voyage de circumnavigation

des Corvettes |’Astrolabe et la Zélée. - L’Institut A.

9, No 399: 279-280.

Moritz M., Fischer S. C. 1975. Die Typen der Myriapoden-

Sammlung des Zoologisches Museums Berlin. |.

Diplopoda. Teil 3: Julida, Spirobolida. - Mitteilungen

aus dem Zoologischen Museum der Humboldt-

Universitat zu Berlin 51, No 2: 199-255.

Pocock R. |. 1893. Report upon the Myriopoda of the

‘Challenger’ Expedition, with remarks upon the

fauna of Bermuda. - Annals and Magazine of

Natural History, Series 6, 11: 121-142.

Pocock R. |. 1903. Remarks upon the morphology and

systematics of certain chilognathous diplopods. -

Annals and Magazine of Natural History, Series 7,

12: 515-532.

Porat O. von 1894. Zur Myriopodenfauna Kameruns.

- Bihang till Kungliga Svenska vetenskaps-

akademiens Handlingar 20, No 4: 3-90.

Preudhomme de Borre A. 1884. Tentamen

catalogi Lysipetalidarum, Julidarum, Archiulidarum,

Polyzonidarum atque Siphonophoridarum

hucusque descriptarum. - Annales de la Société

entomologique de Belgique 58: 46-82.

Silvestri F. 1895. Chilopodi e diplopodi della Papuasia. -

Annali del Museo civico di storia naturale di Genova

34, Series 2: 619-659.

Received: 21.xii.2020.

Accepted: 05.i.2021.

105

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An annotated locality list for morph A (circles):

(7) “Mimika River”, Mimika Regency, Papua, Indonesia,

about S4° 30’, E136 ° 30’; no accurate data are available

for this locality. (8) “Alkmaar”, nowadays a no longer

extant camp of the First South New Guinea Expedition

(1910), located at the place where the Lorentz River

becomes unnavigable, Papua, Indonesia, about S4° 40’,

E138° 43’; no accurate data are available for this locality.

(9) “Tanah Merah Bay”, west of Cyclops Mountains,

Sentani-Jayapura area, Papua, Indonesia, about S2°27’,

E140° 21’; no accurate data are available on this locality.

(10) “Hollandia”, now Port Numbai in Jayapura, Papua,

Indonesia, S2°32’, E140° 42’: this locality is now within

the central Jayapura and does not exist. (11) comprises

two published localities, “between biwak Bronbeek

and biwak Modderlust” (about S2°39’, E140°48’

and S2°41’, E140°50’) and “between bivak Kasawari

and bivak Modderlust” (about S2°37’, E140°35’ and

$2°41’, E140°50’), an area between the southern

coast of Yotefa Bay towards Koya Timur and Tami River,

Papua Indonesia; none of these three Dutch locality

names is in use anymore and the area is mostly covered

by settlements and rice fields, except few scattered still

forested patches on limestone hills. (12) Star Mountains,

13.5 km SSE of Oksibil, Papua, Indonesia, S4°59’33”,

E140°42’02”. (13) Jayapura 32 km SSE, near Indonesia

— Papua New Guinea border crossing point, northeast of

Bougainville Mountains, Papua, Indonesia, S2°37’21”,

E140°58’42”. (14) comprises two published localities,

“Bewani River” area and “bivak Zoutbron” (about S3°,

E141° and S2°41’, E140°59’), nowadays a no longer

existing camp at Begowre River, south of Bougainville

Mountains (on the Papua New Guinea border), Papua,

Indonesia. (16) “Kaiserin-Augusta River”, now Sepik,

East Sepik Province, Paonua New Guinea, about $4°12’,

E142°49’: the locality on the map is at the base camp

of the “Kaiserin-Augusta-Flu8-Expedition” of 1912/13

since no accurate data are available for this locality.

An annotated locality list for morph B (diamonds):

(15) “Umgebung von Kajo Bai, zwischen Njad und

Sekopo” are misspelt Dutch locality names of Njao

and Sekofro, in Sandaun (former West Sepik) Province,

Papua New Guinea, south of Bougainville Mountains

near the boundary with Indonesia, at the headwaters of

one of the right hand side tributaries of Tami River, both

nowadays uninhabited, about S2°40’, E141°01’; this

locality is about 8-10 km straight line from New Guinea’s

northern coast. (16) “Kaiserin-Augusta River”, now Sepik

River, East Sepik Province, Papua New Guinea, about

$4°12’, E142°49’; the locality on the map is at the

base camp of the “Kaiserin-Augusta-Flu8-Expedition” of

1912/13 since no accurate data are available for this

locality. (18) “Astrolabe Bay”, Madang Province, Papua

New Guinea, about S5°, E145°; no accurate data are

available for this locality.

106

Appendix 1. Annotated list of localities.

An annotated locality list for morph C (triangles):

(1) S Bird’s Neck Isthmus, 2-4 km NE of Kaimana, West

Papua, Indonesia, S3°39’, E133°46’. (2) Kaimana 47

km E, Triton Bay, environs of Kamaka (former Warika)

village, Surroundings of Lake Kamakawalar, West Papua,

Indonesia, S3°46’22”, E134°12’02”. (3) “Wammer:

Dobo”, Dobo, Wamar Island, Aru Islands, Indonesia,

about S5°45’, E134°13’. (4) “Wokam: Samang”,

Samang District on SW peninsula of Wokam Island,

Aru Islands, Indonesia, about S5°40’, E134°14’. (5)

“Manoemai, Wokam’”, misspelling for Manoembai village

on Manoembai River splitting Kobroor and Wokam, Aru

Islands, about S6°01’, E134°18’; no accurate data

are available for this locality. (6) “Teranhan: Ngaiguli”,

Ngaiguli, Trangan Island, Aru Islands, Indonesia, about

S6°, E134°. (16) “Kaiserin-Augusta River”, now Sepik

River, East Sepik Province, Papua New Guinea, about

$4°12’, E142°49’: the locality on the map is at the

base camp of the “Kaiserin-Augusta-Flu8-Expedition”

of 1912/13 since no accurate data are available for

this locality. (17) Bosavi Volcano, Western Highlands

Province, Papua New Guinea, about S6°43’, E152°43’.

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GoLovarcH, S. I.: Anew species of the genus Acanthiulus Gervais, 1844 (Diplopoda: Spirobolida: Pachybolidae) ...

pp. 107-110

A new species of the genus Acanthiulus Gervais, 1844

(Diplopoda: Spirobolida: Pachybolidae)

from Papua New Guinea

urn:lsid:zoobank.org:pub:185 /OA4E-4070-4198-86 7 D-ECED55E6522B

SERGE! I. GOLOVATCH

Institute for Problems of Ecology and Evolution, Russian Academy of Sciences, Leninsky prospekt

33, 119071, Moscow, Russia; sgolovatch@yandex.ru

Abstract: A new species, Acanthiulus subcostulatus sp. nov., is described from northern Papua New Guinea,

distinguished from the sole, and type species, A. blainvillei (Leguillou, 1841), polymorphous and widespread across

much of New Guinea and the Aru Archipelago, East Indonesia, by the smaller body (adults 8.5 vs >10 cm long), the

characteristically striate/subcostulate metaterga devoid of any teeth, cones or spines, and the gonopodal structure.

In particular, the sternum of the anterior gonopods is undivided distally (vs divided and diverging apically), and the

apical parts of the posterior gonopods are helmet-shaped (vs much more open and trifid).

Keywords: Millipede, taxonomy, iconography, New Guinea.

Introduction

The present contribution continues the previous

paper devoted to a review of the Papuan millipede

genus Acanthiulus Gervais, 1844 and contained

in the same volume (Golovatch et al. 2021). This

note puts on record a second, new species of

Acanthiulus, which has hitherto been considered a

monospecific genus. The sole, and type species, A.

blainvillei (Leguillou, 1841), is widespread across

much of New Guinea and the Aru Archipelago, East

Indonesia (Golovatch et al. 2021), whereas the new

congener comes from northern Papua New Guinea.

Since Acanthiulus has just been reviewed, and A.

blainvillei shown to be polymorphous (Golovatch

et al. 2021), there is no need to further detail the

status, diversity and distribution of the genus and

species.

Material and methods

The holotype of the new species was taken in

1977 by Galina F. Kurcheva during an expedition

on board the research vessel “Dmitry Mendeleyev”

to several islands and archipelagos in the

southwestern Pacific. It has been deposited in the

collection of the Zoological Museum of the Moscow

State University (ZMUM), Russia.

Images were taken with a Canon EOS 5D

digital camera and stacked using Zerene Stacker

software.

Results

Acanthiulus subcostulatus sp. nov. (Figs 1-15)

Holotype & ZMUM: Papua New Guinea, Maclay

Coast, near Kepoiak, S$5°45’, E146°35’, forest litter,

13.11.1977, G. F. Kurcheva leg.

Derivatio nominis: To emphasize the heavily

striate, subcostulate striations of metazonae

across their entire circumference.

Diagnosis: Differs clearly from A. blainvillei

(Leguillou, 1841), a polymorphous’ species

widespread across much of New Guinea and the

Aru Archipelago, East Indonesia (Golovatch et al.

2021), by the smaller body (adults 8.5 vs >10 cm

long), the characteristically striate/subcostulate

metaterga devoid of any teeth, cones or spines,

and the gonopodal structure, in particular, the

distally undivided sternum of the anterior gonopods

(vs divided and diverging apically) and the helmet-

Shaped apical parts of the posterior gonopods (vs

much more open and trifid).

Description: Body ca. 85 mm long, width or

height of midbody segments 7.2 mm, with 48p+T

segments. Colouration mainly red-brown, pattern

distinct and cingulate due to darker, red-brown

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

metazonae and much lighter, greyish to yellowish

pro- and mesozonae; antennae, legs and most of

head red-yellow; eye patches blackish brown (Figs

1-4). Body cylindrical, postcollum constriction

very faint (Fig. 2). Head as usual, bare, with three

small, but evident, central teeth at fore margin of,

and a short, axial and distinct suture on, labrum,

followed by a superficial, fine, axial, epicranial line,

with ca. 7-8+7-8 labral and 2+2 supralabral setae

(Fig. 2). Eye patches large and suboval (Fig. 1), each

composed of ca. 30 flat ommatidia arranged in six

vertical rows (ca. 4+6+6+6+5+4): interantennal

isthmus >2x diameter of eye patch (Fig. 2).

Antennae short and clavate, curved anteroventrad,

in situ stretchable laterally behind segment 2; in

length, antennomere2>3=4=5>6>1>/7;

distal end of antennomere 4 and antennomeres

5-7 clothed with dense and rather long setae; 8"

with four small apical cones (Figs 1-2). Tegument

nearly smooth to finely leathery, mostly shining

(Figs 1-4). Collum strongly narrowed _ laterad,

broadly rounded and clearly bordered anteriorly,

narrowly rounded and similarly bordered laterally,

gently concave in caudobasal third, very briefly

and slightly bordered at posterior margin (Fig.

1). Ventral corner of segment 2 hidden beneath

collum. Segments/rings 2-6 with increasingly

dense, clear, longitudinal, dorsal striations on

metazonae, these striations on following segments

being already deep, subcostulate, often irregular,

confused and caudally abbreviated (Figs 1-4).

108

Midbody segments/rings with faint sutures only

between meso- and metazonae, these regions

being very poorly constricted and slightly rugulose.

Mesozonae finely and mostly horizontally striolate

across entire circumference, only anteriorly

obliquely striolate dorsad, same as prozonae with

their particularly dense and vertical striolations.

Scobinae absent. Ozopores small, inconspicuous,

round disks lying just in front of and in touch

with suture/line between meso- and metazonae,

starting with segment 6 (Figs 1 & 4). Telson (Fig.

4) as usual, epiproct flat, small, digitiform, barely

projecting behind apical end of paraprocts; the

latter strongly and regularly convex, smooth, clearly

impressed before prominent caudal lips with a small

gutter between both valves; hypoproct nearly semi-

circular, strongly transverse, rounded caudally. Only

3d segment 7 clearly swollen ventrally, a complete

ring with a narrow bridge ventrally. No visible sigilla

on internal surface of meso- and metazonae.

Legs short and slender, more than half as long

as midbody height, each with a spine above claw,

2-4 ventral spines on tarsi, and one strong ventral

seta on each of other podomeres; ventral sole pads

absent; coxae 3-5 somewhat expanded ventrally

(4) (Fig. 2). Gonopods strongly sclerotized (Figs

5-15). Anterior gonopods (Figs 5-8) with a large

and subtriangular sternum showing a prominent,

high and slender central process (s) with its rather

narrowly rounded tip curved caudally, s being

only slightly higher than large, roundly squarish

Figures 1-4. Acanthiulus subcostulatus sp.

nov., 3 holotype. 1-2 - Anterior part of body,

lateral and ventral views, respectively; 3 -

Middle part of body, dorsal view; 4 - Caudal

part of body, lateral view [not to scale] {images

courtesy Kirill V. Makarov).

02-Jun-21 21:48:04

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GoLovarcH, S. I.: Anew species of the genus Acanthiulus Gervais, 1844 (Diplopoda: Spirobolida: Pachybolidae) ...

pp. 107-110

coxae (cx) with a little smaller and subapically at midline through a paramedian pair of strong,

bilobed telopodites (t), each of these being fully chitinized, subhorizontal, sternal halves (s), each

contained inside a hollow on the caudal face of hollow on anterior side, attached basally to a

cx. Posterior gonopods (Figs 9-15) connected partly membranous (m) tracheal apodeme (a), and

Figures 5-10. Acanthiulus subcostulatus

sp. nov., holotype 3. 5-6 - Sternum

and right anterior gonopod, anterior

and posterior views, respectively; 7-8

- Telopodite of left anterior gonopod,

anterior and posterior views, respectively;

9-10 - Left posterior gonopod, anterior

and posterior views, respectively [not to

scale] (images courtesy Kirill V. Makarov).

Designations explained in text.

Figures 11-15. Left posterior gonopod of

Acanthiulus subcostulatus sp. nov., holotype

3, anterior, mesal, posterior, anterior and

posterior views, respectively [11-13 not

to scale; scale bar for figures 14 & 15 - 2

mm] (images courtesy Kirill V. Makarov).

Designations explained in text.

& 109

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TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

supporting a large, boat-shaped and mesally hollow

coxa (cx), the latter clearly and regularly curved,

and showing a low, mesal and very simple lobe

(lo) with a small, midway, apically serrate, squarish

and membranous outgrowth (k) on top (apparently,

marking the end of a seminal groove), and a large,

helmet-shaped and rather complex apical part with

a very small, tongue-shaped and membranous

lobule (Ib) near base.

Differential diagnosis: The new species seems

best to fit within the scope of Acanthiulus, not in

the highly speciose and grossly sympatric genus

Trigoniulus Pocock, 1894. Thus, the collum has

very peculiar paraterga, i.e. long, subtriangular

and strongly drawn ventrad (Fig. 1), just like in

Acanthiulus blainvillei or Zygostroohus Chamberlin,

1920, a small genus from Australia (vs helmet-

Shaped, with short and broadly rounded paraterga

in all Trigoniulus except T. harpagus Attems,

1914). The latter species comes from western New

Guinea, first diagnosed in a key (Attems 1914),

but fully described and illustrated only a little

later (Attems 1917). The strongly subcostulate

metaterga in Acanthiulus subcostulatus sp. nov.

seem to be unique, more reminiscent of the often

ribbed interspaces between the cones, teeth or

spines in A. blainvillei, especially its morph B

(Golovatch et al. 2021), than the metaterga only

very seldom striolate dorsally as observed in

Trigoniulus, e.g., T. densestriatus Attems, 1897

from Borneo (Attems 1897). Usually, the dorsum

in Trigoniulus spp. is completely smooth (Attems

1914; 1932). In addition, neither Acanthiulus

blainvillei nor A. subcostulatus sp. nov. shows the

posterior gonopod equipped with a considerable

and ornamented mesal branch so characteristic

of most of Trigoniulus (Attems 1914; 1932) and its

immediate allies like Arisemolus Hoffman, 1980

(Golovatch et al. 2020). Exceptionally, however,

that branch is underdeveloped, e.g., in the dorsally

smooth Trigoniulus andropygus Attems, 1914,

again a species from western New Guinea which

was first diagnosed in a key (Attems 1914), but fully

described and illustrated only a little later (Attems

1917). That both Acanthiulus blainvillei and A.

subcostulatus sp. nov. come from New Guinea

and tend to be parapatric (near the Astrolabe Bay)

seems to be another argument, albeit an indirect

one, to treat them as being particularly close. The

majority of spirobolidan systematics, including the

Indo-Pacific Pachybolidae, remain badly confused

and chaotic (Jeekel 2001). Trigoniulus taxonomy,

with 80+ species involved, thereby seems to be

particularly imperfect (cf. https://en.wikipedia.or

110

wiki/Trigoniulus and Jeekel (2001)).

Acknowledgements

Special thanks go to Kirill V. Makarov

(Pedagogical State University, Moscow, Russia) for

the help in taking the pictures. | am most grateful

to Thomas Wesener (Zoological Museum, Bonn

University, Germany) for his critical comments on

an early draft of the paper. Jackson Means (Virginia

Natural History Museum, Martinsville, Virginia,

U.S.A.), very helpfully reviewed the manuscript.

The author was partly supported by the

Presidium of the Russian Academy of Sciences,

Program No 41 “Biodiversity of natural systems and

biological resources of Russia”.

References

Attems C. 1897. Myriopoden. In: Kukenthal W. (ed.)

Ergebnisse einer zoologischen Forschungsreise

in der Molukken und Borneo, im Auftrage der

Senckenbergischen naturforschenden Gesellschaft.

Teil 2: Wissenschaftliche Reiseergebnisse, Band

1. - Abhandlungen der Senckenbergischen

naturforschenden Gesellschaft 23, No 3: 473-536.

Attems C. 1914. Die indo-australischen Myriopoden. -

Archiv fur Naturgeschichte 80A, No 4: 1-398.

Attems C. 1917. Myriopoden von Neu-Guinea gesammelt

wahrend der Expedition 1903. - Nova Guinea.

Résultats de |’expédition scientifique néerlandaise

a la Nouvelle-Guinée en 1903 sous les auspices de

Arthur Wichman 5 (Zoologie), Livraison 6: 567-587.

Attems C. 1932. Myriopoden. In: Van Straelen_ V.

(ed.) Résultats scientifiques du voyage aux Indes

Orientales Néerlandaises de LL. AA. RR. le Prince et

la Princesse Léopold de Belgique 3, No 12: 3-34.

Golovatch S. I, Mauriés J.-P., Akkari N. 2020. On

the collections of Indo-Australian Spirobolida

(Diplopoda) kept in the Zoological Museum of the

Moscow State University, Russia. 1. A new species of

Arisemolus Hoffman, 1980 from Papua New Guinea.

- Arthropoda Selecta 29, No 3: 309-315.

Golovatch S. |., Akkari N., Goud J., Telnov D. 2021. Review

of the Papuan millipede genus Acanthiulus Gervais,

1844 (Diplopoda: Spirobolida: Pachybolidae):

91-106. In: Telnov D., Barclay M. V. L., Pauwels O.

S. G. (eds) Biodiversity, biogeography and nature

conservation in Wallacea and New Guinea. Volume

4, The Entomological Society of Latvia, Riga: 443 pp.

Jeekel C. A. W. 2001. A bibliographic catalogue of the

Spirobolida of the Oriental and Australian regions.

— Myriapod Memoranda 4: 5-104.

Received: 30.xii.2020.

Accepted: 10.i.2021.

02-Jun-21 21:48:11

Book4.indd 111

Gree, K.: New species and records of the Papuan Diplommatinidae (Caenogastropoda: Cyclophoroidea)

pp. 111-134

New species and records of the Papuan

Diplommatinidae (Caenogastropoda: Cyclophoroidea)

urn:lsid:zoobank.org:pub:81499D45-368B-4CE5-BB92-EO3AAC1F6EA7/

KRISTINE GREKE

Latvian National Museum of Natural History, K. Barona iela 4, LV-1050, Riga, Latvia;

k.greke@gmail.com

Abstract: Six species of Palaina O. Semper, 1865 (Diplommatinidae L. Pfeiffer, 1856) new to science are described

and illustrated from the Papuan biogeographical region, namely P. (s. |.) cenderawasih, P. (s. |.) evanescens, P. (s. |.)

kagainisi, P. (s. |.) pallgergelyi, P. (s. |.) sarmi and P. (s. |.) Saxatilis spp. nov. New distribution records are provided for

ten poorly known Palaina species. An updated key to the Papuan Diplommatinidae is presented.

Key words: Taxonomy, key, Palaina, Papuan Region, New Guinea, Biak, ecology.

Introduction

The Diplommatinidae L. Pfeiffer, 1856 are

minute (1-10 mm) terrestrial caenogastropod

Snails distributed in subtropical and tropical Asia

(SE Palaearctic and Oriental regions), north-

eastern Australia, the Papuan Region, Pacific

islands, and Central and South America (Egorov

2013). The diplommatinid fauna of the Papuan

biogeographical region and Wallacea was revised

recently by Greke (2017), and is considered highly

diverse and comprised of 121 species in the genera

Arinia H. et A. Adams, 1856 (five species), Diancta

Martens, 1864 (five species), Diplommatina s. l.

Benson, 1849 (22 species), MWoussonia O. Semper,

1865 (12 species), and Palaina O. Semper, 1865

(77 species) (Greke 2017). Approximately 98% of

the Papuan diplommatinid species are considered

local and regional endemics with restricted ranges

usually associated with limestone outcrops in

primary rainforest habitats (Greke 2017).

Recent material provided for current study by

the Natural History Museum in Erfurt (Germany)

and Michael Kesl (Prague, Czech Republic)

contains new data on several poorly known species

from the Central Moluccas, northern New Guinea,

and Biak Island. Six new species are described:

Palaina (s. |.) cenderawasinh, P. (s. |.) evanescens, P.

(s. |.) kagainisi, P. (s. |.) pallgergelyi, P. (s. |.) sarmi

and P. (s. |.) saxatilis spp. nov. As a result of this

study, the total number of Palaina species in the

Papuan Region and Wallacea reaches 83 and the

total number of diplommatinid species in Wallacea

and the Papuan Region - 12/7 species. New

distributional records of ten poorly known species

are presented. Scanning electron micrographs

(SEM) of shell sculpture, especially spiral raised

Striae, is presented for the first time for some of

discussed taxa.

Material and methods

All taxa are listed alphabetically, because a

phylogenetic arrangement is currently impossible.

All label text is reproduced without additions. All

labels are printed. Author’s comments are placed

in square brackets. Type specimens of new species

described in the present publication are provided

with an additional black-framed printed label

“Holotypus” or “Paratypus” on red paper.

Mostspecimens described below are preserved

in 99% ethanol. A Leica S6D stereomicroscope was

used for dissection and study. Shell images were

taken using a Canon EOS 77DSLR camera attached

to the stereomicroscope. CombineZP software was

used for image stacking (Hadley 2010). Further

image manipulations were done in GNU Image

Manipulation program (GIMP). Scanning electron

microscope (SEM) images were taken using SEM

provided by the Latvian University’s Faculty of

Biology. Shells were ultrasonically cleaned in

distilled water for 1-2 minutes, washed through

a graded alcohol series up to 99%, air dried and

placed dorsally onto conductive tape on aluminium

stubs. Shells were imaged using a Hitachi TM-

3000 (Hitachi High Technologies ®) SEM following

methods as described by Geiger et al. (2007).

aes

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

The internal lamellar system was exposed using

entomological pins (size 1 or 3 depending on the

dimensions of the shell).

The morphological terminology introduced

by Liew et al. (2014), supplemented by Neubert

& Bouchet (2015) and Greke (2017) is used for

descriptions. Shell size classes and measurements

follow Greke (2017) except that aperture diameter

was measured instead of peristome width. The

following measurements were taken for all Species:

Shell height, height of the ultimate whorl, maximum

Shell width, and maximum diameter of aperture.

Holotypes of the newly described species

are deposited at the Natural History Museum

(Naturkundemuseum) in mqavlan Germany.

Additional specimens are deposited in the Museum

Zoologicum Bogoriense (see details in the text).

The Papuan Region as considered here is a

zoogeographic term in the sense of Gressitt (1982),

Beehler et al. (1986), Riedel (2002), and Telnov

(2011).

Abbreviations and acronyms used in the text:

AD - maximum diameter of aperture;

env. — environs;

KGC - Collection Kristine Greke, Dzidrinas, Latvia;

H - shell height including the peristome;

HW - height of ultimate whorl;

Is. - island;

juv. - juvenile (not adult or subadult) specimen;

vill. - village or little settlement;

MBBJ - Research Center for Biology, Indonesian Institute

of Sciences, Cibinong, Indonesia;

MKC - Collection Michael Kesl, Prague, Czech Republic;

NME - Naturkundemuseum Erfurt, Germany.

Results

New records

Moussonia strubelli (0. Boettger, 1891)

New material: Indonesia, Maluku, Is. Ambon Halong,

under trees by the stream | 0035 21.1.2012 Igt.M.Kesl

[41 adult, MKC].

Palaina (s. |.) angulata O. Boettger, 1891

New material: Indonesia, Maluku, Is. Ambon Halong,

under trees by the stream | 0003 21.1.2012 Igt.M.Kesl

[5 adults, MKC].

Palaina (s. |.) ascendens (Martens, 1864)

New material: Indonesia, Maluku, Is. Ambon Halong,

under trees by the stream | OO02 21.1.2012 Igt.M.Kesl

[3 adults, MKC].

112

Palaina (s. |.) citrinella van Benthem Jutting,

1963 (Figs 1 & 14)

New material: INDONESIA E, New Guinea, Papua

Prov., Sentani 30 km W, road to Nimbokrang, River

Armo valley, 02°34’26"S, 140°14’21”E, 62-65 m,

312.111.2018, wet limestone rocks at riverbank [29 adults,

6 subadults, 8 juv., MBBJ, NME & KGC] (Map 1).

Measurements: Selected adult specimen H =

3.7 mm, W = 1.9 mm, HW = 2 mm, AD = 1mm.

Additional description: Columella glossy,

Straight, with no knob or callus. Operculum

derivative, with concentric ridges on outer surface.

Shell microstructure as in Fig. 14.

Ecology: Specimens sampled from wet, in part

mossy rocks and from leaf litter in river valley in

disturbed lowland (elevation 62-65 m) rainforest.

Distribution: Northern West New Guinea, Sentani

- Genjem area. First record since the original

description.

Palaina (s. |.) consobrina van Benthem Jutting,

1963 (Fig. 15)

New material: INDONESIA E, New Guinea, Papua

Prov., Jayapura 21 km SSE, Koya Tengah 1-2 km NNW,

02°37'57"S, 140°48’59”E, 50-80 m, 18.11.2018,

secondary lowland rainforest on limestone [NME & KGC,

2 adult & 6 juv. specimens] (Map 1).

Measurements: Selected adult specimen H =

4.5mm, W=3 mm, HW = 2.2 mm, AD = 3.4 mm.

Chirality: One of studied adult specimens is

dextral.

Additional description: Columellar knob large,

somewhat shifted towards base of columella,

turned outwardly in direction of apertural opening.

Shell microstructure as in Fig. 15.

Ecology: Specimens sampled from wet leaf litter

in disturbed lowland (elevation 50-80 m) rainforest

on a limestone ridge.

Distribution: Jayapura & Sentani areas, northern

West New Guinea. First record since the original

description, first record from E of Jayapura.

Considering peculiarities of the landscape in the

boundary area between Indonesian New Guinea

and Papua New Guinea (presence of West-to-East

directed limestone ridges), this species is likely to

occur in Papua New Guinea as well (Vanimo area in

particular).

Palaina (s. |.) cupulifera van Benthem Jutting,

1963 (Figs 2 & 16)

New material: INDONESIA E, Papua _ Prov.,

Cenderawasih Bay, Biak Is. E, Biak ~34 km ENE,

OT *:04'5075:" 1.36°22:08°E. #10-15-- Mice 222018;

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Grete, K.: New species and records of the Papuan Diplommatinidae (Caenogastropoda: Cyclophoroidea)

primary lowland rainforest [NME & KGC, 2 adult

specimens] (Map 2).

Measurements: Adult specimen H = 4 mm, W =

2.5mm, HW = 2.5 mm, AD = 2.2 mm.

Additional description: Shell microstructure as

in fig. 16.

Ecology: Specimens sampled from wet leaf litter

in a small remaining patch of primary lowland

rainforest (elevation 10-15 m) on limestone soil,

immediately near the mangrove zone.

Distribution: Eastern part of Biak Is.,

Cenderawasih Bay Islands. First record since the

original description, fist record from eastern tip of

Biak.

Palaina (s. |.) glabella van Benthem Jutting, 1963

(Figs 3 & 18)

New material: INDONESIA E, Papua _ Prov.,

Cenderawasih Bay, Biak Is. N, Biak ~53.5 km NNW,

OF 44°38" S;, 71:35 75255" E11 50-2608 mi 22 llL2Ors,

primary lowland rainforest on limestone rocks [NME &

KGC, 2 adult specimens]; INDONESIA E, Papua Prov.,

Cenderawasih Bay, Biak Is. N, Biak ~56 km NNW,

07422079; 13525159" E, +95-120%m,. 2212018;

primary lowland rainforest on limestone rocks [NME &

KGC, 2 adult & 1 juv. specimens] (Map 2).

Measurements: Selected adult specimen H =

3.6 mm, W = 2.25 mm, HW = 2 mm, AD = 1.7 mm

Additional description: Spiral raised striae

present, delicate, there are 4-5 ribs per 100 um

(Fig. 18). Columella glossy, straight, no knob or

callus present. Operculum derivative, with low

concentric ridges on outer surface.

Ecology: Specimens sampled from wet leaf litter

and limestone rock cavities in primary lowland

(elevation 95-260 m) rainforest on limestone soil.

Distribution: Southern (“Base Biak”, W of Sorido

(Soredo)) and northern Biak, Cenderawasih Bay

Islands. First records since the original description,

fist record from northern part of Biak.

Palaina (s. |.) inconspicua van Benthem Jutting,

1963 (Figs 4 & 19)

New material: INDONESIA EE, Papua _ Prov.,

Cenderawasih Bay, Biak Is. W, Biak ~16 km W, Urfu

Ville OL: O9° 1255.61 35555 Saks 3-15 mee VN 2O aS:

secondary lowland rainforest on coastal rocks [KGC, 1

adult specimen] (Map 2).

Additional description: Spiral raised striae

present, there are 3-6 very delicate spiral raised

Striae between every two stronger ones (Fig. 19),

there are 3-4 stronger and up to 10 very delicate

pp. 111-134

ribs per 100 um.

Ecology: Specimens sampled from wet leaf litter

and limestone rock cavities in primary lowland

(elevation 50-580 m) rainforest on limestone soil.

Distribution: Southern parts of Biak Is. and Owl

Is. S offshore Biak, Cenderawasih Bay Islands. First

record since the original description.

Palaina (s. |.) liliputana van Benthem Jutting,

1963 (Figs 5 & 22)

New material: INDONESIA E, Papua _ Prov.,

Cenderawasih Bay, Biak Is. N, Biak ~53.5 km NNW,

O° 4438'S, 6135°52'°557E, 150-260 =m. 22 2018;

primary lowland rainforest on limestone rocks [NME &

KGC, 14 adult specimens]; INDONESIA E, Papua Prov.,

Cenderawasih Bay, Biak Is. N, Biak ~60 km NNW,

Of43' 23875; (S52 AGSIES 50-5803 Mig 2? IIL QOAS.

primary lowland rainforest on limestone rocks [KGC,

2 adult specimens]; INDONESIA E, Papua Prov.,

Cenderawasih Bay, Biak Is. N, Biak ~56 km NNW,

0° 42’40"S,- 135°51'59"E, 95-120 -m,. 22.2018,

primary lowland rainforest on limestone rocks [MBBJ,

NME & KGC, 66 adult & 2 juv. specimens] (Map 2).

Additional description: Shell microstructure as

in Fig. 22.

Ecology: Specimens sampled from wet leaf litter

and limestone rock cavities in primary lowland

(elevation 50-580 m) rainforest on limestone soil.

Distribution: Southern and northern parts of

Biak Is., Cenderawasih Bay Islands. First records

since the original description.

Palaina (s. |.) repandostoma van Benthem Jutting,

1963 (Fig. 23)

New material: INDONESIA E, Papua _ Prov.,

Cenderawasih Bay, Biak Is., Biak ~141 km E, at Biak Bird

park, 01° 10’39"S, 136° 10’33”E, 40-70 m, 21.1I1.2018,

primary lowland rainforest on limestone [MBBJ, NME &

KGC, 28 adult specimens] (Map 2).

Measurements: Selected adult specimens H = 3

mm, W = 2 mm, HW = 1.5 mm, AD = 1.5 mm; H =

2.9mm, W = 1.9 mm, HW = 1.7mm, AD = 1.5 mm.

Additional description: Shell microstructure as

in fig. 23.

Ecology: Specimens sampled from wet leaf litter

in primary lowland (elevation 40-70 m) rainforest

at foot of a steep limestone ridge.

Distribution: Southern (“Base Biak”, W of Sorido

(Soredo)) part of Biak Is., Cenderawasih Bay

Islands. First record since the original description.

113

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

New descriptions

Palaina (s. |.) cenderawasih sp. nov. (Figs 6, 12-

13)

http://zoobank.org/EF29B6A0-C1B5-4FB8-91CC-

212F71508C9B

Holotype NME: INDONESIA’ E, Papua _ Prov.,

Cenderawasih Bay, Biak Is., Biak ~11 km E, at Biak Bird

park, 01° 10'39”S, 136° 10’33’E, 40-70 m, 214.111.2018,

primary lowland rainforest on limestone.

Paratypes 2/7 specimens: 9 adult specimens [1

MBBJ, 3 NME & 5 KGC] same data as holotype [one

paratype shell badly broken]; 16 adult & 2 juv. soecimens

[11 NME, 5 adults & 2 juv. KGC] INDONESIA E, Papua

Prov., Cenderawasih Bay, Biak Is. W, Biak ~16 km W, Urfu

VII M01": 09" 12"S. 9135" 5553 Ec Salou, 24 sIbh2Oae:

secondary lowland rainforest on coastal rocks.

Derivatio nominis: Named after Cenderawasih

Bay of New Guinea, where Biak Island (locus

typicus) is situated. In Indonesian “cenderawasih”

means Bird of Paradise, a group of iconic New

Guinean birds. Noun in apposition.

Measurements: Holotype H = 2.5 mm, W =

1.5 mm, HW = 1.5 mm, AD = 1.1 mm. Selected

paratypes H = 2.5 mm, W = 1.5 mm, HW = 1.35

mm, AD = 1.05 mm; H = 2.4 mm, W = 1.5 mm, HW

= 1.4 mm, AD = 1mm; H = 2.4 mm, W = 1.55 mm,

HW = 1.5 mm, AD = 1.1 mm.

Description: Shell small, pale pink to yellowish,

sinistral, subcylindrical with prominent (projecting)

apex. Shell of 5 moderately strongly convex whorls,

1% embryonic whorls are microscopically pitted,

pits arranged in irregular axial rows. Suture is

deeply impressed. Ultimate whorl is slightly wider

than penultimate whorl in apertural view. Umbilicus

is closed in adults. Constriction is moderately well

defined by denser axial rioS and a deep impression

onsuture above constriction. Position of constriction

is above columellar side of aperture. Teleoconch is

sculptured with moderately widely spaced nearly

straight axial ribs. Ribs are twice as dense around

constriction, but gradually becoming more distant

(up to twice as distant as on earlier whorls) on

last half whorl, without abrupt changes in ribbing

pattern. Ribs are not synchronous with those on

previous whorls and are slightly oblique to coiling

axis. There are 6-7 ribs per 0.5 mm on penultimate

whorl in abapertural view. Spiral raised striae of

teleoconch very delicate; at 500x magnification per

100 um there are 1-12 rows of spiral raised striae

present on penultimate whorl but only 4-5 on

ultimate whorl before constriction area. Aperture is

circular, tilted about 10° to coiling axis, apertural

rim is entire. Aperture is slightly shifted left of

114

coiling axis in apertural view. Peristome is double,

in lateral view narrow. Outer peristome is formed

by two nearly same strongly developed lamellae

of previous peristomes (peristome therefore looks

triple in lateral view). Peristome is complex, outer

peristome looks double since promiment lamellar

edge of previous peristome is narrowly attached

to it. Outer peristome slightly expanding beyond

inner peristome. Parietal margin of peristome

is attached to spire. Inner peristome forms a

narrow, continuous, polished callus, is entire and

appressed to penultimate whorl. Outer peristome

is discontinuous along parietal margin. Margins of

peristome are not sinuous, or in some paratypes

only palatal margin of outer peristome is slightly

sinuous. Columella is smooth and straight, without

knob or callus. Operculum is derivative, with low

concentric ridges on outer surface.

Sexual dimorphism: Dimorphism is remarkable

in this species, female shell more globose and less

elongate than male.

Differential diagnosis: The new species is most

similar to Palaina glabella van Benthem Jutting,

1963 (Biak Is.), but is readily differentiated by its

smaller shell size (H = 3.4-3.9 mm in P. glabella),

the sparser spiral raised striae (up to 5-6 spiral

raised striae per 100 um at 300x magnification in

P. glabella) with the two kinds of the spiral raised

striae - stronger sparse and slightly denser (only

one kind of the spiral raised striae in P. glabella),

the axial ribs significantly denser at the constriction

area (notin P. glabella), the outer peristome consists

of two equally developed lamellae of the previous

peristomes (the peristome double in P. glabella,

the outer peristome consists of one or two merged

lamellae), all margins of the outer peristome nearly

equally strongly developed (the columellar and the

palatal margins of the outer peristome stronger,

somewhat protruding from the generally circular

outline of the peristome in P. glabella).

Ecology: Specimens sampled from wet leaf litter

in primary lowland (elevation 40-70 m) rainforest

at foot of a steep limestone ridge and from cavities

in coastal limestone rocks (elevation 3-15 m).

Distribution: South of Biak Island (Map 2).

Palaina (s. |.) evanescens sp. nov. (Figs 7 & 17)

http://zoobank.org/5CA18249-0F99-4730-954C-

DOD580ECC94B

Holotype NME: INDONESIA E, New Guinea, Papua

Prov., Jayapura 21 km SSE, Koya Tengah 1-2 km NNW,

O25 37 57'S, TAO" 46'592E_ 250-804 om Lemk2018"

secondary lowland rainforest on limestone.

Paratype 1 adult KGC: same data as holotype.

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Gree, K.: New species and records of the Papuan Diplommatinidae (Caenogastropoda: Cyclophoroidea)

Derivatio nominis: Named from Latin

“evanescere” [disappear, vanish, evaporate] to

highlight that native habitats at the type locality of

this species are Surrounded and are being replaced

by expanding human settlements.

Measurements: Holotype H = 3.8 mm, W = 2.5

mm, HW = 2.2 mm, AD = 1.7 mm. Paratype H = 3.7

mm, W = 2.3 mm, HW = 1.7 mm, AD = 1.55 mm.

Description: Shell is sinistral, small, whitish with

pale orange to brownish apical whorls, and broadly

cylindrical with prominent (projecting) apex. Shell

with 672 strongly convex whorls, 1% embryonic

whorls are microscopically pitted. Suture is deeply

impressed. Ultimate whorl is about as wide as

penultimate whorl in apertural view. Umbilicus

is closed in adult. Constriction is well defined

by presence of a vague impression on shell wall,

denser pattern of axial ribs, and presence of an

impression on suture above constriction. Position

of constriction is above columellar side of aperture.

Teleoconch is sculptured with moderately widely

Spaced and strongly sinuous axial ribs. Ribs

abruptly become coarser and twice as distant on

last half whorl beyond constriction. Ribs are not

synchronous with those of previous whorls and are

slightly oblique to coiling axis. There are 6-7 ribs

per 0.5 mm on penultimate whorl in abapertural

view. Spiral raised striae not observed at 300x

magnification. Aperture is not tilted to coiling axis,

is circular, with entire apertural rim. Aperture

is shifted left of coiling axis in apertural view.

Peristome is simple in lateral view. Parietal margin

of peristome is attached to spire. Inner peristome

forms narrow, continuous, polished callus, is

entire and appressed to penultimate whorl. Outer

peristome discontinuous along parietal margin.

Peristome margins broadly expanded, not sinuous

except slightly sinuous palatal margin. Columella

is smooth and glossy, without knob or callus.

Operculum unknown.

Differential diagnosis: Palaina evanescens

sp. nov. is readily differentiated from its Papuan

congeners by the combination of the following

features: the peristome is simple, the aperture

shifted left to the coiling axis, the axial ribs becoming

twice as distant beyond the constriction on the last

half whorl, the axial ribs distinctly sinuous, the

columella simple. For similar species see modified

key below.

Ecology: Specimens sampled from wet leaf litter

in disturbed lowland (elevation 50-80 m) rainforest

on a limestone ridge.

Distribution: Northern New Guinea (Map 1).

pp. 111-134

Palaina (s. |.) kagainisi sp. nov. (Figs 8, 20-21)

http://zoobank.org/B/73DD905-488E-4996-AF8B-

7E6E2CE2DFBE

Holotype NME: INDONESIA’ E, Papua _ Prov.,

Cenderawasih Bay, Biak Is. N, Biak ~53.5 km NNW,

0° 44'38'S, 135°52'55"E, 150-260 = m,. 22.11.2018,

primary lowland rainforest on limestone rocks.

Derivatio nominis: Patronymic. This species

named after Dr. Ugis Kagainis (University of Latvia

Faculty of Biology, Riga), well-known acarologist

and collaborative colleague, who is specialized on

advanced microscopy methods.

Measurements: Holotype H = 2.3 mm, W = 1.4

mm, HW = 1.4 mm, AD = 0.75 mm.

Description: Shell small, white, — sinistral,

Subcylindrical with flattened apex. Shell of five

strongly convex whorls, first embryonic whorl is

microscopically pitted. Suture is deeply impressed.

Ultimate whorl is slightly wider than penultimate

whorl in apertural view. Umbilicus is closed in

adults. Constriction is strongly defined by a denser

pattern of axial ribs and presence of a deep

impression on suture above constriction. Position

of constriction is above parietal side of aperture.

Teleoconch is sculptured with widely spaced,

nearly straight, high axial ribs. Ribs become twice

as dense around constriction, but thereafter rapidly

becoming as widely spaced as before constriction.

There are abrupt changes in ribbing pattern at

constriction area. Ribs are not synchronous with

those on previous whorls and are slightly oblique

to coiling axis. Ribs increase in height towards

suture. There are ~4 ribs per 0.5 mm (or 8-9 per

1 mm) on penultimate whorl in abapertural view.

Spiral raised striae of teleoconch very delicate

and barely visible at 500x magnification; there are

about 16-17 delicate spiral raised striae per 100

um on penultimate whorl. Aperture is not tilted to

coiling axis, is circular and with entire apertural rim.

Position of aperture is nearly centered in apertural

view. Peristome is double; outer peristome consists

of three or more nearly equally strongly developed

lamellae of previous peristomes, therefore

peristome looks multilamellate in lateral view.

Outer peristome is more strongly developed than

inner peristome. Parietal margin of peristome is

attached to spire. Inner peristome forms a narrow

continuous polished callus. Inner peristome entire,

appressed to penultimate whorl, outer peristome

discontinuous along parietal margin. Margins of

peristome are not or slightly sinuous. Columella is

smooth and straight, widened towards base, without

knob or callus. Operculum derivative, with 4-5 low,

115

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

concentric ridges and somewhat impressed central

area on outer surface.

Differential diagnosis: The new species is most

close to Palaina glabella van Benthem Jutting,

1963 (Biak Is.), but is readily differentiated by the

smaller shell size (H = 3.4-3.9 mm in P. glabella),

the significantly denser axial ribbing pattern at the

constriction area, the nearly absent spiral raised

Striae, the multilamellate peristome (the peristome

is double in P. glabella), the nearly equally developed

margins of the outer peristome (the columellar and

the palatal margin of the outer peristome stronger

developed, somewhat protruding from generally

circular outline of the peristome in P. glabella).

Ecology: Specimens sampled from cavities of

limestone rock in primary lowland (elevation 150-

260 m) rainforest on limestone.

Distribution: Southern part of Biak Island (Map

Palaina (s. |.) pallgergelyi sp. nov. (Fig. 9)

http://zoobank.org/A/7228052-0184-4030-B2/C-

14F/7009BCDOB

Holotype NME: INDONESIA’ E, Papua _ Prov.,

Cenderawasih Bay, Biak Is. N, Biak ~53.5 km NNW,

0°44'38"S, 135°52'55’E, 150-260 m, 222.111.2018,

primary lowland rainforest on limestone rocks [shell is

damaged as on the images].

Derivatio nominis: Patronymic. This species

honours Barna Pall-Gergely (Department of Biology,

Shinshu University, Japan), famous malacologist

and collaborative colleague, for his work in

malacology.

Measurements: Holotype H = 4.4 mm, W = 2.6

mm, HW = 2.2 mm, AD = 1.7 mm.

Description: Shell large, colourless (discoloured

old shell), sinistral, fusiform with prominent

(projecting) apex. Shell of six moderately strongly

convex whorls, first 1% embryonic whorls are

microscopically pitted. Suture is moderately deeply

impressed. Ultimate whorl is slightly narrower than

moderately bulbous penultimate whorl in apertural

view. Umbilicus is closed in adults. Exact position

of constriction is not determined (the only available

Shell is damaged). Shell with moderately strong

(obsolete angulate) bulb beyond presumable

area of constriction. Post-constriction area with

denser pattern of axial ribs. Presumable position of

constriction is above columellar side of aperture.

Teleoconch is sculptured with dense, strongly

sinuous (on older whorls) to nearly straight (part

of penultimate and ultimate whorls) axial ribs.

Ribs becoming almost twice as dense between

—_—_

it i

116

constriction and bulb, but gradually becoming more

distant (up to twice as distant as on earlier whorls)

on last half whorl, without abrupt changes in ribbing

pattern. Ribs are not synchronous with those of

previous whorls and are strongly oblique to coiling

axis. There are about 9-12 ribs per 0.5 mm on

penultimate whorl in abapertural view. Spiral raised

striae or ribs not present. Aperture is not tilted

to coiling axis, is circular. Apertural rim appears

entire. Aperture is slightly shifted left of coiling axis

in apertural view. Peristome is double, in lateral

view moderately broad and with several delicate,

dense previous peristomes. Outer peristome Is less

strongly developed than inner one. Parietal margin

of peristome is attached to spire. Inner peristome

forms a narrow, continuous, polished callus. Outer

peristome is not entire, broadly discontinuous

parietally. Columellar margin of peristome slightly

sinuous. Columella is smooth and straight with

large, glossy median knob. Operculum primitive,

slightly impressed on outer surface, without ridges.

Differential diagnosis: Hitherto the largest

diplommatinid species on Biak, P. pallgergelyi sp.

nov. is most similar to P. consobrina van Benthem

Jutting, 1963 (Biak Is.) but readily distinguishable

from congeners due to the combination of the

following features: the shell H over 4 mm, the

axial ribbing pattern dense, becoming only slightly

more distant on the last half whorl, the axial ribs

strongly sinuous on the older whorls but nearly

straight on the last two whorls, the outer peristome

is less strongly developed than the inner one, the

columellar knob is large and positioned medially,

the operculum primitive.

Ecology: Specimen sampled from cavities of

limestone rock in primary lowland (elevation 150-

260 m) rainforest on limestone.

Distribution: Northern part of Biak Island (Map

2),

Palaina (s. |.) sarmi sp. nov. (Figs 10 & 24)

http://zoobank.org/BOA0/027-693A-4FB1-990E-

2D4B53E17A

Holotype NME: INDONESIA E, New Guinea, Papua

Prove sari Ls) kine SSEs01° 5871975 -138e51°24E,

50-65 m, 27.III.2018, primary lowland rainforest on

limestone.

Paratype 1 subadult KGC: same data as holotype [the

paratype shell is broken, apertural part missing].

Derivatio nominis: Named after Sarmi, the main

town and the language family in the area, where

this species was first collected. Noun in apposition.

Measurements: Holotype H = 2.6 mm, W = 1.4

mm, HW = 1.3 mm, AD = 1.1 mm.

02-Jun-21 21:48:12

Book4.indd 117

Gree, K.: New species and records of the Papuan Diplommatinidae (Caenogastropoda: Cyclophoroidea)

Description: Shell small, light yellow with darker

yellow protoconch, sinistral, broadly fusiform with

prominent projecting apex. Shell of 572 moderately

strongly convex whorls, 172 embryonic whorls are

microscopically pitted. Suture is deeply impressed.

Ultimate whorl is about as wide as penultimate

whorl in apertural view. Umbilicus is closed in adults.

Constriction is defined by presence of a vague

impression on shell wall, somewhat denser pattern

of axial ribs and presence of a deep impression on

suture above constriction. Constriction is slightly

shifted to columellar side of aperture. Teleoconch

is sculptured with moderately widely spaced,

straight to slightly sinuous axial ribs. Ribs become

coarser and more distant on last half whorl, but

without abrupt changes in ribbing pattern. Ribs are

generally not synchronous with those on previous

whorls and are rather strongly oblique to coiling

axis. There are 5-6 ribs per 0.5 mm on penultimate

pp. 111-134

whorl in abapertural view. Spiral raised striae

moderately strong on earlier whorls of teleoconch;

4-5 rows of spiral raised striae present per 100

um at 300x magnification. Aperture is not tilted to

coiling axis, is circular, with entire apertural rim.

Aperture is more or less centered to coiling axis

in apertural view. Peristome is double, in lateral

view consisting of 1-2 stronger and more distant

and several dense, delicate previous peristomes.

Parietal margin of peristome is attached to spire.

Inner peristome forms narrow continuous polished

callus. Inner peristome entire, appressed to

penultimate whorl, outer peristome discontinuous

along parietal margin. Palatal margin of inner

peristome is sinuous. Columella is smooth, with

small glossy median knob. Operculum unknown.

Differential diagnosis: This species is rather

Similar to several congeners, differential features

as in Table 1.

Table 1. Papuan species of Palaina s. |. similar to P sarmi sp. nov. and their distinctive features.

[Species ——“ s—“‘s;é‘d Feature — —“(SCidC

P. albrechti Greke, 2017 |H = 1.25-1.3 mm, W = 0.7-0.75 mm;

axial ribs dense, ~17-19 ribs per 0.5

mm in abapertural view; more than 25

rows of spiral raised striae per 100 um;

peristome multilamellate (described as

Feature in P. sarmi sp. nov.

H = 2.6 mm, W = 1.4 mm; axial ribs less

dense, ~5-6 ribs per 0.5 mm in abapertural

view; about 4-5 rows of spiral raised striae

per 100 um; peristome double; columella

with small knob

"triple" in Greke (2017)); columella with

inconspicuous median angulation

P. beilanensis Preston,

1913

H =

P. biroi Sods, 1911

H = 1.8-1.9 mm, W = 0.8 mm; ~15 ribs

per 0.5 mm in abapertural view; peristome

triple; peristome not sinuous; distribution

- Obi Islands (North Moluccas

1.3-1.6 mm, W = 0.8-0.9 mm;

abrupt changes in axial ribbing pattern of

teleoconch at place of constriction; axial

ribs dense, ~12-16 ribs per 0.5 mm in

abapertural view; more than 15 rows of

H=2.6 mm, W = 1.4 mm; ~5-6 ribs per 0.5

mm in abapertural view; peristome double;

palatal margin of inner peristome sinuous;

distribution — North New Guinea

H=2.6 mm, W= 1.4 mm; no abrupt changes

in axial ribbing pattern of teleoconch; ~5-6

ribs per 0.5 mm in abapertural view; about

4-5 rows of spiral raised striae per 100 um;

columella with small knob

Spiral raised striae per 100 um; columella

with inconspicuous median ang

H = 1.5-1.7 mm, W = 0.8 mm; axial ribs

nearly straight; axial ribs dense, ~16-18

ribs per 0.5 mm in abapertural view; more

than 27 rows of spiral raised striae per

P. flavocylindrica Greke,

2017

100 um; columella simple

P. iha Greke, 2017 H = 1.35-1.4 mm, W

constriction area is free of axial ribs; axial

ribs dense, ~14-16 ribs per 0.5 mm in

abapertural view; more than 24 rows of

Spiral raised striae per 100 um; outer

peristome more strongly developed than

inner one; columella simple

ulation

H = 2.6 mm, W = 1.4 mm; axial ribs sinuous

at least on their upper and lower edges;

~5-6 ribs per 0.5 mm in abapertural view;

about 4-5 rows of spiral raised striae per

100 um; columella with small knob

H = 2.6 mm, W = 1.4 mm; constriction

area with somewhat denser axial ribs than

surrounding area; ~5-6 ribs per 0.5 mm

in abapertural view; about 4-5 rows of

spiral raised striae per 100 um; both inner

and outer peristomes similarly strongly

developed; columella with small knob

0.7 mm;

TAF

02-Jun-21 21:48:12

Book4.indd 118

TeLNnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

P. insulana Greke, 2017

P. lengguru Greke, 2017

P. minuscularia Greke,

2017

P. psittricha Greke, 2017

P. waigeo Greke, 2017

H = 1.8-1.95 mm, W = 0.95-1 mm;

axial ribS are much coarser and twice

as widely spaced on penultimate and

ultimate whorls; ~8-9 ribs per 0.5 mm

in abapertural view; ~20 rows of spiral

raised striae per 100 um; peristome

multilamellate (described as "triple" in

Greke (2017)); columella simple

H = 1.3-1.4 mm, W = 0.75-0.8 mm;

abrupt changes in axial ribbing pattern of

teleoconch - ribs twice as distant starting

from constriction; ~13-15 ribs per 0.5

mm in abapertural view; peristome thin

in lateral view; palatal margin of inner

peristome not sinuous; columella simple

= 1.3-1.8 mm, W = 0.7-0.9 mm;

constriction is poorly defined by weak

sutural impression above it; ~25-28 ribs

per 0.5 mm in abapertural view; ~12 rows

of spiral raised striae per 100 um; palatal

margin of inner peristome not sinuous;

columella with small basal knob

1.65-1.8 mm, W = 0.9-1 mm;

ultimate whorl is slightly narrower than

penultimate in apertural view; abrupt

changes in axial ribbing pattern of

teleoconch - ribs become twice as distant

beyond constriction; axial ribs dense, ~9-

10 ribs per 0.5 mm in abapertural view;

more than 20 rows of spiral raised striae

per 100 um; aperture is slightly shifted

left to coiling axis in apertural view;

operculum is derivative - high (tuba -like),

with several layers of concentric lines on

outer surface

H = 1.3-1.35 mm, W = 0.7 mm; axial

ribs dense, ~17-19 ribs per 0.5 mm in

abapertural view; more than 15 rows

of spiral raised striae per 100 um; no

columellar knob present

Table 1 (continuation)

H =2.6 mm, W= 1.4 mm; axial ribs not twice

as distant on last whorls; ~5-6 ribs per 0.5

mm in abapertural view; 4-5 rows of spiral

raised striae per 100 um; peristome double;

columella with small knob

H=2.6 mm, W= 1.4 mm; no abrupt changes

in axial ribbing pattern of teleoconch;

~5-6 ribs per 0.5 mm in abapertural view;

peristome moderately broad in lateral view;

palatal margin of inner peristome sinuous;

columella with small knob

H=2.6 mm, W = 1.4 mm; constriction is well

defined by presence of a vague impression,

somewhat denser pattern of axial ribs

and presence of a deep impression on

suture above it; ~5-6 ribs per 0.5 mm in

abapertural view; about 4-5 rows of spiral

raised striae per 100 um; palatal margin

of inner peristome sinuous; columella with

small median knob

H = 2.6 mm, W = 1.4 mm; ultimate whorl

nearly as wide as penultimate in apertural

view; no abrupt changes in axial ribbing

pattern of teleoconch; axial ribs less dense,

~5-6 ribs per 0.5 mm in abapertural view;

about 4-5 rows of spiral raised striae per

100 um; aperture is centered to coiling axis

in apertural view; operculum unknown

H =2.6 mm, W = 1.4 mm; axial ribs sparser,

~5-6 ribs per 0.5 mm in abapertural view;

about 4-5 rows of spiral raised striae per

100 um; columellar knob present

Ecology: Specimens sampled from wet leaf litter

In primary lowland (elevation 50-65 m) rainforest

ona steep limestone ridge.

Distribution: Northern New Guinea (Map 1).

Palaina (s. |.) saxatilis sp. nov. (Figs 11, 25-26)

http://zoobank.org/ 7 D36EDCC-6D8D-48B8-A17A-

5OFIEBABBO27

Holotype NME: INDONESIA E, New Guinea, Papua

Prov., Sentani 30 km W, road to Nimbokrang, River

Armo valley, 02°34’26”"S, 140°14’21”E, 62-65 m,

31.11.2018, wet limestone rocks at riverbank.

Paratypes 30 specimens [5 MBBJ, 13 NME & 12

KGC]: same data as holotype [one paratype shell is badly

118

broken].

Derivatio nominis: Named from Latin “saxatilis”

(living on rocks, rock-dwelling) since this species

was Sampled from mossy rocks on a riverbank.

Measurements: Holotype H = 1.7 mm, W =

0.95 mm, HW = 0.9 mm, AD = 0.5 mm. Selected

paratypes H = 1.6 mm, W=0.8 mm, HW = 0.8 mm,

AD = 0.55 mm; H = 1.65 mm, W = 0.7 mm, HW =

0.6 mm, AD = 0.45 mm; H = 1.65 mm, W = 0.9

mm, HW = 0.9 mm, AD = 0.5 mm.

Description: Shell very small, white, pale yellow

(on older whorls) to pale brownish, sinistral,

elongate cylindrical with flattened apex. Shell of six

02-Jun-21 21:48:13

Book4.indd 119

Gree, K.: New species and records of the Papuan Diplommatinidae (Caenogastropoda: Cyclophoroidea)

moderately strongly convex whorls, 17/2 embryonic

whorl is microscopically pitted. Suture is deeply

impressed. Ultimate whorl is about as wide as

penultimate whorl in apertural view. Umbilicus is

closed in adults. Constriction is vaguely defined

by presence of slight impression on shell wall and

more deeply impressed suture above constriction.

Position of constriction is above columellar side of

aperture. Teleoconch is sculptured with moderately

dense, straight axial ribs. Ribs become coarser

and 2-2.5x more distant on last half whorl, but

without abrupt changes in ribbing pattern. Ribs

are not synchronous with those on previous whorls

and are slightly oblique to coiling axis. There are

10-12 ribs per 0.5 mm on penultimate whorl in

abapertural view. Spiral raised striae of teleoconch

delicate and dense, 10-12 rows of spiral raised

striae present per 100 um at 300x magnification.

Aperture is not tilted to coiling axis, is circular and

with entire apertural rim. Aperture is more or less

pp. 111-134

centered to coiling axis in apertural view. Peristome

is double, in lateral view consisting of 2-4 rather

strong and densely placed previous peristomes.

Parietal margin of peristome is attached to spire.

Inner peristome forms narrow, continuous, polished

callus. Outer peristome is broadly discontinued

parietally. Margins of both inner and _ outer

peristomes are not sinuous. Columella is smooth,

with large median knob, which is curved interiad.

Operculum derivative, with low concentric ridges on

outer surface.

Differential diagnosis: This is one of the

smallest diplommatinids in the Papuan Region

and Wallacea. The new species is rather similar to

several congeners, differential features as in Table

Ecology: Specimens sampled from wet, mossy

rocks and from leaf litter in river valley, in disturbed

lowland (elevation 62-65 m) rainforest.

Distribution: Northern New Guinea (Map 1).

Table 2. Papuan species of Palaina s. |. similar to P. saxatilis sp. nov. and their distinctive features.

Feature in P. saxatilis sp. nov.

P. albrechti Greke, 2017

P. beilanensis Preston,

1913

P. biroi Sods, 1911

P. flavocylindrica Greke,

2017

Paincaorekes 20 ley

P. insulana Greke, 2017

H = 1.25-1.3 mm, W = 0.7-0.75 mm;

axial ribs dense, ~17-19 ribs per 0.5

mm in abapertural view; more than 25

rows of spiral raised striae per 100 um;

peristome multilamellate; columella with

inconspicuous median angulation

shell slender and more regularly cylindrical;

~15 ribs per 0.5 mm in abapertural view;

peristome multilamellate (outer peristome

consists of two nearly same strong lamellae

of previous peristomes); columella not

studied; distribution - Obi Islands (North

abrupt changes in axial ribbing pattern of

teleoconch at place of constriction; axial

ribs dense, ~12-16 ribs per 0.5 mm in

abapertural view; more than 15 rows of

Spiral raised striae per 100 um; columella

with inconspicuous median angulation

axial ribs dense, ~16-18 ribs per 0.5 mm

in abapertural view; more than 27 rows of

Spiral raised striae per 100 um; columella

simple

H = 1.35-1.4 mm, W = 0.7 mm; constriction

area is free of axial ribs; axial ribs dense,

~14-16 ribs per 0.5 mm in abapertural

view; more than 24 rows of Spiral raised

striae per 100 um; outer peristome is

more strongly developed than inner one;

columella simple

axial rips becoming much coarser and twice

as distant on penultimate and ultimate

whorls; ~8-9 ribs per 0.5 mm in abapertural

view; ~20 rows of spiral raised striae

per 100 um; peristome multilamellate;

columella simple

H = 1.6-1.7 mm, W = 0.8-0.95 mm;

~10-12 ribs per 0.5 mm in abapertural

view; ~10-12 rows of spiral raised striae

per 100 um; peristome double; columella

with conspicuous median knob

Shell less slender, tapered to apex;

~10-12 ribs per 0.5 mm in abapertural

view; peristome double; columella with

conspicuous median knob; distribution —-

North New Guinea

no abrupt changes in axial ribbing pattern

of teleoconch; ~10-12 ribs per 0.5 mm in

abapertural view; ~10-12 rows of spiral

raised striae per 100 um; columella with

conspicuous median knob

~10-12 ribs per 0.5 mm in abapertural

view; ~10-12 rows of spiral raised striae

per 100 um; columella with conspicuous

median knob

H = 1.6-1.7 mm, W = 0.8-0.95 mm;

constriction area with axial ribs; ~10-12

ribs per 0.5 mm in abapertural view;

~10-12 rows of spiral raised striae per

100 um; both inner and outer peristomes

similarly strongly developed; columella

with conspicuous median knob

axial ribs not becoming twice as distant

on last whorls except beyond constriction;

~10-12 ribs per 0.5 mm in abapertural

view; ~10-12 rows of spiral raised striae

per 100 um; columella with conspicuous

median knob

119

02-Jun-21 21:48:13

Book4.indd 120

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

P. lengguru Greke, 2017

P. minuscularia Greke,

2017

P, psittricha Greke, 2017

P sarmi sp. nov. (de-

scribed above)

P. waigeo Greke, 2017

Updated key to the Papuan Diplommatinidae

The only existing

H = 1.3-1.4 mm, W = 0.75-0.8 mm;

abrupt changes in axial ribbing pattern of

teleoconch - riobS 2x more distant starting

from constriction; ~13-15 ribs per 0.5

mm in abapertural view; peristome thin in

lateral view; columella simple

H = 1.3-1.8 mm, W = 0.7-0.9 mm; ~25-

28 ribs per 0.5 mm in abapertural view;

columella with small basal knob

operculum is derivative - high (tuba-like),

with several layers of concentric lines on

outer surface; ultimate whorl is slightly nar-

rower than penultimate in apertural view;

abrupt changes in axial ribbing pattern of

teleoconch - ribs becoming twice more dis-

tant beyond constriction; axial ribs dense,

more than 20 rows of spiral raised striae

per 100 um; aperture is slightly shifted left

to coiling axis in apertural view

H = 2.6 mm, W = 1.4 mm; constriction is

well defined by presence of a vague impres-

sion, Somewhat denser pattern of axial ribs

and presence of a deep impression on su-

ture above it; axial ribs less dense, ~5-6

ribs per 0.5 mm in abapertural view; ~4-5

rows of spiral raised striae per 100 um;

palatal margin of inner peristome sinuous;

columella with small median knob

H = 1.3-1.35 mm, W = 0.7 mm; ~17-19

ribs per 0.5 mm in abapertural view; more

than 15 rows of spiral raised striae per 100

um; no columellar knob present

Table 2 (continuation)

H = 1.6-1.7 mm, W = 0.8-0.95 mm; no

abrupt changes in axial ribbing pattern

of teleoconch; ~10-12 ribs per 0.5 mm

in abapertural view; peristome (space

between inner and outer peristome)

moderately broad in_ lateral view;

columella with conspicuous median knob

H = 1.6-1.7 mm, W = 0.8-0.95 mm;

~10-12 ribs per 0.5 mm in abapertural

view; columella with conspicuous median

knob

operculum with low concentric ridges,

lacking tuba; ultimate whorl nearly same

width as penultimate in apertural view; no

abrupt changes in axial ribbing pattern of

teleoconch; ~10-12 rows of spiral raised

striae per 100 um; aperture more or less

centered to coiling axis in apertural view

H = 1.6-1.7 mm, W = 0.8-0.95 mm;

constriction poorly defined; ~10-12 ribs

per 0.5 mm in abapertural view; ~10-12

rows of spiral raised striae per 100 um;

palatal margin of inner peristome not

sinuous; columella with conspicuous me-

dian knob

H = 1.6-1.7 mm, W = 0.8-0.95 mm;

~10-12 ribs per 0.5 mm in abapertural

view; ~10-12 rows of spiral raised striae

per 100 um; columellar knob present

2017: 275) must be read:

key to the Papuan and

Correction: the couplet 71 of the key (Greke

Wallacean diplommatinids species was published

recently (Greke 2017). Herewith | provide

modifications to my Palaina key to include new

Species described in the present paper and one

correction. All plates and figures quoted in the

modified couplets of the key are referring to the

original key of Greke (2017).

Considering the fact that during the present

study a dextral Palaina consobrina specimen was

discovered, users of the key (Greke 2017) must be

aware of this fact and have to assess this species

in both branches of the key - the dextral and the

sinistral ones (couplet 1).

120

71 Shell apex conical, shell is not regularly cylindrical ..

Tes!

The couplet 64 is modified to include P.

cenderawasinh sp. nov.:

64 Constriction situated on same axis with parietal side

of aperture, it is defined by denser axial ribs and a sutural

impression; shell H ~3.8 mm; axial ribs Spaceous, not or

SHE SI WOUS was nrsrstnennt peed tenoncece a enattaseaumonndts P. glabella

- Constriction situated on same axis with columellar

side of aperture, it is defined by denser axial ribs and

a sutural impression; shell H ~2.1-2.3 mm; axial ribs

Spaceous, not or slightly sinuous P. obiensis

- Constriction situated on same axis with columellar

side of the aperture; shell H 4.4 mm; axial ribs dense,

02-Jun-21 21:48:13

Book4.indd 121

Gree, K.: New species and records of the Papuan Diplommatinidae (Caenogastropoda: Cyclophoroidea)

strongly sinuous on older whorls, nearly straight on the

pen- and ultimate whorl ......... P. cenderawasih sp. nov.

The couplet 60 is modified to include P.

eEVvaneSCens Sp. Nov.:

60 Peristome double; outer peristome expands strongly

beyond inner one, broadly encircles aperture and is

WIGElCISCONLINUER | Panielallly. cc: .sacarseveees acarcstataceseers 61

- Peristome double; outer peristome is not or partly

expanding beyond inner one, not encircling aperture (if

encircling, then is not extraordinary broad) 62

- Peristome simple P. evanescens Sp. nov.

The couplet 41 is modified to include P.

kagainisi Sp. nov.:

64 There are ~6 ribs per 1 mm on penultimate whorl,

constriction situated on same axis with parietal side of

aperture, it is defined by slightly denser axial ribs, there

are 4-5 delicate but distinct spiral raised striae per 100

um, peristome double, shell H ~3.8 mm ....... P. glabella

- There are ~8-9 ribs per 1 mm on penultimate whorl,

constriction situated on same axis with parietal side of

aperture, it is defined by twice denser axial ribs, spiral

raised striae or striations almost absent (only tracks

of them visible by 50OOx magnification), peristome

multilamellate, shell H ~2.3 mm ... P. kagainisi sp. nov.

- There are ~8-10 ribs per 1 mm on penultimate whorl,

constriction situated on same axis with columellar side

of aperture, it is defined by denser axial ribs, there

are more than 10 spiral raised striae per 100 um (on

older teleoconch whorls, spiral sculpture not present on

ultimate whorl), peristome double, shell H ~2.1-2.3 mm

P. obiensis

The couplet 41 is modified to include P.

pallgergelyi sp. nov.:

Ai Axial ribs straight or oblique against coiling axis on

penultimate whorl in apertural view, ribs are not strongly

sinuous; shell H not exceeding 3.6 mm 42

- Axial ribs strongly oblique against coiling axis on

penultimate whorl in apertural view and are distinctly

sinuous; shell H ~3.9 mm; spiral raised striae delicate

but distinct on penultimate and ultimate whorls; there

are 5-6 axial ribs per 0.5 mm on abapertural side of

penultimate whorl; columellar knob large, somewhat

shifted down closer to base of columella and turned

outwards towards aperture P. consobrina

- Axial ribs strongly oblique against coiling axis on

penultimate whorl in apertural view; axial ribs stronger

sinuous on older whorls than on penultimate and

ultimate ones; shell H >4 mm; there are 9-12 axial ribs

per 0.5 mm on abapertural side of penultimate whorl;

columellar knob large, positioned medially on columella,

not or indistinctly turned outwards towards aperture ....

P. pallgergelyi sp. nov.

pp. 111-134

Since operculum is not known in P. sarmi sp.

nov., this species is included into two branches of

the modified key:

68 Shell rather slender but not conical, penultimate

and ultimate whorls nearly equally wide in apertural

view (Greke (2017): Plate 31 fFgs 1-3); axial ribs strong

and moderately spaceous, ~5-6 ribs per 0.5 mm on

penultimate whorl; spiral raised striae not present;

aperture slightly shifted left to coiling axis ... PR adelpha

- Shell less slender, fusiform, with maximum width on

penultimate whorl (in apertural view) and with broadly

conical apex (Greke (2017): Plate 45 Figs 1-6); axial

ribs less strong and denser, ~9-10 ribs per 0.5 mm

on penultimate whorl; spiral raised striae delicate and

dense, >20 rows of spiral raised striae per 100 um;

aperture slightly shifted left to coiling axis ..............2.00

P. psittricha

- Shell slender, fusiform, penultimate and ultimate

whorls nearly equally wide in apertural view; axial ribs

moderately spaceous, ~5-6 ribs per 0.5 mm; ~4-5 rows

of spiral raised striae per 100 um; aperture more or less

centered to coiling axis P. sarmi sp. nov.

74 Columella without knob; there are significantly less

than 25 axial ribs per 0.5 mm iD

—- Columella with basal knob; two rather strong lamellae

between inner and outer peristome; 12-13 rows of

spiral raised striae per 100 um (measured above the

aperture); 25-28 axial ribs per 0.5 mm

P. minuscularia

- Columella with delicate median knob; ~5-6 axial ribs

per 0.5 mm; ~4-5 rows of spiral raised striae per 100

um P. sarmi sp. nov.

The minute shell of P. saxatilis sp. nov. has axial

ribbing pattern that is significantly more distant on

the last half whorl and strong median columellar

knob:

69 Columella is straight, slightly curved or twisted,

WITROUEIMECIAN KNOB? Seas sccermaannivatte ssi ietatireseindesattaant 70

- Columella with large, median backwards directed

knob (Fig. 53); shell H 2.7-2.9 mm; ~6-7 axial ribs

per 0.5 mm, ribs becoming denser on ultimate whorl in

GSTLUPAL MEW ccnsvecnencorsptte cote awuuenecun tte sruteaencesne P. doberai

- Columella with a large median knob; shell H 1.6-1.7

mm; ~10-12 axial ribs per 0.5 mm, ribs not becoming

denser on ultimate whorl in apertural VieW ............::0+

P. saxatilis sp. nov.

Acknowledgements

| am indebted to Matthias Hartmann (NME)

and Michael Kesl (MKC) for the specimens provided

for current study and for donating specimens for my

? eiex

121

02-Jun-21 21:48:13

Book4.indd 122

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

comparative collection. | am also thankful to Ugis

Kagainis (University of Latvia Faculty of Biology,

Department of Zoology and Animal Ecology, Riga)

for producing the scanning electron micrographs.

Dmitry Telnov (The Entomological Society of Latvia,

Riga) is thanked for preparing images of the

Shells and for critical review of the manuscript. |

am grateful to John Slapcinsky (Florida Museum

of Natural History, Gainesville, U.S.A.) and Edwin

Telnov (Dzidrinas, Latvia) for English proof of the

manuscript.

References

Beehler B. M., Pratt T. K., Zimmerman D. A. 1986. Birds

of New Guinea. Handbook No 9 of the Wau Ecology

Institute. New Jersey, Princeton University Press: xiii

+ 293 pp.

Egorov R. 2013. A review of the genera of the terrestrial

Pectinibranch molluscs (synopsis mainly based on

published data). Part III. Littoriniformes. Liareidae,

Pupinidae, Diplommatinidae, Alycaeidae,

Cochlostomidae. Treasure of Russian shells.

Supplement 3. Colus - Doverie Itd. Press, Moscow:

62 pp.

Geiger D. L., Marshall B. A., Ponder W. F., Sasaki T.,

Warén A. 2007. Techniques for collecting, handling,

preparing, storing and examining small molluscan

specimens. - Molluscan Research 27, No 1: 1-50.

Greke K. 2017. Taxonomic review of Diplommatinidae

(Caenogastropoda: Cyclophoroidea) from Wallacea

and the Papuan Region: 151-316, pls 19-47. In:

Telnov D., Barclay M. V. L., Pauwels O. S. G. (eds)

Biodiversity, biogeography and nature conservation

in Wallacea and New Guinea. Volume Ill. The

Entomological Society of Latvia, Riga: 658 pp, 172

pls.

122

Gressitt J. L. 1982. General introduction: 3-13. In:

Gressitt J. L. (ed.) Monographiae biologicae 42,

Biogeography and ecology of New Guinea. Dr. W.

Junk / Springer Publishers, the Hague: 983 pp.

Hadley A. 2010. CombineZP. Available from https://

combinezp.software.informer.com/download

[accessed 14 April 2020].

Liew T.-S., Vermeulen J. J., bin Marzuki M. E., Schilthuizen

M. 2014. A cybertaxonomic revision of the micro-

landsnail genus Plectostoma Adam _ (Mollusca,

Caenogastropoda, Diplommatinidae), from

Peninsular Malaysia, Sumatra and Indochina. -

Zookeys 393: 1-107.

Neubert E., Bouchet P. 2015. The Diplommatinidae

of Fiji - a hotspot of Pacific land snail biodiversity

(Caenogastropoda, Cyclophoroidea). - ZooKeys

2015 16, No 487: 1-85.

Riedel A. 2002. Taxonomy, phylogeny, and zoogeography

of the weevil genus Euops (Insecta: Coleoptera:

Curculionoidea) in the Papuan Region. Dissertation

zur Erlangung des Doktorgrades der Fakultat

fur Biologie der Ludwig-Maximilians-Universitat

Munchen: 216 pp.

Telnov D. 2011. Taxonomische Revision der Gattung

Macratria Newman, 1838 (Coleoptera: Anthicidae:

Macratriinae) aus Wallacea, Neuguinea und

den Salomonen: 97-285, pls 17-37. In: Telnov

D. (ed.) Biodiversity, Biogeography and Nature

Conservation in Wallacea and New Guinea. Volume

I. The Entomological Society of Latvia, Riga: 434 pp

+ 92 pls.

Received: 20.viii.2020.

Accepted: 30.ix.2020.

02-Jun-21 21:48:13

Gree, K.: New species and records of the Papuan Diplommatinidae (Caenogastropoda: Cyclophoroidea)

pp. 111-134

Figure 1. Palaina (s. |.) citrinella van Benthem Jutting, 1963, specimens from River Armo valley, Papua. A - Apertural

view; B - Abapertural view; C - Lateral view; D - Apical view; E-G - Operculum, inner (E) and outer surface (F) and

lateral (G) view; H - Columella [not to scale].

123

Book4.indd 123 02-Jun-21 21:48:14

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figures 2-3. Papuan Palaina (s. |.) O. Semper, 1865 species. 2 - P. cupulifera van Benthem Jutting, 1963, specimen

from ~34 km ENE Biak, Biak Is. A - Apertural view; B - Abapertural view; C - Lateral view; D - Columella;

3 - P. glabella van Benthem Jutting, 1963, specimen from ~53.5 km NNW Biak, Biak Is. A - Apertural view; B -

Abapertural view; C - Lateral view; D - Columella; E-G - Operculum, outer (E) and inner surface (F) and lateral (G)

view [not to scale].

124 Zar >

: , —

Book4.indd 124 02-Jun-21 21:48:15

Gree, K.: New species and records of the Papuan Diplommatinidae (Caenogastropoda: Cyclophoroidea)

pp. 111-134

Figures 4-5. Papuan Palaina (s. |.) O. Semper, 1865 species. 4 - P. inconspicua van Benthem Jutting, 1963,

specimen from near Urfu village, Biak Is. A - Apertural view; B - Abapertural view; C - Lateral view; 5 - P. liliputana

van Benthem Jutting, 1963, specimen from 56 km NNW Biak, Biak Is. A - Apertural view; B - Abapertural view;

C - Lateral view; D - Columella [not to scale].

125

Book4.indd 125 02-Jun-21 21:48:16

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figure 6. Palaina (s. |.) cenderawasih sp. nov., paratypes from near Biak Bird park, Biak Is. A - Apertural view;

B - Abapertural view; C - Lateral view; D - Apical view; E-F - operculum, internal surface (E) and

lateral view (F); G - Columella [not to scale].

2s => YF BS oo

Book4.indd 126 02-Jun-21 21:48:17

Grete, K.: New species and records of the Papuan Diplommatinidae (Caenogastropoda: Cyclophoroidea)

pp. 111-134

1 : 4b wes

F >. ¥-4y 4

math \

i 4

pi). 4

LAS

J | > |

” se /

Figure /. Palaina (s. |.) evanescens sp. nov., holotype. A - Apertural view; B - Abapertural view; C - Lateral view;

D - Apical view; E - Columella [not to scale].

127

Book4.indd 127 02-Jun-21 21:48:18

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figure 8. Palaina (s. |.) kagainisi sp. nov., holotype. A - Apertural view; B - Abapertural view; C - Lateral view;

D - Apical view; E - Operculum, outer surface; F - Columella [not to scale].

198 —as- FF &

Book4.indd 128 02-Jun-21 21:48:18

Gree, K.: New species and records of the Papuan Diplommatinidae (Caenogastropoda: Cyclophoroidea)

pp. 111-134

Figure 9. Palaina (s. |.) pallgergelyi sp. nov., holotype. A - Apertural view; B - Abapertural view; C - Lateral view;

D - Apical view; E - Columella [not to scale].

129

Book4.indd 129 02-Jun-21 21:48:19

TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

aS _— a

o ;

J -

Figure 10. Palaina (s. |.) sarmi sp. nov., holotype. A - Apertural view; B - Abapertural view; C-D Lateral view;

E - Apical view [not to scale].

130 Za er <>

Book4.indd 130 02-Jun-21 21:48:19

Grete, K.: New species and records of the Papuan Diplommatinidae (Caenogastropoda: Cyclophoroidea)

pp. 111-134

Figure 11. Palaina (s. |.) saxatilis sp. nov., holotype. A - Apertural view; B - Abapertural view; C-D Lateral view;

E - Apical view [not to scale].

ay & 131

Book4.indd 131 02-Jun-21 21:48:20

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

1 4 300 um 1 5 300 um

a . 300 um 17 | : 7 : aT)

18 300 um 19 300 um

Figures 12-19. Micrographs of the Papuan Palaina (s. |.) O. Semper, 1865 (300x magnification). 12-13 - P

cenderawasih sp. nov., paratype, penultimate (12) and ultimate (13) whorl; 14 - P citrinella van Benthem Jutting,

1963; 15 - P. consobrina van Benthem Jutting, 1963; 16 - P cupulifera van Benthem Jutting, 1963; 17 - P

evanescens sp. nov., holotype; 18 - P. glabella van Benthem Jutting, 1963; 19 - P inconspicua van Benthem

Jutting, 1963.

2 =~ ©

Book4.indd 132 02-Jun-21 21:48:21

Gree, K.: New species and records of the Papuan Diplommatinidae (Caenogastropoda: Cyclophoroidea)

pp. 111-134

Figures 20-26. Micrographs of the Papuan Palaina (s. |.) 0. Semper, 1865. 20-21 - P. kagainisi sp. nov., holotype,

500x (20) and 600x (21) magnification; 22 - P. liliputana van Benthem Jutting, 1963 (500x); 23 - P. repandostoma

van Benthem Jutting, 1963 (300x); 24 - PR. sarmi sp. nov., holotype (300x); 25-26 - P. saxatilis sp. nov., paratype,

300x (25) and 500x (26).

133

Book4.indd 133 02-Jun-21 21:48:22

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

LOR,

ites

1S.)

Map 1. Localities of Palaina (s. |.) O. Semper, 1865 species in northern New Guinea. Circle - P. sarmi sp. nov.;

diamond - P. citrinella van Benthem Jutting, 1963 and P. saxatilis sp. nov.; triangle - P. consobrina van Benthem

Jutting, 1963 and P. evanescens sp. nov. (prepared with ArcGIS 10.3).

Map 2. Localities of Palaina (s. |.) O. Semper, 1865 species on Biak, Cenderawasih Bay Islands. Square and diamonds

(empty and filled) - P. lilioutana van Benthem Jutting, 1963; Diamonds (empty and filled) - P. glabella van Benthem

Jutting, 1963; Filled diamond - P. kagainisi sp. nov. and P. pallgergelyi sp. nov.; Five-point star - P. cenderawasih sp.

nov. and P. inconspicua van Benthem Jutting, 1963; Four-point star - P. cenderawasih sp. nov. and P. repandostoma

van Benthem Jutting, 1963; Circle - PR. cupulifera van Benthem Jutting, 1963 (prepared with ArcGIS 10.3).

134 te

Book4.indd 134 02-Jun-21 21:48:25

Book4.indd 135

GrekE, K. & Stapcinsky, J.: New Taheitia H. et A. Adams, 1863, with revisional notes on the Papuan Truncatellidae ...

pp. 135-184

New TJaheitia H. et A. Adams, 1863, with revisional

notes on the Papuan Truncatellidae (Caenogastropoda:

Truncatelloidea)

urn:lsid:zoobank.org:pub:B /CB17 /C-75EA-47 3E-90EE-6C81/74CC3720

KRISTINE GREKE *, JOHN SLAPCINSKY 2

1 - Latvian National Museum of Natural History, K. Barona tela 4, LV-1050, Riga, Latvia;

k.greke@gmail.com

2 - Florida Museum of Natural History, 1659 Museum Road, Gainesville, Florida, U.S.A.;

Slapcin@flmnh.ufl.edu

Abstract: Surveys of snails from New Guinea and surrounding islands provide new distributional and

ecological data for Truncatellidae from the Papuan Region. We present an annotated checklist of all Papuan

Truncatellidae and the first identification key for the terrestrial genus Taheitia H. and A. Adams, 1863. Unlike

widespread marine and estuarine species of Truncatella, the Papuan Taheitia usually have single island ranges

and are restricted to forested limestone habitats. New anatomical data suggests that many earlier reports

of Taheitia with multi-island distributions appear to be the result of confusion between superficially similar

species. Eight new Taheitia species are described: T. biaka sp. nov. from Biak, T. bifurca sp. nov. from Manus,

T. gebeensis sp. nov. from Gebe, T. gigantea sp. nov. from Waigeo, T. jodiae sp. nov. from New Britain, T.

longpela sp. nov. from New Britain, T. malagan sp. nov. from New Ireland, and T. telnovi sp. nov. from Misool.

Key words: Taxonomy, new species, key, Jaheitia, Truncatella, Papuan Region, ecology, endemism.

Introduction

Truncatellidae Gray, 1840 is a family of

marine, estuarine and terrestrial caenogastropod

Snails distributed throughout the tropics and

subtropics (Clench & Turner 1948; Egorov 2018).

The family is currently divided into two subfamilies:

1) Truncatellinae Gray, 1840, comprised of the

genera Truncatella Risso, 1826 and Taheitia H.

et A. Adams, 1863; 2) Geomelaniinae Kobelt

et Mollendorff, 1897 with the single genus

Geomelania. Truncatella, is cosmopolitan in the

tropics and subtropics in marine and estuarine

habitats, with a few notable exceptions in terrestrial

habitats (Rosenberg 1996). The other two genera

are terrestrial with Taheitia restricted to the Pacific

Basin, and Geomelania, the Atlantic Basin. Taheitia

is morphologically more similar to Truncatella

(Clench & Turner 1948) than Geomelania and the

two terrestrial genera appear to have independently

colonized terrestrial habitats in the Pacific and

Atlantic Basins respectively (Rosenberg 1996).

However, Egorov (2018) places Taheitia in the

Geomelaniinae implying a common origin of the

terrestrial species in both the Pacific and Atlantic

Basins. Unfortunately, independent molecular data

is currently available for only a few taxa from the

Atlantic Basin (Wilke et a/. 2013) and it remains

unclear how many times terrestrial habitats have

been colonized by truncatellids.

The last comprehensive treatment of the

family (Clench & Turner 1948) includes anatomical

and morphological notes, an annotated checklist,

and bibliography of all truncatellids known up to

that time. Two truncatellid genera, Taheitia and

Truncatella, occur in the Papuan Region, defined

as the area from the Maluku to the Solomon

Islands (Gressit 1982; Beehler et a/. 1986; Riedel

2002; Telnov 2011), Papuan truncatellids and

in particular the fully terrestrial genus Taheitia

remain incompletely sampled and even incidental

collections uncover undescribed species (Turner

1959; van Benthem Jutting 1963; Gittenberger

1989). Our surveys allow us to provide an

updated annotated checklist of Truncatellidae

from the Papuan Region with new distributional

and ecological records. New morphological data,

particularly radular and opercular characters allow

us to redescribe Taheitia clathrata (H. Adams

et Angas, 1865), T. foliosocostata van Benthem

135

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Jutting, 1963, T. galactodes van Benthem Jutting,

1963, T. schneideri (|. Rensch, 1937), T. tesselata

Mollendorff, 1897, T. ultima (|. Rensch, 1937), and

T. wallacei (H. Adams, 1865). Our surveys also

uncovered eight new species: Taheitia biaka sp.

nov. from Biak, T. bifurca sp. nov. from Manus, T.

gebeensis sp. nov. from Gebe, T. gigantea sp. nov.

from Waigeo, T. jodiae sp. nov. and T. longpela sp.

nov. from New Britain, T. malagan sp. nov. from New

Ireland, and T. telnovi sp. nov. from Misool. The first

key to Papuan species of Taheitia H. et A. Adams,

1863 is presented. Peculiarities of the exclusively

terrestrial Papuan representatives compared to the

typical Taheitia sensu Adams and Adams (1863)

are briefly discussed.

Material and methods

Newly collected specimens were preserved

as dry shells or whole animals in 75% or 99%

ethanol and were identified by the authors. A Leica

S6D stereomicroscope or Wild M7 were used for

dissection and study. Shell images were taken

using a Canon EOS 77D or Nikon D9O SLR cameras

attached to stereomicroscopes. CombineZP

software was used for image stacking (Hadley

2010). Further image manipulations were done

in GNU Image Manipulation program (GIMP) or

Adobe Photoshop. Micrographs were imaged using

a Hitachi SU5000 Schottky field emission scanning

electron microscope.

A phylogenetic arrangement is currently

impossible and all taxa are listed alphabetically.

All label text for museum specimens is reproduced

without additions. Author’s comments are placed

in square brackets. Text from multiple labels is

separated by a slash. Type specimens of new

Species are provided with an additional black-

framed printed label “Holotypus” or “Paratypus” on

red paper.

Museum acronyms used in the text:

BMNH - Natural History Museum (formerly British

Museum, Natural History), London, United

Kingdom;

KGC - Collection Kristine Greke, Dzidrinas, Latvia;

MBBJ - Museum Zoologicum Bogoriense, Indonesian

Institute of Sciences, Cibinong, Indonesia;

MFNB - Museum fur Naturkunde der Humboldt-

Universitat zu Berlin, Germany;

NME - Naturkundemuseum Erfurt, Germany;

NMNL - Naturalis Biodiversity Center, Leiden,

Netherlands;

the

136

NMS - National Museums Scotland, Natural Sciences,

Edinburgh, United Kingdom;

PNG - Papua New Guinea National Museum and Art

Gallery, Port Moresby, Papua New Guinea

UF - Florida Museum of Natural History, Gainesville,

Florida, USA.

Measurement abbreviations:

AH - height of aperture;

AW - width of aperture;

EW - width of embryonic whorls;

PW - width of penultimate whorl;

SH - decollate adult shell height including the peristome;

SW - shell width including the peristome;

UH - height of ultimate whorl;

UW - width of ultimate whorl.

Other abbreviations:

env. - environs;

juv. - juvenile (not adult or subadult) specimen;

vill. - village or little settlement.

Terminology critical for Truncatellidae species

recognition follows Clench and Turner (1948) and

Turner (1959). The following measures were taken

for all taxa: shell height measured parallel to the

Shell axis, height of the ultimate whorl, maximum

Shell width, and maximum height of the aperture.

For the widespread species only those bibliography

records are given related to the study area.

Holotypes of the newly described species are

deposited at the NME and UF. Additional specimens

are deposited at MBBJ and PNG.

Results

Gastropoda Cuvier, 1795

Caenogastropoda Cox, 1960

Littorinimorpha Golikov et Starobogatov, 1975

Truncatelloidea Gray, 1840

Truncatellidae Gray, 1840

Truncatella Risso, 1826

Type species: Truncatella costulata Risso, 1826

subsequent designation by Lowe (1855).

Truncatella is a genus of caenogastropods,

mostly marine and estuarine but some are

also adapted to living in near shore terrestrial

environments. Five species and one subspecies

are found in the Papuan Region.

02-Jun-21 21:48:25

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GrREkE, K. & Stapcinsky, J.: New Taheitia H. et A. Adams, 1863, with revisional notes on the Papuan Truncatellidae ...

Truncatella (Ss. s.) granum Garrett, 1872

Bibliography: van Benthem Jutting (1963: 440),

record (New Guinea, Doberai Peninsula, “Pasirputih

near Manokwari, on fallen tree trunk”).

Ecology: Salt-tolerant species of drift line.

Distribution: Pacific islands from. Fiji

typicus) to W New Guinea.

(locus

Truncatella (s. s.) guerinii A. Villa et J. B. Villa,

1841 (Fig. 1)

Truncatella aurantia Gould, 1847

Truncatella cristata Crosse, 1868

Truncatella fasciata Tapparone Canefri, 1886

Truncatella ferruginea Cox, 1868

Truncatella pacifica Pease, 1867

Truncatella semperi Kobelt, 1884

Truncatella valida L. Pfeiffer, 1846

Truncatella vitiana Gould, 1847

Truncatella yorkensis Cox, 1868

Bibliography: Tapparone Canefri (1874: 563), as

T. valida, record (Aru Islands: “Wokan”); Tapparone

Canefri (1877: 290), as T. valida, record (New Guinea,

Doberai Peninsula: “Sorong”); Tapparone Canefri

(1883: 280), as T. valida, record (Aru Islands: “Wokan”);

Tapparone Canefri (1886: 192-193), as T. pacifica,

record (New Guinea, Doberai Peninsula: “Tangion Bair,

costa occidentale della Nuova Guinea”), as T. fasciata,

record Aru Islands: “Wokam”); Hedley (1891: 100),

as T. valida, records (New Guinea, Papuan Peninsula:

“Port Moresby”; Milne Bay Province: “Milne Bay,

Samarai” (islet offshore extreme SE New Guinea)); Sods

(1911: 349), as T. valida, records (North New Guinea:

“Berlinhafen” (now Berlin Harbour), “Tamara Island”

(now Tumleo); Leschke (191.2: 144), as T. valida, records

(North New Guinea: “Berlinhafen” (now Berlin Harbour),

“Tamara Island” (now Tumleo); East New Guinea, Huon

Gulf: “Simbang”); Boettger (1922: 392, pl. 22 fig.

27), as T. valida, record (“Aru Inseln”); Adam & Leloup

(1938: 82, pl. 5 fig. 3), as T. valida, records (Banda

Islands: “lle Pisang”; New Guinea, Doberai Peninsula:

“Ile Mansinam, Manokwari”); Rensch I. (1937: 626,

633 & 634), as T. valida, additional description,

genera distribution, records (Bismarck Archipelago,

Duke of York (Neulauenburg); New Britain: “Karlei

(Weite Bucht)”, “Nordkuste (Ulamona)”; New Ireland:

“Ugana”; Vuatom), as Truncatella (Taheitia) semperi,

redescription, records (Bismarck Archipelago, New

Britain (Neupommern), New Ireland (Neumecklenburg);

Solomon Islands, New Georgia: “Batuma, Morawa

Lagune”); Rensch |. (1940: 39), as T. semperi, record

(Bismarck Archipelago: New Britain (Neupommern));

Clench & Turner (1948: 167 & 199), synonymy,

checklist, records (Aru Islands: Wokam (Wokan); North

Moluccas: Bacan (Batjan); Shortland Islands (Papua

New Guinea): Shortland: “Harapa”; Solomon Islands:

Guadalcanal (Guadalcanar); van Benthem Jutting

(1963: 439), as T. guerini [sic], bibliography, records

pp. 135-184

(New Guinea, northern part: “Hollandia” and “near

Base “G” near Hollandia” (now Jayapura area); Doberai

Peninsula: “Manokwari, “Pasirputih near Manokwari,

on fallen tree trunks”; Cenderawasih Bay Islands, Biak

Island: “Ariodbi (Division of Marines)”, “Biak”, “Jodbi”;

“Kombai”, “Warsa”, “mangrove between Parai and Idbi,

W of Bosnik, S coast”, “beach near kampong Mokmer, E

of airport hotel”; “kampong Saba, E part of Biak, under

drift material”; Numfoor Island: “Numfoor”; Raja Ampat

Islands, Waigeo: “Waigeu”); Vermeulen (1996: 108 &

113), as T. guerini [sic!], record (Aru Islands), ecology

(“entirely restricted to areas under immediate influence

of the sea (shores, mangrove vegetation, etc.”); Greke

(2012: 229), general distribution, record (North

Moluccas, Gebe: “on the way from Kacepi to Umera vill.,

secondary rainforest on limestone”).

New material: strand bij kolensteiger (Ambon) 10-6-

1939 [2 adults, NMNL]; INDONESIA E, Prov. Raja Ampat,

Misool S, distr. Misool Utara, Aduwey (Adua) vill. ~1 km

E, Seram Sea shore, in a small stream near the sea,

01°58’41’"S, 129°55'18’E, 28.1I1.2009 [1 adult, KGC];

INDONESIA E, West Papua, S Bird’s Neck, Kaimana E

env., between Triton Bay & Bitsyaru Bay, 3°49’24’S,

133°59’02”E, 6-20 m, 18.IX.2010, primary coastal

rainforest on limestone. First records from southern New

Guinea and Misool Island [1 adult, KGC].

Ecology: Salt-tolerant species of drift line. This

underrecorded species may occur throughout the

Papuan Region.

Distribution: Indo-Pacific islands from

Mozambique, Reunion (locus typicus) through

coastal SE Asia to Japan, the Philippines,

Micronesia, Polynesia, and N Australia. Whole of

Wallacea and Papuan Region (recorded from W,

E, N, S New Guinea, Bismarck Archipelago (New

Britain, New Ireland), Aru Islands, Banda Islands

(Pisang), Bismarck Archipelago (Duke of York, New

Britain, New Ireland, Vuatom), Cenderawasih Bay

Islands (Biak, Numfoor), Milne Bay Islands, North

(Gebe) and Central (Ambon) Moluccas, Raja Ampat

Islands (Misool, Waigeo), and Solomon Islands

(New Georgia)).

Truncatella (s. s.) marginata Kuster, 1855 (Fig. 2)

Truncatella cerea Gassies, 1878

Truncatella ceylanica Pfeiffer, 1856

Truncatella japonica Pilsbry et Hirase, 1905

Truncatella pfeijferi Martens, 1861

Truncatella quadrasi Mollendorff, 1893

Truncatella semicostata Montrouzier, 1862

Truncatella semicostulata Jickeli, 1874

Truncatella teres Pfeiffer, 1856

Bibliography: Hedley (1891 : 100), as T. ceylanica,

records (New Guinea, Papuan Peninsula: “Port

ABY.

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TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Moresby”); Rensch |. (1937: 627, 633 & 634), as T.

quadrasi, additional description, records (Bismarck

Archipelago, New Britain: “Weite Bucht, Karlei”; Vuatom);

van Benthem Jutting (1963: 440), as T. marginata

teres, record (Raja Ampat Islands, Yefman (also Efman,

Jefman) Island: “Jefman I|.: beach of Jefman”).

New material: INDONESIA E, Prov. Raja Ampat, Misool

S, distr. Misool Utara, Aduwey (Adua) vill. ~1 km E, Seram

Sea shore, in a small stream near the sea, 01°58’41S,

129°55'18”E, 28.1I1.2009 [1 adult, KGC]. First record

for Misool.

Ecology: Salt-tolerant species of drift line.

Distribution: Indo-Pacific islands from E Africa

to Japan, Greater Sunda Islands (locus typicus -

Borneo), W New Guinea, coastal Australia, and

New Caledonia. In the Papuan Region known from

Bismarck Archipelago (New Britain, Vuatom) and

Raja Ampat (Misool, Yefman offshore Salawati).

Truncatella (s. s.) thaanumi insularis Clench et

Turner, 1948 (Fig. 3)

New material: INDONESIA E, Papua _ Prov.,

Cenderawasih Bay, Biak Is. W, Biak ~16 km W, Urfu

MiINFOT 09 1275.21.35) 795 SS°Er S215 m2 Vl 2O1S:

coastal limestone cliff, in cavities with vegetation and

debris [1 adult, KGC]. First record from the Papuan

Region and the Island of Biak.

Notes: Considering the high variability of

conchological features in Truncatella (as of

Clench & Turner 1948) and the relatively minor

morphological differences that separate these

two taxa we question the subspecific status of

T. thaanumi insularis but do not propose a new

synonymy, since type material was not available for

the present study. Both subspecific taxa occur in

the Papuan Region (see below).

Distribution: Tonga (locus typicus) to W New

Guinea. In the Papuan Region only known from

Cenderawasih Bay Islands (Biak).

Truncatella (s. s.) thaanumi thaanumi Clench et

Turner, 1948

Bibliography: van Benthem Jutting (1963: 440),

record (New Guinea, Doberai Peninsula: “Pasirputih

near Manokwari, on fallen tree trunk”); Egorov (2018:

31, fig. 501), record (New Guinea, Doberai Peninsula:

“Manokwari, near Uriami River”).

Distribution: Caroline Islands (locus typicus) to W

New Guinea. Only two records are known from the

Papuan Region, one from New Guinea's Doberai

Peninsula and the other from Atoll Fragment near

Manus Island, Admiralty Islands (USNM ID 835181;

—_—_

it i

138

GUID http://n2t.net/ark:/65665/3b22ad929-

Ob8f-4921-92b6-3d94b0/b555d), the Manus

specimens were not available for this study.

Taheitia H. et A. Adams, 1863

Type species: Truncatella porrecta Gould, 1847

original designation.

Taheitia is a genus of terrestrial Truncatellidae

(Turner 1959) with twenty one species in the

Papuan Region. Taheitia was originally erected

by Adams & Adams (1863) to include a single

Pacific islands species, 7. porrecta (Gould, 1847),

the small species (height of adult decollated

Shell distinctly under 10 mm). Adams (1865) was

the first to include a new species of large and

exclusively terrestrial Papuan species in Taheitia,

when describing T. wallacei. No justification for

the generic placement was indeed provided. This

was soon followed by various authors placing their

new species from the Papuan Region in the genus

Taheitia (Adams & Angas 1865; Mollendorff 1897;

Rensch 1937). Clench and Turner (1948) on their

catalogue of the Truncatellidae made some new

combinations and generally agreed with generic

placement of previously described Papuan Taheitia.

Later, van Benthem Jutting (1958; 1963), Turner

(1959), and Gittenberger (1989) continued placing

new terrestrial species in the genus Taheitia with

only short discussion of Taheitia morphology by

Turner (1959).

In fact, exclusively terrestrial, large Papuan

species hitherto and in the present paper

arranged to Taheitia appear strongly differentiated

conchologically from the T. porrecta (Fig. 4), the

type of the genus. In the present sense, Taheitia is

a strongly dimorphic group. The authors currently

refrain from erecting a new genus for terrestrial

Papuan Taheitia as well as from proposing new

combinations, since comparative material of T.

porrecta required to study its anatomy, radular

features, and operculum are currently unavailable.

Taheitia biaka sp. nov. (Fig. 5)

http://zoobank.org/A8C /6309-BBB9-4BC0-9E60-

39B21700A/04

Holotype NME: INDONESIA’ E, Papua _ Prov.,

Cenderawasih Bay, Biak Is. N, Biak ~53 km NNW,

0°44'°19"S, 135°52’56”E, 70-80 m, 22.1I1.2018,

primary lowland rainforest.

Derivatio nominis: Toponymic from the locus

typicus, Biak Island.

Measurements: Holotype H = 11.3 mm, W = 2.8

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Greke, K. & Stapcinsky, J.: New Taheitia H. et A. Adams, 1863, with revisional notes on the Papuan Truncatellidae ...

mm, HW = 4 mm, AH = 2.2 mm.

Description: Shell is decollate, whitish

transparent, dextral, very narrow, almost parallel-

sided, slightly and gradually narrowing apicad.

Decollate adult shell with 642 moderately strongly

convex whorls, number of embryonic whorls is

unknown. Suture is deeply impressed, oblique to

the coiling axis (angle at ~70°). Ultimate whorl is

nearly as wide as penultimate whorl in apertural

view (maximum width 2.6 mm vs 2.5 mm), 6

whorls, the first non-decollate whorl is 1.7 mm

wide. Umbilicus is closed in adult. Teleoconch is

sculptured with moderately dense, nearly straight

(in apertural view), and more or less regular

axial ribs. Ribs slightly curved and wider below

the suture (Fig. 5). Ribs do not always run from

suture to suture. There are no abrupt changes

in axial ribbing pattern, but ribs on the last half

whorl widen to nearly twice the width of ribs on

penultimate whorl. Ribs are not synchronous with

those of previous whorls and are parallel to coiling

axis. There are 5 ribs per 0.5 mm on penultimate

whorl in abapertural view and 50-51 ribs on

ultimate whorl. Axial ribs of ultimate whorl evenly

rounded basally, basal ridge is therefore absent

(ultimate whorl rounded, not angled, basally and

laterally). Spiral striae or ribs not observed at 80x

magnification. Last part of ultimate whorl protruding

slightly anteriad from shell outline in apertural and

apical view. Parietal gap present, moderately wide

and comparatively shallow. Aperture is obliquely

ovoid (narrowed in parietal-palatal angle), not tilted

to coiling axis, positioned vertically, apertural rim

entire, and with shortly reflected, not flattened

apertural lip. Aperture is shifted right to the coiling

axis in apertural view. Peristome is simple in lateral

view. Parietal margin of peristome is detached from

ultimate whorl, peristome is therefore continuous.

Margins of peristome not sinuous. Operculum

(Fig. 5D-F) irregularly ovoid, height 1.9 mm,

maximum width 1.2 mm, strongly concave on outer

surface (therefore convex internally). Its chitinous,

paucispiral base bears a calcareous plate with

a high erect thin ridge along peripheral margin,

and at the opposite margin with 9 strong, apically

pointed and slightly curved lamellar ridges radiate

from apophysis.

Differential diagnosis: Taheitia biaka sp. nov.

is readily differentiated from congeners by the

combination of the following features: the shell high

and slender, slowly narrowing apicad, ~50-51 axial

ribs on the ultimate whorl, the base of the ultimate

whorl not angled, the aperture not produced into the

spur, the broadly rounded at the basal-columellar

pp. 135-184

angle, the axial ribs only slightly increasing in the

shape towards the suture, the operculum with the

conspicuous, radial, raised and pointed lamellae.

Ecology: Specimen sampled from wet leaf litter in

primary lowland (elevation 70-80 m) rainforest on

a limestone ridge.

Distribution: The Papuan Region. Cenderawasih

Bay Islands, Biak (Map 2).

Taheitia bifurca sp. nov. (Figs 6, 27 & 30)

http://zoobank.org/EDD40E8/-80D9-4688-9C10-

FEOF5BCF49D1

Holotype UF 563275: PAPUA NEW GUINEA, Manus

Province, Manus Island, 0.4 km W of Loniu, -2.0749°,

147.3328°, John Slapcinsky, Cindy Bick, 1 December

2012.

Paratypes: UF 475/732 (4 whole in 75% ethanol),

476354 (1 whole in 75% ethanol) as holotype; UF

475765 (3 whole in 75% ethanol), UF 475766 (2 shells):

Papua New Guinea, Manus Province, Los Negros Island,

3.8 km E of Bubi, -2.0732° , 147.3719 °, John Slapcinsky,

Cindy Bick, 2 December 2012.

Derivatio nominis: From the Latin “bifurcus”

meaning two forked or bifurcate for the pattern of

v-shaped ridges on the shell surface.

Measurements: Holotype H = 14.09 mm, W =

5,25, AH = 4.66, AW = 3.22.

Description: Shell is dextral, conical, gradually

expanding, and not umbilicate. The total number

of whorls is unknown because only decollate

adults were found and collected. Suture is slightly

shouldered and not deeply impressed, oblique to

the coiling axis atan angle of ~70°. Embryonic shell

unknown. Adult shell is decollate, with 5-5% broadly

rounded whorls, moderate sized, SH=12.17+1.01,

SW=4.60+0.33, AH=4.21+0.25, AW=2.90+0.23,

N=7. Width of first non-decollate whorl is 2.40+0.20

mm, penultimate whorl is 3.57+0.24 mm, and

ultimate whorl is 3.85+0.27 mm. Aperture not

deflected downward or peripherally and remains

attached to penultimate whorl at sutural margin.

Aperture is almost rectangular, somewhat flattened

at periphery, and base. Junction of basal and flat

columellar margin of peristome Is angled. Peristome

is pale tan, entire, only slightly reflected near

columellar margin, and not flattened. Teleoconch

whorls are light brown, contrasting with tan ribs and

surface sculpture. There is no other colour pattern.

Non-decollate teleoconch whorls are sculptured

with regular slightly arched axial ribs, each about

¥% the width of the rib-interspaces. Some ribs do

not extend to the sutures and other ribs widen at

apical suture forming weak nodules. There are also

chevron-shaped ridges that point in the direction of

139

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

growth. These are most prominent on penultimate

and ultimate whorls. Ribs are often thicker and

nodulate where ridges cross them. Ribs are angled

basally forming basal ridge. Spiral sculpture not

observed at 80x magnification. Operculum (Fig. 6E)

irregularly ovoid, height 3.28, maximum width 1.91

mm, slightly concave on outer surface. Its chitinous,

paucispiral base bears a calcareous plate with

large semicircular apophysis with radiating lamellar

ridges that extend to the peripheral margin. Radula

(Fig. 30) formula 2-1-1-1-2. Rachidian with

large central cusp flanked on each side by three

small near-basal ectocones. Lateral teeth wide,

asymmetric, with poorly defined ectocones. Inner

marginal with 4-5 finger like cusps. Outer marginal

with 3 short broad, rounded cusps.

Differential diagnosis: Taheitia bifurca sp. nov.

is unique in posessing shell sculpture of chevron

shaped ridges and thin axial ribs that are thickened

and nodulose where the chevrons cross them. The

flattened shell whorls and irregularly oval aperture

with flattened base and nearly parallel edges are

superficially similar to the smaller T. foliosocostata

van Benthem Jutting, 1963 from Biak which differs

in having stronger and more regular ribbing and

T. tessellata from New Britain which has sinuous

ribs on the ultimate and penultimate whorls and a

thicker lip. The operculum is similar to Fijian species

T. arcasiana abbotti Clench et Turner, 1948 and T.

soluta Clench et Turner, 1948 as well as Papuan

Region species T. biaka from Biak and T. wallacei

from Waigeo Island in having several long lamellae

that radiate from a single point but unlike these

species the origin of the lamellae in 7. bifurca sp.

nov. is from a semicircular apophysis.

Ecology: On limestone rocks and in leaf litter

among rocks in tropical rainforest

Distribution: The Papuan Region. Manus Island

(Map 4).

Taheitia bismarckiana (I. Rensch, 1937) (Fig. 7)

Bibliography: Rensch I. (1937: 627, 633 &

634), original description as Truncatella avenacea

bismarckiana, records (Bismarck Archipelago, Duke of

York (Neulauenburg), New Ireland (Neumecklenburg);

Clench & Turner (1948: 191 & 196), new combination,

checklist.

Syntypes 4 specimens MFNB, stored in two capsules.

2 specimens: Lauen [upper side of the label] burg

[underside of the label] [both sides handwritten]; 2

specimens: Neu Mecklen [upper side of the label] burg

[underside of the label] [both sides handwritten]. Both

capsules stored in same box with two additional labels:

Truncatella avenacea bismarckiana [upper side of the

140

label] leg. P. Schneider [underside of the label] [both

sides handwritten] / Zoolog. Museum Berlin [printed]

97961 Truncatella avenacea - bismarckiana I. Rensch

Neu - Lauenb. P. Schneider [handwritten] [upper side of

the label] Neu-Lauenbg. [handwritten] [underside of the

label].

Notes: Lectotype is not designated at this time.

This small species appears to be a typical Taheitia

sensu Adams and Adams (1863), with strongly

double peristome, or Truncatella. We provide

images of this species for the first time.

Distribution: The Papuan Region. Known only

from Bismarck Archipelago (Duke of York (locus

typicus), New Ireland).

Taheitia calcarata van Benthem Jutting, 1963

(Fig. 8)

Bibliography: van Benthem Jutting (1963: 442, fig. 8),

original description, records (Cenderawasih Bay Islands,

Biak: “subdistrict South-Biak, base of the Marines”;

“base of the Marines”).

New material: INDONESIA E, Papua _ Prov.,

Cenderawasih Bay, Biak Is. N, Biak ~56 km NNW,

O° 42'40'S, .. 135*51'59°E, 95-120" mM, 422.2018;

primary lowland rainforest on limestone cliffs [2 adults,

NME & KGC]. First record since the original description,

first record from northern part of Biak.

Additional description: The studied decollate

adult specimen has ~5 whorls, shell height is 10

mm, maximum width is 3.8 mm. There are 35 axial

ribs on the ultimate whorl.

Distribution: The Papuan Region. Known only

from Biak, Cenderawasih Bay Islands (Map 2).

Taheitia clathrata (H. Adams et Angas, 1865)

(Fig. 9)

Bibliography: Adams H. & Angas (1865: 54, pl. 2 fig.

2), original description as Truncatella (Taheitia), record

(Solomon Islands); Clench & Turner (1948: 191 & 197),

checklist.

Syntypes 3 specimens NMS: TRUNCATELLA (TAHEITIA)

CLATHRATA AD. + ANG. [underlined] SOLOMON IS. [label

handwritten].

Notes: The syntypes have accession No

NMS.Z.1961.61.163. Lectotype is not being

designated at this time. This small species seems

to be true Taheitia in sense of Adams & Adams

(1863). We provide images of this species for the

first time.

Additional description based on the syntypes:

Shell is decollate, creamy-yellowish. Shell high

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conical. Adult (decollate) shell of 542-6 rather flat

whorls. Suture deep. Axial ribs strong and high,

ridge-like, straight to nearly straight, widely spaced,

rib interspaces twice as wide as ribs. Ribs are not

synchronous with those of previous whorls and are

generally parallel to the coiling axis. Ribs becoming

significantly less high and denser on last quarter

whorl. Spiral striae or ribs not observed. Parietal

gap distinct. Aperture is half-moon shape, nearly

flat along the parietal and columellar margin and

broadly rounded along palatal margin. Aperture

not tilted to coiling axis, positioned subvertically,

apertural rim entire, with flattened apertural lip,

which is reflected slightly basally. Aperture is shifted

right to coiling axis in apertural view. Peristome is

simple in lateral view. Parietal margin of peristome

is detached from ultimate whorl, peristome is

therefore continuous. Margins of peristome not or

barely sinuous. Operculum unknown.

Distribution: The Papuan Region. Known only

from type series of syntypes from the Solomon

Islands (locus typicus) in National Museum Scotland

(Edinburgh, United Kingdom), without exact locality.

Taheitia foliosocostata van Benthem Jutting,

1963 (Fig. 10)

Bibliography: van Benthem Jutting (1963: 443, fig. 9),

original description, records (Cenderawasih Bay Islands,

Biak: “Biak”, “Bosnik, coral reef near sago swamp”,

“subdistrict South-Biak, base of Marines”).

New material: INDONESIA E, Papua _ Prov.,

Cenderawasih Bay, Biak Is. W, Biak ~16 km W, Urfu vill.,

01°09'12"S, 135° 55’53’E, 3-15 m, 21.11.2018, coastal

limestone cliff, in cavities with vegetation and debris

[20 adults, 2 subadult & 2 juv., NME, MBBJ & KGC];

INDONESIA E, Papua Prov., Cenderawasih Bay, Biak Is.

E, Biak ~34 km ENE, 01.°04’50’S, 136°22’08’E, 10-15

m, 22.11.2018, primary lowland rainforest [1 adult & 1

juv., KGC]. First record since the original description, first

record from eastern tip of Biak.

Additional description: Total number of whorls

is no less than 12, the number of embryonic whorls

is two. The studied decollate adult soecimens have

444-5 whorls, shell height varies from 5.6 mm

(Smallest adult shell from Urfu vill. env.) to 12 mm

(adult shell form E Biak), minimum width (on first

non-decollate whorl) is 1.6 mm, maximum width (on

ultimate whorl) to 3.8 mm. The subadult specimen

from E Biak has 8 whorls, is 7.6 mm high. Except on

microscopically pitted embryonic whorls, axial ribs

are strong, with acute blades. Basal outline of the

peristome is subtruncate to slightly angular in this

species.

pp. 135-184

Distribution: The Papuan Region. Known only

from Biak, Cenderawasih Bay Islands (Map 2).

Taheitia galactodes van Benthem Jutting, 1963

(Fig. 11)

Bibliography: van Benthem Jutting (1963: 442, fig. 7),

original description, record (Cenderawasih Bay Islands,

Biak: “Bosnik, coral reef near sago swamp”).

New material: INDONESIA E, Papua _ Prov.,

Cenderawasih Bay, Biak Is. W, Biak ~16 km W, Urfu

Vill, “ODO 971225, 135: 55;03%E, Se. 5am, 21 Wlz2018;

coastal limestone cliff, in cavities with vegetation and

debris [18 adult, 2 subadult & 4 juv., MBBJ, NME & KGC].

Additional description: The largest adult

decollate shell from W Biak (NME) is 9.5 mm high

and 3.4 mm wide. Total number of whorls is 12,

number of embryonic whorls is two.

Distribution: The Papuan Region. Known only

from Biak, Cenderawasih Bay Islands (Map 2).

Taheitia gebeensis sp. nov. (Fig. 12)

http://zoobank.org/A3/79A407-837C-4F34-8A5C-

CA6F6985D962

Holotype NME: INDONESIA E, Maluku Utara Prov.,

Gebe Island, on the way from Kacepi to Umera vill., 29.X-

03.XI.2011, secondary lowland rainforest on limestone,

leg. L.Wagner.

Paratype: 1 adult KGC: same as holotype.

Same specimens were previously reported as 7.

gracilenta (Greke 2012).

Derivatio nominis: Toponymic from the locus

typicus, Gebe Island.

Measurements: Holotype H = 10.4 mm, W = 3

mm, HW = 4.1 mm, AH = 2.1 mm. Paratype shell is

Slightly shorter.

Description: Shell is decollate, creamy-pink

with whitish axial ribs, dextral, very narrow, almost

parallel-sided, slightly and gradually narrowing

apicad. Decollate adult shell with 6 slightly convex

whorls, number of embryonic whorls is unknown.

Suture is moderately deeply impressed, oblique

to coiling axis (angle ~80°). Ultimate whorl is

as wide as penultimate whorl in apertural view

(maximum width 2.6 mm vs 2.55 mm), decollate

Shell is 6 whorls, first non-decollate whorl is 1.9

mm wide. Umbilicus is closed in adult. Teleoconch

is sculptured with widely spaced nearly straight (in

frontal view) to slightly irregularly sinuous and more

or less regular axial ribs. Ribs are slightly curved,

higher and wider near upper suture (Fig. 12D-E).

Some ribs do not run from suture to suture. There

are no abrupt changes in axial ribbing pattern, ribs

141

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

on last half whorl more widely spaced than ribs on

penultimate whorl. Ribs are not synchronous with

those of previous whorls and are generally parallel

to coiling axis. There are 3 ribs per 0.5 mm on

penultimate whorl in abapertural view and 33-35

ribs on ultimate whorl. Axial ribs on ultimate whorl

obtuse angulate basally; basal ridge not prominent

(ultimate whorl angled basally and laterally). Spiral

striae or ribs not observed at 80x magnification.

Last part of ultimate whorl projects slightly anteriad

from shell outline in apertural view. Parietal gap

not present. Aperture is obliquely ovoid (narrowed

in parietal-palatal angle), not tilted to coiling axis,

positioned subvertically, apertural rim entire, and

with flattened apertural lip, which is reflected

slightly basally. Aperture is shifted right to coiling

axis in apertural view. Peristome is simple in lateral

view. Parietal margin of peristome is detached from

ultimate whorl, peristome is therefore continuous.

Margins of peristome slightly sinuous palatalo-

basally. Operculum unknown.

Differential diagnosis: Taheitia gebeensis sp.

nov. is conchologically similar to T. biaka sp. nov.

(Biak; described above), T. gracilenta (E.A. Smith,

1897) and T. mansueta van Benthem Jutting, 1963

(both last - Doberai Peninsula of New Guinea) but

is readily differentiated by the combination of the

following features: the shell almost parallel-sided,

slowly narrowing apicad, ~33-35 axial ribs on the

ultimate whorl (50-51 in T. biaka sp. nov., 35-36

in T. gracilenta, 28-30 in T. mansueta), the base of

the ultimate whorl slightly angled (not angled in T.

biaka sp. nov. and T. mansueta, slightly angled in T.

gracilenta), the axial ribs significantly less dense on

all whorls (except the ultimate) than in T. biaka sp.

nov. and T. gracilenta, there are no abrupt changes

in the ribbing pattern on the ultimate whorl (axial

ribs on ultimate whorl twice as widely spaced as

rest of teleoconch in T. gracilenta), the decollated

spire of 6 whorls (6% in T. biaka sp. nov., 7 in T.

gracilenta, 7-8 in T. mansueta), the parietal gap

not present (Short gap present in T. mansueta), the

shell generally less high and less slender than in T.

biaka sp. nov. and T. gracilenta, the penultimate and

the ultimate whorl of same wide (the penultimate

whorl slightly narrower than the ultimate in T. biaka

sp. nov. and T. mansueta).

Ecology: Specimen sampled from wet leaf litter in

disturbed lowland rainforest on limestone.

Distribution: The Papuan Region.

Moluccas, Gebe (Map 1).

North

142

Taheitia gigantea sp. nov. (Fig. 13)

http://zoobank.org/8A0183A9-C86A-42F4-8B5B-

313465F3D5F2

Holotype NME: INDONESIA E, Raja Ampat, Waigeo

Island, Majalibit Bay, Waisai 19 km NE, River Werabiai

valley, 00° 18’02”S, 130° 56’00’E, 40-60 m, 20.11.2012,

primary lowland rainforest on limestone.

Derivatio nominis: Hitherto the largest

truncatellid, this species named from Latin

“giganteus” (giant) for its extraordinarily large shell.

Measurements: Holotype H = 31 mm, W = 8.1

mm, HW = 13.2 mm, AH = 6.2 mm.

Description: The holotype is in poor condition,

decollate, white, dextral, conical, gradually

narrowing apicad. Decollate adult shell with 6%

rather flat whorls, number of embryonic whorls is

unknown. Suture is slightly impressed, oblique

to coiling axis (angle at ~80°). Ultimate whorl in

apertural view is 6.9 mm wide, penultimate whorl

- 6.3 mm, topmost remaining (first non-decollate

of almost 7 remaining whorls) whorl is 3.7 mm

wide. Umbilicus is closed in adult. Teleoconch

is sculptured with very dense sinuous (in frontal

view), rather delicate and in part irregular axial

ribs. Some ribs increase in width at suture. Ribs

do not always run from suture to suture. There are

no abrupt changes in axial ribbing pattern, ribs on

the last half whorl nearly as dense as those on

the penultimate whorl. Ribs are not synchronous

with those on previous whorls and are slightly

oblique to coiling axis. There are 5-6 ribs per 0.5

mm on penultimate whorl in abapertural view and

about 110-115 ribs on ultimate whorl. Axial ribs

of ultimate whorl evenly rounded basally, basal

ridge is therefore absent (ultimate whorl rounded,

not angled, basally and laterally). Spiral striae or

ribs not observed at 80x magnification (possibly,

because of poor condition of shell); considering how

dense axial ribbing pattern is, there is literally no

space left for spiral sculpture). Last part of ultimate

whorl protruding slightly anteriad from shell outline

in apertural view. Parietal gap present, broad and

very deep. Aperture is obliquely ovoid (narrowed in

angular corner), not tilted to coiling axis, positioned

vertically, apertural rim entire, and with broadly

reflected, flattened apertural lip. Aperture is shifted

right to coiling axis in apertural view. Peristome is

simple in lateral view. Parietal margin of peristome

is detached from ultimate whorl, peristome is

therefore continuous. Angular corner of peristome

broadly detached from penultimate whorl (Fig. 13).

Palatal margin of peristome sinuous. Operculum

unknown.

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Differential diagnosis: Taheitia gigantea sp.

nov. is undoubtedly closely related to T. heinrichi

Gittenberger, 1898 (Doberai Peninsula, Salawati

& Warir islands), 7. telnovi sp. nov. (Misool),

and T. wallacei (H. Adams, 1865) (Waigeo). The

only available shell of the new species is in poor

condition, however, differs from aforementioned

species in the more delicate, denser and sinuous

axial ribbing pattern, the stronger reflexed apertural

lip and in the significantly larger shell (it seems

unlikely that the same Taheitia species, even in

disparate natural conditions, develops a shell 1/3

higher than average for the species). Discovering

more specimens and studying the operculum

should provide additional differential features of T.

gigantea sp. nov.

Ecology: Primary lowland rainforests on limestone,

wet leaf litter.

Distribution: The Papuan Region. Raja Ampat

Islands, Waigeo, eastern peninsula (Map 6).

Taheitia gracilenta (E. A. Smith, 1897) (Fig. 14)

Bibliography: Smith E.A. (1897: 289, pl. 17 figs 16-17),

original description as Truncatella, record (New Guinea,

Doberai Peninsula: “Andai”); Clench & Turner (1948:

191 & 200) new combination, checklist; Turner (1959:

182, pl. 34 fig. 4), lectotype designation, redescription,

records (New Guinea, Doberai Peninsula: “Andai”;

Bismarck Archipelago: New Britain, New Ireland); van

Benthem Jutting (1963: 440), records (New Guinea,

Doberai Peninsula: “Teminabuan”; Cenderawasih Bay

Islands, Biak: “Bosnik, coral reef near sago swamp”,

“vicinity of Nica camp, in deserted garden overgrown with

grass”, “S Biak, hospital cave above kampong Parai”).

Type material: lectotype [here designated] & 5

paralectotypes BMNH, Truncatella gracilenta Smith

Type Andai, New Guinea (W.D.) 1897. 98.10.25.16-

21. [handwritten] / Type. gracilenta, Smith. New

Guinea. [handwritten]. The first label supplemented

with additional text in different colour and different

handwritting: [Type] "?" "(See Turner 1959, Occ. Papers

on Malacology, 2,pp.182-5)".

Ecology: This species may be salt-tolerant.

Distribution: The Papuan Region. Known from

Doberai Peninsula (locus typicus - Andai) (Map 7)

and Cenderawasih Bay Islands (Biak). Record from

North Moluccas (Gebe) by Greke (2012: 29) refer to

a new species, laheitia gebeensis sp. nov. Here we

agree with Turner (1959) who already questioned

the Bismarck Archipelago records of this species

by Rensch |. (1937: 629, 633 & 634, fig. 54), as

Truncatella (Taheitia), New Britain (Neupommern):

“Cormoranhuk(Luwelon)”, “Karlei”, “Malkongbach”,

“Wattokgebiet”; New Ireland (Neumecklenburg):

pp. 135-184

“Mittel-Neu-Mecklenburg”). The record from Biak

needs confirmation and may refer to T. biaka sp.

nov. (this locality is not mapped) (Map 7).

Taheitia heinrichi Gittenberger, 1989

Bibliography: Gittenberger (1989: 24, figs 1-2, 4),

original description, records (New Quinea, Doberai

Peninsula: “5.5 km ENE of Klamono at the drilling

site Klamoekoek, 131°33’E 1°06.5’S; 6 km ENE of

Klamono at the drilling site Klawilis, 131°34’E 1°07’S;

16 km NNW of Klamono at the drilling site Klamesin,

131 °26’E 0°59’S”; Raja Ampat Islands, Salawati; Warir:

“island Warir off the east coast of the island Salawati, 7

km SE of Samate, 131°07’E 1°01’S”).

Distribution: The Papuan Region. Known from

southern lowlands of New Guinea’s' Doberai

Peninsula and Raja Ampat Islands (Salawati (locus

typicus), Warir (locus typicus)). It is likely the record

of Taheitia wallacei from Salawati by van Benthem

Jutting (1963: 441 “Waileh district”) is this species

(Map 7).

Taheitia jodiae sp. nov. (Figs 15, 28 & 31)

http://zoobank.org/46 /OCD4A-E5F8-4833-9188-

BD6B8CC884D4

Holotype UF 549960: PAPUA NEW GUINEA, East New

Britain Province, large cave 11 km N of Marmar Village

on E side of trail to Pakia Village, -5.444°, 151.465°

John Slapcinsky, 24 February 2005.

Paratypes: UF 366599 (45 adults), UF 366600 (13

whole in 75% ethanol) as holotype.

Derivatio nominis: Matronymic. Named for Jodi

Slapcinsky, wife of the second author. Without her

support long field excursions in remote areas would

not have been possible.

Measurements: Holotype SH = 15.25 mm, SW =

5.06 mm, AH=4.91, AW=3.65.

Description: Shell is dextral, conical, gradually

expanding, not umbilicate, with approximately

11*/,-12°/, total whorls, half of which are lost in

adults. Suture is not deeply impressed, oblique to

the coiling axis at an angle at ~70°. Embryonic

shell of approximately 2 #4 smooth, cylindric whorls,

EW=1.10+0.03 mm, N=10; first whorl descends

slowly and has a moderately deep suture. Adult

shell is decollate, with 5*/,-6*/, broadly rounded

whorls, large, SH=14.88+1.00, SW=5.09+0.38,

AH=4.81+0.33, AW=3.47+0.32, N=21. Width

of first non-decollate whorl is 2.28+0.20 mm,

the penultimate whorl is 3.944+40.31 mm, and

the ultimate whorl is 4.32+0.27 mm. Aperture is

deflected slightly downward and peripherally and

detaches from the penultimate whorl. Aperture is

143

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half-moon shape, flat along parietal and columellar

margin and broadly rounded along palatal margin.

Peristome dirty white and entire, reflected slightly

palatally, and broadly basally and along columellar

margin and then narrowing parietally. Teleoconch

whorls dirty white, after third whorl shell marked

with alternating irregular dark brown axial stripes.

Stripes strongest near sutures and_ usually

broken into zig-zag lines between sutures. First

four teleoconch whorls sculptured with regular

slightly arched axial ribs, about same width as

rib-interspaces. After fifth whorl ribs become

progressively more sinuous, pustulose and weaker.

After peripenultimate whorl ribs are only clearly

defined near suture where some are also thicker

and whiter. Rest of middle of penultimate and

ultimate whorls are sculptured with elongate

pustules that are oriented axially or diagonally

(Fig. 15D). Ribs thicken and then weaken abruptly

creating a well-defined basal ridge. Spiral sculpture

not observed at 80x magnification. Operculum (Fig.

15E) irregularly ovoid, height 3.144 mm, maximum

width 1.64 mm. Chitinous, paucispiral base bears

calcareous plate with strong, unicuspid, hook-

Shaped apophysis. Outer margin slightly upturned.

Between periphery and apophysis there is irregular

and often interrupted lamellar ridge terminating in

triangular projection with narrow base almost in

centre of operculum. Radula (Fig. 31) formula 2-1-

1-1-2. Rachidian with large central cusp flanked

on each side by three small near-basal ectocones.

Lateral teeth wide, asymmetric, with poorly defined

ectocones. Inner marginal with 6 finger like cusps.

Outer marginal with 3 bluntly pointed cusps.

Differential diagnosis: Taheitia jodiae sp. nov.

has a colour pattern of alternating dark and light

rectangular markings below the suture and irregular

dark markings on the rest of the whorls similar only

to T. schneideri (|. Rensch, 1937) from New Britain

and T. orrae Turner, 1959, from Biak and unlike

most Papuan Taheitia which are either unicoloured

or marked with axial bars. Taheitia jodiae has a

peristome that is free of the penultimate whorl,

expanded but not recurved abaperturally unlike

the peristome of T. orrae which is appressed to the

penultimate whorl, or T. schneideri the free margin

of which is abaperturally recurved.

Ecology: Found in only one location, but locally

abundant in tropical rainforest in a ravine under

piles of mossy limestone rocks outside the entrance

of a large deep cave that emitted cool moist air.

Distribution: The Papuan Region. New Britain

Island, Jacquinot Bay area (Map 3).

144

Taheitia longpela sp. nov. (Fig. 16)

http://zoobank.org/ACF86BB4-56 /A-44CC-8791-

1B9C2FAGB5DB

Holotype UF 366528: PAPUA NEW GUINEA, East New

Britain Province, large cave 11 km N of Marmar Village

on E side of trail to Pakia Village, -5.444°, 151.465°

John Slapcinsky, 24 February 2005.

Derivatio nominis: Longpela means tall in Tok

Pisin, the most widely spoken language in Papua

New Guinea.

Measurements: Holotype SH = 10.54 mm, SW =

2.69, AH=2.46, AW=1.68 mm.

Description: Shell is dextral, conical, gradually

expanding, not umbilicate. Total number of whorls

is unknown because only a single decollate adult

is known. Suture is moderately impressed, oblique

to coiling axis at angle at ~70°. Embryonic shell is

unknown. Adult shell is decollate, with 8 broadly

rounded whorls. Width of first non-decollate

whorl is 1.45 mm, penultimate whorl is 2.15

mm, and ultimate whorl is 2.29 mm. Aperture is

not deflected downward and peripherally. Palatal

margin of peristome is attached to penultimate

whorl. Aperture is almost rectangular, flat along

palatal and basal margins. Parietal and columellar

margins are sinuous recessed. Parietal margin

curves into aperture. Shell worn and no colour

description is possible. Sculptured with regular,

slightly arched axial ribs, that are slightly narrower

than rib-interspaces. Ribs maintain their strength

and consistency throughout teleoconch whorls (Fig.

16). Ribs are not thickened at sutures. No basal

ridge is present. Spiral sculpture was not observed

at 80x magnification although shell is chalky

and fine shell sculpture is not visible. Operculum

unknown.

Differential diagnosis: Taheitia longpela sp.

nov. has a tall narrow decollate shell of more than

seven very gradually expanding whorls like the shell

of 7. gracilenta (E. A. Smith, 1897) from Doberai

Peninsula of New Guinea. The two species differ in

the shape of their whorls which are flattened in T.

gracilenta and rounded in T. longpela sp. nov. The

apertures of both species also differ. The parietal

edge of the peristome in T. gracilenta meets the

outer lip at the opening of the aperture whereas the

parietal edge of the peristome of T. longpela meets

the outer lip well inside the aperture.

Ecology: Single long dead shell found in leaf litter

among moss covered limestone rocks outside the

entrance of a large deep cave that emitted cool

moist air.

Distribution: The Papuan Region. New Britain

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GrEkE, K. & Starcinsky, J.: New Taheitia H. et A. Adams, 1863, with revisional notes on the Papuan Truncatellidae ...

Island, Jacquinot Bay area (Map 3).

Taheitia malagan sp. nov. (Fig. 17)

http://zoobank.org/31D101FE-OF1D-414E-8648-

94279398893B

Holotype UF 475675: PAPUA NEW GUINEA, New

Ireland Province, Nusa Island off Kavieng -2.5822°,

150.7772°, overgrown coconut plantation, John

Slapcinsky and Cindy Bick, 27 November 2012.

Paratype: UF 563274 (1 whole in 75% ethanol) as

holotype.

Derivatio nominis: Malagan is from the Nalik

language of northwestern New Ireland and refers

both to funerary carvingS and the ceremony

where they are displayed. Named for this species

operculum, which resembles an intricately carved

malagan shield.

Measurements: Holotype SH = 8.03 mm, SW =

3.11 mm, AH=2.66 mm, AW=1.93 mm.

Description: Shell is dextral, conical, gradually

expanding, not umbilicate, with approximately 9

total whorls, half of which are lost in adults. Suture

moderately impressed, oblique to coiling axis at an

angle at ~70°. Embryonic shell of approximately

2¥%2 smooth, cylindric whorls, EW=0.93 mm; first

whorl descends slowly and has a moderately

Shallow suture. Adult shell is decollate, with 4*/,

broadly rounded whorls, large, SH=8.31+0.40,

SW=3.22+0.15, AH=2.75+0.13, AW=2.00+0.10,

N=2. Width of first non-decollate whorl is 1.74+0.08

mm, penultimate whorl is 2.64+0.06 mm, and

ultimate whorl is 2.8740.04 mm. Aperture

is deflected peripherally and detaches from

penultimate whorl. Aperture is teardrop shaped,

angled and narrow at detached sutural margin and

broadening and slightly flattened basally. Peristome

white and entire, reflected slightly palatally and

basally but notalong columellar and parietal margin.

Teleoconch whorls dirty white, final three whorls

also marked with irregular narrow axial bands of

grey. Stripes extend from suture to midpoint of whorl

and less commonly to lower suture. Teleoconch

whorls are sculptured with regular slightly arched

axial ribs, about same width as rib-interspaces

which extend from suture to suture. On final three

whorls some ribs are thickened and white at suture

making whorl shoulders nodulate (Fig. 17D). Base

somewhat angled but not clearly ridged. Spiral

sculpture not observed at 80x magnification.

Operculum (Fig. 17E) irregularly ovoid, height

1.56 mm, maximum width 1.04 mm. Chitinous,

paucispiral base bears calcareous plate with

strong, unicuspid, hook shaped apophysis. Outer

pp. 135-184

Between periphery and apophysis there is single

row of triangular projections, which extend from

base nearly to parietal margin. Each projection

has two wing shaped tips and row of projections

resembling a line of birds in flight.

Differential diagnosis: Taheitia malagan sp.

nov. with a height of approximately 8 mm is one

of the smallest Taheitia species in the Papuan

Region, larger only than T. bismarkiana (I. Rensch,

1937) (approximately 5 mm) from Duke of York

Island which differs in having the parietal edge of

the peristome appressed, and similar in size to T.

galactodes van Benthem Jutting, 1963 from Biak

which differs from T. malagan sp. nov. in having the

basal-columellar edge of the peristome expanded

forming a wing-like projection. The operculum of T.

malagan sp. nov. has a row of raised projections

the tops of which are bifurcated, each tip acutely

pointed and flattened parallel to the operculum

surface, forming gull wing like structures. While

several species have rows of projections on the

outer surface of the operculum these usually

consist of columns with oval crossections as seen

in 7. ultima (|. Rensch, 1937), and T. jodiae sp.

nov. or form lamellar ridges as seen in P. heinrichi

Gittenberger, 1989 from Salawati and Warir islands

and Doberai Peninsula of New Guinea.

Ecology: Under limestone rocks and coconut leaf

litter in abandoned coconut plantation.

Distribution: The Papuan Region. Nusa Island off

New Ireland Island (Map 3).

Taheitia mansueta van Benthem Jutting, 1963

(Fig. 18)

Bibliography: van Benthem Jutting (1963: 445, fig.

10), original description, records (New Guinea, Doberai

Peninsula: “Manokwari, along the beach near Uriami

River”, “Pasirputin near Manokwari, on fallen tree

trunks”); Egorov (2018: 34, fig. 57E), as T. mansuata

[sic!], record (New Guinea, Doberai Peninsula:

“Manokwari, Pasirputih”).

Distribution: The Papuan Region. Known from

northern part of New Guinea’s Doberai Peninsula

(locus typicus) (Map 7).

Taheitia orrae Turner, 1959

Bibliography: Turner (1959: 186, pl. 31 figs 1-2),

original description, record (Cenderawasih Bay Islands,

Biak: “under fern roots in a large sink-hole cave behind

the air strip, Biak”); van Benthem Jutting (1963: 441),

records (New Guinea, Doberai Peninsula: “Sorong,

Cape Nuijew”, “region of the Klawilu canyon”, “along

road from Ajamaru to Teminabuan”; “New Guinea”;

Cenderawasih Bay Islands, Biak: “under fern roots in a

145

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

large sink-hole cave behind the air strip”; Raja Ampat

Islands, Salawati: “Waileh district”); Gittenberger (1989:

24, fig. 3), holotype image.

Note: The type specimens are 12.4-12.8 mm

high and 4 mm wide (Turner 1959; Gittenberger

1989). Van Benthem Jutting (1963) gives a wide

variety of heights for her presumably decollate

specimens from Klawili env. (14.5-21.5 mm),

Sorong (18-20 mm) and Salawati (17.6-22 mm).

We had no opportunity to study these specimens

but, considering the large shell height, it is not

impossible that aforementioned specimens are

aberrant Taheitia heinrichi, which seems not

uncommon in the same area.

Distribution: The Papuan Region. Known from

lowlands of New Guinea’s Doberai Peninsula,

Cenderawasih Bay Islands (Biak (locus typicus)) and

Raja Ampat Islands (Salawati). However, all records

by van Benthem Jutting (1963) from outside Biak

considered dubious and need further confirmation

(Map 2).

Taheitia scalariformis (Reeve, 1842)

Note: Solem (1960) mentioned this species

from Nggela (Florida) Island, Solomon Islands,

commenting that identification is uncertain and

studied available specimen is dissimilar to those

of typical T. scalariformis (originally described from

the Tuamotu Archipelago). We therefore ignore this

otherwise highly interesting record.

Taheitia schneideri (1. Rensch, 1937) (Figs 19-20

& 32)

Bibliography: Rensch I. (1937: 628 & 633, figs

51-52), original description as Truncatella, records

(Bismarck Archipelago, New Britain (Neupommern):

“Jacquinotbucht (Malekur, Mailmail)”); Clench & Turner

(1948: 191 & 196), new combination, checklist.

Holotype MFNB by original designation: Mailmail

[handwritten] / Mailmail Jacquinotbucht [handwritten]

[upperside of the label] leg. P. Schneider Truncatella

schneideri |. R. [handwritten] [underside of the label] /

Zoolog. Museum Berlin [printed] Truncatella schneideri

|. Rensch Mailmail 97962 P.Schneider [handwritten].

Supposed holotype shell is broken, kept in a separate

capsule without any specific marking. On the fig. 54 in

the original description (Rensch 1937: 628) the holotype

shell is figured unbroken.

Paratypes 1/7 specimens MFNB: same labels as

holotype. Rensch (1937) mentions two series of type

specimens, from Malekur and Mailmail. Malekur

specimens were not available for the present study.

However, Rensch (1937: 629) selected the specimen

146

from mailmail as the "type" (here considered holotype).

New material: UF 366522 (24 adult shells), UF

546812 (8 juvenile shells) UF 366527 (18 whole in 75%

ethanol): Papua New Guinea, East New Britain Province,

4 km NW of Marmar Village along trail to Pakia Village,

-5.49651, 151.49027, John Slapcinsky, 4 March 2005.

Measurements: Holotype (decollated shell) H =

15.7 mm, W = 6.9 mm, from original description.

Adult decollated paratypes H = 14-18.3 mm.

Redescription: Shell is dextral, conical, gradually

expanding, not umbilicate, with approximately

11-12 total whorls, half of which are lost in

adults. Suture is not deeply impressed, oblique

to coiling axis at angle at ~70°. Embryonic shell

IS approximately 2% smooth, cylindric whorls,

EW=1.2840.08 mm, N=3; first whorl is planar or

even sunken with very deep sutures. Adult shell is

decollate, with 5-6 broadly rounded whorls, large,

SH=22.1841.09, SW=8.7240.37, AH=8.31+0.44,

AW=6.91+0.46, N=23. Width of first non-decollate

whorl is 3.3840.32 mm, penultimate whorl is

6.29+0.30 mm, and ultimate whorl is 6.87+40.34

mm. Aperture is deflected downward and

peripherally and detaches from penultimate whorl.

Aperture is half-moon shape angled at sutural

margin and flattened along columellar margin.

Peristome is entire, broadly reflected, flattened

and dark grey from periphery towards base where

it is slightly curved abaperturally. Peristome is

only slightly reflected along columellar margin.

Teleoconch whorls light brown, after third whorl

shell marked with alternating irregular dark brown

axial stripes. Stripes strongest near sutures and

usually broken into zig-zag lines between sutures.

First four teleoconch whorls are sculptured with

regular slightly arched axial ribs, each about

one-third width of rib-interspaces. After fifth

whorl ribs become progressively more sinuous,

pustulose and weaker. After peripenultimate

whorl ribs are only clearly defined near suture.

Ultimate and penultimate whorls sculptured

with elongate pustules, that are oriented axially

but also less commonly diagonally (Fig. 20D). In

many individuals the surface sculpture is weak or

worn on the last two whorls and surface is nearly

smooth. Basal ridge is absent and spiral sculpture

not observed at 80x magnification. Operculum (Fig.

20E) irregularly ovoid, height 5.14 mm, maximum

width 2.82 mm. Chitinous, paucispiral base bears

calcareous plate with strong, unicuspid, triangular

apophysis at base. Outer margin rimmed with thin

raised lamellar ridge. Between peripheral ridge and

apophysis there is strong irregular lamellar ridge

02-Jun-21 21:48:26

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Greke, K. & Starpcinsky, J.: New Taheitia H. et A. Adams, 1863, with revisional notes on the Papuan Truncatellidae ...

with several poorly defined peaks. Radula (Fig. 32)

formula 2-1-1-1-2. Rachidian with large central

cusp flanked on each side by 4 small near-basal

ectocones. Lateral teeth wide, asymmetric, with

poorly defined ectocones. Inner marginal with 4-5

finger like cusps. Outer marginal with 3 short bluntly

pointed cusps.

Differential diagnosis: Taheitia schneideri

has a colour pattern of alternating dark and light

rectangular markings below the suture and irregular

dark markings on the rest of the whorls similar

only to T. jodiae sp. nov. from New Britain and T.

orrae from Biak, and unlike most Papuan Taheitia

which are either unicoloured or marked with axial

bars. Taheitia schneideri has a peristome that is

free of the penultimate whorl, darkly pigmented

and broadly expanded and recurved abaperturally

unlike the peristome of T. orrae which is white, not

expanded, and attached to the penultimate whorl

or T. jodiae sp. nov. which is white, not as expanded,

and not abaperturally recurved.

Ecology: Rainforest under limestone rocks and in

leaf litter among limestone rocks.

Distribution: The Papuan Region. New Britain

Island, Jacquinot Bay area.

Taheitia telnovi sp. nov. (Figs 21 & 33)

http://zoobank.org/OBD /42EF-03B6-45AF-BCOF-

9128050CDB9E

Holotype NME: INDONESIA E, Raja Ampat, Misool

Island (central), River Gam upstream, Gamta vill. 12-

14 km NW, 01°57’20”S, 130°11’04’E, 105-325 m,

04-06.11.2012, primary lowland rainforest on limestone

ridge. Paratypes 20 specimens. Two adults MBBJ, 8

adults & 1 subadult NME, 7 adults & 41 juv., KGC: same

as holotype; 1 adult NME & 2 adults, 1 juv., 1 adult shell

fragment KGC: INDONESIA E, Raja Ampat, Misool Island

(central), River Gam upstream, Gamta vill. 12-14 km

NW, 01.°57’45"S, 130° 10’59”E, 80-85 m, 03.IV.2009,

primary lowland rainforest on limestone, leaf litter; 1

adult KGC: INDONESIA E, Raja Ampat, Misool Island

S, Biga vill. ~7,5 km W, River Biga valley, 02°01’23"S,

130°12’38’E, 45-78 m, 03.11.2012, primary lowland

rainforest on limestone.

Derivatio nominis: Patronymic. Named in honour

of Dr. Dmitry Telnov (The Entomological Society of

Latvia, Riga), famous coleopterist, taxonomist,

biogeographer and experienced specialist on the

Papuan Region and Wallacea, the chief editor of

this book series.

Measurements: Holotype H = 18 mm, W =

5.2 mm, HW = 7.5 mm, AH = 4.5 mm. Selected

paratypes 15.2 x 4.9 mm (River Biga valley), 17.4 x

5.1 mm, 18.5 x 5. Imm, 19 x 5.1 mm (data same

pp. 135-184

as holotype).

Description: Shell is decollate, cream coloured,

dextral, conical, gradually narrowing apicad.

Decollate adult shell with 512-7 rather flat whorls,

number of embryonic whorls is about two. Total

number of whorlsis about 12%. Suture is moderately

deeply impressed, oblique to coiling axis (angle at

~70-80°). Ultimate whorl in apertural view is 4.6

mm wide, penultimate whorl - 4.2 mm wide. Of

seven remaining whorls the topmost remaining

whorl is 2.4 mm wide. Umbilicus is closed in adult.

Teleoconch is sculptured with dense nearly straight

to slightly arched or sinuous (in frontal view) and

in part irregular axial ribs. Some ribs increase in

width at the sutures, especially the upper suture

(Fig. 21D). Ribs do not always run from suture to

suture. There are no abrupt changes in axial ribbing

pattern, ribs on last half whorl slightly less dense

than those on penultimate whorl. Ribs are not

synchronous with those of previous whorls and are

slightly oblique to coiling axis. There are 4-5 ribs

per 0.5 mm on penultimate whorl in abapertural

view and about 90-95 ribs on ultimate whorl. Axial

ribs on ultimate whorl evenly rounded basally, basal

ridge is therefore absent (ultimate whorl rounded,

not angled, basally and laterally). Spiral striae or

rips not observed at 80x magnification. Last part

of ultimate whorl protruding slightly anteriad from

Shell outline in apertural view. Parietal gap present,

variable wide and deep. Aperture is obliquely ovoid

(narrowed in angular corner), not tilted to the coiling

axis, positioned vertically, apertural rim entire,

and with moderately broadly reflected, flattened

apertural lip. Aperture is shifted right to coiling

axis in apertural view. Peristome is simple in lateral

view. Parietal margin of peristome is detached from

ultimate whorl, peristome is therefore continuous.

Angular corner of peristome broadly detached

from penultimate whorl (Fig. 217A). Margins of

peristome not or slightly sinuous; parietal margin

is slightly obtusely angulate. Operculum (Fig.

21E-G) irregularly ovoid, height 3 mm, maximum

width 1.9 mm, slightly concave on outer surface. Its

chitinous, paucispiral base bears small calcareous

plate with unicuspidate apophysis provided with

strong cow-horn-shaped-curved projection, which

is longitudinally grooved on inner margin (one

directed inside the shell), without lamellar ridges.

Radula (Fig. 33) formula 2-1-1-1-2. Rachidian

with large central cusp flanked on each side by

three small near-basal ectocones. Lateral teeth

wide, asymmetric, with relatively well defined short

pointed ectocones. Inner marginal with 3 short

finger like cusps. Outer marginal with 3 short broad,

Ree

147

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

rounded cusps.

Differential diagnosis: Taheitia telnovi sp. nov.

is undoubtedly closely related to T. gracilenta (E.A.

Smith, 1897) (Doberai Peninsula, Biak, Gebe), T.

heinrichi Gittenberger, 1898 (Doberai Peninsula,

Salawati & Warir islands) and T. wallacei (H. Adams,

1865) (Waigeo) but readily differentiated primarily

by operculum morphology, which has the large

horn-like apophysis and has no raised ridges (not

like in T. heinrichi and T. wallacei), being generally

larger than T. gracilenta (15-18 vs 11.5-14 mm),

having rounded, non-angulate shell base (the shell

base slightly angulate in T. gracilenta) and the axial

ribs not or slightly sparser on the last half whorl (the

axial rioS sparser on the last half whorl than on the

rest of teleoconch in T. gracilenta).

Ecology: Lowland rainforests on limestone and

vegetated limestone cliffs. On Misool, this species

is recorded far inland. Some specimens were found

at foot of the limestone ridge, but most - at the

elevation 300-350 m directly on the watershed on

a dry limestone cliff with small patches of soil and

leaf litter (Fig. 29).

Distribution: The Papuan Region. Raja Ampat

Islands, Misool (Map 5). We consider records of T.

wallacei from Misool (“Fakal”, “between Waigama

and Waima”, “Waima”) by van Benthem Jutting

(1958: 322; 1963: 441) as referring to the new

species as well.

Taheitia tesselata Mollendorff, 1897 (Fig. 22)

Bibliography: Modllendorff (1897: 32), original

description, record (Bismarck Archipelago); Rensch

|. (1937: 630, 633 & 634), additional description as

Truncatella (Taheitia), records (Bismarck Archipelago,

Massait (Masahet); New Britain (Neupommern):

“Ghoghuwulou”, “Jacquinotbucht (Pomeo, Mailmail)”,

“Kap Dampier”, “Matong”, “Nangurup”; New Ireland

(Neumecklenburg): “Ugana”, “Ulaputur”); Clench &

Turner (1948: 192 & 208), checklist, in part (p. 192) as

T. tessellata [sic!] Quadras [sic!] and v. Mollendorff.

Holotype: Not studied.

Studied material: TrumeateHa aheitia [corrected by

hand] tesselata (BItg. Ms) Mollendorff [sic!] Nachricht.

1897 New Pommern Bismark [sic!] Archipelago near New

Guinea 94.11.23.13-16 B. Strubell Esq. [handwritten]

/ tesselata, Bitg. Bismark [sic!] Arch. B.Strubell Esq

[handwritten].

Notes: The studied specimen is possible syntype

exchanged from M6llendorff's collection. We

provide images of this species for the first time.

Additional description based on the specimen

from "New Pommern" (now New Britain): Shell is

—_—_

148

decollate, high conical. Adult decollate shell of 5

rather flat whorls. Suture deep. Axial ribs strong

and dense, generally sinuous on penultimate and

ultimate whorl, more regularly straight on older

whorls, dense, generally separated by own width.

Ribs are not synchronous with those of previous

whorls and are generally parallel to coiling axis. No

changes in ribbing pattern on ultimate whorl. Spiral

Striae or ribs not observed. Parietal gap not present.

Aperture is drop-shaped, rounded along parietal

and palatal margin. Aperture not tilted to coiling

axis, positioned subvertically, apertural rim entire,

and with flattened apertural lip, which is reflected

slightly basally. Aperture is shifted right to the coiling

axis in apertural view. Peristome is simple in lateral

view. Parietal margin of peristome is attached to

ultimate whorl, peristome is continuous. Palatal

margins of peristome slightly sinuous. Operculum

unknown.

Distribution: The Papuan Region. Known from

Bismarck Archipelago (Massait, New Britain, New

Ireland). Locus typicus Is not specified exactly (“in

archipelago Bismarckiano”).

Taheitia ultima (I. Rensch, 1937) (Figs 23-24 &

34)

Bibliography: Rensch |. (1937: 629 & 633, fig.

53), original description as Truncatella, records

(Bismarck Archipelago, New Britain (Neupommern):

“Cormoranhuk”, “Jacquinotbucht”); Clench & Turner

(1948: 191 & 209), new combination, checklist.

Holotype MFNB: Not studied.

Paratype 1 specimen MFNB: Zoolog. Museum Berlin

[printed] Truncatella ultima |. Rensch Jacquinot Bucht

Neu Pommern 97959 Pater Schneider [handwritten].

According to Rensch (1937: 629), this species was

originally described from nine specimens from two

localities.

New material: Papua New Guinea, East New Britain

Province: UF 366523 (10 whole in 75% ethanol), UF

366525 (15 adult shells) UF 539697 (18 juvenile

shells), Renis Peninsula, E of Galowe Village, 5.5207 °

S, 151.4833° E, 1 meter, John Slapcinsky, 6.111.2005,

primary coastal forest; UF 366524 (3 shells), Marmar

Village, 1 km W of Pomio, 5.5183° E, 151.5067° E.,

20 meters, John Slapcinsky, 21-Il-2005, hill forest and

gardens; UF 366526 (5 shells), W bank of Pomio River, 1

km E of Marmar Village, 5.5153° S, 151.5111° E, 0-65

meters, primary and secondary hill forest.

Measurements: Holotype H = 17 mm, W = 6.1

mm, from the original description. Studied paratype

H = 16.5 mm.

Redescription: Shell is dextral, conical, gradually

expanding, not umbilicate, with approximately

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11-12 total whorls, half of which are lost in

adults. Suture is not deeply impressed, oblique

to coiling axis at angle of ~70°. Embryonic shell

IS approximately 2% smooth, cylindric whorls,

EW=1.01+0.05 mm, N=8; first whorl is planar with

deep sutures. Adult shell is decollate, with 5-6

broadly rounded whorls, large, SH=14.53+1.00,

SW=5./70+0.37, AH=5.08+40.34, AW=3.84+0.28,

N=14. Width of first non-decollate whorl is

2.52+0.26 mm, penultimate whorl is 4.19+0.26

mm, and ultimate whorl is 4.59+0.30 mm. Aperture

is deflected peripherally and detaches from

penultimate whorl. Aperture is half-moon shape,

nearly flat but sinuous along parietal and columellar

margin and broadly rounded along palatal margin.

Peristome dirty white and entire, reflected slightly

palatally, and broadly basally and along columellar

margin and then narrowing parietally. Teleoconch

whorls light brown, some individuals show very

occasional grey lateral stripes or blotches after

third whorl. Teleoconch whorls sculptured with

slightly arched axial ribs, each about same width

as rib-interspaces. After fifth whorl ribs become

progressively more sinuous, sometimes merging

with neighbouring ribs especially on penultimate

and ultimate whorls. Ribs usually extend from

suture to suture (Fig. 24D). Ribs weaken basally

especially on ultimate whorl but a basal ridge is not

clearly defined. Spiral sculpture not observed at

80x magnification. Operculum (Fig. 24E) irregularly

ovoid, height 3.28 mm, maximum width 1.87 mm.

Chitinous, paucispiral base bears calcareous plate

with strong, hook-shaped apophysis with basal

projection. Outer margin rimmed with thin raised

lamellar ridge. Between peripheral ridge and

apophysis there is row of several strong usually

triangular projections with hollowed tips. Radula

(Fig. 34) formula 2-1-1-1-2. Rachidian with

large central cusp flanked on each side by three

small near-basal ectocones. Lateral teeth wide,

asymmetric, with relatively well defined ectocones.

Inner marginal with 4-5 finger like cusps. Outer

marginal with 3 bluntly pointed cusps.

Differential diagnosis: The basal-columellar

edge of the peristome of Taheitia ultima sp. nov.

is expanded into a wing-like projection and the

parietal edge is free of the penultimate whorl like

T. jodiae and T. schneideri from New Britain as

well as 7. calcarata and T. galactodes from Biak.

However, Taheitia jodiae and T. schneideri differ in

having complex colour patterns and T. calcarata

and T. galactodes differ in being smaller and having

regular ribbing that does not become sinuous in

the ultimate and penultimate whorls.

pp. 135-184

Ecology: Under limestone rocks and leaf litter

among rocks in primary and secondary coastal and

hill forest at low elevation.

Distribution: The Papuan Region. New Britain

Island, Jacquinot Bay area. The record from

Cenderawasih Bay Islands (Yapen) by van

Benthem Jutting (1963: 441: “Serui, in gardens of

agricultural school” and “near brackish water lake

of Sarawandori, W of Serui”) is considered highly

doubtful given the very spotty distribution of most

Taheitia and their specialized habitat.

Taheitia wallacei (H. Adams, 1865) (Figs 25-26)

Bibliography: Adams H. (1865: 416, pl. 21 figs 13-

14), original description as Truncatella (Taheitia), record

(Raja Ampat Islands, Waigeo (Waigiou)); Wallace (1865:

414), record (Raja Ampat Islands, Waigeo: “Waigiou, on

limestone rocks”) [this record is not considered a nomen

nudum since the original description is published in

the very same paper a few pages later, see Adams H.

1865]; Tapparone Canefri (1883: 281), as Truncatella,

record (Raja Ampat Islands, Waigeo: “Waigheu, trovata

sugli scogli calcai”); Clench & Turner (1948: 192 & 209),

checklist; Turner (1959: 184, pl. 31 fig. 3), redescription,

general distribution; van Benthem Jutting (1963: 441),

bibliography.

Type material: Type material was probably never

properly designated, therefore only topotypes were

studied, 1 specimen BMNH - “Waigiou Island” /

1903. 7.11.38 [purchased off MRE. Gerrard]; 1 specimen

BMNH - “Waigiou, Molucca Is” / 1904.12.15.56-57

[collected by Mr Waterhast, purchased off Gerrard].

New material: INDONESIA E, Raja Ampat, Waigeo

Island, Waisai 3 km W, 00°26’04”S, 130°47’41’E,

40-50 m, 17.11.2012, secondary lowland rainforest

on limestone [3 adults (one is broken), NME & KGC];

INDONESIA E, Raja Ampat, Waigeo Island, Waisai 4-6 km

W (around Waisai airport), 00°24’46”"S, 130°44'11’E,

10-100 m, 17.11.2012, primary lowland rainforest on

limestone & karst [2 adults, NME & KGC]; INDONESIA

E, Raja Ampat, Waigeo Island, Waisai 10 km NWW,

00° 24’46"S, 130°44'11”E, 70-200 m, 19.11.2012,

primary lowland rainforest on limestone [3 adults, NME

& KGC].

Turner (1959) based her redescription of T.

wallacei on topotypic subadult specimens from

BMNH (Fig. 25), since the types were not allocated

(the types are not in BMNH - J. Ablett, personal

communication, 18.vill.2020). However, these

specimens are atypical or belong to a different

species, because they are significantly smaller than

mentioned in the original description (9.5-10.5 mm

vs 18 mm) and have denser axial ribbing pattern

on the ultimate whorl (not mentioned by Adams H.

1865). Recent specimens from Waigeo in the NME

oes

149

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collection perfectly match the original description

of this species. A neotype is not being designated

since the syntype(s) may still be present in BMNH.

Redescription based on specimen from Waisai

env., S Waigeo, Raja Ampat Islands (see Annotated

checklist below for details). H = 17.9 mm, W = 5.5

mm, HW=8.3 mm, AH =5.3 mm;selected specimen

from Waisai airport area H = 12.4 mm, W= 4.1mm,

HW =5.9 mm, AH=3.1mm. Shell is decollate, cream

coloured, dextral, conical, gradually narrowing

apicad. Decollate adult shell with 6-6% rather flat

whorls, number of embryonic whorls is unknown.

Suture is moderately deeply impressed, oblique to

coiling axis (angle at ~70-75°). Ultimate whorl in

apertural view is 3.9 mm wide, penultimate whorl

- 3.5 mm, topmost remaining (first non-decollate

of totally 6 whorls) is 1.65 mm wide. Umbilicus

is closed in adult. Teleoconch is sculptured with

dense nearly straight to slightly arched (in frontal

view) and in part irregular axial ribs. On ultimate

whorl riobS become slightly sinuous basally. Some

ribs are distinctly wider in their upper part at Suture

(Fig. 26C). Ribs do not always run from suture

to suture. There are no abrupt changes in axial

ribbing pattern, ribs on the last half whorl are as

dense as those on penultimate whorl. Ribs are not

synchronous with those of previous whorls and are

oblique to coiling axis. There are about 5 ribs per

0.5 mm on penultimate whorl in abapertural view

and about 80-93 ribs on ultimate whorl. Axial ribs

of ultimate whorl evenly rounded basally, basal

ridge is therefore absent (ultimate whorl rounded,

not angled, basally and laterally). Spiral striae or

ribs not observed at 80x magnification. Last part

of ultimate whorl protruding slightly anteriad from

shell outline in apertural view. Parietal gap present,

rather wide and comparatively deep. Aperture is

obliquely ovoid (narrowed in angular corner), not

tilted to coiling axis, positioned vertically, apertural

rim entire, and with moderately broadly reflected,

flattened apertural lip. Aperture is shifted right to

coiling axis in apertural view. Peristome is simple

in lateral view. Parietal margin of the peristome

is detached from ultimate whorl, peristome is

therefore continuous. Angular corner of peristome

attached to or broadly detached from penultimate

whorl (Fig. 26A). Margins of peristome not or

barely sinuous; parietal margin is slightly obtusely

angulate. Operculum (Fig. 26D-F) irregularly ovoid,

height 2.4 mm, maximum width 1.7 mm, flattened

on outer surface. Its chitinous, paucispiral base

bears calcareous plate with slightly raised ridge

along one margin and on opposite margin with

strong, unicuspid apophysis and 5 strong lamellar

150

elevations of variable size raising radially from

apophysis.

Distribution: The Papuan Region. Only known

from Waigeo, Raja Ampat Islands (Map 6).

Key to Papuan Taheitia H. et A. Adams, 1863

The present key is an attempt to summarize

critical features of the Papuan Region species

presently arranged to Taheitia in broad sense (see

short discussion above). Considering our generally

poor present knowledge on the Papuan fauna, it

is likely further species will be recorded from the

area in the future and, therefore, this key is only

valid for the Papuan taxa mentioned in the present

paper. This key is generally based on conchological

features. Shape and structure of operculum is a

very useful feature, but opercula are known only

for limited number of Papuan Taheitia, therefore

we generally avoided usage of opercula for treating

groups of species in the key except if opercula

known in all closely related species and proper

comparison was possible. Taking into account high

variability in shell height among some Papuan

Taheitia, some features used in this key may not

work sufficiently for extreme sized (smallest and

largest) specimens of same species. Taheitia is a

difficult group thanks to high variability of shell size

(as above), shape and ribbing pattern and usage of

comparative specimens are mandatory for certain

identification.

1 Shell H of adult decollated specimen 10 mm or more;

adult decollated spire of 5 or more whorls; peristome

not thickened, single, layers of previous peristomes not

visible in apertural view

— Adult decollate shell small, H distinctly under 10 mm;

adult decollated spire of less than 4 whorls; peristome

thickened, distinctly double with layers of previous

peristomes visible in apertural view (Fig. 7A-B)

T. bismarckiana [this is either Truncatella

or typical Taheitia sensu Adams and Adams (1863), with

strongly double peristome. Strongly dissimilar species

from exclusively terrestrial Papuan congeners]

2 Shell slender; decollate adult shell subcylindrical to

nearly elongate cylindrical (Figs 5,12, 14, 16 & 18); for

five last whorls the ratio of shell height to width is 3.5

or more; teleoconch whorls widen slightly and gradually;

ratio of maximum width of ultimate whorl (measured

above aperture) to maximum width of fourth preceding

whorl is 1.3 or less 3 (the ratio is 3.1-3.2 in T.

foliosocostata, this species treated in the next couplet)

- Shell less slender; decollate adult shell conical to

strongly subcylindrical (Figs 6, 8-11, 13, 15, 17, 19-

26); for five last whorls the ratio of shell height to width

is 3 or less; teleoconch whorls widen more rapidly; ratio

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of maximum width of ultimate whorl (measured above

aperture) to maximum width of fourth preceding whorl is

EE OEOIAIMOL Ce ease vcs aranamumne es cusamen nes aml term watarcas mua 7

3 In adult shell axial ribbing pattern is nearly same dense

or slightly sparser on the last half whorl as on the rest of

teleoconch

— In adult shell axial ribbing is significantly sparser on

the last half whorl than on the rest of teleoconch (Fig.

14B-C); shell SIENCErer .......c ce eeeeeeseneeeeeees T. gracilenta

4 Less than 40 axial ribs on ultimate whorl, counting both

complete and incomplete ribs; operculum is unknown ..

— Fifty or more axial ribs on ultimate whorl counting both

complete and incomplete ribs (Fig. 54-C); operculum on

inner side with 9 lamellar elevations (Fig. 5D & E) .........

T. biaka sp. nov.

5 Aperture more or less regularly ovoid, rounded along

the palatal and basal margins; the parietal margin not

curves into the aperture

- Aperture nearly rectangular, flat along palatal and

basal margins (Fig. 16A & D); parietal margin bent into

the aperture T. longpela sp. nov.

6 Axial ribs on ultimate whorl rounded basally (Fig. 18);

ridge at shell base not present (ultimate whorl rounded

basally and laterally); 28-30 axial ribs on the ultimate

whorl T. mansueta

- Axial ribs on ultimate whorl obtuse angulate basally;

weak ridge is present at shell base (ultimate whorl

angled basally and laterally) (Fig. 12); 30-35 axial ribs

on the ultimate whorl T. gebeensis sp. nov.

7 Shell height ~30 mm; axial ribbing pattern very dense;

axial ribs slightly sinuous, comparatively delicate; about

110-120 axial ribs on the ultimate WNOFI ........ ee eereeee

T. gigantea sp. nov.

- Shell height under 25 mm; axial ribbing pattern

moderately dense; ribs straight to slightly arched (only

become slightly sinuous at shell base); up to 95 ribs on

the ultimate whorl

8 Teleoconch with axial ribs longitudinally grooved (with

one or more delicate groves) along their crests, as if they

have been composed on two merged lamellae (Fig. 8D);

columellar-basal corner of peristome obtusely protruding

in a spur in adult shell (Figs 8A & 11A)

- Axial ribs of teleoconch not grooved longitudinally

along their crests; columellar-basal corner of peristome

OTe OK OUGUIGIN Se circcceceemeneennte ss beecmaaundiductuae cue tiicencemauseo ee 10

9 About 22-25 axial ribs on ultimate whorl; teleoconch

whorls comparatively stronger convex (Fig. 11)

T. galactodes

- About 32-34 axial ribs on ultimate whorl; teleoconch

whorls comparatively less convex (Fig. 8) ... T. calcarata

10 After peripenultimate whorl axial ribs are only clearly

defined near the suture, the rest of penultimate and

ultimate whorls is sculptured with axially or diagonally

directed elongate pustules (Figs 15 & 19-20);

operculum on inner side with a strong unicuspid hook-

shaped apophysis at the base (Figs 15E & 20E) ....... 11

- Axial ribs clearly defined; operculum (where known)

MOtaS MNS RO AOE ae hates uantas gilt crenluen Sea tals

pp. 135-184

11 Peristome darkened; peristome — recurved

abaperturally (Figs 19A-20A); operculum on inner side

with unicuspid triangular apophysis at the base (Fig.

JOE we iter eGericaiieht ict: Sete clas nee aae tra tke ce T. schneideri

- Peristome whitish; peristome not recurved

abaperturally (Fig. 15A); operculum on inner side with

unicuspid hook-shaped apophysis at the base (Fig.

ETE) ee) re dans eats eaatcs La aieesPls- eka ctnh auricnte haat cae a T. jodiae sp. nov.

12 Parietal margin of peristome attached to wall of

ultimate whorl (Figs 6, 10 & 22); parietal gap not present

OL TAS ISG ANCLY. FRAN NOW at ek tat, ee tat os ee cess wtrdeaate taste 13

- Parietal margin of peristome broadly detached from

wall of ultimate whorl (Figs 9, 21, 24-26); parietal gap

CUSENAC tar ite oat cantrnntnaretiines ties ee acanenanituaie oak eennticrsnanmsiaeations ils,

13 Axial ribbing pattern generally denser, more than 35

ribs on the ultimate whorl; operculum different (not known

in T. clathrata); aperture less wide and more vertically-

oriented (Figs 6 & 22): decollated shell generally shorter

and less slender (Figs 6 & 22); distribution: Bismarck

AICNIPCla SO" svcvesiscccrccoreeaiiitenccceecoete nae eerecteeits 14

- There are ~25-27 ribs on ultimate whorl; operculum

apophysis cow-horn-shaped, unicuspidate, not lamellate

(cf. van Benthem Jutting (1963, fig. 9b)); aperture

stronger expanded, comparatively wider (Fig. 10A); shell

generally slenderer (Fig. 10OA-C); distribution: Biak,

Cenderawasih Bay Islands ............::000 T. foliosocostata

14 Shell sculptured with chevron-shaped ridges as well

as axial ribbing (Fig. 6); ridges and ribs separated by

more than own width (Fig. 6); palatal margin of peristome

nearly straight in lateral view (Fig. 6C) ............008 T. bifurca

— Shell with axial ribbing but lacking raised ridges (Fig.

22); ribs denser, separated by about their own width;

palatal margin of aperture sinuous in lateral view (Fig.

T. tesselata

15 Some of axial ribs on teleoconch markedly increasing

in shape or height towards suture, or at least on ultimate

whorl axial ribs are grouping or merging together to form

short stronger ridges at suture and at shell base (Figs

PLD Wee S| OEE te EERE tom Re Ot hea Pe Ret Ae ee 16

- Axial ribs on teleoconch not increasing in shape or

height towards suture; axial ribs are not merging together

LOTORMe CNICKER MOSS: aes ss eraseiwutioo tera sscovecinencengenntetad de

16 Operculum with erect ridge along one margin,

with bicuspidate apophysis on opposite margin and

5-6 subparallel irregularly shaped partly strongly

high transverse lamellae close to the erect ridge (cf.

Gittenberger (1989, fig. 4)); on ultimate whorl some

axial ribs are grouping (merging together to form short

stronger blades) at suture (see Gittenberger (1989, figs

1-2); shell uniformly coloured, without darker markings

T. heinrichi

- Operculum with slightly erect ridge along one of

margins; apophysis unicuspidate, with five strong

lamellar elevations of variable size raising radially from

it (Fig. 26D & F); on ultimate whorl some axial ribs

are grouping (merging together to form short stronger

ridges) at suture (Fig. 26C); shell not uniformly coloured

but without darker brown marking ............000 T. wallacei

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—- Operculum without erect ridge; apophysis cow-horn-

shaped unicuspidate, not lamellate (Fig. 21E & G);

axial ribs not merging together to form stronger ridges

at suture or shell base, slightly increasing in shape at

suture (Fig. 21D); shell uniformly coloured, without

CGarker Drown Marking ......ccceceeeeeeeeeeeeees T. telnovi sp. nov.

- Operculum apophysis massive, somewhat laterally

curved, truncate apically (cf. Turner (1959, figs 2-5));

at least on ultimate whorl some axial ribs are grouping

(merging together to form short stronger ridges) at

suture and shell base; shell with irregular darker brown

markings on generally paler background T. orrae

17 Axial ribs on teleoconch dense, strongly sinuous (Figs

23-24); shell stronger obese (Figs 23-24) ..... T. ultima

- Axial ribs on teleoconch straight, very spaceous (Fig.

9); shell comparatively slenderer (Fig. 9) .... T. clathrata

Discussion

The two truncatellid genera in the Papuan

Region, Truncatella and Taheitia, differ markedly

in their distributions. The five Truncatella taxa: T.

granum, T. guerini, T. marginatum, T. thaanumi, and

T. thaanumi insularis, are saltwater tolerant Species

living in the extreme upper intertidal and supratidal

edges of saltwater habitats under mats of cast up

marine vegetation. This habitat likely facilitates

long-distance transport on floating mats during

storm events, and all are distributed beyond the

Papuan Region (Clench & Turner 1948). In contrast,

Taheitia are found in terrestrial habitats and our

study suggests are generally narrowly distributed.

The eight new species described here are all found

on single islands: Taheitia biaka sp. nov. from Biak,

T. bifurca sp. nov. from Manus, T. gebeensis sp.

nov. from Gebe, T. gigantea sp. nov. from Waigeo,

T. malagan sp. nov. from New Ireland, T. telnovi sp.

nov. from Misool, T. jodiae sp. nov. and T. longpela

sp. nov. from New Britain. Some reports of species

with multi island ranges appear to be based on

misidentifications. Taheitia wallacei records from

Misool and Salawati (van Benthem Jutting, 1958;

1963) are likely the subsequently named T. telnovi

sp.nov. and T. heinrichi Gittenberger, 1989. Records

of Taheitia gracilenta (E.A. Smith, 1897) from New

Britain (Rensch |. 1937) and Biak (Benthem Jutting

1963: 440) likely refer to morphologically similar

Taheitia longpela sp. nov. and T. biaka sp. nov.,

respectively. Other records of T. gracilenta from

New Ireland (Rensch I. 1937: 629 & 633-634)

and of Taheitia ultima (|. Rensch 1937) from Yapen

(Benthem Jutting 1963: 441) need confirmation.

Remaining species with multi island ranges are not

separated by deep water, for example, T. heinrichi

152

is found in Salawati, Warir, and adjacent lowlands

of New Guinea’s Doberai Peninsula, which were

likely contiguous during low water stands.

The Papuan Region is geologically complex

(Baldwin et al. 2012), and many areas are

experiencing rapid uplift of karst habitats (Lindley

2008), an extreme example of this is the Nakanai

Mountains of eastern New Britain, where rapid

uplift has resulted in extensive and very deep cave

systems (Gabriel et a/. 2018). Taheitia species

appear to be calciphiles and are not generally

distributed over wide areas, instead occurring in

discontinuous habitats where weathered limestone

is exposed at the soil surface. All eight new species

of Taheitia were found in rainforest on karst, usually

under or on limestone rocks within rock piles, or in

leaf litter among rocks. These new species increase

the Papuan Region’s Taheitia fauna by more than

50% bringing the total number of species to 21. It

is likely other unexplored karst areas in the Papuan

Region are inhabited by additional undiscovered

Species.

The habitat specificity of Taheitia species

makes them particularly vulnerable to both

limestone extraction and deforestation. Species

restricted to karst habitats are often found

where limestone is exposed at the soil surface,

unfortunately these are also the conditions that

are optimal for commercial extraction for cement

and other industrial applications. Entire limestone

exposures can be mined away before endemic

species are discovered (Oheimb et al. 2019;

Schilthuizen et al. 2005). Even in areas where

limestone is not extracted, forest loss can lead to

loss of soil moisture, which is likely to negatively

impact moisture dependent snails. The island of

New Guinea harbours the third largest tracts of

primary rainforest remaining on the planet, behind

only the Amazon and Congo basins. As remaining

uncut forests decline in Australasia, particularly in

Kalimantan and other parts of Indonesia, logging

has increased in primary forests of the provinces

of Papua and West Papua (Nasendi 2000). In

Papua New Guinea, rates of deforestation between

1972 and 2002 increased from 1.4 to 1.7% per

year, surpassing deforestation rates in the Amazon

Basin. In only 30 years, 15% of the country’s

tropical forests have been felled and another 8.8%

degraded (Shearman 2009). Terrestrial snails are

the faunal group most likely to be impacted by this

forest loss, as these moisture dependent animals

are among the most endangered species globally,

and approximately 50 percent of all extinctions

recorded since 1500 have been mollusks

02-Jun-21 21:48:26

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particularly land snails (Regnier et a/. 2015). This

high rate of extinction coupled with loss of habitat

due to deforestation emphasizes the importance

and urgency of increased sampling to identify and

describe species before their extinction (Richling &

Bouchet 2013) and while conservation efforts are

Still possible.

Acknowledgements

Our special thanks to Jonathan Ablett (BMNH),

Sankurie Pye (NMS), and Christine Zorn (MFNB)

for providing specimens and images from the

collections under their care. We are indebted to

Matthias Hartmann (NME) for material provided

for current study and for donating specimens for

authors collections. Roman Egorov (Moscow, Russia)

is thanked for images of the paratype Taheitia

mansueta and Dmitry Telnov (The Entomological

Society of Latvia, Riga) - for preparing numerous

images for the present paper and for critical review

of the manuscript.

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Received: 21.ix.2020.

Accepted: 15.xii.2020.

02-Jun-21 21:48:27

Greke, K. & Stapcinsky, J.: New Taheitia H. et A. Adams, 1863, with revisional notes on the Papuan Truncatellidae ...

pp. 135-184

Figures 1-2. Papuan Truncatella Risso, 1826 species. 1 - T. guerinii A. Villa et J. B. Villa, 1841; 2 - T. marginata

Kuster, 1855. A - Apertural view; B - Abapertural view; C - Lateral view [scale bars 5 mm].

155

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TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figures 3-4. Papuan Truncatellidae species. 3 - Truncatella thaanumi insularis Clench et Turner, 1948; 4 -

Taheitia porrecta (Gould, 1847), holotype (USNM 5527) from the National Museum of Natural History, Smithsonian

Institution, Washington, D.C., U.S.A. (image 3 licensed under http://creativecommons.org/publicdomain/zero/1.0,

out of copyright). A - Apertural view; B - Abapertural view; C - Lateral view; D - Apical view; E - Umbilical view [scale

bars 5 mm].

156 oo

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GrekeE, K. & Starcinsky, J.: New Taheitia H. et A. Adams, 1863, with revisional notes on the Papuan Truncatellidae ...

pp. 135-184

Figure 5. Taheitia biaka sp. nov., holotype. A - Apertural view; B - Abapertural view; C - Lateral view; D-F -

Operculum, inner (D) and outer surface (E), lateral view (F) [figures A-C scale bar 5 mm; figures D-F not to scale].

r=,

caaa

157,

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Figure 6. Taheitia bifurca sp. nov., holotype. A - Apertural view; B - Abapertural view; C - Lateral view; D - Axial

ribbing pattern, close view; E - Operculum, inner surface [figures A-C scale bar 5 mm; figures D-E not to scale].

158 emer

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GrekeE, K. & Starcinsky, J.: New Taheitia H. et A. Adams, 1863, with revisional notes on the Papuan Truncatellidae ...

pp. 135-184

Zoolog. Museum Berlin

Figure 7. Taheitia bismarckiana (|. Rensch, 1937), two syntypes (syntype 1 - A, C, D; syntype 2 - B, E, F). A & B-

Apertural view; D & E - Abapertural view; C & F - Lateral view; G - Original labels [scale bar 2 mm].

159

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figure 8. Taheitia calcarata van Benthem Jutting, 1963, specimen from ~56 km NNW Biak, Biak Island. A - Apertural

view; B - Abapertural view; C - Lateral view; D - Axial ribbing pattern, close view [figures A-C scale bar 5 mm].

160

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Greke, K. & Stapcinsky, J.: New Taheitia H. et A. Adams, 1863, with revisional notes on the Papuan Truncatellidae ...

pp. 135-184

Figure 9. Taheitia clathrata (H. Adams et Angas, 1865), three syntypes (syntype 1 - A, D-F, syntype 2 - B, syntype

3 - C). A-C - Apertural view; D - Abapertural view; E - Lateral view; F - Axial ribbing pattern, close view [figures A-E

scale bars 2 mm] (images courtesy Sankurie Pye).

161

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figure 10. Taheitia foliosocostata van Benthem Jutting, 1963, specimen from Urfu village, Biak Island. A - Apertural

view; B - Abapertural view; C - Lateral view; D - juvenile specimen, apertural view [figures A-C scale bar 5 mm].

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GrekeE, K. & Starpcinsky, J.: New Taheitia H. et A. Adams, 1863, with revisional notes on the Papuan Truncatellidae ...

pp. 135-184

Figure 11. Taheitia galactodes van Benthem Jutting, 1963, specimen from Urfu village, Biak Island. A - Apertural

view; B - Abapertural view; C - Lateral view; D - juvenile specimen, apertural view [figures A-C scale bar 5 mm].

163

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figure 12. Taheitia Sebeensis sp. nov., holotype. A - Apertural view; B - Abapertural view; C - Lateral view;

D-E - Axial ribbing pattern, close view [figures A-C scale bar 5 mm; figures D-E not to scale].

164 oe

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GrekeE, K. & Starcinsky, J.: New Taheitia H. et A. Adams, 1863, with revisional notes on the Papuan Truncatellidae ...

pp. 135-184

Figure 13. Taheitia gigantea sp. nov., holotype. A - Apertural view; B - Abapertural view; C - Lateral view

[scale bar 5 mm].

165

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ee. da 1 CO y eke vt te

ae f we ‘3

Cf Orie »

Yi dette ae

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bi Tecra, EUS 2 2, pltxS)|

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Figure 14. Taheitia gracilenta (E. A. Smith, 1897), lectotype. A - Apertural view; B - Lateral view; C - Abapertural

view; D - Apical view; E - Umbilical view; F-G - Original labels [figures A-E scale bar 5 mm] (images courtesy

Jonathan Ablett).

166 = \ i> |

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pp. 135-184

Figure 15. Taheitia jodiae sp. nov., holotype. A - Apertural view; B - Abapertural view; C - Lateral view; D - Axial

ribbing pattern, close view; E - Operculum, inner surface [figures A-C scale bar 5 mm; figures D-E not to scale].

MP a

167

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figure 16. Taheitia longpela sp. nov., holotype. A - Apertural view; B - Abapertural view; C - Lateral view; D -

Aperture, close view [figures A-C scale bar 5 mm; figure D not to scale].

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pp. 135-184

Figure 17. Taheitia malagan sp. nov., holotype. A - Apertural view; B - Abapertural view; C - Lateral view; D - Axial

ribbing pattern, close view; E - Operculum, inner surface [figures A-C scale bar 5 mm; figures D-E not to scale].

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Figure 18. Taheitia mansueta van Benthem Jutting, 1963, paratype (collection Zoological Museum Moscow State

University) from Pasirputih near Manokwari, Doberai Peninsula, New Guinea. A - Apertural view; B - Abapertural

view; C - Lateral view [scale bar 5 mm] (images courtesy Roman Egorov).

170 a es . > >

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pp. 135-184

Zoolog. Museum: Berlin

Figure 19. Taheitia schneideri (|. Rensch, 1937), paratype from Mailmail village, New Britain. A - Apertural view; B

— Lateral view; C - Apical view); D - Original labels [scale bar 5 mm].

AA:

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Figure 20. Taheitia schneideri (|. Rensch, 1937), specimen UF366522 from ~ 4 km NW of Marmar village, E New

Britain. A - Apertural view; B - Abapertural view; C - Lateral view; D - Axial ribbing pattern, close view; E - Operculum,

inner surface [figures A-C scale bar 5 mm; figures D-E not to scale].

Tha

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pp. 135-184

Figure 21. Taheitia telnovi sp. nov. A-G - Holotype. A - Apertural view; B - Abapertural view; C - Lateral view;

D - Axial ribbing pattern, close view; E-G - Operculum, inner (E) and outer surface (F), lateral view of the opercular

figures A-C scale bar 5 mm; figures D-H not to scale].

Fe ,

173

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TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Vaieetaee lee ad Lelalr Silly

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Grehetis Lake tay fs ae

Fd Kecte : Ge, a too a -R

ures arisen Pp PIERS Ma re ees: a

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Figure 22. Taheitia tesselata Mollendorff, 1897, specimen from Bismark Archipelego (B. Strubell collection, BMNH).

A - Apertural view; B - Lateral view; C - Abapertural view; D - Apical view; E - Umbilical view; F-G - Original labels

[scale bar 5 mm] (images courtesy Jonathan Ablett).

174 aG- Y

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GrekeE, K. & Siarcinsky, J.: New Taheitia H. et A. Adams, 1863, with revisional notes on the Papuan Truncatellidae ...

pp. 135-184

LAS AKeual E

Figure 23. Taheitia ultima (|. Rensch, 1937), paratype from Jacquinot Bay, New Britain. A - Apertural view; B -

Lateral view; C - Dorsal (top) view); D - Original label [scale bar 5 mm].

175

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figure 24. Taheitia ultima (|. Rensch, 1937), specimen UF366525 from E of Galowe village, E New Britain. A -

Apertural view; B - Abapertural view; C - Lateral view; D - Axial ribbing pattern, close view; E - Operculum, inner

surface [figures A-C scale bar 5 mm; figures D-E not to scale].

176

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GrekeE, K. & Stapcinsky, J.: New Taheitia H. et A. Adams, 1863, with revisional notes on the Papuan Truncatellidae ...

pp. 135-184

Figure 25. Taheitia wallacei (H. Adams, 1865), topotype (possible syntype) BMNH. A - Apertural view; B - Lateral

view; C - Abapertural view; D - Apical view; E - Umbilical view; F - Original labels [scale bar 5 mm] (images courtesy

Jonathan Ablett).

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Figure 26. Taheitia wallacei (H. Adams, 1865), specimen from 3 km W of Waisai, Waigeo. A - Apertural view;

B - Lateral view; C - Axial ribbing pattern, close view; D-F - Operculum, inner (D) and outer (E) surface,

lateral view (F) [figures A-B scale bar 5 mm; figures C-F not to scale].

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GrREekE, K. & Stapcinsky, J.: New Taheitia H. et A. Adams, 1863, with revisional notes on the Papuan Truncatellidae ...

pp. 135-184

Figure 27. Taheitia bifurca sp. nov., paratype, live specimen.

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TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figures 28-29.

Live specimen and

microhabitat of

Papuan Taheitia.

28 - Taheitia jodiae

sp. nov., paratype,

live specimen; 29A

& B - Microhabitat

of Taheitia telnovi

sp. nov., River Gam

valley.

180

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GrekE, K. & Siarcinsky, J.: New Taheitia H. et A. Adams, 1863, with revisional notes on the Papuan Truncatellidae ...

pp. 135-184

Figures 30-34. SEM micrographs of radulae of the Papuan Taheitia H. et A. Adams, 1863. 30 - T. bifurca sp. nov.,

paratype UF4/6354; 31 - T. jodiae sp. nov., paratype UF366600; 32 - T. schneideri (|. Rensch, 1937), specimen

UF366527 from 4 km NW of Marmar village, E New Britain; 33 - T. telnovi sp. nov., paratype UF542950; 34 - T.

ultima (Il. Rensch, 1937), specimen UF366523 from E of Galowe village, E New Britain [scale bars 0.05 mm].

circle) on Gebe, North Moluccas (prepared with ArcGIS 10.3).

> ee Fen |

=—> @Qd é& 181

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

x: a

J ctl

Map 2. Localities of Taheitia H. et A. Adams, 1863 on Biak, Cenderawasih Bay Islands. Four-pointed stars (filled

and empty) - T. calcarata van Benthem Jutting, 1963; Circle - T. biaka sp. nov.; Diamonds (filled and empty) and

an empty four-pointed star - T. foliosocostata van Benthem Jutting, 1963; Empty four-pointed star - T. orrae Turner,

1959; Empty diamonds - T. galactodes van Benthem Jutting, 1963 (prepared with ArcGIS 10.3).

Map 3. Localities of Taheitia H. et A. Adams, 1863 on Bismarck Archipelago. Four-pointed star - T. malagan sp. nov.;

circle - T. jodiae and T. longpela spp. nov. (prepared with ArcGIS 10.3).

2 =< ©

02-Jun-21 21:48:56

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GrREkE, K. & Starcinsky, J.: New Taheitia H. et A. Adams, 1863, with revisional notes on the Papuan Truncatellidae ...

pp. 135-184

Map 4. Localities of Taheitia bifurca sp. nov. (circles) on Manus, Admiralty Islands (prepared with ArcGIS 10.3).

4 a

pepe aif

ee I,

Map 5. Localities of Taheitia H. et A. Adams, 1863 on Misool, Raja Ampat Islands. Filled eirciee Type localities of T.

telnovi sp. nov.; empty circles - Unverified localities of T. telnovi sp. nov. published as those of T. wallacei (H. Adams,

1865) by van Benthem aes (1958) (prepared with ArcGIS 10.3).

183

Book4.indd 183 02-Jun-21 21:48:59

TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

~\ =>

= rd

dir

z fi

, a asi

Map 6. Localities of Taheitia H. et A. Adams, 1863 on Waigeo Island. Circles - Localities of T. wallacei (H. Adams,

1865); Four-pointed star - T. gigantea sp. nov. (prepared with ArcGIS 10.3).

A ae eee am 4

Map 7. Localities of Taheitia H. et A. Adams, 1863 on Salawati and Warir, Raja Ampat Islands, and on Doberai

Peninsula of New Guinea. Circles - Type localities of T. heinrichi Gittenberger, 1989; Squares - T. gracilenta (E. A.

Smith, 1897); Four-pointed star - T. mansueta van Benthem Jutting, 1963 (prepared with ArcGIS 10.3).

184 seme

Book4.indd 184 02-Jun-21 21:49:02

Greke, K., TELNOv, D., KAGAINIS, U. & TELNov, E.: The possible functional role of periostracal processes in Japonia saetigera ...

pp. 185-198

The possible functional role of periostracal processes

in Japonia saetigera (van Benthem Jutting, 1958)

(Caenogastropoda: Cyclophoridae) based on SEM

micrograph analysis and numerical calculations

urn:lsid:zoobank.org:pub:ED2 /O4BD-BFOC-4D /6-9B4B-E6FF53BB23A9

KRistiINE GREKE *, Duitry TeLNov~:'**, Uais Kacainis *'° & Epwin TELNov °

1 - Corresponding author: Latvian National Museum of Natural History, K. Barona iela 4, LV-

1050, Riga, Latvia; k.greke@gmail.com

2 — Natural History Museum, Department of Life Sciences, Cromwell Road, SW7 5BD, London,

United Kingdom; anthicus@gmail.com (ORCID: O0O00-0003-3412-0089)

3 - Institute of Life Sciences and Technology, Daugavpils University, Vienibas iela 13, LV-5401,

Daugavpils, Latvia

4 — Institute of Biology, University of Latvia, O. VacieSa iela 4, LV-1004, Riga, Latvia;

ugis.kagainis@lu.lv (ORCID: OO00-0002-9921-2331)

5 - Faculty of Biology, University of Latvia, Jelgavas iela 1, LV-1004, Riga, Latvia

6 - Darza iela 10, Stopinu novads, LV-2130, Dzidrinas, Latvia

Abstract: A possible explanation for the functional importance of shell appendages in Japonia saetigera (van

Benthem Jutting, 1958), an endemic New Guinean terrestrial cyclophorid gastropod, is given based on a morphology

and calculation-based model and scanning electron microscopy. Results demonstrate that the periostracal processes

increase the perimeter length of the ultimate whorl by nearly 42%, the net dorsal shell surface area - by about 20%,

and the shell volume by only around 1% in the studied specimen. Possible adaptive importance of periostracal

processes in J. saetigera is hypothesised and discussed. New distributional record for J. saetigera is presented.

Book4.indd 185

Key words: Shell morphology, adaptations, adhesive properties, floating, air bubbles, ecology, Papuan Region.

Introduction

In molluscs, the calcium carbonate (calcareous)

Shell is covered externally by a thin proteinic

layer composed of conchiolin, the periostracum,

which is in part mineralized and, thus, involved

in shell formation (de Paula & Silveira 2009). The

periostracum of some pulmonate species is of

taxonomic interest Since microscopical structures

often derive from it. Periostracal setae or hairs are

the most common and extensively studied type of

such structures, but Some species develop extreme

clava-, Sooon- or feather-like periostracal structures

(cf. Figs 1, 25-26). Allgaier (2011) described

such morphology and Suvorov (1999) and

Pfenninger et al. (2005) discussed the functional

role and evolutionary importance of periostracal

setae in selected terrestrial European Helicidae,

Helicodontidae and Hygromiidae taxa. Both studies

found that the possession of periostracal setae

| (EE ss

facilitates the adherence of the shells to wet

surfaces and highlighted correlation of setation

loss with a shift from humid to dry habitats in

terrestrial molluscs.

Japonia (Mylicotrochus) saetigera (van

Benthem Jutting, 1958) (Figs 1-3) is terrestrial

cyclophorid (Cyclophoridae Gray, 1847) species

endemic to Misool Island (Raja Ampat Islands) off

the coast of western New Guinea. Prior to this study

this species was known only from the type series

collected in the northern part of Misool and is

peculiar among hitherto known Papuan terrestrial

molluscs due to specific spoon-like “bristles”

which are regularly, moderately widely spaced

along the flat peripheral carina of the teleoconch

(van Benthem Jutting 1958). The morphology and

function of these “bristles” (referred to further in

the text as periostracal processes) have not been

previously studied in detail. Acquiring additional

specimens of this species made our studies

J > af

ti.

(ee ee

>. ea

185

02-Jun-21 21:49:02

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

on the morphology of periostracal processes in

Japonia saetigera possible, the results of which are

presented and discussed further.

The primary aim of the present paper is to

initiate a discussion on the possible adaptive role

of non-hair-like periostracal processes in terrestrial

molluscs living under humid conditions, as well as

an attempt to estimate contribution of periostracal

processes to the net surface area and volume of

the studied molluscan shell. The present study

does not pretend to be comprehensive and is to be

considered as an initial part of a larger scale study.

Material and methods

The first author identified the specimens used

for the present research. Studied specimens were

fixed in 99% ethanol but all were empty shells with

no body. The holotype is also only a dry shell and is

stored dry. Only one of the four studied specimens

(inclusive of the holotype) has most of the

periostracal processes present, therefore we based

ourresearch and calculations on this best-preserved

specimen only. A Leica S6D stereomicroscope was

used for species identification. For micrograph

imaging a Scanning electron microscope (SEM) was

used facilitated by the Faculty of Biology, University

of Latvia. To prepare specimens for imaging, shells

were ultrasonically cleaned in distilled water for 1-2

minutes, washed through a graded alcohol series

up to 99%, air-dried and then placed dorsally onto

conductive tape on aluminium stubs. Selected

samples (Figs 12-22) were coated with 20 nm Au

in an ion coater Eiko IB-3 before observations were

made. All shells were imaged using a Hitachi TM-

3000 (Hitachi High Technologies ®) SEM following

methods as described by Geiger et a/. (2007).

Data from labels are reproduced verbatim,

without additions. Labels (if there are multiple

labels on a specimen) are separated by a slash.

Authors' comments are placed in square brackets.

The Papuan Region as considered here is

a zoogeographical term in the sense of Gressitt

(1982), Beehler et a/. (1986), Riedel (2002), and

Telnov (2011).

The studied material is stored at the following

collections:

KGC - Collection Kristine Greke, Riga, Latvia;

NME - Naturkundemuseum Erfurt, Germany;

NMNL - Naturalis Biodiversity Center, Leiden,

Netherlands.

the

186

Measurements and calculations

Measurements of periostracal projections

taken from SEM micrographs. Measurements of

Shell taken with a stereomicroscope (see above)

using a reticule.

The model

The surface area and volume of the shell (not

including the periostracal processes) is modelled

separately from the surface area and volume

of the periostracal projections on it, so that their

contribution can be compared to the overall value.

As such, the periostracal processes of the shell

and the shell itself are treated separately, and

both can be modelled as solids of revolution (for

description of this model see Harris & Stocker

(1998) and Weisstein (2020a)). The perimiter

length of a single periostracal process was found

by calculating the arc length of the generatrix

function of the said surface of revolution (Weisstein

2020b) and multiplying it by 2 since the generatrix

is rotated 360° and is thus mirrored along the axis

of symmetry. The areas were modelled as surfaces

of revolution (for description of this model see

Weisstein (2020c)). These models are based on

several assumptions, which were observed to hold

true for both the periostracal projections and the

shell (together and separately called the object) on

magnification of up to 200x:

The object is radially symmetrical along its axis

of symmetry;

The surface of the object is smooth and

continuous on every freely selected segment;

The surface of the object has a negligible

thickness.

Since these conditions hold only for a certain

scale (for instance, surface of shell whorls is rarely

entirely smooth under magnification since axial

ribbing patterns are present and _ periostracal

projections are not entirely smooth under high

magnification) and are not “truly” satisfied, the

model is expected to produce minor errors,

dependent on the extent to which these conditions

are not met.

The formulae used

The formulae for double arc length, the surface

area and volume of a solid of revolution are:

02-Jun-21 21:49:03

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Greke, K., TELNOv, D., KAGAINIS, U. & TELNov, E.: The possible functional role of periostracal processes in Japonia saetigera ...

Vih)=0 / r?(h)dh

respectively, where r is the generatrix of the solid

of revolution, h is the distance along the axis of

symmetry (or, for case of mollusk shell, its dorso-

ventral axis) and (a, 6) are the boundaries of the

freely selected region.

The uncertainties calculated in this work

are results of propagating potential errors

in measurements, and do not account for

possible deviations from the model, uncertainty

in regression, rounding errors or variations in

statistical distributions.

The following software was used for the

calculations: OriginPro data analysis tool v. 9.8,

Wolfram Alpha (httos:/7/www.wolframalpha.com).

Results

Taxonomy and morphology

Japonia (Mylicotrochus) saetigera (van Benthem

Jutting, 1958) (Figs 1-24)

Holotype NMNL: Holotype en Paratypen [handwritten]

Zodlogisch Museum. Amsterdam. [printed] Lagochilus

(Mylicotrochus) saetigerum v.B.Jutting Fakal, 0-75 m

hoog Centraal Misool 7 Oct.1948 leg. M.A. Lieftinck

[handwritten, the Latin name is underlined] / Holotype

[printed] / holotype 3.58.00 1 dr ZOOLOGISCH MUSEUM

AMSTERDAM ZMA Moll. 135638 Cyclophoridae 0767.0

Lagochilus saetigerum Van Benthem Jutting, 1958

INDONESIA Papua Misodl, Fakal, 0-75 m 1948 10 07

sta. Leg.M.A. Lieftinck Ex coll. Original publication: Van

Benthem Jutting, W.S.S., 1958 Non-marine Mollusca of

the island of Misool. Nove Guinea, new ser., 9(2): 293-

338 Autoref: Van Benthem Jutting, W.S.S., 1958e: 304-

306, fig. 3 Add.publ. Notes: Current Scientific Name:

Lagochilus saetigerum Van Benthem Jutting, 1958 Det.

/ Naturalis Biodiversity Center 40380 - Cyclophoridae

Lagochilus saetigerum van Benthem Jutting, 1958 -

HOLOTYPE Lieftinck, M.A., c. Indonesia, Papua, Misool,

Pulau, Fakal; depth [sic! altitude] 0-75 m 7.x.1948

[printed].

Additional studied material, 3 specimens NME

& KGC: INDONESIA E, Prov. Raja Ampat, distr. Misool

Barat, Lilinta (Lelintah) vill. ~13.5 km NW, Gamta vill.

pp. 185-198

~11 km NWW, River Gam valley in the middle of course,

01°57’46"S, 130° 10’54’E, 03.IV.2009, primary lowland

rainforest, limestone creek [of the three specimens, one

bears complete periostracal projections and two other

lack them or only have fragmentary processes present].

Note: Assuming that “saetigerum” (as originally of

van Benthem Jutting (1958)) is a Latinized Greek

adjective and not a noun, we consider it appropriate

to change the ending of the specific epithet to

“saetigera” when combined with a feminine generic

name Japonia.

Taxonomy: This taxon was originally described

as Lagochilus saetigerum in the subgenus

Mylicotrochus (van Benthem Jutting 1958).

Lagochilus Fischer, 1885 and of later authors is an

unjustified emendation of Lagocheilus Theobald in

Blanford, 1864 and ajunior homonym of Lagochilus

Loew, 1860 (Diptera), now a subgenus of Japonia

A. Gould, 1859 (Egorov 2009). Mylicotrochus P.

Sarasin et F. Sarasin, 1899 was originally erected

as a subgenus of Lagochilus for a single Sulawesi

species Japonia (M.) celebensis on the basis of

peculiarities of the radula (Sarasin & Sarasin

1899). However, the radula of Japonia saetigera

was not described by van Benthem Jutting

(1958) and remains unknown since no preserved

anatomical material is available. Therefore, the

original placement of Japonia saetigera in the

Subgenus Mylicotrochus by van Benthem Jutting

(1958) was not fully justified. However, we do not

intend to change the initial subgeneric placement

at this point.

Morphology of periostracal processes:

In Japonia saetigera, periostracal processes are

absent on the protoconch but are present on

older parts of the teleoconch (Figs 4-5) which

follow the microscopically pitted protoconch (Fig.

5). On the teleoconch, every fifth or sixth radial

growth line of the periostracum is accentuated

with a lamellar periostracal extension (Figs 1, 4,

6), which is stronger-built than the surrounding,

less prominent, radial periostracal growth lines.

These stronger, lamellar growth lines each end in

a conspicuous periostracal process (Figs 1 & 4).

On the first 1-1%% teleoconch whorls, periostracal

processes are shorter and clavate (e.g., like a

finger or bowling pin) (Figs 4, 7-11), with a suture

on ventral side (Figs 7, 10-11). On later whorls

the clavae “open” along this suture and develop

a conspicuous spoon-like shape, with the convex

side facing upwards (Figs 1, 4, 12, 18). The

external surface of each process (both clavate and

spoon-shaped) has gentle irregular, transverse

187

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concentric ridges (growth lines?) and somewhat

lower, irregular, longitudinal interconnecting ridges

between the concentric ones (Figs 7-8, 10, 12-16).

Concentric ridges represented by different layers of

periostracum partially overlap each other with the

posterior margin of each layer (Figs 14-17). Clavate

processes on the first teleoconch whorl are empty

inside (filled with an air bubble). The underside

of the spoon-shaped process (in fact - the inner

side of the clavate process of the earlier whorls) is

nearly smooth and glabrous, with a few irregularly

spaced, gentle, circular to slightly ovoid pores (Figs

20 & 22) of approximate 0.5 to 1,5 um in diameter

(measured from one process, not statistically

proven). Spoon-shaped periostracal processes of

the last teleoconch whorls (the pen- and ultimate

whorls of the shell) are about 3-4x as large as the

smaller clavate processes of earlier whorls (Table

1). Both clavate and spoon-shaped processes

are strongly attached to periostracum and hard

to remove. In subfossil calcareous J. saetigera

Shells without periostracum there are no traces of

peripheral processes or their points of articulation.

Shells from living J. saetigera specimens do not

become dirty from its microhabitat (a mixture of

soil and litter). Length of periostracal processes:

small (50-100 wm), mid-sized (100-300 um),

large (400-700 um) (measured from one shell

specimen only, not statistically proven).

Ecology: Occurs in wet leaf litter in primary lowland

rainforest on limestone.

Distribution: Endemic to Misool Island, Raja

Ampat Islands. The record provided in this paper

is the first record of J. saetigera since the original

description and the first locality in southern Misool.

Calculations for the periostracal processes

For the periostracal processes of the shell,

the generatrix ras a function of distance from the

origin along the axis of symmetry h was proven

to be too complex to be described by a single

continuous function, and as such was. found

numerically by dividing it into several segments.

The functions describing these segments, as well

as their corresponding domain and R? value are

listed below:

r(h) = -5- 10h? + 0.0022h? - 0.0706h + 34.547

h € (0;200) R?= 0.9980

r,(h) = 0.2928h + 14.689

h € (200;459) R?= 0.9991

188

r,(h) = —1.582 - 10°*h® — 4.040808 - 10*h? +

0.6317584237h — 40.5058825844

h € (459;725) R? = 0.9839

r(h) = -9.0701134413047 - 10-°h* +

0.207883829564922h? — 158.884236111089h

+ 40632.8636822398

h € (725;868) R? = 0.9964

The significant figures given above are

important, as since the value of h grows to

increasingly larger values in the latter equations,

the precision to which the said equations are

defined must also be increased. Thus, the general

function of the generatrix r(h) and its R? value are:

ri(h) -h € (0; 200)

ay = 4 tah) he € (200; 459)

TMT) rs(h) bh € (459; 725) (4)

ra(h) —-h € (725; 868)

R? = 0.9937

Note that this is a general function describing

any periostracal process, and that the domain of h

is O to 868. This means that any such periostracal

process Is defined to be 868 units in length exactly.

Inserting the above equation (4) into the formula

for double arc length (1) returns the result for

the perimiter length of periostracal process P

in terms of u, which is the unit length along the

axis of symmetry (i.e. the length of one A unit, or

from equation (4): 4/s6s" of the length of the

periostracal process).

PDP St 8895 71074 (5)

Doing the same for surface area and volume

with equations (2), (3) results in net surface area

and volume of the periostracal process, S and V

respectively:

S=YS = 6.4406 - 10° uv?

V= DV. = 4.8695 -107 U8

(6)

(7)

Since not all of the periostracal processes are

geometrically equal, as the majority of them are

open on the underside (not clavate-like closed)

and possess a “gap” in the otherwise perfect radial

symmetry (for instance, Fig. 18), the model must be

corrected. For perimiter length it was numerically

determined that the quality had insignificant effect

on the value. For surface area it can be accounted

for by removing some angle 8, which corresponds

to the missing circular sector. This can be done by

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Greke, K., TELNOv, D., KAGAINIS, U. & TELNov, E.: The possible functional role of periostracal processes in Japonia saetigera ...

instead of multiplying by 2m in equation (2) using

a factor of 2m — 8, which can be brought out of the

summation sign and inserted instead in equation

(6). The volume of the periostracal process however

stays mostly the same in the case of a gap since they

are comparatively small and the value of interest is

that of the extended circular segment rather than

the sector. Therefore correction for the volume of

the periostracal process was only performed for

cases where the value of 8 was comparable to rt

radians, and ignored otherwise.

Since the length of the entire periostracal

process Is defined to be 868 u (see equation 4), itis

easy to equate this value to one actually measured,

and hence get the value of u for the given case.

Through inspection, these periostracal processes

were counted and categorized according to their

missing angle 8, length /, and amount present, N

(Table 1).

Using this information, the values of u, u? and

u> were calculated and inserted into equations

(6) and (7), which were then multiplied by a factor

pp. 185-198

which accounted for the value of 8 and the amount

of periostracal processes N. The values were then

Summed up for the three categories to produce

the net surface area S and volume V from the

contribution of the periostracal processes:

S = 6.0298 + 1.2729 mm? (8)

= 0.2010 + 0.0420 mm? (9)

process

For the case of contour length however, the only

periostracal processes of interest are those that

sit on the ultimate whorl, as that is the presumed

area of contact with water. Through inspection it

was noted that in this soecimen the ultimate whorl

contained 20 processes of "large" class. As such the

net contour length contributed by the periostracal

processes was obtained by multiplying equation (5)

by a factor that accounted for the corresponding

u-value and the 20 present processes, and was

found to be

P = 3.2653 + 0.2177 mm

process

(10)

Table 1. Size classes of periostracal processes in studied Japonia saetigera specimen.

Size class of periostracal processes

Legends: | - length; N - quantity.

Small (present on earlier whorls) 1/6 +7/6 | 0.050-0.100 +0.005 0.075 +0.005

Midsize (on intermediate whorls) 1/3 + 11/6 | 0.100-0.300 +0.005 | 0.200 +0.005

Large (on pen- and ultimate whorls) 0.400-0.700 +0.005 | 0.550 +0.005

Calculations for the smooth shell and results

The surface area and volume of the smooth

Shell without the periostracal processes were found

analogously to those of periostracal processes.

First, the length of periphery of shell's spire L was

defined to be equal to 18068 u and measured to

be 24.3003 + 2.51638 mm (resulting in a single

u-value of 0.00134491 + 0.00014987 mm). The

relationship between the distance along the outer

periphery of the shell (treating the shell apex as

origin) L and the radius at that point R, and its R?

value were then numerically determined to be:

R(L) = 1024.432514 - 967.7655137¢'0.00004819931 1)

R? = 0.9975

Treating the volume of the spire and the shell

surrounding its area as a Solid of revolution that

has been "rolled-up" into a spiral permits the usage

of equations (2) and (3) to again find the surface

area and volume of the smooth shell, which were

calculated to be equal to

S = 4.3101 - 10’u? (11)

and

V = 9.6035 - 10%u? (12)

respectively. Very much like the case with the

periostracal processes, not all the surface should

be considered, as due to the solid being “rolled-

up” (Fig. 4), some of the surface is shared on the

intersections between whorls, and due to the main

interest pertaining to the visible surface of the shell

(in view from above, as shown in Fig. 1); the area of

the umbilical channel should not be counted either.

Thus, through inspection (for instance, Fig. 2) it is

estimated that the dorsal surface area of the shell

is 28 +3 % of the net surface area. Thus, multiplying

equations (11) and (12) by this percentage and the

calculated u-value results in dorsal surface area

and volume of the smooth shell and were shown

to be

S,,, = 21.8296 + 5.2125mm? (13)

V,, = 23.3634 + 7.8102 mm (14)

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For the case of perimiter however only the

length of the ultimate whorl is of interest, as

mentioned before. It was separately measured and

found to be equal to

P_ = 4.5337 + 0,4695 mm

shell

(15)

By summing the perimiter length, surface area

and volume of the smooth shell (15), (13), (14)

with the perimiter length, surface area and volume

of the shell periostracal processes (10), (8), (9),

the net perimiter length, dorsal surface area and

volume can be calculated:

see 7.7990 + 0.5175 mm

Ss 27.8594 + 0.3187 mm?

VA 23.5644 + 0.3941 mm?

The calculation results (cae mane Sse wa

Sand Votes /V.-) lllustrate that the periostracal

processes of the studied Japonia saetigera shell

contribute to 41.87 + 9.41% of the ultimate whorl

perimiter length, 21.64 + 4.58% of the net dorsal

surface area of the shell and 0.85 + 0.18 % to the

net volume of the shell.

Discussion

As it was demonstrated by Pfenninger et al.

(2005), the presence of periostracal hairs increases

adherence of terrestrial gastropod shells to wet

substrates. Moreover, haired shells are proved to

be an ancestral character state (Pfenninger et al.

2005). Suvorov (1999) pointed out that the only

cause of the phenomenon of clean shells of litter-

dwelling gastropods is due to the hydrophobic

properties of their microsetose periostracum.

The terrain in Misool is complex at the

study area (River Gam valley), with several steep

subparallel limestone anticlines and deep ravines

in-between. Japonia saetigera inhabits the wet floor

of lowland rainforests in Misool, this species was

found at low elevations at the base of limestone

anticlines but not reported from the anticlines

itself. Soecimens are known to dwell in leaf litter. At

the study area in the River Gam valley, it rains 1-3

times a day in February, March and April (Second

author, personal observation), but the soil remains

constantly wet to very wet (Fig. 23) throughout

the year with an annual rainfall about 2500 mm

(Prentice & Hope 2007). Frequent showers ensure

a continuous water flow from the anticlines down to

the creeks and larger rivers. Thus, rainforest floor

190

gastropods are under a continuous risk of being

flushed or sunk (Fig. 23).

We here hypothesise that the presence of

periostracal processes in Japonia saetigera shell

1) is an adaptation to increase adherence of

the shell to wet substrates (e.g. fallen leaves) due

to shell area increment (Fig. 24);

2) may act as a floating adaptation in wet

conditions: air bubbles can be "trapped" by spoon-

Shaped processes and can theoretically ensure

floating (keeping the shell on the waters’ surface);

3) the processes may serve to increase the

perimeter of the shell allowing for more force to be

applied before surface tension is broken. Under

standard conditions, the ~42% increase in length

translates to ~0.02 grams of additional mass that

can be supported afloat.

The mechanism by which the increase in

adherence presumably works is ensured by the

water adhesion. The mechanism of adherence

of air bubbles to a solid surface of a periostracal

process is ensured by the capillary bridge-force

(Hotta et al. 1974; Derjaguin et a/. 1987; Myshkis

et al. 1987). In its microhabitat (which is in fact a

three-dimensional environment consisting of litter,

soil, logs and fallen twigs), the conical shell of

Japonia saetigera shell is nearly always in contact

with two solid surfaces that are constantly wet - the

first is the surface of the substrate to which snail’s

foot is attached (usually a fallen leaf), the second

is another fallen leaf in the leaf litter, contacting

the shell from above or from the sides, meaning

that the snail is constantly covered from above

and below (Fig. 24). Under these circumstances,

the periostracal processes will adhere to both

solid surfaces thanks to the conical shape of the

Shell. Indeed, these connections will be ensured by

dorsal and ventral sides of processes depending on

which side stays in a contact with a solid surface.

Further experiments on living specimens involving

appropriate equipment is planned to test the

hypothesis.

By increasing the net dorsal surface area of

the shell by over 20%, the periostracal processes

can be considered to provide a significant increase

in shell surface area (in terms of both hypotheses -

to increase shell adherene or ensure shell floating).

Contrary, contribution by the periostracal processes

to the net volume of the shell (increase for <1%)

considered not significant.

The periostracal processes, as discussed in

the present paper for Japonia saetigera, appear in

02-Jun-21 21:49:03

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Greke, K., TELNOv, D., KAGAINIS, U. & TELNov, E.: The possible functional role of periostracal processes in Japonia saetigera ...

Similar or even more dramatic forms in some other

terrestrial tropical cyclophorid molluscs from the

Papuan Region, like in the enigmatic “Mychopoma’”

pennatum van Benthem Jutting, 1958 (Figs 25-

26), which is endemic to Misool Island and shares

its the habitat with J. saetigera and is only known

from the holotype stored in NMNL.

Acknowledgements

We are indebted to Bram van der Bijl (NMNL)

and Matthias Hartmann (NME) for the type and

recent material provided for the present study. We

also thank to Valters Gobins (Faculty of Biology,

University of Latvia, Riga) for providing a technical

Support during the SEM procedures. Jonathan

Ablett (Natural History Museum, London, United

Kingdom) is thanked for English proof.

Results of this study were presented at the 79"

International Scientific Conference of the University

of Latvia (Riga, 28.01.2021).

References

Allgaier C. 2011. A hairy business - periostracal

hair formation in two species of helicoid snails

(Gastropoda, Stylommatophora, Helicoidea). -

Journal of Morphology 272: 1131-1143.

Beehler B. M., Pratt T. K., Zimmerman D. A. 1986. Birds

of New Guinea. Handbook No. 9 of the Wau Ecology

Institute. New Jersey, Princeton University Press: xiii

+ 293 pp.

Derjaguin B. V., Churaev N. V., Muller V. M. 1987. Surface

Forces. Consultants Bureau, New York & London:

440 pp.

De Paula S. M., Silveira M. 2009. Studies on molluscan

Shells: Contributions from microscopic and

analytical methods. - Micron 40: 669-690.

Egorov R. 2009. A review of the genera of the recent

terrestrial pectinibranch molluscs (Synopsis mainly

based on published data). Part II. Littoriniformes.

Hainesiidae, Aciculidae, Cyclophoridae,

Craspedopomatidae. Treasure of Russian Shells.

Supplement 3. Colus-Doverie Print, Moscow: 58 pp.

Geiger D. L., Marshall B. A., Ponder W. F., Sasaki T.,

Warén A. 2007. Techniques for collecting, handling,

preparing, storing and examining small molluscan

specimens. - Molluscan Research 27, No 1: 1-50.

Gressitt J. L. 1982. General introduction: 3-13. In:

Gressitt J.L. (ed.) Monographiae biologicae 42,

Biogeography and ecology of New Guinea. Dr. W.

Junk / Springer publishers, the Hague: 983 pp.

Harris J. W., Stocker H. 1998. 4.10. Solids of Rotation:

(eds)

111-113. In: Harris J. W., Stocker H.

pp. 185-198

Science. Springer, New York: xxviii + 1028 pp.

Hotta K., Takeda K., linoya K. 1974. The capillary binding

force of a liquid bridge. - Powder Technology 10,

No 4/5: 231-242.

Myshkis A. D., Babskii V. G., Kopachevskii N. D.,

Slobozhanin L. A., Tyuptsov A. D. 1987. Low-Gravity

Fluid Mechanics: Mathematical Theory of Capillary

phenomena. Springer, Berlin & Heidelberg: xx +

584 pp.

Pfenninger M., Hrabakova M., Steinke D., DéprazA. 2005.

Why do snails have hairs? A Bayesian inference of

character evolution. - BMC Evolutionary Biology 5:

1-11.

Prentice M. L., Hope G. S. 2007. 2.3. Climate of Papua:

177-195. In: Marshall A. J., Beehler B. M. (eds)

The Ecology of Papua. Part one. The Ecology of

Indonesia Series, Volume VI. Periplus Editions,

Singapore: 749 pp.

Riedel A. 2002. Taxonomy, phylogeny, and zoogeography

of the weevil genus Euops (Insecta: Coleoptera:

Curculionoidea) in the Papuan Region. Dissertation

zur Erlangung des Doktorgrades der Fakultat

fur Biologie der Ludwig-Maximilians-Universitat

Munchen: 216 pp.

Sarasin P., Sarasin F. 1899. Materialien zur

Naturgeschichte der Insel Celebes. Band 2. Die

Land-Mollusken von Celebes. Kreidel’s Publishing

House, Wiesbaden: vili + 248 pp, 31 pls.

Suvorov A. N. 1999. Some mechanisms of adaptation to

the wet microhabitats in higher Geophila (Mollusca,

Pulmonata). — KypHaa o6Otueu Ouoaoruu 6O, No 2:

177-188.

Telnov D. 2011. Taxonomische Revision der Gattung

Macratria Newman, 1838 (Coleoptera: Anthicidae:

Macratriinae) aus Wallacea, Neuguinea und

den Salomonen: 97-285, pls 17-37. In: Telnov

D. (ed.) Biodiversity, Biogeography and Nature

Conservation in Wallacea and New Guinea. Volume

I. The Entomological Society of Latvia, Riga: 434 pp

+ 92 pls.

van Benthem Jutting W. S. S. 1958. Non-marine Mollusca

of the island of Misool. - Nova Guinea: A Journal

of Botany, Zoology, Anthropology, Ethnography,

Geology and Palaeontology of the Papuan Region

9, No 1: 293-338.

Weisstein E. W. 2020a. Solid of Revolution. MathWorld

- A Wolfram Web Resource. https://mathworld.

wolfram.com/SolidofRevolution.html [accessed 30

December 2020].

Weisstein E. W. 2020b. Arc Length. MathWorld-A Wolfram

Web Resource. https://mathworld.wolfram.com/

ArcLength.html [accessed 02.iv.2021].

Weisstein E. W. 2020c. Surface of Revolution. MathWorld

- A Wolfram Web Resource. https://mathworld.

wolfram.com/SurfaceofRevolution.html [accessed

30 December 2020].

Received: 09.xi.2020.

Accepted: 06.i.2021.

@e € 1

02-Jun-21 21:49:03

TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figures 1-3. Japonia saetigera (van Benthem Jutting, 1958). 1 - Studied and measured specimen from River Gam

valley, Misool; 2-3 - Holotype (NMNL), periostracal processes absent (fallen), apertural (2) and umbilical (3) view.

2 , eal 7

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Greke, K., TELNOv, D., KAGAINIS, U. & TELNov, E.: The possible functional role of periostracal processes in Japonia saetigera ...

pp. 185-198

5 500 um

Figures 4-5. SEM micrographs of Japonia saetigera (van Benthem Jutting, 1958) specimen from River Gam valley,

Misool. 4 - Apical view; 5 - Embryonic whorls (consider absence of periostracal processes).

Gy & 193

Book4.indd 193 02-Jun-21 21:49:06

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

50 um

Figures 6-11. SEM micrographs of Japonia saetigera (van Benthem Jutting, 1958) specimen from River Gam valley,

Misool. 6 - Lamellar periostracal extensions; 7 - Clavate (closed, empty inside) periostracal process, ventral view

(consider presence of a median "suture"); 8 - Clavate periostracal process, dorsal view; 9 - Arising point of clavate

periostracal processes on shell surface; 10 - Clavate periostracal process, ventral view (consider presence of a

median "suture"); 11 - Basal (tubular) part of clavate periostracal process, ventral view (consider presence of a

median "suture").

194

Book4.indd 194 02-Jun-21 21:49:09

Greke, K., TELNOv, D., KAGAINIS, U. & TELNov, E.: The possible functional role of periostracal processes in Japonia saetigera ...

pp. 185-198

16 = ; | 100 um

17 50 um

Figures 12-17. SEM micrographs of Japonia saetigera (van Benthem Jutting, 1958) specimen from River Gam

valley, Misool. 12 - Amputated "open" periostracal process, dorsal view; 13 - Basal part of amputated "open"

periostracal process; 14 - Median part of amputated "open" periostracal process, dorsal view;

15-17 - ditto; 17 - lateral margin of amputated "open" periostracal process, dorsal view.

195

Book4.indd 195 02-Jun-21 21:49:10

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

50 um

Figures 18-22. SEM micrographs of Japonia saetigera (van Benthem Jutting, 1958) specimen from River Gam

valley, Misool. 18 - Amputated "open" periostracal process, ventral view; 19 - Basal part of amputated "open"

periostracal process; 20-22 - Lateral margins (20-21) of amputated "open" periostracal process, ventral view;

22 - ditto but with visible median (nearly smooth) area.

196

Book4.indd 196 02-Jun-21 21:49:11

Greke, K., TELNOv, D., KAGAINIS, U. & TELNov, E.: The possible functional role of periostracal processes in Japonia saetigera ...

pp. 185-198

Figures 23-24. 23 - Habitat of Japonia saetigera (van Benthem Jutting, 1958) in River Gam valley, Misool (consider

constant water level and flow); 24 - Model of possible adhesive role of periostracal processes in Japonia saetigera

(van Benthem Jutting, 1958) (those processes potentially in a contact with leaf litter marked red).

197

Book4.indd 197 02-Jun-21 21:49:13

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figures 25-26. "Mychopoma" pennatum van Benthem Jutting, 1958, holotype (NMNL), apical (25) and apertural

(26) view (consider extreme feather-like periostracal processes on teleoconch, with possible adhesive or floating

function). This species shares its habitat with Japonia saetigerum (van Benthem Jutting, 1958) [not to scale].

198 Sa

Book4.indd 198 02-Jun-21 21:49:18

Book4.indd 199

Hava, J.: Orohinus (Orphinus) terminalis (Sharp in Blackburn & Sharp, 1885) (Coleoptera: Dermestidae) from ...

p. 199

SHort NotTE

Orphinus (Orphinus) terminalis (Sharp in Blackburn &

Sharp, 1885) (Coleoptera: Dermestidae) from

the Solomon Islands and Vanuatu

urn:lsid:zoobank.org:pub:A /9F855E-/7A45-4A68-B 7 FA-DD16221FE5/0

Jiri HAVA

Forestry and Game Management Research Institute, Strnady 136, CZ-156 OO, Praha 5 - Zbraslav,

Czech Republic; jh.dermestidae@volny.cz

Abstract: The species Orphinus (Orphinus) terminalis (Sharp in Blackburn & Sharp, 1885) is newly recorded from

Solomon and Vanuatu Islands.

Key words: Dermestidae, new records, Pacific.

Introduction

The dermestid genus Orphinus Motschulsky,

1858 is distributed worldwide with 128 known

species (Hava 2015). The present faunistic

contribution newly records one species as new for

the Solomon Islands and Vanuatu.

Material and methods

Following abbreviations refer to the collections,

in which the examined material is deposited:

JHAC - Jifi Hava, Private Entomological Laboratory and

Collection, Unétice u Prahy, Praha-zdpad, Czech

Republic;

NMPC - National Museum, Prague, Czech Republic.

Results

Orphinus Motschulsky, 1858

Subgenus Orphinus Motschulsky, 1858

Orphinus (0O.) terminalis (Sharp in Blackburn &

Sharp, 1885)

New records: 14° & 19 NMPC: SOLOMON ISLAND,

Guadalcanal, Honaira (gardens), 19.xi-13.xii.2013, Jiri

Hajek leg.; 19 JHAC: VANUATU, Efate, Tagabe Agr. Stn,

22 Nov-2 Dec 1983, malaise trap, P.A. Maddison leg.

Distribution: This species is known from the

islands of Hawaii, the Philippines, Cook, Gilbert,

South Mariana, Marshall, Samoa and Upolu (Beal

1961; Hava 2015). New to the Solomon Islands

and Vanuatu.

Acknowledgements

| would like to express my thanks to Jiri Hajek

(NMPC) for the Solomon Islands material loan and

to Dmitry Telnov (The Entomological Society of

Latvia, Riga) for proofreading the manuscript.

The research was supported by the Ministry

of Agriculture of the Czech Republic, institutional

support MZE-ROO118.

References

Beal R. S. 1961. Coleoptera: Dermestidae. Insects of

Micronesia, Volume 16. Bernice P. Bishop Museum,

Honolulu: 109-131.

Hava J. 2015. World Catalogue of Insects. Volume 13.

Dermestidae (Coleoptera). Brill, Leiden & Boston:

xxvi + 419 pp.

Received: 14.v.2020.

Accepted: 19.vi.2020.

199

02-Jun-21 21:49:18

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

200

Book4.indd 200 02-Jun-21 21:49:18

Book4.indd 201

HAva, J.: Anew synonym in the genus Orphinus Motschulsky, 1858 (Coleoptera: Dermestidae) from ...

pp. 201-202

SHort NOTE

A new synonym in the genus Orphinus Motschulsky,

1858 (Coleoptera: Dermestidae) from Papua New

Guinea

urn:lsid:zoobank.org:pub:6B 7C5/ 79-91D3-43C9-BB9/7-DE952AEC65D0

Jiri HAVA

Forestry and Game Management Research Institute, Strnady 136, CZ-156 OO, Praha 5 - Zbraslav,

Czech Republic; jh.dermestidae@volny.cz

Abstract: The species Thanmaglossa testaceipes Pic, 1915 syn. nov., is herewith synonymized with Orphinus

fulvipes (Guérin-Méneville, 1838).

Key words: Taxonomy, new synonym, Coleoptera, Dermestidae, Orphinus, Papua New Guinea.

Introduction

The dermestid genus Orphinus Motschulsky,

1858 is distributed worldwide with 128 known

species (Hava 2015). The present taxonomic

contribution newly synonymizes one species from

Papua New Guinea.

Material and methods

Labels (if there are multiple labels on a

specimen) are separated by a slash. Following

abbreviations refer to the collections, in which the

examined materials are deposited:

JHAC - Jiff Hava, Private Entomological Laboratory and

Collection, Unétice u Prahy, Praha-zdpad, Czech

Republic;

MNHN - Museum National d “Histoire Naturelle, Paris,

France.

Results

Orphinus Motschulsky, 1858

Subgenus Orphinus Motschulsky, 1858

Orphinus (0.) fulvipes (Guérin-Méneville, 1838)

= Thanmaglossa [sic!] testaceipes Pic, 1915: 6 syn.

nov.

Orphinus testaceipes: Pic (1916: 3); Hava (2015: 181).

Holotype 4 MNHN: Thanmaglossa testaceipes Pic:

,»otephansort, Astrolabe B., D.N.Guinea, (Kunzmann

1894)“ / ,Orphinus probabil“ / ,Thaumaglossa

testaceipes Pic“ / ,, Type“ / , TYPE“ / HOLOTYPE Orphinus

testaceipes Pic, Jiri Hava det. 2019 / Orphinus fulvipes

(Guér.) Jiff Hava det. 2019.

New record: 192 JHAC: N.-Guinea, Biré 1900 /

Stephansort, Astrobe Bay [det. J. Hava].

Figure 1. Thanmaglossa testaceipes Pic, 1915,

holotype 4 (MNHN).

Ree

201

02-Jun-21 21:49:19

TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Acknowledgements

ef (yrnins Ak edau.

Cah wht fi LA | would like to express my thanks to Azadeh

‘ Taghavian (MNHN) for the type material loan and to

Dmitry Telnov (The Entomological Society of Latvia,

Riga) for proofreading the manuscript.

The research was supported by the Ministry

of Agriculture of the Czech Republic, institutional

support MZE-ROO118.

References

Hava J. 2015. World Catalogue of Insects. Volume 13.

Dermestidae (Coleoptera). Brill, Leiden & Boston:

xxvi + 419 pp.

Pic M. 1915. Nouvelles especes de diverses familles. —

Mélanges Exotico-Entomologiques 15: 2-14.

Pic M. 1916. Notes et descriptions abrégées diverses. -

Mélanges Exotico-Entomologiques 17: 2-8.

Received: 22.ix.2020.

Accepted: 30.x.2020.

Figure 2. Thanmaglossa testaceipes Pic, 1915,

holotype, labels.

Remarks: The species Thanmaglossa testaceipes

was originally described by Pic (1915) and later

transferred to the genus Orphinus (Pic 1916).

All morphological characters of this species are

identical with those of O. fulvipes. Orphinus

testaceipes is therefore considered a new junior

synonym of O. fulvipes.

202

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Book4.indd 203

INGRISCH, S.: New species and records of Tettigoniidae Krauss, 1902 from the Papuan Region (Orthoptera)

pp. 203-214

New species and records of Tettigoniidae Krauss, 1902

from the Papuan Region (Orthoptera)

urn:lsid:zoobank.org:pub:D51D90E5-CB20-419B-BEC/7-433FBBCBFE52

SIGFRID INGRISCH

Zoological Research Museum Alexander Koenig, Leibniz Institute for Animal Biodiversity,

Adenauerallee 160, D-53113, Bonn, Germany; s.ingrisch@leibniz-zfmk.de

Abstract: The article is based on Tettigoniidae collected during expeditions to West Papua and Misool Island in

2009, 2012 and 2018, deposited in the Naturkundemuseum Erfurt. Three species of the tribe Sexavaini within

the subfamily Mecopodinae, are described as new: Phrictaetypus brevispina sp. nov. from Tamarau Mountains,

Huona inarmata sp. nov. from Biak Island, and Mossula striatifrons sp. nov. from the Star Mountains. New records

for six species of Agraeciini within the subfamily Conocephalinae are reported, of which PSeudonicsara (Cercana)

spinibranchis Ingrisch, 2009 and Rhytidaspis camela Ingrisch, 2019 are new records for Misool Island. The female

of R. camela is described for the first time.

Key words: Mecopodinae, Sexavini, Conocephalinae, Agraeciini, new species, new records, New Guinea, Misool,

Biak.

Introduction

The Tettigoniidae Krauss, 1902 fauna of New

Guinea is still fragmentary known although many

species of this family are rather large insects. A

search for “Tettigoniidae from Papuasia” (including

New Guinea, Bismarck Archipelago and the

Solomon Islands) in the online database Orthoptera

Species File (Cigliano et a/. 2020) revealed a list

of 454 species, of which more than half belong to

the Conocephalinae Burmeister, 1838 for which a

few modern revisions exist (239 species with 16

species in Conocephalini Burmeister, 1838, 223 in

Agraeciini Redtenbacher, 1891). The Mecopodinae

Walker, 1871 were listed with 44 species. Other

subfamilies which were reported but not dealt

with in this publication are: Meconematinae

Burmeister, 1838 with 59 species, mostly Phisidini

Jin, 1987 that have been revised by Jin & Kevan

(1992), Phaneropterinae Burmeister, 1838 with 44

species, Phyllophorinae Stal, 1874 with 42 species,

Pseudophyllinae Burmeister, 1838 with 18 species,

and Hexacentrinae Karny, 1925 with eight species.

Thus only seven of the currently recognized 19

extant subfamilies have been reported from that

area. At least for some of these subfamilies equally

high numbers of species as in Conocephalinae or

even higher numbers could be expected, which

shows the poor knowledge on the tettigoniid fauna

of the area.

The recent expeditions to New Guinea and

Misool Island took place in 2009, 2012, and 2018.

The Tettigoniidae specimens studied comprised

species of two taxonomic groups: genus group

Mossulae Willemse, 1961 of the tribe Sexavaini

Karny, 1924 within Mecopodinae and the tribe

Agraeciini Redtenbacher, 1891 of Conocephalinae.

The last revisionary work on the Mossulae group

to which the new species belong was finished

by Willemse (1961), while for some of the New

Guinean Agraeciini, revisions of several genera

had been made by Willemse (1959) and the author

(Ingrisch 2008; 2009; 2015; 2019; 2020). In the

last-mentioned publication also a new species from

the collection of D. Telnov has been described,

Salomona_ lanigera Ingrisch, 2020, as it was

reasonable to publish it together with other species

of the same genus.

For the Mecopodinae from South East

Asia, it has been recently shown that acoustic

communication by stridulation and the stridulatory

apparatus with the stridulatory file on the underside

of the left male tegmen and the scrapper and mirror

of the right male tegmen are important taxonomic

characters (Liu et a/. 2020). For the Agraeciini

more traditional characters as differences in the

male and female abdominal appendages like male

cerci and sclerotized parts of the male phallus,

the titillators, and the female subgenital plate are

useful characters to differentiate between species.

203

Tan

€

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TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Material and methods

The specimens were dry pinned or affixed when

received. They were relaxed and the wings spread

when necessary to study the stridulatory apparatus

in Sexavini or to extract the male titillators in

Agraeciini. The specimens of the Mossulae group

studied for this publication do not have slerotized

titillators on the phallus. Documentation of the

specimens studied was done by photography using

a Canon D500 mounted to a copy stand for habitus

images and a Canon D6 mounted to the photo

adapter of a Motic M5 for microscopic images.

The images were processed with CaptureOne and

stacked with Zerene Stacker. Habitus images were

photographed together with a piece of scale paper.

Microscopic images were done at one of four fixed

magnifications. For each magnification also a

micrometer was photographed that allowed to add

scales to the finished images.

Types of the new species are deposited at

the Naturkundemuseum Erfurt, Germany (NMEGQ).

Voucher specimens for the new records that are not

types are deposited in the same collection.

Results

New descriptions

Tettigoniidae Krauss, 1902

Mecopodinae Walker, 1871

Sexavaini Karny, 1924

Mossulae group

Phrictaetypus Brunner von Wattenwyl, 1898

Four species were previously known (Cigliano et

al. 2020): P. viridis Brunner, 1898 from New Guinea

without exact locality, P. nigroornatus Willemse,

1940 from “Solomon Is., Makira, Kirakira”, P.

flavoornatus Willemse, 1953 from “Bougainville

Island, Buin”, and P. aberrans Willemse, 1961 from

“Bougainville”.

Phrictaetypus brevispina sp. nov. (Figs 1, 16-19)

Holotype 2 (3261563) NMEG: Indonesia E, West

Papua, Tamarau Mts., ca 12 km NW Fef vill., 0°46’5’’S,

132°19’6”E, 1050 m, 12-13.ii1.2012, primeval lower

montane rain forest, MV light.

Derivatio nominis: The name of the new species

refers to the short spines at the lateral angles of

the pronotum compared to the long spines in other

204

species of the genus; from Latin “brevis” (Short)

and “spina” (spine).

Measurements, holotype 9: Body length with

wings 68 mm, body length without wings 37 mm,

pronotum length 6.2 mm, tegmen length 59 mm,

tegmen width 10 mm, hind femur length 28 mm,

antenna length 90 mm, ovipositor length 16 mm.

Description. Long and slender species with long,

narrow tegmen almost reaching tip of stretched

hind tibia. Head rather small with eyes strongly

protruding laterally. Fastigium verticis narrow,

furrowed at tip, Separated by a narrow furrow

from fastigium frontis; lateral ocelli inserted at

lateral margins of fastigium verticis, medial ocellus

below fastigium frontis; antennal scrobae with

inner margin moderately expanded dorsad (Fig.

17). Pronotum divided by two deep furrows into

three sections, the last section with an additional

furrow restricted to disc; disc strongly rugose and

with angular and slightly projecting lateral margins;

lateral margins provided with four short blunt

spines: one in first section, two in second section,

and one at beginning of third section; median keel

rather large but blunt, interrupted by the main

transverse furrows, reaching from anterior margin

to just behind the additional furrow in metazona and

interrupts that furrow in middle (Fig. 1). Prosternum

with a pair of spine-like projections, little curved

laterad; mesosternum with parallel, band-shaped

projections that towards end are curved latero-

dorsad; metasternum with ventral surface nearly

smooth but little wavy, lateral margins forming

elevated rims becoming higher towards end (Fig.

16). Tegmen long and narrow, in basal and central

areas faintly narrower than in subapical area. Tibial

tympana open on both sides. Knee lobes of all legs

spinose. Fore coxa with a spine on dorsal margin;

fore femur with 3 spines on ventro-internal margin,

external margin mute; mid femur with 5 spines on

ventro-external margin, internal margin mute; hind

femur with 10-12 ventro-external and 7-8 ventro-

internal spines; knee lobes of all femora spinose on

both sides, hind knee lobes with one spine.

Male is unknown.

Female: Subgenital plate simply rounded; surface

membranous, marginal area sclerotized and

broadly bulging but interrupted at end in middle by

a narrow membranous zone (Fig. 18). Ovipositor

rather short, upcurved with acute tip (Fig. 19).

Colouration: Body and tegmen_ uniformly

yellowish to reddish brown, probably discolored.

Hind femur with a pre-genicular black spot that is

more expressed on ventral than on dorsal side. The

posterior spines on hind femur and most spines

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INGRISCH, S.: New species and records of Tettigoniidae Krauss, 1902 from the Papuan Region (Orthoptera)

of hind tibia black with light tip. Tegmen sub-

transparent with distinct and little darkened veins

and cross veins; resulting cells filled with minute

uncolored veinlets. Hind wings transparent, along

anterior margin yellowish.

Differential diagnosis: The new species is

characteristic due to the presence of small but

distinct spines at the lateral margins of pro-,

meso-, and metazona of pronotum, the diagnostic

characters of the genus Phrictaetypus. Also, the

globose eyes and the long and narrow tegmen

are typical for the genus. With 59 mm length, the

tegmen is however distinctly longer than in all other

species of the genus described so far (22-48 mm

according to Willemse (1953; 1961)). In general

habitus and with regard to the rather short lateral

spines on lateral angles of pronotum, the new

species agrees best with P aberrans Willemse,

1961. It differs by the absence of black spots on

tegmen and by the longer ovipositor (16 against

12 mm). The tegmen is of almost uniform width

throughout in the new species (greatest subapical

width 10.5 mm, narrowest subbasal width 10.0

mm) while it is markedly wider in subapical than

in Subbasal area in P. aberrans (12 against 9 mm).

The black subapical ring on the hind femur and

the black spines with light tips on hind tibia and

in apical area of hind femur are also characteristic

for the new species. From Pseudophrictaetypus

uniformis Willemse, 1961 the new species differs

by the possession of minute but distinct spines

at the lateral angles of all three sections of the

pronotum, the distinctly higher lateral lobes of

pronotum, and the elongate tegmen with parallel-

sided margins to nearly tip.

Huona Kuthy, 1910

Only one species was formerly known (OSF

2020): H. variegata Kuthy, 1910 from New Guinea,

Huon Golf, Sattelberg. That species was also

recorded by Naskrecki & Rentz (2011) from East

New Britain, Nakanai Mountains, Lamas. The

collection received from Dmitry Telnov contains

an adult male from Biak Island that differs in

diagnostic details to be worth the description of

a new species and two photos of a nymph from

Misool Island. The photos show enough details to

identify the individual as belonging to the genus

Huona, but not to assign it to a distinct species. The

latter record extends the known distribution of the

genus even further west. The known distribution of

the genus Huona thus extends from Misool Island

in the west to New Britain in the east.

pp. 203-214

Huona inarmata sp. nov. (Figs 2, 9-12 & 20)

Holotype 3 (3261564) NMEG: Indonesia E, Papua

Prov., Cenderawasih Bay, Biak Is. W, Biak ca 16 km W,

between Urfu & Yendidori, approx. 1°8’S, 135°5/7’E,

65-35 m, 21.iii.2018, secondary lowland rainforest on

limestone, night collecting.

Derivatio nominis: The name of the new species

refers to the absence of a crest with elevated teeth

at the lateral margins of the metazona of pronotum;

from Latin inarmatus, inarmata unarmed.

Measurements, holotype 4: Body length with wings

51 mm, body without wings 30 mm, pronotum

length 6 mm, tegmen length 41 mm, hind femur

length 33 mm, tegmen width near base 9.5 mm,

in middle 8.5 mm, antenna broken, longest rest 77

mm.

Description: Fully winged species with narrow

tegmen that little surpasses the hind knees (Fig.

2). Head globular, rather small, with eyes little

projecting lateral; fastigium verticis narrow and

deeply furrowed in midline, separated by an

angular incision from fastigium frontis; lateral ocelli

inserted at the lateral surfaces of fastigium verticis,

median ocellus at tip of fastigium frontis. Antennal

scrobae with internal margin strongly elevated (Fig.

11). Mandibles with external surface markedly

impressed. Pronotum rather short with anterior

margin convex in middle, concave at both sides,

hind margin faintly convex, ventral margins rounded,

without humeral sinus; with two transverse furrows;

dorsal surface in pro- and mesozona convex and

with rounded lateral margins, metazona flat with

a stout median keel and angular lateral margins.

Tegmen rather narrow, in more than basal half

little wider than in apical area. Legs rather long

and thin; fore coxa with a spine at dorsal margin;

anterior tibia with tympana conchate at external,

open at internal side; mid tibia in more than basal

third wider than thereafter; fore femur with 4-5

spines on ventro-internal margin, external margin

mute; mid femur with 5-6 spines on ventro-external

margin, internal margin mute; hind femur with 13

ventro-external and 9 ventro-internal spines; knee

lobes of all femora spinose on both sides, hind

knee lobes with one spine.

Male: Stridulatory file on underside of left tegmen

about 4.3 mm long, with over 250 teeth that are

at base extremely dense and narrow and hardly

countable, then gradually widening and increasing

in size to about apical third and getting little denser

again in about apical quarter; from behind middle

to before apical quarter of file with 40 teeth per

mm, in second quarter from base with about 56

205

02-Jun-21 21:49:20

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

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Figures 1-4. Habitus in dorsal view with left wings spread; inserts: head and pronotum, enlarged. 1 - Phrictaetypus

brevispina sp. nov., holotype 2; 2 - Huona inarmata sp. nov., holotype <4; 3-4 - Mossula striatifrons sp. nov.,

holotype 3 (3) and paratype @ (4) [scale bars 10 mm].

206 tee

Book4.indd 206 02-Jun-21 21:49:25

Book4.indd 207

INGRISCH, S.: New species and records of Tettigoniidae Krauss, 1902 from the Papuan Region (Orthoptera)

teeth per mm increasing in size towards middle

(Fig. 9). Mirror at base of right tegmen roughly oval,

at internal side behind basal third partly covered

by a rather low elevated frame that performs

a U-shaped curve behind the mirror and then

continuous as a low rim along the external margin

of the mirror; scrapper little elevated and darkened,

internal margin proximal of scrapper concave and

then convex and strongly thickened, afterwards of

normal shape (Fig. 10). Tenth abdominal tergite

short, apart from the very basal area convexly

projecting behind and incised in middle by an

obtuse notch. Cerci moderately stout, not very

long, straight with slightly converging margins from

base, in about apical quarter curved mediad and

strongly narrowed to obtuse tip that carries a small

compressed tooth that is slightly curved proximad

(Fig. 20). Subgenital plate elongate, in basal area

moderately wide with convex lateral margins that

are prolonged at end into stout, parallel, beam-

like projections; styli strong but thinner than

those projections and little shorter than the gap

between the apical projections (Fig. 12). Phallus

membranous.

Colouration: Pale yellowish to brownish. Face of

general color, with a pair of black bands in ventral

area of antennal scrobae; with pairs of small

irregular black spots on lateral sides of fastigium

verticis, below antennal scrobae, on clypeus, in

lateral areas of mandibles, and on both sides of

antennal scapus; on dorsal side of head with a

black band to and behind antennal scrobae; furrow

in fastigium verticis and a spot behind also black;

pronotum of light color with black ornaments.

Tegmen semi-transparent brownish with scattered

black spots mainly behind radius and in apical area.

Hind wings transparent with brownish veins. Legs

of rather light color; fore and mid femora and tibiae

with rather faint brown spots of various size, hind

femur in about basal half on external, dorsal and

internal sides with distinctly outlined black spots,

partly of hieroglyphic appearance; posterior half

and hind tibia with brownish spots as in fore legs.

Most tergites brownish with black hind margins, the

extension of the black color increases towards end

of abdomen; male tenth abdominal tergite with two

black spots in basal area and dark brown along hind

margin and in midline; epiproct light with a black

spot in middle; cerci of light color. Male subgenital

plate at dorsal side blackish brown, at ventral side

including styli lignt brownish, a large area at base

of disc of light color with an oval black spot on both

sides of base, the area behind the light spot dark

brown with light dots, later black, lateral ascending

pp. 203-214

areas black but not at apical projections.

Differential diagnosis: The new_ species

resembles H. variegata Kuthy, 1910 in shape and

coloration. It differs by larger size and longer wings

from H. variegata. The metazona of pronotum has

the anterior transverse bulge more expressed than

in H. variegata, but the lateral angles of disc form an

indistinct and weak carinula that carries at anterior

angle a small tubercle instead of forming a crest

with 2-3 spikes in H. variegata. Moreover, the hind

femur at ventral margin carries numerous rather

small spines of regular size that only faintly increase

in size towards tip, instead of less numerous spines

that are in about mid-length of large and partly very

large size and decrease in length towards tip; the

hind knees terminate into a short spine instead of a

long spine, and the wart-shaped black spots on the

external and dorsal sides of the hind femur follow

a different pattern. The male subgenital plate has

an apical incision which is distinctly longer than

wide (about 7:5) rather than about quadrate, and

male styli which are shorter than the apical incision

instead of roughly equal length.

Huona sp. (Figs 21-22)

Image of a nymph, @ about third-last instar:

Indonesia E, Raja Ampat, Misool Island, Lilinta (Lelintah)

vill. env., O0O2°02'’54’S, 130°16'19”E, 0O1.IV.2009,

secondary lowland rainforest on limestone, observed by

D. Telnov.

This is currently the westernmost known record

of Huona Kuthy, 1910.

Mossula Walker, 1869

Nine species have so far been assigned to this

genus (OSF 2020): M. caudelli Karny, 1924 from

Central Moluccas (Seram Island), M. intermedia

Willemse, 1940 from New Britain, M. kiriwina

Hebard, 1922 from Trobriand Islands (Kiriwina

Islands), M. loriae Griffini, L908 from New Guinea,

Mt. Astrolaba, M. purarica Griffini, 1908 from

Papua New Guinea, on Purari River, M. salomonis

Kirby, 1891 from Solomon Islands (Malaita, New

Georgia, Bougainville), M. toxopei Karny, 1926

from Central Moluccas (Buru Island), M. variopicta

Willemse, 1961 from the Solomon Islands (Malaita

and Auki), and M. vitticollis Walker, 1869 without

known locality data. According to the key in

Willemse (1961), the new species is most similar

to M. variopicta Willemse, 1961 from Malaita.

207

02-Jun-21 21:49:25

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TeLNnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Mossula striatifrons sp. nov. (Figs 3-8 & 13-15)

Holotype 4 (3261565) NMEG: Indonesia E, New

Guinea, Papua Prov., Star Mountains, Oksibil 13.5-19.5

kit SSE, 04459" 3375, 2140" 42 20° st0- 05"03,02"S,

140°43'16’E, elev. 780-380 m, 14.iii.2018, primary

lowland rain forest, at base of young Pandanus’s leaves.

Paratypus 19 (3261566) NMEG: same data as

holotype.

Derivatio nominis: The name of the new species

refers to the striped face; from Latin stria stripe,

band and frons face.

Measurements, holotype 4: Body length with

wings 59 mm, body without wings 41 mm, pronotum

length 6.7 mm, tegmen length 47.5 mm, hind femur

length 37 mm, tegmen width 11.5-12.0 mm in

basal area, 9.5-10.0 mm in apical area, antenna

length 135 mm. Measurements, paratype 2: Body

length with wings 70 mm, body without wings 42

mm, pronotum length 8.5 mm, tegmen length 59

mm, hind femur 41 mm, mean tegmen width 12.5

mm (12.8 mm near base, 11.9 mm in subapical

area), antenna length 100 mm, ovipositor length

38 mm.

Description: Fully winged species with tegmen

Surpassing the hind knees, in male less so

than in female; in female almost reaching tip of

ovipositor. Male tegmen in basal half distinctly

wider than in apical third, female tegmen faintly

and more gradually narrowing to tip; surface of

tegmen appears rugose due to a fine meshwork

of minute veinlets, but main veins distinct (Figs

3-4). Head large, eyes globular, little projecting

laterad; fastigium verticis triangular with narrow,

furrowed tip, separated by an angular step from

fastigium frontis; lateral ocelli inserted at lateral

surfaces of fastigium verticis, median ocellus

below middle of fastigium frontis; antennal scrobae

with internal margin strongly elevated; mandibles

with external surface markedly depressed with

concave surface (Fig. 7). Pronotum with anterior

margin convex in middle, faintly concave at both

sides, hind margin faintly convex, ventral margin

convex in middle, humeral sinus simply curved;

with two deep transverse furrows; dorsal surface

in pro- and mesozona coarsely rugose with lateral

margins rounded into paranota, in metazona finely

rugose with lateral margins rounded-angular;

metazona with indication of a faint medial carina.

Auditory spiracle long oval, in situ hidden under

pronotum. Prosternal lobes triangularly projecting

with acute tip; meso- and metasternal lobes wide,

flattened, with a minute angular projection at apico-

lateral angles; the apico-internal margins of the

208

mesosternal lobes deviating posteriorly while those

of the metasternal lobes are expanded on internal

side leaving only a narrow oval gap between them

with the rounded internal angles almost touching

each other at end (Fig. 15). Legs rather long

and narrow; anterior coxa with a spine on dorsal

margin; anterior femur with 3-5 internal but no

external spines, mid femur with 5-6 external but

no internal spines, hind femur with 7-10 internal

and 10-11 external spines; all tibiae with spines

on both ventral angles, fore and mid tibiae also with

small spines on dorsal posterior angle, hind tibiae

with large spines on both dorsal angles; all spines

at femora and tibiae black; tibial tympana semi-

conchate leaving the greatest part of tympanum

uncovered (Fig. 13); mid tibia of both sexes

markedly thickened in basal third.

Male: Stridulatory file on underside of left tegmen

about 6.56 mm long, with about 210 teeth that are

very dense and narrow at base, increasing in size in

about basal quarter and getting little denser again in

about apical quarter; in middle of file with 20 teeth

per one mm (Fig. 5). Mirror at base of right tegmen

long oval, at internal side covered by a curved

frame that is wider in anterior half than in posterior

half; scrapper thickened, black, internal margin

proximal of scrapper concave and then convex and

strongly thickened (Fig. 6). Tenth abdominal tergite

for the most part black, rather short; hind margin

except near lateral angles convexly projecting, in

middle notched with rounded bottom; epiproct with

rounded apical margin; cerci elongate with faintly

converging margins, in about apical third more

strongly narrowed and slightly curved mediad, apex

rounded with acute, recurved tooth at top. Epiproct

about triangular with widely rounded tip, surface

grooved. Phallus membranous. Subgenital plate

elongate, in about basal half wider with convex

lateral margins, in apical half narrow elongate

and parallel-sided with a medial furrow, margins

heightened throughout, apex incised; styli about as

long as apical incision (Fig. 8).

Female: Subgenital plate with convex, upcurved

lateral margins, continued seamlessly into bilobate

apical margin; baso-lateral angles with a small

depression (Fig. 14). Ovipositor about as long as

body, substraight, in about apical half very faintly

upcurved, tip acute (Fig. 13).

Colouration: General color reddish brown. Face

whitish brown with six vertical dark brown bands,

the central bands continued at clypeus; part of

ventral and internal marginal area of antennal

scrobae black; scapus with two large black spots,

pedicellus with basal and apical margins black,

02-Jun-21 21:49:26

INGRISCH, S.: New species and records of Tettigoniidae Krauss, 1902 from the Papuan Region (Orthoptera)

pp. 203-214

. SOPH 9

A ; af

oone8

Figures 5-12. Stridulatory file on underside of left tegmen (5, 9), mirror at base of right tegmen (6, 10), face (7, 11)

and subgenital plate (8, 12) of males of: 5-8 - Mossula striatifrons sp. nov., holotype; 9-12 - Huona inarmata sp.

nov., holotype [scale bars 1 mm (figures 5 & 9) and 5 mm (figures 6-8, 10-12)].

@qg ze 209

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Book4.indd 210

TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

flagellum fully black; mandibles whitish with a

black spot in depressed lateral area, apical area

brown, teeth black but at rest hidden under labrum;

vertex medium to dark brown, in male with three

longitudinal black bands: in midline and behind

eyes; in female black bands incomplete; abdominal

tergites dark brown, male tenth abdominal tergite

black, in female last tergite black in central area;

pronotum dark reddish brown, disc with apical rim

black, paranota with a light yellowish brown band in

upper half; tegmen brown, dorsal area little darker,

with black spots along hind margin and black

dots along fore margin; hind wings greyish semi-

transparent, anterior area brownish; legs reddish

brown but coxa, trochanter and basal area of hind

femur yellowish brown, knees blackish; coxae and

trochanter with few dark spots; fore and mid femora

with indication of fine striation; hind femur with

distinct striation of parallel dark strokes in about

basal half and a black stroke along ventral margin;

spines of all legs black. Abdominal tergites with a

small black spot in lateral areas, especially distinct

in male. Female subgenital plate medium brown,

in apical area light brown; ovipositor brown, little

lighter in basal area with a black band at base of

ventral valves. Male tenth abdominal tergite black,

little lighter in lateral areas; epiproct black with light

margin; subgenital plate light to medium brown

with furrow at dorsal side dark brown, at ventral

side black and with two black spots in basal area;

Styli black.

Differential diagnosis: The new species

resembles M. variopicta Willemse, 1961 in general

Shape and coloration. In both species, males have

Shorter wings than females, but in M. striatifrons

sp. nov. the male tegmen is rather suddenly

narrowed in mid-length, and surpasses the hind

knees instead of being gradually narrowed to tip,

and does not reach the hind knees in M. variopicta.

The female tegmen is gradually narrowed from

base to tip and almost reaches the end of the

long ovipositor in M. striatifrons sp. nov., instead

of being suddenly narrowed before tip and little

Surpassing the hind knees in M. variopicta. The

best diagnostic character is however the male

cercus that markedly differs between both species.

In M. variopicta, the male cercus consists of a

rather short and stout trunk that narrows in apical

area to the subacute tip [“process” as of Willemse

(1961)], and it releases a blunt and narrow internal

process dividing rectangularly from the cercus stem

before it narrows. In contrast, in the new species,

the male cercus is elongate with apical area little

curved towards middle and with blunt tip, at which

210

a small black hooked tooth is inserted. The female

subgenital plate of the new species has the widely

rounded hind margin only faintly incised in middle

instead of with “a large triangularly rounded incision

in middle” (Willemse 1961) in M. variopicta.

New records

Tettigoniidae Krauss, 1902

Conocephalinae Burmeister, 1838

Agraeciini Redtenbacher, 1891

Eulobaspis dehaani (Karny, 1920)

Examined material: Indonesia: West Papua, Fak-Fak

Peninsula [sic! Correct name is Onin Peninsula], Fak

Fak 10 km E, Sakarteman River valley, elev. 100-300

m (2°56’54"S, 132° 22’23’E), 26.ix.2010 (14 NMEG).

Remark: The species is known from several

localities in West Papua (Ingrisch 2015).

Eulobaspis variata Ingrisch, 2015

Examined material: Papua [New Guinea NW],

Cenderawasih Bay, Biak Island, Biak W - 16 km W,

between Urfu & Yendidori, elev. 35-65 m (1°8’S,

1357572), 21:1 2048410 NMEG).

Remark: The species has been recently described

from Biak Island (Ingrisch 2015). This is the second

record for that species.

Paramacroxiphus rufus Ingrisch, 2008

Examined material: Indonesia: Papua (New Guinea),

Prov. Jayapura, Jayapura 21 km SSE, Koya Barat ~ 8

km NE; Skouw vill. env., elev. 25-50 m (2°38'34’S,

140°52’6”E), 17.iii.2018 (12 NMEG).

Remarks: The species was so far known from

several localities along the Sepik River in Papua

New Guinea (Ingrisch 2008). This is the first record

for the Jayapura area of Papua Province, Indonesia.

Pseudomacroxiphus szentia (Willemse, 1958)

Examined material: Indonesia: Papua, Star

Mountains, Oksibil 13.5-19.5 km SSE, elev. 380-780

m (4°59’33"S, 140°42’2”E), 14.ii1.2018 (1¢ & 19

NMEG).

Remarks: That species is obviously widely

distributed throughout New Guinea. It has been

found along the central mountain ranges and at

the south coast in both, Papua (Indonesia) and

Papua New Guinea (Ingrisch 2008). Surroundings

of Oksibil is a new locality record, although it had

been recorded from the Star Mountains.

02-Jun-21 21:49:29

INGRISCH, S.: New species and records of Tettigoniidae Krauss, 1902 from the Papuan Region (Orthoptera)

pp. 203-214

Figures 13-20. Habitus in lateral view (13), female subgenital plate (14, 18), thoracic sternites (15-16), face (17),

ovipositor (19), male abdominal apex in dorsal view (20). 13-15 - Mossula striatifrons sp. nov., paratype 9; 16-19

- Phrictaetypus brevispina sp. nov., holotype 2; 20 - Huona inarmata sp. nov., holotype ¢ [scale bars 5 mm (figures

14-18 & 20) and 10 mm (figures 13 & 19)].

Book4.indd 211 02-Jun-21 21:49:35

Book4.indd 212

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Pseudonicsara (Cercana) spinibranchis Ingrisch,

2009

Examined material: Indonesia: Prov. Raja Ampat,

Distr. Misool Barat, Lilinta (Lelintah) vill. env., (2°2’54’’S,

130°16'19’E), 1.iv.2009 (19 NMEG).

Remarks: The species has been described from

the Bintuni Bay area, Maccluer Gulf in West Papua

Province (Ingrisch 2009). The new specimen is the

first record for the Misool Island and Raja Ampat

Islands.

Rhytidaspis camela Ingrisch, 2019 (Figs 23-26)

Examined material: Indonesia: E. Prov. Raja Ampat,

Misool SW, Aduwey (Adua) vill. 2-5 km NNW, valley

of River Hakau, (1°58’46"S, 129°54’37’E), 25-27.

iii.2009 (146 & 19 NMEG); Indonesia: E. Prov. Raja

Ampat, Misool SW, Aduwey (Adua) vill. 5 km NNW, valley

of River Hakau, (1°58’46"S, 129°54’37”E), 26-29.

iii.2009 (24 & 12 nymph, NMEG); Indonesia, West

Papua, Tamarau Mts., Bamus Bama vill. env., 730 m

(0°45'19’S, 132°15’48"E), 11-12. 11.2012 (14 NMEG).

Remarks: The genus Rhytidaspis Redtenbacher,

1891 comprises so far seven species and two

Subspecies, all with rather local distribution in

the Indonesian part of New Guinea including one

species from Waigeo Island. R. camela has been

known so far from only two males from two localities

in West Papua Province (Sorong and Komara)

(Ingrisch 2019). The specimens from the valley of

River Hakau are the first record of a species of the

genus Rhytidaspis from Misool Island and Raja

Ampat Islands while those from Bamus Bama fall

within the previously known range of distribution.

One of the three males from River Hakau shows

an abnormal modification of the tenth abdominal

tergite. The pair of beam-like apical lobes are fused

to a single inflated cyst, while cerci and titillators

show the typical shape of R. camela. The other two

males from the same locality are fully normal.

The material from River Hakau also contains one

adult female, which was formerly unknown. This is

described below.

Description of the female: The subgenital

plate of R. camela is similar to that of R. arfak

Ingrisch, 2019 in ventral view but appears little

stouter. As in the latter species, it has a faint angle

of the lateral margins in about mid-length at the

transition from a more strongly converging basal

half to a less strongly converging apical half; at

end, it is only shortly divided into two angular and

faintly upcurved apical lobes with blunt tips (Fig.

26). The lateral areas of the subgenital plate are

nearly angularly curved dorsad and especially in

HAL,

about basal half very high. In contrast to most other

species of the genus these lateral lobes are nearly

fully sclerotised and darkened and only little curved

laterad in dorsal area (Fig. 25).

Colouration: Head, pronotum and abdomen red

brown; abdominal tergites along lateral margins

black. Tegmen yellow with black spots. Legs

medium brown; femora with rows of black dots or

strokes; spines of all legs with black tips or fully

black. Subgenital plate with ventral surface black

with a yellowish stroke in midline of apical half

before apical excision and a medium brown spot at

lateral angles in basal half; upcurved lateral lobes

blackish brown along midline, medium brown along

anterior and posterior margins but black at tip of

lateral lobes. Ovipositor brown, darker towards end.

Rhytidaspis sp. (Figs 27-29)

Examined material: Indonesia: West Papua, South

Bird’s Neck, Kaimana 35-40 km East, Triton Bay, River

Lengguru valley upriver from Oray vill. (3°43’26’S,

134°6’6’E), 200-300 m, 13.ix.2010 (19 NMEG);

Kaimana 40 km East, Triton Bay, Lobo vill. env.

(3°44’8”"S, 134°5’40”E), 200-300 m, 15-17.ix.2010

(12 NMEG).

Remarks: The two females have been found in

a locality that lies between the areas of R. picta

Redtenbacher, 1891 in the north and west and

R. variata Ingrisch, 2019 in the east. Although of

general similar shape, the female subgenital plate

shows small but clear differences to those in both

other species (Figs 28-29). Compared to R. picta,

the subgenital plate of the female from Triton Bay

is in lateral view missing the short dorsal lobe in

about half-length of the subgenital plate and the

two apical lobes are shorter and blunt instead of

narrow-elongate, closer together, and less strongly

upcurved. Compared to R. variata, the subgenital

plate is more elongate and less strongly curved

in the two females from Triton Bay. They might

represent an undescribed species. However,

females of the genus Rhytidaspis show only weak

differences in the diagnostic characters between

species in contrast to the strong differences found

in the male cerci and titillators, which are of high

diagnostic value. It is thus reasonable to wait with

the full identification until males from that locality

become available.

Acknowledgements

Curator-in-charge from NMEG kindly provided

the specimens for study.

02-Jun-21 21:49:35

INGRISCH, S.: New species and records of Tettigoniidae Krauss, 1902 from the Papuan Region (Orthoptera)

pp. 203-214

Figures 21-29. Habitus in dorsal view (21-23), habitus in lateral view (24, 27), female subgenital plate in lateral view

(25, 28) and in ventral view (26, 29). 21-22 - Huona sp., nymph in about third last instar from environs of Lilinta

village sitting on a tree trunk (21), close-up image, the arrows aim at the nymphal ovipositor and the lateral wing

pads (22); 23-26 - Rhytidaspis camela Ingrisch, 2019, 2 from River Hakau valley, Misool: 27-29 - Rhytidaspis

sp. 2 from Triton Bay, southern Bird's Neck, West Papua (images 21-22 courtesy Dmitry Telnov, the Entomological

Society of Latvia, Riga).

213

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Brunner von Wattenwyl C. 1898. Orthopteren des

Malayischen Archipels, gesammelt von _ Prof.

Dr. W. Kukenthal in den Jahren 1893 und

1894. - Abhandlungen der Senckenbergischen

Naturforschenden Gesellschaft 24: 193-288, pls

16-20.

Cigliano M. M., Braun H., Eades D. C., Otte D. 2020.

Orthoptera Species File. Version 5.0/5.0 http://

Orthoptera.SpeciesFile.org [accessed: 02 August

2020].

Griffini A. 1908. Sopra alcuni Stenopelmatidi e sopra

alcune Mecopodidi Malesi ed Austro-Malesi. - Atti

della Societa Italiana di Scienze Naturali 46: 271-

288.

Hebard M. 1922. Studies in Malayan, Melanesian and

Australian Tettigoniidae (Orthoptera). - Proceedings

of the Academy of Natural Sciences of Philadelphia

74: 121-299, pls 11-22.

IngrischS. 2008. Revisionofthe genera Paramacroxiphus

C. Willemse 1961 and Pseudomacroxiphus

C. Willemse 1961 (Orthoptera: Tettigoniidae:

Conocephalinae: Agraeciini). - Zootaxa 1755:

1-34

https://doi.org/10.11646/zootaxa.1755.1.1

Ingrisch S. 2009. Revision of the genus Pseudonicsara

Karny, 1912 (Orthoptera: Tettigoniidae:

Conocephalinae: Agraeciini). - Zootaxa 2185:

1-122

https://doi.org/10.11646/zootaxa.2185.1.1

Ingrisch S. 2015. A revision of the Axylus group

of Agraeciini (Orthoptera: Tettigoniidae:

Conocephalinae) and of some other species

formerly included in Nicsara or Anthracites. -

Zootaxa 4046: 1-308

https://doi.org/10.11646/zootaxa.4046.1.1

Ingrisch S. 2019. Revision of the genus Rhytidaspis

Redtenbacher, 1891 including the description of a

new genus Hauadrhytidaspis gen. nov. (Orthoptera:

Tettigoniidae: Conocephalinae). - Zootaxa 4661:

343-370

https://doi.org/10.11646/zootaxa.4661.2.5

Ingrisch S. 2020. New species and genera of Agraeciini

(Orthoptera, Tettigoniidae, Conocephalinae) from

New Guinea, Solomon Islands and Sulawesi. -

Zootaxa 4821, No 1: 1-48

https://doi.org/10.11646/zootaxa.4821.1.1

Jin X. & Kevan D. K. McE. 1992. Taxonomic revision and

phylogeny of the tribe Phisidini (Insecta: Grylloptera:

Meconematidae). - Theses Zoologicae 18: 1-360.

Karny H. H. 1924. Beitrage zur Malayischen

Orthopterenfauna. VIII. Die Mecopodinen des

Buitenzorger Museums. - Treubia 5: 137-160.

214

Karny H. H. 1926. Fauna Buruana. Orthoptera, Fam.

Tettigoniidae. - Treubia 7: 145-215, pl. 4.

Kirby W. F. 1891. Notes on the Orthopterous family

Mecopodidae. - Transactions of the Royal

Entomological Society of London 1891, No 3: 405-

AND:

Kuthy D. 1910. Orthoptera nonnulla nova in Nova-Guinea

a Lud. Biré collecta. - Annales Historico-Naturales

Musei Nationalis Hungarici, Pars Zoologica 8:

213-216.

Liu C. X., Heller K. G., Wang X. S., Wu C., Liu F., Zhang

T. 2020. Taxonomy of a katydid genus Mecopoda

Serville (Orthoptera: Tettigoniidae, Mecopodinae)

from East Asia. - Zootaxa 4758, No 2: 296-310.

https://doi.org/10.11646/zootaxa.4/758.2.5

Naskrecki P., Rentz D. C. F. 2011. Chapter 11. A rapid

assessment survey of katydids and relatives of the

Muller Range (Insecta: Orthoptera: Ensifera): 168-

174. In: Richards S. J., Gamui B. G. (eds) Rapid

Biological Assessments of the Nakanai Mountains

and the upper Strickland Basin: surveying the

biodiversity of Papua New Guinea’s_ sublime

karst environments. - RAP Bulletin of Biological

Assessment 60: 2+260 pp.

Redtenbacher ap 1891. Monographie der

Conocephaliden. - Verhandlungen der Zoologisch-

Botanischen Gesellschaft in Wien 41: 315-562,

pls 3-4.

Walker F. 1869. Catalogue of the specimens of

Dermaptera Saltatoria in the collection of the

British Museum. Part Il. Catalogue of Locustidae.

London (British Museum of Natural History): iv +

225-423.

Willemse C. 1940. On a collection of Indo-Australian

Tettigoniidae. - Natuurhistorisch Maandblad 29:

60-64.

Willemse C. 1953. On a collection of Orthoptera from

the Solomon Islands, chiefly from the Institut Royal

des Sciences Naturelles de Belgique. - Bulletin de

I’Institut Royal des Sciences Naturelles de Belgique

29: 1-27.

Willemse C. 1959. Notes on the genus Salomona

Blanchard (Orthoptera, Tettigonioidea, subfam.

Agraecinae). - Publicaties van het Natuurhistorisch

Genootschap in Limburg 11: 1-118, pls 1-43.

Willemse C. 1961. Tettigonoidea of the Papuan

subregion (Orthoptera). 1. Mecopodidae. - Pacific

Insects 3: 93-116.

Received: 06.viii.2020.

Accepted: 12.xii.2020.

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KALASHIAN, M. Yu.: A new species of Endelus Deyrolle, 1864 from Sulawesi, Indonesia, with description of ...

pp. 215-220

A new species of Endelus Deyrolle, 1864 (Coleoptera:

Buprestidae) from Sulawesi, Indonesia, with

description of Endelus (Hexagonodelus) subgen. nov.

urn:lsid:zoobank.org:pub:4A66161B-O9CE-40F 7-A1C8-OEA8/7 DCCAAFB

Mark Yu. KALASHIAN

Scientific Center of Zoology and Hydroecology, National Academy of Sciences of Armenia, 7

Paruyr Sevak Street, 0014, Yerevan, Armenia; mkalashian1@gmail.com

Abstract: Endelus (Hexagonodelus) subgen. nov. is described and reviewed. Endelus (H.) caelestis sp. nov. from

Sulawesi, Indonesia, is described and illustrated. Besides the new species, the new subgenus also includes E.

difformis Deyrolle, 1864, E. brutus Deyrolle, 1864, and E. imperator Obenberger, 1937, all from Greater Sunda

Islands, E. brutus also from the Moluccas, E. violaceipennis Fisher, 1921, from the Philippines, and E. coomanianus

Descarpentries, 1948 and E. coeruleoniger Descarpentries et Villiers, 1963, both from Vietnam. A key to the species

of the subgenus is provided. Lectotypes of Endelus difformis Deyrolle, 1864 and E. imperator Obenberger, 1937 are

designated.

Key words: Sunda Islands, the Philippines, Vietnam, Endelus (Hexagonodelus) subgen. nov., Endelus (Hexagonodelus)

caelestis sp. nov., lectotype designation.

Introduction

The present work continues the author’s study

of the buprestid genus Endelus Deyrolle, 1864

(Kalashian 1995; 1997; 1999a & b; 2007; 2011;

2013; 2017; 2019) and comprises the description

of anew species from Sulawesi, Indonesia. The new

species belongs to a new subgenus, also described

and reviewed in the present paper.

Material and methods

The labels are cited using the following

abbreviations: h (handwritten text), p (printed

text). Additional comments or interpretations are

included in square brackets. The data of individual

labels are separated by a slash.

Acronyms of repositories:

BMNH - Natural History Museum (formerly British

Museum, Natural History), London, United

Kingdom;

MKCY - Collection Mark Kalashian, Yerevan, Armenia;

MNHN - Muséum national d’Histoire naturelle, Paris,

France;

NMPC - Narodni Muzeum, Prague, Czech Republic;

USNM - National Museum of Natural History,

Washington, U.S.A.

Results

Subgenus Hexagonodelus subgen. nov.

Type species: Endelus difformis Deyrolle, 1864

by present designation.

Derivatio nominis: The subgeneric name is

formed from the Latin “hexagon” referring to the

peculiar body shape of the species included, and

“-delus” - the terminal part of the generic name

“Endelus”.

Description: Body rather robust, 1.9-2.1 time

as long as wide, with somewhat hexagonal general

Shape, widened from head to humeral tubercles,

then subparallel approximately to posterior 2/5

of elytra and narrowed towards elytral apices.

Surface rather shiny, without or with very fine

microreticulation. Head large, moderately

transverse, only slightly wider than long, slightly

narrowed anteriorly. Frontovertex deeply concave,

oculofrontal margins subangulate, eyes large,

convex, clearly visible in dorsal aspect, pointed

laterally. Temples clearly visible in dorsal aspect,

moderately divergent backwards, approximately

half the length of eyes. Pronotum distinctly

narrower than elytra, widest approximately near

posterior 1/3, where sides are angular, strongly

convergent forwards and very delicately sinuate or

219

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TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

nearly straight backwards to obtuse basal angles.

Pronotum rather narrowly unevenly flattened along

sides and medially of posterior angles, disc convex,

with one or two slightly pronounced transverse

gibbosities. Scutellar shield rather large, triangular.

Elytra with sides subparallel in approximately

anterior 3/5, then abruptly convergent to apices.

Apices viewed together almost irregularly rounded.

Elytra with distinct humeral tubercles, behind them

rather widely flattened along sides to apical 2/5 of

elytra, elytral disc uneven with more or less deep

depressions and distinct elevations.

Differential diagnosis: The most characteristic

feature of the new subgenus is general body shape

which never occurs in other Endelus species.

Besides, in all other Endelus the head is much

wider, distinctly wider than long. In the deeply

concave frontovertex and uneven elytral surface,

the new subgenus shows some resemblance

to subgenera Kubaniellus Kalashian, 1997 and

Papuadelus Kalashian, 2017, and species-group E.

(Ss. Str.) bakeri Kerremans, 1914 (Kalashian, 2017)

but differs from them by structure of pronotum - in

Kubaniellus pronotum medially with a pair of large

and deep transversely oval, rounded or irregular

foveae, in representatives of E. (Ss. str.) bakeri

group the pronotal disc has distinct transverse

elevations and in Papuadelus there is only one

distinct elevation. Besides, in Papuadelus the eyes

are pointed to the front, temples much longer than

eyes, angularly projecting behind them; punctuation

of head much rougher than in the new subgenus; in

the species of the E. (Ss. str.) bakeri group, head Is

distinctly widened anteriorly.

The new subgenus includes the following species:

Endelus difformis Deyrolle, 1864, E. brutus Deyrolle,

1864, E. imperator Obenberger 1937, E. caelestis

sp. nov., all from Greater Sunda Islands (E. brutus

also from Makian, Moluccas), E. violaceipennis

Fisher 1921, from the Philippines, E. coeruleoniger

Descarpentries & Villiers 1963 and E. coomanianus

Descarpentries 1948, both from Vietnam.

Though both genera of the subtribe Aphanisticina

(Aphanisticus Latreille, 1810 and Endelus) are

presented on both sides of Wallace’ line (Bellamy,

2008), as far as it is known there are only few

groups below the generic level crossing the line

(e.g., Aphanisticus chloris Obenberger, 1928

species-group, A. endeloides Carter, 1924 species

group, E. marseuli Deyrolle, 1864 species-group -

Kalashian, 2011, Kalashian, Kuban, 2014), and

Hexagonodelus subgen. nov. iS another group

distributed in the same manner.

216

Endelus (Hexagonodelus) caelestis sp. nov. (Fig. 7)

Holotype 9 BMNH, INDONESIA: Sulawesi, Utara,

Dumoga-Bone N.P. [Bogani Nani Wartabone National

Park], March 1985 [p] / Lowland forest, ca 200 m [p]

/ Malaise trap [p] / R. Ent. Soc. Lond. Project Wallace,

B.M. 1985-10 [p].

Derivatio nominis: From the Latin “caelestis” -

“celestial” to emphasise the bright blue coloration

of a new species.

Description: Body rather wide, approximately

twice as long as wide, of hexagonal shape as

described above, dorsal surface bright blue,

without microreticulation, shiny; ventral surface

black with very fine bronzy lustre. Body length 5.85

mm, width - 2.9 mm. Head large, narrower than

anterior margin of pronotum. Clypeus turned back,

situated nearly horizontally, transversally triangular,

shallowly arcuately concave distally. Frons barely

widened posteriorly, frontovertex deeply nearly

regularly arcuately concave, concavity posteriorly

not reaching anterior margin of pronotum. Frons

posteriorly of clypeus shortly keel-shaped, behind

this keel with deep, transverse, arcuate split,

between antennal cavities and inner margin of

eyes with well-developed longitudinal keels visible

in frontal aspect as distinct tubercles. Oculofrontal

margins subangulate, eyes large and convex,

protruded from head outline, clearly visible in dorsal

aspect, in lateral aspect unevenly reniform with

large upper portion and small and narrow lower

one. Temples about half as long as eyes, slightly

diverging backward. Frons with rather sparse large

flat round punctures, toward vertex and laterally

with rather small and sparse simple punctures.

Antennomers 1 and 2 large, longitudinal, slightly

enlarged distally, antennomeres 3-4 small,

longitudinal, subcylindrical, very barely enlarged

distally, antennomere 5 slightly longitudinal, finely

enlarged distally, antennomeres 6-11 serrate,

antennomere 6. slightly longitudinal, following

antennomeres strongly transverse. Pronotum

distinctly narrower than elytra, 1.95 times as wide

as long, widest approximately near posterior 1/3

where sides angular, strongly convergent nearly

Straight forwards and very finely sinuate backwards

to obtuse basal angles. Lateral margins very

indistinctly irregularly serrate, bordered with fine

edging shortly continued along posterior margin of

pronotum. Anterior margin of pronotum moderately

bisinuate with protruded medial portion, bordered

with entire narrow edging, posterior margin strongly

bisinuate with wide trapezoidal medial lobe.

Pronotum rather widely irregularly depressed along

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KALASHIAN, M. Yu.: A new species of Endelus Deyrolle, 1864 from Sulawesi, Indonesia, with description of ...

sides and posterior margin, medially of posterior

angles depressions enlarged forwards, disk

convex, with shallow transverse depressions at the

level of approximately anterior 1/4 and posterior

1/3 separating two fine transverse elevations.

Surface in posterior portion (about up to the level

of posterior 1/3 of pronotum) with irregular rather

dense, large, flat punctures, anteriorly with small,

Sparse and irregular simple punctures. Scutellum

large, slightly transversally triangular, nearly

glabrous. Elytra approximately 1.45 times as long

as wide, their sides slightly diverging near humeral

tubercles where elytra are widest, then very slightly

and barely sinuously converging to approximately

posterior 2/5, then nearly straight and very apically

delicately sinuously narrowed to apices. Apices

together almost irregularly rounded. Sides of elytra

finely irregularly serrate, bordered with distinct thin

edging in anterior 3/4, elytral apices with several

fine teeth. Elytra with very uneven structure,

laterally rather widely flattened along anterior

3/5, each elytron with distinct humeral tubercle

continued backwards with slightly pronounced

oblique elevation reaching posterior 1/4 of elytra

and with rather small tubercle laterally at the

level of approximately posterior 1/3 of elytra, with

slightly pronounced elevation behind scutellum and

convex apical portion, with rather deep irregular

depressions medially of humeral tubercle and near

suture at anterior 1/3 and posterior 2/5-1/6 of

elytra. Elytral surface with rather dense irregular

small punctures with hyphen-like bottom very

slightly smoothed backwards, anteriorly surface

with few very fine and short irregular transverse

wrinkles. Prosternal process with rather dense

and rough somewhat rasp-shaped punctures,

metasternum medially before transverse sulcus

with small and superficial sparse punctures,

behind sulcus with denser and larger more distinct

punctures, laterally with superficial arcuate wrinkles

forming net of cells, abdominal sternites with

punctures medially and wrinkles laterally as well,

this structure smoothed backwards. Anal ventrite in

female slightly arcuately notched distally.

Sexual dimorphism: Male unknown.

Differential diagnosis: See key below.

Distribution: Indonesia, North Sulawesi.

Endelus (Hexagonodelus) difformis Deyrolle,

1864 (Fig. 2)

Deyrolle (1864: 236, pl. Ill, fig. 9); Théry (1932: 12).

Lectotype sex unknown, here designated, MNHN:

difformis H. Deyr., Borneo [h, bottom label] / TYPE [p,

pp. 215-220

red paper] / SYNTYPE [p, red paper] / SYNTYPE, Endelus

difformis Deyrolle, 1865 [p] / MNHN, EC 10581 [p] /

TRANSCRIPTIO M. KALASHIAN, 2019: difformis H. Deyr.,

Borneo [p] / Lectotypus, Endelus difformis Deyrolle,

1864, M. Kalashian design., 2019 [p, red paper].

Total body length 4.66 mm, maximum width 2.25

mm.

Distribution: Borneo (unstated whether Indonesia

or Malaysia).

Notes: Described from an unstated number of

specimens from “Borneo” (without indication of

exact locality). A Specimen deposited at the MNHN

has been selected as lectotype and the designation

is made to maintain nomenclatural stability.

Endelus (Hexagonodelus) brutus Deyrolle, 1864

(Fig. 1)

Deyrolle (1864: 236); Théry (1932: 11).

Holotype, sex unknown, MNHN: brutus H. Deyr., Makian

[h, bottom label] / TYPE [p, red paper] / HOLOTYPE [p,

red paper] / Museum Paris, 1952, Coll. R. Oberthur

[p] / HOLOTYPE, Endelus brutus Deyrolle, 1865 [p] /

MNHN, EC 10582 [p] / TRANSCRIPTIO M. KALASHIAN,

2019: brutus H. Deyr., Makian [p] / Holotypus, Endelus

brutus Deyrolle, 1864, M. Kalashian design., 2019 [p,

red paper].

Additional material: 19 NMPC: Celebes [Sulawesi,

Indonesia], Dr. Kuhr.

Distribution: Indonesia: Makian Island (North

Moluccas), Sulawesi (Théry 1932; Bellamy 2008).

Notes: Described from Makian from an unstated

number of specimens, but there is a feature

mentioned by H. Deyrolle (absence of head) which

is evidence that he had only one specimen at his

disposal; thus, the headless specimen in MNHN

can be considered as a holotype according to

Article 73.1.2. of International Code of Zoological

Nomenclature (ICZN 1999). Besides the type one

specimen originated from Sulawesi is mentioned by

Théry (1932) [labelled “Pangie, Célébes (C. Ribbe,

1882)”]. | have had the possibility to study another

single specimen also from Sulawesi.

Endelus (Hexagonodelus) violaceipennis Fisher,

1921 (Fig. 6)

Fisher (1921: 443).

Holotype, sex unknown, USNM: [The Philippines]

Davao, Mindanao, Baker/ Type No [p] 51522 [hl]

U.S.N.M [p] 7 Endelus violaceipennis Fisher [h, pencil,

not the original museum label] / Holotypus Endelus

violaceipennis Fisher 1921 M. Kalashian design. 2019

[p, red paper].

217

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Book4.indd 218

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Distribution: The Philippines: Mindanao. paper] / Endelus imperator m. Type [h] det. Dr. Obenberger

Note: Described from “Davao, Mindanao (Baker)”. [p] / Lectotypus, Endelus imperator Obenberger, 1937,

So far known only from the holotype. M. Kalashian design., 2019 [p, red paper].

Endelus (Hexagonodelus) imperator Obenberger, D'Stribution: Indonesia: Java.

1937 (Fig. 3) Note: Described from an unknown number of

Obenberger (1937: 21). specimens from “Java or.: Kediri”. | designate as

the lectotype the male specimen deposited at the

Lectotype <, here designated, NMPC: [Indonesia] NMPC. A lectotype is designated in order to maintain

Pandan Aroe M, res. Kediri, E. Java [h] / Typus [p, red nomenclatural stability.

Figures 1-7. Endelus

(Hexagonodelus subgen.

nov.) spp., dorsal view. 1

- F, (H.) brutus Deyrolle,

1864, holotype (MNHN);

2 - E. (H.) difformis

Deyrolle, 1864, lectotype

(MNHN); 3 - E. (H.)

imperator Obenberger

1937, lectotype (NMPC);

4 - E. (H.) coeruleoniger

Descarpentries

et Villiers, 1963,

holotype (MNHN); 5 -

F. (H.) Ccoomanianus

Descarpentries, 1948,

holotype (MNHN); 6 -

F. (H.) violaceipennis

Fisher, 1921, holotype

(USNM); 7 - E. (H.)

caelestis Sp. nov.,

holotype (BMNH) [not to

scale] (images 1-2 and

4-5 courtesy Christophe

Rivier, MNHN).

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KALASHIAN, M. Yu.: A new species of Endelus Deyrolle, 1864 from Sulawesi, Indonesia, with description of ...

Endelus (Hexagonodelus) coomanianus

Descarpentries, 1948 (Fig. 5)

Descarpentries (1948: 96); Descarpentries & Villiers

(1963: 159-160).

Holotype 92 MNHN: [North Vietnam] TONKIN, Hoa-

Binh, leg. Ade Cooman [p] / Endelus coomanianus mihi

[h] TYPE [p, red] unique [h], A. Descarpentries det. [p] /

TYPE [p, red paper] / HOLOTYPE [p, red paper] / MNHN,

EC 10598 [p].

Distribution: Vietnam.

Note: Described from one female from “Tonkin,

environs de Hoa-Binh (R. P. de Cooman)”. So far

known only from the holotype.

Endelus (Hexagonodelus) coeruleoniger

Descarpentries et Villiers, 1963 (Fig. 4)

Descarpentries & Villiers (1963: 159-160).

Holotype 9 MNHN: [North Vietnam] Tonkin, Hoa-Binh,

Coll. J. Clermont [p] / TYPE [p, red paper] / HOLOTYPE [p,

red paper] / Endelus coeruleoniger n. sp. nob. Holotype,

Q, A. Descarpentries et [h] A. Villiers det. [p] 1963 [h] /

MNHN, EC 10599 [p].

Additional material: 2¢ & 12 MKCY: VIETNAM bor.

Quang Ninh prov., 10 km SE Tien Yen, 1.-14.4.2004, H.

Muhle leg.

Distribution: North Vietnam.

Notes: Described from one female from “Tonkin:

Hoa-Binh (R. P. de Cooman)” [sic!, does not

corresponds to the original labels]. After the

description, a series of this species including about

40 specimens was collected in North Vietnam by

Hans Muhle. Beetles were collected on the fern

plants on the shore of aswamp near forest. Thanks

to Muhle’s kindness | have had the possibility to

study some specimens from this series. According

to his personal communication, the specimens

discussed will be deposited in several European

Museums including BMNH, MNHN, NMPC,

Naturhistorisches Museum in Basel, Switzerland,

and in Zoologische Staatssammlung in Munich,

Germany.

A key to species of Endelus (Hexagonodelus)

subgen. nov.

1 Body above distinctly bicolorous with pronotum golden

with greenish reflection and elytra bright blue. Elytral

surface less uneven, with elevations and depressions

less pronounced. Body 4.42-5.25 mm. Indonesia:

Makian; Sulawesi FE. brutus Deyrolle, 1864 (Fig. 1)

- Body above unicolorous or slightly bicolorous. Elytral

surface strongly uneven, with well pronounced elevations

pp. 215-220

ANGHACPIESSIONS. cesessterscAgaarencccgeadgariddetaceattagseedtazaceane 2

2 Body above bright blue; 5.85 mm. Indonesia:

SUIAWES Iz: toensscrsepeasnearieeeotent F. caelestis sp. nov. (Fig. 7)

—- Body above dark violet with purplish reflection, or dark

bronzy with reddish-cupreous reflection or black with

violet reflection

3 Body rather robust, 1.90 as long as wide, dorsally

slightly bicolorous, dark violet with purplish reflection

distinctly more pronounced on elytra than on pronotum.

Body 5.25 mm. The Philippines: Mindanao

F. violaceipennis Fisher 1921 (Fig. 6)

- Body comparatively slender, 2.0-2.1 times as long as

wide, nearly unicolorous sometimes with less pronounced

difference of tint between elytra and pronotum

4 Dark bronzy with fine reddish-cupreous reflection,

sometimes more pronounced on pronotum than on

elytra. Species from Greater Sunda Islands ............06 5

— At least elytra black with violet reflection. Species from

VLU Gd Nb achicoesatict ean tuamaratananimsdaateatyasstseaaadeseahisia ee ameaatar setae 6

5 Reddish-cupreous reflection on pronotum distinctly

more pronounced than on elytra. Pronotum with more

distinct and dense big flattened punctures, with slightly

pronounced transversal elevation near approximately

anterior portion of pronotum. Body 4.8 mm. Indonesia:

DAVE wircccrdes cena F. imperator Obenberger 1937 (Fig. 3)

- Body above nearly unicolorous. Punctuation of

pronotum less distinct than in previous species, smooth,

transverse elevation situated approximately near middle

portion of pronotum. Body 4.75 mm. Borneo ............+++++

E. difformis Deyrolle, 1864 (Fig. 2)

6 Sides of pronotum behind anterior angles slightly

arcuate, then distinctly sinuate before angularity at

the widest portion of pronotum. Pronotum with fine but

distinct transverse elevation near anterior 1/3. Elytral

sides behind posterior 2/5 nearly straight, slightly

sinuate just before apex. Body 5.75 mm

F. coomanianus Descarpentries 1948 (Fig. 5)

- Sides of pronotum nearly straight, very finely arcuate

behind anterior angles, then very slightly sinuate before

angularity at the widest portion of pronotum. Pronotum

near anterior 1/3 convex but without separated

elevation. Elytral sides behind posterior 2/5 distinctly

sinuate, then nearly straight towards apex. Body 4.6-5.3

mm .... E. coeruleoniger Descarpentries & Villiers 1963

(Fig. 4)

Acknowledgements

lam glad to express my deepest gratitude to

Dr. Maxwell V. L. Barclay (BMNH), Dr. Svatopluk

Bily, Drs Jiri Hajek and Lukas Sekerka (all NMPC),

and Dr. Jean Menier (MNHN, retired) for their

hospitality during my visits to their beautiful

countries and for giving me the opportunity to work

in the collections of their institutions, and to Dr.

Steven Lingafelter (USNM) for the loan of materials

for study. | am also very grateful to Dr. Antoine

? mie

219

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Mantilleri and Christophe Rivier (both MNHN)

and Dr. Eduard Jendek (Bratislava, Slovakia) who

provided me with the photographs for this study,

either included here or used for comparison. |

am especially thankful to Hans Muhle (Nuf8dorf

am Inn, Germany) for providing me with some

materials, photographs and information on Endelus

coeuleoniger. Special thanks to Dr. Dmitry Telnov

(BMNH) for his continuous efforts in promoting the

study of the outstanding biodiversity of Wallacea

and New Guinea and for his hard work on the

beautiful book series “Biodiversity, Biogeography

and Nature Conservation in Wallacea and New

Guinea”. Finally, |!am very thankful to reviewers, Dr.

Mark G. Volkovitsch (Zoological Institute RAS, St.

Petersburg, Russia) and an anonymous reviewer,

whose comments helped very much to improve the

manuscript

References

Bellamy C. L. 2008. A World Catalogue and Bibliography

of the Jewel Beetles (Coleoptera: Buprestoidea).

Volume 4: Agrilinae: Agrilina through Trachyini.

Pensoft Series “Faunistica’ No 79. Pensoft

Publishers, Sofia & Moscow: 1932-2684.

Descarpentries A. 1948. Buprestides nouveaux

d’Indochine frangaise (Col. Buprestidae). - Notes

d’Entomologie Chinoise 12, No 9: 87-98.

Descarpentries A., Villiers A. 1963. Catalogue raisonné

des Buprestidae d’Indochine. IV. Aphanisticini. -

Revue Francaise d’Entomologie 30, No 3: 153-163.

Deyrolle H. 1864. Description des Buprestides de la

Malaisie recueillés par M. Wallace pendant son

voyage dans cet Archipel. - Annales de la Société

entomologique de Belgique 8: 1-280, 4 plates (i-iii

in colour).

Fisher W. S. 1921. New Coleoptera from the Philippine

Islands. Family Buprestidae, tribe Agrilini. - Philippine

Journal of Science, 18, No 4: 349-447.

ICZN 1999. International Code of Zoological

Nomenclature. Fourth edition. The International

Trust for Zoological Nomenclature, London: 106

pp. Available from https://www.iczn.org/the-

code/the-international-code-of-zoological-

nomenclature/the-code-online [accessed: 10

August 2020]

Kalashian M. Yu. 1995. Material on the fauna of buprestid

beetles (Coleoptera, Buprestidae) of Vietnam.

Ill. New species of the genus Endelus Deyrolle

from Vietnam and China. - Entomologicheskoe

Obozrenie 74, No 2: 376-382 (in Russian, English

abstract).

Kalashian M. Yu. 1997. New subgenus of Endelus

(Coleoptera: Buprestidae) with description of three

new species from Vietnam and China. - Folia

220

heyrovskyana 5, No 2: 73-81.

Kalashian M. Yu. 1999a. New species of buprestid genus

Endelus Deyrolle (Coleoptera, Buprestidae) from

Southeast Asia. - Entomologicheskoe Obozrenie

78, No 3: 629-635 (in Russian, English abstract).

Kalashian M. Yu. 1999b. New species of Aphanisticus

and Endelus (Coleoptera: Buprestidae) from

South-East Asia. - Folia heyrovskyana 7, No 5,

293-300.

Kalashian M. Yu. 2007. New species of buprestid genus

Endelus Deyrolle (Coleoptera, Buprestidae) from

China, India and Laos. - Entomologicheskoe

Obozrenie 86, No 3: 655-664 (in Russian, English

abstract).

Kalashian M. Yu. 2011. New species of Endelus

Deyrolle, 1864 (Coleoptera: Buprestidae) from

Sulawesi, Indonesia: 83-87, pl. 16. - In: Telnov

D. (ed.) Biodiversity, Biogeography and Nature

Conservation in Wallacea and New Guinea. Volume

I. The Entomological Society of Latvia, Riga: 434

pp, 92 pls.

Kalashian M. Yu. 2013. Two new species of the genus

Endelus Deyrolle, 1864 (Coleoptera: Buprestidae)

from Malaysia and the Philippines with notes on

the synonymy of some species of the subgenus

Endelus s. str. - Caucasian Entomological Bulletin

9, No 1: 83-88 (in Russian, English abstract).

Kalashian M. Yu., Kuban V. 2014. New species

of Aphanisticus Latreille, 1810 (Coleoptera,

Buprestidae) from Sulawesi and Australia: 213-

218, Pl. 29-30. In: Telnov D. (ed.) Biodiversity,

Biogeography and Nature Conservation in Wallacea

and New Guinea. Volume Il. The Entomological

Society of Latvia, Riga: 658 pp, 172 pls.

Kalashian M. Yu. 2017. A new subgenus and three new

species of Endelus Deyrolle, 1864 (Coleoptera:

Buprestidae) from Indonesia and the Philippines,

with a review of the Endelus bakeri Kerremans,

1914 species group: 349-356, pl. 51. In: Telnov D.,

Barclay M. V. L., Pauwels O. S. G. (eds) Biodiversity,

Biogeography and Nature Conservation in Wallacea

and New Guinea. Volume Ill. The Entomological

Society of Latvia, Riga: 658 pp, 172 pls.

Kalashian M. Yu. 2019. Two new species of the buprestid

genus Endelus Deyrolle, 1864 (Coleoptera,

Buprestidae) from Vietnam. - Entomologicheskoe

Obozrenie 98, No 1: 152-158 (in Russian, English

abstract).

Obenberger J. 1937. De novis generis Endelus H. Deyr.

speciebus tribus (Col. Bupr.). Tri nové druhy rodu

Fndelus H. Deyr. (Col. Bupr.). - Acta Entomologica

Musaei Nationalis Pragae 15: 21-22.

Théry A. 1932. Contribution a l’etude des espéces du

genre Endelus H. Deyr. (Coleopt. Buprestidae). -

Novitates Entomologicae 2: 1-23.

Received: 10.viii.2020.

Accepted: 09.ix.2020.

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KaLNINS, M.: Contribution to the knowledge of dragonflies (Odonata) from the Raja Ampat (Indonesia), with notes ...

pp. 221-238

Contribution to the knowledge of dragonflies (Odonata)

from the Raja Ampat (Indonesia), with notes on their

ecology

urn:lsid:zoobank.org:pub:B63F3548-/740D-4F43-970A-85E0E 7/O09068C

MARTINS KALNINS * 2

1 - Department of Biosystematics, Institute of Life Sciences and Technology, Daugavpils

University, Vienibas iela 13, LV-5401, Daugavpils, Latvia; martins.kalnins@biology.|v

2 — Joint Stock Company “Latvia's State forests”, Vainodes iela 1, LV-1004, Riga, Latvia

Abstract: This article contains information on 103 Odonata species from Raja Ampat - Waigeo, Misool, Salawati,

and Batanta Islands. No information was found on the species composition of Kofiau dragonflies. Eleven Odonata

species are new to the fauna of Misool Island, collected or observed by the author and five Odonata species found

by other researchers. Three Odonata species are new records for the fauna of Waigeo Island. Additional information

on ecology and behaviour of several species is also given.

Key words: Odonata, dragonfly, fauna, Raja Ampat, Wallacea, Papua, Indonesia.

Introduction

The Raja Ampat, or the Four Kings, is an

archipelago comprising over 1500 small islands,

cays, and shoals surrounding the four main

islands of Misool, Salawati, Batanta, and Waigeo,

and the smaller island of Kofiau. Administratively,

the archipelago is part of the province of West

Papua (formerly Irian Jaya) in Indonesia. The

Raja Ampat archipelago straddles the Equator

and forms part of the Coral Triangle. The islands

are part of the Vogelkop-Aru lowland rainforests

ecoregion. Generally, this ecoregion exhibits low

to moderate richness and endemism compared

with those of other ecoregions in Indo-Malaysia.

The surface geology of this ecoregion is composed

predominantly of sedimentary rock and recent

alluvium, with some large areas of limestone or

ultramafic near Sorong and on Waigeo and Misool

islands (Wikramanayake et al. 2002).

Waigeo, also known as Amberi, or Waigiu,

has an area of 3155 km? and its highest point

is 958 m. The town of Waisai in the east of the

island is the capital of the Raja Ampat Regency.

According to remote sensing data (cf. Google Earth,

Sentinel), most of the island is covered by primary

lowland rainforests. Due to relief hardly accessible,

Waigeo is still 70-80% covered by dense pristine

rainforests. The largest settlements and areas of

human impact are on the north coast of the island.

Misool, formerly spelled Mysol (Dutch: Miso6l)

or Misol has an area of 2034 km2. In some sources

Misool is also called Batanme (Anonymous 2004).

The highest point is 561 m and the main towns are

Len Malaas, located on the island’s northern coast

and Lilinta (Lelintah), located on the island’s south

coast (Some maps, such as the Mapcarta https://

mapcarta.com/15621866, incorrectly show the

location of Lilinta as about 20 km northwest).

According to remote sensing data (cf. Google Earth,

Sentinel), most of the island is covered by primary

lowland rainforests. Hardly accessible, Misool is

still 80-90% covered by dense pristine rainforests

and there are almost no roads or active airstrips

there. The largest settlements and areas of human

impact are on the north coast of the island. While

the Western part of the island is mostly formed by

basalt formations the eastern part lies on limestone.

Salawati has an area of 1623 km?. Salawati

is separated from New Guinea to the southeast

by the Sele Strait (Galowa Strait, Revenges Strait),

and from Batanta to the north by the Pitt Strait

(Sagewin Strait). According to remote sensing data

(cf. Google Earth, Sentinel), most of the island is

covered by primary lowland rainforests, although

the island, including its central part, has a relatively

well-developed road network and a relatively large

number of settlements.

Batanta has an area of 453 km? and its

highest point is 1184 m. The Pitt Strait Separates it

221.

Tan

€

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from Salawati, while the Dampier Strait (Augusta’s

Strait) separates it from Waigeo. According to

remote sensing data (cf. Google Earth, Sentinel),

most of the island is covered by primary lowland

rainforests. Hardly accessible, Batanta is still

80-90% covered by dense pristine rainforests

and there are almost no roads or active airstrips

there. The largest settlements and areas of human

impact are on the coasts of the island.

Kofiau has an area of ~180 km? and its highest

point is 255 m. The island is primarily raised coral

limestone with some volcanic hills. According to

remote sensing data (cf. Google Earth, Sentinel),

at least 70% of the island is covered by primary

lowland rainforests, but the W part of the island,

the N coast and the small islands are affected by

human economic activities - coconut plantations

have been established there.

Information about dragonflies in Raja Ampat

can be found mainly in Lieftinck’s publications

(Lieftinck 1948; 1971 etc.). The first work to provide

details of all the dragonflies and damselflies found

in Batanta, Misool, Salawati and Waigeo is the work

of John Michalski (2012), although a checklist of

dragonflies for each island was not provided in

this work. Over the last few years, two works have

been published with a checklist of damselflies

(Kalkman & Orr 2013) and dragonflies (Orr &

Kalkman 2015) recorded from New Guinea and

the adjacent islands. Among these 17 Zygoptera

species and 12 Anisoptera species are identified

for the Odonata fauna of Waigeo, 13 Zygoptera

species and 16 Anisoptera species are identified

for Misool, and 9 Zygoptera species and 12 (one

more species mentioned as possible) Anisoptera

species were listed for Batanta and Salawati

(together). Tibor Kovacs has published four works

(Kovacs et al. 2015a & b; 2016; 2021), in which 24

Zygoptera species and 37 Anisoptera species (and

Metagrion sp. not identified to species level) have

been identified for Batanta Island (including Arefi

and Birie Islands).

New data became available on the Misool and

Waigeo dragonfly fauna since the publication of

the above-mentioned checklists, both published

(cf. Theischinger & Kalkman 2014a & b) and

unpublished; the purpose of the present paper is

to provide an updated checklist of the Odonata of

Raja Ampat Islands.

Material and methods

Data on dragonflies were obtained mainly from

222

the author’s personal observations on dragonflies

in Misool (including a single observation from

adjacent Polee islet) and Dmitry’ Telnov’s

observations from Misool and Waigeo. Both field

and hand identification was performed. The

processed material covers several observation

sites in Misool and three in Waigeo (Maps 1-3).

The following abbreviations (key words of localities

of the species) are used in the description of the

material:

Aduwey - 28.03.2009 and 30.03.2009 Misool SW,

Aduwey (Aduai) vill., 01°59’4"S, 129°54’32’E,

village, gardens, bushes & small ditches or pools;

Biga - 03.02.2012 Misool S, Biga vill. ~7,5 km W, River

Biga valley, 02°01’23”"S, 130°12’38’E, primary

lowland forest on limestone;

Gam - 03.04.2009 Misool SE, Lilinta (Lelintah) vill. ~13

km NW, Gamta vill. ~1.4 km NWW, River Gam valley,

01°57’49"S, 130° 11’10’E, primary lowland forest;

Gam* - 04.-05.02.2012 Misool (central), Gamta

vill. 12-14 km NW, River Gam valley, 01°57’50’S,

130°11’09”E, primary lowland rainforest on

limestone;

Hakau 1 - 25.03.2009 and 27.03.2009 Misool SW,

Aduwey (Aduai) vill. ~3 km NNW, valley of River

Hakau, 01°57’22”"S, 129°54’43’E, near river, ina

primary lowland forest;

Hakau 2 - 29.03.2009 and 31.03.2009 Misool SW,

Aduwey (Aduai) vill. ~1,5 km NNW, valley of River

Hakau, 01°58'21’"S, 129°54’30”E, in lowland

forest/forest gardens (Fig. 1);

Ifeylo - 26.03.2009 Misool SW, Aduwey (Aduai) vill.

~4 km NNE, valley of River Ifeylo, 01°57’49’S,

129°56'17’E, near river in primary lowland forest;

Lilinta - 01.-02.04.2009 Misool SE, Lilinta (Lelintah) vill.

env., 02° 02’47"S, 130° 16’7”E, secondary lowland

forest;

Polee - 02.04.2009 Misool S, Lilinta (Lelintah) vill. ~16

km S, Polee Island, 02°11’56’S, 130°16’19’E,

primary lowland forest;

Vanlal - 03.04.2009 Misool SE, Lilinta (Lelintah) vill. ~14

km NW, Gamta vill. ~2 km NWW, River Vanlal valley

near the waterfall, 01°57’'17”S, 130°13’47’E;

Waisai 1 - 16.-17.02.2012 Waigeo, Waisai 2-3 km W,

00°25’40"S, 130°47’36”E, secondary lowland

rainforest on limestone;

Waisai 2 - 16.02.2012 Waigeo, Waisai 10-13 km NE,

00°21'17"S, 130°54’37’”E, primary lowland

forest, limestone creek;

Waisai 3 - 17.02.2012 Waigeo, Waisai 4-6 km (around

Waisai_ airport), O0°24’46’S, 130°44'11’E,

coastal lower bog.

Dragonflies from Raja Ampat region were also

searched for on the insect trader website, Insect-

Trade.EU (https://www.insect-trade.eu). Soecimens

were not purchased, but the information available

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on the website was used - in particular, the image

(for species identification) and its locality.

The term “Papuan Region” (or in the strict

zoogeographic sense - the Papuan subregion of

the Australian Region) is often listed in zoological or

zoogeographic literature, but sometimes extended

to different areas. Therefore, for the purposes of this

work, the definitions as explained in the literature

(e.g., Greke & Telnov 2014) - Papuan Region sensu

lato and sensu stricto - have been used (Map 4).

Results

Brackets after the reference indicate the

location of the species only in the Raja Ampat

Archipelago. Papers indicating that the species is

found in the surrounding islands of New Guinea,

without specifying specific sites, are not cited.

Zygoptera Selys, 1854

Argiolestidae Fraser, 1957

Argiolestes australis Guerin, 1832

Kalkman & Orr (2013) Waigeo; Kovacs et al. (2016;

2021) Batanta; Kalnins (2017) Waigeo.

Note: An unidentified specimen of Argiolestes sp.

also known from Misool (Kalnins 2014).

Argiolestes varga Kovacs et Theischinger, 2021

Kovacs et al. (2021) Batanta.

Metagrion coartans Lieftinck, 1956

Kalkman & Orr (2013) Waigeo.

Metagrion ochrostoma Lieftinck, 1949

Kalkman & Orr (2013) Waigeo.

e Metagrion postnodale Selys, 1878

Kovacs et al. (2015b; 2016) Batanta.

Note: All former records of M. postnodale from

Batanta now refer to Metagrion sp. with a remark

“more study is needed for the delimitation of this

taxon” (Kovacs et al. 2021).

Podopteryx selysi Forster, 1899

Kalkman & Orr (2013) Batanta, Salawati, Waigeo.

Calopterygidae Selys, 1850

Neurobasis australis australis Selys, 1897

Kalkman & Orr (2013) Batanta, Salawati, Waigeo;

pp. 221-238

Kovacs et al. (2015b) Batanta, as Neurobasis australis

s. |., subspecies identification not provided.

Neurobasis australis misoolensis Lieftinck, 1955

Michalski (2012) Misool; Kalkman & Orr (2013) Misool.

Chlorocyphidae Cowley, 1937

Rhinocypha tincta sagitta Lieftinck, 1938

Michalski (2012) Misool; Kalkman & Orr (2013) Batanta,

Misool, Salawati; Kovacs et al. (2015b; 2016; 2021)

Batanta, as Rhinocypha tincta cf. sagitta.

New records: Hakau 1 (124 & 29), Ifeylo (43 & 29),

Vanlal (2).

Note: Common near rivers in primary forest on

Misool, one observation - a female laying eggs on

a dead branch together with a male guarding them

(Fig. 2); only few females observed while males

were abundant.

Rhinocypha tincta tincta Rambur, 1842

Kalkman & Orr (2013) Waigeo.

Coenagrionidae Kirby, 1890

Agriocnemis femina Brauer, 1868

Kovacs et al. (2015b; 2016; 2021) Batanta.

New records: Aduwey (634), Waisai 1 (3¢ & 69),

Waisai 2 (4¢ & 19).

First records for Misool and Waigeo.

Note: Numerous specimens observed in Aduwey

village, at small ditches with dense, partially

submerged aquatic vegetation. During high tide,

seawater enters the ditches (Figs 10-11).

Archibasis mimetes Tillyard, 1913

Kalkman & Orr (2013) Batanta, Misool, Salawati,

Waigeo.

Argiocnemis rubescens Selys, 1877

Kovacs et al. (2016; 2021) Batanta.

Ceriagrion aeruginosum Brauer, 1869

Kovacs et al. (2015b; 2021) Batanta.

Ischnura senegalensis Rambur, 1842

Kovacs et al. (2015b; 2021) Batanta.

Papuagrion magnanimum Selys, 1876

Kovacs et al. (2021) Batanta.

Pseudagrion civicum Lieftinck, 1932

Kovacs et al. (2015b; 2016; 2021) Batanta.

223

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figures 1-5. Odonata from Raja Ampat. 1 - Downstream of River Hakau - mosaic landscape with gardens and secondary

lowland rainforest and partly dried riverbed - the habitat of adult Agrionoptera insignis, Camacina gigantea, Hydrobasileus

brevistylus, Neurothemis stigmatizans, and Orthetrum villosovittatum; 2 - Rhinocypha tincta sagitta in Aduwey, Misool,

a Q (on the left) ovipositing on dead branches partly submerged in the water in a small forest river (stream) and a @ (on

the right) guarding the 9; 3 - Place of oviposition of Agrionoptera longitudinalis - a small hole in a stone filled with water

and some dead leaves; 4 - Camacinia gigantea ¢ in Aduwey, Misool. Individual specimens were resting in sunny places

between bushes in a mosaic landscape; 5 - Hydrobasileus brevistylus in Aduwey, Misool. Pair of specimens observed in

copula, resting in a sunny place in a partly dry riverbed between bushes in a mosaic landscape - river valley with gardens

and secondary lowland rainforest patches.

224

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Pseudagrion starreanum Lieftinck, 1949

Kalkman & Orr (2013) Batanta, Salawati; Kovacs et al.

(2015b; 2016; 2021) Batanta.

Teinobasis michalskii Theischinger et Kalkman

2014

Theischinger & Kalkman (2014a) Misool.

Teinobasis prothoracica Selys, 1877

Michalski (2012) Misool; Kalkman & Orr (2013) Misool,

Waigeo.

Teinobasis rufithorax Selys, 1877

Kalkman & Orr (2013) Batanta, Misool, Salawati, Waigeo;

Theischinger & Kalkman (2014a) Misool; Kovacs et al.

(2015b; 2016; 2021) Batanta.

New records: Ifeylo (23), Lilinta (19).

Teinobasis superba (Hagen in Selys, 1877)

Theischinger & Kalkman (2014a) Misool, as Teinobasis

cf. superba; Kovacs et al. (2016; 2021) Batanta.

Teinobasis wallacei Campion, 1924

Theischinger & Kalkman (2014a) Misool.

Xiphiagrion cyanomelas Selys, 1876

Kovacs et al. (2015b; 2021) Batanta.

Isostictidae Fraser, 1955

e Selysioneura cf. cervicornu Forster, 1900

Kovacs et al. (2015b) Batanta.

Note: Kovacs et al. (2021) stated that all previous

records of S. cf. cervicornu from Batanta refer to

Selysioneura cornelia.

Selysioneura cornelia Lieftinck, 1953

Michalski (2012) Misool; Kalkman & Orr (2013) Misool;

Kovacs et al. (2021) Batanta.

New record: Gam (62).

Tanymecosticta jejuna Lieftinck, 1959

Michalski (2012) Misool; Kalkman & Orr (2013) Misool.

Lestidae Calvert, 1901

Indolestes albicaudus McLachlan, 1895

Michalski (2012) Misool; Kalkman & Orr (2013) Misool.

Platycnemididae Yacobson et Bianchi, 1905

Idiocnemis bidentata Selys, 1878

Ris (1913) Waigeo; Gassmann (2000) Batanta, Salawati,

pp. 221-238

Waigeo; Kovacs et al. (2015b; 2016; 2021) Batanta.

Idiocnemis dagnyae Lieftinck, 1958

Gassmann (2000) Misool; Michalski (2012) Misool;

Kalkman & Orr (2013) Misool.

Idiocnemis fissidens Lieftinck, 1958

Gassmann (1999) Waigeo; Kalkman & Orr (2013)

Waigeo.

Idiocnemis inornata Selys, 1878

Kovacs et al. (2015b; 2016; 2021) Batanta.

Idiocnemis strumidens Lieftinck, 1958

Gassmann (1999) Misool; Michalski (2012) Misool;

Kalkman & Orr (2013) Misool; Kovacs et al. (2016;

2021) Batanta.

Nososticta atrocyana Lieftinck, 1960

Kalkman & Orr (2013) Waigeo; Theischinger & Kalkman

(2014b) Waigeo.

Nososticta aurantiaca Lieftinck, 1938

Kalkman & Orr (2013) Batanta, Salawati; Theischinger &

Kalkman (2014b) Salawati; Kovacs et al. (2015b; 2016;

2021) Batanta.

Nososticta dora Kovacs et Theischinger, 2016

Kovacs et al. (2016; 2021) Batanta.

Nososticta erythroprocta Selys, 1886

Kalkman & Orr (2013); Theischinger & Kalkman (2014b)

Waigeo.

Nososticta evelynae Lieftinck, 1960

Kalkman & Orr (2013) Waigeo; Theischinger & Kalkman

(2014b) Waigeo.

Nososticta nigrifrons (Ris, 1913)

Theischinger & Kalkman (20146) Misool.

Note: Theischinger & Kalkman (2014b: 221)

considered Nososticta lorentzi (Lieftinck, 1938)

to be a junior synonym of N. nigrifrons (Ris, 1913).

However, in the abstract of the same paper

Theischinger & Kalkman (2014b: 217) listed

"Nososticta nigripes (Ris, 1913)" presumably for

N. nigrifrons. The name ‘Nososticta nigripes’ was

never published and is therefore a nomen nudum.

Michalski (2012: 154) stated that “N. /orentzi

appears to be morphologically identical with

N. nigrifrons (which was originally described from

incomplete specimens)”. Later on, the same author

(Michalski 2012: 155-156) listed both N. nigrifrons

nigrifrons and N. nigrifrons lorentzi as separate

oes

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taxa and provided description and drawings of

their (rather distinctive!) differential features such

as thoracic pattern, colouration of abdominal

segments and inconspicuous differences in male

abdominal appendages.

Nososticta plagiata Selys, 1886

Michalski (2012) Misool; Kalkman & Orr (2013) Misool;

Theischinger & Kalkman (2014b) Misool.

New records: Hakau 1 (3d 19), Ifeylo 1 (5¢ 19),

Vanlal (24 19), Biga (1¢ & 19).

Note: The identification of females is tentative and

based on specimens sampled in copula, because

no female descriptions are available for any species

of this genus.

Nososticta pyroprocta Lieftinck, 1960

Michalski (2012) Misool; Kalkman & Orr (2013) Misool;

Theischinger & Kalkman (2014b) Misool.

Nososticta rufipes Theischinger et Kalkman,

2014

Theischinger & Kalkman (2014b) Misool.

Nososticta xanthe Lieftinck, 1938

Kalkman & Orr (2013) Batanta, Salawati; Theischinger &

Kalkman (2014b) Misool, Salawati.

Palaiargia charmosyna Lieftinck, 1932

Kovacs et al. (2016; 2021) Batanta.

Palaiargia melidora Lieftinck, 1953

Kalkman & Orr (2013) Waigeo.

Palaiargia micropsitta Lieftinck, 1957

Michalski (2012) Misool; Kalkman & Orr (2013) Misool.

New record: Gam* (12).

Palaiargia nasiterna Lieftinck, 1938

Kalkman & Orr (2013) Waigeo.

Palaiargia susannae Kovacs et Theischinger,

2015

Kovacs et al. (2015b; 2021) Batanta.

Platystictidae Kennedy, 1920

Drepanosticta auriculata Selys, 1878

Kovacs et al. (2015b; 2016; 2021) Batanta.

Drepanosticta batanta Kovacs et Theischinger,

2015

Kovacs et al. (2015b; 2016; 2021) Batanta.

226

Drepanosticta inconspicua Lieftinck, 1938

Kalkman & Orr (2013) Waigeo.

Drepanosticta misoolensis van Tol, 2007

van Tol (2007) Misool; Michalski (2012) Misool.

New records: Hakau 1 (44 & 19), Ifeylo (1¢ (teneral)),

Vanlal (19).

Note: The identification of females is tentative

since one female is a teneral specimen and in bad

condition, the second is in normal condition. The

original species’ description was based on a single

teneral specimen (female) with last abdominal

segments damaged and distorted (van Tol 2007).

Anisoptera Selys, 1854

Aeshnidae Leach, 1815

Agyrtacantha dirupta (Karsch, 1889)

Orr & Kalkman (2015) Misool; Kovacs et al. (2021)

Batanta.

New records: Hakau 1 (19), Ifeylo (10).

Note: The female was flying ~0.4 m above a partly

dried-up riverbed in dusk. The identification of the

female is tentative, since no female descriptions

are available for any species of this genus.

Agyrtacantha microstigma (Selys, 1878)

Orr & Kalkman (2015) Waigeo.

New record: Vanlal(1<'). One-two specimens observed

patrolling over a small river (Stream) in a shady place.

The stream is fast-flowing (rhithral), but soecimens were

observed in the slower flowing section of it.

First record for Misool.

Anax fumosus Hagen 1867

Orr & Kalkman (2015) Batanta, Salawati?.

Anax maclachlani Forster, 1898

Orr & Kalkman (2015) Misool; Kovacs et al. (2021)

Batanta.

Gynacantha calliope Lieftinck, 1953

Orr & Kalkman (2015) Waigeo.

Gynacantha kirbyi Kriger, 1899

Michalski (2012) Misool; Orr & Kalkman (2015) Misool,

Waigeo.

Gynacantha mocsaryi Forster, 1898

Orr & Kalkman (2015) Batanta, Salawati, Waigeo;

Kovacs et al. (2015b; 2016; 2021) Batanta.

02-Jun-21 21:49:43

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pp. 221-238

Figures 6-9. Odonata from Raja Ampat. 6 - Orthetrum villosovittatum, ¢ in Aduwey, Misool. The single observed specimen

was resting on the ground in a sunny place between bushes in a mosaic landscape; 7 - Prothorthemis coronata, 9° in

Aduwey, Misool Island. Adult individuals were resting or flying in shadowy places along a small river in a primary

lowland rainforest; 8 - Place of oviposition of Prothorthemis coronata - a small hole in a tree trunk filled with water

and some dead leaves; 9 - Rhyothemis phyllis chloe, ° from Waigeo Island.

227,

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Gynacantha rosenbergi Brauer, 1867

Kovacs et al. (2021) Batanta.

Gomphidae Rambur, 1842

Ictinogomphus lieftincki (Schmidt, 1934)

Orr & Kalkman (2015) Batanta, Salawati.

Synthemistidae Tillyard, 1911

Palaeosynthemis cervula (Lieftinck, 1938)

Michalski (2012) Misool; Kovacs et al. (2021) Batanta,

as Palaeosynthemis cf. cervula.

New record: Vanlal (146 & 19).

N ote: The Misool record of this species was omitted

by Orr & Kalkman (2015).

Macromiidae Needham, 1903

Macromia euphrosyne Lieftinck, 1952

Orr & Kalkman (2015) Waigeo; Kovacs et al. (2015b;

2021) Batanta.

Macromia hermione Lieftinck, 1952

Michalski (2012) Misool; Orr & Kalkman (2015) Misool.

Macromia sophrosyne Lieftinck, 1952

Orr & Kalkman (2015) Waigeo.

Libellulidae Leach, 1815

Agrionoptera insignis (Rambur, 1842)

Orr & Kalkman (2015) Batanta, Salawati; Kovacs et al.

(2015b; 2021) Batanta.

New record: Hakau 2 (19).

First record for Misool.

Note: The specimen corresponds to the subspecies

Agrionoptera insignis papuensis, although

according to Michalski (2012) the status of this

Subspecies is not clear.

Agrionoptera longitudinalis Selys, 1878

Orr & Kalkman (2015) Batanta, Salawati, Waigeo;

Kovacs et al. (2015b; 2016; 2021) Batanta.

New records: Hakau 2 (10), Ifeylo (24 & 19), Vanlal

(19).

First record for Misool.

Notes: All Soecimens correspond to the subspecies

Agrionoptera longitudinalis biserialis. ln one case,

a female was ovipositing in a small (~LOx10x10

cm) hole filled with water and some dead leaves in

a stone. The stone was located in a shaded place

228

on the bank of a small forest river (Fig. 3).

Brachydiplax duivenbodei (Brauer, 1866)

Orr & Kalkman (2015) Batanta, Salawati; Kovacs et al.

(2015b; 2016; 2021) Batanta.

Camacinia gigantea (Brauer, 1867)

Orr & Kalkman (2015) Waigeo; Kovacs et al. (2015b;

2021) Batanta.

New records: Hakau 2 (10), Gam (1).

First record for Misool.

Notes: Two specimens - one in the gardens of

a river valley, one near a village - were resting in

sunny places between bushes (Fig. 4). The image

of C. gigantea (two specimens, image of one male)

from Misool Island available on the Insect-Trade.

EU website (https://www.insect-trade.eu/?catego

ry=odonata&eshop=catalogue&page=4, accessed

ZEAL 2ZO20).

Diplacina cyrene Lieftinck, 1953

Michalski (2012) Misool; Orr & Kalkman (2015) Misool.

Diplacina erigone Lieftinck, 1954

Michalski (2012) Misool; Orr & Kalkman (2015) Misool.

Diplacina cf. ismene Lieftinck, 1933

Kovacs et al. (2016) Batanta.

Diplacina micans Lieftinck, 1953

Orr & Kalkman (2015) Waigeo.

Diplacina olahi Theischinger et Kovacs, 2015

Kovacs et al. (2015b; 2016) Batanta.

Diplacina paula Ris, 1919

Michalski (2012) Misool, as Diplacina paula lethe; Orr &

Kalkman (2015) Misool.

Diplacina smaragdina Selys, 1878

Note: As stated by Kovacs et al. (2021), all records

of Diplacina olahi by Kovacs et al. (2016) in fact

refer to Diplacina smaragdina.

Diplacodes trivialis (Rambur, 1842)

Kovacs et al. (2015b; 2016; 2021) Batanta.

New record: Waisai 3 (49).

First record for Waigeo.

Huonia epinephela Forster, 1903

Kovacs et al. (2015b; 2016; 2021) Batanta.

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KaLNIns, M.: Contribution to the knowledge of dragonflies (Odonata) from the Raja Ampat (Indonesia), with notes ...

pp. 221-238

ill)

cll

Figures 10-11. The habitat of Agriocnemis femina in Aduwey village, Misool Island, Raja Ampat. 10 - Small ditches along

the foothpath, with dense, partially submerged aquatic vegetation (image courtesy Dmitry Telnov); 11 - Ditches near the

backyard garden during high tide, with seawater overflowing the ditches.

229

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Huonia silvicola Lieftinck, 1942

Michalski (2012) Misool; Orr & Kalkman (2015) Misool.

Huonia thais Lieftinck, 1953

Michalski (2012) Misool; Orr & Kalkman (2015) Misool;

Kovacs et al. (2015b; 2021) Batanta.

Huonia thisbe Lieftinck, 1953

Michalski (2012) Misool; Orr & Kalkman (2015) Misool.

Hydrobasileus brevistylus (Brauer, 1865)

New record: Hakau 2 (1¢ & 19).

First record for Misool and Raja Ampat Islands.

Note: Specimens observed in copula near a small

pool in a river valley (Fig. 5).

Hydrobasileus vittatus Kirby, 1889

Orr & Kalkman (2015) Batanta; Kovacs et al. (2015b;

2021) Batanta.

Lyriothemis hirundo Ris, 1913

Orr & Kalkman (2015) Batanta, Salawati.

Lyriothemis meyeri (Selys, 1878)

Orr & Kalkman (2015) Batanta, Salawati, Waigeo.

Nannophlebia amnosia Lieftinck, 1955

Michalski (2012) Misool; Orr & Kalkman (2015) Misool;

Kovacs et al. (2015b; 2016; 2021) Batanta.

Nannophlebia amphycillis Lieftinck, 1933

Kovacs et al. (2021) Batanta.

Note: Records of Nannophlebia amnosia by Kovacs

et al. (2016) now refer to Nannophlebia amphycillis

(see Kovacs et al. 2021).

Nannophya pygmaea Rambur, 1942

Kovacs et al. (2016; 2021) Batanta, as Nannophya cf.

pygmaea.

New record: Aduwey (1).

First record for Misool.

Nesoxenia mysis (Selys, 1878)

Michalski (2012) Misool; Orr & Kalkman (2015) Misool,

Waigeo; Kovacs et al. (2021) Batanta.

New records: Hakau 2 (19 (teneral)), lfeylo (2<),

Aduwey (13), Lilinta (54 & 32), Gam (23).

Note: All Soecimens correspond with the nominate

subspecies N. mysis mysis.

Neurothemis ramburii (Brauer, 1866)

Kovacs et al. (2015b; 2016; 2021) Batanta.

230

Neurothemis stigmatizans (Fabricius, 1775)

Michalski (2012) Salawati, subspecies manadensis; Orr

& Kalkman (2015) Batanta, Misool, Salawati, Waigeo;

Kovacs et al. (2015b; 2016; 2021) Batanta.

New records Neurothemis stigmatizans bramina:

Hakau 1 (1), Hakau 2 (1), Ifeylo (1), Aduwey (3),

Lilinta (14 (teneral) 22), Gam (23), Gam* (363), Waisai

1 (13).

New records Neurothemis stigmatizans manadensis:

Hakau 2 (14), Polee (14 & 19).

Notes: According to Michalski (2012), N.

stigmatizans manadensis is only known so far from

western New Guinea and Maluku. The female from

Polee was a heterochrome (hyaline) form. According

to Michalski (2012), only homeochrome form

females are known for this subspecies. Neurothemis

stigmatizans manadensis (two females, an image

of one specimen) from Misool Island are available

on the Insect-Trade.EU website (https://www.

insect-trade.eu/?category=odonata&eshop=catalo

sue&page=3, accessed 27.11.2020). Although the

Subspecies have been described and illustrated

(Michalski 2012), relatively large differences in

colouring have been found in the material already

analysed. Therefore, in the present paper, not all of

these subspecies are recognised as separate taxa.

Orthetrum serapia Watson, 1984

Kovacs et al. (2015b; 2021) Batanta.

New records: Hakau 2 (1), Aduwey (2).

First record for Misool.

Orthetrum villosovittatum (Brauer, 1868)

Orr & Kalkman (2015) Batanta, Salawati; Kovacs et al.

(2015b; 2021) Batanta.

New records: Hakau 1 (2¢ & 19), Hakau 2 (4).

First record for Misool.

Note: One male patrolling over a small pool in

gardens in a river valley (Fig. 6).

Pantala flavescens (Fabricius, 1798)

Kovacs et al. (2021) Batanta.

New records: Hakau 2 (19), Aduwey (6¢ & 29).

First record for Misool.

Protorthemis coronata (Brauer, 1866)

Orr & Kalkman (2015) Batanta, Salawati, Waigeo;

Kovacs et al. (2015b; 2021) Batanta.

New records: Hakau 1 (29), Hakau 2 (4¢' & 19), Ifeylo

(23 & 19), Lilinta (1¢), Vanlal (10).

First record for Misool.

Notes: Common species in river valleys, but

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occasionally observed in villages. In two cases

females were observed ovipositing in small (~15

x 50 x 7 cm and ~10 x 15 x 5 cm) holes in tree

trunks filled with water and some dead leaves.

One of the tree trunks was located on a small

forest path, relatively far from watercourses, and

the second one was on the bank of a small forest

river, both in shaded places. In several cases

males were observed, patrolling or perching near

Shallow puddles in shaded parts of a forest (Figs

7-8). An image of a female P. coronata from

Waigeo Island was available on the Insect-Trade.

EU website (https://www.insect-trade.eu/?catego

ryv=odonata&eshop=catalogue&page=5, accessed

27.11.2020).

Raphismia bispina (Hagen, 1867)

Orr & Kalkman (2015) Misool; Kovacs et al. (2015b;

2016; 2021) Batanta.

Rhodothemis nigripes Lohmann, 1984

Kovacs et al. (2021) Batanta.

Rhyothemis phyllis Sulzer, 1776

Kovacs et al. (2015b; 2016; 2021) Batanta.

New records: Waisai 3 (19).

First record for Waigeo.

Notes: The Waisai specimens correspond to R.

phyllis chloe Kirby, 1894 (Fig. 9) so far known only

from Aru Islands and North Australia to Brisbane

(Theischinger & Hawking 2006; Michalski 2012).

An image of R. phyllis chloe (male) from Waigeo

Island was available on the Insect-Trade.EU

website __ (https://www.insect-trade.eu/?category

=odonata&eshop=catalogue&page=/, accessed

ZL 112020).

Rhyothemis regia (Brauer, 1867)

Kovacs et al. (2021) Batanta.

Rhyothemis resplendens Selys, 1878

Orr & Kalkman (2015) Batanta, Salawati; Kovacs et al.

(2016; 2021) Batanta.

Rhyothemis rita Kovacs et Theischinger, 2016

Kovacs et al. (2016) Batanta.

Tetrathemis irregularis Brauer, 1868

Orr & Kalkman (2015) Misool; Kovacs et al. (2021)

Batanta.

Tramea eurybia eurybia Selys, 1878

Orr & Kalkman (2015) Misool; Kovacs et al. (2015b;

pp. 221-238

2021) Batanta, reported as Tramea eurybia s. l.,

subspecies identification not provided.

Tramea transmarina propinqua Lieftinck, 1942

Kovacs et al. (2021) Batanta.

Zyxomma multinervorum Charpenter, 1897

Kovacs et al. (2021) Batanta.

Discussion

There are about 6000 species of dragonflies

known worldwide. Of these, about 620 species are

found in New Guinea, Maluku and the Solomon

Islands (Michalski 2012) and about 490 in New

Guinea itself (Orr & Kalkman 2015). As several new

species are described from this region each year,

the total number of species is already significantly

higher. A total of 103 species of dragonflies are

currently known from Raja Ampat, 50 Zygoptera

and 53 Anisoptera (Table 11).

Information about four species of dragonflies

was found on and insect trader website. According

to the information on the website, all insects are

accompanied with complete collecting data, often

including altitude, latitude, and longitude. As no

specimens were purchased, the exact locality

details are not known, but it is known that some

commercial specimens can have unreliable or

incomplete, or even incorrect data. However, such

specimens are also used in research, including

in this case; the information given on the website

does not contradict the data obtained from other

research.

Summarizing published and _— previously

unpublished data, 21 Zygoptera species and 27

Anisoptera species are currently known for the

fauna of Misool, for a total of 48 species. Although

the number of species has increased, the Misool

fauna can still only be considered as satisfactorily

studied, and it is likely that a relatively large number

of additional species remain to be discovered on

the island. This is Supported by the small number

of research expeditions and the limited time spent

on research, as well as the absence on Misool

of various species or groups of species found on

nearby islands. The current proportion of species

in the suborders, Zygoptera 52% and Anisoptera

48%, indicates an incomplete knowledge of the

Zygoptera fauna in Misool and Waigeo, too. For

example, in New Guinea - Zygoptera represent

62% and Anisoptera 38% of dragonfly diversity

(Kalkman & Orr 2013; Orr & Kalkman 2015).

aes

231i

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The small number of species found for each

observation site, confirms that the studies of all

islands are incomplete. For example, in a ~2.5

km? area in a lower montane forest of Papua New

Guinea, 61 species were found during an 8-9

month observation period; however, most species

occurred in small numbers (Oppel 2005). In

another study, which compared natural (~2.5 km?)

and modified (~4 km?) rainforest areas, 78 species

were found, with a very high ratio of Zygoptera vs

Anisoptera; 73-76% and 24-27% respectively

(Oppel 2006).

Table 1. The currently known dragonflies in Raja Ampat with notes on the level of endemism.

Legends: E - endemic species for the Papuan region in strict sense; E* - endemic species, recorded only from

one island; ? - species marked with "?" by previous authors).

Zygoptera

00,

ray

australis

varga

coartans

ochrostoma

Argiolestes

Metagrion

Podopteryx

Calopterygidae

Neurobasis

Chlorocyphidae

Rhinocypha

Coenagrionidae

Agriocnemis

Archibasis

Argiocnemis

Ceriagrion

Ischnura

Papuagrion

Pseudagrion

Teinobasis

Xiphiagrion

Isostictidae

Selysioneura

Tanymecosticta

Lestidae

io)

1°)

selysi

australis australis

australis misoolensis

| be

incta sagitta

incta tincta

[Riiiioeyeie |

femina

mimetes

rubescens

aeruginosum

senegalensis

magnanimum

civicum

starreanum

michalskii

prothoracica

rufithorax

superba

wallacei

cyanomelas

albicaudus

232

Guerin, 1832

Kovacs et Theischinger, 202

Lieftinck, 1956

Lieftinck, 1949

Forster, 1899

Selys, 1897

Lieftinck, 1955

Batanta & Salawati

Raja Ampat

Lieftinck, 1938

Rambur, 1842

Brauer, 1868

Tillyard, 1913

Selys, 1877

Brauer, 1869

Rambur, 1842

Selys, 1876

Lieftinck, 1932

Lieftinck, 1949

Theischinger et Kalkman, 2014.

Selys, 1877

(Hagen in Selys, 1877)

Campion, 1924

Selys, 1876

Lieftinck, 1953

Lieftinck, 1959

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pp. 221-238

Table 1 (continuation)

Idiocnemis _—_|dagnyae i Lieftinck, 1958 | Cf | || x |

jIdiocnemis _ifissidens——[Lieftinck, 1958 tee | ||| x

Idiocnemis _ijinornata sels 1878 CT | Ct eT x

Idiocnemis _—i|strumidens—|Lieftinck, 1958 | Cf | UT | Cx |

|Nososticta i fatrocyana ——[Lieftinck, 1960, tex | | | | x

[Nososticta _—‘[dora_———SSCS*«dKovoS et Theischinger 2006 [|| [ex| | x |

|Nososticta —_ferythroprocta ss [Selys, 18860 te | | CT | x

|Nososticta | nigrifrons (R893) | Ct | || x

[Nososticta—_—i(plagiata—“‘C;S*#*dS WS TBBGSCOCSC~*~sSC“‘<‘iPC*|~SC*T~SCdT~Cd Cx

[Nososticta__—|pyroprocta——=S~=siLiefinck, 7960———<C~*‘“‘TSCédT ir «*(| S| SCT Sid x |

[Nososticta__[rufipes——=SCS*«sTHeischinger et Kalkan, 2004 | Te | | | | x |

[Nososticta_—i[xanthe—SSCS~*~=*dL ein T9B™SCSC*~iESC‘iC ee Te | | = x

[Palaiargia____—_|charmosyna__——=s(Lieftinok, 7982_———=—C*é~SC<‘*tS*~‘iS:C‘i CWT Sid (Cx

[Palaiargia __—|melidora———S—=|Lieftinck, 1953 —SSCi em | | || Cd x

[Palaiargia___[micropsifta——=s(Lieftinck, 1967—SSC*TSsee* | | | Sd x

[Palaiargia___—_—‘[nasiterna———SSSC«LiencK, 038 | S| Cd] CT Cd x

[Drepanosticta_[auriculata——s—=~=~«‘i SWS TBSC=“‘COCSC#@CSSOCSOCdSSCOCdC | OC

[Drepanosticta_|batanta——=SCS*«Kovics et Theischinger 2075 [|| [ex| |x|

[Drepanosticta_|inconspicua—_—=i(Lieftinck, 938 ——S—C~s | S| | S| Cd x

[Drepanosticta[misoolensis [van ol, 2007 |e] [OT x

|Agyrtacantha—|dirupta, si (Karsch 1889) | tx | UT x | CT x |

Says fers) SSS Ses

jAnax si maclachlani_ (Forster, 1898 | tx | x | x

|Gynacantha—|mocsaryi si Forsten 1898 x | lx | x | x

(Schmick, 1934) ee ee

(Wefinek, 1938) SiS Ses

|Macromia [hermione Lieftinck, 1952, | te] | | x

(Rambur, 1842) Mckee en

Selys, 1878 Mths han

(Brauer, 1866) mrs yiaee

|Camacinia [gigantea s[(Brauern 867) xk tT x | Tl x | lx |

|Diplacina——cyrene i ieftinck 1953) | fe | CC

|Diplacina——siferigone—s[tieftinck, 1954 | fe | | | CT x |

|Diplacina——siismene Ss Lieftinck, 1933 | |e |

|Diplacina——|micans—siieftinck, 19530 fee | | CT | x |

Diplacina olahi Theischinger et Kovacs, 2015 Ex x

—

233

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Diplacina

Diplacodes

Huonia

Hydrobasileus

Lyriothemis

Nannophlebia

Nannophya

Nesoxenia

Neurothemis

Orthetrum

Pantala

Protorthemis

Raphismia

Rhodothemis

Rhyothemis

Tetrathemis

Tramea

paula

smaragdina

trivialis

epinephela

Silvicola

thais

thisbe

brevistylus

vittatus

hirundo

meyeri

amnosia

amphicyllis

pygmaea

mysis

ramburii

stigmatizans

serapia

villosovittatum

flavescens

coronata

bispina

nigripes

phyllis

regia

resplendens

rita

irregularis

eurybia eurybia

Kirby,

Ris, 1919

Selys, 1878

(Rambur, 1842)

Forster, 1903

Lieftinck, 1942

Lieftinck, 1953

(Brauer, 1865)

1889

Ris, 1913

(Selys, 1878)

Lieftinck, 1955

Lieftinck, 1933

Rambur, 1942

(Selys, 1878)

(Brauer, 1866)

(Fabricius, 1775)

Watson, 1984

(Brauer, 1868)

(Fabricius, 1798)

(Brauer, 1866)

(Hagen, 1867)

Lohmann, 1984

Sulzer, 1776

(Brauer, 1867)

Selys, 1878

Kovacs et Theischinger 2016

Brauer, 1868

Selys, 1878

Cettinok, 1942 Lees

Table 1

—

continuation)

X X

[Zaomma____|muftinervorum [Carpenter 7897 | | | fe | | x

Totalnumberofspecies = —C—‘“‘;é‘—~*CSC*ir 8 a8 | 22 | 2 | Op | 203

Of all Raja Ampat Odonata species, 62 species

(60%) are endemic to the Papuan Region in a

Strict sense. Twenty-three of these species can be

considered as local endemics, as these species are

known from only one of the Raja Ampat islands, and

there are no records from other islands, including

New Guinea (Table 1). It is possible that as the level

of research in the region increases, some of the

locally endemic species may be found elsewhere

over time. However, given the relatively small

ranges of most species in this area, these species

are likely to remain endemic to the Papuan region

in a strict sense.

As a result of comprehensive research

(Polhemus et al. 2004) 40 areas of freshwater

endemism have been defined on New Guinea and

nearby islands, grouped into 6 broad regions. The

Raja Ampat Archipelago is one of the above regions

234

of broad endemism. Waigeo, Batanta, and Misool

are defined as separate areas of endemism. At

the same time, there is no strong justification for

the division of Salawati, with the southern part of

Salawati being included in the Vogelkop Lowlands

area, while the northern part (Wagon Mountains)

could potentially be included in the Batanta area.

Kofiau is not included in any area.

The same study mentions taxa defining areas

of endemism. Although it is not clearly defined

why only Zygoptera species have been used to

characterize areas of endemism, this may be

explained by the fact that Anisoptera as a whole

have better dispersal abilities. In the light of current

knowledge on the distribution of Odonata species

in the Papuan Region, clarifications are proposed

to the list of taxa defining areas of endemism,

including Anisoptera species (Table 2).

02-Jun-21 21:49:50

KaLNins, M.: Contribution to the knowledge of dragonflies (Odonata) from the Raja Ampat (Indonesia), with notes ...

pp. 221-238

Table 2. Proposals for the list of taxa defining areas of endemism.

Legends: Species are listed in alphabetical order, first mentioning Zygoptera, and then Anisoptera. The left

column of the table shows the species names used by Polhemus et al. (2004), and the right column shows

current species names.

Taxa defining areas of endemism as proposed by

Proposals for updating the list of taxa defining areas

Polhemus et al. (2004 of endemism

Metagrion coartans

Metagrion ochrostoma

Idiocnemis fissidens

Nososticta atrocyana

Nososticta erythroprocta

Nososticta evelynae

Palaiargia nasisterna [sic! nasiterna] Exclude from list. Palaiargia nasiterna also reported

from West New Guinea

pf Ptlaciargia meliciora

Teinobasis prothoracica Exclude from list. Species also reported from West New

Guinea and Misool

Gynacantha calliope

| Macromia sophrosyne

Diplacina micans

Batanta area (Salawati not included

Not suggested

COOCOCOCOCOCOCOC‘“‘(‘(CN_*AiOTe'STE’S WEE

J Brepanosticta batanita

Nos osticta dora

Palani susannae

ip iacina fai

FR ythemis rita

‘Argiolestes pyroprocta' The name ‘Argiolestes pyroprocta’ was never published

and is nomen nudum

CCSCSCSCSCSCSCSSS=C Drepanostcta misoolensis

Nososticta pyroprocta Exclude from list. Species also reported from West New

Guinea

Palaiargia micropsitta Palaiargia micropsitta

Teinobasis michalskii

Macromia hermione

Vogelkop Lowlands area

(Di

Jian paula

| t=‘! VogeilkopLowilandsarea—i“(wstsd|CCC™C~i“‘TCTOC#(U ;Sailawatiarea” st —CisCisz

Argiolestes connectens Metagrion connectens not yet found in Raja Ampat

Huonia thisbe

Argiolestes fontinalis sic! fontinale Metagrion fontinale not yet found in Raja Ampat

Book4.indd 235

Acknowledgements

| am greatly indebted to Dr. Dmitry Telnov

(The Entomological Society of Latvia, Riga) for

valuable suggestions for the preparation of the

final version of the manuscript, for the material and

data provided, as well as to Laszlo Wagner (http://

www.east-indonesia.info) for guidance and help in

expeditions, and to Prof. Gunther Theischinger for

reviewing the manuscript. | want to pass my friendly

thanks to the particularly hospitable people in the

villages of Aduwey, Lilinta, and Gamta.

References

Anonymous 2004. Pub. 164, Sailing Directions (Enroute)

New Guinea, Ninth Edition. National Geospatial-

Intelligence Agency, Bethesda, Maryland: 263 pp.

ESRI Data & Maps 2006. ArcG/S 9 ESRI Data & Maps

Media Kit (5-CD-disk set). Redlands, U.S.A.

Gassmann D. 1999. Taxonomy and distribution of the

inornata species-group of the Papuan genus

Idiocnemis Selys, 1878 (Odonata: Zygoptera:

Platycnemididae). - Invertebrate Taxonomy 13:

977-1005.

Gassmann D. 2000. Revision of the Papuan Idiocnemis

235

02-Jun-21 21:49:50

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

bidentata-group (Odonata: Platycnemididae). -

Zoodlogische Mededelingen 74, No 23: 375-402.

Greke K., Telnov D. 2014. Review and assessment of

the literature on recent non-marine molluscs of

the Papuan biogeographical region: 61-112, pls

In: Telnov D. (ed.) Biodiversity, biogeography and

nature conservation in Wallacea and New Guinea.

Volume Il. The Entomological Society of Latvia,

Riga: 458 pp, 126 pls.

Kalkman V. J., Orr A. G. 2013. Field Guide to the

damselflies of New Guinea. - Brachytron 16

Supplement: 3-120.

Kalnins M. 2014. Argiolestes zane sp. nov. from New

Guinea (Odonata: Argiolestidae): 221-224, pls 31-

34. In: Telnov D. (ed.) Biodiversity, biogeography

and nature conservation in Wallacea and New

Guinea. Volume Il. The Entomological Society of

Latvia, Riga: 458 pp, 126 pls.

Kalnins M. 2017. Argiolestes spungisi sp. nov. (Odonata:

Argiolestidae) from New Guinea: 357-362, pls

52-55). In: Telnov D., Barclay M. V. L., Pauwels O.

S. G. (eds) Biodiversity, biogeography and nature

conservation in Wallacea and New Guinea. Volume

Ill. The Entomological Society of Latvia, Riga: 658

pp, 172 pls.

Kovacs T., Horvath R., Juhasz P. 2015a. Szitakot6k és

tegzesek (Insecta: Odonata, Trichoptera) kutatasa

Batanta szigeten (Indonézia, Nyugat-Papua). Study

of dragonflies and caddisflies (Insecta: Odonata,

Trichoptera) on Batanta Island (Indonesia, West

Papua). - Annales Musei _historico-naturalis

hungarici 107: 269-288.

Kovacs T., Theischinger G., Juhasz P., Danyik T. 2015b.

Odonata from Batanta (Indonesia, West Papua)

with description of three new species. - Folia

historico-naturalia Musei Matraensis 39: 17-29.

Kovacs T., Theischinger G., Danyik T. 2016. Odonata from

Batanta (Indonesia, West Papua) with description

of two new species. - Folia historico-naturalia

Musei Matraensis 40: 27-37.

Kovacs T., Theischinger G., Horvath R., Juhasz P. 2021.

Odonata from Batanta (Indonesia, West Papua)

with description of one new species. - Opuscula

zoologica 52 (in press).

Lieftinck M. A. 1949. The dragonflies (Odonata) of New

Guinea and neighbouring islands. Part VII. Results

of the Third Archbold expedition 1938-1939 and

of the Le Roux Expedition 1939 to Netherlands

New Guinea (Il. Zygoptera). - Nova Guinea (New

Series) 5: 1-271.

Lieftinck M. A. 1971. Studies in Oriental Corduliidae

(Odonata), |. - Tijdschrift voor Entomologie 114:

1-63.

Michalski J. 2012. A Manual for the Identification of the

Dragonflies & Damselflies of New Guinea, Maluku

& the Solomon Islands. Kanduanum_ Books,

Morristown, New Jersey: 561 pp.

236

Oppel S. 2005. Habitat associations of an Odonata

community in a lower montane rainforest in Papua

New Guinea. - International Journal of Odonatology

8, No 2: 243-257.

Oppel S. 2006. Comparison of two Odonata communities

from a natural and a modified rainforest in Papua

New Guinea. - International Journal of Odonatology

9, No 1: 89-102.

Orr A. G, Kalkman V. J. 2015. Field Guide to the

dragonflies of New Guinea. - Brachytron 17,

Supplement: 3-156.

Polhemus D. A., Englund R. A., Allen G. R. 2004.

Freshwater biotas of New Guinea and nearby

islands: analysis of endemism, richness, and

threats. - Bishop Museum Technical Report 31: 62

pp.

Riedel A. 2002. Taxonomy, phylogeny, and zoogeogra-

phy of the weevil genus Euops (Insecta: Coleoptera:

Curculionoidea) in the Papuan region. Disserta-

tion zur Erlangung des Doktorgrades der Fakultat

fur Biologie der Ludwig-Maximilians-Universitat

Munchen: 216 pp.

Ris F. 1913. Die Odonata von Dr H.A. Lorentz’ Expedition

nach Sudwest-Neuguinea 1909 und einige

Odonaten von Waigéu. - Nova Guinea 3: 476-478.

Theischinger G., Hawking J. 2006. The complete field

guide to dragonflies of Australia. CSIRO Publishing,

Collingwood: 366 pp.

Theischinger G., Kalkman V. J. 2014a. The genus

Teinobasis on the Bird’s Head Peninsula and

the Raja Ampat Islands, Indonesia (Odonata:

Coenagrionidae). - Odonatologica 43, No 3/4:

143-168.

Theischinger G., Kalkman V. J. 2014b. Nososticta rufipes

sp. nov. from Misool, with notes on the other

Nososticta species on the Bird’s Head Peninsula

and the Raja Ampat, Indonesia (Zygoptera:

Platycnemididae). - International Journal of

Odonatology 17, No 4: 217-222.

The Times Atlas of the World 1994. 4" Revised Concise

Edition. London.

van Tol J. 2007. The Platystictidae of the Moluccas and

Misool (Odonata). - Deutsche Entomologische

Zeitschrift 54: 3-26.

Wikramanayake E., Dinerstein E., Loucks C. J., Olson

D. M., Morrison J., Lamoreux J., McKnight M.,

Hedao P. 2002. Terrestrial Ecoregions of the Indo-

Pacific: A Conservation Assessment. Island Press,

Washington, DC: 643 pp.

Received: 06.xii.2020.

Accepted: 19.xii.2020.

02-Jun-21 21:49:51

KaLNiIns, M.: Contribution to the knowledge of dragonflies (Odonata) from the Raja Ampat (Indonesia), with notes ...

pp. 221-238

_Batanta: 3,24

Wp i Gy

Be Fig A A

i aie ae

ee ae

Hakau 2 Aduwe

Map 1. The Raja Ampat Islands and localities mentioned in the text on the imagery and shaded relief map (ESRI

Data & Maps 2006) (prepared with ArcGIS 10.3).

Misool

Map 2. Misool and its adjacent islands and localities mentioned in the text on the imagery and shaded relief map

(ESRI Data & Maps 2006) (prepared with ArcGIS 10.3).

237

Book4.indd 237

02-Jun-21 21:49:55

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Cablesego) =

Map 3. Waigeo and its adjacent islands and localities mentioned in the text on the imagery and shaded relief map

(ESRI Data & Maps, 2006) (prepared with ArcGIS 10.3).

Wallace's line | — | ag

i Weber's line je™ J?

Map 4. The Papuan region in strict and broad senses and the surrounding regions, showing important

zoogeographic lines (redrawn from the Times Atlas of the World (1994), with modifications by Riedel (2002)).

238

Book4.indd 238 02-Jun-21 21:49:57

Book4.indd 239

Katnins, M.: Contribution to the knowledge of true bugs (Hemiptera: Heteroptera) from Wallacea and New Guinea ...

pp. 239-244

Contribution to the knowledge of aquatic bugs

(Hemiptera: Heteroptera) from Wallacea

and New Guinea, with ecological notes

urn:lsid:zoobank.org: pub: DED53FD6-4231-45FF-B188-4D31BCO2A/7EB

MARTINS KALNINS * 2

1 - Department of Biosystematics, Institute of Life Sciences and Technology, Daugavpils

University, Vienibas iela 13, LV-5401, Daugavpils, Latvia; martins.kalnins@biology.|v

2 — Joint Stock Company “Latvia's State forests”, Vainodes iela 1, LV-1004, Riga, Latvia

Abstract: The present paper contains information on 18 taxa, including 13 species and 5 genus-rank identifications

of Heteroptera from the Wallacea region and New Guinea based on personal observations and museum material.

When possible, supplementary information on habitat is given, and morphological variation in Ranatra_ biroi

Lundblad, 1933 is discussed.

Key words: Heteroptera, aquatic bugs, faunistics, distribution, ecology, morphology.

Introduction

Over time, the author has accumulated

observations on Heteroptera Latreille, 1810

from Wallacea and adjacent areas. The material

discussed in the present paper is not the result

of extensive Heteroptera-targeted research, but,

considering the present relatively low knowledge

level of research in the region, the author believes

that publishing even a relatively small amount of

precise data and sharing them with the scientific

community is welcome. As demonstrated for some

other invertebrates, small amounts of random data

form a larger dataset over time and contribute well

to regional studies and the general knowledge of

the taxonomic group.

Material and methods

Data on aquatic bugs were obtained by

processing material from the author’s and museum

collections of Heteroptera. The origin of the studied

material is New Guinea (Doberai Peninsula: Aitinyo;

Bomberai Peninsula: Fak-Fak; Bird’s Neck Isthmus:

Kaimana, Kamakawalar, Oray; mainland Papua:

Oksibil, Setani), Misool in Raja Ampat Islands

(Aduwey) and Halmahera (Lina) in North Moluccas

(Map 1). Specimens were identified by Dan A.

Polhemus in 2011-2020, except for Lethocerus

insulanus identified by Dmitry Telnov in 2010.

Aquatic bugs were mainly sampled with

a hydrobiological net in the littoral zone of

various water bodies. The only exception is the

Lethocerus insulanus attracted to white light. The

material discussed is mainly deposited at the

Naturkundemuseum Erfurt, Germany.

The term “Papuan Region” (or in the strict

zoogeographic sense - the Papuan subregion of

the Australian Region) is often listed in zoological or

zoogeographic literature, but sometimes extended

to include some adjacent areas. Therefore, for the

purposes of this paper, the terms used by Greke &

Telnov (2014) - Papua sensu lato and Papua sensu

Stricto - are used (Map 2).

Results

Heteroptera Latreille, 1810

Belostomatidae Leach, 1815

Diplonychus eques (Dufour, 1863) (Fig. 1)

Studied material: 1 (immature) INDONESIA, West

Papua Prov., Kaimana 4/7 km E, Triton Bay, Kamaka

(former Warika) village environs, Lake Kamakawalar

and surroundings, 3°46’22”S, 134°12’02”E, 60-310

m, little ditch in primary lowland forest on limestone,

08.09.2010; 3 specimens INDONESIA E, West New

Guinea, Doberai Peninsula, Ayamaru vill. 27 km SE, Aitinyo

(Uter) lakeside, 1°26’01”S, 132° 23'10”E, ~175-177 m,

from aquatic vegetation, 04.09.2015; 29 INDONESIA

E, New Guinea, Papua Prov., Star Mountains, Oksibil

239

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

23 km SSE, Beringin vill., 05°05’01”S, 140°43'27’E,

310 m, edge of secondary lowland rainforest, MV light,

T2OS*201.8.

Note: Widespread in Australia and New Guinea.

May be conspecific with the very widespread

D. rusticus (Fabricius 1781) (India to Australia)

(Andersen & Weir 2004).

Lethocerus insulanus (Montandon, 1898)

Studied material: 14 INDONESIA E, Prov. Raja Ampat,

Misool SW, Aduwey (Aduai) vill. ~2 km NNW, valley of

River Hakau, 01°58’21’"S, 129°54’30’E, in lowland

rain forest, on UV light, 25.03.2009.

Note: The species inhabits relatively wide areas

of Australia and also recorded from surroundings

of Port Moresby in Papua New Guinea (Insecta in

GBIF Secretariat 2019), Maprik, E Sepik province

(Buzzetti et al. 2006), from New Caledonia

(Damgaard & Zettel 2014) and _ Philippines

(Andersen & Weir 2004). First record for the fauna

of Misool and Raja Ampat Islands.

Gerridae Leach, 1815

Limnogonus darthulus (Kirkaldy, 1901)

Studied material: 1¢ & 12 INDONESIA, West

Papua Prov., Kaimana 47 km E, Triton Bay, Kamaka

(former Warika) village environs, Lake Kamakawalar

and surroundings, 3°46’22”S, 134°12’02”E, 60-310

m, little ditch in primary lowland forest on limestone,

08.09.2010.

Limnogonus fossarum skusei (Torre-Bueno, 1926)

Studied material: 34 INDONESIA, West Papua Prov.,

Kaimana 47 km E, Triton Bay, Kamaka (former Warika)

village environs, Lake Kamakawalar and surroundings,

3°46’22"S, 134°12’02”E, 60-310 m, little ditch in

primary lowland forest on limestone, 08.09.2010.

Limnometra kallisto (Kirkaldy, 1899)

Studied material: 73 & 139 INDONESIA, West

Papua Prov., Kaimana 4/7 km E, Triton Bay, Kamaka

(former Warika) village environs, Lake Kamakawalar,

3°45'33"S, 134°12’05’E, 90 m, 2nd pool in primary

lowland rainforest, 09.09.2010; 2¢ & 49 INDONESIA,

West Papua Prov., Kaimana 7-9 km NW, sago swamp,

32 35:02"S 41'33°49' 58°F, 05.09!2010; 2ar & AY

Figure 1. Diplonychus eques (Dufour, 1863), specimen from Lake Aitinyo, Doberai Peninsula, New Guinea, dorsal

view (image courtesy Dmitry Telnov, Riga, Latvia).

240

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KaLnins, M.: Contribution to the knowledge of true bugs (Hemiptera: Heteroptera) from Wallacea and New Guinea ...

INDONESIA, West Papua Prov., Kaimana 4/7 km E,

Triton Bay, Kamaka (former Warika) village environs,

Lake Kamakawalar and surroundings, 3°46’22’S,

134°12’02”E, 60-310 m, little ditch in primary lowland

forest on limestone, 08.09.2010.

Limnometra lipovskyi Hungerford et Matsuda,

1958

Studied material: 1¢ & 209 INDONESIA, West

Papua Prov., Kaimana 30-40 km E, Triton Bay, River

Lengguru valley upriver from Oray village, 3°43’26’S,

134°06’06’”E, ~150 m, limestone creek, 13.09.2010;

24 INDONESIA, West Papua Prov., Kaimana 47 km E,

Triton Bay, Kamaka (former Warika) village environs,

Lake Kamakawalar and surroundings, 3°46’22’S,

134°12’02”E, 60-310 m, little ditch in primary lowland

forest on limestone, 08.09.2010.

Tenagogonus sp.

Studied material: 14 (brachypterous) INDONESIA,

West Papua Prov., Kaimana 30-40 km E, Triton Bay, River

Lengguru valley upriver from Oray village, 3°43’26’S,

134°06'06’E, ~150 m, limestone creek, 13.09.2010.

Hydrometridae Billberg, 1820

Hydrometra sp.

Studied material: 3 specimens INDONESIA E, W New

Guinea, Doberai Peninsula, Ayamaru vill. 27 km SE,

Aitinyo (Uter) lakeside, 1°26’01”S, 132° 23'10’E, ~175-

177 m, from aquatic vegetation, 04.09.2015.

Nepidae Latreille, 1802

Ranatra biroi Lundblad, 1933 (Figs 2-8)

Studied material: 14 INDONESIA E, West New

Guinea, Doberai Peninsula, Ayamaru vill. 27 km SE,

Aitinyo (Uter) lakeside, 1°26’01”S, 132° 23710’E, ~175-

177 m, from aquatic vegetation, 04.09.2015.

Notes: First record from Doberai Peninsula,

hitherto the westernmost record for this species.

The studied specimen of this transpapuan species

is slightly different in external morphology from

what is known from the literature (Lundblad

1933; Lansbury 1972). Frons and vertex convex,

distinctly raised over compound eye in lateral view

(Fig. 3). Compound eyes comparatively small (Figs

2-4), not as in Lansbury (1972, figs 200-201).

Prothorax on either side of the flat median carina

is feebly longitudinally emarginate, margins of the

emargination not carinate (Fig. 4). Mesoventrite

is either subtrapezoid (Fig. 5), not circular.

Metaventrite slightly convex medially (in lateral

aspect), not “raised” as in Lansbury (1972). The

variability considered not sufficient for delimiting

pp. 239-244

a new subspecific taxon, since shape of male

genitalia in studied specimen (Figs 6-8) agrees

with those in typical R. biroi.

Ranatra megalops Lansbury, 1972

Studied material: 34 & 22 INDONESIA, West

Papua Prov., Kaimana 30-40 km E, Triton Bay, River

Lengguru valley upriver from Oray village, 3°43’26’S,

134°06’06”E, ~150 m, ditch in the primary forest on

limestone and sago swamp, 13.09.2010.

Notonectidae Leach, 1815

Anisops sp.

Studied material: several specimens INDONESIA,

West Papua Prov., Kaimana 47 km E, Triton Bay, Kamaka

(former Warika) village environs, Lake Kamakawalar,

3°45'33"S, 134°12’05’E, 90 m, 2nd pool in primary

lowland rainforest, 09.09.2010.

Anisops stali Kirkaldy, 1904

Studied material: 12 INDONESIA, West Papua Prov.,

Kaimana 47 km E, Triton Bay, Kamaka (former Warika)

village environs, Lake Kamakawalar and surroundings,

3°46’22"S, 134°12’02”E, 60-310 m, little ditch in

primary lowland forest on limestone, 08.09.2010.

Enithares sp.

Studied material: several specimens INDONESIA,

West Papua Prov., Kaimana 47 km E, Triton Bay, Kamaka

(former Warika) village environs, Lake Kamakawalar,

3°45'33"S, 134°12’05’E, 90 m, 2nd pool in primary

lowland rainforest, 09.09.2010.

Enithares bakeri Brooks, 1948

Studied material: 14 INDONESIA E, North Moluccas,

Halmahera N, Lake Lina, SW lakeside, 1°31’38’N,

127°50'15’”E, 205 m, lake & sago swamp, 04.07.2013.

Note: A species distributed in the Philippines and

on several Indonesian islands such as Sulawesi

etc. (Insecta in GBIF Secretariat 2019). According

to Polhemus (2020) this is perhaps the most

widespread of any Enithares species occurring on

the insular systems of Southeast Asia.

Enithares elongata Lansbury, 1974

Studied material: 24 INDONESIA E, New Guinea,

Papua Prov., Sentani, Cyclops Mts., 02°32’04’S,

140°30’47”E, 550 m, perennial stream, headwater,

03.04.2018.

Enithares megalops Lansbury, 1968

Studied material: 1 INDONESIA, West Papua

Prov., Onin Peninsula, Fak-Fak 5-7 km N, 2°53’26’S,

132°18’22”E, 300-400 m, creek, 23.09.2010; 10 &

241

ari

02-Jun-21 21:49:59

Book4.indd 242

242

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

32 INDONESIA, West Papua Prov., Kaimana 30-40 km

E, Triton Bay, River Lengguru valley upriver from Oray

village, 3°43’26"S, 134°06’06’”E, ~150 m, limestone

creek, 13.09.2010; 3 specimens INDONESIA, West

Papua Prov., Kaimana 4/7 km E, Triton Bay, Kamaka

(former Warika) village environs, Lake Kamakawalar

and surroundings, 3°46’22”S, 134°12’02”E, 60-310

m, little ditch in primary lowland forest on limestone,

08.09.2010.

Veliidae Amyot et Serville, 1843

Rhagovelia transbintuni D. Polhemus et J.

Polhemus, 2011

Studied material: 2¢ & 22 INDONESIA, West

Papua Prov., Kaimana 30-40 km E, Triton Bay, River

Lengguru valley upriver from Oray village, 3°43’26’S,

134°06’06’E, ~150 m, limestone creek, 13.09.2010.

Figures 2-8. Ranatra biroi Lundblad, 1933, 4 from Lake Aitinyo, Doberai Peninsula, New Guinea. 2 - Forebody,

dorsal view; 3 - Head and anterior part of thorax, lateral view; 4 - Thorax, ventral view; 5 - Meso- and metaventrite,

ventral view; 6-8 - Male accessory genitalia and paramere (8), different views [not to scale] (images courtesy

Dmitry Telnov, Riga, Latvia).

02-Jun-21 21:49:59

Book4.indd 243

KaLnins, M.: Contribution to the knowledge of true bugs (Hemiptera: Heteroptera) from Wallacea and New Guinea ...

Strongylovelia sp.

Studied material: 4 specimens INDONESIA, West

Papua Prov., Kaimana 30-40 km E, Triton Bay, River

Lengguru valley upriver from Oray village, 3°43’26’S,

134°06’06’E, ~150 m, limestone creek, 13.09.2010.

Discussion

The present paper discusses new distributional

data on 18 taxa, including 13 species and 5 genus-

rank identifications of Heteroptera. According

to the GBIF’s species datasets (Insecta in GBIF

Secretariat 2019) and the most recent available

publications on particular taxonomic groups

(Andersen 1975; Polhemus & Polhemus 1998;

2011; Polhemus 2020), several of the discussed

species are not yet reported outside the Papuan

Region and are considered Papuan _ regional

endemics in a_ strict (Enithares elongata, E.

megalops, Limnogonus darthulus, Ranatra_ biroi,

R. megalops, and Rhagovelia transbintuni) or a

broad sense (Limnometra kallisto) (consider map

2 for zoogeographic definitions). However, none of

these taxa are classified among the taxa that define

Papuan areas of endemism (Polhemus et al. 2004;

Polhemus 2020).

Acknowledgements

lam greatly indebted to Dr. Dmitry Telnov (The

Entomological Society of Latvia, Riga) for sharing

the data used for the present research and for

valuable suggestions during the preparation of the

final version of the article, and to Laszlo Wagner

(http://www.east-indonesia.info) for field-guidance

and help during the expeditions. Dan A. Polhemus

(Bernice P. Bishop Museum, Honolulu, Hawaii,

U.S.A.) is thanked for providing identifications. and

valuable comments on the general distribution of

the species discussed.

References

Andersen N. M. 1975. The Limnogonus and Neogerris

of the Old World. - Entomologica Scandinavica,

Supplement 7: 1-96.

Andersen N. M., Weir T. A. 2004. Australian Water

Bugs. Their Biology and Identification (Hemiptera-

Heteroptera, Gerromorpha & Nepomorpha).

Entomograph volume 14, Apollo Books & CSIRO

Publishing: 344 pp.

Buzzetti F. M., Nieser N., Damgaard J. 2006. Notes on

water bugs from South East Asia and Australia

(Heteroptera: Nepomorpha & Gerromorpha). - Atti

pp. 239-244

della Accademia Roveretana degli Agiati a. 256,

Series VIII. Volume VI, B: 31-45.

Damgaard J., Zettel H. 2014. The water bugs

(Hemiptera-Heteroptera: Gerromorpha and

Nepomorpha) of New Caledonia: Diversity, ecology

and biogeographical significance: 219-238. In:

Guilbert E., Robillard T., Jourdan H., Grandcolas

P. (eds) Zoologia Neocaledonica 8. Biodiversity

studies in New Caledonia. Mémoires du Muséum

national d’Histoire naturelle, Paris: 315 pp.

Greke K., Telnov D. 2014. Review and assessment of

the literature on recent non-marine molluscs of the

Papuan biogeographical region: 61-112. In: Telnov

D. (ed.) Biodiversity, biogeography and nature

conservation in Wallacea and New Guinea. Volume

Il. The Entomological Society of Latvia, Riga: 458

pp, 126 pls.

Insecta in GBIF Secretariat. 2019. GBIF Backbone

Taxonomy. Checklist dataset https://doi.

org/10.15468/390mei accessed via_ GBIF.org

[accessed 25 January 2021].

Lansbury |. 1972. A review of the Oriental species

of Ranatra Fabricius (Hemiptera-Heteroptera:

Nepidae). - Transactions of the Royal Entomological

Society of London 124, No 3: 287-341.

Lundblad O. M. 1933. Zur Kenntnis der aquatilen und

semiaquatilen Hemipteren von Sumatra, Java und

Bali. - Archiv fur Hydrobiologie, Supplement Band

12: 263-488.

Polhemus D. A., Polhemus J. T. 1998. A review of the

genus Limnometra Mayr in New Guinea, with the

description of a very large new species (Heteroptera:

Gerridae). - Journal of the New York Entomological

Society 105: 24-39.

Polhemus D. A., Polhemus J. T. 2011. Seven new species

of Rhagovelia (Heteroptera: Veliidae) from the

Raja Ampat Islands, Eastern Indonesia. - Aquatic

Insects 33, No 1: 41-74.

Polhemus D. A., Englund R. A., Allen G. R. 2004.

Freshwater biotas of New Guinea and nearby

islands: analysis of endemism, richness, and

threats. Bishop Museum Technical Report 34: 62

Ppp.

Polhemus D. A. 2020. Nine new species of Enithares

(Heteroptera: Notonectidae) from New Guinea, with

distributional notes on other species and an updated

world checklist. - Zootaxa 4772, No 1: 132-182.

https://doi.org/10.11646/zootaxa.4/772.1.5

Riedel A. 2002. Taxonomy, phylogeny, and zoogeogra-

phy of the weevil genus Euops (Insecta: Coleoptera:

Curculionoidea) in the Papuan region. Dissertation

zur Erlangung des Doktorgrades der Fakultat

fur Biologie der Ludwig-Maximilians-Universitat

Munchen: 216 pp.

The Times Atlas of the World 1994. 4" Revised Concise

Edition. London.

Received: 27.1.2021.

Accepted: 06.ii.2021.

? eiex

243

02-Jun-21 21:49:59

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

NG kcamakawalar,

Oksibil S

National Geographic, Esri, Garmin, HERE, UNEP-WCMC USGS NoAAMincrement P Garp:

Map 1. The localities mentioned in the present paper shown on the National Geographic World Map (National

Geographic, Esri, Garmin, HERE, UNEP-WCMC, USGS, NASA, ESA, METI, NRCAN, GEBCO, NOAA, increment P Corp.).

Wallace's line] :

Map 2. The Papuan region in strict and broad senses and the surrounding regions, showing important

zoogeographic lines (redrawn from the Times Atlas of the World (1994), with modifications by Riedel (2002)).

i =~ &

Book4.indd 244 02-Jun-21 21:50:01

Book4.indd 245

KazANTSEV, S. V.: New Plateros Bourgeois, 1879 from New Guinea and the Moluccas, with a checklist of species ...

pp. 245—252

New Plateros Bourgeois, 1879 from New Guinea and

the Moluccas, with a checklist of species of the region

(Coleoptera: Lycidae)

urn:lsid:zoobank.org:pub:C5/73A/7D5-81BE-4654-AF36-B2C84/7F72DF1

SerGeEY V. KAZANTSEV

Insect Centre, Donetskaya 13-326, 109651, Moscow, Russia; kazantss@mail.ru

Abstract: Three new species of net-winged beetles of the genus Plateros Bourgeois, 1879, namely P. bacchusi

sp. nov., P. doberaiensis sp. nov. and P. halmaheraensis sp. nov., are described from New Guinea and Halmahera

(North Moluccas). Descriptions of P. grootaerti Kazantsev, 2011, P. holynskiorum Kazantsev, 2011, P. papuaensis

Kazantsev, 2011, P. rufosuturalis Kazantsev, 2011 and P. villosopenis Kazantsev, 2011 from New Guinea are

complemented with photos of the adults habitus and aedeagus. A checklist of Plateros species of the region with

information on their distribution is provided.

Key words: Platerotini, new species, checklist, Papuan Region.

Introduction

The genus Plateros Bourgeois, 1879, one of the

largest in the family of net-winged beetles and one

of the most widespread, includes over 800 species

distributed in all biogeographic realms, mostly in

the Palaeotropics (e.g. Kleine 1933; Kazantsev

2011). The number of its species has been recently

steadily rising in all them, and the Papuan region is

no exception. First Plateros species were described

in New Guinea in the twenties and thirties of the

XXth century (Pic 1921; 1923; Kleine 1939), with

their number in the New Guinea area sky-rocketing

from just three to seventy three within the last

twenty years (Bocakova 1997; Kazantsev 2011;

2015a; Kazantsev & Telnov 2017).

The present study is a further contribution

to the knowledge of Plateros of New Guinea and

the surrounding islands, including the Moluccas.

Examination of the recently collected material

allows describing another three new species;

additional illustrations are provided for some of the

previously described ones, along with a checklist of

all known species of the region.

Material and methods

The studied specimens were glued on

cardboard mounting plates. For detailed

examination they were relaxed in water; then the

detached ultimate abdominal segments were

treated for several hours in 10% KOH at room

temperature, then, with the extracted genitalia,

placed in microvials with glycerin; or glued back on

mounting plates, along with the extracted genitalia,

without the KOH treatment.

MSP-1 zoom _ stereoscopic dissecting

microscope with x8-x80 magnification range was

used. Photographs were taken with Canon EOS 6D

camera and Canon MP-E 65 mm lens.

Acronyms for scientific collections used in the text:

BMNH — Natural History Museum (formerly British

Museum, Natural History), London, United

Kingdom;

ICM — Insect Center, Moscow, Russia;

NME — Naturkundemuseum Erfurt, Germany.

Results

Platerotini Kleine, 1929

Plateros Bourgeois, 1879

Plateros Bourgeois, 1879: xix

Type species: Eros brasiliensis Lucas, 1857

subsequsent designation by Zaragoza (1999).

Distribution: All biogeographic regions; absent

only in western Palaearctic, Greater Antilles,

Madagascar, New Zealand and Melanesia /

Polynesia (Kazantsev 2011).

ey &e

245

a ha

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Plateros bacchusi sp. nov. (Figs 1—3)

Holotype 4 BMNH: New Guinea, Madang. Distr.,

Finisterre Mts., Damanti, 3550 ft, 2-11.X.1964; Stn No.

46; M.E. Bacchus, B.M. 1965-120.

Paratype 12 BMNH: same label as in holotype.

Derivatio nominis: The new species is named

after the collector of its type series, M. E. Bacchus.

Description: Length 5.2-5.9 mm. Width across

humeri 1.2-1.4 mm. Dark brown to black.

Antennomere 2, trochanters and proximal fourth

of femurs testaceous to light brown. Vertex densely

punctate, with conspicuous round impression

behind antennal prominence. Eyes large,

interocular distance ca. 1.6 times shorter than eye

diameter. Labrum small, transverse, moderately

convex anteriorly; epistoma truncate. Palps

Slender; ultimate palpomeres small, elongate,

almost parallel-sided, flattened and oblique at

apex. Antennal sockets separated by minute

lamina. Antennae long, attaining to elytral five

sixths, feebly serrate; antennomere 3 ca. 4.3 times

longer than antennomere 2 and 1.3 times shorter

than antennomere 4; antennomeres 3-11 with long

erect pubescence (Fig. 1). Pronotum transverse,

ca. 1.4 times wider than long, with conspicuously

convex and explanate sides, bisinuate basally

and strongly triangularly produced anteriorly, with

small acute posterior and obsolete anterior angles.

Scutellum transverse, slightly narrowing distally

and emarginate at apex (Fig. 1). Elytra long, ca. 3.4

times longer than wide at humeri, slightly concave

at sides, shining, with four equally developed

primary costae; interstices with double rows of

small round cells; pubescence scarce, short and

decumbent (Fig. 1). Legs long, slender; tibiae and

femoris straight, narrow; tibiae subequal in length to

femoris (Fig. 1). Male aedeagus with conspicuously

widened and curved medially median lobe, with two

longitudinal ridges in median portion, two hooked

distal projections ventrally and two rounded distal

dents dorsally (Figs 2-3).

Sexual dimorphism: Female similar to male,

but eyes smaller, filiform antennae shorter, their

pubescence decumbent.

Differential diagnosis: Plateros bacchusi Sp. nov.

IS quite similar in the structure of male genitalia to

P. maai (Bocakova, 1997), easily separable by the

Shape of its pronotum with conspicuously convex

and explanate sides and obsolete anterior angles

(vs emarginate sides and pronounced anterior

angles in P. maail) (Figs 1—3).

Distribution: Known only from the Finisterre

Mountains in Madang Province, Papua New Guinea.

—_—_

246

Plateros doberaiensis sp. nov. (Figs 4—6)

Holotype 4 NME: East Indonesia, W New Guinea,

Doberai Peninsula, 15.5-14 km N Ayamaru_ vill.,

01°08'04’’S, 132 1:0 5OF E to Ol" O9229"S;

132°11’30”E, 275-250 m, primary lowland rainforest

on limestone, 2.1X.2015, leg. D.Telnov.

Derivatio nominis: The name of the new species

is derived from the locality, ‘Doberai Peninsula’,

where the unique specimen was collected.

Description: Length 4.5 mm. Width across humeri

1.0 mm. Dark brown. Coxae, trochanters, proximal

half of femoris and tarsomeres 5 testaceous to

light brown. Elytra, except narrow distal margin,

whitish yellow; elytral narrow distal margin gray.

Vertex densely punctate, with conspicuous deep

round impression behind antennal prominence.

Eyes relatively large, interocular distance ca. 1.1

times shorter than eye diameter. Labrum small,

transverse, moderately convex anteriorly. Palps

slender; ultimate palpomeres small, elongate,

securiform, widest in distal third. Antennal sockets

separated by minute lamina. Antennae _ long,

attaining to elytral five sixths, filiform; antennomere

3 ca. 2.5 times longer than antennomere 2 and 1.3

times shorter than antennomere 4; antennomeres

3-11 with short sub-erect pubescence (Fig. 4).

Pronotum transverse, ca. 1.4 times wider than

long, with almost parallel sides, bisinuate basally

and semi-circularly produced anteriorly, with

conspicuous acute posterior and blunt rounded

anterior angles. Scutellum sSubquadrate, slightly

emarginate at apex (Fig. 4). Elytra long, ca. 3.3

times longer than wide at humeri, parallel-sided,

with four equally developed primary costae;

interstices with double rows of small subquadrate

cells; pubescence moderately dense, short and

decumbent (Fig. 4). Legs long, slender; tibiae and

femoris straight, narrow; tibiae inconspicuously

Shorter than femoris (Fig. 4). Male aedeagus with

narrow, conspicuously curved, first inwardly, then

outwardly, median lobe, tapering and hooded

distally, with several barbs (Figs 5—6).

Sexual dimorphism: Female unknown.

Differential diagnosis: Plateros doberaiensis sp.

nov. is somewhat similar in the structure of male

genitalia to P. simbaensis (Bocakova, 1997), easily

separable by the elytral coloration (with yellow

anterior fourth in RP simbaensis) and details of the

more narrow median lobe of the aedeagus (Figs

4—6).

Distribution: Known only from lowlands of Doberai

Peninsula in West New Guinea.

02-Jun-21 21:50:01

KazANTSEV, S. V.: New Plateros Bourgeois, 1879 from New Guinea and the Moluccas, with a checklist of species ...

pp. 245—252

Figures 1—12. Habitus and aedeagi of Plateros. 1-3 - P. bacchusi sp. nov.; 4-6 - P. doberaiensis sp. nov.; 7-9 - P.

halmaheraensis sp. nov.; 10-12 - P. grootaerti Kazantsev, 2011; 1-9 - Holotypes; 10-12 - Paratype; 1, 4, 7, 10

- Habitus, dorsal view; 2-3, 5-6, 8-9 & 11-12 - Aedeagi: 2, 5,8 & 11 - Dorsal view; 3, 6, 9 & 12 - Lateral view

[figures 1, 4, 7 & 10 not to scale; scale bars for figures 2-3, 5-6, 8-9 & 11-12 - 0.5 mm].

Ce 247

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Book4.indd 248

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Plateros halmaheraensis sp. nov. (Figs 7—9)

Holotype 4 NME: East Indonesia, North Moluccas,

Halmahera SE, Patani W, Sibenpopo vill. E, 0°20’31’’N,

128°32’02”"E, 15 m, sago swamp and stream,

10.VII.2013, leg. D.Telnov.

Derivatio nominis: The name of the new species

is derived from the island, ‘Halmahera’, where the

unique specimen was collected.

Description: Length 4.3 mm. Width across humeri

1.0 mm. Dark brown. Antennomere 2 and distal

half of antennomere 11, narrow pronotal margins,

trochanters, bases of femoris and tarsi testaceous

to light brown. Vertex with deep longitudinal

impression behind antennal prominence. Eyes large,

interocular distance ca. 1.8 times shorter than eye

diameter. Labrum small, transverse, slightly convex

anteriorly. Palos slender; ultimate palpomeres

small, relatively short, slightly narrowing distally.

Antennal sockets separated by minute lamina.

Antennae attaining to elytral three fourths, from

antennomere 3 flattened, serrate; antennomere 3

ca. 2.8 times longer than antennomere 2 and 1.1

times shorter than antennomere 4; antennomeres

3-11 with relatively long erect pubescence (Fig. 7).

Pronotum transverse, ca. 1.6 times as wide as long,

trapezoidal, concave basally and semi-circularly

produced anteriorly, with conspicuous long acute

posterior and obsolete anterior angles. Scutellum

transverse, truncate at apex (Fig. 7). Elytra long, ca.

3.2 times longer than wide at humeri, parallel-sided,

but somewhat widening posteriad humeri, with

four equally developed primary costae; interstices

with double rows of small Subquadrate cells;

pubescence dense, concealing reticulation, short

and decumbent, with longer scarce erect bristles

(Fig. 7). Legs relatively short; tibiae and femoris

Straight, subequal in length (Fig. 7). Male aedeagus

with narrow, more or less straight median lobe,

bent and hooked distally, with ventro-distal ladle

and broadened inner sac in distal half (Figs 8-9).

Sexual dimorphism: Female unknown.

Differential diagnosis: Plateros halmaheraensis

sp. nov., being somewhat similar in general

appearance to P. doberaiensis sp. nov., may be

distinguished by the broader body, larger eyes,

denser elytral pubescence, serrate antennae,

narrow light brown pronotal margins and completely

different structure of the aedeagus (Figs 7-9).

Distribution: Known only from Halmahera, North

Moluccas.

248

Plateros grootaerti Kazantsev, 2011 (Figs 10—12)

Plateros grootaerti Kazantsev, 2011: 177

Paratype < ICM: Papua New Guinea, Canopy Mission,

Madang prov., Baiteta, light, I1V.1996, O. Missa leg.

Note: The description of Plateros grootaerti

Kazantsev, 2011 is complemented with photos of

the habitus and aedeagus of a paratype.

Plateros holynskiorum Kazantsev, 2011 (Figs

13—15)

Plateros holynskiorum Kazantsev, 2011: 177

Holotype <& ICM: Papua New Guinea, 10 km N Madang,

2 m NNW Riwo, 9-11.11.(19)89, M. & R. Holynski leg.

Note: The description of Plateros holynskiorum

Kazantsev, 2011 is complemented with photos of

the habitus and aedeagus of the holotype.

Plateros papuaensis Kazantsev, 2011 (Figs 16—

18)

Plateros papuaensis Kazantsev, 2011: 179

Holotype <@ ICM: Papua New Guinea, 10 km N Madang,

2 m NNW Riwo, 9-11.11.(19)89, M. & R. Holynski leg.

Note: The description of Plateros papuaensis

Kazantsev, 2011 is complemented with photos of

the habitus and aedeagus of the holotype.

Plateros rufosuturalis Kazantsev, 2011 (Figs 19—

21)

Plateros rufosuturalis Kazantsev, 2011: 180

Holotype 4 ICM: Papua New Guinea, 38 km N Madang,

3 km NW Matucar, 27.1.1989, M. & R. Holynski leg.

Note: The description of Plateros rufosuturalis

Kazantsev, 2011 is complemented with photos of

the habitus and aedeagus of the holotype.

Plateros villosopenis Kazantsev, 2011 (Figs 22—

24)

Plateros villosopenis Kazantsev, 2011: 180

Paratype @ ICM: Papua New Guinea, Canopy Mission,

Madang prov., Baiteta, light, 20.VI.1996, O. Missa leg.

Note: The description of Plateros villosopenis

Kazantsev, 2011 is complemented with photos of

the habitus and aedeagus of a paratype.

02-Jun-21 21:50:02

KazANTSEV, S. V.: New Plateros Bourgeois, 1879 from New Guinea and the Moluccas, with a checklist of species ...

pp. 245—252

Figures 13-24. Habitus and aedeagi of Plateros. 13-15 - P. holynskiorum Kazantsev, 2011; 16-18 - P. papuaensis

Kazantsev, 2011; 19-21 - P. rufosuturalis Kazantsev, 2011; 22-24 - P. villosopenis Kazantsev, 2011; 13-19 -

Holotypes; 22-24 - Paratype; 1, 16, 19, 22 - Habitus, dorsal view; 14-15, 17-18, 20-21 & 23-24 - Aedeagi: 14,

17, 20, 23 - Dorsal view; 15, 18, 21 & 24 - Lateral view [figures 13, 16, 19 & 22 not to scale; scale bars for figures

14-15, 17-18, 20-21 & 23-24 - 0.5 mm].

€ »

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TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

The list of the net-winged beetles of the genus

Plateros Bourgeois, 1879 known from New

Guinea and the surrounding islands, including

the Moluccas

The list now includes seventy-six species.

admirabilis (Bocakova, 1997): 196 (Melaneros). Papua

New Guinea: Wau, 1250-1300 m; Wau, Big Creek,

1200 m; Karimui, 1080 m.

aduncus (Bocakova, 1997): 201 (Melaneros). Papua

New Guinea: Wau, 1200-1300; Mt. Missim; E Mt.

Albert Edward, 1450-1800 m.

albofasciatus (Bocakova, 1997): 195 (Melaneros). Irian

Jaya: Mokwam, Kwau, 1300-1650 m.

anulifer Kazantsev, 2015: 27. East Indonesia: Raja

Ampat, Waigeo Island.

bacchusi sp. nov. Papua New Guinea: Madang, Finisterre

Mts., 3550 ft.

baloghi (Bocakova, 1997): 194 (Melaneros). Papua New

Guinea: Nondung, 750 m; Wau, Eddy Creek, 2050

bocakovae Kazantsev, 2011: 189 (replacement name

for Plateros prominens (Bocakova, 1997), nec

Plateros prominens Kleine, 1940). Papua New

Guinea: Mt. Giluwe, 2550 m.

= Melaneros prominens Bocakova, 1997: 208

brandti (Bocakova, 1997): 209 (Melaneros). Papua New

Guinea: Finisterre Range, Saidor, Matoko.

bruggei Kazantsev, 2011: 177. ‘Neth. New Guinea,

Hollandia’, 250 feet.

bulolensis (Bocakova, 1997): 210 (Melaneros). Papua

New Guinea: Wau, 1200-1300 m; Wau, Bulolo R.,

800-1100 m; Wau, 1150 m; Wau, Big Creek, 300

caligulus Kazantsev, 2015: 28. Papua New Guinea:

Madang, Baiteta.

candidus Kazantsev, 2017: 369. West New Guinea:

Doberi Peninsula, Arfak Mts., 1900-2480 m.

chimbuensis (Bocakova, 1997): 207 (Melaneros). Papua

New Guinea: Chimbu V., 1800 m; West Highlands,

Kamans nr Minj, 1840 m.

ciconius Kazantsev, 2015: 28. West Papua: Bird’s Neck,

150-200 m.

clissoldi (Bocakova, 1997): 216 (Melaneros). Papua

New Guinea: Wau, Nakate Ridge, 1800 m.

consuetus (Bocakova, 1997): 209 (Melaneros). Papua

New Guinea: Western Highlands, Minj.

cordatus (Bocakova, 1997): 216 (Melaneros). Papua

New Guinea: Baiyer R.

corniger Kazantsev, 2015: 30. East Indonesia: Raja

Ampat, Waigeo Island.

cornutus (Bocakova, 1997): 199 (Melaneros). Papua

New Guinea: Madang.

decoctus (Bocakova, 1997): 207 (Melaneros). Papua

New Guinea: Bismarck Range, Simbai.

deflectus (Bocakova, 1997): 204 (Melaneros). Papua

New Guinea: Wau, 1200 m.

250

dimidiatus (Bocakova, 1997): 217 (Melaneros). Papua

New Guinea: Wau, Morobe distr., Mt. Missim, 880-

1050 m; Wau, 1150 m.

doberaiensis sp. nov. West New Guinea: Doberai

Peninsula, 250-275 m.

excelsus (Bocakova, 1997): 220 (Melaneros). Papua

New Guinea: Yaibos, 2030-2180 m.

falcatus (Bocakova, 1997): 205 (Melaneros). Papua

New Guinea: Maprik, 160 m; Irian Jaya: Cyclops

Mts., Sentani, 300-450 m.

filum (Bocakova, 1997): 202 (Melaneros). Papua New

Guinea: W Okapa, 1870 m; Tambul, 2200-2250 m.

flavofasciatus (Bocakova, 1997): 193 (Melaneros). Irian

Jaya: Baliem Tal, 1700 m; Jayawijaya, Bime 1600-

1900 m.

flavohumeralis (Bocakova, 1997): 198 (Melaneros).

Papua New Guinea: Big Wau Creek, 1050-1200 m;

Bulolo R., 850 m.

gravis (Bocakova, 1997): 215 (Melaneros). Papua New

Guinea: Wau, 1150-1600 m.

grootaerti Kazantsev, 2011: 177. Papua New Guinea:

Madang, Baiteta.

guineensis (Pic, 1921): 2 (Graciloplateros). Irian Jaya;

Humbold Bay; Cyclops Mts.; Wisselmeren, Kamo,

1500-1700 m.

= Plateros bicolor Kleine, 1939: 112

halmaheraensis sp. nov. North Moluccas: Halmahera,

15 m.

holynskiorum Kazantsev, 2011: 177. Papua New Guinea:

Madang, Baiteta.

humboldi Pic, 1923: 35. ‘New Guinea’.

ibaensis (Bocakova, 1997): 197 (Melaneros). Irian Jaya:

Iba, 1300 m.

imbricatus (Bocakova, 1997): 219 (Melaneros).

Bismarck Arch., Manus Island, 1-75 m.

inclinatus (Bocakova, 1997): 200 (Melaneros). Irian

Jaya: Wisselmeren, Obano, 1770 m.

indentatus Kazantsev, 2011: 177. Papua New Guinea:

Madang, Baiteta.

inflexus (Bocakova, 1997): 218 (Melaneros). Papua

New Guinea: Wau, Morobe distr., 1200 m; Wau,

McAdam Park.

infusus (Bocakova, 1997): 220 (Melaneros). Papua New

Guinea: Wau, Bulolo R., 850-900 m.

laditatus Kazantsev, 2011: 190 (replacement name for

Plateros dilatatus (Bocakova, 1997), nec Plateros

dilatatus (Pic, 1926) (Calleros)). Papua New

Guinea: Western Highlands, Minj.

= Melaneros dilatatus Bocakova, 1997: 221

languidulus (Bocakova, 1997): 221 (Melaneros).

Papua New Guinea: Wewak, Sepik distr., 30 m;

Fimschnatou; Stephansort, Astrolab Bai, 1900 m.

maai (Bocakova, 1997): 198 (Melaneros). lrian Jaya:

S Hollandia, Waris, 450-500 m; Cyclops Mts., W

Sentani, 150-250 m.

madangensis Kazantsev, 2011: 178. Papua New

Guinea: Madang, Baiteta.

makarovi Kazantsev, 2011: 178. Papua New Guinea:

Madang, Baiteta.

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KazANTsEV, S. V.: New Plateros Bourgeois, 1879 from New Guinea and the Moluccas, with a checklist of species ...

miladae Kazantsev, 2011: 190 (replacement name for

Plateros gressitti (Bocakova, 1997), nec Plateros

gressitti Nakane, 1971). Papua New Guinea:

Kokoda, 400 m.

= Melaneros gressitti Bocakova, 1997: 203

montanus (Bocakova, 1997): 210 (Melaneros). Papua

New Guinea: Mt. Wilhelm, 3000 m; Giluve, 2850

m; Epilam.

nigronotatus (Bocakova, 1997): 208 (Melaneros). Papua

New Guinea: Mt. Giluwe, 2550 m.

nigropallescens (Bocakova, 1997): 211 (Melaneros).

lrian Jaya: Diuremma, 1900-2100 m.

novaguineensis Kazantsev, 2011: 178. Papua New

Guinea: Madang, Baiteta.

pallescens (Bocakova, 1997):

Jaya: Baliem tal, 1700 m.

papuaensis Kazantsev, 2011: 179. Papua New Guinea:

Madang, Baiteta.

211 (Melaneros). Irian

perticatus (Bocakova, 1997): 218 (Melaneros). Irian

Jaya: Wisselmeren, Kamo V., 1500 m.

praecipuus (Bocakova, 1997): 197 (Melaneros). Irian

Jaya: Hollandia-Binnen, 100 m.

proprius (Bocakova, 1997): 199 (Melaneros). Irian Jaya:

Biak Island, 50-100 m.

quateorum (Bocakova, 1997): 195 (Melaneros).

Jaya: Vogelkop, Kebar V., 550 m.

riedeli (Bocakova, 1997): 200 (Melaneros). Irian Jaya:

Jayawijaya, Bommela, 1750 m.

rigidus (Bocakova, 1997): 214 (Melaneros). Papua New

Guinea: Central Mts., Archbold Lake, 760 m.

rufosuturalis Kazantsev, 2011: 180. Papua New Guinea:

Madang, Baiteta.

samuelsoni (Bocakova, 1997): 201 (Melaneros). Papua

New Guinea: Wau, 1200-1300 m; Wau, Morobe

distr., 1200-1300 m; Mt. Missim, 1300-2100 m;

SW U Watut, 1500-1800 m; Kunai Ck., 1500 m.

sedlaceki (Bocakova, 1997): 192 (Melaneros). Tambul,

2250 m; Mt. Giluwe, 2200-3280 m; Wau, Nami

Creek, 2600 m; Wau, Edie Ck, 2050-2300 m;

Aiyura, 1700; V. Watut, WS 1100-1600 m; Jimmi V.

- Baier R., 1750 m; Mt. Otto, 2200 m; Lake Sirunki,

25/70 m; Daulo Pass, 2400 m; Mt. Kaindy, 2100-

2350 m; S Mt. Hagen, 2000-2300 m; Yaibos, 2080

m; E Mt. Albert Edward, 1450 m; Buolo, 200 m;

lrian Jaya, Baliem Tal, 1700 m.

semiapertus (Bocakova, 1997): 215 (Melaneros). Papua

New Guinea: Bismarck Range, Simbai, 1900

sentus (Bocakova, 1997): 196 (Melaneros). Papua New

Guinea: Wewak, Sepik distr.

serratus (Bocakova, 1997): 194 (Melaneros). lrian Jaya:

Baliem Tal, 1700 m; Jayawijaya, Bime, 1600-1900

m; Bombela, 1750 m.

simbaiensis (Bocakova, 1997): 205 (Melaneros). Papua

New Guinea: Bismarck Range, Simbai, 1900 m.

simplex (Bocakova, 1997): 203 (Melaneros). Papua New

Guinea: Wau, Morobe distr., 1100 m; Wau, Kilolo

Creek.

sollemnitex (Bocakova, 1997): 206 (Melaneros). Papua

New Guinea: Rubia, Markham, 50 m; Morobe distr.,

lrian

pp. 245—252

Miudik, 1200-1600 m; Wewak, 20 m; Irian Jaya:

Waris, 450 m; Hollandia-Binnen, 100 m.

spinosus (Bocakova, 1997): 217 (Melaneros). Papua

New Guinea: Bulolo R., 900-1100 m; Garaina, 800

supinus (Bocakova, 1997): 212 (Melaneros). lrian Jaya:

Nabire, 5-50 m.

tenuissimus (Bocakova, 1997) (Melaneros): 202. Papua

New Guinea: Kepilam, 2420-2490 m; Mt. Sinonki,

2580 m.

turritus (Bocakova, 1997): 213 (Melaneros). Papua New

Guinea: Wau, Bulolo R., 850-900 m; Bulolo, 730 m.

uncinatus (Bocakova, 1997): 213 (Melaneros). Papua

New Guinea: Morobe distr., 1200-1300 m; Morobe,

Aseki, 1000-1300 m.

usitatus (Bocakova, 1997): 214 (Melaneros). lrian Jaya:

Cyclops Mts, Ifar, 400 m; Jayapura, Cyclops Mts.,

Sentani, 950-1450 m.

villosopenis Kazantsev, 2011: 180. Papua New Guinea:

Madang, Baiteta.

waigeoacus Kazantsev, 2015: 30. East Indonesia: Raja

Ampat, Waigeo Island; West Papua: Bird’s Neck,

9-30 m.

wauensis (Bocakova, 1997): 204 (Melaneros). Papua

New Guinea: Wau, 1200-1300 m; Mt. Missim,

1200 m; Wau, 1150-1600 m; Mt. Kaindi, 1550 m.

Discussion

There are three major types of colour

pattern in New Guinea Plateros: with mostly black

upperside, sometimes with narrowly testaceous

margined pronotum (as in Figs 1, 4, 7), with

testaceous upperside (as in Fig. 22) and with

black elytra, sometimes with narrow testaceous

margins, and testaceous pronotum (as in Figs

13, 16, 19), sometimes brightened with yellow

elytral band or whitish or yellowish anterior

part of the elytra (Bocakova 1997). Although

seemingly diverse, this variety is noticeably poorer

compared to some of the other New Guinea lycid

lineages, such as Metriorrhynchini, which display

a much greater number of colour patterns that

include blue, green, grey, yellow upperside (é.g.,

Kazantsev 2015b; 2017; Kazantsev & Telnov

2017) and form apparently more still little explored

mimetic assemblages, which include Cantharidae,

Chrysomelidae (Kazantsev & Telnov 2017) and

possibly many other insect groups.

Acknowledgements

It is my pleasant duty to express gratitude to

Dr. Matthias Hartmann (Naturkundemuseum Erfurt,

Ye se

251

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252

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Germany) and Dr. Dmitry Telnov (Natural History

Museum, London, United Kingdom), through whose

courtesy | was able to study the Lycidae collections

under their care.

References

Bocakova M. 1997. Revision and phylogenetic analysis of

the genus Melaneros Fairmaire, 1877 (Coleoptera,

Lycidae) from New Guinea. - Entomologica

basiliensia 20: 165-222.

Kazantsev S. V. 2011. New and little known taxa of

Platerotini, with a note on biogeography of the tribe

(Lycidae, Coleoptera). - Russian Entomological

Journal 20, No 2: 151-187.

Kazantsev S. V. 2015a. Six new species of Plateros

Bourgeois from New Guinea (Coleoptera, Lycidae).

— Eurasian Entomological Journal 14 No 1: 27-31.

Kazantsev S. V. 2015b. New species of Metriorrhynchus

Gemminger et Harold, 1869 and Porrostoma

Laporte, 1838 from New Guinea (Coleoptera:

Lycidae). - Russian Entomological Journal 24 No

3: 211-233.

Kazantsev S. V. 2017. New metriorrhynchine taxa from

New Guinea (Coleoptera: Lycidae): 371-402, pls

62-79. In: Telnov D., Barclay M. V. L., Pauwels O.

S. G. (eds). Biodiversity, Biogeography and Nature

Conservation in Wallacea and New Guinea. Volume

Ill. The Entomological Society of Latvia, Riga: 658

pp, 172 pls.

Kazantsev S. V., Telnov D. 2017. A mimetic assemblage

of net-winged beetles (Coleoptera: Lycidae) from

West Papua: 363-370, pls 56-61. In: Telnov D.,

Barclay M. V. L., Pauwels O. S. G. (eds) Biodiversity,

Biogeography and Nature Conservation in Wallacea

and New Guinea. Volume Ill. The Entomological

Society of Latvia, Riga: 658 pp, 172 pls.

Kleine R. 1933. Coleopterorum Catalogus auspiciis

et auxilio W. Junk editus a Schenkling. Pars 123:

Lycidae. W. Junk, Berlin: 145 pp.

Kleine R. 1939. Neue Brenthiden und Lyciden die Miss

Cheesman 1936 in Hollaendisch Neu Guinea

gesammelt hat. - Nova Guinea. Journal of Botany,

Zoology, Anthropology, Ichnography, Geology and

Palaeonthology of the Papuan Region. New Series,

3: 106-112.

Pic M. 1921-1922. Contribution a l'étude des Lycides. -

L’Echange, hors texte 37-38, No 404/410: 1-28.

Pic M. 1923. Contribution a |’étude des Lycides (suite

et fin). - L’Echange, hors texte 39, No 411/413:

29-40.

Received: 13.iv.2020.

Accepted: 08.vii.2020.

02-Jun-21 21:50:03

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KazanTseV, S. V.: New metriorrhynchine species (Coleoptera: Lycidae) from Sulawesi

pp. 253—263

New metriorrhynchine species (Coleoptera: Lycidae)

from Sulawesi

urn:lsid:zoobank.org:pub:B423EF4 /7-C6F4-4CEE-BED9-900/7 28479685

SERGEY V. KAZANTSEV

Insect Centre, Donetskaya 13-326, 109651, Moscow, Russia; kazantss@mail.ru

Abstract: Nine new species of metriorrhynchine net-winged beetles, Mangkutanus uluwayensis sp. nov.,

Metriorrhynchus longeolivaceus sp. nov., Sulabanus bicolorinotus sp. nov., S. brunneifasciatus sp. nov., S.

nitidifasciatus sp. nov., S. torajaensis sp. nov., Xylobanus latimojongensis sp. nov., X. pangoensis sp. nov. and X.

Sinajiensis sp. nov., are described from Sulawesi. A key to species of the endemic Sulawesi genus Mangkutanus

Kubecek, Dvorak et Bocak, 2011 is provided.

Key words: Metriorrhynchini, new taxa, Mangkutanus key, Wallacea.

Introduction

The Metriorrhynchini are by far the most

diverse and species-rich group of net-winged

beetles in the Papuan region, where the relatively

modest number of higher taxonomic groups is set

off by the diversity of Lycidae at alpha-taxonomic

level (Kleine 1933; Masek et al. 2018). Although

the first metriorrhynchine taxa from the island

of Sulawesi, which is part of this region, were

described back in the second half of the nineteenth

century, only recently has its fauna started to reveal

its actual richness (Bocak & Matsuda 1997; Bocak

1999; 2000; 2001; Bocak & Jass 2004; Bocak et

al. 2006; Dvorak & Bocak 2007; 2009; Kubecek et

al. 2011).

An opportunity to study the Lycidae material

collected in Sulawesi recently allows adding

nine new metriorrhynchine species to the genera

Mangkutanus Kubecek, Dvorak et Bocak, 2011

(one species), Metriorrhynchus Gemminger et

Harold, 1869 (one species), Sulabanus Dvorak

and Bocak, 2007 (four species) and Xylobanus

Waterhouse, 1879 (three species). Mangkutanus

and Sulabanus, described just several years

ago, externally can hardly be distinguished from

Xylobanus, but their aedeagi suggest they belong

in different, and not even closely allied lineages.

Most of the new species described below come

from three localities in South Sulawesi: vicinity of

Puri Rimba Resort near Palopo, at 750-800 meters

above sea level, Mountain Pangopango near

Makale, at 1740-1780 meters above sea level, and

the Latimojong Mountains, in most cases at 2000

meters above sea level (Figs 1-2).

Material and methods

The studied specimens were glued on

cardboard mounting plates. For detailed

examination they were relaxed in water; then the

detached ultimate abdominal segments were

treated for several hours in 10% KOH at room

temperature, then, with the extracted genitalia,

placed in microvials with glycerin; or glued back on

mounting plates, along with the extracted genitalia,

without the KOH treatment.

MSP-1 zoom _— stereoscopic dissecting

microscope with x8-x80 magnification range was

used. Photographs were taken with Canon EOS 6D

camera and Canon MP-E 65 mm lens.

Acronyms for scientific collections used in the text:

ICM — Insect Center, Moscow;

ZMUU — Zoological Museum of Moscow University.

Results

Metriorrhynchini Kleine, 1926

Mangkutanus Kubecek, Dvorak et Bocak, 2011

Mangkutanus Kubecek, Dvorak & Bocak 2011: 648

Type species: Sulabanus tenggahensis Dvorak et

Bocak, 2007 (by original designation).

Distribution: Endemic of Sulawesi.

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figures 1-2. Lycidae habitats on Sulawesi. 1 - Puri Rimba Resort, 780 m above sea level, habitat of Sulabanus

bicolorinotus sp. nov. and S. brunneifasciatus sp. nov.; 2 - Slope of Mountain Sinaji, Latimojong Mountains, 2000 m

above sea level, habitat of Mangkutanus uluwayensis sp. nov., Metriorrhynchus longeolivaceus sp. nov., Xylobanus

latimojongensis sp. nov. and X. sinajiensis sp. nov.

254

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Book4.indd 255

KazantTseV, S. V.: New metriorrhynchine species (Coleoptera: Lycidae) from Sulawesi

Mangkutanus uluwayensis sp. nov. (Figs 3-8)

Holotype < ICM: Indonesia, S Sulawesi, Latimojong

Mts, N slopes [of] Sinaji Mt., env. Uluway, 3°18'36”"S

120°01’40’”E, 2000 m, 19-20.01.2020, S. Kazantsev

leg.

Paratypes 184 ICM & ZMUU: same label as holotype.

Derivatio nominis: The name of the new species

is derived from ‘Uluway’, a village at the foothills

of the Latimojong Mts in South Sulawesi, where its

type series was collected.

Description: Length 7.0-9.0 mm. Width across

humeri 1.5-2.0 mm. Dark brown to black. Pronotal

margins and elytra, except around scutellum and

along suture in proximal third, reddish light brown

(Fig. 3). Vertex finely and sparsely punctate, with

two inconspicuous small round impressions behind

antennal prominence. Eyes relatively small, eye

diameter to interocular distance ratio ca. 0.65.

Labrum small, transverse, slightly emarginate

medially at anterior margin. Palps slender; ultimate

palpomeres small, elongate, almost parallel-

sided, somewhat narrowed, flattened and oblique

at apex. Antennal sockets separated by narrow

lamina. Antennae moderately long, attaining to

elytral half, strongly serrate; antennomere 3 ca. 6.6

times longer than antennomere 2 and subequal

in length to antennomere 4; antennomeres 3-11

with short dense sub-erect pubescence (Fig. 3).

Pronotum only ca. 1.1 times wider than long, with

straight, almost parallel sides, bisinuate basally

and triangularly produced anteriorly, with small

acute posterior and inconspicuous anterior angles.

Scutellum transverse, narrowing distally, concave

at sides, deeply semi-circularly emarginate at apex

(Fig. 3). Elytra long, ca. 4 times longer than wide at

humeri, slightly widening distally, with four equally

developed primary costae; interstices with one row

of subquadrate cells, but at base and apices with

two rows of roundish cells; pubescence dense,

short and decumbent, present on costae (Fig. 3).

Legs relatively long and slender; tibiae and femurs

Straight, narrow; tibiae subequal in length to femurs

(Fig. 3).

Male: Aedeagus with elongate straight median

lobe, in apical two fifths with noticeably widened

inner sac bearing a pair of slender inconspicuously

curved spines (Figs 4-5).

Sexual dimorphism: Female unknown.

Differential diagnosis: Mangkutanus

uluwayensis sp. nov. is readily distinguishable from

congeners in the coloration and the shape of its

aedeagus (Figs 3-8).

Variation: The ratio of light and dark brown in

pp. 253—263

pronotum and elytra can vary from mostly reddish

light brown (as in the holotype, fig. 3) to almost

entirely dark brown to black, with only anterior

pronotal margin and elytral apices reddish light

brown (as in one of the paratypes, fig. 6). The

aedeagus of M. uluwayensis sp. nov. can vary from

slightly curved in dorsal view (as in the holotype,

Figs 4-5) to completely straight (as in one of the

paratypes, Figs 7-8).

Distribution: Known only from the Latimojong

Mts., South Sulawesi.

A key to species of Mangkutanus

1 Elytra black, pronotum reddish brown

M. utarensis (Dvorak et Bocak, 2007)

- Elytra bicolour, pronotum black, at most with reddish

brown margins (e.g. Figs 3 & 6)

2 Humeral three fifths of elytra lighter than apical two

fifths M. tenggahensis (Dvorak et Bocak, 2007)

- Elytra black with reddish apices or reddish testaceous

with infuscated scutellar and sutural area (Figs 3 &

6); aedeagus with narrow straight median lobe and

relatively broad, ovally widened inner sac (Figs 4-5 &

Os Sop ae a he oe, M. uluwayensis sp. nov.

Metriorrhynchus Gemminger et Harold, 1869

Metriorrhynchus Gemminger & Harold 1869: 1629

Type species: Lycus parallelus Guerin-Meneville,

1838 (Subsequsent designation by Waterhouse,

1878).

Distribution: Oriental, Papuan and Australian

regions; one species occurs in the south-east of the

Palaearctic Region.

Metriorrhynchus longeolivaceus sp. nov. (Figs

9-10)

Holotype 9 ICM: Indonesia, S Sulawesi, Latimojong

Mts, N slopes [of] Sinaji Mt., env. Uluway, 3°18’36’S

120°01’40"E, 1900-2000 m, 19-20.1.2020, S.

Kazantsev leg.

Derivatio nominis: The name of the new species

is derived from the Latin for ‘long and olive’, alluding

to the shape of its body and the coloration of its

upperside.

Description: Length 23.5 mm. Width across

humeri 5.0 mm. Olive. Pronotal margins brownish.

Abdomen dark brown; ultimate ventrite and

tergite orange testaceous (Fig. 9). Vertex densely

pubescent, with conspicuous transverse impression

behind antennal prominence. Eyes relatively small,

interocular distance ca. 1.3 times greater than eye

diameter. Labrum large, with its length subequal to

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TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figures 3-10. Habitus and aedeagi of Mangkutanus and Metriorrhynchus. 3-8 - Mangkutanus uluwayensis

sp. nov.; 9-10 - Metriorrhynchus longeolivaceus sp. nov.; 3-5 - Holotype male; 6-8 - Paratype males; 9-10 -

Holotype female; 3, 6 & 9 - Habitus, dorsal view; 4-5 & 7-8 - Aedeagi: 4 & 7 - Dorsal view; 5 & 8 - Lateral view;

10 - Female genitalia, ventral view [figures 3, 6 & 9 not to scale; scale bars for figures 4-5 & 7-8 & 10 - 0.5 mm].

256

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KazanTseV, S. V.: New metriorrhynchine species (Coleoptera: Lycidae) from Sulawesi

width, semi-oval, medially emarginate at anterior

margin; epistoma concave. Palps slender; ultimate

palpomeres small, elongate, almost parallel-sided,

narrowed and glabrous in distal half, rounded

at apex. Antennal sockets separated by narrow

lamina. Antennae long, narrow, attaining to elytral

half, feebly serrate; antennomere 3 ca. 10 times

longer than antennomere 2 and ca. 1.1 times longer

than antennomere 4; antennomeres 3-11 with

short decumbent pubescence (Fig. 9). Pronotum

transverse, ca. 1.3 times wider than long, with

conspicuously slightly concave sides, bisinuate

basally and noticeably produced anteriorly, with

prominent acute posterior and obsolete anterior

angles; transverse rib hardly noticeably near

median areole. Scutellum transverse, parallel-

sided and triangularly emarginate at apex (Fig.

9). Elytra long, ca. 4 times longer than wide

at humeri, slightly narrowing distally, with four

equally developed primary costae; interstices with

double rows of small very narrow transverse cells;

pubescence dense, short and decumbent (Fig. 9).

Legs very long, slender; tibiae and femurs straight,

narrow; tibiae subequal in length to femurs; femurs

noticeably swollen near apex (Fig. 9).

Female: External female genitalia with elongate,

ca. 3.2 times longer than wide, styli, relatively short

membranous coxites and relatively long, slightly

bent in the middle and truncate at base, free

sclerotised valvifers (Fig. 10).

Sexual dimorphism: Male unknown.

Differential diagnosis: Metriorrhynchus

longeolivaceus Sp. nov. iS easily separable from its

Sulawesi congeners by the coloration, it may also be

distinguished from M. thoracicus (Fabricius, 1801)

and allied species (Bocak & Matsuda, 1997) by

the longer styli and shorter and differently shaped

valvifers (Figs 9-10).

Distribution: Known only from the Latimojong

Mts. (South Sulawesi).

Sulabanus Dvorak et Bocak, 2007

Sulabanus Dvorak & Bocak 2007: 3

Type species: Sulabanus mamasensis Dvorak

and Bocak, 2007 (by original designation).

Distribution: Endemic of Sulawesi.

Sulabanus bicolorinotus sp. nov. (Figs 11-13)

Holotype <4 ICM: Indonesia, S Sulawesi, W Palopo,

Puri Rimba Resort, 2°57’'33”"S 120°05’12”E, 780 m, at

light, 7.1.2020, S. Kazantsev leg.

Derivatio nominis: The name of the new species

pp. 253—263

is derived from the Latin for ‘noted for two colours’,

alluding to the coloration of its antennae and

upperside.

Description: Length 6.3 mm. Width across

humeri 1.4 mm. Dark brown to black. Antennomere

11, pronotal sides and posterior pronotal margin,

prosternum, mesoventrite, trochanters, femurs

proximally and relatively narrow band at elytral

two fifths testaceous (Fig. 11). Vertex glabrous,

shining, feebly convex. Eyes small, interocular

distance ca. 1.5 times greater than eye diameter.

Labrum transverse, moderately convex anteriorly,

inconspicuously medially emarginate. Palps

slender; ultimate palpomeres elongate, securiform,

oblique at apex. Antennal sockets separated by

minute lamina. Antennae moderately long, slightly

Surpassing elytral middle, serrate; antennomere 3

ca. 10 times longer than antennomere 2 and 1.2

times shorter than antennomere 4; antennomeres

3-11 with short semi-erect pubescence (Fig. 11).

Pronotum subquadrate, about as wide as long,

with slightly concave sides, bisinuate basally and

semi-circularly produced anteriorly, with small

rounded posterior and blunt anterior angles.

Scutellum transverse, parallel-sided, semi-

circularly emarginate at apex (Fig. 11). Elytra long,

ca. 3.8 times longer than wide at humeri, parallel-

sided, with four equally developed primary costae;

interstices with a row of regular sSubquadrate cells;

pubescence scarce, short and decumbent (Fig.

11). Legs moderately long; tibiae and femoris

Straight, relatively broad; tibiae subequal in length

to femoris (Fig. 11).

Male: Aedeagus with relatively narrow, moderately

widened in the middle and feebly emarginate

apically median lobe, with short erect hairs in the

middle portion (Figs 12-13).

Sexual dimorphism: Female unknown.

Differential diagnosis: Sulabanus bicolorinotus

sp. nov. Somewhat resembles S. nitidifasciatus sp.

nov., separable by the distinctly less narrow body,

orange testaceous antennomere 11 and bicoloured

pronotum with more acute posterior angles, as well

as by the more narrow, less widened in the middle

and feebly emarginate apically median lobe of the

aedeagus (Figs 11-13).

Distribution: Known only from the mountains

near Palopo (South Sulawesi).

Sulabanus brunneifasciatus sp. nov. (Figs 14-16)

Holotype <@ ICM: Indonesia, S Sulawesi, W Palopo, trail

nr. Puri Rimba Resort, 2°58’00”S 120°05'13’E, 720-

850 m, 23-27.|.2020, S. Kazantsev leg.

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Derivatio nominis: The name of the new species

is derived from the Latin for ‘with a brown band’,

alluding to its elytral coloration.

Description: Length 9.8 mm. Width across

humeri 2.0 mm.

Dark brown to black. Antennomere 3 proximally,

thorax, trochanters, proximal third to half of femurs

and broad elytral band below humeri brownish

testaceous (Fig. 14). Vertex densely and finely

punctate at sides and anteriorly and glabrous

in the middle, with two inconspicuous round

impressions behind antennal prominence. Eyes

relatively small, interocular distance ca. 1.4 times

greater than eye diameter. Labrum transverse,

with broad prominent median incision; epistoma

noticeably medially incised. Palps slender; ultimate

palpomeres broad, securiform, oblique at apex.

Antennal sockets separated by narrow lamina.

Antennae moderately long, slightly surpassing

elytral middle, serrate; antennomere 3 ca. 6.3

times longer than antennomere 2 and subequal in

length to antennomere 4; antennomeres 3-11 with

short semi-erect pubescence (Fig. 14). Pronotum

transverse, ca. 1.3 times wider than long, with short,

almost parallel sides, slightly bisinuate basally and

strongly triangularly produced anteriorly, with small

acute posterior and obsolete anterior angles; with

conspicuous round punctures at posterior angles

and anteriorly. Scutellum transverse, short, slightly

narrowing distally and broadly emarginate at apex

(Fig. 14). Elytra narrow and long, ca. 3.9 times longer

than wide at humeri, widest below humeri, with four

equally developed primary costae; interstices with

a row of regular Subquadrate cells; pubescence

dense, short and decumbent, present only on

longitudinal and transverse costae (Fig. 14). Legs

long; tibiae and femoris straight, relatively broad;

tibiae slightly shorter than femoris (Fig. 14).

Male: Aedeagus with relatively broad and strongly

emarginate apically median lobe, with short erect

hairs in the middle portion dorsally (Figs 15-16).

Sexual dimorphism: Female unknown.

Differential diagnosis: Sulabanus

brunneifasciatus sp. nov. iS quite similar in general

appearance and aedeagal structure to S. brancuccil

Dvorak et Bocak, 2007, separable by the distinctly

more narrow elytra (only 3.3 times longer than wide

at humeri in S. brancuccii), as well as by the more

parallel-sided median lobe of the aedeagus with

conspicuous vestiture dorsally in the middle part

(Figs 14-16).

Distribution: Known only from the mountains

near Palopo (South Sulawesi).

258

Sulabanus nitidifasciatus sp. nov. (Figs 17-19)

Holotype < ICM: Indonesia, S Sulawesi, N slopes [of]

Mt. Pangopango, 3°09’46”S 119°49’45’E, 1740-1780

m, 9-11.1.2020, S. Kazantsev leg.

Paratype 19 ICM: same label as holotype.

Derivatio nominis: The name of the new species

is derived from the Latin for ‘narrow’ and ‘with a

band’, alluding to its shape and elytral coloration.

Description: Length 6.3-6.8 mm. _ Width

across humeri 1.1-1.2 mm. Dark brown to black.

Antennomere 2 and narrow elytral band at elytral two

fifths brownish testaceous (Fig. 17). Vertex glabrous,

shining, feebly convex, with fine punctuation near

eyes. Eyes relatively small, interocular distance ca.

1.4 times greater than eye diameter. Labrum small,

transverse, slightly medially incised anteriorly;

epistoma conspicuously medially incised. Palps

slender; ultimate palpomeres small, slightly longer

than wide, broadly securiform, widest at base.

Antennal sockets separated by narrow lamina.

Antennae long, attaining to elytral two thirds,

serrate; antennomere 3 ca. 6.7 times longer

than antennomere 2 and 1.1 times shorter than

antennomere 4; antennomeres 3-11 with long erect

pubescence (Fig. 17). Pronotum subquadrate, only

ca. 1.1 times wider than long, with almost straight

sides, bisinuate basally and slightly semi-circularly

produced anteriorly, with small acute posterior

and conspicuous blunt anterior angles. Scutellum

transverse, short, rounded and emarginate at apex

(Fig. 17). Elytra narrow and long, ca. 4.1 times

longer than wide at humeri, parallel-sided, with four

equally developed primary costae; interstices with

a row of elongate rectangular cells; pubescence

scarce, short and decumbent (Fig. 17). Legs long,

slender; tibiae and femoris straight, narrow; tibiae

Subequal in length to femoris (Fig. 17).

Male: Aedeagus with relatively narrow, moderately

widened in the middle and rounded apically median

lobe, with short erect hairs in the middle portion

dorsally (Figs 18-19).

Sexual dimorphism: Female similar to male, but

eyes slightly smaller, antennae somewhat shorter

and less serrate.

Differential diagnosis: Sulabanus

nitidifasciatus sp. nov. is Somewhat similar in the

aedeagal structure to S. ambangensis Dvorak et

Bocak, 2007, easily separable by the testaceous

band on the elytra, as well as by the distinct central

widening of the median lobe of the aedeagus (Figs

17-19). S. nitidifasciatus sp. nov. also resembles

S. bicolorinotus sp. nov., but is separable by the

distinctly narrower body, uniformly dark brown

02-Jun-21 21:50:05

KazantTseV, S. V.: New metriorrhynchine species (Coleoptera: Lycidae) from Sulawesi

pp. 253—263

Figures 11-22. Habitus and aedeagi of Sulabanus, holotype males. 11-13 - S. bicolorinotus sp. nov.; 14-16 - S.

brunneifasciatus sp. nov.; 17-19 - S. nitidifasciatus sp. nov.; 20-22 - S. torajaensis sp. nov.; 11, 14, 17 & 20 -

Habitus, dorsal view; 12-13, 15-16, 18-19 & 21-22 - Aedeagi: 12, 15, 18 & 21 - Dorsal view; 13, 16, 19 & 22 -

Lateral view [figures 11, 14, 17 & 20 not to scale; scale bars for figures 12-13, 15-16, 18-19 & 21-22 - 0.5 mm].

ee 259

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Book4.indd 260

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

to black antennomere 11 and pronotum, acute

posterior pronotal angles, as well as by the rounded

apically median lobe of the aedeagus.

Distribution: Known only from Mt. Pangopango in

Tana Toraja (South Sulawesi).

Sulabanus torajaensis sp. nov. (Figs 20-22)

Holotype < ICM: Indonesia, S Sulawesi, N slopes [of]

Mt. Pangopango, 3°09’46’S 119° 49’45’E, 1740-1780

m, 9-11.1.2020, S. Kazantsev leg.

Derivatio nominis: The name of the new

species is derived from ‘Tana Teraja’, a locality in

South Sulawesi where, on Mt. Pangopango, its type

specimen was collected.

Description: Length 5.4 mm. Width across humeri

1.0 mm. Uniformly dark brown to black (Fig. 20).

Vertex glabrous, shining, feebly convex, with fine

punctuation near eyes and a pair of small round

impressions behind antennal prominence. Eyes

small, interocular distance ca. 1.8 times greaterthan

eye diameter. Labrum small, transverse, truncate

anteriorly; epistoma greatly medially incised. Palps

slender; ultimate palopomeres small, about as long

as wide, widest at base. Antennal sockets separated

by minute lamina. Antennae long, attaining to elytral

three fifths, serrate; antennomere 3 ca. 6.2 times

longer than antennomere 2 and 1.1 times shorter

than antennomere 4; antennomeres 3-11 with

short semi-erect pubescence (Fig. 20). Pronotum

subquadrate, only ca. 1.1 times wider than long,

with almost straight and parallel sides, bisinuate

basally and semi-circularly produced anteriorly,

with small acute posterior and conspicuous blunt

anterior angles. Scutellum transverse, rounded and

emarginate at apex (Fig. 20). Elytra narrow and long,

ca. 4.1 times longer than wide at humeri, parallel-

sided, with four equally developed primary costae;

interstices with a row of regular elongate cells;

pubescence scarce, short and decumbent, present

only on longitudinal and transverse costae (Fig.

20). Legs long, slender; tibiae and femoris straight,

narrow; tibiae subequal in length to femoris (Fig.

20).

Male: Aedeagus with robust, almost cylindrical

in ventral view and triangularly produced apically

median lobe, without short erect hairs in the middle

portion (Figs 21-22).

Sexual dimorphism: Female unknown.

Differential diagnosis: Sulabanus torajaensis

Sp. nov. iS Similar in the aedeagal structure to S.

similis Dvorak et Bocak, 2007, separable by the

uniformly black elytra (with bright yellow band in

S. similis), as well as by the details of the distal

260

opening and inner sac structures of the median

lobe of the aedeagus (Figs 20-22). Unlike in S.

bicolorinotus sp. nov., S. brunneifasciatus sp.

nov. and S. nitidifasciatus sp. nov., in Sulabanus

torajaensis sp. nov. the outer surface of the median

lobe does not have hairs dorsally.

Distribution: Known only from Mt. Pangopango in

Tana Toraja (South Sulawesi).

Xylobanus Waterhouse, 1879

Xylobanus Waterhouse 1879: 38

Type species: Lycus costifer Walker, 1858 (by

original designation).

Distribution: Afrotropical, Oriental and eastern

Palaearctic regions.

Xylobanus latimojongensis sp. nov. (Figs 23-25)

Holotype < ICM: Indonesia, S Sulawesi, Latimojong

Mts, N slopes [of] Sinaji Mt., env. Uluway, 3°18’36’S

120°01’40”E, 2000 m, 19-20.01.2020, S. Kazantsev

leg.

Paratype 10 ICM: same label as holotype.

Derivatio nominis: The name of the new species

is derived from the ‘Latimojong Mountains’, a

locality in South Sulawesi where its type series was

collected.

Description: Length 9.1-9.5 mm. Width across

humeri 1.8-1.9 mm. Uniformly black (Fig. 23).

Vertex finely punctate, with conspicuous oblique

impressions behind antennal prominence. Eyes

small, interocular distance ca. 1.7 times greater

than eye diameter. Labrum small, transverse,

slightly convex anteriorly; epistoma very slightly

concave. Palps slender; ultimate palpomeres

small, elongate, almost parallel-sided, flattened

and oblique at apex. Antennal sockets separated

by minute lamina. Antennae moderately long,

slightly surpassing elytral middle, antennomeres

3-11 flattened, almost parallel-sided; antennomere

3 ca. 10 times longer than antennomere 2 and 1.1

times longer than antennomere 4; antennomeres

3-11 with short sub-erect pubescence (Fig. 23).

Pronotum transverse, ca. 1.4 times wider than

long, with conspicuously concave sides, bisinuate

basally and semi-circularly produced anteriorly,

with prominent acute posterior and rounded

anterior angles; median cell oval, narrow, going

half-way to anterior margin; lateral and antero-

lateral ribs almost obsolete. Scutellum transverse,

narrowing distally, broadly emarginate at apex

(Fig. 23). Elytra long, ca. 3.8 times longer than

wide at humeri, somewhat widening distally and

02-Jun-21 21:50:05

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KazanTseV, S. V.: New metriorrhynchine species (Coleoptera: Lycidae) from Sulawesi

slightly dehiscent in distal third, with four equally

developed primary costae; interstices with a row

of regular subquadrate cells; pubescence scarce,

short and decumbent (Fig. 23). Legs long, slender;

tibiae and femoris straight, narrow; tibiae subequal

in length to femoris; tarsomeres narrow (Fig. 23).

Male: Aedeagus with relatively robust, gradually

widened distally and rounded at apex median lobe,

with a pair of relatively straight inner sac spines

(Figs 24-25).

Sexual dimorphism: Female unknown.

Differential diagnosis: Xylobanus

latimojongensis sp. nov. is easily separable

from almost all Xylobanus species, and not only

from Sulawesi, by the reduction of lateral and

anterolateral pronotal ribs, as well as by the

gradually widened distally median lobe of the

aedeagus, with rounded apex (Figs 23-25).

Distribution: Known only from the Latimojong

Mts. (South Sulawesi).

Xylobanus pangoensis sp. nov. (Figs 26-29)

Holotype ICM: Indonesia, S Sulawesi, N slopes [of]

Mt. Pangopango, 3°09’46”S 119°49’45’”E, 1740-1780

m, 9-11.1.2020, S. Kazantsev leg.

Paratypes 53 & 39 ICM: same label as holotype.

Derivatio nominis: The name of the new species

is derived from ‘Pangopango’, a mountain in South

Sulawesi where its type series was collected.

Description: Length 6.7-9.6 mm. Width across

humeri 1.4-2.1 mm. Dark brown to black. Pronotal

margins narrowly brownish; elytral vestiture olive

(Fig. 26). Vertex densely and finely punctate,

with conspicuous trapezoidal impression behind

antennal prominence with distinct narrow median

ridge. Eyes large, interocular distance ca. 1.1

times shorter than eye diameter. Labrum semi-

oval; epistoma convex. Palos slender; ultimate

palpomeres small, elongate, narrow, narrowing

distally, pointed and glabrous at apex. Antennal

sockets separated by minute lamina. Antennae

moderately long, slightly surpassing elytral middle,

relatively broad, serrate; antennomere 3 ca. 5

times longer than antennomere 2 and subequal in

length to antennomere 4; antennomeres 3-11 with

Short sub-erect pubescence (Fig. 26). Pronotum

transverse, ca. 1.3 times wider than long, with

conspicuously concave sides, bisinuate basally and

prominently semi-circularly produced anteriorly,

with long acute posterior and obsolete anterior

angles; median cell long and narrow, going half-way

to anterior margin; lateral ribs noticeably reduced.

Scutellum transverse, narrowing distally, broadly

pp. 253—263

emarginate at apex (Fig. 26). Elytra narrow and

long, ca. 4.1 times longer than wide at humeri,

parallel-sided, with four equally developed primary

costae; interstices with a row of regular strongly

transverse cells; pubescence scarce, short and

decumbent (Fig. 26). Legs long, slender; tibiae and

femoris straight, narrow; tibiae subequal in length

to femoris (Fig. 26).

Male: Aedeagus with conspicuously widened

in distal half and narrowed at apex median lobe,

with short apical spike and a pair of asymmetrical,

noticeably narrowed in distal half inner sac spines

(Figs 27-28).

Sexual dimorphism: Female similar to male,

but larger, eyes smaller, antennae distinctly shorter

and broader (Fig. 29).

Differential diagnosis: Xylobanus pangoensis

sp. nov. may be distinguished from the somewhat

similar in general appearance X. panensis Kubecek,

Dvorak et Bocak, 2011 by the only slightly concave

pronotal sides and shorter and more robust apex

of the median lobe of the aedeagus (Figs 26-29).

Distribution: Known only from Mt. Pangopango in

Tana Toraja (South Sulawesi).

Xylobanus sinajiensis sp. nov. (Figs 30-32)

Holotype < ICM: Indonesia, S Sulawesi, Latimojong

Mts, N slopes [of] Sinaji Mt., env. Uluway, 3°17’48’S

119°59’55"E, 1100-1300 m, 18-20.01.2020, S.

Kazantsev leg.

Derivatio nominis: The name of the new species

is derived from ‘Sinaji’, amountain in the Latimojong

mountain range in South Sulawesi where its type

specimen was collected.

Description: Length 5.5 mm. Width across

humeri 1.2 mm. Dark brown to black. Pubescence

on pronotal ribs and margins, as well as on elytral

costae and elytral margins brownish (Fig. 30). Vertex

finely and densely punctate, with conspicuous round

impression behind antennal prominence and a pair

of minute pits at its bottom. Eyes large, interocular

distance ca. 1.2 times shorter than eye diameter.

Labrum small, semi-oval; epistoma truncate. Palps

slender; ultimate palpomeres small, elongate,

narrow, pointed and glabrous at apex. Antennal

sockets separated by minute lamina. Antennae

moderately long, slightly surpassing elytral middle,

feebly serrate; antennomere 3 ca. 6.7 times longer

than antennomere 2 and 1.1 times shorter than

antennomere 4; antennomeres 3-11 with short sub-

erect pubescence (Fig. 30). Pronotum transverse,

ca. 1.4 times wider than long, trapezoidal, bisinuate

basally and semi-circularly produced anteriorly, with

261

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figures 23-32. Habitus and aedeagi of Xylobanus. 2-25 - X. latimojongensis sp. nov.; 26-29 - X. pangoensis sp.

nov.; 30-32 - X. sinajiensis sp. nov.; 23-28 & 30-32 - Holotype males; 29 - Paratype female; 23, 26, 29 & 30

—- Habitus, dorsal view; 24-25, 27-28 & 31-32 - Aedeagi: 24, 27 & 31 - Dorsal view; 25, 28 & 32 - Lateral view

[figures 23, 26, 29-30 not to scale; scale bars for figures 24-25, 27-28 & 31-32 - 0.5 mm].

262

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KazanTseV, S. V.: New metriorrhynchine species (Coleoptera: Lycidae) from Sulawesi

moderately long acute posterior and noticeable

blunt anterior angles. Scutellum transverse, slightly

narrowing distally and emarginate at apex (Fig. 30).

Elytra narrow and long, ca. 3.8 times longer than

wide at humeri, parallel-sided, with four equally

developed primary costae; interstices with a row of

regular subquadrate to elongate cells; pubescence

scarce, short and decumbent, present mostly on

costae (Fig. 30). Legs long, slender; tibiae and

femoris straight, narrow; tibiae subequal in length

to femoris (Fig. 8).

Male: Aedeagus with conspicuously widened in

distal half and narrowed at apex median lobe, with

elongate apical spike and a pair of symmetrical,

evenly acuminate inner sac spines (Figs 31-32).

Sexual dimorphism: Female unknown.

Differential diagnosis: Xylobanus sinajiensis

sp. nov. is Somewhat similar to X. pangoensis sp.

nov., distinguishable by the uniformly dark brown

pronotum, noticeably more narrow antennomeres,

subquadrate to elongate cells in the middle part of

elytra and narrower apical spike of the less elongate

median lobe of the aedeagus (Figs 30-32).

Distribution: Known only from the Latimojong

Mts. (South Sulawesi).

References

Bocak L. 1999. New taxa of the subtribe Hemiconderina

(Coleoptera: Lycidae) from Indonesia and New

Guinea. - Entomologische Blatter 95, No 2/3:

166-170.

Bocak L. 2000. Revision of the genus Wakarumbia

(Coleoptera: Lycidae). - European Journal of

Entomology 97: 271-278.

Bocak L. 2001. New species of the genus Wakarumbia

from Sulawesi (Coleoptera: Lycidae). - The Raffles

Bulletin of Zoology 49, No 2: 259-267.

pp. 253—263

Bocak L., Jass R. 2004. Revision of the genus Broxylus

(Coleoptera: Lycidae). - Deutsche Entomologische

Zeitschrift 51: 65-75.

Bocak L., Matsuda K. 1997. A_ review of the

“Metriorrhynchus thoracicus group” (Coleoptera:

Lycidae). - Giornale Italiano di Entomologia 8:

409-415.

Bocak L., Matsuda K., Yagi T. 2006. A revision of

Metriorrhynchus from the Philippines with

molecular evidence of an Australian origin of

the Oriental Metriorrhynchus fauna (Coleoptera:

Lycidae). - European Journal of Entomology 103:

115-126.

Dvorak M., Bocak L. 2007. Sulabanus gen. nov., a new

genus of Lycidae (Coleoptera) from Sulawesi. -

Zootaxa 1611: 1-24.

DvorakM., BocakL. 2007. Tennewspecies of Wakarumbia

Bocak, 1999 from Sulawesi (Coleoptera: Lycidae),

with a key to males of the genus. - Zootaxa 2282:

51-61.

Kleine R. 1933. Pars 123: Lycidae. In: Schenkling S. (ed.)

Coleopterorum Catalogus. W. Junk, Berlin: 145 pp.

Kubecek V., Dvorak M., Bocak L. 2011. The phylogenetic

structure of Méetriorrhynchini fauna Sulawesi

(Coleoptera: Lycidae) with descriptions of a new

genus, Mangkutanus, and three new species. -

Zoological Studies 50, No 5: 645-656.

Masek M., Motyka M., Kusy D., Bocek M., Li Y., Bocak

L. 2018. Molecular Phylogeny, Diversity and

Zoogeography of Net-Winged Beetles (Coleoptera:

Lycidae). - Insects 9, No 154: 1-18.

Received: 02.vi.2020.

Accepted: 31.x.2020.

263

02-Jun-21 21:50:06

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

264

Book4.indd 264 02-Jun-21 21:50:06

Book4.indd 265

LésL, I.: Contribution to the knowledge of the Scaphisomatini (Coleoptera: Scaphidiinae) of Mindanao, Philippines

pp. 265-272

Contribution to the knowledge of the Scaphisomatini

(Coleoptera: Staphylinidae: Scaphidiinae) of Mindanao,

Philippines

urn:lsid:zoobank.org:pub:CE53/7389-2F24-4F92-A661-/D9OF2ZC5E2D46

IVAN LOBL

Muséum d’histoire naturelle, Case postale 6434, CH-1211, Genéve 6, Switzerland;

lvan.lobl@bluewin.ch

Abstract: An account of Scaphisomatini Casey, 1893 species found in two sites of Mindanao (the Philippines)

is given. Baeocera candalagensis sp. nov., B. glabra sp. nov., B. onerosa sp. nov., Scaphobaeocera lata sp. nov.

and Scaphoxium shavrini sp. nov. are described, and Baeocera wolfgangi LObl| is reported for the first time from

Mindanao. A key to the Philippine species of Scaphoxium LObI is given.

Key words: Scaphisomatini, Philippines, taxonomy, new species, key.

Introduction

Recently, the Philippine Scaphisomatini Casey,

1893 have been dealt with in a few papers (LObI

2011a & b; Lobl & Ogawa 2016; LObI 2018) that

point to unexpected richness of the group on the

archipelago biotas. Nevertheless, the knowledge of

these mycophagous and myxomycetophagous rove

beetles remains still quite inadequate, at least as far

as the fauna of most Philippine Islands concerned.

This is also highlighted by a new collection of

Mindanao Scaphisomatini received from Alexey V.

Shavrin. The material comprises eleven species

sampled in two sites, with five of the species being

new to science and described below.

Material and methods

The material studied is deposed in the following

collections:

MHNG - Muséum d’histoire

Switzerland;

PCAS - Collection Alexey Shavrin, Daugavpils, Latvia.

naturelle, Geneve,

The locality data of the type specimens are

reproduced verbatim. Data from different labels

are separated by a slash. The body-length is

measured from the anterior pronotal margin to the

posterior of inner angles of elytra. As interocular

width is given the shortest interval between the

eyes, seen in dorsal view. The length / width ratios

of the antennomeres are measured on slide-

mounted antennae. The abdominal microsculpture

refers to the exposed segments, and not to the

intersegmental membranes. The sides of the

aedeagi refer to their morphological side with

the ostium situated dorsally, while it is in resting

position rotated 90°. The dissected body-parts are

embedded in Euparal and fixed on a separate card

on the same pin as the respective specimen.

Results

Baeocera Erichson, 1845

Currently, 20 species of Baeocera are known

to occur in the Philippines, only four of them have

been described or reported from Mindanao. Three

of them are in the new collections, together with

additional three new species.

Baeocera brunnea (Lobl, 1972)

Material examined. 2¢ & 42 PCAS & MHNG:

Mindanao, Araibo, Pantukan, Compostela Valley, 900

m;. Gandalaga /- Mis;..'7 716'85.3°N,-126510'12.8".E,

4.5.2019 Shavrin A. V. leg.

Distribution: This species has yet been reported

from the Philippine islands Leyte, Luzon, Mindanao

and Siargao.

265

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TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Baeocera candalagensis sp. nov. (Figs 1-4)

Holotype @ MHNG: Philippines Mindanao. Araibo.

Pantukan. Compostela Valley. 900 m, Candalaga / Mts.,

#216'35:.3"'N, 226°10'12:8"E,.4.5:20179 Shavrin AcV.

Derivatio nominis: The species epithet is an

adjective derived from the name of the mountains

where it was found.

Description: Length 1.48 mm, width 1.10 mm.

Frons and body very dark brown, almost blackish,

apex of abdomen ochraceous, femora and tibiae

reddish-brown, tarsi and antennomeres | to VI

ochraceous, following antennomeres light brown.

Body strongly convex dorsally. Eyes in dorsal view

slightly larger than half of interval between them.

Length/width rations of antennomeres as: III 25/8:

IV 25/8: V 30/8: VI 26/8: VIl 45/11: VII 44/11: IX

45/14: X 44/14: Xl 51/14. Pronotum and elytra not

microsculptured, with lateral contours separately

arcuate. Pronotum very finely punctate, with lateral

margins rounded, lateral margin carinae concealed

in dorsal view. Apex of scutellum exposed. Elytra

not overlapping abdominal apex, moderately

narrowed apically, lateral margin carinae not visible

in dorsal view, Sutural striae entire, parallel in basal

third of sutural length, converging posteriad, bent

along pronotal lobe and extended along bases

to form complete basal striae joined with lateral

Striae. Sutural striae distinctly punctate, except

along apical third, punctures along lateral striae

almost indistinct. Basal striae deep and well visible

near lateral margins. Punctation on basal 0.15 to

0,20 mm, on 0.15 mm broad stripe along sutural

striae, on narrow stripe along lateral margins

and on apical halves of elytra fine, very shallow

and not well delimited, with punctures distinctly

larger than pronotal punctures. Conspicuous

patch of coarse and sharply delimited punctures

situated in anterior half of each elytron; intervals

between coarse punctures about 1.5 to two times

puncture diameters. Hind wings not reduced.

Hypomera extremely finely punctate. Mesepimera

about four times as long as wide and four times

as long as intervals to mesocoxae. Metaventrite

convex and impunctate in middle, with several

coarse punctures delimiting smooth centre; lateral

parts of metaventrite extremely finely punctate.

Mesocoxal lines parallel, with coarse puncture

row extended laterally almost to mid-length of

mesepimera; mesocoxal areas about 0.03 mm, as

fifth of shortest intervals to metacoxae. Exposed

part of metanepisterna large, near anterior angles

about 0.10 mm wide, narrowed posteriad, with

distinct, impunctate suture. Metepimera each with

266

longitudinal stria. Protibiae straight, mesotibiae

and metatibiae slightly bent. Propygidium and

ventrite | not microsculptured, pygidium and

ventrites Il to VI with hardly visible punctulate

microsculpture. Basal half of propygidium densely

and rather coarsely punctate, with punctures well

delimited, smaller than coarse elytral punctures,

to part about as large as puncture intervals. Apical

part of propygidium and pygidium very finely and

sparsely punctate. Ventrite | without wrinkles,

with basal row of coarse elongate punctures not

interrupted in middle; punctation extremely fine

posterior of basal row, as on following ventrites.

Male protarsomere | strongly widened, somewhat

narrower than apices of protibiae, Protarsomere II

and Ill slightly widened. Mesotarsomeres | slightly

widened. Ventrite VI shallowly emarginate at each

side of triangular, about 0.10 mm long and acute

median process. Aedeagus (Figs 1-4) 0.87 mm

long.

Differential diagnosis: This new species falls

in the key to Philippine species (LObI, 2012) under

the couplet 5, to B. alticola LObl, 2012 and B.

fortis LObl, 2012, its aedeagal characters suggest

relationships with B. fortis. The new species is

readily distinguishable from B. fortis by its smaller

size, the darker body colour, the antennomeres

lll as long as the antennomeres IV and the

antennomeres V clearly longer than antennomeres

IV or VI, the shape of the apical margin of the male

ventrite VI, the parameres widened apically and

lacking lobes or processes, the blunt and in dorsal

view not incurved tip of the median lobe, and the

shape of the sclerotized pieces of the internal sac.

Diagnostic for the new species are also tufts of

elongate spine-like and lamellar structures within

the internal sac.

Baeocera glabra sp. nov. (Figs 5-7)

Holotype 3 MHNG: Mindanao, Araibo, Pantukan,

Compostela Valley, 900 m, Candalaga / Mts.,

7°16'35.3"'N, 126°10'12.8’E, 4.5.2019 Shavrin A.

Paratypes 36 PCAS & MHNG: data as the holotype.

Derivatio nominis: The species epithet is a Latin

adjective meaning glabrous.

Description: Length 0.88-0.91 mm, width 0.63-

0.65 mm. Frons and most of body blackish, apex

of abdomen, femora and tibiae reddish-brown,

tarsi and antennae ochraceous to yellowish. Body

strongly convex dorsally. Bode strongly convex

dorsally. Eyes in dorsal view slightly smaller than

half of interval between them. Length/width

rations of antennomeres as: Ill 16/5: IV 17/5:

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Book4.indd 267

LdsL, I.: Contribution to the knowledge of the Scaphisomatini (Coleoptera: Scaphidiinae) of Mindanao, Philippines

V 18/6: VI 17/5: VII 24/8: VIIl 18/5: IX 25/10:

X 25/10: Xl 34/10. Pronotum and elytra not

microsculptured, with lateral contours continuously

arcuate. Pronotum very finely punctate, with

lateral margins rounded, lateral margin carinae

concealed in dorsal view. Scutellum concealed.

Elytra not overlapping abdominal apex, moderately

narrowed apically, lateral margin carinae not visible

in dorsal view, Sutural striae entire, parallel, bent

along pronotal lobe and extended along bases to

form complete basal striae joined to lateral striae.

Adsutural areas and sutural striae with few rather

distinct fine puncture, discal punctation very

fine, as that on pronotum, hardly visible at 100

times magnification. Hind wings fully developed.

Hypomera appearing impunctate. Mesepimera

about three times as long as wide and somewhat

more than two times as long as intervals to

mesocoxae. Metaventrite convex and impunctate

in middle, with coarse punctures delimiting smooth

centre; lateral parts of metaventrite smooth near

metacoxae, coarsely and irregularly punctate on

Why.

GIT Z

EMT G TEL

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27)

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pp. 265-272

remaining surface. Mesocoxal lines slightly convex,

with coarse puncture row extended laterally almost

to mesepimera; mesocoxal areas about 0.02

mm, as third of shortest intervals to metacoxae.

Exposed part of metanepisterna narrow, parallel,

about 0.04 mm wide, with distinct, punctate suture.

Metepimera each with longitudinal stria. Tibiae

Straight. Ventrite | without wrinkles, with basal

row of coarse, to part slightly elongate punctures

not interrupted in middle, punctation posterior of

basal row extremely fine, hardly visible at LOO times

magnification, as that on following ventrites. Male

protarsomere | to Ill slightly widened. Aedeagus

(Figs 5-7) 0.21-0.24 mm long.

Differential diagnosis: This new species is a

member of the B. lenta species-group and is similar

to B. danielae Lobl, 2012. It may be distinguished

by the elytra entirely very finely punctate, in

combination with the minute body-size and the

internal sac lacking membranous denticles and

bearing a proximally widened sclerite (Seen in

lateral view).

Figures 1-4. Baeocera candalagensis sp. nov. 1 - Aedeagus in lateral view [scale bar 0.2 mm]; 2 - Apical part of

aedeagus [scale bar 0.1 mm]; 3 - Internal sac in dorsal view [scale bar 0.1 mm]; 4 - Parameres in ventral view,

[scale bar 0.1 mm].

267

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Baeocera hypomeralis Lobl, 2012

Material examined. 3 ¢ & 3 @ PCAS & MHNG:

Mindanao, Araibo, Pantukan, Compostela Valley, 900

m, Candalaga / Mts., 7°16’35.3’"N, 126°10'12.8”E,

4.5.2019 Shavrin A. V. leg.

Comments. This species is known only from

Luzon and Mindanao.

Baeocera onerosa sp. nov. (Figs 8-10)

Holotype d MHNG: Mindanao, Araibo, Pantukan,

Compostela Valley, 900 m, Candalaga / Mts.,

7 °16’35.3’N, 126°10'12.8’E, 4.5.2019 Shavrin A. V.

leg.

Paratypes 8d & 69 PCAS & MHNG: same data as in

holotype.

Derivatio nominis: The species epithet is a Latin

adjective meaning difficult.

Description: Length 1.08-1.18 mm, width 0.71-

0.81 mm. Frons and body blackish, with apical

abdominal segments light, ochraceous to yellowish.

Femora and tibiae dark reddish-brown, tarsi and

antennae light, nearly yellowish. Body strongly

convex dorsally. Eyes in dorsal view as large as half

of interval between them. Length/width rations

of antennomeres as: Ill 28/6: IV 21/6: V 30/6: VI

28/6: VII 34/8: VIII 33/7: IX 35/10: X 35/13: Xl

40/13. Pronotum and elytra not microsculptured,

with lateral contours nearly continuously arcuate.

Pronotum very finely punctate, with lateral margins

rounded, lateral margin carinae concealed or hardly

visible in dorsal view. Scutellum concealed. Elytra

not overlapping abdominal apex, strongly narrowed

apically, lateral margin carinae visible only near

bases in dorsal view, sutural striae entire, parallel

in basal half of sutural length, converging posteriad

mid-length, bent along pronotal lobe and extended

along bases to form complete basal striae joined

with lateral striae. Anterior third of Sutural and lateral

Striae distinctly punctate. Basal striae very shallow

and hardly visible near lateral margins. Punctation

coarse and sparse on basal third of elytral disc,

except on narrow band along base and near

sutural striae, intervals between coarse punctures

mostly about 1.5 to 2 times as large as puncture

diameters. Punctation on remaining surface of

elytra very fine and sparse. Hind wings not reduced.

Hypomera appearing impunctate. Mesepimera

about four times as long as wide and three times

as long as intervals to mesocoxae. Metaventrite

flattened in middle, with impunctate mesal stripe,

remaining mesal and entire lateral surfaces

coarsely and very densely punctate, punctures

mostly larger than puncture intervals. Mesocoxal

lines parallel, mesocoxal areas about 0.02 mm, as

—_—_

268

fourth of shortest intervals to metacoxae. Exposed

part of metanepisterna very narrow, with suture

indicated by outer coarse punctures. Metepimera

without longitudinal striae. Tibiae straight. Ventrite

| without wrinkles, with coarse row of basal

punctures narrowly interrupted in middle and

several scattered rather large punctures on sides;

remaining abdominal punctation very fine. Male

protarsomere | to Ill slightly widened. Aedeagus

(Figs 8-10) 0.33-0.35 mm long.

Differential diagnosis: This new species is

also a member of the B. lenta species-group and

falls under the couplet 13 in the key to Philippine

Baeocera (LObI 2012). Its aedeagal characters,

especially the complex sclerotized parts of the

internal sac, differ drastically from both species

under this couplet, B. /ouisi Lobl, 2012 and B.

danielae Lobl, 2012. Baeocera onerosa sp. nov.

may be readily distinguished from these two

species by the much dense punctation on lateral

parts of the metaventrite and by several rather

large punctures on lateral parts of the abdominal

ventrite |, situated posterior of basal puncture row.

The internal sac of B. onerosa sp. nov. is similar to

that of B. boettcheri (L6bl, 1972) while the shape

of the parameres and the elytral punctation differ

drastically.

Baeocera wolfgangi Lobl, 2012

Material examined. 2 PCAS & MHNG: Mindanao,

Davao Prov., Mt. Talomo (Mt. Apo) Catigan, / 800-1000

Mm, 7 OL 27°0" Nj) 125-°29430'5."E, 29;4.-1.5. 2019, AN.

Shavrin leg.; 64 & 49 PCAS, MHNG: Mindanao, Araibo,

Pantukan, Compostela Valley, 900 m, Candalaga / Mts.,

f-16°35.37 N/126°10'12°8E-4.5.2019,-Shavrin-A. V.

leg.

Distribution: The species was known only from

the island Leyte.

Birocera Lobl, 1970

This genus comprises three species, with B.

derougemonti Lobl, 1983 appearing restricted to

Sulawesi (Greater Sunda Islands, Indonesia), B.

punctatissima (Reitter, 1880) known both from the

Philippines and Sulawesi, and B. basicollis LObl,

2011 described from the Philippine specimens.

Birocera basicollis Lobl, 2011

Material examined. 19 PCAS: Mindanao, Araibo,

Pantukan, Compostela Valley, 900 m, Candalaga / Mts.,

7°16'35.3’'N, 126°10'12.8”E, 4.5.2019 Shavrin A. V.

leg.

02-Jun-21 21:50:07

LésL, I.: Contribution to the knowledge of the Scaphisomatini (Coleoptera: Scaphidiinae) of Mindanao, Philippines

pp. 265-272

Distribution: This species is currently known (2011a). Only one of them, B. rufescens LObl, 1972,

from Luzon and Mindanao, the Philippines. is Known to occur in Mindanao, the Philippines.

Bironium Csiki, 1909 Bironium rufescens Lobl, 1972

Material examined. 1d & 19 PCAS & MHNG:

Mindanao, Davao Prov., Mt. Talomo (Mt. Apo) Catigan,

800-1000 m, 7°01’21.0’N, 125° 22'30.5”E, 29.4.-1.5.

2019, A.V. Shavrin leg.

This genus is with 31 species widely distributed

in the Oriental region and in Melanesia. The four

Philippine species of Bironium were keyed in LObl

Figures 5-10. Aedeagi in Baeocera. 5-6 - B. glabra sp. nov., dorsal (5) and lateral (6) view [scale bar 0.1 mm];

7 - ditto, apical part of aedeagus [scale bar 0.05 mm]; 8-10 - B. onerosa sp. nov.; 8 & 10 - Aedeagus, dorsal (8)

and lateral (10) view [scale bar 0.1 mm]; 9 - Internal sac [scale bar 0.05 mm].

Yo & »~

02-Jun-21 21:50:07

Book4.indd 269

Book4.indd 270

TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Distribution: This species is reported from Leyte

and Mindanao in Lobl et a/. (2020). Unlike other

examined specimens of B. rufescens, both are

black.

Scaphisoma Leach, 1815

Scaphisoma comprises 112 species reported

from the Philippines (Lobl & Ogawa 2016) and is

the far most species-rich genus of the scaphidiines.

A single species is present in the recent material.

Scaphisoma laminatum Lobl, 1972

Material examined. 14 & 29 PCAS & MHNG:

Mindanao, Davao Prov., Mt. Talomo (Mt. Apo) Catigan, /

800-1000 m, 7°01’21.0’N, 125° 22’30.5’E, 29.4.-1.5.

2019, A.V. Shavrin leg.; 3' PCAS & MHNG: Mindanao,

Araibo, Pantukan, Compostela Valley, 900 m, Candalaga

7 Miss. e7 M6353" NY 26510 12:6 Ey “4.52019

Shavrin A.V. leg.

Distribution: The species has been reported

from Leyte, Luzon, Mindanao and Panay.

Scaphobaeocera Csiki, 1909

The Philippine species were dealt with and

keyed in LOobl (2011b). From the 18 currently

recognized Philippine species, only four are known

to occur on Mindanao. An additional, new species

iS present in the examined collections.

Scaphobaeocera /ata sp. nov. (Figs 11-12)

Holotype MHNG: Mindanao, Araibo, Pantukan,

Compostela Valley, 900 m, Candalaga / Mts.,

7°16’35.3’N, 126°10'12.8"E, 4.5.2019 Shavrin A.V.

leg.

Paratype @ PCAS: Mindanao, Davao Prov., Mt. Talomo

(Mt. Apo) Catigan, / 800-1000 m, 7°01’'21.0’N,

125° 22%30.5’E, 29.4.-1.5. 2019, A.V. Shavrin leg.

Derivatio nominis. The species epithet is a

Latin adjective meaning wide, referring to the wide

parameres.

Description: Length 1.35 mm, width 0.69

mm, dorsoventral diameter 0.71 mm. Head and

body dark brown with somewhat reddish shine,

apical abdominal segments and appendages

lighter, reddish-brown. Length/width rations of

antennomeres as: Ill 17/7: IV 22/5: V 27/5: VI

22/7: Vil 33/9: Vill 18/8: IX 30/19: X 33/14: XI

40/14. Thorax not microsculptured, pronotum very

finely punctate. Tip of scutellum exposed. Elytra

with strigulate microsculpture hardly visible at 100

times magnification, not iridescent, with punctation

270

similar with that of pronotum; sutural striae starting

near pronotal lobe, parasutural striae hardly visible.

Hypomeron without longitudinal stria. Middle part

of metaventrite impressed, lacking stria, densely

and finely punctate and bearing short pubescence

on posterior half, impunctate in middle of anterior

half. Sides of metaventrite sparsely, extremely fine

punctate, with fairly long pubescence. Mesocoxal

lines with row of fine punctures not exceeded

laterally along mesepimera; mesocoxal areas about

0.04 mm long, almost as fourth of shortest intervals

to metacoxae. Metanepisterna flat, 0.06 mm wide,

narrowed apically, with curved suture. Tibiae

straight. Abdomen with strigulate microsculpture

and very finely punctate. Basal punctures of ventrite

| rather coarse. Male segments | to III of protarsi

distinctly widened, segment | narrower than apex

of protibia. Aedeagus (Figs 11-12) 0.42 mm long.

Differential diagnosis: The aedeagal characters

Suggest relationships of this new species with the

Philippine S. episternalis Lob|, 2011, though it would

fall under the couplet 18 in the key to the Philippine

species (LObI, 2011b). It differs drastically from S.

episternalis by the much narrower metanepisterna,

darker body, not iridescent elytra, and much wider

parameres. The two species given under the

couplet 18, S. pseudotenella Lobl, 2011 and S.

data Lobl, 2011, may be easily distinguished by

the shape of the internal sac. Prominent articular

processes are present in several congeners, such

as S. cognata Lobl, 1984, S. difficilis Lobl, 1979, S.

jirkai LOblI, 2018, S. ornata (Pic), S. papuana Csiki,

1909, S. stipes Lobl, 1971, S. tenella Lobl, 1990

and S. uncata Lobl, 1990. These species differ

conspicuously from S. lata sp. nov. either by the

Shape of the apical process of the median lobe or

by the narrow basal section of the parameres (Seen

in lateral view).

Scaphoxium Lobl, 1979

Currently, two species of Scaphoxium are

known from the Philippines, S. alesi Lobl, 2011

from Luzon and S. taylori Lobl, 1981 described

from Sarawak and subsequently reported from the

Philippine islands Luzon, Mindanao and Palawan.

The new collections yielded an additional species.

Key to the Philippine species of Scaphoxium

1 Antennomeres XI about 1.5 times as long as

antennomeres X. Sides of metaventrite with coarse

punctures

02-Jun-21 21:50:07

Book4.indd 271

LdsL, I.: Contribution to the knowledge of the Scaphisomatini (Coleoptera: Scaphidiinae) of Mindanao, Philippines

- Antennomeres XI about as long or slightly longer

than antennomeres X. Sides of metaventrite very finely

UNI CLANG: s sssiccsstesgsasanmeassandeeenesebeosacatessenavetbes S. taylori LObl|

2 Sides of metaventrite with few coarse punctures.

Antennomeres VIIl about 4 times as long as wide.

Internal sac of aedeagus lacking sclerotized denticles ..

Jaber ALLELE tsntacnee reese NN ateee eeerd Mane NOM Mehderd S. alesi LObl

- Sides of metaventrite with dense patch of coarse

punctures. Antennomeres VIII about 2.5 times as long as

pp. 265-272

wide. Internal sac of aedeagus with strongly sclerotized

GS NGI SS a sick atrristandtantaeindidunneruaeduaaennt S. shavrini sp. nov.

Scaphoxium shavrini sp. nov. (Figs 13-15)

Holotype 6 MHNG: Mindanao, Araibo, Pantukan,

Compostela Valley, 900 m, Candalaga / Mts.,

7°16'35.3’"N, 126°10'12.8”"E, 4.5.2019 Shavrin A.V.

leg.

Figures 11-15. Scaphidiid aedeagi. 11-12 - Scaphobaeocera lata sp. nov., dorsal (11) and lateral (12) view [scale

bar 0.1 mm]; 13-15 - Scaphoxium shavrini sp. nov. 13 - Aedeagus, dorsal view [scale bar 0.1 mm]; 14 - Parameres,

ventral view [scale bar 0.1 mm]; 15 - Internal sac [scale bar 0.1 mm].

201

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Derivatio nominis: Patronymic. The species

is dedicated to its collector Alexei V. Shavrin

(Daugavpils, Latvia), a well-known expert of rove

beetles.

Description: Length 1.54 mm, width 0.77 mm,

dorsoventral diameter 0.86 mm. Head and most

of body evenly very dark reddish-brown, apical

abdominal segments, femora and tibiae lighter,

tarsi lighter than tibiae, antennae brown. Length/

width rations of antennomeres as: III 20/6: IV

19/62 V° 26/62 4.22772 NILSO/LI= Vill) 25/222. (xX

38/13: X 33/13: Xl 48/15. Pronotal punctation

very fine, hardly visible at 50 times magnification.

Scutellum concealed. Elytral punctation distinct,

fine and dense, sutural striae starting about

0.20 mm posterior of level of pronotal lobe.

Mesoventrite not striate, impressed and with

low mesal ridge narrowed apically. Median part

of metaventrite slightly convex, with irregular

transverse row of extremely fine punctures. Lateral

parts of metaventrite appearing impunctate

near metacoxae, each with dense patch of

conspicuously coarse punctures and oblique ridge.

Mesocoxal areas about 0.05 mm, almost as halves

of shortest intervals to metacoxae. Metanepisterna

slightly convex, about 0.06 mm wide, with curved,

deep, apically shortened sutures. Ventrite | with

few distinct basal punctures, not microsculptured.

Following ventrites and exposed tergites with

punctulate microsculpture and extremely fine

punctation. Male segments | to Ill of protarsi slightly

widened. Aedeagus (Figs 13-15) 0.68 mm long.

Differential diagnosis: This new_ species

is likely related with S. alesi LObl, 2011. It may

be distinguished by the dark body-colour, the

shorter sutural striae of the elytra, the sides of the

metaventrite bearing dense and coarse punctures,

the shape of the angular subapical parameral lobes

and the presence of strongly sclerotized denticles

in the internal sac of the aedeagus.

272

Acknowledgements

My cordial thanks are due to Alexey Shavrin

(Institute of Life Sciences and_ Technology,

Daugavpils University, Daugavpils, Latvia) for

providing the studied material and for his generous

gift of soecimens to the MHNG.

References

Lobl |. 2011a. On the Scaphisomatini (Coleoptera:

Staphylinidae: Scaphidiinae) of the Philippines. -

Studies and Reports, Taxonomic Series 7: 301-

314.

Lobl |. 2011b. On the Scaphisomatini (Coleoptera:

Staphylinidae: Scaphidiinae) of the Philippines, II.

— Revue suisse de zoologie 118: 695-721.

Lobl |. 2012. On the Scaphisomatini (Coleoptera:

Staphylinidae: Scaphidiinae) of the Philippines, III:

the genus Baeocera Erichson. - Revue suisse de

zoologie 119: 351-383.

Lobl |. 2018. Supplement to the knowledge of

the Philippine Scaphisomatini (Coleoptera:

Staphylinidae: Scaphidiinae). - Baltic Journal of

Coleopterology 18, No 1: 97-102.

Lobl |, Ogawa R. 2016. On the Scaphisomatini

(Coleoptera: Staphylinidae: Scaphidiinae) of the

Philippines, IV: the genera Sapitia Achard and

Scaphisoma Leach. - Linzer biologische Beitrage

48, No 2: 1339-1492.

Lobl |., Leschen R. A. B., Kodada J. 2020. Review of

Asian species and cladistic analysis of Bironium

Csiki (Coleoptera: Staphylinidae: Scaphidiinae) with

comments on biogeography. - Annales Zoologici

70, No 4: 711-734.

Received: 16.xi.2019.

Accepted: 30.x.2020.

02-Jun-21 21:50:07

Book4.indd 273

Owen, |. L. & Boray, J. C.: Distribution of Orientogalba viridis, the snail intermediate host of Fasciola hepatica ...

pp. 273-280

Distribution of Orientogalba viridis (Quoy & Gaimard,

1832) (Gastropoda: Lymnaeidae), the snail

intermediate host of Fasciola hepatica Linnaeus,

1758, in Papua New Guinea

urn:lsid:zoobank.org:pub:F D566DA1-F1FD-45FC-A46E-B2FOADAEB1A6

IFoR L. Owen & Joserpo C. Boray *

c/o National Veterinary Laboratory, National Agriculture, Quarantine and Inspection Authority,

P.O. Box 741, Port Moresby, Papua New Guinea, now 14 Cwrt Sant Tudno, Clarence Rd., LL30O

1BZ, Llandudno, United Kingdom; iforlowen@gmail.com

* — Deceased

Abstract: In surveys carried out to determine the distribution of Orientogalba viridis (Quoy et Gaimard, 1832) in rivers

of mainland Papua New Guinea, the snail was found only at altitudes above about 600 metres where temperatures

below 25°to 26°C enabled the snail to reproduce. It was common in the Lamari, Asaro and Wahgi rivers that drain

the highland provinces of Eastern Highlands, Simbu and Jiwaka, and in a part of Morobe Province drained by the

Watut River. Its distribution elsewhere was sporadic, being found at isolated sites in other parts of Morobe Province,

and in Western and Central provinces. The snail was not present in streams of Western Highlands, Enga, Hela,

Sandaun or Southern Highlands provinces. Animal infection with the liver fluke, Fasciola hepatica, was common

where the snail had a long-established contact with infected cattle and sheep. Local conditions in the Ramu River

valley led to snails being present at an elevation below 600 meters and infection of cattle with liver fluke, while

unknown factors limit the presence of the snail in the upper reaches of the Wahgi River.

Key words: Host, parasites, cattle, sheep, surveys, river systems, Papua New Guinea.

Introduction

Interest in the distribution of the semi-aquatic

snail, Orientogalba viridis (Quoy et Gaimard 1832)

in Papua New Guinea (further in text - PNG) lies

in its role as intermediate host of the digenetic

trematode, Fasciola hepatica Linnaeus, 1758, the

liver fluke of herbivorous mammals and humans.

Ruminants were unknown in PNG until the late

19th century when cattle were imported to coastal

plantations and some sheep and goats into upland

areas. The majority of the animals were killed

during World War 2 (WW2) (Anderson 1962).

There are no authentic records of F. hepatica being

present in the country before WW2, but staff at the

Catholic Mission at Fane, Central Province, when

enquiries were made in the 1970s, said that liver

fluke was seen in sheep at Ononge Mission prior

to the war. The sheep had been imported from

eastern Australia.

In 1949 attempts were made to establish

sheep from Australia at Nondugl, a highland site

in the then Western Highlands District, now in

Jiwaka Province (Anderson 1972). They were soon

found to be carrying F. hepatica which, due to the

presence of the snail intermediate host, became a

well-established disease. This, together with other

issues, led to the abandonment of the enterprise In

1962 (Anderson 1972) and led to the decline of the

pioneer sheep industry (Talbot 1972).

The PNG administration realised that before

another attempt was made at establishing a sheep

industry, more information was needed about

the snail intermediate host. A preliminary survey

carried out in 1964 by the second author (J.C.B.)

to assess the extent of the snail’s distribution

showed that the snail was not found below about

600 metres. The disease of fascioliasis, therefore,

would be a highland problem. Experimental

work on the PNG snail in Australia found it was a

relatively efficient host of F. hepatica but that it

was unable to reproduce at temperatures above

25°to 26°C, although able to survive temporarily

In temperatures up to 35°C (Boray 1964; 1978).

273

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Up to this time, the PNG snail (Fig. 1) was

regarded as being the same species as occurred in

eastern Australia, Lymnaea tomentosa (L. Pfeiffer,

1855) - now placed in the genus Austropeplea

Cotton, 1942. Later, the PNG snail was shown to bea

separate species, L. viridis Quoy et Gaimard, 1832,

common in east Asian and Pacific countries (Boray

1978; 1982), and was subsequently placed in the

genus Austropeplea. Classification of lymnaeid

Snails, however, has been problematic (Puslednik

et al. 2009), being based largely on morphological

features. Recently, a clearer relationship has

been established within the lymnaeids by means

of molecular taxonomic studies (Aksenova et al.

2018) which place the PNG snail in the genus

Orientogalba Kruglov in Kruglov et Starobogatov,

1985 as O. viridis (Quoy et Gaimard, 1832)

(Aksenova et a/. 2018: Vinarski et al. 2020).

In 1975 the government established a sheep

farm at Menifo, Eastern Highlands Province, to

serve as a distribution centre for village-based

sheep projects. It became apparent that liver fluke

was a problem at the farm as well as in many of the

projects. There was a need for more information on

the distribution of the snail.

Figure 1. Orientogalba viridis (Quoy et Gaimard, 1832)

shell (height 10 mm) from Menyamya, Morobe Province,

Papua New Guinea, apertural view (schematic).

Material and methods

Surveys of rivers in mainland PNG were carried

out over many years (Fig. 2). They focused on parts

of the highlands where cattle or sheep were present

or where there was potential for establishing

projects. Examination centred mainly on shallow

274

margins of streams and creeks, seepages, drains

and swampy ground, the favoured habitats of the

snail. The altitude, hydrogen ion concentration

(OH) and temperature of water were recorded for

most sites. Snails were collected and stored in 10

percent formal saline and sent to the late Dr Boray

in Australia for identification. Information on the

presence of F. hepatica in animals is based on data

contained in Annual Reports of the Department of

Agriculture, Stock and Fisheries, later Department

of Primary Industry, abattoir reports, National

Veterinary Laboratory records and field trials (Owen

1984; 1989).

Results

Although O. viridis was reported to prefer fast-

running streams and deep water of rocky creeks to

Shallow aquatic habitats in PNG (Boray 1969), this

was not experienced during the current surveys. Its

presence at a site, however, could vary over time;

it might be common on one visit but absent on

another, the result, possibly, of flash-flooding. This

could mean the presence of the snail was missed

in some locations where only one or two sites were

examined on a single occasion.

Spatial distribution of O. viridis was

discontinuous (Table 1). It was widespread in the

eastern part of the central highlands, comprising

of Eastern Highlands, Simbu and Jiwaka provinces,

drained by branches of the Lamari, Asaro and

Wahgi rivers that eventually join the south-flowing

Purari River. The snail was common also in that

part of Eastern Highlands Province drained by

headwaters of the Ramu River, that flows north,

and the Wanton River that flows east to join the

Markham River in Morobe Province. A limit in the

distribution of the snail was found in the valley of

the Wahgi River in the Minj / Banz area, Jiwaka

Province. Extensive examination upstream and to

the immediate west of sites in Malenban, Fatima

and Kurumul properties, failed to find the snail. It

is absent, therefore, from the upper reaches of the

Wahgi River that drains much of Western Highlands

Province.

No snails were found near the source of the

Sepik River at Telefomin or in the headwaters of its

tributaries, the Lai, Baiyer and Trauna rivers. The

Snail was absent from waters of the south-flowing

Kikori and Erave rivers. An early report of O. viridis

occurring in the Kagua area of Southern Highlands

Province, drained by the Erave River, is considered

to be a case of mis-identification as extensive

02-Jun-21 21:50:08

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Owen, |. L. & Boray, J. C.: Distribution of Orientogalba viridis, the snail intermediate host of Fasciola hepatica ...

Bismarck Sea

Biaru

Gulf of Papua

Angabanga—"§

Dilava

pp. 273-280

Solomon Sea . t)

Figure 2. Map of the River Systems of mainland Papua New Guinea whose headwaters were examined for the

presence of the snail, Orientogalba viridis (Quoy et Gaimard, 1832).

Legends: PNG provinces: 1 - Sandaun; 2 - East Sepik; 3 - Southern Highlands; 4 - Hela; 5 - Enga; 6 - Western

Highlands; 7 - Jiwaka; 8 - Simbu; 9 - Eastern Highlands; 10 - Madang; 11 - Morobe; 12 - Western; 13 - Gulf;

14 - Central.

searches of the area at different times failed to

find the snail. Hela, Enga and Southern Highlands

provinces, therefore, do not have the snail.

The only record of O. viridis in the Fly /

Strickland river system of Western Province was

at Bultem village near the source of Ok Tedi that

drains the southern slopes of the Star Mountains.

The Tauri River at Menyamya was the only river of

those that enter the Gulf of Papua to the east of the

Purari, to have the snail. Orientogalba viridis was

common in Morobe Province, in branches of the

Watut River, a southern tributary of the Markham

River. It was found also at Wantoat on the Leron

River, a northern tributary of the Markham River,

but present only in the Kwama River at Kabwum

of the fast-flowing streams of the Huon Peninsula.

Records of cattle slaughtered at a highland

abattoir in the 1970s showed that 100 percent of

livers of animals originating from both large and

Small projects in the eastern part of the central

highlands were condemned due to fascioliasis. The

records also revealed that cattle born and bred at

two sites, Gusap and Dumpu, located at elevations

below where the snail would be expected to occur,

were infected with liver fluke. Examination of sites

at the two farms discovered the snail in a shallow

backwater of the main Ramu River at Gusap

(altitude 432 m) (See location in Fig. 3). Examination

of small streams and creeks in the vicinity of both

Gusap and Dumpu failed to find other examples of

the snail.

275

02-Jun-21 21:50:09

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

WAL i

4 Qnonge/kodigeg |

sh cmc 2 . saan

Figure So: Map showing the distribution of the snail, Orientogalba viridis (Quoy et Gaimard, 1832), and the digenetic

trematode, Fasciola hepatica Linnaeus, 1758 in mainland Papua New Guinea. Provinces are numbered as in fig. 2.

Table 1. River systems and locations in mainland Papua New Guinea

examined for the presence of the snail Orientogalba viridis (Quoy et Gaimard, 1832).

Legends: + = present; - = none seen / not present; nd = no data; * = sites visited twice or more times.

River(s) or Locality No. of Altitude, Water Orien- | Other molluscs

tributaries | creek(s) into sites m (ap- temp., °C | togalba | (provisional iden-

which sites examined | prox.) viridis | tification)

drain (No. with

A. viridis

Arau Highlands Planorbinae spp.

Markham | Watut Isano / Sites Morobe 4 1153- 6.0-7.6 | 18.0-20.0 Physastra Sp.

Langimar between 1263

Menyamya &

Aseki

Markham | Watut Snake / Zenag, Mu- Morobe 14% (4) 525- 6.9-7.5 | 19.0-30.8 + Physastra sp.,

i a Cl ll ld

Thiara sp.

1200 Thiara sp.

Markham | Leron Leron Wantoat Morobe 11(6) 1050- 7.5-8.0 | 19.8-22.5 + Physastra sp.,

i a a = dS

Markham | Erap Selep Kisengan Morobe is’ 1518 7.5-8.5 | 20.0-21.0 Physastra sp.,

ia a a el a Ad

Markham | Markham | Markham Waterais, Ut- | Morobe ae 441 (2-0 | 23.5-272 Physastra Sp.,

Pr pee ere [ee

Gabbia sp.

: a

14 (4)

Mindik a ieial Thiara sp.

Kwama Pumune Usugem Kabwum, Morobe 1072- 7.5-8.0 | 17.5-21.0 + Physastra Sp.,

Tipsit 1547 Planorbinae sp.,

Succineidae spp.

276

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Owen, |. L. & Boray, J. C.: Distribution of Orientogalba viridis, the snail intermediate host of Fasciola hepatica ...

pp. 273-280

Table 41 (continuation)

Physastra sp.,

Planorbinae sp.

Thiara sp.,

Planorbinae sp.,

Potamopyrgus sp.

Bullastra lessoni

Physastra sp.

Thiara sp.

Timbe el es 1412- | 8.0-8.5 | 15.5-21.5

1480

Busu ee ici 6 575-909 | 7.5-8.5 | 19.0-22.5

|Morobe__| -27,

Waria Bu-Bu Morobe 19 598-767 | 5.0-6.0 | 20.6-27.0

Ramu Buru / Gusap 15* (1) [432-450] 7.5-7.6 | 23.7-32.5

i sane ila. i al

Bumpu

Ramu Macin 1880

Sele)

sam Highlands 1350

Aiyura Highlands 1734

Physastra sp.

Physatra sp.,

Planorbinae sp.

Unantu Highlands 1680

Highlands

Obura Highlands

susa, Lufa Highlands

Highlands

coe Pee el

3) 1720- | 7.2-7.4 | 17.4-22.0 Physastra sp.

2452

(

Physastra sp.

4) 1403- | 7.5-8.0 | 15.4-31.5

igor

1853

Physastra sp.,

°Physa sp.

Imani Highlands

Henganofi Highlands

Chuave

ole co El

3) 1660- | 7.5-8.0 | 19.0-25.4

1848

3) 1891 7.48.5 | 23.2-31.4

Physastra sp.

Gembogl

Gumine

Physastra sp.

3) APs REST) SRSA AO ne |Z ORG Eis 25)

Oat

iket Planorbinae

sp., limpets

Purari

(Pettancylus or

Ferrissia s

2) 1772- 19.5-20.0

1965

7 S37- 6.0-7.0 | 20.0-22.0

1067

ifs

Purari Wahgi/Tua | Wahgi 2) 1518- 7.0-7.3 | 25.0-26.0

Miunde, Kup 1788

Purari Wahgi/Tua | Minj/Kimil Minj/Banz 30* (10) 1603- | 6.0-7.3 | 19.0-25.0

area 1607

Purari Wahgi/Tua | Aiba Nondugl, iwaka 6*(2) 1588- 6.0-7.0 | 22.6-28.5

Kudjip, Kind- 1676

iwaka

ing Ei

Purari Wahgi/Tua | Minj Jiwaka 19 *(4) 1560- | 6.5-7.5 | 19.0-25.5

Kugark 1570

J Physastra sp.

+ Physastra sp.

limpets

(Pettancylus or

Ferrissia sp.)

Purari Wahgi/Tua | Kilip Kurumul J 18* (7) 1550- 6.5-7.4 | 19.2-30.0

Fatima, 1603

ae a jal ea el

Kempis

Purari Kimil Aviamp, Ko- 6* 20.0-22.0

bam, CLTC 1589

Purari Wahgi/Tua | Gumants / Mt Hagen, Western 40* 1591- | 6.0-7.5 | 18.0 -29.0

lke Kum / Ku- Mt Ambra, Highlands 1818

man Gumants

Purari Erave Ka Kandep, Lon- | Enga 14 2402- 6.0-8.0 | 15.4-28.4

Fee

Valle

Highlands

Physastra sp.

Physastra sp.,

Thiara sp.

Physastra sp.,

Bullastra lessoni

Physastra sp.

TBOS 55.2 cL Ome ly Osos:

Physastra sp.

Purari Erave Kombia Mendi area, | Southern 16* 1495- | 5.5-8.5 | 12.5-31.0

Mendi

Purari Erave Erave Erave Southern 2 1096 6.0-6.5 | 23.5-24.5

a a a a Dl

Sugu Highlands

Purari laro laro lalibu area Southern 2061 5.5-6.5 | 16.0-20.0 Physastra sp.

Highlands

Purari Nama / Poru pect Se ear ere |e 1624 | 6.5-7.0 | 20.0-23.5 ea Physastra sp.

Highlands

Tua

Nebylier / Togoba, Western 4 2567 6.0-7.2 | 11.5-24.5 -

Kaugel Tomba Highlands

Purari

Book4.indd 277 02-Jun-21 21:50:12

Book4.indd 278

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Sepik Yuat Lai / Baiyer / | Baiyer, Western

Trauna Highlands

Sepik Yuat Lai Wapenaman-

da, Tsak Val-

Fl Logaiyu /

Tumbudu /

hlan

Tapini,

Erume, Ke-

rau, Kosipe

maipa

Discussion

As PNG is a tropical country and temperatures

Suitable for O. viridis to reproduce are found only

in the highlands, the only possible explanation

for the presence of the snail at Gusap, is that it is

transported downstream in waters of the Ramu

River from its normal habitat at more than 1260

m in Eastern Highlands Province, to an altitude of

441 m on the main valley floor. Flash-flooding, that

occurs in the highlands, can remove alluvial mud,

and the snails, leading to passive migration and

introducing snails into new environments (Boray

1969). Snails are known also to float and drift with

currents for long distances (Boray 1963).

The temperature of the Ramu River after its

rapid descent from 1260 m at Yonki to 441 m

at the main valley floor was 23°C, while 10 km

278

9 1206

2 ee

Ss 2274 | 6.5-7.0 | 13.0-25.2

2200- | 5.6-7.0 | 14.2-26.0

2666

4 635- 6.0-7.4 | 19.0-21.5

1320

| = =4. oT 1608 | 5.5-6.5 | 22.0-30.0

LZ ;

1240- | 6.0-7.5 | 17.5-22.5

1263

6.8-7.

1335 2-6 mero. 0-23.25

- 750- 0-7.6 | 15.0-21.0

1359

1099- | 7.2-7.6

1464

(1)

(

nea ee ae)

iors s |ModGhr ee i Msp ori. 0)

2063

ie iota i

Avele, Iride 1362

Ononge 1694

Table 41 (continuation)

6.5-7.4 | 18.5-28.0 - Physastra Sp.,

Planorbinae s

Physastra sp

| Physastra sp.__|

Planorbinae sp.

Sastra Sp.

ysastra sp.

ysastra Sp.,

ivalves

1728- | 6.0-7.4 | 17.2-31.0

2217

2036

| 2396 | 7.0-8.0_ | 22.0-22.4 |

es ES Eee

72)

is)

72)

AN)

650 n

aia Physastra sp.,

; en Physastra sp

1209

a a |

eae |

Physastra sp.

Physastra sp.,

Planorbinae sp.

1528

+ Physastra Sp.,

Planorbinae sp.

Physastra sp.,

Thiara sp.,

°Physa sp.

Physastra sp

Thiara sp.

6.0-6.5

Physastra sp.,

Physastra sp.

16.0

19.6

5.5-6.5 | 22.5-28.0

downstream the mid-morning water temperature

in the shallow Gusap backwater, was 25°C under

shade and 27°C in the open. Water temperature

of small creeks flowing into the Ramu River at both

Gusap and Dumpu had a range of 26°C to 32°C;

none carried O. viridis. The presence of empty O.

viridis shells attached to shaded sides of rocks out

of the water at Gusap suggests an environment

not conducive to survival. Nevertheless, as cattle

at Gusap and nearby Dumpu can be infected

with liver fluke, the snail is able to survive and

have the opportunity to liberate the parasite’s

metacercariae on to pasture. A high proportion of

metacercariae are viable and infective for 36 days

at +25°C and for 14 days at +30°C (Boray 1969).

Cattle could also become infected while drinking;

Morley (2015) states that floating metacercariae,

buoyed by small trapped air bubbles, may have an

02-Jun-21 21:50:13

Book4.indd 279

Owen, |. L. & Boray, J. C.: Distribution of Orientogalba viridis, the snail intermediate host of Fasciola hepatica ...

important epidemiological role particularly under

Still or stagnant water conditions, as might occur in

a backwater.

An explanation remains to be found why

O. viridis does not occur further upstream in the

Wahgi River beyond the Minj / Banz area of Jiwaka

Province. The situation has not changed since

monitoring of the area began. There are no physical

barriers and the pH levels and the temperature

range are similar on either side of the Minj / Banz

boundary. It is possible that waters of the upper

Wahgi River in Western Highlands Province contain

mineral elements that inhibit migration of the snail

upstream.

Experimental work conducted in Australia

found the host-parasite relationship between O.

viridis from PNG and the Australian strain of F.

hepatica was less well adapted when compared

with the taxon Austropeplea tomentosa (L. Pfeiffer,

1855) and the fluke (Boray 1978). Infection was

found to curtail the growth of the PNG snail from

the normal size of 10-12 mm to 7 mm, under

experimental conditions (Boray 1978). However, the

relationship is sufficiently well adapted under field

conditions in PNG for pasture to be contaminated

with enough larvae for cattle, known to have

greater resistance against F. hepatica than sheep

(Boray 1969; Rickard & Howell 1982), to become

heavily infected, as seen from abattoir records and

field trials (Owen 1984). Preventing the spread of

F. hepatica beyond its current distribution (Fig. 3)

necessitates selecting animals only from fluke-

free locations for distribution to sites such as

Menyamya, where the snail occurs. Treatment of

fluke-infected animals with a course of fasciolicide

before movement is, under PNG conditions, a less

certain way of ensuring transferred animals are

free of the fluke.

No animals at Ononge Mission (Vanapa River

system), Central Province (Fig. 3), have been

found infected with F. hepatica after WW2, but

unconfirmed reports in later years of infected cattle

from nearby Kodige (Dilava River system) in the

Fane area of the province, where O. viridis was not

found, are unresolved, and the provenance of the

animals could not be fully ascertained.

There are no records of humans or of goats,

pigs and other susceptible animals in PNG, being

infected with liver fluke.

Bullastra lessoni (Deshayes, 1830), another

lymnaeid snail encountered in the surveys, is an

unsuitable intermediate host for F. hepatica (Boray

1978).

pp. 273-280

Acknowledgements

Grateful thanks go to departmental livestock

officers and villagers for the assistance given to ILO

while undertaking the surveys and to staff at

Fane, Kamulai and Menyamya_ Catholic

Missions (PNG) for their cooperation in supplying

information.

References

Aksenova O. V., Bolotov |. N., Gofarov M. Y., Kondakov

A. V., Vinarski M. V., Bespalaya Y. V., Kolsova Y. S.,

Palatov D. M. Sokolova S. E., Spitsyn V. M., Tomilova

A. A. Travina O. V., Vikhrev I. V. 2018. Species

richness, molecular taxonomy and biogeography of

the radicine pond snails (Gastropoda: Lymnaeidae)

in the Old World. - Scientific Reports 8: (11199)

1-17.

Anderson J. L. 1962. The development of the cattle

industry in the Territory of Papua and New Guinea.

—- Papua and New Guinea Agricultural Journal 14:

133-140.

Anderson J. L. 1972. Livestock: 645-646. In: Talbot

N. T. (ed.) Encyclopaedia of Papua and New

Guinea. Volume 2. Melbourne University Press, in

collaboration with the University of Papua and New

Guinea, Victoria, Australia: 589-1231.

Boray J. C. 1963. The ecology of Fasciola hepatica with

particular reference to the intermediate host in

Australia. - Proceedings of the 17 World Veterinary

Congress, Section 6: 709-715.

Boray J. C. 1964. Studies on the ecology of Lymnaea

tomentosa, the intermediate host of Fasciola

hepatica. 1. History, geographical distribution, and

environment. - Australian Journal of Zoology 12:

217-230.

Boray J. C. 1969. Experimental fascioliasis in Australia.

- Advances in Parasitology 7: 96-210.

Boray J. C. 1978. The potential impact of exotic Lymnaea

spp. on fascioliasis in Australasia. - Veterinary

Parasitology 4: 127-141.

Boray J. C. 1982. Molluscan hosts and trematodes in the

Pacific Basin: 81-106. In: Symons L. E. A., Donald

A. D., Dineen J. K. (eds) Biology and Control of

Endoparasites. Academic Press, Sydney, London &

New York: 416 pp.

Cotton B. C. 1942. Some Australian freshwater

Gastropoda. - Transactions of the Royal Society of

South Australia 66: 75-82.

Deshayes G. P. 1830. Lymnaeus lessoni. - Magasin de

Zoologie 1: Classe V. Mollusques: 46.

Kruglov N. D., Starobogatov Y. |. 1985. The volume of the

subgenus Galba and of the other similar subgenera

of the genus Lymnaea (Gastropoda, Pulmonata). -

Zoologicheskiy Zhurnal 64: 24-35 (in Russian).

279

02-Jun-21 21:50:13

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Linnaeus C. 1758. Systema naturae per regna tria

naturae, secundum classes, ordines, genera,

species, cum characteribus, differentiis, synonymis,

locis. 10“ edition, 1, Holmiae: 823pp.

Morley N. J. 2015. Ecology of free-living metacercariae

(Trematoda). - Advances in Parasitology 89: 1-78.

Owen |. L. 1984. Production effects of the liver fluke

Fasciola hepatica on weaner cattle in Papua New

Guinea. - Tropical Animal Health and Production

16: 158-160.

Owen |. L. 1989. The epidemiology of fascioliasis in

Papua New Guinea. - Australian Veterinary Journal

66: 58-60 & 127.

Pfeiffer L. K. G. 1855. Descriptions of fifty-seven new

species of Helicea from Mr. Cumings’ collection.

- Proceedings of the Zoological Society London

1854: 286-298.

Puslednik L., Ponder W. F., Dowton M., Davis A. R.

2009. Examining the phylogeny of the Australasian

Lymnaeidae (Heterobranchia: Pulmonata:

Gastropoda) using mitochondrial, nuclear and

morphological markers. - Molecular Phylogenetics

and Evolution 52: 643-659.

280

Quoy J. R., Gaimard J. 1832. Zoologie, Mollusques.

Volume 2. Voyage de Découverte de |’Astrolabe

exécuté par Ordre du Roi, Pendant les Années

1826-1829, sous le commandement de M.J.

Dumont-d’Urville. J. Tastu, Paris: 320 pp.

Rickard M. D., Howell M. J. 1982. Comparative aspects

of immunity in fascioliasis and cysticercosis in

domesticated animals: 343-373. In: Symons L.

E. A., Donald A. D., Dineen J. K. (eds) Biology and

Control of Endoparasites. Academic Press, Sydney,

London & New York: 416 pp.

Talbot N. T. 1972. Livestock pests: Internal parasites:

649-650. In: Talbot N. T. (ed.) Encyclopaedia of

Papua and New Guinea. Volume 2. Melbourne

University Press, in collaboration with the University

of Papua and New Guinea, Victoria, Australia: 589-

12734.

Vinarski M. V., Aksenova O. V., Bolotov I. N. 2020.

Taxonomic assessment of genetically-delineated

species of radicine snails (Mollusca, Gastropoda,

Lymnaeidae). - Zoosystematics and Evolution 96:

577-608.

Received: 23.v.2020.

Accepted: 10.vii.2020.

02-Jun-21 21:50:13

Owen, I. L.: A brief review of animal parasites recorded in Western Province, Papua New Guinea

pp. 281-327

A brief review of animal parasites recorded in Western

Province, Papua New Guinea

urn:lsid:zoobank.org:pub:BF49EOFO- 7256-415 /7-88CC-O4A62BAEF 719

IFOR L. OWEN

c/o National Veterinary Laboratory, National Agriculture Quarantine and Inspection Authority,

P.O. Box 741, Port Moresby, Papua New Guinea, now 14 Cwrt Sant Tudno, Clarence Rd., LL3O

1BZ, Llandudno, United Kingdom; iforlowen@gmail.com

Abstract: Papua New Guinea’s Western Province has a rich biodiversity of both animal hosts and parasites. In the

text, hosts are arranged into three groups: non-native domestic animals, long-established domestic animals, and

wildlife, while they are listed systematically in a table. The range of animals in the first and second groups is small

and harbours mainly parasites commonly found in domestic animals. The wildlife group contains an extensive range

of host animals and parasites. Rusa deers are unusual, as herding animals, in having no nematode parasites. Wild

pigs, besides having nematodes commonly found in suids, carry a zoonotic nematode in the musculature and one,

unique to the country, in the tongue. Digenetic trematodes are almost the only parasites recorded from dugongs.

Amongst 12 species of bats, virtually no endoparasites are recorded in frugivores but ectoparasites were present

on both frugivores and insectivores. Fleas were the main ectoparasites of the 24 species of rodents identified to

species, particularly in the north of the province, while nematodes were the predominant endoparasites. Twenty-one

species of marsupials, identified to species, were examined. Nematodes formed the majority of the endoparasites

of wallabies, while the smaller marsupials (bandicoots, dunnarts and possums) harboured a relatively small range

of ecto- and endo- parasites. Thirty-six bird species were examined, with ectoparasites (chewing or feather lice in

particular) being more frequently found than endoparasites. One case of brood parasitism by a bird is recorded,

as well as two species of arthropods (a louse and a mite), normally parasites of mammals, infesting a bird (a

cassowary). Parasites of lower vertebrates have received little study and consequently only a small number are

recorded from reptiles, amphibia and fishes. Some parasites and parasitoids, mostly insects and brood parasitizing

birds, with no named hosts, are included in a separate table. Some species of rodents, marsupials and reptiles,

together with their parasites, occur in both Western Province and northern Australia and is a legacy of the physical

connection that linked the localities in the past.

Key words: Hosts, domestic animals, wildlife, ectoparasites, endoparasites, helminths, arthropods, annelids, New

Guinea.

Book4.indd 281

Introduction

Parasites are an integral part of nature.

They can play a part in the process of evolution

through their influence on their hosts and their

own adaptation to the environment which they

have adopted. Although parasites of wildlife, under

natural conditions, seldom appear to have obvious

harmful effects on their hosts, nevertheless, they

probably participate in the ‘survival of the fittest’

by helping to weed-out the weakest members of

the host species. As can be seen in this account,

scarcely any of the animals examined, with the

exception of a few marsupials, harboured sufficient

numbers of parasites that would likely affect their

well-being. The exceptions were wallabies and

pademelons in which large numbers and a variety

of species of worms were found. Such numbers are

most often associated with domestic animals when

they are confined to limited space which becomes

heavily contaminated with infective stages of

internal parasites or when animals are kept in close

contact which facilitates the spread of parasitic

arthropods. The habit, at least of agile wallabies,

of grazing in groups may be part of the explanation,

but the host-parasite association suggests it to be a

well-established and well-tolerated association that

has no obvious ill-effect on the health of the host

animals.

Western Province (further in text - WP) lies in

the southwest corner of Papua New Guinea (further

in text - PNG), bordered in the north and east by

Sandaun (West Sepik), Hela, Southern and Gulf

provinces, in the south by Torres Strait and in the

west by the Indonesian province of Papua (Fig.

1). It is by far the largest province in PNG, with

281

€

02-Jun-21 21:50:13

Book4.indd 282

282

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

an area of 99300km2?, and is sparsely populated

- a population of about 200000 with an average

density of 2.1 persons per square km - with poor

infrastructure. Most of the province is low lying with

much swamp-land but rises in the north-northeast

in a series of steep-sided ranges, that include

the Star Mountains, Hindenburg, Victor Emanuel,

Blucher and Muller ranges with an average height

of 3000 meters (Fig. 2). These form the source of

two major WP river systems, the Fly River and its

4 as .

@ Ok Tedi iain, a eaten

. Bi 3 Feramin

e HINDERGURG Cogs i i

_S WG EL

Olsobip. Finimterr AS

3 a

Paap cond \ Bafunmin SARBADIN Tonda Wildlife Management Area

Fae” 23/ Tifalmin OVINCE ,

a af ee “eee -——- comes Provincial Boundaries

=O S\ S~ MIDDLE FLY ey

*2 é : =, * :

: x o > %

iy : 2. : mh

SS i é <8, \

Na: \ ye

pe ly Kavienanga y , <

i | sc i SS

S H a te ¥ XN

y | suki ee Re AS River ~ :

2 Awaba - -— Sa :

: niver in|

: TD

+ TRANS FLY PS 5 & OF

| eVWeam : S I.

Heeb z Rouku a M

TORRES

main tributary, the Strickland. A large open-cast

copper and gold mine is located at Ok Tedi in the

Star Mountains, that has had a profound effect on

the immediate environment of the mine site and,

downstream, on the character of Ok Tedi and Fly

rivers.

WP has a wet climate with tropical rainforest

cover, other than in the southern Trans Fly portion

which has distinct wet and dry seasons with tropical

savannah woodlands dominated by Melaleuca

+ ~+ International Boundaries

® Provincial Headquarters

A District Headquarters

@ Villages

SOUTHERN

HIGHLANDS

PROVINCE

ne rm ee ee, | |

GULF PROVINCE

STRAIT

02-Jun-21 21:50:14

Book4.indd 283

Owen, I. L.: A brief review of animal parasites recorded in Western Province, Papua New Guinea

Linnaeus, 1767 and Acacia Linnaeus, 1773

species. Much of this area can be flooded in the

wet season (approximately December - April) while

experiencing drought conditions in the dry period

(Fig. 3). The province has a rich fauna, due in part,

probably, to the sparseness of the population and

consequently a lack of intensive hunting, although

this has increased in recent years in parts of Trans

Fly. The remote south-west Bula Plain / Bensbach

River area is noted for its variety of permanent and

migratory birds, is designated as a wildlife protected

zone, the Tonda Wildlife Management Area, and

was declared a Ramsar Site in 1993.

This account is a compilation based on

Surveys carried out by the author, mostly in the

Tonda Wildlife Management Area (Fig. 1), and on

parasite collections and data published by other

researchers, notably Drs |. Beveridge, L. Smales

and M. Hastriter, from this area and elsewhere

in the province, including locations straddling

the border of WP and Sandaun Province. The

catalogue, probably, comprises of only a fraction of

the parasite diversity; it is likely that much remains

to be disovered in the rich wildlife of the province.

pp. 281-327

Hosts and their parasites are listed in Table

1 together with host localities and the locations

of parasites in/on their hosts. Parasites of

domestic animals are included but form only a

small part of the account. Parasites that have no

named hosts are included in Table 2. As will be

noted, many parasites discovered remain to be

identified. Host nomenclature is based largely on

the following sources: rodents and marsupials -

Flannery (1995); bats - Bonaccorso (1998); birds

- Beehler & Pratt (2016); varanids - Pianka, King

& King (2004); chelids - Kennett et al. (2014);

amphibians - Menzies (2006); freshwater fishes -

Allen (1991). Parasites are arranged systematically

to ordinal level and the generally accepted mode of

classification is followed.

A local artist, John Siune, from Simbu Province

(PNG), has depicted examples of PNG / WP

parasites in Fig. 4 - using much artistic liberty -

and, with similar liberty, produced illustrations of

host animals (Figs 5-6). Although not scientific, the

illustrations convey the uniqueness of the country

and its culture.

Figure 2. The mountainous north of Western Province: part of the Hindenburg Wall at the border with Sandaun

Province.

Pa.

| foray

283

02-Jun-21 21:50:16

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figure 3. A dust cloud raised by the vehicle draws attention to the bone-dry condition of Bula Plain in the south west

of Western Province during the dry season; the stranded canoe signifies the mode of transport required in the wet

season when the area can be under a meter of water (image courtesy Eric Lindgren, October 1969).

Results

Table 1. Parasites of animals recorded in Western Province, Papua New Guinea.

Western Province host

Mammals Horse (Timor Pony) Bula Plain/Bens- Nematoda (Spirurida Habronema sp. (larvae Associated with

(equids) Equus Caballus (Lin- bach ‘swamp cancer’

naeus, 1758 lesions

Bula Plain/Bens- Insecta (Diptera Chrysomya bezziana Ville- Wounds

bach neuve, 1914 (larvae

)

a | Nematoda (Spirurida) Dirofilaria immitis (Leidy, 1856) | Pulmonary ar-

tery, ventricle

Pe) Bultem village near | Insecta (Phthiraptera) Heterodoxus spiniger (Ender- Body surface

Tabubil lein, 1909

(

)

i )

Mammals Dog (Domestic/Feral) Bula Plain/Bens- Nematoda (Strongylida) |Ancylostoma caninum (Ercol- Small intestine

(canids) Canis familiaris Lin- bach ani, 1859)

naeus, 1758

)

[| [SR eee aac (Pc a rs

bach neuve, 1914 (larvae

bach Bouche, 1835

Mammals Pig (Domestic/Wild) Bula Plain/Bens- Protozoa (Ciliophora) Balantidium coli (Malm- Caecum, colon

(Suids) Sus scrofa (Linnaeus, sten,1857)

1758

bach lidea ?’sparganum system

bach 1975

)

Bula Plain/Bens- Nematoda (Enoplida) Trichinella papuae Pozio et al Adults in s.

bach 1999 int., larvae in

muscle

)

bach caecum

bach Vostokov, 1905

(

)

Bula Plain/Bens- Nematoda (Strongylida) | Oesophagostomum quadrispi- {Large intestine

bach nulatum (Marcone, 1901

)

Bula Plain/Bens- Nematoda (Strongylida) | Stephanurus dentatus Diesing, | Perirenal fat,

bach 1839 thoracic fascia

284

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Owen, I. L.: A brief review of animal parasites recorded in Western Province, Papua New Guinea

pp. 281-327

Table 1 (continuation)

bach

bach sui, 1925

bach Sandosham, 1954 cavit

Coe

bach Molin, 1860

pie ag and

bach 1864

Acanthocephala (Oliga- Small intestine

bach canthorhynchida naceus (Pallas, 1781

Haematopinus suis (Linnaeus, | Body surface

bach 1758

Ctenocephalides f. felis (Bou- | Body surface

bach ché, 1835

Sarcoptes scabiei (Linnaeus, Skin

bach 1758

)

Mammals Cattle (European/Asian) | Oriomo Insecta (Diptera) Chrysomya bezziana Ville- Wounds

(ruminants) Bos Taurus / Bos indi- ie a neuve, 1914 (larvae)

cus Linnaeus, 1758

Haematobia irritans exigua Body surface

Meijere, in Schat, 1903

)

Awaba, Balimo, Acari (Ixodida

Daru, Oriomo, Rum-

sinae, Suki

Goat Samagos village Nematoda (Rhabditida Strongyloides sp. Small intestine

Capra hircus (Linnaeus, | near Kiunga

1758

Haemonchus contortus (Rudol- | Abomasum

near Kiunga phi, 1803

near Kiunga num Curtice, 1890

Nematoda (Strongylida) Small intestine

near Kiunga

Rusa Deer Bula Plain/Bens- Protozoa (Piroplas- Theileria mutans (Theiler, Red blood cells

Cervus timorensis Blain- morida) 1906)

ville, 1822

Fischoederius elongatus (Po- | Rumen

bach irier, 1883

bach 1883

sab

bach neuve, 1914 (larvae

Rhipicephalus (Boophilus) Body surface

australis (Fuller, 1899)

Rhipicephalus (Boophilus) Body surface

bach australis (Fuller, 1899

bach 1914

Mammals Dugong/Sea Cow Daru Is., Gulf of Trematoda (Digenea) Indosolenorchis hirudinaceus Large intestine,

(Sirenia) Dugong dugon (Miller, | Papua in caecum

1776

Daru Is., Gulf of Trematoda (Digenea) Opisthotrema dujonis (Leuck- Eustachian

Papua art, 1874 tube, middle

ear, oesopha-

Papua et Gupta 1971 sages, lungs

ane

Papua 1981 of stomach

Papua Blair, 1981 ducts

Trematoda (Digenea) Foliotrema jecoris Blair, 1981 | Gall bladder,

Papua bile ducts

Daru Is., Gulf of Nematoda (Ascaridida) | Paradujardinia halicoris (Owen, | Cardiac gland

Papua 1833 of stomach

Mammals Greater Bare-backed Bula Plain/Bens- Acari (Mesostigmata) Meristaspis jordani (Radford, Body surface

(bats, Megachi- | Bat bach 1947)

Bula Plain/Bens- Acari (Mesostigmata) Neolaelaps vitzthumi Domrow, | Body surface

bach 1961

Big-eared Flying Fox Bula Plain/Bens- Acari (Mesostigmata) Neolaelaps vitzthumi Domrow, | Body surface

Pteropus macrotis epu- | bach 1961

larius Ramsay, 1878

)

roptera) Dobsonia moluccensis

Quoy et Gaimard, 1830

Spectacled Flying Fox Bula Plain/Bens- Diptera (batflies Cyclopodia (C.) sp. (sykesii Body surface

Pteropus conspicillatus | bach group)

Gould, 1850

Oy 285

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Mammals

(bats, Microchi-

roptera)

Mammals (ro-

dents, Muridae

Hydromyini

286

Common Rousette Bat

Rousettus amplexicau-

datus (Geoffroy, 1810

Common Blossom Bat

Syconycteris australis

Peters, Siphonaptera

Large-eared Sheath-

tailed Bat

Emballonura dianae

fruhstorferi Flannery,

1994

Raffray’s Sheath-tailed

Bat

Emballonura r. raf-

Lesser Sheath-tailed

Bat

Mosia nigrescens (Gray,

1843

Diadem Leaf-nosed Bat

Hipposideros diadema

sriseus (Meyen, 1833

Fawn Leaf-nosed Bat

Hipposideros cervinus

Gould, 1854

Bat

Rhinolophus euryotis

timidus Anderson

Bent-winged Bat

Miniopterus sp.

Northern Hydromyine Bafunmin Insecta (Siphonaptera Acanthopsylla enderleini (Wag- | Body surface

Paraleptomys rufilatus ner, 1933

Osgood, 1945

aie eee Insecta (Siphonaptera Striopsylla vandeuseni Holland, | Body surface

1969

Short-haired Hydro-

myine Paraleptomys

wilhelmina Tate et

Archbold, 19

Hydromyine Paralepto-

mys Sp

Black River, Blucher

Ok Tedi

Diptera (battflies)

Protozoa (Haemospo-

rida

Table 1 (continuation)

Nycteribia p. parilis Walker,

1861

Hepatocystis sp. + a malaria-

like blood parasite

Body surface

Red blood cells

Nematoda (Spirurida

Bula Plain/Bens- | Diptera (batflies)

bach

Bula Plain/Bens- Acari (Mesostigmata) Meristaspis sp. nr. lateralis Body surface

bach Kolenati, 1856

Ok Tedi Diptera (batflies) Cyclopodia s. sycophanta Maa, | Body surface

1971

Cestoda

Acari (Mesostigmata)

Trematoda (Digenea)

Species not identified

(

Meristaspis calcarata (Hirst,

1923)

? Paralecithodendrium sp.

Body surface

Small intestine

A Pteropus

sp. mite - an

accidental

infestation

Small intestine

Black River, Blucher | Nematoda 2 species not identified Stomach/small

Range intestine

Black River, Blucher | Nematoda 2 species not identified Stomach/small

Range intestine

lack River, Blucher } Acari (Mesostigmata) Species not identified

Range

New Guinea Horseshoe | Black River, Blucher | Trematoda (Digenea)

Black River, Blucher

Range

Cestoda

Parabascus sp

Species not identified

Body surface

Stomach/duo-

denum

Range intestine

bach mum (Mehlis, 1831

bach

1969

Black River, Blucher

Range

Bula Plain/Bens-

bach

Bafunmin, Feramin

Nematoda (Ascaridida)

Nematoda (Spirurida)

Insecta (Siphonaptera)

)

Acari (Mesostigmata)

)

Trematoda (Digenea)

Brachylaima sp.

Species not identified Stomach/small

intestine

Small intestine

Heterakis fieldingi Smales, Large intestine

1996

Protospirura kaindiensis

Smales, 2001b

Traubia (Kuruopsylla) kuru Hol-

land, 1969

Body surface

Insecta (Siphonaptera Rectidigitus spooneri Body surface

Rothschild, 1934

' :

Lake Louise

min area

Bafunmin

Insecta (Siphonaptera

“

Orthopsylloides (O.) uncinata

Hastriter, 2014

)

(Telefo- | Insecta (Siphonaptera) | Acanthopsylla eudromiciae Body surface

Holland, 1969

)

Body surface

Striopsylla vandeuseni Holland, | Body surface

1969

02-Jun-21 21:50:19

Owen, I. L.: A brief review of animal parasites recorded in Western Province, Papua New Guinea

pp. 281-327

Table 1 (continuation)

Mammals (ro- | Menzies’ Mouse Abeom- | Telefomin Nematoda (Strongylida) | Montistrongylus karungi Small intestine

dents, Muridae | elomys sevia (Tate et Smales, 2012

Uromyini Archbold, 1935a

| :

Nematoda (Strongylida) | Odilia mackerrasae (Mawson,

1961

)

Nematoda (Ascaridida

Nematoda (Ascaridida Ophidascaris larvae Small intestine

Uneven-toothed Rat An- | Telefomin Insecta (Siphonaptera Rectidigitus szentivanyi Hol- Body surface

isomys imitator Thomas, land, 1969

1904a

Lake Louise (Telefo- | Insecta (Siphonaptera Traubia (T.) durdeni Hastriter, Body surface

min area 2015

(

Insecta (Siphonaptera Rectidigitus traubi Body surface

Holland, 1969

Lake Louise (Telefo- | Insecta (Siphonaptera Traubia (T.) egregia Smit, 1953 | Body surface

min area

N.E. of Telefomin Insecta (Siphonaptera Traubia (T.) traubi Hastriter Body surface

2015

Star Mountains Insecta (Siphonaptera Striopsylla bifurcata Hastriter & | Body surface

Easton, 2013

Star Mountains Insecta (Siphonaptera Tiflovia stellalpestris Traub Body surface

1977

“Giant rat” Star Mountains Insecta (Siphonaptera Acanthopsylla richardsoni Smit, | Body surface

1953

Insecta (Siphonaptera Rectidigitus szentivanyi Hol- Body surface

land, 1969

Eastern White-eared N.E. of Telefomin Insecta (Siphonaptera Traubia (T.) traubi Hastriter, Body surface

Giant Rat 2015

Hyomys goliath (Milne-

Edwards, 1900

Acari (Ixodida) Ixodes goliath Apanaskevich et | Body surface

Lemon, 2018

Long-footed Tree Mouse | Star Mountains Nematoda (Strongylida) | Hasanuddinia sp. Small intestine

Jentink, 1911a

2010.

Smales, 2010

Rothschild’s Woolly Rat | Bafunmin Insecta (Siphonaptera) Rectidigitus spooneri Body surface

Mallomys rothschildi a ae Rothschild, 1934 ca

Thomas, 1898

Large-scaled Mosaic- Tifalmin Nematoda (Strongylida) | Cyclodontosomum purvisi Small intestine

tailed Rat Adams, 1933

Mammelomys lanosus

Thomas, 1922a

Nematoda (Strongylida) | Odilia mackerrasae (Mawson, | Small intestine

1961

he ee puedes Be

Smales, 2017

Nematoda (Ascaridida

Unidentified larvae Small intestine

Insecta (Siphonaptera) | Acanthopsylla enderleini (Wag- | Body surface

ner, 1933

)

) it,

Ee i ) iter,

N.E. & N.W. of Insecta (Siphonaptera) Traubi (Kuruopsylla) kuru Hol- Body surface

Telefomin land, 1969

N.W. of Telefomin Acari (Ixodida Ixodes priscicollaris Schulze, Body surface

1932

) ;

eat

)

Rummler’s Mouse Star Mountains Insecta (Siphonaptera Acanthopsylla eudromiciae Body surface

Coccymys rummleri Holland, 1969

Tate et Archbold, 1941

Bafunmin, Lake Insecta (Siphonaptera Striopsylla vandeuseni Holland, | Body surface

Louise (Telefomin 1969

area

——— a eS Insecta (Siphonaptera) | Orthopsylloides (O.) uncinata

Hastriter, 2014

LN ea ee a ae aa

triter, 2012

Papua Grassland Melo- | Bula Plain/Bens- Cestoda (Cyclophyllidea) | Raillietina celebensis (Janicki, | |ntestine

la mys Melomys lutillus bach + Star Moun- pat ee 1902)

Thomas, 1913 tains (Bafunmin

A cr a na

bach

| le see en Nematoda (Strongylida) | Odilia mackerrasae (Mawson,

bach 1961

Bula Plain/Bens- Nematoda (Strongylida) | Odilia melomyos (Mawson, Small intestine

bach 1961)

28/7

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Table 1

Bula Plain/Bens- Nematoda (Oxyurida) Syphacia darwini Hugot et Intestine

bach Quentin, 1985

Bula Plain/Bens- Nematoda 2 species not identified Intestine

bach

Bula Plain/Bens- Acari (Ixodida) Haemaphysalis humerosa Body surface

bach Warburton et Nuttall, 1909

nymphs

Bula Plain/Bens- Acari (Mesostigmata) Laelaps rothschildi Hirst, 1914 | Body surface

bach

Dokfuma Nematoda (Strongylida) | Hughjonestrongylus mirzai Small intestine

Smales, 2009

Dokfuma Nematoda (Strongylida) | Melomystrongylus sepikensis Small intestine

Smales, 2009

Bula Plain/Bens- Nematoda Species not identified Duodenum

bach

Bula Plain/Bens- Insecta (Phthiraptera) Hoplopleura sp. Body surface

bach

Trans Fly (Eramoe) |} Insecta (Siphonaptera) Orthopsylloides (Mirzapsylla) Body surface

andersoni George et

Beaucournu, 1995

Telefomin Insecta (Siphonaptera) Body surface

Hastriter, 2014

Telefomin Insecta (Siphonaptera) Body surface

land, 1969

Telefomin Insecta (Siphonaptera) Body surface

Holland, 1969

Bula Plain/Bens- Acari (Ixodida) Haemaphysalis humerosa Body surface

bach Warburton et Nuttall, 1909

nymphs

Bula Plain/Bens- Acari (Mesostigmata) Laelaps rothschildi Hirst, 1914 | Body surface

bach

Bula Plain/Bens- Acari (Trombidiformes) Body surface

bach

Bula Plain/Bens- Acari Species not identified Body surface

bach

Lorentz’s Paramelomys_ | Star Mountains Nematoda (Enoplida) Capillaria s. |. Duodenum,

Paramelomys lorentzii caecum

Jentink, 1908

Bafunmin, Tifalmin, | Nematoda (Strongylida) Small intestine

Telefomin Smales, 2017

Bafunmin Nematoda (Strongylida) Small intestine

2017

—

continuation)

Melomys Melomys spp.

Bafunmin Nematoda (Strongylida) | Equilophos similis (Smales, Small intestine

2009

Bafunmin Nematoda (Strongylida) Small intestine

Smales, 2008

| ——— Bafunmin Nematoda (Strongylida) | Odilia mackerrasae (Mawson, | Small intestine

1961

ee ee eee ee oy

Smales, 2009

Pp O—i‘iT ST Barfunmin sd Nematoda (Strongylida) | Heligmonellidspp. SSS Small intestine

a i saad

Adams, 1933

Pp —“‘i‘idS CS Baafurnmin Body cavit

a es ee es canara ce oak a ue

2001a

SSS ES Tar oud

Smales, 2001b

ee a ee sie a ncaa

ner, 1933

7 Bafunmin Insecta (Siphonaptera) Body surface

triter, 2016

Star Mountains Insecta (Siphonaptera) Body surface

Hastriter, 2016

Lowland Paramelomys | Bafunmin Insecta (Siphonaptera) Orthopsylloides (O.) uncinata Body surface

Paramelomys platyops Hastriter, 2014

Thomas, 1906

Telefomin Insecta (Siphonaptera) Orthopsylloides (O.) orthodacty- | Body surface

lus Holland, 1969

)

Telefomin Insecta (Siphonaptera) Traubia (Kuruopsylla) kuru Hol- | Body surface

land, 1969

Acari (Ixodida) Ixodes priscicollaris Schulze, Body surface

1932

Mountain Mosaic-tailed | Mabiomskin (near | Cestoda (Cyclophyllidea) | Railietina celebensis (Janicki, | Small intestine

Rat Ok Tedi Mine) 1902)

Paramelomys rubex

(Thomas, 1922a)

- Ds i ~~

288 Si ol e>

Telefomin

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Owen, I. L.: A brief review of animal parasites recorded in Western Province, Papua New Guinea

pp. 281-327

Table 1 (continuation)

min valle

Ofektaman (Telefo- | Cestoda (Cyclophyllidea) | Rodentolepis fraterna (Stiles, Small intestine

min valle 1906

near Ok Tedi Mine, | Nematoda (Strongylida) | Mawsonema mokwanensis Small intestine

min valle

Ee oe ee

Dokfuma et Heinrich, 2010

Tifalmin, Finimterr, | Nematoda (Strongylida) | Hughjonestrongylus ennisae Small intestine

Telefomin Valle

Finimterr, Tifalmin, | Nematoda (Strongylida) | Parasabanema szalayi Smales | Small intestine

near Feramin, two et Heinrich, 2010

sites in Telefomin

area

Dokfuma, Tifalmin, | Nematoda (Spirurida) Protospirura kaindiensis Stomach

Telefomin valle

Ee EG a eo Lael

mysi Smales et Heinrich, 2010

Ok Tedi Mine identified

2001a

triter, 2014

Rothschild, 1934

min Rothschild, 1934

min land, 1969

5 - -

) : i

)

Bafunmin Insecta (Siphonaptera Rectidigitus traubi Body surface

Holland, 1969

Bafunmin Insecta (Siphonaptera Traubia (T.) durdeni Hastriter, Body surface

2015

N.E. & N.W. of Insecta (Siphonaptera Traubia (Kuruopsylla) kuru Hol- | Body surface

Telefomin land, 1969

Telefomin, Tifalmin | Acari (Ixodida)

Ixodes priscicollaris Schulze, Body surface

1932

Acanthopsylla bisinuata Hol-

land, 1969

Mountain Paramelomys | Bafunmin

or Mountain Mosaic-

tailed Rat

Paramelomys cf. rubex

Insecta (Siphonaptera

Body surface

min ner, 1933

Bafunmin, Lake Insecta (Siphonaptera) | Acanthopsylla eudromiciae Body surface

area

Star Mountains Acanthopsylla richardsoni Smit, | Body surface

1953

Telefomin Orthopsylloides (O.) uncinata Body surface

Hastriter, 2014

min area 1978

min area 1969

)

= Paramelomys sp. Telefomin Insecta (Siphonaptera Wilsonipsylla spinicoxa Has- Body surface

triter, 2012

Shaw Mayer’s Pogonom- | Star Mountains

elomys Pogonomelomys

mayeri (Rothschild et

1932

Champion’s Tree-mouse | Tifalmin Cestoda (Cyclophyllidea) | Bertiella musasabi Yamaguti, Small intestine

Flannery, 1988

1876

min valle Smales, 2015

Telefomin valle Smales, 2014

: ( )

Ofektaman (Telefo- | Nematoda (Strongylida Nippostrongylinae: species not | Small intestine

min valley) identified

Insecta (Siphonaptera Acanthopsylla enderleini (Wag-

ner, 1933)

Body surface

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Table 1 (continuation)

Tifalmin, Ofektaman | Nematoda (Enoplida) Trichuris germani Smales

Telefomin valle 2013

Chestnut Tree-mouse Telefomin Insecta (Siphonaptera) Orthopsylloides (O.) uncinata Body surface

Pogonomys macrourus Hastriter, 2014

Milne-Edwards, 1877

Holland, 1969

Gray-bellied Tree-mouse | Bafunmin Nematoda (Ascaridida) | Heterakis sp. Large intestine

Thomas, 1920

2015

Insecta (Siphonaptera) | Rectidigitus traubi

Holland, 1969

5 :

Black-tailed Giant-rat Tifalmin Nematoda (Strongylida) | Nippostrongylus sembeli Small intestine

Uromys anak Thomas, 5 Hasegawa et Tarore, 1995 ‘aad |

1907

Tifalmin Nematoda (Strongylida) | Odilia carinatae Smales, 2008 | Small intestine

a, [Nippostryngylinae incertae

sedis acc. to Durette-Desset &

Digiani, 2015

Nematoda (Strongylida) | Hughjonestrongylus implexus | Small intestine

Smales, 2008

Nematoda (Strongylida) | Odilia melomyos (Mawson, Small intestine

1961

Tifalmin Nematoda (Strongylida) | Odilia uromyos (Mawson, 1961) | Small intestine

nl [Nippostrongylinae incertae

sedis acc. to Durette-Desset &

Digiani, 2015

Smales, 2007

1876

2001a

Mottled-tailed Giant-rat | Star Mountains Nematoda (Ascaridida) | Subulura andersoni (Cobbold, | Caecum

Uromys caudimaculatus 1876)

Krefft, Siphonaptera

Tifalmin Nematoda (Strongylida) | Odilia carinatae Smales, 2008 | Small intestine

[Nippostryngylinae incertae

sedis acc. to Durette-Desset &

Digiani, 20

Smales, 2008

min ner, 1933

: :

) ini (Wag-

)

Hastriter, 2014

Mammals (ro- | House Mouse Telefomin Insecta (Siphonaptera Acanthopsylla enderleini Body surface

Murini naeus, 1758

Mammals (ro- | Cape York Rat Rattus Bula Plain/Bens- Cestoda (Cyclophyllidea) | Hymenolepis diminuta (Rudol- | Small intestine

dents, Muridae | leucopus (Gray, Siphon- fo | pence phi, 1819)

Rattini aptera

Bula Plain/Bens- Cestoda (Cyclophyllidea) | Raillietina (R.) sp. Small intestine

bach

woo || Bula Plain/Bens- Nematoda (Ascaridida) | Heterakis soumosa (Schneider,

bach 1866

— a a | Bula Plain/Bens- Nematoda (Ascaridida) |Subulura andersoni (Cobbold, | Large intestine

bach 1876

[| Bula Plain/Bens- Nematoda (Enoplida) Trichuris muris (Schrank, 1788)

bach

_— | Bula Plain/Bens- Acanthocephala Species not identified Small intestine

bach

a oa Bula Plain/Bens- Insecta (Siphonaptera) | Xenopsylla vexabilis Jordan, Body surface

bach 1925

a el Bula Plain/Bens- Acari (Mesostigmata) Laelaps sedlaceki Strandt- Body surface

bach mann et Mitchell, 1963

Moss-forest Rat Tifalmin, Dokfuma, | Cestoda (Cyclophyllidea) | Hymenolepis cf. diminuta Small intestine

Rattus niobe (Thomas, | Ofektaman (Rudolphi, 1819)

1906) (Telefomin valley),

Kamptamen (S of

Hindenburg Wall

Beveridge, 2008

hh )

Tifalmin, Ofektaman | Cestoda (Cyclophyllidea) | Dilepididae: species not identi- | Small intestine

(Telefomin valley), fied

290

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pp. 281-327

Table 1 (continuation)

Dokfuma, Tifalmin _]| Nematoda (Enoplida Small intestine

Nematoda (Enoplida Trichuris muris (Schrank, 1788) | Caecum

Ofektaman (Telefo- | Nematoda (Ascaridida) Toxocara mackerrasae (Sprent, | Stomach

min valle 1957

)

Dokfuma, Tifalmin, | Nematoda (Ascaridida Heterakis sirawii Smales, Caecum

Ofektaman 2016b

(Telefomin valley),

Ok Tedi Mine

min valle 2016a

Smales, 2016a

Dokfuma, Tifalmin, | Nematoda (Strongylida) | Nippostrongylinae: species not | Small intestine

Telefomin valle

Dokfuma, Nematoda Juveniles not identified Caecum

Ofektaman

(Telefomin valley), on

Kamptamen (s. of

Hindenburg Wall

[

_ a

Dokfuma, Tifalmin, | Nematoda (Oxyurida) Syphacia (S.) niobe Smales, Caecum

Ofektaman (Tele- 2016a

lf fomin valley), s. of

Hindenburg Wall,

Ok Tedi Mine

Dokfuma, Tifalmin, | Nematoda (Spirurida) Protospirura kaindiensis Stomach

Telefomin valle

Protospirura muricola Gedo- Stomach

elst, 1916

Mabiomskin (near | Acanthocephala Moniliformis moniliformis Small intestine

Ok Tedi Mine) (Bremser, 18414 in Rudolphi,

1819)

Telefomin Hastriter, 2016

min Rothschild, 1934

land, 1969

min Holland, 1969

min area 2015

min area

1932

Moss-forest Rat Telefomin, Bafun- Insecta (Siphonaptera Acanthopsylla bisinuata Hol- Body surface

Thomas, 1906

)

Insecta (Siphonaptera) | Acanthopsylla enderleini (Wag- | Body surface

ner, 1933

)

Lake Louise Insecta (Siphonaptera Acanthopsylla eudromiciae Body surface

(Telefomin area), Holland, 1969

Bafunmin, Star

Mountains

—

Lake Louise Insecta (Siphonaptera) | Acanthopsylla richardsoni Smit, | Body surface

Bafunmin

4 Lake Louise (Tele- Insecta (Siphonaptera) Striopsylla bifurcata Hastriter Body surface

Mountains

mene 64 Lake Louise (Telefo- | Insecta (Siphonaptera) Body surface

min area 1969

Small Spiny Rat Bafunmin Nematoda (Enoplida) Capillaria s. |. Stomach

Rattus steini Rummler,

1935

ls Telefomin, Feramin | Nematoda (Enoplida Small intestine

SSS Telefomin, Bafun- Nematoda (Ascaridida) | Heterakis soumosas (Schnei- | Caecum

min, Feramin der, 1866

(

pO Basfunmin_ sd Nematoda (Ascaridida) | Subulura sp. Caecum

a eee Telefomin, Bafun- Nematoda (Ascaridida) | Toxocara makerrasae (Sprent, | Stomach

min 1957

iS is | Telefomin, Bafun- Nematoda (Strongylida) | Cyclodontostomum purvisi Caecum

min Adams, 1933

“a )

Telefomin, Bafun- Nematoda (Strongylida Nippostrongylinae species 1 & | Small intestine

min, Feramin 2 not identified

291

Book4.indd 291 02-Jun-21 21:50:21

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Table 1 (continuation)

Bafunmin Nematoda (Strongylida Unidentified heligmonellid Small intestine

Telefomin, Bafun- Nematoda (Oxyurida) Syphacia australasiensis Caecum

i Smales, 2004

Telefomin, Bafun- Nematoda (Spirurida) Mastophorus muris (Gmelin, Stomach

min, Feramin 1790

sites sold

Smales, 2001b

| | Bafunmin Acanthocephala (Echino- Small intestine

rhynchida

a ee a ee

ner, 1933

a | lS ce uae | Insecta (Siphonaptera) | Orthopsylloides (O.) uncinata Body surface

min Hastriter, 2014

a eke mim pre ee ee ea

1932

New Guinea Slender Rat | Bafunmin Nematoda (Enoplida) Capillaria s. |. Duodenum

Rattus (Stenomys)

verecundus (Thomas,

1904b

a (a Bafunmin, Mt. Nematoda (Ascaridida) | Heterakis soumosa (Schneider, | Caecum

Fubilan (Ok Tedi 1866

= | Mt. Fubilan (Ok Nematoda (Ascaridida) |Subulura andersoni(Cobbold, | Caecum

i 1876

Mt. Fubilan (Ok Nematoda (Spirurida) Protospirura kaindiensis Stomach

Smales, 2001b

is (

oa Bafunmin Nematoda (Strongylida) Small intestine

Smales, 2018

— Mt. Fubilan (Ok Species not identified Caecum

land, 1969

ner, 1933

Rattus sp. bach tum

ARTEL UMELE

2 2.

Bula Plain/Bens- Acari (Mesostigmata) Laelaps sedlaceki Strandt- Body surface

bach mann et Mitchell, 1963

Bula Plain/Bens- Acari (Mesostigmata) Laelaps nuttalli Hirst, 1915 Body surface

bach

Mammals New Guinea Quoll Bula Plain/Bens- Protozoa (Piroplas- Sarcocystis sp Musculature

(marsupials, Dasyurus albopunctatus | bach morida)

Dasyuridae Schlegel, 1880

Bula Plain/Bens- Species not identified Small intestine

bach

i | Star Mountains Insecta (Siphonaptera) | Acanthopsylla richardsoni Smit, | Body surface

1953

—— a Bula Plain/Bens- Insecta (Phthiraptera) Hoplopleura pacifica Ewing, Body surface

bach 1924

land, 1969

Holland, 1969

Habbema Dasyure Star Mountains Insecta (Siphonaptera) | Acanthopsylla richardsoni Smit, | Body surface

Tate et Archbold, 1941

Short-furred Dasyure Bafunmin Insecta (Siphonaptera) | Acanthopsylla enderleini ( Body surface

Schlegel, 1866

Speckled Dasyure Bafunmin Insecta (Siphonaptera) Rectidigitus spooneri Body surface

Jentink, 1911b

| )

Narrow-striped Dasyure | Bafunmin nsecta (Siphonaptera Acanthopsylla enderleini (Wag- | Body surface

Phascolosorex dorsalis ner, 1933)

(Wag-

Bula Plain/Bens- Nematoda (Enoplida) Trichuris muris (Schrank, 1788) | Caecum

bach

Wag-

brevicaudata (Roths-

child et Dollman, 1932

Red-cheeked Dunnart Bula Plain/Bens- Protozoa (Piroplas- Sarcocystis sp. Musculature

Sminthopsis virginiae bach morida)

Tarragon, 1847

Bula Plain/Bens- Cestoda (Cyclophyllidea) | Hymenolepis sp. nov. Small intestine

bach

Bula Plain/Bens- Nematoda (Strongylida) | Mackerrastrongylus sp. Stomach

bach

Ss Bula Plain/Bens- Nematoda Species not identified Stomach, small

bach intestine

——

\ |

292 bl

Book4.indd 292 02-Jun-21 21:50:22

Owen, I. L.: A brief review of animal parasites recorded in Western Province, Papua New Guinea

pp. 281-327

Table 1 (continuation)

Dunnart Bula Plain/Bens- Cestoda (Cyclophyllidea) | taenid sp. not identified Small intestine

Sminthopsis sp. bach

Bula Plain/Bens- Nematoda (Strongylida Patricialina spp Small intestine

bach

Bula Plain/Bens- Nematoda (Strongylida) |? Mackerrastrongylus sp Small intestine

bach

)

) ;

Mammals Common Spiny Bandi- Bula Plain/Bens- Nematoda (Ascaridida) | Heterakis soumosa Schneider, | Large intestine

(marsupials, coot Echymipera kalubu | bach 1866

Peroryctidae Fischer, 1829

( ida)

)

Smales, 1997

Johnston et Mawson, 1938

Mawson, 1960

Species not identified Small intestine

Bula Plain/Bens- Acanthocephala Species not identified Small intestine

bach

Acari (Ixodida Species not identified Body surface

Acari (Mesostigmata Species not identified Body surface

Bandicoot Insecta (Siphonaptera) | Parastivalius novaeguineae Body surface

Echymipera sp. Rothschild, 1904

)

Striped Bandicoot Telefomin, Bafun- Insecta (Siphonaptera Parastivalius chelaformis Has- | Body surface

Microperoryctes longi- min) triter, 2014

cauda (Peters et Doria,

Rothschild, 1934

land, 1969

——

a a , |

Si | a

min Holland, 1969

et Easton, 2013

Bafunmin, Lake Insecta (Siphonaptera) | Striopsylla vandeuseni Holland, | Body surface

area

PC Star Mountains [Insecta (Siphonaptera) | Traubia (T.) egregia Smit, 1953 [Body surface

Bafunmin, Lake Insecta (Siphonaptera) Traubia (Kuruopsylla) kuru Hol- | Body surface

area

Microperoryctes sp. 1978

Raffray’s Bandicoot Telefomin Insecta (Siphanoptera) Parastivalius papillatus Has- Body surface

Milne-Edwards, 1878

Lake Louise (Telefo- | Insecta (Siphonaptera) Body surface

min area 1969

Bandicoot “Peroryctes” Insecta (Siphonaptera) Body surface

sp. 1969

bach morida

bach Warburton et Nuttall, 1909

Mammals Northern Brown Bandi- | Bula Plain/Bens- Cestoda (Cyclophyllidea) | Paralinstowia semoni (Zs- Small intestine

(marsupials, coot Isoodon macrourus oe eae ae) chokke, 1896)

Peramelidae Gould, 1842a

bach Bullen, 1902

bach tall et Warburton, 1915

bach Warburton & Nuttall, 1909

)

Mammals Gray Dorcopsis Middle Strickland Nematoda (Strongylida) | Labiostrongylus redmondi Stomach

(marsupials, Dorcopsis luctuosa River area Smales, 1982b

Macropodidae D’Albertis, 1874

)

Paralabiostrongylus bicollaris || Stomach

River area Smales, 1982b

Nematoda (Strongylida) | Dorcopsinema dorcopsis (Bay- | Stomach

River area lis, 1940

Middle Strickland Nematoda (Strongylida) | Dorcopsistrongylus labiacarina- | Stomach

River area tus Smales, 1982a

Agile Wallaby Bula Plain/Bens- Trematoda (Digenea Gemellicotyle wallabicola Prud- | Fore stomach

Macropus agilis (Gould, | bach hoe, 1975

1841

“e (

Bula Plain/Bens- Trematoda (Digenea) Macropotrema pertinax Caecum

bach Blair, Beveridge, Speare, 1979

293

Book4.indd 293 02-Jun-21 21:50:22

Book4.indd 294

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

294

“a

“

Red |

ed-legged Pademelon

Thylogale stigmatica

oriomo (Tate et Arch-

[oy

[or

(ce)

(e)

‘ o

Bula Plain/Bens- Cestoda (Cyclophyllidea

bach

Progamotaenia fellicola (Nybe-

lin, 1917

Table 1 (continuation)

Gall bladder,

bile ducts

bach Fuhrmann, 1933

bach 1980

(

Bula Plain/Bens- Nematoda (Strongylida)

bach

Cloacina cornuta (D

Wood, 1938

avey et

bach 2002a

Bula Plain/Bens- Nematoda (Strongylida) | Globocephaloides macropodis

bach Yorke et Maplestone, 1926

Bula Plain/Bens- Nematoda (Strongylida) | Hypodontus macropi Ménnig,

bach 1929

Bula Plain/Bens- Nematoda (Strongylida)

bach

(

Labiostrongylus (L.)

gylus Yorke et Maplestone,

1926

Large intestine,

caecum

labiostron-

bach Baylis, 1927

bach dis Yorke et Maplestone, 1926

bach Beveridge, 1986

bach Yorke et Maplestone, 1926

bach

)

bach et Baccam, 1985

)

Woodwardostrongyl

dorfi Mawson, 1976

Breinlia (B.) boltoni

Varughese, 1975

us oben- Stomach

(Spratt et Body cavities

Bula Plain/Bens- Nematoda (Spirurida Breinlia (B.) robertsi Johnston — | Thoracic cavity

bach et Mawson, 1938

Bula Plain/Bens- Insecta (Phthiraptera

bach

Heterodoxus spiniger (Ender-

lein, 1909

Body surface

Bula Plain/Bens- Insecta (Phthiraptera Body surface

bach

Bula Plain/Bens- Insecta (Phthiraptera Species not identified Body surface

bach

Bula Plain/Bens- Acari (Ixodida) Haemaphysalis bancrofti Nut- | Body surface

bach tall et Warburton, 1915

Bula Plain/Bens- Acari (Mesostigmata) Laelaps sedlaceki Strandmann | Body surface

bach et Mitchell, 1963

Bula Plain/Bens- Acari (Mesostigmata) Alliphis sp. Body surface

bach

)

Bula Plain/Bens- Nematoda (Spirurida) Breinlia (B.) sp.

bach

)

Bula Plain/Bens-

bach

Bula Plain/Bens-

bach

Coleoptera (Scarabaei-

dae)

Bula Plain/Bens-

bach

Coleoptera (Scarabaei-

dae)

Bula Plain/Bens-

bach

Protozoa (Piroplas-

morida)

Blanchard, 1853

Onthophagus symbioticus (Ar-

row, 1920)

Sarcocystis sp.

Acari (Trombidiformes) Eutrombicula sp. (larvae) Body surface

Onthophagus parvus

Temporarily

attached to fur

around cloacal

Temporarily

attached to fur

around cloacal

opening

Musculature

bach Ortlepp, 1922

bach sis Beveridge, 1985

bach Beveridge, 1979

bach 1998

Bula Plain/Bens-

bach

Bula Plain/Bens-

bach

Nematoda (Strongylida

)

iE Cloelia dahli (Linstow, 1898

)

Coronostrongylus jo

Beveridge, 2002b

hnsoni Stomach

02-Jun-21 21:50:23

Owen, I. L.: A brief review of animal parasites recorded in Western Province, Papua New Guinea

pp. 281-327

Table 1 (continuation)

Bula Plain/Bens- Nematoda (Strongylida) | Foliostoma macropodis Bev- Oesophagus

bach eridge et Johnson, 1981

Bula Plain/Bens- Nematoda (Strongylida) | Macropostrongylinea: genus Stomach

bach not identified

Bula Plain/Bens- Nematoda (Strongylida) | Monilonema lacunosum Bev- Stomach

bach eridge et Johnson, 1981

Bula Plain/Bens- Nematoda (Strongylida) | Pharyngostrongylus thylogale Stomach

bach Beveridge, 2016

Bula Plain/Bens- Nematoda (Strongylida) | Pharyngostrongylus setosus Stomach

bach Beveridge, 1982

Bula Plain/Bens- Nematoda (Strongylida) | Popovastrongylus thylogale Stomach

bach Beveridge, 1986

) .

)

bach Beveridge et Andrews, 1993

bach eridge et Durette-Desset, 2004

bach Beveridge, 1983

bach 1981b stomach

bach eridge, 1981b stomach

bach mum Beveridge, 1981a stomach

bach

Bula Plain/Bens- Nematoda (Oxyurida) Small intestine

bach

)

Mammals Southern Common Cus- | Baiyer River Sanctu- | Nematoda (Enoplida Trichuris sp. Caecum

(marsupials, cus Phalanger mimicus |} ary ex Western

Phalangeridae Thomas, 1922b Province

Feramin

Stein’s Cuscus Bafunmin Insecta (Siphonaptera) | Acanthopsylla enderleini (Wag- | Body surface

Phalanger vestitus ner, 1933)

Milne-Edwards, 1877

) (

Mammals Long-tailed Pigmy Pos-

(marsupials, sum Cercartetus cau-

Burramyidae) datus (Milne-Edwards,

Insecta (Siphonaptera

Acanthopsylla enderleini (Wag-

ner, 1933)

Body surface

Mammals Great-tailed Triok Bafunmin

(marsupials, Dactylopsila megalura

Petauridae) Rothschild et Dollman,

1932

Striped Possum Bula Plain/Bens- Nematoda (Spirurida) Breinlia (B.) oweni Spratt, 2011 | Thoracic &

Dactylopsila trivirgata J. | bach . oe ee. eae abdominal

E. Gray, 1858 cavities

Bula Plain/Bens- Nematoda (Spirurida) Stomach

bach ca Bennett, 2003

a a Bula Plain/Bens- Protozoa (Piroplas- Musculature

Dactylopsila sp. bach morida

Sugar Glider Bafunmin Insecta (Siphonaptera) | Acanthopsylla enderleini (Wag- | Body surface

Waterhouse, 1838

triter, 2014

land, 1969

( )

Mammals Painted Ringtail Bafunmin Insecta (Siphonaptera Acanthopsylla enderleini (Wag- | Body surface

(marsupials, Pseudochirulus forbesi ner, 1933)

Pseudocheiri- (Thomas, 1887)

Pygmy Ringtail Telefomin Insecta (Siphonaptera) | Acanthopsylla enderleini (Wag-

Pseudochirulus mayeri ner, 1933)

(Rothschild et Dollman, ~o

1932

Birds (Casuari- | Southern Cassowary Bula Plain/Bens- Nematoda (Spirurida) Habronema casuarii Maple- Stomach wall &

Linnaeus, 1758

bach thighs

Amblyomma papuanum Hirst, | Body surface

Balimo Acari (Sarcoptiformes) Sarcoptes scabiei (Linnaeus, Skin of head &

1758) neck

Insecta (Siphonaptera) | Acanthopsylla enderleini (Wag- | Body surface

ner, 1933)

Body surface

295

Book4.indd 295 02-Jun-21 21:50:23

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Table 1 (continuation)

——k———

a |

ied Village near Kiunga | Nematoda (Spirurida Oxyspirura mansoni (Cobbold, | Under eyelids

1879

| Village near Kiunga | Nematoda (Strongylida) | Trichostongylus tenuis (Mehlis | Small intestine

in Creplin, 1846

— a | Village near Kiunga | Acanthocephala (Gigan- | Mediorhynchus gallinarum Small intestine

torhynchida Bhalarao, 1937

Birds (Gallifor- | Domestic Chicken Village near Kiunga | Trematoda (Digenea) Prosthogonimus ovatus (Rudol- | Oviducts, bursa

mes) Gallus gallus (Linnaeus, phi, 1803)

1758

stow, 1872

aa | ee ered |e ka cae | pee eno

1910

PC SC Village near Kiungar | Cestoda (Cyclophyllidea) |Hymenolepis s.l.,sp.n._____—s| Smalll intestine _|

Megnin, 1881

1788

( ida)

Village near Kiunga | Insecta (Phthiraptera) Menopon gallinae (Linnaeus,

1758

Village near Kiunga | Insecta (Phthiraptera) Cuclotogaster heterographus Head feathers

Nitzsch in Giebel,1866

Acari (Sarcoptiformes) Knemidokoptes mutans (Robin

et Languetin, 1859

Orange-footed Scrub- Bula Plain/Bens- Cestoda (Cyclophyllidea) | Megacirrus leptophallus (Kot- | Small intestine

fowl bach lan, 1923)

Megapodius reinwaradt

reinwardt Dumont,

Insecta (Phthiraptera) Lipeurus latifasciatus Piaget,

bach 1890b

Yellow-legged Brush- Bula Plain/Bens- Acari (Ixodida) Amblyomma papuanum Hirst, | Body surface

turkey bach 1914

Talegalla cf. fuscirostris

Salvadori, 1877

Birds (Anseri- Magpie Goose Bula Plain/Bens- Insecta Phthiraptera) Acidoproctus hilli (Harrison, Underside of

Latham, 1798

Bula Plain/Bens- Insecta (Phthiraptera) Holomenopon goliath Clay, Underside of

bach 1961 wing feathers

Wandering Whistling Bula Plain/Bens- Trematoda (Digenea) Apatemon sp. Small intestine

Duck bach

Dendrocygna arcauata

australis (Reichenbach,

1850

bach merman, 1962

bach Burmeister, 1838

bach

5 —

)

ce)

Bula Plain/Bens- Insecta (Phthiraptera Species not identified

bach

Raja Shelduck Bula Plain/Bens- Trematoda (Digenea) Apatemon (A.) gracilis (Rudol- | Small intestine

Tadorna radjah (Lesson, | bach phi, 1819)

1828

is ey Bula Plain/Bens- Trematoda (Digenea) Small intestine

bach

waa a Bula Plain/Bens- Acanthocephala Polymorphus sp. near marilis Small intestine

bach Echynorhynchida Van Cleave, 1939

cen 0 aia coe

bach

a Bula Plain/Bens- Insecta (Phthiraptera Holomenopon obscurum

bach Piaget, 1880

Birds (Columbi- | Pinon’s Imperial Pigeon | Bula Plain/Bens- Nematoda (Spirurida) Tetrameres sp. Proventriculus

formes) Ducula pinon (Quoy et bach

Gaimard, 1824

Torresian Imperial Pi- Bula Plain/Bens- Trematoda (Digenea) Brachylecithum sp. Small intestine

geon Ducula spilorrhoa | bach

(G. R. Gray, 1858)

296

Book4.indd 296 02-Jun-21 21:50:23

Book4.indd 297

Owen, I. L.: A brief review of animal parasites recorded in Western Province, Papua New Guinea

| Bula Plain/Bens- Trematoda (Digenea)

bach

Black-billed Cuckoo-

Dove

Macropysgia nigrirostris

Salvadori, 1875a

—— Bula Plain/Bens- Acari (Mesostigmata)

bach

Orange-fronted Fruit-

Dove

Ptilinopus aurentiifrons

G. R. Gray, 1858

Pink-spotted Fruit-Dove

Ptilinopus perlatus

zonurus (Salvadori,

Birds (Ciconi-

iformes)

Little Pied Cormorant

Microcarbo

melanoleucos (Vieillot,

1817

Australian Pelican

Temminck, 1824

Australian White Ibis

Threskiornis molucca

Cuvier, 1829

Birds (Charadri-

iformes)

Brown Noddy

Anous Stolidus (Lin-

naeus, 1758

Sharp-tailed Sandpiper

Calidris acuminata

Horsfield, 1821

Curlew Sandpiper

toppidan, 1763

“a

Pelecanus conspicilatus

Bula Plain/Bens- Trematoda (Digenea Bolbophorus confusus (Krause,

bach 1914)

)

Bula Plain/Bens- Nematoda (Ascaridida Contracaecum micropapillatum | Gizzard

bach Stossich, 1890

)

Bula Plain/Bens- Trematoda (Digenea Nephrostomum sp. Small intestine

bach

) i

Bula Plain/Bens- Cestoda (Cyclophyllidea) | Paradilepis sp.

bach

Bula Plain/Bens- Acanthocephala Small intestine

bach

Bula Plain/Bens- Insecta (Phthiraptera) Ardeicola ibis Le Souéf &

bach Bullen, 1902

Bula Plain/Bens- Insecta (Phthiraptera) Colpocephalum aethiopicae

bach Price & Beer, 1965

Bula Plain/Bens- Insecta (Phthiraptera) Ibidoecus dianae Tandan,

bach 1958

)

Bula Plain/Bens- Insecta (Phthiraptera Plegadiphilus threskiornis

bach Bedford, 1939

Bula Plain/Bens- Nematoda (Ascaridida Contracaecum variegatum

bach Piaget, 1880

Bula Plain/Bens- Insecta (Phthiraptera) Actornithophilus umbrinus

oe eee [See

Carduiceps zonarius (Nitzsch,

bach 1866

bach Chapman, 1899

Calidris ferruginea (Pon-

: :

Bula Plain/Bens- Insecta (Phthiraptera) Carduiceps sp.

bach

Bula Plain/Bens- Insecta (Phthiraptera)

bach

Bula Plain/Bens- Insecta (Phthiraptera) Saemondssonia (S.) sp.

bach

Red-necked Stint Bula Plain/Bens- Insecta (Phthiraptera Actornithophilus umbrinus

Calidris ruficollis (Pallas, | bach (Burmeister, 1838)

1776

pp. 281-327

Table 1 (continuation)

Skrjabinus sp. Small intestine

Coloceras spp Feathers

Species not identified

Bula Plain/Bens-

bach

Insecta (Phthiraptera)

Feathers

Bula Plain/Bens- Feathers

bach

Insecta (Phthiraptera)

Bula Plain/Bens-

bach

Nematoda (Spirurida) Ceratospira ?sp. n. Under eyelids

Bula Plain/Bens- Intestine

bach

Nematoda (Ascaridida) | Contracaecum spiculigerum

(Rudolphi, 1809)

? Gizzard

Bula Plain/Bens- Small intestine

Trematoda (Digenea) Mesostephanus sp. prob. pele-

bach cani Cribb, Barker et Beuret,

1995

Bula Plain/Bens-

bach

Kidney

Bula Plain/Bens- Trematoda (Digenea Duodenum

bach

Parastrigea sp

Duodenum

Feathers

Bula Plain/Bens-

bach

Insecta (Diptera) Icostia (Ardmoeca) sp. ‘N’ Maa, | Body surface

1969

Proventriculus

Feathers

Bula Plain/Bens- Feathers

bach (Burmeister, 1838)

Feathers

: Feathers

d Feathers

Feathers

Bula Plain/Bens- Feathers

bach

Bula Plain/Bens-

bach

Insecta (Phthiraptera) Carduiceps zonarius (Nitzsch,

1866

Insecta (Phthiraptera) Saemondssonia (S.) sp. Feathers

297

02-Jun-21 21:50:24

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Table 1 (continuation)

Oriental Plover Bula Plain/Bens- Insecta (Phthiraptera) Quadriceps assimilis (Piaget, Feathers

Charadrius veredus bach 1890a)

Gould, 1848

Whiskered Tern Bula Plain/Bens- Insecta (Phthiraptera) Quadraceps anagrapsus Feathers

Chlidonias hybrida bach (Nitzsch in Giebel, 1866)

javanicus (Horsfield,

bach Giebel, 1874

Beach Stone-curlew Bula Plain/Bens- Insecta (Phthiraptera) Quadriceps sp. near annulatus | Feathers

Esacus magnorostris oes (Denny, 1842)

Vieillot, 1818b

Black-winged Stilt Bula Plain/Bens- Cestoda (Cyclophyllidea) | Gyrocoelia crassa (Fuhrmann, | Duodenum

Himantopus himanto- bach a ead 1900)

ous (Linnaeus, 1758

Wardium tsengi (Joyeux et Small intestine

bach Baer, 1940

Little Curlew Bula Plain/Bens- Insecta (Phthiraptera) Austromenopon sp. (nymphs) Feathers

Numenius minutus bach igi nap

Gould, 1840

bach

Bula Plain/Bens- Insecta (Phthiraptera) Saemondssonia (S.) scolopaci- | Feathers

bach lial al sphaeopodis (Schrank, 1803)

SrOUp

Birds (Accipitri- | Whistling Kite Bula Plain/Bens- Trematoda (Digenea) Neodiplostomum spathula Small intestine

Vieillot, 1818a

Bula Plain/Bens- Cestoda (Cyclophyllidea) | Paradilepis urceus (Wed, Small intestine

bach I USSENS)

Bula Plain/Bens- Cestoda (Cyclophyllidea) | Parvitaenia milvi (Singh, 1952) | Small intestine

bach

axe

(

Bula Plain/Bens- Nematoda (Ascaridida) Small intestine

bach

Bula Plain/Bens- Insecta (Phthiraptera) Colpocephalum turbinatum Body surface &

bach Denny, 1842 feathers

Bula Plain/Bens- Insecta (Phthiraptera) Laemobothrion (L.) maximum Feathers

bach Scopoli, 1763

)

Birds (Coraci- Rufous-bellied Bula Plain/Bens- Insecta (Phthiraptera Emersoniella sp. Feathers

iformes) Kookaburra bach

Dacela gaudichaud

Bula Plain/Bens- Insecta (Siphonaptera) | Xenopsylla vexabilis Jordan, Body surface

bach 1925

Spangled Kookaburra Bula Plain/Bens- Insecta (Phthiraptera Alcedoeécus sp. Feathers

Dacelo tyro G. R. Gray, bach

1858

)

Bula Plain/Bens- Insecta (Phthiraptera) Species not identified Feathers

bach

)

ie)

tna

G@)

ret)

fos

Birds (Falconi- | Brown Falcon Bula Plain/Bens- Insecta (Phthiraptera Craspedorhynchus sp. Feathers

formes) Falco berigora novae- bach

Suineae (Meyer, 1894

EE, Bula Plain/Bens- Insecta (Phthiraptera) Degeeriella rufa (Burmeister, Head feathers

bach 1838

Bula Plain/Bens- Insecta (Phthiraptera Species not identified Feathers

bach

Australian Hobby Bula Plain/Bens- Nematoda (Spirurida) Serratospiculum tendo Air sacs

Falco longipennis bach (Nitzsch, in Giebel, 1857)

Swainson, 1838

Birds (Passeri- | Little Strikethrush Bula Plain/Bens- Acari (Sarcoptiformes) Trouessartia sp. Feathers

formes) Colluricincla megarhyn- | bach

cha (Quoy et Gaimard,

Black-faced Cuckoo- Bula Plain/Bens- Nematoda (Spirurida) Diplotriaena tridens (Molin, Body cavity

shrike bach 1858)

Coracina novaehollandi-

Bula Plain/Bens- Species not identified ?

bach

Bula Plain/Bens- Insecta (Phthiraptera) Feathers

bach

Bula Plain/Bens- Insecta (Phthiraptera) Philopterus sp. Body surface,

bach feathers

Bula Plain/Bens- Insecta (Phthiraptera) Species not identified Feathers

bach

Bula Plain/Bens- Acari (Sarcoptiformes) Gabucinia sp. Feathers

bach

je)

ie)

298

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Owen, I. L.: A brief review of animal parasites recorded in Western Province, Papua New Guinea

pp. 281-327

Table 1 (continuation)

Torresian Crow Bula Plain/Bens- Insecta (Phthiraptera) Brueelia sp. Feathers

Corvus orru Bonaparte,

1850

ac a arc al) rend Eo

bach 1842

SS lial a

bach

ae eee | ee |e ge fl

bach

bach tall et Warburton, 1915

(NL OS aed anind cue |e nce OE) chee Be)

bach

Hooded Butcherbird Bula Plain/Bens- Cestoda (Cyclophyllidea) | dilepid species not identified Small intestine

sicus Boddaert, 1783

a rae | aaa Se geiaycne sic oan | ee cea

bach et Mawson, 1940

Spangled Drongo Bula Plain/Bens- Insecta (Phthiraptera) Philopterus sp. Body surface

Gould, 1842b

) :

bach

Common Cicadabird Bula Plain/Bens- Insecta (Phthiraptera) Philopterus sp. Body surface,

Jardine, 1831

Large-billed Gerygone Ok Tedi, Kiunga, Aves (Cuculiformes) Chalcites minutillus (Gould, Eggs in nest

Gould, 1842b

Brown Oriole Bula Plain/Bens- Cestoda Species not identified ?

Madarasz, 1900

Tawny-breasted Hon- Bula Plain/Bens- Nematoda (Spirurida) Diplotriaena sp.

eyeater bach

Xanthotis flaviventer

saturatior (Rothschild &

Hartert, 1903

Reptiles (vara- | Gould’s Monitor/Go- Bula Plain/Bens- Cestoda (Proteocephali- | Kapsulotaenia sp. Intestine

nids) anna Varanus gouldii bach dea)

Gray, 1838

bach lidea

bach

| | Bula Plain/Bens- Acari (Ixodida) Amblyomma fimbriatum Koch, | Body surface

bach 1844

|| cs Bula Plain/Bens- Acari (Ixodida) Amblyomma sp. Body surface

bach

( )

)

Mangrove Monitor/Go-_ | Bula Plain/Bens- Acari (Ixodida Amblyomma trimaculatum Body surface

Daudin, 1802

Emerald Tree Monitor/ | Daru Is. Acari (Ixodida Amblyomma fimbriatum Koch, | Body surface

Goanna 1844

Varanus prasinus Schle-

gel, 1839

Papua Monitor/Goanna | Bula Plain/Bens- Cestoda (Proteocephali- | Proteocaphalus sp. prob. sac- Intestine

ters et Doria, 1878

bach lidea ter

1844

tomida

Varanus Sp. bach

Reptiles (croco- | New Guinea Crocodile Daru Is., Lake Trematoda (Digenea) Cyathocotyle (Suchocyathocoty- | Intestine,

diles) Crocodylus novaeguin- } Murray eae le) crocodili Yamaguti, 1954 rectum

eae Schmidt, 1928

Lake Murra Trematoda (Digenea Exotidendrum sp. Large intestine

Lake Murray Nematoda (Spirurida) Micropleura sp. Body cavity

299

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Book4.indd 300

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Crocodile

Crocodylus sp.

Northern Snake-necked

Turtle

Chelodina rugosa

Reptiles (che-

lonids)

Ogilby, 1890

Arafura File Snake Bensbach River

Acrochordus arafurae

McDowell, 1979

Angiana Tree Frog

Litoria angiana (Bou-

lenger, 1915

Beck’s tree frog

a Litoria becki Loveridge,

1945

Sandy Big-eyed Tree

[ Frog Nyctimystes kubori

Zweifel, 1958

Common Big-eyed Tree

= Frog Nyctimystes nari-

nosa Zweifel, 1958

Mountain Swamp Frog

Papurana grisea (van

Kampen, 1923

Catfish

Arius sp.

Reptiles

(snakes)

Amphibia Star Mountains

(frogs)

Star Mountains

Fishes (fresh- Bensbach River

water, migra-

tor

Barramundi Bensbach River

Lates calcarifer (Bloch,

1790

Bensbach River

——— ||

Pp rr—“—*éi‘idSCSC“‘(NC CC Bensshach River

Echynorhynchida

Po Bernsbacch River Crustacea (Copepoda

Trematoda (Digenea)

ioe

Monogenea

Papuan Blackbass

a Lutjanus goldiei (Ma-

cleay, 1882

Sawfish

——— Pristis microdon

Latham, 1794

————EEE=|

(SS

Saratoga

Scleropages jardinii

Saville-Kent, 1892

Lake Murray

Lake Murra

Fly River

Bensbach River

Torres Strait near

Kaviopo village

Torres Strait near

marine fish’

Kaviopo village

Fishes (marine,

migrator

‘tuna’ Torres Strait near

| Kaviopo village

Eleutheronema sp. Kaviopo village

Kaviopo village

a: King Threadfin Gulf of Papua

Polydactylus macrochir

Gunther, Siphonaptera

Insects (Hyme-

noptera)

Asian Honeybee

Apis cerana Fabricius,

1793

Kiunga

300

Nematoda (Ascaridida

Annelida (Hirudinida)

Protozoa (Piroplas-

morida

Aramia River

Cestoda (Trypanorhyn-

cha)

Crustacea (Copepoda)

)

Nematoda (Ascaridida

Nematoda (Spirurida)

Nematoda (Ascaridida

Nematoda (Ascaridida)

)

Crustacea (Copepoda

Crustacea (Copepoda)

Cestoda (Trypanorhyn-

cha

Cestoda (Trypanorhyn-

cha

Acanthocephala

Cestoda (Trypanorhyn-

cha)

Cestoda (Trypanorhyn-

cha

Nematoda (Ascaridida)

Acari (Mesostigmata)

Table 1 (continuation)

Stomach

Body surface

Red blood cells

Dujardinascaris sp.

Placobdella sp.

‘haemogregarines’

Polystomoides sp. Bladder

Tanqua ophidis Johnston et Stomach

Mawson, 1948

Leiobdella singularis (Richard-

son, 1975)

Beneath skin,

body cavity

Beneath skin,

body cavity

Leiobdella singularis (Richard-

son, 1975)

Leiobdella singularis (Richard-

son, 1975)

Beneath skin,

body cavity

Leiobdella singularis (Richard-

son, 1975)

Beneath skin,

body cavity

Leiobdella singularis (Richard-

son, 1975)

Beneath skin,

body cavity

Poecilacanthum oweni Palm, Musculature

1995 (plerocerci)

Ergasilus sp.1 Gills

Cucullanus hians (Dujardin, Intestine

1845)

Mesenteries

Ovaries

Dujardinascaris sp. (larvae

Philometra ?lateolabracis

Yamaguti, 1935

Species not identified

Longicollum edmonds Golvan,

1969

Ergasilus sp.2

‘didymozoid’, possibly Metane-

matobothrium sp.

Mesenter

Intestine

Gills

Coiled in dorsal

muscle

Nonacotyle pristis Ogawa, 1991 } ? Gills

Pristonchocotyle papuensis ? Gills

Ogawa, 1991

Hysterothylacium sp.

Species not identified

Brevimulticaecum scleropagi

Khalil, 1984

ntestine

Stomach

Lernaea papuensis Boxshall, Gill arches

‘unidentified tangled mass of

larvae’

Ergasilus sp. 2

Body cavity

mesenteries

Gills

Connective

tissue

Body cavity,

muscle

Intestine

‘plerocerci’

Species not identified

Poicilancixtrum sp. ?caryophyl- | Musculature

lum (Diesing, 1850) (plerocerci)

‘plerocerci’ Gill canals

Goezia sp. (larvae) Within trypano-

rhynch plero-

cerci cysts

Body surface,

worker & drone

broods

Varroa jacobsoni Oudemans,

1904

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Owen, I. L.: A brief review of animal parasites recorded in Western Province, Papua New Guinea

Review of parasites of animals in Western

Province, Papua New Guinea

Parasites of non-native domestic animals

Domestic animals, other than pigs, dogs and,

in some localities, chickens have been present in

the country only for the last hundred years or so.

Ruminants, horses and cats

At present, cattle are few in numbers in WP.

Most, it seems, are no longer under control and

have become feral in localities such as Suki and

Balimo. Goats have diminished in number also,

and there are no sheep currently recorded in the

province. A few horses (Timor ponies), that are

mostly feral, occur in the Wando area of Bula Plain/

Bensbach. Cats are found in most communities

other than in some remote villages. They are few

in numbers, some having become feral but usually

remaining close to human habitation. Their effect

as hunters on local fauna is possibly significant

(Flannery & Seri 1990).

The only endoparasites recorded from

the domestic ruminants are nematodes from

goats; Haemonchus contortus (Rudolphi, 1803),

Oesophagostomum columbianum Curtice, 1890,

Strongyloides sp. and Trichstrongylus sp. There

are no records of horses being examined for

endoparasites but the presence of ‘swamp cancer’,

a fungal disease, in an animal at Wando indicates

that the nematode Habronema sp. is present as

its larvae are associated with the fungal infection.

Likewise, there are no records of parasites of cats

in WP, although elsewhere in the country they have

some of the same parasites as dogs.

Three ectoparasites are known to occur on

some of these imported domestic animals. The tick,

Rhipicephalus (Boophilus) australis (Fuller, 1899)

that infests cattle, was confused until recently

with R. (B.) microplus (Canestrini, 1888) (Estrada-

Pena et al. 2012). The dipteran flies, Haematobia

irritans exigua (Meijere in Schat, 1903) occurs on

cattle, and Chrysomya bezziana Villeneuve, 1914

on cattle and horses (as well as other domestic

and wild animals). The latter insect, commonly

known as the Old World screw-worm fly, unlike the

other two ectoparasites, is long-established in the

country. It found a ready source of suitable hosts

with the arrival of new types of imported livestock.

The wounds its larvae (maggots) produce are

debilitating and can be fatal.

pp. 281-327

Parasites of long-established domestic animals

Dogs

Dogs have been in PNG, and probably WP,

for at least a couple of thousand years; no one

knows for sure when they were brought into the

country. They form an essential part of most village

families and are especially important to hunters

and, increasingly, for security. They are often poorly

fed, sometimes having to fend for themselves and

consequently can be in poor condition, particularly

when present in excessive numbers as, for example,

a village in Middle Fly region was recorded at one

time as having 300 dogs, 23 of them with one

owner. It means that many are likely to have more

problems with parasites than they otherwise might

have. A number of feral dogs inhabit Bula Plain/

Bensbach area that predate on young fawns in

particular (Hitchcock 2004), and wild dogs, or

what are known as New Guinea singing dogs, are

recorded by Flannery & Seri (1990) in the subalpine

grassland of Dokfuma in the Star Mountains.

Few dogs in WP have been examined for

parasites but they are known to have a hookworm,

Ancylostoma sp., and the heartworm, Dirofilaria

immitis (Leidy, 1856) is common. Heartworm (Fig.

7), when present in large numbers in the heart

or pulmonary artery, can block blood flow to the

lungs that may lead to the collapse of the dog, and

sometimes death. Ectoparasites found on dogs

include the chewing louse, Heterodoxus spiniger

(Enderlein, 1909), the flea, Ctenocephalides f.

felis (Bouché, 1835), and the screw-worm fly

Chrysomya bezziana. The absence of the dog tick,

Rhipicephalus sanguineus (Latreille, 1806) and

mange producing mites, is likely to reflect the low

number of hosts examined, as they are recorded

almost universally in the country.

Pigs

Domestic pigs are an integral part of most

village families in PNG, other than where wild pigs

are plentiful, as in much of the southern part of

WP where, if kept, they are caught as wild piglets

and usually penned. Pigs are “cultural superfood”

for the people of Bula Plain/Bensbach according

to Hitchcock (2004). Wild and village pigs often

cross-breed and are considered to be of the same

lineage (Groves 1981; Ayalaw et a/. 2011). The time

of arrival of pigs into the country is uncertain but

generally believed to be 3000 to 4000 years ago

(Hide 2003). As the parasites of domestic and wild

301

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TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

pigs are similar, and as many more wild pigs have

been examined than domestic ones, their parasites

are considered when discussing those of wild pigs.

Chickens

Some coastal areas in the country are known

to have had chickens for a few centuries, but they

did not become widespread until the 20" century.

Today they are found probably in most WP villages.

They harbour both ecto- and endo-parasites.

Although no protozoan parasites have been

recorded from WP chickens, the various coccidial

species (Eimeria) are ubiquitous in PNG and

almost certainly present in WP. The tapeworms,

Hymenolepis exigua (Yoshida, 1910), Raillietina

echinobothrida (Megnin, 1881) and Amoebotaenia

cuneata (Linstow, 1872) are common in the

province, the latter species, possibly, being the most

harmful. A small flatworm, Prosthogonimus ovatus

(Rudolphi, 1803), that occurs in oviducts, can

affect egg production. Nematodes found include

Capillaria sp., the eyeworm Oxyspirura mansoni

(Cobbold, 1879), Trichostrongylus tenuis (Mehlis in

Creplin, 1846) and Heterakis gallinarum (Schrank,

1788). The only acanthocephalan recorded is

Mediorhynchus gallinarum (Bhalarao, 1937).

A few ectoparasites have been recorded

from WP chickens, including the chewing lice

Cuclotogaster heterographus (Nitzsch in Giebel,

1866) and Menopon gallinae (Linnaeus, 1758).

Heavy infestations are seldom seen as village

chickens are not kept in conditions of close

contact. The only mite recorded is the scaly leg

mite, Knemidocoptes mutans (Robin et Lanquetin,

1859).

Parasites of wildlife

Investigations were conducted into the

parasites of WP wildlife over several decades,

primarily in the Bula Plain / Bensbach River section

of the Tonda Wildlife Management Area. This south

western locality has distinct wet and dry seasons,

with much of the land under water for part of the

year, alternating often with drought conditions (Fig.

S)e

Deer

Rusa deer (Cervus timorensis Blainville, 1822)

occur in southern WP and were first observed in

the Bula Plain / Bensbach area during late 1930s,

having spread across the border from adjacent West

302

Papua. They have thrived to become a major food

source for local people and have spread eastwards,

crossing the Fly River. A Survey conducted on the

grasslands of Bula Plain / Bensbach area in 1987

recorded an estimate of 23700 animals (Fraser-

Stewart 1988). Numbers have diminished since

due to increased hunting and poaching; severe

drought conditions, as occurred in 1997/8, can

lead to a high mortality (Hitchcock 2004). The

presence of the deer has had a “profound impact

on the local environment” (Hitchcock 2004).

The deer (65 were autopsied over 32 years)

had no nematode parasites, which is unusual for

herding ruminants. They were parasitized by two

species of digenean flatworms, Homalogaster

paloniae Poirier, 1883, and _ Fischoederius

elongatus (Poirier, 1883). Large numbers of the

latter species were frequently found in the rumen

with no apparent harmful effect on the host (Fig.

8). Homalogaster, on the other hand, was always

few in numbers, and was not seen following the

severe drought of 1997/8 which may have had a

deleterious effect on its molluscan intermediate

host. The apparently benign protozoan Theileria

mutans (Theiler, 1906) parasitized the red blood

cells of most animals.

The cattle tick, Rhipicephalus (Boophilus)

australis (Fuller, 1899), was acommon ectoparasite

(78% parasitism), but animals carried more larvae

and nymphs than adults. Rusa deer is a supple

animal with a bristly coat, whose hair is readily

erected - factors that may help remove adult

ticks during self-grooming. Deer sometimes cover

themselves with mud that is sloughed off when dry,

an action which may also help remove large adult

ticks. Infestation tended to be heaviest in the dry

season, particularly for males and non-pregnant

females (Owen 1977). The ability of the deer to

carry the cattle tick presents a problem for cattle

farmers. Another tick, Amblyomma papuanum

Hirst, 1914, was recorded on one occasion. Three

cases of screw-worm fly lesions were seen.

Wild (and village) pigs

About 80 pigs (mostly wild) have been examined

for parasites, almostall from the southern part of the

province. The few ectoparasites recorded comprise

of the louse Haematopinus suis (Linnaeus, 1758),

the flea Ctenocephalides felis (Bouché, 1835) and

the mite Sarcoptes scabiei (Linnaeus, 1758).

Common nematodes found in both wild and

village pigs included the lungworm, Metastrongylus

pudendotectus (Vostokov, 1905), the gut worms

eee Bar

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Owen, I. L.: A brief review of animal parasites recorded in Western Province, Papua New Guinea

Ascaris suum (Goeze, 1782), Gnathostoma

doloresi (Tubangui, 1925), Oesophogostomum

quadrispinulatum Marcone, 1901, Physocephalus

sexalatus (Molin, 1860) and the long, threadlike

Setaria thomasi (Sandosham, 1954) found in the

body cavity. A nematode of the tongue, Capillaria

papuensis Copland, 1975, so far, reported only

from PNG, was present in 29% of wild pigs on Bula

Plain.

A zoonotic nematode, Trichinella papuae Pozio,

Owen, La Rosa, Sacchi, Rossi et Corona, 1999, was

first detected in village and wild pigs in 1988 and

later, with serological testing, humans were found

to be infected (Owen et al. 2001; 2005). It is an

unusual worm in that both adults and larvae infect

the same host, with the minute adults living in the

intestine and larvae in striated muscle fibres (Fig.

9). It is unusual, also, in being able to parasitize

both warm and cold-blooded (crocodile) hosts.

About 11% of wild pigs on Bula Plain were infected

and 10% of villagers in Morehead District tested

positive. Domestic pigs become infected by being

fed contaminated wild pig meat, while eating

uncooked or undercooked infected pig meat leads

to human infection. Another zoonotic parasite

found in pigs was the protozoan Balantidium coli

(Malmsten, 1857).

The large acanthocephalan or thorny-headed

worm, Macracanthorhynchus hirudinaceus (Pallas,

1781) is a common intestinal parasite of wild pigs

(Fig. 10). It may be a problem for the host if present

in appreciable numbers as it can damage and

penetrate the gut wall.

Dugongs

Parasites of the dugong, Dugong dugon

(Muller, 1776), which lives in the shallow coastal

waters of WP, include at least six species of

flatworms, occupying locations ranging from the

middle ear and Eustachian tubes, to the lungs,

stomach, intestine, liver, and pancreas, but with

only one nematode species reported (Blair 1981).

Bats

Bats have been examined for parasites from

three localities, the Bula Plain / Bensbach area,

Ok Tedi, and Black River in the Blucher Range.

Those from the latter two localities were museum

specimens preserved and stored in 70% alcohol.

None of the fruit bats, flying foxes and blossom

bats (Megachiroptera) (34 individuals examined)

contained endoparasites, but Peters (1957)

pp. 281-327

recorded Hepatocystis sp., a malaria-like parasite,

and microfilariae in the blood of WP flying foxes.

Amongst the insectivorous bats (Microchiroptera)

(24 individuals examined), most had at least one

type of endoparasite. This points to the role insects

may play in the lifecycles of bat endoparasites that

require intermediate hosts. Ectoparasites, on the

other hand, were found on both insectivores and

frugivores, but less often on the former than the

latter group. Mites were present on both groups but

dipteran batflies occurred only on Megachiroptera.

None was heavily parasitized with either of the

parasite types. On one occasion an ectoparasitic

mite, Meristaspis calcarata (Hirst, 1923) was found

attached to the wing rib of a microchiropteran,

Emballonura r. raffrayana (Dobson, 1879) - an

accidental infestation probably, as it is a parasite of

Pteropus sp., a megachiropteran. Many of the bat

parasites have yet to be identified.

Rodents

Rodents have been examined from the extreme

south and the extreme north of the province.

Included in this account of the northern group are

animals from sites in the Star Mountains region,

such as Tifalmin, Bafunmin, Telefomin and Feramin,

that lie adjacent to, but on the Sandaun side of,

the border with WP. This group contained a greater

range of host species than did the southern group

from Bula Plain/Bensbach. The northern rodents

also harboured a larger range of parasites; Rattus

niobe (Thomas, 1906) and Paramelomys lorentzii

(Jentink, 1908) had 14 and 11 species (discounting

larvae and juveniles) of endoparasites respectively,

dominated largely by strongylid nematodes,

while the highest numbers in the southern hosts

were seven endoparasites in Melomys_ lutillus

(Thomas, 1913) and six in Rattus leucopus (Gray,

Siphonaptera). Five parasite species were present

in both northern and southern hosts, namely, the

cestodes Hymenolepis diminuta (Rudolphi, 1819)

and Raillietina celebensis (Janicki, 1902), and the

nematodes Odilia mackerrasae (Mawson, 1961),

O. melomyos (Mawson, 1961) and Trichuris muris

(Schrank, 1788).

Ectoparasites of rodents have been little

studied. So far, it seems, fleas are the only ones

reported from the northern based rodents other

than for two species of tick. Twenty-three species

of fleas were recorded by Hastriter & Easton

(2013) and Hastriter (2014; 2015; 2016) on a

range of murid hosts. Acanthopsylla enderleini

(Wagner, 1933) was the most common, occurring

303

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

on 10 host species, followed by Orthopsylloides

uncinata Hastriter, 2014 on eight species. The ticks

comprise of Ixodes priscicollaris Schulze, 1932,

found on three species of rodents and |. goliath

Apanaskevich et Lemon, 2018 on one. Only two

flea species are recorded from the southern region;

Xenopsylla_ vexabilis Jordan, 1925 on Rattus

leucopus from the Bula Plain/Bensbach area and

Orthopsylloides andersoni George et Beaucournu,

1995 on Melomys sp. from Trans Fly area of the

province. The lice Hoplopleura pacifica Ewing, 1924

and Hoplopleura sp. were found on Rattus sp. and

Melomys sp. respectively, and nymphal stages

of the tick Haemphysalis humerosa Warburton

et Nuttall, 1909 were collected from Melomys

spp. Mites (Laelaps spp.) occurred on species of

Melomys and Rattus, with L. rothschildi Hirst, 1914

a common parasite of Melomys lutillus (Thomas,

1913).

Marsupials

Parasites have been recorded from about ten

species of marsupials. One of the iconic animals

of the Bula Plain savannah is the agile wallaby,

Macropus agilis (Gould, 1841). As with deer and

wild pigs, and for the same reasons, there has been

a marked decline in numbers during recent times,

despite the area being part of the Tonda Wildlife

Management Area.

A large range of parasites is recorded from the

wallaby (24 were examined), including two species

of flatworms, two tapeworm species, 16 nematode

species (of which 13 were strongylids), three

species of lice, one tick species and four species

of mites. The most common endoparasites were

the nematodes Labiostrongylus labiostrongylus

Yorke et Maplestone, 1926 and Cloacina cornuta

(Davey et Wood, 1938) found in 11 and 10

individuals respectively. The most frequently seen

ectoparasite was the tick Haemaphysalis bancrofti

Nuttall et Warburton, 1915 recovered from six

animals. In addition, two species of dung beetles,

Onthophagus parvus Blanchard, 1853 and O.

symbioticus (Arrow, 1920), have an association

with the wallaby but were encountered only on

animals seen at night. The nocturnal beetles are

firmly attached around the host cloacal opening

with the aim of acquiring dung pellets as these

are passed out. The two species have slightly

different adaptations of the claws for grasping the

host’s fur (Matthews 1972). The beetles should be

viewed probably as pseudoparasites or, possibly, as

practicing commensalism as depicted by Combes

304

(2005). The presence of these beetles can explain

finding the mite Aliphis sp. on a wallaby as the mite

is often found on scarabid dung beetles (Krantz &

Walter 2009).

A much less frequently seen marsupial was

the pademelon or red-legged wallaby, Thylogale

stigmatica oriomo (Tate et Archbold, 1935b) (3

were examined). It is shy and more solitary than

the agile wallaby, preferring a mixed woodland/

savannah than an open savannah environment

and is restricted to the southern Trans Fly area

of New Guinea. Its range of nematode species

(21) is even greater than that of the agile wallaby.

One, apparently healthy, animal contained 32270

nematode worms, comprising of 17 species in the

stomach alone.

Agile wallabies and pademelons occur in the

same locality but they harbour different parasites.

Worms of the same genus may be found in both,

such as, the nematode genera Cloacina von

Linstow, 1898, Pharyngostrongylus Yorke et

Maplestone, 1926, Popovastrongylus Mawson,

1977 and the tapeworm Progamotaenia Nybelin,

1917, but no species of these genera was found

common to both hosts. This may reflect, in part, on

a difference in diet; the agile wallaby is primarily a

grazer, while the pademelon is more of a browser,

reported to feed mainly on fruit and understorey

plants (Flannery 1995).

Another wallaby, Dorcopsis /uctuosa (D’Albertis,

1874) (wrongly identified in the past as D. veterum

(Lesson, 1827)), from the Middle Strickland

area, had species of nematodes that are largely

confined to this host, such as, Dorcopsistrongylus

labiocarinatus Smales, 1982a, Labiostrongylus

redmondi Smales, 1982b, Paralabiostrongylus

bicollaris Smales, 1982b and Dorcopsinema

dorcopsis (Baylis, 1940).

The smaller marsupial species from the

southern part of the province had relatively few

parasites. Of the 23 dunnarts (Smynthopsis spp.)

examined, only six harboured endoparasites

comprising of possibly four nematode species, two

species of tapeworms and a protozoan (Sarcocystis

Sp.); no ectoparasites were seen. The striped

possum (Dactylopsila trivirgata J. E. Gray, 1858)

and Dactylopsila sp. had two nematode species

and a species of Sarcocystis. Only a muscle

biopsy, that revealed a Sarcocystis sp. infection,

was possible with the carnivorous quoll (Dasyurus

albopunctatus Schlegel, 1880). Two bandicoot

species were examined from the Bula Plain/

Bensbach area; Echymipera kalubu (Fischer, 1829)

had four nematode species, an acanthocephalan,

02-Jun-21 21:50:25

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Owen, I. L.: A brief review of animal parasites recorded in Western Province, Papua New Guinea

a tick and a mite, while Isoodon macrourus (Gould,

1842a) had a tapeworm, a louse and two species

of ticks.

Small marsupials from elsewhere in the

province have not been studied for endoparasites,

but 14 species of fleas have been recorded by

Hastriter & Easton (2013) and Hastriter (2014;

2015; 2016) from about 14 host species at

locations near the northern border; these were

examined for ectoparaites only. As with rodents,

Acanthopsylla enderleini (Wagner, 1933) was the

most prevalent, being recorded on eight different

marsupial species. The largest number of flea

species reported on a host species was eight on

the bandicoot Microperoryctes longicauda (Peters

& Doria, 1876).

Birds

The Bula Plain / Bensbach area is a focal point

for a variety of birds. It is virtually the only locality

of the province where birds have been examined

for parasites. Thirty-six species were examined

during the surveys - 10 visitors or migrants, nine

endemics, while 17 were resident species, of which

possibly nine may have included both resident and

visitor populations. Forty-seven of 49 individual

birds examined had one or more species of parasite.

They were far more likely to have ectoparasites than

endoparasites, amounting to 56 and 32 species

respectively. Levels of parasitism with ecto- and

endo- parasites amongst resident and migratory/

visitor Species were similar. The two individual birds

that had no parasites were an Australian pratincole

(Stiltia isabella (Vieillot, 1816)) and a sharp-tailed

sandpiper (Calidris acuminata Horsfield, 1821).

Feather or chewing lice were the most

frequently seen ectoparasites. At least 49 species

were recorded, of which 31 have been identified

to species, 18 to genus, while the remainder

await identification. The Australian white ibis,

Threskiornis molucca (Cuvier, 1829), the Torresian

crow, Corvus orru Bonaparte, 1850, the wandering

whistling duck, Dendrocygna arcuata australis

(Reichenbach, 1850) and the curlew sandpiper,

Calidris ferruginea (Pontoppidan, 1763) each had

four different species of chewing lice. No avian host,

however, was particularly heavily infested. Of those

lice identified to species, Actornithophilus umbrinus

(Burmeister, 1838) was the only one found on

three host species, namely, shorebirds (Calidris

spp.). These shorebirds (order Charadriiformes)

were parasitized also by members of the same

genera of other lice but, with current identification,

pp. 281-327

usually different species. One louse, Holomenopon

leucoxanthum (Burmeister, 1838), recovered

from a wandering whistling duck, has a reputation

overseas of rendering feathers of water birds no

longer waterproof, a condition that can lead to

pneumonia.

The double-wattled or southern cassowary

(CasuariuS casuarius (Linnaeus, 1758)) Is

parasitized in the Bensbach area by a species of

chewing louse, Therodoxus oweni Clay, 1971, which

is considered unusual as members of the family

Boopidae are parasites of mammals. Its recovery

on more than one occasion and the presence of

both sexes, precludes the possibility of accidental

infestation. Also exceptional was the discovery of

the mammalian scabies mite, Sarcoptes scabiei

(Linnaeus, 1758) infesting the head and neck region

of a cassowary in the Balimo area. Again, finding

a similar case in Southern Highlands Province

precludes accidental infestation. The only ticks

found on birds were Amblyomma papuanum Hirst,

1914 on a cassowary and a megapode (Talegalla cf.

fuscirostris Salvadori, 1877), and Haemaphysalis

bancrofti Nuttall et Warburton, 1915 on a Torresian

crow. Two insects were recovered from birds: a flea

Xenopsylla vexabilis Jordan, 1925 on a rufous-

bellied Kookaburra (Dacelo gaudichaud (Quoy et

Gaimard, 1824) and a louse fly (Icostia sp.) on an

Australian white ibis.

Nematod species (16) were the most

numerous bird endoparasites, followed by cestodes

(12), digeneans (12) and acanthocephalans

(three). No endoparasites were recorded in 17

host species but, as the examinations were carried

out under field laboratory conditions, very small

or single worms might have been overlooked.

The nematodes included three (possibly four)

species of Contracaecum Railliet et Henry, 1912

and two (possibly three) of Diplotriaena Railliet et

Henry, 1909. It is noteworthy that only one of nine

species of shorebirds (Charadriiformes) contained

a nematode parasite. The most common cestodes

were members of the family Dilepididae, that

included Paradilepis Hsu, 1935, Parvitaenia Burt,

1940, and Megacirrus Beck, 1951. Diverse species

of digeneans were recorded in different hosts, with

members of only one genus (Apatemon) occurring

in more than one host species (ducks). Few blood

samples were collected and no parasites were

detected. This reflects on the low number examined

probably, as Ewers (1967) found 37% parasitism in

the blood of birds in another PNG lowland locality.

Cuckoos and koels are well known for

parasitizing other birds. About 12 species

ay & 305

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

(disregarding vagrants) have been recorded in

WP, of which at least eight are resident/endemic

practicing brood parasitism (Table 2). The remainder

are non-breeding visitors (Beehler & Pratt 2016).

Most sightings come from the northern half of the

province where the avifauna is most studied by

bird watchers. In literature available to the author,

it appears there is only one published record of

cuckoo eggs being found in a host-bird nest in WP

- namely, in nests of the Large-billed Gerygone or

warbler (Gerygone magnirostris Gould, 1842) in

the Middle Fly area (Rand & Gilliard 1967). The

Little Bronze Cuckoo, Chalcites (Chrysococcyx)

minutillus (Gould, 1859) was the likely culprit as it

was the only cuckoo species seen in the area at

the time. There is another record in PNG of this

cuckoo parasitizing a species of honeyeater near

the coastal town of Vanimo, Sandaun Province of

New Guinea (GOmez & Garza 1993).

Reptiles

Four species of monitor lizards or goannas

(Varanus spp.) (nine individuals were examined)

inhabit Bula Plain; all carry ticks of the genus

Amblyomma. Of the seven species of nematodes

recovered, only one, Tanqua tiara (Linstow, 1879),

has been identified to species. Sparganum, the

larval stage of a tapeworm, Spirometra sp., occurred

in the muscle of V. gouldii (Gray, 1838), and V.

Salvadorii (Peters et Doria, 1878). An unidentified

pentastomid parasite was present in the lungs of

V. salvadoril.

Crocodiles of WP have been little examined

for parasites but the freshwater Crocodylus

novaeguineae Schmidt, 1928, of Lake Murray

is known to have two nematode parasites

(Dujardinascaris sp. and Mlicropleura_ sp.), a

digenean (Exotidendrum sp.) and an ectoparasitic

leech (Placobdella sp.). A flatworm (Cyathocotyle

crocodili Yamaguti, 1954) has been recorded in

animals from Lake Murray and the Daru area, and

a protozoan infecting red blood cells in crocodiles

from the latter locality.

There is a dearth of knowledge of parasites

of other reptiles in WP. A monogenetic trematode,

Polystomoides sp. is recorded from the bladder

of a freshwater turtle, Chelodina rugosa Ogilby,

1890 at Awaba, Aramia River (Fairfax 1990), and

a nematode, Tanqua ophidis Johnston et Mawson,

1948 in the stomach of an Arafura filesnake,

Acrochordus arafurae McDowell, 1979, from the

Bensbach River. The stomach of a filesnake from

the Fly River at Suki had large numbers of, probably,

306

the same nematod species (Pinney 1976).

Amphibians

There are few records also of parasites of WP

frogs. Leeches, Leiobdella singularis (Richardson,

1975), are reported as internal parasites of

Papurana grisea (van Kampen, 1923), Nyctimystes

kubori (Zweifel, 1958), N. narinosa Zweifel, 1958,

Litoria angiana (Boulenger, 1915) and L. becki

(Loveridge, 1945) in the Star Mountains (Mann

& Tyler 1963; Van der Lande 1994). They occupy

either the space between the skin and body wall

(dermal lymph sacs), or the body cavity; one or two

may be present ina frog. The means of entering and

leaving the host is through the cloaca, according to

Tyler et al. (1966).

Fishes

Parasites of fishes, likewise, have received

little study. Barramundi (Lates calcarifer (Bloch,

1790)) harbour four species of nematodes, one

being the larvae of Dujardinascaris Baylis, 1947

which, as adults, live in crocodiles. Parasitic

copepods belonging to the genera Ergasilus

Nordmann, 1832 and Lernaea Linnaeus, 1758

occur on barramundi, saratoga (Scleropages

jardinii (Saville-Kent, 1892)) and catfish (Arius

sp.) of the Bensbach River. A Papuan black bass

(Lutjanus goldiei Macleay, 1882) caught at Balimo

had an elongate, thread-like didymozoid flatworm

coiled in the host’s musculature. Two species of

ectoparasites (monogenean trematodes) were

recorded on the sawfish, Pristis microdon Latham,

1794, in Lake Murray, as well as two nematode

species (Ogawa 1991).

Marine fishes, including a species of tuna, a

king threadfin salmon (Polydactylus macrochir

(Gunther, Siphonaptera)) and a_- salmonid

(Eleuthronema sp.) from the WP coast, contained

trypanorhynch plerocerci and the latter host, an

unidentified acanthocephalan. The musculature

of an unidentified marine fish contained a

tangled mass of nematode larvae that could not

be identified, and a copepod (Ergasilus sp.) was

attached to the gills.

Insects

Lepidoptera in PNGare known to have parasites

and parasitoids, but none has been described

from WP. Nevertheless, several butterfly species

known to be infected elsewhere in the country,

02-Jun-21 21:50:25

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Owen, I. L.: A brief review of animal parasites recorded in Western Province, Papua New Guinea

occur in the province, such as, Delias aruna inferna

Butler, 1871, D. callima satura Jordan, 1930,

Hypochrysops arronica (Felder et Felder, 1859),

Candalides helenita dimorphus (Rober, 1886),

Zizula hylax dampierensis Rothschild, 1915, and

Euploea batesii rotunda Van Eecke, 1915 (Parsons

1999). Amongst the parasites and parasitoids

found associated with them, and likely to occur

in WP, is a midge, Forcipomyia papuensis Lane et

Cotman, 1986, that feeds on the haemolymph of

adult males of high altitude Delias spp. by tapping

the anal vein on the underside of the hindwings. Its

legs have structures adapted to grip the host’s wing

scales. The butterflies are prone to these attacks

while drinking. The most common parasitoids of

eggs, larvae and pupae of these butterflies are

small chalcid wasps, and tachnid flies. In some

cases, they can lead to a 50% or higher mortality of

the host (Parsons 1999).

The Asian hive bee, Apis cerana Fabricius,

1793, entered the northern part of PNG from Papua

Province (Indonesia) in the 1980s and, since then,

it has spread to all mainland provinces, including

WP. The bees brought with them a mite, Varroa

jacobsoni Oudemans 1904, which reproduces in

the drone brood cells of the bee colony. It infests

worker brood cells also but does not reproduce

in them. Nevertheless, the mite can lead to a

mean worker brood mortality of 38% in PNG, thus

affecting the wellbeing of an Asian bee colony.

Parasites with no named hosts (Table 2)

The limestone country of the Star Mountains

has an extensive network of caves, where

speleologists reported finding an _ unidentified

temnocephalid (a commensal) attached to the

carapace of freshwater crabs (Chapman 1976).

It was speculated that they fed either on small

cyclopoid copepods or on the haemolymph of the

host crabs. The caves also had leeches, including

a ‘pink coloured’ species that may, possibly, be

Leiobdella jawarerensis Richardson, 1974, which

occurs in caves, feeding on bats, elsewhere in PNG,

and another unidentified ‘pimply skinned’ terrestrial

leech. None was attached to a host. Lukwi caves on

Ok Ma, a tributary of Ok Tedi, had rows of leeches

on the roof of a cave, about 50 metres in from the

entrance, which the cavers believed were waiting

for the return of roosting bats to get their meal

of blood. Unidentified ticks were present on leaf

platforms in caves used as sleeping quarters by

cuscus (Phalanger sp.) (Chapman 1976).

Aulicid wasps (Hymenoptera) are

pp. 281-327

endoparasitoids of wood-boring wasps and beetles;

one species, Pristaulacus kiunga Jennings et

Austin, 2006, has been reported near Kiunga, but

no host was identified (Jennings & Austin 2006).

Female mosquitoes are temporary parasites

requiring at least one blood meal before they can

lay eggs. The anthropophilic species are vectors of

human malaria and filariasis in the province. Other

species are more zoophilic, preferring non-human,

domestic animal or wildlife hosts such as the agile

wallaby and flying foxes (Charlwood et al. 1985;

Van den Hurk et al. 2003). Some are opportunistic

feeders; Colless (1959) reported that females of the

nominal species of the Culex sitiens Wiedemann,

1828 subgroup fed mainly on birds and pigs, but

were also known to bite cattle, dogs and man.

The Flavivirus that causes Japanese encephalitis

in humans was isolated from members of this

Subgroup of mosquitoes at Lake Murray, Balimo

and Abam (inland of Daru) (Johansen et a/. 2000).

The virus can be transmitted to certain wading

birds (Ardeidae) that act as important reservoirs,

and to pigs that serve as the primary amplifying

host (Hanna et al. 1996). Culex annulirostris Skuse,

1889 is a vector of Dirofilaria immitis (Leidy, 1856)

of dogs.

Amongst anopheline mosquitoes recorded in

the province, Anopheles bancroftii Giles, 1902, in

particular, is zoophilic (Keven et al. 2017) while

A. hinesorum Schmidt, Foley, Hartel, Williams et

Bryan, 2001 and A. /ongirostris Brug, 1928 are

considered to be primarily zoophilic (Cooper et al.

2009) or, with other species, such as A. farauti

s. s. Laveran, 1902 and A. punctulatus Do6nitz,

1901, opportunistic feeders (Keven et al. 2017).

Provincial surveys of anophelines by Cooper et al.

(1997; 2002) recorded 10 species that differed

in their distribution. A. farauti s. s., for example,

was confined to the coast, A. longirostris to the

mountainous NW corner, and A. bancroftii largely

to the Middle Fly region. Only A. hinesorum had

a province-wide distribution. The presence of the

filarial nematode Setaria thomasi Sandosham,

1954 in pigs indicates that its mosquito vector

(probably an anopheline) exists in WP.

Tabanid flies (horseflies), feed on animal blood

and, when doing so, inflict painful bites, leading

to avoidance actions by many animals. Some act

as carriers of protozoan blood parasites. At least

20 tabanid species have been recorded in WP

(Mackerras 1964) (Table 2).

307

Ree

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Table 2. Bird, arthropod and annelid parasites and parasitoids with no named hosts, recorded

in Western Province, Papua New Guinea.

Legends: Key to bird residency status: E = endemic; R = resident; R/V = resident & visitor;

V/°R = visitor & possibly resident].

Birds (Cuculiformes, Cuculidae Black-capped Cuckoo or Dwarf Koel Microdynamis Ok Menga, Ok Tedi, Kiunga, Fly

parva (Salvadori, 1875b) River

Eastern or Common Koel Kiunga, Tabubil hill forest, Fly

Eudynamys subcyanocephalus Mathews, 1912 River, Morehead

V/OR

Long-billed Cuckoo Kiunga

Rhamphomantis megarhynchus (G. R. Gray, 1858)

White-eared Bronze-Cuckoo Ok Tedi hill forest

Chalcites meyerii (Salvadori, 1874)

Channel-billed Cuckoo Kiunga, Fly River

Scythrops novaehollandiae Latham, 1790

V/2R

White-crowned Cuckoo Ok Tedi hill forest

Caliechthrus leucolophus (Muller, 1840

( )

Chestnut-breasted Brush-Cuckoo Tabubil, Fly River

Cacomantis castaneiventris Gould, Siphonaptera

Fan-tailed Cuckoo Higher altitudes of Ok Tedi

Cacomantis flabelliformis (Latham, 1801)

R/V

Grey-breasted Brush-Cuckoo Kiunga, low/mid-levels of Ok Tedi

Cacomantis variolosus variolosus Vigors et Horsfield,

1827

R/V

Feramin (Star Mountains

a ae —aa en oo ea any

YT rrrr—<“sSOC—S—C‘is

_ SSS

——————!

——SSSS——————S—SSS=S=i

i al F

fF tsi‘; CL iDasybasis Sermanica (Ricardo, 1915 Rouku, (Morehead District

1926

SSS

Laz

aT a. ee

ly River

Mesomyia (Perisilvius) demeijerei (Ricardo, 1913) Upper Fly River, Lake Daviambu,

Oriomo

[ParasiteGroup ss C—“‘dCPavasite == C“‘C‘C‘éd WLoweality,

ade (Cacali | Tey

i Anopheles hinesorum Schmidt, 2001 (previously A.

farauti No 2

id Anopheles torresiensis Schmidt, 2004 (previously A.

farauti No 3 River

a ce a Pe

rivers

Sn a ae St eae sae

tributaries

Anopheles sp. nr. punctulatus

Lake Murra

Anopheles longirostris Brug, 1928

Anopheles meraukensis Venhuis, 1932

Anopheles novaguinensis Venhuis, 1933

Culex annulirostris Skuse, 1889 Lake Murray, Balimo, Daru,

Wando

308

Book4.indd 308 02-Jun-21 21:50:25

Book4.indd 309

Owen, I. L.: A brief review of animal parasites recorded in Western Province, Papua New Guinea

pp. 281-327

Table 2 (continuation)

Culex sitiens Wiedemann, 1828 subgroup

Insects (Diptera: Hymenoptera Pristaulacus kiunga Jennings et Austin, 2006

Discussion

Western Province was connected by land with

northern Australia until about 8000 years ago

when a rise in Sea level Severed the connection and

Torres Strait was formed (Chappell 2005). It follows

that some animals, as well as their parasites,

are common to both localities. For example, 19

helminths, identified to species, are recorded from

the agile wallaby in WP, of which 17 are found in

animals in Australia (see Spratt et al. 1991) but, of

arthropods, only one species (a tick) is known to be

shared (see Speare et a/. 1983) - with the addition

of the two species of ‘commensal’ dung beetles.

Monitor lizards on either side of the Torres Strait,

also, are known to share nematode parasites, such

as Tanqua tiara (von Linstow, 1879). With more

studies, particularly in WP, more species common

to both countries may be found.

A notable feature of the parasite fauna of rusa

deer, wild pigs and agile wallabies on Bula Plain

is that, although large numbers graze together,

there is no interchange of ecto- or endo-parasites

between the three groups. While many parasites,

like these, demonstrate strong host specificity,

utilizing only one or closely related species, others

have a broad range of hosts. The parasite with the

widest range of hosts within the country as a whole

is the cat flea, Ctenocephalides f. felis (Bouché,

1835), recorded on five different domestic animals,

a rodent, a bat, a marsupial, a bird, as well as

man (Owen 2011), but it has been found only on

dogs and pigs in WP. This could reflect on a lack

of observation. The fleas, Acanthopsylla enderleini

(Wagner, 1933) and Orthopsylloides uncinata

Hastriter, 2014 have a broad range of hosts in WP’s

northern border area, the former being recorded

on 10 species of rodents and eight species of

marsupials and the latter on eight species of

rodents. The protozoan, Sarcocystis spp. used

five different marsupials as intermediate hosts in

southern WP, while a tick, Haemaphysalis bancrofti

Nuttall et Warburton, 1915, parasitized a bird and

three marsupial species. The larval (Sparganum)

stage of Spirometra erinacei (Rudolphi, 1819),

a tapeworm of dogs and cats, was found in two

varanid species and possibly wild pigs but, so far,

the adult worm has not been discovered.

The parasite fauna of many animal groups in

WP, particularly amongst lower vertebrates and

invertebrates, have received little attention, and

the low number of parasites recorded for many

other host species in this account is due, probably,

to the limited number of individuals examined. No

doubt, many parasites, remain to be discovered.

This is exemplified by the report of a biodiversity

assessment survey of the Hindenburg Wall (part

of the Hindenburg Range in the Star Mountains

region) (Fig. 2) undertaken in 2013 by a team of

international and PNG scientists (Post Courier

2013; Richards & Whitmore 2015). (The area forms

part of a proposed series of UNESCO World Heritage

Sites in PNG.) Of 656 animal species documented,

an estimated 66 species are reported as probably

(or Suspected to be) new to science, potentially

serving as hosts to a range of parasites. WP can be

a fruitful ground of study for future biologists and

parasitologists.

Over-exploitation of the environment by man,

however, is a potential problem. Reports in a local

newspaper (Post Courier 2012) of the depletion

of wildlife due to over-hunting in the Bula Plain /

Bensbach area and, more recently (Post Courier

2020), the uncovering of a poaching ring with a

cache of 250 deer antlers and a “huge amount”

of dugong bones at Weam, do not auger well for

the future if the current trend continues. Planned

expansion of mining in the Star Mountains, and

logging elsewhere in the province, can also be a

concern.

WP has a high biodiversity factor, and its

rich wildlife is partly due to the extensive, largely

undisturbed, forest cover, and partly to the

sparseness of human population. As the parasites

of WP fauna form part of this rich biodiversity, it

begs the question, when considering conservation

of species, should parasites be given the same

consideration as their hosts, aS advocated by

309

02-Jun-21 21:50:25

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

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02-Jun-21 21:50:26

Book4.indd 310

Owen, I. L.: A brief review of animal parasites recorded in Western Province, Papua New Guinea

pp. 281-327

Legends for figure 4: 1 - Head end of a hookworm (nematode); 2- A larva (cercaria) of a flatworm (digenetic

trematode); 3 - A flea; 4 - A tick, such as a cattle tick; 5 - An egg capsule containing eggs of Dipylidium caninum

(tapeworm); 6 - A ciliated protozoan, such as Balantidium; 7 - A long, slender, filarial nematode, such as Setaria;

8 - An egg of a thorny-headed (acanthocephalan) worm; 9 - A thorny-headed worm, such as Macracanthorhynchus;

10 - Amite; 11 - Alouse, such as Haematopinus; 12 - A mite, Sarcoptes, in its burrow in the skin of its host; 13 -A

monogenean (ectoparasitic trematode); 14 - A flagellate protozoan parasite; 15 - A bedbug; 16 - A leech; 17 - An

egg of Ascaris (nematode); 18 - A thick-walled oocyst of a coccidian (protozoan) parasite of the gut, such as Eimeria

bukidnonensis of cattle; 19 - A parasitic larva (or maggot) of an insect, such as the screw-worm fly; 20 - The typical

form of a parasitic nematode infecting the gut of a host; 21 - A nematode egg containing cells divided to form a

larva; 22 - The typical form of a digenetic flatworm, such as the liverfluke, Fasciola hepatica; 23 - A pentastomid,

a lung parasite, particularly of snakes; 24 - Trichinella papuae larvae (nematode) in the muscle fibres of a host; 25

- An egg of an insect parasite, such as a botfly of horses, attached to a host’s hair; 26 - Atypical tapeworm; 27 -A

copepod, such as Lernaea, that parasitizes the gills or skin of fishes; 28 - Protozoan parasites, such as Theileria,

inside red blood cells; 29 - A typical infective larval stage of a nematode gut parasite, such as Oesophagostomum.

vy ee Soll

ss tn Es : ¥ }

“a > 4 6 —/_

pi 5 fon Ve '

A -_ ‘- 7 y Gn i

eee 2) Sis 7

y ae a

H a

- 7. _

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h r -

Figure 5. Some Papua New Guinea / Western Province domestic animals illustrated, with artistic liberty, by John

Siune (PNG) [artist’s Tok Pisin captions: dok (dog), meme (goat), hos (horse), bullmacau (bull/cow), pik (pig), pato

(duck), kKakaruk (cockerel/chicken), sipsip (Sheep), pwsiket (cat)].

& 311

Book4.indd 311 02-Jun-21 21:50:28

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figure 6. A range of Papua New Guinea / Western Province fauna illustrated, with artistic liberty, by Jonn Siune

PNG) [artist’s Tok Pisin captions: bilakbokis (fruit bat/flying fox), taragau (bird of prey), domu siaburu (long-beaked

echidna - a Simbu language name), kapul ( cuscus), rokrok (frog), paradais pisin (bird of paradise), traisel (turtle),

geko (gecko), pukpuk (crocodile), sinek (snake), makani (wallaby - magani in Motu language), guria (goura

pigeon), muruk (cassowary)].

312

Book4.indd 312 02-Jun-21 21:50:29

Book4.indd 313

Owen, I. L.: A brief review of animal parasites recorded in Western Province, Papua New Guinea

pp. 281-327

Figures 7-10. Parasites from the Western Province, Papua New Guinea. 7 - A population of the heartworm, Dirofilaria

immitis, recovered from the heart and pulmonary artery of a dog in Papua New Guinea; 8 - A population of the

digenetic flatworm, Fischoederius elongatus, attached to the lining of the rumen (1° stomach) of a Bula Plain rusa

deer, Cervus timorensis; 9 - A stained section of muscle tissue of a wild pig from Bula Plain showing a Trichinella

papuae larva (arrowed) within an individual muscle fibre. The absence of a cyst around the larva in this species can

make detection difficult; 10 - A portion of the small intestine of a wild pig from Bula Plain opened to show thorny-

headed (acanthocephalan) worms, Macracanthorhynchus hirudinaceus, attached to the gut lining.

Smales (1994), Windsor (1995), and Durden &

Keirans (1996)? Also, to quote Dunn et al. (2009),

in the context of coextinction - “Although, in the

popular view, parasites tend to lack charisma, their

loss bears consequences perhaps just as great as

those of the loss of their hosts”.

Acknowledgements

Limited resources in PNG has meant that

much of the identification of parasites was done

by specialists in overseas institutions. My thanks,

in particular, go to the staff of the Parasitic Worms

Section of the British Natural History Museum,

London, namely, David Gibson, Rod Bray, Eileen

Harris and the late Stephen Prudhoe, and to

lan Beveridge of the Faculty of Veterinary and

Agricultural Sciences, University of Melbourne,

for their assistance over many years. Grateful

acknowledgement is made also of the assistance

given by the staff of other sections of the Natural

History Museum, London, United Kingdom, as well

as parasitologists, such as, the late Nixon Wilson

and the late T. C. Maa, then of the Bishop Museum,

Honolulu, Hawaii, U.S.A., and specialists in other

institutions in Australia, New Zealand, Germany,

Canada and U.S.A.

In PNG, the technical assistance of the late

Columba Awui, National Veterinary Laboratory, is

acknowledged as is the assistance given during

the surveys by Eric Lindgren, Geoff O’Leary and

Geoff Sadler of the then Wildlife Section of DASF,

Asmo Pisau, Health Extension Officer, Morehead,

villagers of the Bula Plain and staff of the Bensbach

Lodge. Thanks also go to Frank Bonaccorso, James

Menzies and Lester Seri, for allowing access to

examine specimens in their respective institutions

or collections, to Eric Lindgren for the use of Fig. 3,

and John Siune’s unique artistry is acknowledged.

The great help received from staff of the

Science Reading Room, British Library, London,

313

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Book4.indd 314

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

in obtaining and accessing obscure references

during the preparation of this paper, is gratefully

acknowledged. Thanks go also to Lesley Smales and

Simon Reid in Australia for supplying references.

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S27

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TELNov, D. & PotHemus, D. A.: A new Tanycricos La Rivers, 1971 (Heteroptera: Naucoridae) from Cyclops Mountains ...

pp. 329-344

A new Tanycricos La Rivers, 1971 (Hemiptera:

Heteroptera: Naucoridae) from the Cyclops Mountains,

New Guinea, with a key to species

urn:lsid:zoobank.org:pub:5D5D 7/F1C-5CAF-42BF-9216-2B3908/7CC7/53

Dmitry TELNov 2:2? & Dan A. PoLHemus *

1 - Department of Life Sciences, Natural History Museum, SW7 5BD, London, United Kingdom;

anthicus@gmail.com (ORCID: O0O00-0003-3412-0089)

2 - Institute of Biology, University of Latvia, O. VacieSa iela 4, LV-1004, Riga, Latvia

3 - Coleopterological Research Center, Institute of Life Sciences and Technology, Daugavpils

University, Vienibas iela 13, LV-5401, Daugavpils, Latvia

4 —- Department of Natural Sciences, Bernice P. Bishop Museum, 1525 Bernice Street, Honolulu,

96817, Hawaii, U.S.A.; bugman@bishopmuseum.org (ORCID: OO00-0001-6132-6475)

Abstract: Tanycricos cyclops sp. nov. from Cyclops Mountains, northern New Guinea, is described and illustrated.

An updated key and annotated checklist to the species of Tanycricos La Rivers, 1971 is also provided. The present

range of Tanycricos is discontinuous, with seven of the nine known species inhabiting New Guinea’s 1300 km long

and about 150 km wide fold-and-thrust belt (Central Cordillera), one species occuring on a microcontinent (Arfak

Mountains on Doberai Peninsula), and yet another one on an accreted island arc terrane (Cyclops Mountains of

northern New Guinea).

Key words: Taxonomy, new species, area of endemism, New Guinea, annotated checklist.

Introduction

Tanycricos La Rivers, 1971 is a genus of

creeping water bugs, or Naucoridae, and a

member of the tribe Tanycricini La Rivers, 1971,

which is endemic to New Guinea. The genus was

originally proposed to hold four species from

the Central Cordillera of New Guinea (La Rivers

1971) and considered endemic to the montane

perennial streams of this largest tropical island

(D. Polhemus & J. Polhemus 1986; D. Polhemus

1999a). Tanycricini was established by La Rivers

(1971: 3) within the Cheirochelinae Montandon,

1897 based on a set of external morphological

features, including the absence of an “antennal

groove,” the head with the “posterior interocular

head space a simple, variably-depthed curve”, the

“elongated shape” of the body, the presence of an

“oval, static sense organ along the ventral lateral

margin of prothorax near or at the anterior angle”,

and the “middle tibia armed with at least of four

terminal transverse rows of spines”. In fact, most

of these features are plesiomorphic within the

Naucoridae, as subsequently noted by D. Polhemus

(1999b). Tanycricini currently contains three

Papuan endemic genera - Idiocarus Montandon,

1897, Nesocricos La Rivers, 1971 and Tanycricos

(La Rivers 1971; Schuh & Slater 1995).

Tanycricos was long considered endemic to

Central Highlands of New Guinea (D. Polhemus

& J. Polhemus 1986; D. Polhemus 1999a), until

recently when a new species was described from

the Doberai (or Vogelkop) Peninsula (Sites & Suputa

2008) of far western New Guinea. Up to the present

publication, eight species of Tanycricos were known,

seven of these from the Central Cordillera and one

from Arfak Mountains of the Doberai Peninsula (La

Rivers 1971; D. Polhemus & J. Polhemus 1986; D.

Polhemus 1999a; Sites & Suputa 2008).

In the present paper, another new species,

Tanycricos cyclops sp. nov., iS described and

illustrated based on material from the Cyclops

Mountains, an accreted island arc terrane lying in

the central coastal sector of northern New Guinea,

and a new record for T. acumentum La Rivers, 1971

is presented. An updated, annotated checklist

and updated key to Tanycricos are also presented

and certain biogeographical peculiarities in the

distribution of Tanycricos species are discussed.

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TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Material and methods

All type label text for specimens from

the Naturkundemuseum Erfurt (Germany) is

reproduced with no additions. Each of the type

specimens of the newly described species in the

Naturkundemuseum Erfurt is provided with a

black-framed label on red paper as “HOLOTYPUS”

or “PARATYPUS”, respectively. Authors’ comments

are placed in square brackets.

For specimens collected by the second author

(DP), complete locality data is provided. These

data are contained on labels for alcohol-preserved

specimens, with a more restricted set of data

present on labels for dry-pinned specimens. The

collection number accompanying these data refers

to a collection locality numbering system used by

the second author to cross reference specimens,

label data, photographs, and collection field notes.

For morphological studies, a Leica S6D

binocular stereomicroscope was used. Images were

taken with Canon EOS 77D camera and Canon MP-E

65 mm lens. Helicon Focus ® software was used for

image stacking. Further image manipulations were

conducted using GNU Image Manipulation program

(GIMP). For proper examination the specimens were

relaxed in water. Aodomens were detached and

cleaned for several hours in room temperature 10%

KOH. After the study, genitalia and terminalia were

mounted on mounting cards near their respective

Specimens.

The Papuan Region as considered here is a

zoogeographic term in the sense of Gressitt (1982),

Beehler et al. (1986), Riedel (2002), and Telnov

(2011).

Acronyms for scientific collections:

BPBM - Bernice P. Bishop Museum, Honolulu, Hawaii,

U.S.A.;

NME - Naturkundemuseum Erfurt, Germany;

USNM - United States National Museum of Natural

History, Smithsonian Institution, Washington, DC,

U.S.A.

Results

Tanycricos acumentum La Rivers, 1971 (Figs 1-5)

Material studied: 1 brachypterous 5 NME, INDONESIA

E, New Guinea, Papua Prov., Star Mountains, Oksibil

8.6 km WNW, road to Serambakot vill., 4°52’53”S,

140° 33'22”E, 1500 m, 114.111.2018, perennial stream

midreach, sandy ground / small rocks (NME).

330

Note: This species is reported from most of New

Guinea’s Central Cordillera at elevation over 1000

m (D. Polhemus 1999a). The new locality is the third

one in Star Mountains, located about 30 km south

of the nearest hitherto known locality, “Juliana

Bivak”, which is the locus typicus for this species

(cf. La Rivers 1971). The gradient of the stream on

a road to Serambakot is less than 30%, therefore

we consider it the midreach of a perennial stream

as Classified by D. Polhemus & Allen (2007b). The

habitus and images of the male genitalia of this

species are provided here for the first time.

In Papua New Guinea, introduced rainbow trout

have been documented to prey upon T. acumentum,

with unknown impact on the population of this

Papuan endemic (D. Polhemus & J. Polhemus,

1986).

Tanycricos cyclops sp. nov. (Figs 6-17)

http://zoobank.org/B369E6F2-73C2-45CE-833F-

2C7CC6EDC648

Holotype 4 NME: INDONESIA E, New Guinea, Papua

Prov., Sentani, Cyclops Mts., 02°32’04’S, 140° 30’47’E,

550 m, O3.IV.2018, perennial stream, headwater

[printed] [macropterous specimen].

Paratypes 15 specimens: INDONESIA, New Guinea:

2 macropterous 99 NME: same label as _ holotype;

33, 32 & 6 immatures BPBM, 1 brachypterous 3 &

1 brachypterous 2 USNM, Cyclops Mountains, rocky

stream above Pos Tujuh. NW of Sentani, 260-300 m.,

02°32'26"S, 140°30’47’E, water temp. 22.5° C., 18

September 2000, 14:00-16:30 hrs., CL 7145, D. A.

Polhemus [printed] (1 brachypterous 3, 1 brachypterous

3, 1 macropterous 9, 3 4" instar immatures, 3 3% instar

immatures USNM, all preserved in alcohol) [all of the

above records refer to the same stream, with collections

taken at slightly different elevational ranges along the

catchment].

Derivatio nominis: Toponymic. Named after

Cyclops Mountains, locus typicus of this species.

Measurements: Holotype, macropterous male,

total body length 15.2 mm; head length 2.6 mm,

head width across eyes 3.3 mm, pronotal length 2.5

mm, maximum length from base to anterolateral

angle 3.5 mm, pronotal width across midlength

4.9 mm maximum pronotal width at base 5.8

mm, length from base of the scutellum to the tip

of abdomen length 10 mm, combined maximum

width of wings 6.7 mm.

Description, macropterous male (Fig. 6): Head

and pronotum dorsally dirty yellowish, dark

pattern on head medially covering most of vertex

except narrow area at posteromedial corner of

compound eye, and strongly constricting anteriad

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and subparallel narrow dark line on mesal margin

of compound eye, pronotal disc with dark V-shaped

median and paired parenthesis-shaped sublateral

lines. Scutellum black-brown with dirty yellowish

apex and paired irregular markings on anterior

margin (interconnected or broadly interrupted by

dark colouration). Forewings dark-brown to black-

brown, each with an irregularly shaped, dirty-

yellowish median spot and less prominent pale

spot on each wing at the apex of the scutellum.

Abdominal segments brown dorsally and ventrally,

paler yellowish-brown on terga and sterna VI-VII.

Legs dirty-yellowish to yellowish-brown, profemora

narrowly darkened meso-longitudinally on dorsal

surface. Venter of head, pro- and mesonotum dirty-

yellowish. Head subopaque dorsally and ventrally,

with very large and prominent compound eyes.

Head anterior margin obtusely pointed, projecting

beyond compound eyes 26-27% of head length.

Head basal margin strongly convex between non-

to hardly convergent compound eyes. Compound

eye subtruncate on the inner, irregularly rounded

on the outer lateral margin. Frons slightly raised

dorsally, vertex flattened. Shallow, rather broad

dorsal longitudinal impression at inner margin

of each compound eye. Anteclypeus anterior to

compound eyes gradually converging anteriad; apex

narrowly rounded, dorsally shallowly longitudinally

impressed on either side along slightly convex

midline. Labrum directed anteroventrad, recessed.

Antennae short and slender, most of terminal

antennomere extending beyond anterior margin

of compound eye. Dorsal setae inconspicuous,

mainly arranged in longitudinal impressions along

inner margin of compound eyes. Pronotum

dorsally slightly convex but flattened anteromesad

and broadly along basal margin. Maximum width

across posterolateral angles. Anterior margin

deeply concave, basal margin truncate. Lateral

margins delicately depressed, slightly narrowing

anteriad. Anterolateral angles acutely angulate,

posterolateral angles rather narrowly rounded.

Scutellum basally 4.6 mm wide, exposed part

medially 2.9 mm long, with slightly sinuate lateral

margins, evenly flattened dorsally. Hemelytra

broadly rounded apically, extending towards

genital operculum, not exceeding end of abdomen.

Embolium broad, embolar suture distinct. Clavus

present, claval suture complete, distinct. Faint

venation of hemelytron visible with transmitted

light. Metathoracic wing not folded, four longitudinal

veins and two cells present. Prosternum medially

carinate. Proepimeron with elongate-ovoid, golden

sense organ lateral to procoxal acetabulum, its inner

pp. 329-344

margin touching prosternum, with thin extension

over a deep pit at posterior margin behind procoxa.

Abdomen with terga densely dark setose medially,

glabrous laterally. Terga III-VIIl with marginal row

of short, stout, dirty-yellow, spine-like setae and

Ssubmarginal row of long, golden, suberect setae.

Posterolateral angles of terga Il-IV nearly squared,

V nearly squared on left but acutely angulately

produced on the right angle, VI produced as slightly

acute lobe, VII-VIIl produced as elongate, narrowly

rounded lobes (Figs 6 & 11). Posterior margin of

tergum V distinctly asymmetrically sinuous, with

short and rather deep median emargination,

expanded posteriorly, covering most of tergum

VI in dorsal aspect. Posterior margin of tergum

VI slightly asymmetrically sinuous, of tergum VII -

symmetrical, broadly emarginate medially. Sterna

Il-VIl with spiracle 1/5 distance from _ lateral

margin to midline. Posterior margin of sternum

V subtruncate, VI - sinuous, Vil - emarginate

with strongly projecting apically rounded median

process (Fig. 12). Genital capsule (Figs 15-17)

yellowish-brown, with delicate short and dense

golden setation on exposed areas, broadly rounded

on posterior margin; posterior margin with 3-4

inconspicuous flat transverse ridges. Left paramere

oriented to right, overlapping right paramere and

median lobe, moderately broad and stout in basal

half, strongly narrowed in apical half, slightly angled

anteriad, with slightly concave ventral surface,

rounded apically (Figs 15-17). Right paramere

somewhat Lshaped, broad and stout, narrowed

towards obtusely rounded apex, broadly excavated

dorsolaterally (except at basal part) to embrace

cylindrical median lobe (Figs 15-17). Legs strong.

Profemur delicately tuberculate dorsally, inner

margin with thick pad of short, stiff golden setae,

outer margin all along with long and dense brownish-

golden setae and a row of small black tubercles

posterior of setae. Protibia and single tarsus slightly

curved, dorsally delicately impressed, inner margin

in upper part with thick pad of short, golden setae.

Protarsal claw single, slightly curved. Meso- and

metafemora delicately spinulose on both anterior

and posterior margin; spines are larger and denser

on outer margins. Upper part of posterior margin

of meso- and metatibiae with a row of long dense

golden swimming setae. Meso-and metatibia

dorsally delicately spinulose. Mesotibia dorsally at

distal margin with profuse, long, golden swimming

setae. Metatibia dorsally with row of very long,

dense, brown, suberect swimming setae. Meso-

and metatarsomeres on inner margins with a row

of dense golden swimming setae. Pretarsal claws

Bee

331

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figure 1. Tanycricos acumentum La Rivers, 1971, brachypterous ¢@ from WNW of Oksibil, Star Mountains,

New Guinea, habitus, dorsal view.

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TELNov, D. & PotHemus, D. A.: A new Tanycricos La Rivers, 1971 (Heteroptera: Naucoridae) from Cyclops Mountains ...

pp. 329-344

Figures 2-5. Tanycricos acumentum La Rivers, 1971, 3 from WNW of Oksibil, Star Mountains, New Guinea.

2-3 - Terminal abdominal segments, dorsal (2) and ventral (3) view; 4-5: Genital capsule and aedeagus,

ventral (4) and lateroventral (5) view [not to scale].

3935

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

non-appendiculate, strongly curved, with setiform

parempodia. Length of mesotarsomere 1 - 0.25

mm, 2 - 0.75 mm, 3 (pretarsal claw excluded) - 1

mm, length of metatarsomere 1 - 0.32 mm, 2 -

0.7 mm, 3 - 12 mm.

Macropterous female (Fig. 7): Body length

14.7-15.8 mm. Generally more robust than

macropterous male, abdominal segments wider

than those of male. Posterolateral angles of

terga II-IV nearly squared, without asymmetry,

tergum V - subsquared, less than right angled

as on previous tergum, VI - symmetrical, acutely

angulately produced, VIl - symmetrical, acutely

angulately produced and _ slightly emarginate

preapically on lateral margin (Figs 7 & 13).

Posterior margin of sterna V-VI slightly sinuous to

subtruncate, symmetrical, of sternum VII - slightly

asymmetrically sinuous (Fig. 14). Subgenital plate

broadly subtriangular, sinuous on lateral margins,

with deep, moderately broad, U-shaped median

incision (Fig. 14).

Brachypterous male (Fig. 8): Similar to

macropterous male in general structure and

coloration, with following exceptions: body length

16.5 mm, maximum width (across posterior margin

of hemelytra) 7.6 mm.; head length 2.9 mm, head

width across eyes 3.6 mm, pronotal length 2.5 mm,

maximum pronotal width at base 5.7 mm, scutellum

length 2.7 mm, width across base 4.4 mm, length

of hemelytra along commissure 7.6, length along

outer margin 7.3 mm. Pronotum less swollen and

campanulate than in winged form, posterolateral

angles not as broadly flared; center of scutellum

flat, not swollen; hemelytra extending onto anterior

margin of 6" visible abdominal tergum, coloration

predominantly dark brown with embolar margins

narrowly yellowish-brown, posterior margins slightly

concave anteriorly.

Brachypterous female: Similar to macropterous

male in general structure and coloration, with

following exceptions: Body length 16.7 mm,

maximum width (across posterior margin of

hemelytra) 7.3 mm.

3 instar (Fig. 9): Similar in general form to

adult in regard to mesosternal scuplture and leg

armature, but with wings and scutellum not fully

developed; body length 13 mm, maximum width

(across metathorax) 4.9 mm, head width/length

2.6/1.5 mm, pronotum width/length 4/1.4 mm,

mesonotum width/length 4.7/1.4 mm, metanotum

width/length 4.9/1.2 mm, abdomen width (base)/

length 4.7/4.6 mm, lengths of abdominal terga

I-VIll 0.4/0.8/0.9/0.8/0.5/0.5/0.3/0.4 =mm

respectively. Lateral portions of mesonotum

334

somewhat produced as _ incipient wing pads,

reaching two-thirds of length along lateral margins

of metanotum, lateral mesonotum/lateral

metanotum lengths 1.8/0.8 mm_ respectively;

posterior margin of mesonotum sinuate, produced

into a broad, obtusely convex process medially,

anteriorly depressed basomedially, with triangular

form of scutellum faintly indicated. Suture

between abdominal terga Il and Ill with pair

(1+1) of transversely elongate sulci bearing thin,

golden cuticle. Elongate fovea on ventral side of

anterolateral pronotal angle above fore acetabula

only incipiently developed. Venter of abdomen

completely covered with fine, tightly appressed,

shining golden hydrofuge pile, with round glabrous

patches at spiracles and as paired, longitudinally

elongate patches on each abdominal segment on

both sides of midline.

4‘ instar (the description of the 4‘ instar only

includes differences from that of the 3% instar)

(Fig. 10): Similar in general form to 3” instar, but

larger, with lateral mesonotal wing pads more fully

developed; body length 14.5 mm, maximum width

(across posterior thorax) 6.7 mm, head width/length

3.7/3.3 mm, pronotum width/length 5.0/2.2 mm,

mesonotum width/length 6.7/1.7 mm, metanotum

width/length 6.7/1.3 mm, abdomen width (base)/

length 6.7/6.0 mm, lengths of abdominal terga

I-VIIl 0O.3/0.9/1.1/1.0/0.8/0.6/0.3/1.0 mm

respectively. Lateral portions of mesonotum

produced posteriorly as a pair (1+1) of flaps

representing incipient forewing development,

covering most of lateral metanotum, lateral

mesonotum/lateral metanotum lengths 3.5/0.4

mm respectively. Suture between abdominal terga

ll and III lacking transversely elongate sulci bearing

thin, golden cuticle. Elongate fovea on ventral side

of anterolateral pronotal angle above fore acetabula

partially developed, bearing golden pruinosity.

Differential diagnosis: Tanycricos cyclops sp.

nov. is readily different from congeners primarily

in the shape of male genitalia (parameres, in

particular) and male and female abdominal

terminalia.

Ecology: Specimens were taken in the foothills of

the Cyclops Mountains, at several points between

260 and 550 m elevation along the course of

a clear, steeply descending stream in a bed of

very large boulders, with a high waterfall at the

upstream end of the reach sampled (for holotype).

Specimens were dwelling on boulders at depths of

1-3 cm. Specimens collected by the second author

were taken in kick samples from swift water runs,

occurring amid cobbles and large rocks overlying

02-Jun-21 21:50:34

TELNov, D, & PotHemus, D. A.: A new Tanycricos La Rivers, 1971 (Heteroptera: Naucoridae) from Cyclops Mountains ...

pp. 329-344

Figure 6. Tanycricos cyclops sp. nov., holotype macropterous <, habitus, dorsal view.

& 335

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

a base substrate of sand and gravel, at depths of

5-20 cm. The second author collected this species

in company with other New Guinea endemic

Naucoridae in the genera Idiocarus and Cavocoris

La Rivers, 1971 while the first author - together

with Idiocarus as well.

Distribution: Currently known only from Cyclops

Mountains, northern New Guinea.

Annotated checklist of Tanycricos La Rivers, 1971

species

Species arranged alphabetically in the present

list Since a phylogenetic arrangement is not yet

possible.

Naucoridae

Cheirochelinae Montandon, 1897

Tanycricini La Rivers, 1971

Tanycricos La Rivers, 1971

La Rivers (1971: 4).

Type species: Tanycricos usingeri La Rivers, 1971 by

original designation.

Tanycricos acumentum La Rivers, 1971

La Rivers (1971: 5, 80, 103-104 & 107) original

description, checklist, habitat photographs; Polhemus

& Polhemus (1986: 165) key, additional description,

records; D. Polhemus (1999a: 178) records; D. Polhemus

et al. (2004: 40) note on predators; D. Polhemus & Allen

(2007b: 891) note on predators.

Tanycricos binarius La Rivers, 1971

La Rivers (1971: 8, 80 & 108) original description,

checklist; D. Polhemus & J. Polhemus (1986: 165-166)

key, additional description, records.

Tanycricos cyclops sp. nov.

Described in the present paper.

Tanycricos froeschneri D. Polhemus et J. Polhemus,

1986

D. Polhemus & J. Polhemus (1986: 165 & 171) key,

original description; D. Polhemus et al. (2004: 25)

distribution.

Tanycricos inequalis Sites, 2008

Sites & Suputa (2008: 24) original description.

Tanycricos jaetipi D. Polhemus, 1999

D. Polhemus (1999a: 171) original description.

Tanycricos longiceps La Rivers, 1971

La Rivers (1971: 10, 80 & 109) original description,

checklist; D. Polhemus & J. Polhemus (1986: 165 &

168) key, additional description, records; D. Polhemus

(1999a: 179) record.

336

Tanycricos usingeri La Rivers, 1971

La Rivers (1971: 13, 80 & 110) original description,

checklist; D. Polhemus & J. Polhemus (1986: 165-166)

key, additional description, records.

Tanycricos ziwa D. Polhemus, 1999

D. Polhemus (1999a: 176) original description.

Updated key to Tanycricos La Rivers, 1971 species

The present key is partially based on La Rivers

(1971) and D. Polhemus & J. Polhemus (1986),

incorporating four additional species described

subsequently to their publications.

1 Female left posterolateral angle of abdominal tergum

V in dorsal view distinctly produced and asymmetrical;

posterolateral angle of tergum VI not spinose; male

abdominal tergum VI in ventral view with lateral

projections overreaching beyond lateral margins of

corresponding sternum

- Female abdominal tergum V not or _ weakly

asymmetrical, if slight asymmetry present, then tergum

VI spinose on posterolateral angles; male abdominal

tergum VI in ventral view with lateral projections not

overreaching lateral margins of corresponding sternum

2 Anteclypeus distinctly sinuate on anterior margin,

slightly indentate on either side of apex; inner margin

of mesotibia with erect slender spines; medial hump on

male abdominal tergum VI not dentate apically; male left

paramere strongly narrowed distally; female subgenital

plate bifurcate for half its length, incision narrowing

basally acumentum La Rivers, 1971

- Anteclypeus rounded on anterior margin, without

lateral subapical indentations; inner margin of mesotibia

not spinose; medial hump on male abdominal tergum VI

dentate apically; male left paramere blunt, not strongly

narrowed distally; female subgenital plate bifurcate for

two thirds of its length, incision widening basally

binarius La Rivers, 1971

3 Female abdominal tergum V with posterolateral angles

acutely produced; male abdominal tergum VI with lateral

projections rounded

—- Female abdominal tergum V with posterolateral angles

not produced, squared, nearly squared or rounded; male

abdominal tergum VI with lateral projections squared

off, acute or Subacute

4 Female abdominal tergum VI with posterolateral angles

acutely pointed; male abdominal tergum VI with distinct

median hump on posterior margin; male left paramere

acute angulate apically; female subgenital plate tapering

evenly from base toward apex, with basally widened

incision longiceps La Rivers, 1971

- Female abdominal tergum VI with posterolateral angles

rounded; posterior margin of male abdominal tergum VI

not humped; male left paramere broad, blunt, not acute

angulate apically; female subgenital plate distinctly

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TELNov, D. & PotHemus, D. A.: A new Tanycricos La Rivers, 1971 (Heteroptera: Naucoridae) from Cyclops Mountains ...

narrowing on apical half, very narrowly bifurcate

froeschneri D. Polhemus et J. Polhemus, 1986

5 Female abdominal tergum V with posterolateral angles

squared to rounded; male abdominal tergum VI with

posterolateral angles squared

- Female abdominal tergum V with posterolateral angles

squared to subsquared (slightly less than right-angled);

male abdominal tergum VI with posterolateral angles

acute to subacute (broadly pointed as in the original

description (Sites & Suputa 2008))

6 Total body length under 15 mm; male left paramere

comparatively longer and more slender, lateral tab not

indicated; female subgenital plate with very broad and

deep incision ziwa D. Polhemus, 1999

- Total body length over 17 mm; male left paramere

comparatively shorter and less slender, lateral tab more

or less strongly indicated

7 Total body length 19 mm or over; male left paramere

with distinct lateral tab present; female subgenital plate

with narrow, short, V-shaped incision

BR REE E Sr ne nS ae ob jaetipi D. Polhemus, 1999

- Female abdominal tergum V with posterolateral angles

not produced, squared to rounded; male abdominal

tergum VI with posterolateral angles squared; body large

and robust (total body length over 17 mm); male left

paramere moderately slender, comparatively short, with

marginal lateral tab present, not curved, apex rounded,

hot. bent with regard*to itSiaxiS. ic. etes.h le thee oe

usingeri La Rivers, 1971

8 Female abdominal tergum V with posterolateral

angles subsquared (slightly less than right-angled); male

abdominal tergum VI with posterolateral angles acute;

body less robust, total length under 16 mm; male left

paramere slender, comparatively longer, lateral tab

absent, irregularly curved, with apex rounded and bent

interiad towards median lobe with regard to its axis

cyclops sp. nov.

- Female abdominal tergum V with posterolateral angles

squared; male abdominal tergum VI with posterolateral

angles subacute; body total length under 15 mm; male

left paramere short and broad, lateral tab absent,

obtusely pointed apically inequalis Sites, 2008

Discussion

Tanycricos was long considered to be

restricted to the Central Cordillera of New Guinea

(La Rivers 1971; D. Polhemus & J. Polhemus 1986;

D. Polhemus 1999a; D. Polhemus & Allen 2007b).

Based on the data available at that time (Six Species

of Tanycricos) D. Polhemus (1996) postulated that

ancestors of this taxonomic group likely arrived

via a “Papuan arc”, a terminology introduced by

Kroenke (1984) but not widely utilized by later

authors (for instance, Baillie et al. 2004; Cloos

et al. 2005; Baldwin et a/. 2012; Hall 2012), who

presented alternative terminologies for the same

pp. 329-344

set of terranes accreted in the Eocene (for a review

of the competing models of New Guinea’s tectonic

assembly see D. Polhemus 2007). However, with

discovery of T. inequalis Sites, 2008 in the Arfak

Mountains of the Doberai Peninsula (Sites &

Suputa 2008) and T. cyclops sp. nov. from the

Cyclops Mountains, central northern New Guinea, it

has became evident that the biogeographic history

of Tanycricos is more complex than previously

appreciated. It is possible that Tanycricos may have

evolved on an offshore oceanic arc now accreted

to northern New Guinea as a series of terrains

including the Finisterre, Adelbert, Bewani-Toricelli,

and Cyclops mountains, although the genus is at

present known from only the latter north coast

terrane. Alternately, the original hypothesis of D.

Polhemus (1996) may be correct, with the genus

having evolved on a set of Eocene terranes now

incorporated into the New Guinea central highlands,

with subsequent dispersal to other terranes, such

as Doberai and Cyclops, that were accreted in the

Miocene or later.

Whatever the underlying processes

responsible, the present range of Tanycricos is

discontinuous, with seven of the nine known

species inhabiting New Guinea’s 1300 km long and

about 150 km wide fold-and-thrust belt (Central

Cordillera), one species inhabiting a microcontinent

(Arfak Mountains of the Doberai Peninsula), and

yet another one on an accreted island arc terrane

(Cyclops Mountains).

As earlier postulated by D. Polhemus (1999b),

the endemic complex of Papuan naucorid genera

appears to represent a monophyletic radiation from

a single ancestral stock, and forming an evolutionary

continuum, with the monophyly or paraphyly of the

closely related Sagocorini La Rivers, 1971 with the

Tanycricini not firmly established. For instance,

the Sagicorini and Tanycricini were both identified

aS non-monophyletic groups within the recent

comprehensive molecular phylogenetic study of

aquatic Hemiptera (Ye et al. 2020), although this

was based on limited taxon sampling.

As stated by earlier authors, Tanycricos species

occur ataltitudes over 1000 m (D. Polhemus 1999a;

Polhemus & Allen 2007a & b; Sites & Suputa 2008)

although D. Polhemus (2000) provided a wider

interval of lower altitudes of 500-1000 m for two

unspecified species of Tanycricos corresponding

to T. jaetipi and T. ziwa (D. Polhemus 1999a), from

upper course of the Wapoga River in Indonesian

New Guinea. The present records from Cyclops

Mountains, by contrast, come from 260 to 550 m

elevation, which represents a new lower elevational

337

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pp. 329-344

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bound for the genus. Nevertheless, considering the

extremely steep gradient of the Cyclops Mountains,

which along certain reaches exceeds 75% at the T.

cyclops sp. nov. locus typicus and is nearly 90% a

short distance uphill from this locality (large vertical

waterfall), it should not be excluded that the type

specimens were accidentally washed from higher

altitudes of the mountains, although the collection

of several instar stages of immatures at elevations

as low as 260 m Suggests that this species may be

capable of breeding at such low elevations.

Tanycricos, and members of the Tanycricini

in general, appear to be restricted to perennial

streams (D. Polhemus & Allen 2007b; Sites &

Suputa 2008; current publication). The present

distribution of the genus comprises the Arfak

Mountains (Doberai, Vogelkop, or Bird’s Head,

Peninsula), Central Cordillera and Cyclops

Mountains (mainland New Guinea). It is likely that

orogeny in the Central Range (tectonically defined

as Central Range fold belt in Baillie et a/. 2004)

began in either the Late Cretaceous (100.5-66

Ma) or Early Eocene (54-58 Ma) (Cloos et al.

2005; Davies et al., 1996). Later Melanesian arc

accretion in the Late Oligocene (27.8-24.5 Ma) -

Miocene (23.03-5.333 Ma) (Baldwin et al. 2012)

resulted in emplacement of the Cyclops Mountains

(Cyclops ophiolite belt). The history of the orogeny

in the Arfak Mountains of the Doberai Peninsula is

complicated, involving collision between a fragment

of the Australian craton-stable platform and the

Arfak terrane (Toussaint et al. 2014). Considering

present distribution of Tanycricos and different

geological ages of the Central Cordillera, Arfak and

Cyclops mountains, the sequence by which the

genus colonized these disparate geological units is

not yet fully understood.

Acknowledgements

For the loan of material used in this study we

are highly indebted to Matthias Hartmann (NME).

Special thanks to Paul Brock (Brockenhurst,

United Kingdom) for proofreading the manuscript.

Two referees, Leonidas-Romanos Davranoglou

(Oxford, United Kingdom) and Robert W. Sites

(Columbia, U.S.A.), are also warmly thanked for

improving general quality of the manuscript. The

second author thanks the Indonesian Institute of

Sciences (LIPI) for permission to conduct research

in that country, and Conservation International for

providing logistical assistance in accessing the

Cyclops Mountains.

340

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341

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figures 9-10. Tanycricos cyclops sp. nov., paratypes instars, dorsal view. 9 - 3" instar immature;

10 - 4" instar immature.

0 =~ o

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pp. 329-344

Figures 11-14. Tanycricos cyclops sp. nov., terminal abdominal segments. 11-12 - Holotype 4, dorsal (141) and

ventral, genital capsule dissected (12) view; 13-14 -Paratype 9 (including subgenital plate), dorsal (13) and ventral

(14) view [not to scale].

343

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figures 15-17. Tanycricos cyclops sp. nov., holotype 3’, genital capsule and aedeagus,

ventral (15) and lateroventral (16-17) view [not to scale].

344

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WEIGEL, A. & SKALE, A.: On the taxonomy, synonymy and faunistics of the Apomecynini of the Asian-Australian region ...

pp. 345-362

On the taxonomy, synonymy and faunistics

of the Apomecynini of the Asian-Australian region

(Coleoptera: Cerambycidae: Lamiinae). Part 8

urn:lsid:zoobank.org:pub:445F662C-8431-4C2F-9 74B-1822C39E/78F6

ANDREAS WEIGEL * & ANDRE SKALE 2

1 - Am Schlofggarten 6, D-O7381, Wernburg, Germany; rosalia~aw@gmx.de

2 - Gemeindeweg 6, D-07546, Gera, Germany; andre.skale@online.de

Abstract: In the present work, taxonomic changes are made to a total of 31 species of the tribes Apomecynini and

Desmiphorini. The genus Atelais Pascoe, 1867, currently contains four valid species. The two species Mimosybra

gebeensis Breuning, 1961 and Sybra semilunaris Breuning, 1939 are transferred to this genus. Two new synonyms

are also proposed: Atelais longicornis (Breuning, 1938) = Sybra quadristicta Breuning & de Jong, 1941 syn. nov. and

Atelais semilunaris (Breuning, 1939) comb. nov. = Sybra (Atelais) papuensis Breuning, 1982 syn. nov. Cristosybra

Breuning, 1950 stat. rev. is considered a valid genus within the Apomecynini, with the single species C. cristipennis

Breuning, 1950 comb. nov., and the synonym Mimosybra luzonica Breuning, 1957 syn. nov. is proposed. The

following three Sybra species are junior synonyms to species from the tribe Desmiphorini: Cornallis gracilipes

Thomson, 1864 = Hyagnis sybroides Breuning, 1939 syn. nov. = Sybra longipes Breuning & de Jong, 1941 syn. nov.;

Falsostesilea perforata (Pic, 1926) = Sybra coomani Breuning, 1956 syn. nov.; Mimectatina singularis Aurivillius,

1927 = Sybra apicesignata Breuning et de Jong, 1941 syn. nov. = Mimectatina fuscoapicata (Breuning, 1964) syn.

nov. In this context the synonymy of two additional Mimectatina Aurivillius, 1927 species could be established:

Mimectatina celebica Breuning, 1975 = Mimectatina celebensis Breuning, 1975 syn. nov. The monotypic genus

Laosepilysta Breuning, 1965 (Apomecynini) syn. nov. and the subgenus Trizotale Breuning, 1975 syn. nov. of the

genus Mimozotale Breuning, 1951 (Desmiphorini) are congeneric with Mimozotale. To Mimozotale trivittata (Pic,

1931) the synonyms Laosepilysta flavolineata Breuning, 1965 syn. nov. and furthermore Mimozotale tonkinea

Breuning, 1969 syn. nov. are proposed. The genus Plocia Newman, 1842 currently contains 7 valid taxa, Parepilysta

puncticollis Breuning, 1965 is transferred to this genus. The genus Plociella Breuning is an available name for

Plocia mixta Newman, 1842, because its only species Plociella conspersa (Aurivillius, 1927) is conspecific with

it. The monotypic genus Sybroplocia Breuning, 1959 is also considered synonymous with the species Sybroplocia

sybroides (Schwarzer, 1931). Furthermore, a total of 14 synonyms are proposed for nine Sybra species. Sybra

amboinica Breuning, 1965 is transferred to the genus Orinoeme Pascoe, 1867. Sybra triflavomaculata Breuning,

1975 as synonym to Mimosybra trimaculata Breuning, 1953 and Sthenias leucothorax Breuning, 1938 as synonym

to Xylariopsis mimica Bates, 1884 are proposed. The given “nomina nova” for three Sybra species by Slipinski &

Escalona (2013) are not necessary because of unjustified generic synonymizations, and consequently are placed

in synonymy.

Key words: Apomecynini, Desmiphorini, faunistics, new synonymy, new combinations, revalidation, Oriental Region,

Australian Region.

Book4.indd 345

Indroduction

In the Oriental Region of Southeast Asia,

including New Guinea, the Apomecynini Thomson,

1860 are a very complex tribe for which the

genus system is still unsatisfactory, and in many

parts unclear and confused. In the current paper

we present additional new data within the tribe

Apomecynini (in some cases also for Desmiphorini)

based on recent studies. We propose a total of 34

new synonymies, 6 new combinations and one

revalidation for a genus for a total of 26 listed valid

species. Some of the species dealt with here are

already mentioned in earlier works (Skale & Weigel

2012; 2014; Weigel & Skale 2009; 2011; 2017).

Among the existing material, a large part,

particularly from New Guinea, Malaysia and Borneo,

has not yet been identified to species level. Although

there are probably many undescribed species, we

will not describe any this time. The confusion in the

related groups Is still too great. A number of species

are also described in other tribes (e.g. Desmiphorini

345

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TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Thomson, 1860, Pteropliini Thomson, 1860), some

are listed in the present paper.

Material and methods

For the present study, 36 type specimens

and 98 additional specimens were examined. The

genitalia for most of the male specimens (penis with

endophallus, tegmen, 8th tergite) were examined.

For type material, genitalia were examined only if

the material was in good condition. The dissected

genitalia were fixed onto a white card together with

the specimen.

Photographs of Figs 1 and 18 were taken witha

LM Macroskop / Canon EOS 80D and subsequently

processed and measured wirh Helicon Software

6.8.0. Photographs of figures 2, 5-9, 16, 19, 22,

26-28 and 31-34 were taken with a Leica Z6

APOA stereomicroscope using a Planapo lens 1.0

and subsequently processed and measured with

the Leica Applications software 4.0. Photographs

of Figs. 17, 20, 24, 25, 29 and 30 were taken

with an Olympus Tough 4 digital camera. The final

processing made with Photoshop 7.0.

Total body length is measured from the anterior

border of the clypeus to the apex of elytra and does

not include partially exposed abdominal segments.

Label data of types cited verbatim, using single

Slash to separate rows of the same label and double

slash for separation of different labels. Additional

remarks by the authors are cited in Square brackets.

For non-type specimens, no special delimitation is

given for rows on the labels.

Acronyms of scientific collections:

BM - Bernice P. Bishop Museum, Honolulu, Hawaii,

U.S.A.;

BMNH - Natural History Museum (formerly British

Museum, Natural History), London, United

Kingdom;

CBT - Collection Shinichi Befu, Tokyo, Japan;

CCS - Collection Jim Cope, Sacramento, California,

U.S.A.;

CDB - Collection Lubos Dembicky, Brno, Czech Republic;

CHH - Collection Daniel Heffern, Houston, Texas, U,S,A.;

CHS - Collection Herbert Schmidt, Vienna, Austria;

CLS - Collection Loong Fah Cheong, Singapore;

CMI - Collection Hiroshi Makihara, Ibaraki, Japan;

CSG - Collection André Skale, Gera, Germany;

CTO - Collection Tomas Tichy, Ostrava, Czech Republic;

CVB - Collection Eduard Vives, Barcelona, Spain;

CWW - Collection Andreas Weigel, Wernburg, Germany;

FREY - Collection G. Frey, Basel, Switzerland;

HNHM - Hungarian Natural History Museum, Budapest,

346

Hungary;

JMU - Julius—-Maximilians

Germany;

MHNL - Musée d’Histoire Naturelle de Lyon, France;

MNHN - Muséum National d’Histoire Naturelle, Paris,

France;

NHRS - Naturhistorika Riksmuseet, Stockholm, Sweden;

RBINS - Royal Belgian Institute of Natural Sciences,

Brussels, Belgium;

University, Wurzburg,

RMNH - "Naturalis" Biodiversity Centre, Leiden, the

Netherlands;

SMF - Senckenberg Museum, Frankfurt am Main,

Germany;

SMNS - Staatliches Museum fur Naturkunde Stuttgart,

Germany;

SNSD - Senckenberg Naturhistorische Sammlungen

Dresden, Germany;

USNM - United States National Museum, Washington

D.C., U.S.A.;

ZFMK - Zoologisches Forschungsmuseum Alexander

Konig, Bonn, Germany;

ZMB - Zoologisches Museum der Humboldt Universitat

zu Berlin, Germany;

ZSM- = Zoologische

Germany.

Staatssammlung, Munchen,

Additional abbreviations used:

QO - Female;

3 - Male;

HT - Holotype;

LT - Lectotype;

PLT - Paralectotype;

PT - Paratype;

spm(s) - specimen(s).

Results

Remarks on the genus Atelais Pascoe, 1864

Catalogue of the species

Atelais Pascoe, 1867: 457

Type species: Atelais illaesa Pascoe, 1867 by

original designation.

Atelais gebeensis (Breuning, 1961) comb. nov.

(Fig. 1)

Mimosybra gebeensis Breuning, 1961a: 545

Atelais illaesa Pascoe, 1867: 457 (Weigel & Skale

2009)

Atelais longicornis (Breuning, 1938) (Figs 2-3)

= Sybra (Atelais) longicornis Breuning, 1938b: 61

= Sybra quadristicta Breuning et de Jong, 1941: 105

syn. nov. (Fig. 3)

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WEIGEL, A. & SKALE, A.: On the taxonomy, synonymy and faunistics of the Apomecynini of the Asian-Australian region ...

Atelais semilunaris (Breuning, 1939) comb. nov.

(Fig. 4-6)

Sybra semilunaris Breuning, 1939: 248

= Sybra (Atelais) papuensis Breuning, 1982a: 10 syn.

nov.

Notes:According to the present state of knowledge,

the genus Atelais revalidated by Weigel & Skale

(2009) includes the four species mentioned above,

with two species being transferred to this genus.

There are specimens in collections representing at

least two other Atelais species from West Papua

(Indonesia), which are still undescribed and will be

treated in future studies. Tavakilian & Chevillotte

(2020) list six species in Atelais. The species Sybra

modesta Pascoe, 1865, S. putida Pascoe, 1865

and S. surigaonis Breuning, 1943 do not belong to

this genus. S. modesta is a typical Sybra species,

as already mentioned in Weigel & Skale (2009).

For S. putida, Breuning (1964a) has created its

own subgenus Paratelais. According to a colour

photograph of the HT (BMNH), this specimen

belongs neither to Atelais nor to Sybra (s. str.).

For the time being the subgenus Paratelais will be

retained:

Sybra subgenus Paratelais Breuning, 1964a: 306

Type species: Sybra putida Pascoe, 1865 by

original designation and monotypy.

Sybra putida Pascoe, 1865: 206

Sybra (Atelais) putrida Breuning, 1960: 154 [incorrect

spelling]

Sybra_ (Paratelais) putrida Breunng,

[incorrect spelling]

Atelais putida Tavakilian & Chevillotte (2020)

1964a: 306

Sybra surigaonis also belongs to the genus Sybra

because of its morphology. The mentioned male

specimen from Surigao has already been labelled

by Aurivillius as “Sybra sp.”

Sybra surigaonis Breuning, 1943

Sybra (Atelais) surigaonis Breuning, 1943: 45

Sybra (Atelais) surigaonis Breuning, 1964a: 305

Atelais surigaonis Lingafelter et al. 2014: 330

Type material examined: HT 3 USNM: Surigao /

Mindanao // Type // Sybra / (Sg. Atelais) / surigaonis

/ mihi Typ [all handwritten] / det. Breuning // BLNO /

000985; preservation: completely preserved; length:

7.5 mm (s.a. Lingafelter et al. 2014).

Other studied material: 14 CSG: Surigao, Mindanao,

Baker, 191, Sybra sp. Auriv 27; length: 8 mm.

pp. 345-362

Atelais gebeensis (Breuning, 1961) comb. nov.

(Fig. 1)

Type material examined: colour photograph of the

HT in MNHN.

Other studied material: 19 CWW: Indonesia E,

Maluku Utara Prov., Gebe Island, on the way from Kacepi

to Umera vill., 29.X.-03.XI.2011, secondary & old growth

rainforest on limestone, leg. L. Wagner; length: 12 mm.

Note: The species was described from one

specimen from the Indonesian Moluccan island

of Gebe. The type specimen, which was compared

with the mentioned female specimen (Fig. 1) from

Gebe Island, corresponds to the characteristics of

the genus Atelais and is therefore transferred to

this genus.

Atelais longicornis (Breuning, 1938) (Figs 2-3)

Type material examined: A. longicornis HT 3 RBINS:

Nouv. Guinée / Buolay [handwritten, ?] [glued at the

following label] Coll. R. |. Sc. N. B. 7 Nouvelle Guinée /

ex coll. Le Moult [printed on] // nov. ent. 1938, 8: / 61

[white label, with two glued labels] Atelais / longicornis

/ mihi Typ / det. Breuning // Holotype [red, black

bordered]; preservation: left antennae with eigth and a

half segments only; length: no data (Breuning 1938b);

S. quadristicta: HT 4 (RMNH): Rosenberg / Andai / N.

G. [round label] // 171 // Sybra Pasc. quadri- / sticta

mihi Typ ! / det. Breuning [handwritten] // quadristicta

/ Breun. v. de Jong [handwritten] // Museum Leiden /

(NNM) // Type [red, handwritten], preservation: well

preserved, without left protarsus claw, left antennae

with nine and right with ten segments only; length: 9.7

mm.

Other studied material (25 spms): 1 som CWW:

INDONESIA Irian Jaya, Biak NE, 10km N _ Bosnik,

10.11.1998, prim. Urwald, leg. A. Weigel, KL [umbrella];

12 CWW: INDONESIA Irian Jaya, Biak, 10km N

Bosnik, 136°20’E, 01°05’S, UWP [secondary forest],

13.11.1998, leg. A. Weigel; 16’ CWW: INDONESIA Irian

Jaya, Biak, 10km NW Sjaba, 01°03’S, 135°57’E,

31.X11.1998, Sek.-Wald, leg. A. Weigel, KL [umbrella]; 1

spm CWW: INDONESIA Irian Jaya, Biak, 1O0km NE, Saba

E, 01°0O7’S, 136°18’E, 30.XII.1998, leg. A. Weigel; 2

spms CWW: ditto, 17.1.1999, leg. A. Weigel; 6 soms CSG,

CWW & SMNS: IRIAN JAYA, Biak IsI., Korim, Nernu, 100-

150m, 12.-14.Xll.2000, leg. A. RIEDEL; 1 som SMNS:

ditto, 1-4.1I.2001, leg. A. RIEDEL; 24 & 19 CSG: W-

Papua, Manokwari Prov., L£8km NE Ransiki, 01°21.05’S,

134°12.46’E, 02.-06.II|.2007, leg. A. Skale, cutting

area; 12 CWW: W-Papua, Manokwari Prov., 18km NE

Ransiki, 01°21.05’S, 134°12.46’E, 02.-06.III.2007,

leg. A. Weigel, cutting area; 16° CWW: W-Papua,

Manokwari Prov., 14km NE Ransiki, Warbiati (Oransbari),

01°18.25’S, 134°14.14’E, 02.1I1.2007, leg. A. Weigel,

cutt. area; 2 soms CWW: INDONESIA Irian Jaya, Japen SE,

15km W Serui, 03.1.1999, KL [umbrella], leg. A.Weigel,

UWP [primary forest]; 14° CSG: Warkapi, Manokwari

lrian Jaya, Indonesia, 10.-15.March 2008, leg. Y. Yokoi;

347

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TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

32 CWW: INDONESIA Irian Jaya, Wapoga River E Asori,

km 64 Kwadewa Camp, 2°49’S, 136°28’E, 10.1.1999,

leg. A. Weigel KL [umbrella]; 1 CWW: INDONESIA Irian

Jaya, Asori E, km 60 Kwadewa Camp, nr. Wapoga River,

02°49’S, 136°28’E UWP [primary forest], 10.1.1999,

leg. A. Weigel; length: 8 - 11 mm.

Note: The examination of the two type specimens

has shown that they are conspecific, so S.

quadristicta is a junior synonym of A. longicornis.

Atelais semilunaris (Breuning, 1939) comb. nov.

(Figs 4-6)

Type material examined: colour photographs of the

HT soms of Sybra semilunaris [BMNH] and S. (Atelais)

papuensis 2 in MNHN.

Other studied material: 14 & 19 [CWW)]: Papua

Neuguinea, Morobe prov., Garaina 10km S, 15-1800m,

23.-27.|I1.1998, leg. A. Riedel, length: 6 11.3 mm, 2

12.2 mm.

Notes: The comparison of the two holotype

specimens has shown that both species are

conspecific, so S. papuensis is a junior synonym

of S. semilunaris. Both species are described

from Papua New Guinea (Orori (?) = possibly Oro

province and Mailu Island). Due to morphological

characteristics (not dotted rows on the elytra, long

antennae, pronounced shoulder etc.) the species

is transferred to the genus Atelais. The yellowish

macules on the elytra and pronotum are variable.

Cornallis gracilipes Thomson, 1864: 47

= Hyagnis sybroides Breuning, 1939: 206 syn. nov.

= Sybra longipes Breuning et de Jong, 1941: 104 syn.

nov. (Fig. 7)

= Hyagnis bootangensis Breuning, 1968 (Weigel 2006:

synonymy)

= Parasybrodiboma sikkimensis Breuning, 1969 (Weigel

2006: synonymy)

Type material examined: Sybralongipes HT @ RMNH:

Sikkim / Marz, April / H. Fruhstorfer [black bordered] //

250 // Museum Leiden / Ex. coll. / G. van Roon // Mus.

Leiden // type [red label] // Sybra Pasc. / longipes mihi

/ Typ! det. Breuning [handwritten] // longipes / Breun. &

de Jong [handwritten, black bordered]; preservation: left

antenna with six and right antenna with nine segments

only, right protarsal claw missing, elytra apex shrinked;

length: 6.3 mm.

Notes: The synonymy of Hyagnis sybroides and

Cornallis gracilipes, which was mentioned among

others in Lobl & Smetana (2010) and Tavakilian

& Chevillotte (2020), has not been published yet.

This is possibly due to Weigel (2006) "probably

all three species are junior synonyms of Cornallis

———

Z ~ i, i

= aE :

Sek jie

348

gracilipes ..." This taxonomic act is hereby valid,

Hyagnis sybroides is a junior synonym of Cornallis

gracilipes. The examination of the male type

specimen of Sybra longipes has shown that the

species is conspecific with Cornallis gracilipes and

is, therefore, also a junior synonym.

Cristosybra Breuning, 1950 stat. rev.

Type species: Sybra cristipennis Breuning, 1950,

by original designation.

Sybra (Cristosybra) Breuning, 1950: 268

Sybra Breuning, 1964a: 146 (synonymy)

Cristosybra cristipennis Breuning, 1950 comb.

nov.

Sybra (Cristosybra) cristipennis Breuning, 1950: 268

Sybra (Cristosybra) cristipennis Breuning, 1960: 154

Sybra (s.str.) cristipoennis Breuning, 1964a: 239

= Mimosybra luzonica Breuning, 1957: 10 syn. nov.

(Skale & Weigel 2012)

= Sybra luzonia (Breuning, 1957) (Skale & Weigel 2012)

= Sybra medioflavomaculata Breuning, 1966 (Skale &

Weigel 2012: synonymy)

= Sybra lingafelteri Skale & Weigel 2012 (Weigel & Skale

2017: nom. nov.)

Type material examined: Sybra cristipennis HT

NHRS: Heightsplan / N. Luzon // Typus [red label] // 69

[red] // Sybra / cristipennis / mihi Typ [all handwritten]

/ det. Breuning; preservation: completely preserved;

length: 5.5 mm (Breuning 1950); Mimosybra luzonica

HT (Skale & Weigel 2012); Sybra medioflavomaculata

HT (Skale & Weigel 2012).

Notes: Colour photographs of this species are

given already in Skale & Weigel (2012). Breuning

(1950) has described the subgenus Cristosybra

for Sybra cristipennis: “Par la présence d’une

créte élytrale postbasilaire, surmontée d’un

fascicule de poils cette espéce différe de toutes

les autres espéces du genre et formera le type

d’un nouveau Sous-genre: Cristosybra nov.

Le sous-genre est caractérisé en plus par une

téte rétractile et les fémurs et tibias dépourvus

de poils dressés”. In Breuning’s revision of the

Apomecynini (Breuning 1964a), he synomyzied

this subgenus with Sybra (s. str.), but Tavakilian

& Chevillotte (2020) did not follow. Due to the

characteristics described by Breuning, especially

the presence of a small postbasal crest covered

with a tuft of hair, Cristosybra stat. rev. is regarded

as an independent, hitherto monotypic genus. A

comparison of the types of S. cristipennis and M.

luzonica shows that both species are conspecific,

so M. luzonica is a junior synonym for C. cristipennis

02-Jun-21 21:50:42

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WEIGEL, A. & SKALE, A.: On the taxonomy, synonymy and faunistics of the Apomecynini of the Asian-Australian region ...

comb. nov. A further specimen of S. cristipennis

labeled as PT (NHRS) with the same information

as the HT, does not belong to the type series

according to the description. Sybra bifasciculosa

Breuning, 1956 from Australia, which is also listed

in Breuning (1960) and also Tavaklian & Chevillotte

(2020) under the subgenus Cristosybra, cannot be

assessed at this time, therefore this taxon is still

listed under Sybra according to Breuning (1964a).

Falsostesilea perforata (Pic, 1926)

Stesilea perforata Pic, 1926: 9

= Sybra coomani Breuning, 1956: 686 syn. nov.

= Mimosybra melli Breuning, 1964a (Weigel et al. 2013:

synonymy)

Note: Gérard Chemin (Champigny-sur-Marne,

France) was kind enough to provide a type image

of Sybra coomani (preserved in MHNL) in the

longhorn beetle forum (http://www.cerambycoidea.

com). According to this colour picture it can be seen

that both species are conspecific and accordingly

Sybra coomani is a junior synonym of Falostesilea

perforata. Both species were described from Hoa

Binh (Vietnam).

Mimectatina singularis Aurivillius, 1927 (Fig. 8)

= Sybra apicesignata Breuning & de Jong, 1941: 104

syn. nov. (Fig. 8)

= Sybra fuscoapicata Breuning, 1964b: 304 syn. nov.

(see also Weigel & Skale 2011: new combination)

Type material examined: M. singularis LT USNM:

Island / Samar / Baker // 22776 // 36 // Typus [red,

black bordered] // Mimectatina / singularis / Auriv

"27 Auriv [handwritten] // BLNO 000650 [blue label];

preservation: left antenna with ten and right antenna

with six segments only; length: 14.8 mm; PLT (NHRS):

Island / Sibuyan / Baker // 18473 [handwritten] //

Typus [red, black bordered] // Mimectatina / singularis

/ Auriv. [handwritten] // 134 [bright red] // Collectio

Baker. Riksmuseum in Stockholm ; preservation: well

preserved, complete; S. apicesignata HT 2 (RMNH):

S.Celebes. / Bonthain. / C. Ribbe 1882. [white, black

bordered] // 13 // Museum Leiden / (NNM) // type

[red label] // Sybra Pasc. apice- / signata mihi / Typ !

dét. Breuning [handwritten] // apicesignata / Breun. &

de Jong. [handwritten, black bordered]; preservation:

toment slighly rubbed, left metatarsus missing, left

antenna with nine and right with seven segments only;

length: 10 mm.

Other studied material (6 spms) (Weigel & Skale

2011): 14 CWW: Philippinen, Romblon/Sibuyan, 1988,

local coll.; 19 CDB: Indonesia; Sulawesi-C; 38km SE

Pendolo vill. (pass); 120°46,55’E, 2°14,03’S; Bolm

leg.; 17.vii.1999, 1200m; Collectio - L. Dembicky,

Cerambycidae, Invt. No.: 2250; 1¢ & 19 CDB &

pp. 345-362

CWW: Philippines; Mindoro occid., Amnay river valley;

Z5kKM~ SE” OT, santa Cruze 19" Sf N12 0" 56: EAA

Dembicky.; 17.iv.2000; 180m, Collectio - L. Dembicky,

Cerambycidae, Invt. No.: 2248 and 2256; 19 SMF:

Vivac, S.O Luzon, Senckenberg-Museum Frankfurt/

Main; 1 spm CWW: Filipines-N, Calayan/N-Luzon,

VI.2005, local coll.

Notes: The specimen mentioned above was

designated as LI (preserved in USNM)_ by

Lingafelter, Nearns & Bondi in 2013, but the

corresponding label (Lingafelter et al. 2014: fig.

156m) is not present on this specimen. In addition

Aurivillius (1927) wrote as type locality "Philippines:

Sibuyan. Samar (C. F. Baker)". Two further PLT

exist in the USNM (Lingafelter et a/. 2014). The

examined type specimen in the NHRS is also to be

evaluated as PLT. The examination of the female

type specimen of Sybra apicesignata has shown

that the species is conspecific with Mimectatina

singularis. After studying further material, the

synonymy of Mimectatina fuscoapicata with M.

singularis could also be confirmed as already

Suggested in Weigel & Skale (2011). Both species

are therefore junior synonyms of M. Singularis.

Breuning (1975a) described two other species of

Mimectatina Aurivillius, 1927 from Sulawesi, which

probably also belong to this species. After colour

photographs of the holotypes of both species

(RBINS), kindly provided by Florence Trus (RBINS),

the following synonymy results:

Mimectatina celebica Breuning, 1975 (Figs 10-

15)

Mimectatina celebica Breuning, 1975a: 33 (Figs 10-12)

= Mimectatina celebensis Breuning, 1975a: 33 syn.

nov. (Figs 13-15)

Type material examined: Mimectatina celebica HT

3 RBINS: Celebes / Kintabaroe Paloe / J.P.CH.Kalis. /

600° XII.1936 [glued, black bordered] // Coll. R. I. Sc.

N. B. / Iles Moluques // Mimectatina / celebica mihi /

Breuning dét. Typ // Holotype [red label, black bordered]

// {both labels are glued on:] cf.: Bull.Soc ent. Mulhouse

/ 1976: 512; preservation: left antenna missing,

rigth antenna with four segments only, right protarsus

missing, toment at the pronotum slightly rubbed; length:

13.2 mm; Mimectatina celebensis HT 3 RBINS: Celebes

/ Mendao [= Manado] [glued, handwritten in blue] //

Coll. R. |. Sc. N. B. 7 Iles Moluques // Mimectatina /

celebensis / mihi Typ / Breuning dét. // Holotype [red

label, black bordered] // [both labels are glued on:] cf.:

Bull. Soc. ent. / Mulhouse 1976: 52; preservation: well

preserved, only rigth antenna missing; length: 12.6 mm.

349

02-Jun-21 21:50:42

TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

7 8 9

Figures 1-9. Asian-Australian Apomycinini, habitus in dorsal view. 1 - Atelais gebeensis (Breuning, 1961), 2

from Gebe Island; 2 - Sybra (Atelais) longicornis Breuning, 1938, HT 3 (RBINS); 3 - Sybra quadristicta Breuning

et de Jong, 1941, HT 4 (RMNH); 4 - Sybra semilunaris Breuning, 1939, HT (BMNH); 5 - Atelais semilunaris, 3

(CWW) from Gariana, Papua New Guinea; 6 - ditto, 9 (CWW) from Garaina, Papua New Guinea; 7 - Sybra longipes

Breuning et de Jong, 1941, HT 3 (RMNH); 8 - Sybra apicesignata Breuning et de Jong, 1941, HT 2 (RMNH); 9 -

Mimosybra trimaculata Breuning, 1953, 3 (CWW) from Morobe Province, Papua New Guinea (image 4 courtesy

Lubos Dembicky) [not to scale].

350

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WEIGEL, A. & SKALE, A.: On the taxonomy, synonymy and faunistics of the Apomecynini of the Asian-Australian region ...

Mimosybra trimaculata Breuning, 1953: 105 (Fig.

= Sybra triflavomaculata Breuning, 1975b: 162 syn. nov.

Type material examined: M. trimaculata HT ¢@

HNHM: N. Guinea / Biro 1899. // Sattelberg / Huon

Golf. // Holotypus // 1952 // Mimosybra / trimaculata

Breuning // Mimosybra / trimaculata mihi / Typ

Breuning dét.; preservation: toment well preserved,

complete, only head a bit detached; length: 6-8 mm

(HT + 1 PT, Breuning 1953); S. triflavomaculata HT ¢

HNHM: NEU GUINEA / NE / Lae 4-6.1X.1968. // No.NG-

L-C.16. // leg. Dr. J. Balogh // Holotypus 1974. / Sybra

/ triflavomaculata Breuning [red bordered] // TYPE //

Sybra triflavo- / maculata mihi / Typ Breuning dét.;

preservation: toment well preserved, right antenna with

nine segments only; length: 8 mm, 3 mm wide (Breuning

1975b).

Other studied material: 14 (Fig. 9) & 129 CWW:

Papua Neuguinea, Morobe Prov., Pindiu E, Kobau,

1250-1400m, 24.V1.1998, leg. A. Riedel; length: 3 8.6

mm, 2 8.9 mm.

Notes: The generic assignment of this species

is still unclear. There is a group of similar species

distributed in Papua New Guinea which can not be

assigned to any described genus at present. Such

species with distinct, interspersed strong white

bristles on all femora and tibiae, no teeth on the

internal side of the middle tibia and very scarred

elytral surface have nothing to do with Mimosybra

Breuning, 1939. This species remains for the

time being in the genus Mimosybra, where it was

originally placed by Breuning.

Mimozotale Breuning, 1951: 137 (Desmiphorini)

= Laosepilysta Breuning, 1965a: 31 syn. nov.

(Apomecynini)

Type species: L. flavolineata Breuning, 1965 by

original designation and monotypy.

= subgenus Trizotale Breuning, 1975a: 41 syn. nov.

Type species: Z. trivittata Pic, 1931 by original

designation.

Mimozotale trivittata (Pic, 1931) (Fig. 16)

Zotale trivittata Pic, 1931: 14

= Laosepilysta flavolineata Breuning, 1965a: 31 syn.

nov.

= Mimozotale tonkinea Breuning, 1969: 666 syn. nov.

Mimozotale (Trizotale) trivittata Breuning, 1975a: 41

Type material examined: colour photograph of:

Zotale trivittata HT 2 in MNHN: Hoa Binh (North-

Vietnam), length: 7.5 mm, 1.75 mm wide (Breuning

1975a); Mimozotale tonkinea: HT 2 MNHN: Hoa Binh

(North-Vietnam), length: 9.5 mm, 3 mm wide (Breuning

1969); Laosepilysta flavolineata: HT 3 BM: Laos:

pp. 345-362

Vientiane, length: 7-8 mm (Rondon & Breuning 1970).

Other studied material: 12 CWW: N-Vietnam, Bac

Kan Prov., Ba Be National Park (entry), 16.-20.V.2014,

22°25'07’"N, 105° 38’09’E, 180-220m, leg. A. Weigel,

length: 7.6 mm (Fig. 16); 12 CWW: N-VIETNAM, Thai

Nguyen Prov., vic. Ngoc Thanh, Le Minh (IEBR station),

12.V.2012, 21°23'3.43"N, 105° 42'43.77"E, 60m, leg.

A. Weigel. length: 8.6 mm.

Notes: According to the colour photographs of

the holotype specimens of these three species, L.

flavolineata and M. tonkinea are conspecific with

M. trivittata and are therefore junior synonyms of

M. trivittata. Breuning (1975a) has described the

subgenus Trizotale. The differentiation compared

to the nominal Mimozotale (s. str.): “3rd antenna

segment longer than the scape, as long or a

little longer than the 4th antenna segment” is

completely insufficient and not comprehensible.

Accordingly, we consider the subgenus Trizotale to

be congeneric with Mimozotale, which is therefore a

Junior synonym. The monotypic genus Laosepilysta

is therefore also a junior synonym of Mimozotale.

Breuning (1965a) placed Laosepilysta among the

Apomecynini, but only the species L. flavolineata

has divaricate claws.

Orinoeme amboinica (Breuning, 1965) comb.

nov. (Figs 17-18)

Sybra amboinica Breuning, 1965b: 88

Type material examined: HT 3 ZSM: Amboina /

Felder // 38282 // Holotypus [red label] // Sybra (s. s.)

/ amboinica / mihi Typ [handwritten] / Breuning dét.;

preservation: toment a bit rubbed, left antenna with one

and right with nine segments only, right mesotarsus with

one segment only and without right metatarsus claw;

length: 14 mm.

Other studied material: 29 CWW: Indonesia:

Moluccas, See Buru Isl., 200-350m, Ilat vill. env.,

Remaja Mts., 5.-18.1.2013, leg. S. Jaki. length: 13-14.7

mm.

Notes: Due to morphological characteristics, long

antennae, shape of the prosternal and mesoventral

process, all femora covered with short, single,

adjacent white hairs, elytra mostly irregularly

punctured, this species belongs to the genus

Orinoeme Pascoe, 1867.

Genera Plocia Newman and Plociella Breuning

An original anotated checklist of species is

presented below.

351

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Figures 10-15. Mimectatina species, habitus and labels. 10-12 - Mimectatina celebica Breuning, 1975, HT 4

(RBINS); 10 - Habitus, dorsal view; 11 - ditto, lateral view; 12 - Original labels; 13-15 - Mimectatina celebensis

Breuning, 1975, HT 4 (RBINS); 13 - Habitus, dorsal view; 14 - ditto, lateral view; 15 - Original labels (photographs

courtesy Florence Trus, RBINS).

352

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WEIGEL, A. & SKALE, A.: On the taxonomy, synonymy and faunistics of the Apomecynini of the Asian-Australian region ...

Plocia Newman, 1842: 292

Type species: Plocia notata Thomson, 1864,

Subsequent designation by Thomson, 1864.

= Mimoplocia Breuning, 1949: 19

Tavakilian & Jiroux (2015): 78, synonymy).

Plocia notata Newman, 1842: 292 (Tavakilian &

Jiroux 2015: new combination).

Mimoplocia notata Breuning, 1949

= FEpaphra albicornis Heyrovsky, 1935: 19 (Breuning

1949: 19, synonymy).

Plocia diverseguttata Heller, 1924: 205 (Tavakilian

& Jiroux 2015: 78, new combination).

m. diversenotata Breuning, 1964: 302 nomen nudum

m. plurinotata Breuning, 1964: 302 nomen nudum

Plocia diverseguttata albopunctata Breuning,

1982b: 141 (Tavakilian & Jiroux 2015: 78,

new combination).

Plocia diverseguttata boholensis Breuning,

1982b: 141 (Tavakilian & Jiroux 2015: 78,

new combination).

Plocia diverseguttata mindanaonis Breuning,

1982b: 142 (Tavakilian & Jiroux 2015: 78,

new combination).

Plocia puncticollis (Breuning, 1965) comb. nov.

(Fig. 19)

Parepilysta puncticollis Breuning, 1965c: 182

Plocia splendens (Hudepohl, 1995) (Tavakilian &

Jiroux 2015: 78, new combination).

Mimoplocia splendens Hudepohl, 1995: 285

Plociella Breuning, 1949: 18

Type species: Plocia conspersa Aurivillius, 1927

by original designation.

= Sybroplocia Breuning, 1959: 77 syn. nov.

Plociella mixta (Newman, 1842) comb. nov. (Figs

20-22)

Plocia mixta Newman, 1842: 292

= Plociella conspersa (Aurivillius, 1927) syn. nov.

Plocia conspersa Aurivillius, 1927: 564

= Sybroplocia sybroides (Schwarzer, 1931) syn. nov.

Plocia sybroides Schwarzer, 1931: 73

Type material examined: Plocia mixta: colour

photograph of the HT in BMNH; Plocia conspersa: ST

3 NHRS: Island / Samar / Baker // Typus [red label]

// Plocia / conspersa / Auriv. [handwritten] // Plocia

/ conspersa / Auriv’ 27 / Auriv. [transversely written];

preservation: well preserved, right protarsus with one

pp. 345-362

segment only, left antennae with seven segments only;

length: 12-14 mm (Aurivillius 1927); Plocia sybroides:

HT SME; Philippines / Polillo // Typus [red label, black

bordered] // Plocia / sybroides / det. Schwarzer 1931

/ Typus !; preservation: complete preserved; length: 16

mm (Schwarzer 1931); Parepilysta puncticollis: HT °

ZMB: Luzon. / Jagor. [yellow label] // 39305 // ? Plocia

/ mixta / det. Schultze [handwritten] // Parepilysta / (s.

s.) / puncticollis mihi / Breuning dét. Typ; preservation:

tomentation rubbed, left antennae with four segments

only; length: 12.6 mm.

Other studied material (9 spms): Plocia mixta (=

Plociella): 1 spm CVB: Filipines-S, Mindanao, local

collector, [bottom side] Miganis Or., 1.2011; 19 CDB:

Philippines; N Luzon 2000m; Mountain Prov.; Bontoc

region, NW of Barlik; 17°03’N 121°04’E; L. Dembicky.;

O09.iv.2000; Collectio - L. Dembicky, Cerambycidae, Invt.

No.: 2982; 14 CDB: Philippines; Mindoro occid., Amnay

river valley; 25km SE of Santa Cruz; 12°57’N 120° 56’E;

L. Dembicky.; 17.iv.2000; 180m, Collectio - L. Dembicky,

Cerambycidae, Invt. No.: 2214; 19 CDB: Philippines;

Mindoro occid., Amnay river valley; 25km SE of Santa

Cruz; 12°57’N 120°56’E; L. Dembicky; 17.iv.2000;

180m, Collectio - L. Dembicky, Cerambycidae, Invt.

No.: 3010; 16° CWW: N-Philippinen, Laguna South,

V.20038, leg. |. Lumawig; 23 CSG: Philippines, Romblon,

Sibuyan, loc. collector, 1988; 14 CSG: Philippines,

Ambaguio, Nueva Vizcaya Prov., loc. coll., 11.2011; 12

CSG: Philippines: Luzon, Aurora Prov., 7.2015, loc. coll.;

length: 12-15.8 mm.

Notes: The examination of the HT of Plocia mixta,

one ST (NHRS) of Plocia conspersa and the HT

of Plocia sybroides, leads to the conclusion that

the three taxa are conspecific, so P. conspersa

and P. sybroides are junior synonyms of P. mixta.

Schwarzer (1931) himself wrote “P. sybroides

scheint mir nach der Beschreibung mit der mir

unbekannt gebliebenen P mixta Newm. am

nachsten verwandt zu sein ... auch mit conspersa

Auriv. 1927 ist diese Art durch die auf dem

Apikalteil angehauften weiS8lichen Flecke verwandt,

aber die Deckenspitzen sind nicht bedornt”. He is

already pointing out that this species is very near

to P. mixta. The last feature is wrong, the elytral

apex is extended to a spine at the outer edge by

the type specimen, and not unarmed as mentioned

in the description. The syntype of P conspersa

deposited in the USNM and shown in Lingafelter

et al. (2014) with the label ,Butuan / Mindanao /

Baker“ according to the description doesn’t belong

to the type series.

Tavakilian & Jiroux (2015) synonymized Mimoplocia

with Plocia, as Thomson (1864) defined the type

species Plocia notata for it, as this was not done

by Newman (1842). Plocia mixta Is very different

from P. notata and does not belong to this genus.

aes

S133)

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Because Plocia mixta and Plociella conspersa are

conspecific, Plociella is an available genus name

for this taxa. A characteristic feature of Plociella

mixta comb. nov. is the very broad mesoventral

process (Fig. 21), which is also referred to in the

descriptions by Aurivilllus (1927), Schwarzer

(1931) and Breuning (1949; 1959). Furthermore

the species is characterised by very close antennal

insertions and a strongly trapezoidal frons. It

remains unclear why two genera, Plociella and

Sybroplocia, were established by Breuning (1959)

for the same species, but this is not the only case

for this author.

A characteristic feature of the genus Plocia is the

conspicuous white tomentation of the 7th antennal

joint, which is not the case with Plociella and also

related genera Falsepilysta Breuning, 1939 and

Parepilysta Breuning, 1939.

The size and shape of the yellow or white macules

or lines show a large variation, and sometimes

are partly united, and therefore not useful for

differentiations.

The status of the three subspecific taxa to P.

diverseguttata cannot be clarified at present, further

studies are necessary. It is possible that this taxon

is very variable and widespread in the Philippines.

The two morphs plurinotata and diversenotata

described by Breuning (1964b), according to article

12 (ICZN 2000) are infrasubspecific, and therefore

not available. The corresponding specimens

are possibly within the variation spectrum of P.

diverseguttata.

Plocia puncticollis Breuning, 1965 comb. nov.

belongs to the genus Plocia, due to the general

habitus and especially the conspicuous white

tomentation at the 7* and basal half of the 8"

antennal segments.

Sybra biguttata Aurivillius, 1927: 573 (Fig. 23)

= Sybra negrosensis Breuning, 1947 (Weigel & Skale

2009: synonymy)

= Sybra flavoguttata medioalbomaculata Breuning,

1970 (Skale & Weigel 2012: synonymy)

= Sybra bisignatoides Breuning, 1980: 161 syn. nov.

= Sybra parabisignatoides Breuning, 1980: 162 syn.

nov.

Type material examined: Sybra biguttata: LT 3 USNM:

see also Weigel & Skale (2009, 2011); PLT 2 (NHRS):

see also Weigel & Skale (2011); Sybra bisignatoides: HT

MNHN: colour photograph of the HT (type picture also in

Chemin & Vives (2017)); Sybra parabisignatoides: PT ¢

ZMB: Insel / Mindanao [handwritten] // PARATYPE [red

label] // Sybra para- / bisignatoides / P.T. [handwritten]

/ Breuning dét. // Sybra para [handwritten] bisig- /

————

. |

354

natoides nov.; preservation: toment well preserved,

without left foreleg, right protarsus with one segment

only; right middle leg without tarsus, aedeagus is blown

out, no fibula could be found; length: 10.5 mm; PT CHS:

lle Mindanao / Philippinen [handwritten] /7/ PARATYPE

[red label] // Sybra para- / bisignatoides [handwritten]

/ Breuning dét. PT; preservation: toment somewhat

damaged, left fore- and middle tarsus missing, right

median tarsal claw and right hind tarsus missing, both

antennae with ten remaining segments; length: 12 mm.

Other studied material (5spms): 14 CTO: May 2013,

Philippines, Central Visayas, col. Ismael, Negros; 16' CTO:

Nov 2013, Philippines, Mt. Apo, Davao, SE Mindanao,

col. Ismael, Mindanao; 1 2 (CTO): JAN 2013; Philippines,

Cotabato SW, Mindanao, col Ismael, Mindanao; 1< CSG:

Philippines, |. 2014, San Fernando, Bukidnon; 14 CSG:

Philippines, Negros oriental, i. 2014.

Notes: According to HUdepohl (1983) the type

specimen of S. bisignatoides is heavily damaged,

another specimen is heavily rubbed off and also

damaged. The latter (deposited in MNHN) could

be examined according to a colour photograph

and is labeled as type (See also Chemin & Vives

2017). The species was described by Breuning

after a specimen from Mindanao (Philippines). We

consider this specimen to be conspecific with S.

biguttata. According to Breuning (1980) this species

should be similar to S. bisignata Schwarzer, 1931.

After examination of the male type specimen of S.

bisignata (SMF) this species rather does not belong

to the genus Sybra. The true genus assignment of

S. bisignata could not be clarified yet.

According to research by HUdepohl (1983) no type

specimen exists for S. parabisignatoides. However,

the above mentioned two PT could be investigated.

One PT (ZMB) is also mentioned in Breuning

(1980). S. parabisignatoides is also considered to

be conspecific with S. biguttata, therefore the two

species S. bisignatoides and S. parabisignatoides

are junior synonyms of S. biguttata.

Sybra conicollis Aurivillius, 1927: 567 (Fig. 24)

= Sybra densealbomarmorata Breuning, 1966: 238 syn.

nov.

Sybra obliquealbovittata Breuning, 1970: 465 syn.

nov.

Falsoropica albopunctata Breuning et Villiers, 1983

(Weigel & Skale 2011: synonymy)

Type material examined: Sybra_ conicollis HT

NHRS: Philippinen / n. Luzon / Mt. Polis // Typus [red

label] // 9947 / 22 [blue label]; preservation: well

preserved; length: 114 mm (Aurivillius 1927); Sybra

densealbomarmorata HT 92 (ZMB): s.a. Weigel & Skale

2011; Sybra obliquealbovittata HT MNHN: coulor

photograph of the HT.

————“a

02-Jun-21 21:50:47

WEIGEL, A. & SKALE, A.: On the taxonomy, synonymy and faunistics of the Apomecynini of the Asian-Australian region ...

pp. 345-362

Figures 16-25. Asian-Australian Apomycinini, habitus in dorsal and sternum in ventral view. 16 - Mimozotale trivittata

(Pic, 1931), 2 (CWW) from N Vietnam; 17 - Sybra amboinica Breuning, 1965, HT 3 (ZSM); 18 - ditto, 9 (CWW) from

Buru, Central Moluccas, Indonesia; 19 - Parepilysta puncticollis Breuning, 1965, HT 2 (ZMB); 20 - Plocia mixta

Newman, 1842, HT (BMNH); 21 - Plociella mixta (Newman, 1842), 4 (CWW) from Laguna Province, N Philippines,

meso- and metasternum, ventral view; 22 - ditto, habitus; 23 - Sybra parabisignatoides Breuning, 1980, PT (CHS)

from Mindanao, Philippines; 24 - Sybra conicollis Aurivillius, 1927, HT (NHRS); 25 - Sybra constricticollis Aurivillius,

1927 HT (NHRS) (photograph 23 courtesy Herbert Schmidt, Vienna, Austria) [not to scale].

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Other studied material: 22 CSG & SMF: Philippines:

Mt Polis, Luzon.

Notes: The examination of the HT of S. conicollis

(NHRS) has shown that the two species S.

densealbomarmorata and S. obliquealbovittata

are conspecific with S. conicollis and are therefore

Junior synonyms.

Sybra constricticollis Aurivillius, 1927: 569 (Fig.

25)

= Sybra postalbomaculata Breuning, 1964b: 308 syn.

nov.

Type material examined: S. constricticollis HT NHRS:

St. Thomas / Luzon // Typus [red label] // 9937 / 783

[blue label]; preservation: right profemur and -tarsus

missing, right hind tarus without claw; length: 10 mm,

3.5 mm wide (Aurivillius 1927); S. postaloomaculata HT

3 SNSD: Luzon, Pl. / Benguet Haights Place // Coll. W.

Schultze / Ankauf 1942 // Staatl. Museum fur Tier- /

kunde, Dresden // Typus // Sybra / postalbomaculata

mihi / Typ / Breuning det.; preservation: completely

preserved, only toment somewhat rubbed; length: 9

mm, 3 mm wide (Breuning 1964b).

Other studied material (2 spms): 14 SMF: St.

Thomas, Luzon, Coll. B. Schwarzer; 14 CDB: Philippines:

N Luzon; 2000m; Mountain Prov.; Bontoc region; NW of

Barlig; 17°03’N 121°04’E; L. Dembicky leg.; 9.iv.2000;

Collection - L. Dembicky, Cerambycidae, Invt. No.: 3718.

Note: The two HT specimens are morphologically

identical and their taxa therefore considered

conspecific, accordingly S. postalbomaculata is a

Junior synonym of S. constricticollis.

Sybra epilystoides Breuning et de Jong, 1941:

105 (Fig. 26)

= Parepilysta (Spinepilysta) ochreoguttata Breuning,

1961b: 246 syn. nov.

Type material examined: Sybra epilystoides: HT 9

RMHN: J.D. / Pasteur / Toegoe / Java / occ. // 220 //

Mus. Leiden [underlined] // type [red label] // Sybra

Pasc. epilyst- / toides mihi Pasc. / typ det. Breuning [all

handwritten]; preservation: left antenna with 4,5 and

right with nine segments only, at the legs missing: left

hind tarsus, right fore tarsus, middle tibia and tarsus;

length: 10.7 mm; Parepilysta ochreoguttata: coulor

photograph of the HT 9 FREY; preservation: left antenna

with one and right with 4,5 segments only, left hind

tarsus with one segment only; length: 10 mm, 3,5 mm

wide (Breuning 1961b).

Other studied material: 2 RMHN: Java occident.

/ Sukabumi / 2000’ 1893 / H. Fruhstorfer [black

bordered] // 221 // Mus. Leiden [underlined] // type

[red label] // Sybra Pasc. epilyst- / toides mihi Pasc. /

typ det. Breuning [all handwritten]; length: 10.7 mm.

356

Notes: Breuning & de Jong (1941) have described

the species based on one specimen (type locality:

Java: Toegoe), therefore the 2 mentioned above

and labeled with “type” (RMNH) does not belong to

the type material. Judging by a colour photograph

of the HT of Parepilysta ochreoguttata, this species

is conspecific with Sybra epilystoides and therefore

a junior synonym.

Sybra patruoides Breuning, 1939: 254 (Fig. 27)

= Sybra sumatrana Breuning, 1939: 268 syn. nov. (Fig.

27)

= Sybra sumbawana Breuning, 1959 (Weigel & Skale

2011: synonymy)

Type material examined: Sybra patruoides (HT

Weigel & Skale 2011; Sybra sumatrana HT 4 (NHRS):

Sumatra. // Staudinger. // Sybra sumatrana mihi Typ

[handwritten] det. Breuning // 3058 E 93 +. preservation:

well preserved, right antenna with nine segments only;

length: 10.1 mm.

Other studied material (2 spms, see also Weigel

& Skale 2011): 12 (NHNL): Bali - Drescher, 1919,

Djemb...ana [handwritten, in part unreadable], coll. F.C.

DRESCHER, RMNH ex. collection ZMAN; 14 (NHNL):

Nederl. Indis, A. Koller, 7, Banda [Banda Isl., Indonesia,

Moluccas], 1906, coll. F.C. DRESCHER, RMNH ex.

collection ZMAN.

Notes: The examination of the 3 HT of Sybra

Sumatrana showed that it is morphologically

identical to Sybra patruoides in all characteristics.

Sybra sumatrana is therefore a junior synonym

of Sybra patruoides. The species has been found

on the Indonesian islands of Sumatra, Java ?,

Bali, Lombok, Sumbawa, Sulawesi and Timor. The

species is very similar to Sybra destituta Pascoe,

1865, but in Sybra patruoides, the tomentum of

the elytra is whitish and covered with numerous

white macules (especially in the apical region),

whereas in Sybra destituta, the tomentum is always

yellowish and only a few macules are tomentated

whitish. Additionally between the upper eye lobes

in Sybra patruoides, the absence of punctures is

particularly remarkable.

Sybra peraffinis Breuning, 1942: 147 (Fig. 28)

= Sybra_ flavostictipennis Breuning, 1966: 239

syn.n.

Type material examined: Sybra peraffinis HT 4

NHRS: Butuan / Mindanao / Baker // Sybra / peraffinis

/ mihi Typ / det. Breuning // peraffinis Breun. // 9973

E92 +.; preservation: left middle and hind leg, and right

fore leg missing; length: 9.9 mm; Sybra flavostictipennis

02-Jun-21 21:50:51

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WEIGEL, A. & SKALE, A.: On the taxonomy, synonymy and faunistics of the Apomecynini of the Asian-Australian region ...

HT 4 ZMB: Philippinen / Leyte ... // Sybra flavo- /

Sstictipennis / mihi Typ / Breuning det. // Holotypus;

preservation: well preserved, only right antenna with

tree segments; length: 10 mm.

Notes: Sybra peraffinis is a species of the very

homogeneous Sybra_ incana-group, which is

represented by several species in the Philippines

(Skale & Weigel 2014). By examining the fibula, itcan

be clearly established that Sybra flavostictipennis

iS a Junior synonym of Sybra peraffinis. Besides

Sybra luzonica Breuning, 1939 and Sybra sibuyana

Aurivillius, 1927 (Weigel & Skale 2011), Sybra

peraffinis is currently the only Philippine species of

the incana-group without an apical elytral macule.

Sybra plagiata Aurivillius, 1927: 569 (Figs 29-30)

= Sybra plagiatoides Breuning, 1950: 268 syn. nov.

Sybra plagiata plagiatoides Breuning, 1964a: 230

Type material examined: Sybra plagiata HT ¢@

NHRS: Philippinen / N Luzon [N = handwritten] / Mt.

Data [handwritten] // Typus [red label, black bordered]

// 8838 [light blue label]; preservation: well preserved,

only the left middle claw is missing; length: 11 mm;

Sybra plagiatoides HT 9 NHRS: Mt. Polis / Luzon. //

Typus [red label, black bordered] // Sybra plagia- /

toides mihi / Breuning det. Typ [except ,,Breuning det.“

all handwritten]; preservation: well preserved, left

metatarsus missing, right antenna with six segments

only; length: 15 mm.

Other studied material (6 spms): 19 CWW: the

Philippines, N Luzon, CAR border Abra/Kalinga, E of

Melibeong, Basiwag, 17°30.191’N, 120°58.942’E,

1660m 3+4. Apr. 2008, JH Lourens, leg., Coll. ISRNB,

Achat J. A. Lourens, |: G.: 30.852; length: 13.8 mm; 1)

USNM: Filippines N, Apayao, local collector, IX-2011

[bottom side]; length: 10.4 mm; 1¢ CSG: Mt. Makiling,

Luzon, Baker, 4139, 197, Sybra sp., Auriv ’27; length:

9.4 mm; 14 SMF: Mt. Polis, Luzon, Coll. B., Schwarzer,

Sybra plagiata Aur., det. Schwarzer 1938; length: 8.9

mm; 19 ZSM: Philippines, Luzon Mountain Province,

VIII.87; length: 10.7mm; 12 CWW: Philippinen, Luzon,

Mt. Prov., April 1990; length: 15 mm.

Notes: The specimen described by Breuning as

S. plagiatoides is a large female of Sybra plagiata.

Both type specimens are morphologically identical,

in particular the differentiating characteristics

listed by Breuning (1950) are not applicable or

correspond only to minor individual variations. The

lower eye lobes are slightly larger in the female

specimens than in examined males. The light

tomentum at the base of the last 4-5 antennal

segments is not different in both type specimens,

as Breuning (1964a) notes. Both species are

conspecific, S. plagiatoides is therefore a junior

pp. 345-362

synonym of S. plagiata.

Sybra porcellus Pascoe, 1865: 211 (Figs 31-32)

= Sybra obliquevittata Breuning, 1939 (Weigel & Skale

2011, synonymy)

= Sybra pseudirrorata Breuning, 1939: 261 syn. nov.

(Fig. 31)

= Sybra laterivitta Breuning, 1940b: 428 syn. nov. (Fig.

32)

Sybra proximata Breuning, 1942 (Weigel & Skale 2017:

synonymy)

Sybra proximatoides Breuning, 1966 (Weigel & Skale

2011: synonymy)

Sybra submodesta Breuning, 1970 (Weigel & Skale

2011: synonymy)

= Sybra canoides Breuning, 1980: 161 syn. nov.

Type material examined: Sybra porcellus (Weigel &

Skale 2011); Sybra pseudirrorata HT 4 NHRS: Aroroy /

Philippin. // Sybra / pseudirrorata / mihi Typ [handwritten]

/ det. Breuning // 9948 E 92 +; preservation: well

preserved, only left antenna with ten and rigth with

eight segments; length: 8.5 mm; Sybra laterivitta HT 9

NHRS: Maratoea- / island // O. Borneo / Mjéberg //

Typus // Sybra / laterivitta / mihi Typ [handwritten] / det.

Breuning // 3028 E 93 +; preservation: right fore leg

missing, right antenna with tree segments only; length:

11.2 mm; Sybra canoides HT MNHN: coulor photograph

of the HT and labels (Chemin & Vives 2017) [labels

without location, according Breuning (1980) described

from Mindanao].

Other studied material (Weigel & Skale 2011; 2017):

1 som NHRS: same data as HT, but bears PT label.

Notes: Sybra porcellus is a widely distributed

and well-known species in Indonesia and the

Philippines. The examination of the holotypes of

Sybra pseudirrorata and S. laterivitta has shown

that they are conspecific with S. porcellus and

therefore both are junior synonyms. Judging by the

colour photograph of the poorly preserved HT of

S. canoides, this species is also conspecific with

S. porcellus, and therefore a junior synonym of S.

porcellus too. The aforementioned PT (NHRS) of S.

laterivitta does not belong to the the type material

according to Breuning (1940b).

Sybra propinqua Breuning, 1939: 265 (Figs 33-

34)

= Sybra sumatrensis Breuning, 1943: 45 syn. nov. (Fig.

34)

= Sybra variefasciata Breuning, 1973: 41 syn. nov.

Type material examined: Sybra propinqua HT 3

BMNH: Singapore. / H.N.Ridley. / 1900—242 // Sybra /

propinqua mihi Typ / det. Breuning // Type [white label,

red bordered]; preservation: toment well preserved, right

357

02-Jun-21 21:50:51

TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

32 33 34

Figures 26-34. Asian-Australian Apomycinini, habitus in dorsal view. 26 - Sybra epilystoides Breuning et de Jong,

1941, HT 2 (RMHN); 27 - Sybra sumatrana Breuning, 1939, HT 4 (NHRS); 28 - Sybra peraffinis Breuning, 1942,

HT 4 (NHRS); 29 - Sybra plagiata Aurivillius, 1927, HT 3 (NHRS); 30 - Sybra plagiatoides Breuning, 1950, HT

(NHRS); 33 - Sybra propinqua Breuning, 1939, HT 3 (BMNH); 34 - Sybra sumatrensis Breuning, 1943, HT 2

(NHRS) [not to scale].

358

Book4.indd 358 02-Jun-21 21:50:53

Book4.indd 359

WEIGEL, A. & SKALE, A.: On the taxonomy, synonymy and faunistics of the Apomecynini of the Asian-Australian region ...

antennae with one segment only; length: 7.7 mm; Sybra

sumatrensis HT 2° NHRS: Sumatra [handwritten] //

Typus [red label, black bordered] // Sybra / sumatrensis

mihi Typ / det. Breuning // 3056 E93 +; preservation:

toment well preserved, right middle leg missing; length:

8.6 mm; Sybra variefasciata HT 9 ZMB: S.E. Borneo //

TYPE // Holotypus // Sybra / variefasciata mihi Typ /

Breuning det. // Zool. Mus. Berlin; preservation: well

preserved; length: 9.3 mm.

Other studied material (26 spms): 19 CMI: Benakat,

S. Sumatra Indonesia, 9. vi. 1995, H. Makihara; 19

ZSM: Sumatra, Acjeh, III-89, Sybra sumatrensis Br.,

Hudepohl det. 1993; MALAYSIA-Borneo: 192 CCS:

Malaysia, Sabah, Crocker Range 12-III-2005 Cope

Collection; 1¢ & 12 CSG & CCS: Malaysia, Sabah,

Crocker Range, 12-IV-2005, Cope Collection; 14 CCS:

Indonesia, Sumatra Langkat, Pulau Buluh Rambung,

30-X-2002, Cope Coll.; 24’ & 19 CMI: Bunsit Park Nr.

Keningau, Sabah, 29.1I|.1992, Nobou Kanie leg.; 16

CMI: Bukit Soeharto Kalimantan Timur Indonesia, April

— Juli 1999, leg. |. Makihara; 14 CMI: Bukit Soeharto,

East Kalimantan, Borneo Indonesia, 26 - 28. x. 1996,

H. Makihara & H. Kinuura / Light Trap; 14’ CSG: 10 miles

point Keningau, Sabah, 21. Mar. 1989, A. Nishiyama

leg.; 19 JMU: Bergil, Mv SW3 N6 17.204 E 116 42.305,

V. pinnata, B8 F2, leg. A. Floren, 10.3.1997; 1 spm JMU:

TOPOU 12.3.97, leg. A. Floren; 1¢ JMU: Kinabalu Park,

Sorinsim SW II 15Years, Bergil, 10 2.3.97, leg. A. Floren;

14 CSG: Kinabalu Park, 6°5’N,116°33’E, Sorinsim III,

AOyr, Bergil 6, 7.3.97, leg. A. Floren; 14 CHH: Malaysia,

Sabah Crocker Range, vic. Trus Madi, III-20-2000, local

coll., DJHC Acc. # 98.5193; 14 CHH: Malaysia, Sabah

Ranau, IV-11-2008, local coll; 19 CHH: Malaysia,

Sabah, Mt. Trus-Madi, 700m, IV-6-2009, local coll.; 19

CBT: Malaysia, Pahang, Raub Alt. 250m, 30 Nov. 2008,

Shinichi Befu leg., Shinichi Befu Collection 05008; 19°

FREY: Singapor, Sybra propinqua mihi, det. Breuning,

Paratyp [red]; 23’ BMNH: Singapore., H.N.Ridley, 1903 -

353.; 19 USNM: Singapore Coll. Baker, 175, Sybra sp.,

Auriv 27; 14 CSG: Singapore., H.N.Ridley, 1903 - 237.;

19 CLS: SINGAPORE, dead tree, Nee Soon pipeline, CLF

& CYW, 2 Nov 2007, night.

Notes: Based on the examination of the three

HT specimens we consider S. sumatrensis and

S. variefasciata as conspecific with S. propinqua,

therefore both species are junior synonyms of

S. propinqua. Breuning (1939) described S.

propinqua based on one specimen from Singapore,

the mentioned 2 PT (FREY) therefore does not

belong to the type material.

Sybra umbratica Pascoe, 1865: 203

= Sybra petulans Pascoe, 1865 (Weigel & Skale 2009:

synonymy)

= Sybra albertisi Breuning, 1939 (Skale & Weigel 2012:

synonymy)

= Sybra lineatipennis Breuning, 1939 (Skale & Weigel

pp. 345-362

2012: synonymy)

= Sybra flavipennis Breuning, 1942 (Weigel & Skale

2009: synonymy)

Sybra umbratica m. flavescens Breuning, 1948: 15

nomen nudum

= Sybra bitriangularis Breuning, 1953 (Skale & Weigel

2012: synonymy)

Sybra umbratica m. alorensis Breuning, 1956: 687

nomen nudum

Type material examined: Sybra umbratica (Skale

& Weigel 2012); Sybra umbratica m. flavescens HT 2

NHRS: Key / Insel // Sybra / umbratica / m flavescens /

mihi Typ / det. Breuning.

Notes: As already pointed on by Skale & Weigel

(2012), the specimen designated Sybra umbratica

m. flavescens as type is conspecific with S.

umbratica. The taxon “flavescens” established

by Breuning (1948) is infra-subspecific. Later

Breuning (1960: 145; 1964a: 194) considered it

as subspecific. However, since he was not named

as the author of this taxon, the rank increase

according to article 10.2. or 45.5. ICZN (2000) is

invalid and the name is therefore not available. The

taxon “alorensis”, also listed in Breuning (1960:

145, 1964a: 195) as a subspecies of S. umbratica,

iS correspondingly infrasubspecific. This specimen

from the Indonesian island of Alor (MHNL) has not

yet been investigated. S. umbratica is a common

and widespread species, which varies more in size

and tomentation.

Xylariopsis mimica Bates, 1884: 247

= Sthenias (Albosthenias) leucothorax Breuning, 1938a:

370 syn. nov.

Type material examined: colour image of Xylariopsis

mimica HT in BMNH: Sapporo (Japan), lenght: 12 mm

(Bates 1884); Sthenias leucothorax: HT SNSD: Jiangsu

(China), length: 13 mm, 3.5 mm wide (Breuning 1938a).

Other studied material (3 spms): 14 CWW: Japan,

Fukairi, Togouchi, Hiroshima, 10/20 VI 2000, M.

Yasuda; 13' & 19 CWW: China-Shaanxi, Lueyang, 23.-

30.5.2009, leg. E. KuCera; length: 11.2 - 12.7 mm.

Note: According to the colour photographs of

the holotype specimens of these two species,

we consider X. mimica and S. leucothorax as

conspecific and therefore S. leucothorax is a junior

synonym of X. mimica. This is another example

that Breuning has described the same species in

different tribes.

New names given by Slipinski & Escalona

(2013) are not necessary because of incorrect

309.

02-Jun-21 21:50:53

Book4.indd 360

TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

synonymizations already correct in Weigel & Skale

(2014). The following names consequently are

placed in synonymy:

Sybra albovittata Breuning, 1943: 45

= Sybra gressitti Slipinski & Escalona, 2013: 231 [nom.

nov.] syn. nov.

Mimosybra continentalis Breuning, 1960: 140

= Sybra arnhemensis Slipinski & Escalona, 2013: 229

[nom. nov.] Syn. nov.

Mimosybra uniformis Breuning, 1940a: 166

= Sybra gaindahensis Slipifski & Escalona, 2013: 299

[nom. nov.] Syn. nov.

Acknowledgements

We sincerely thank the curators of the

various institutions and private individuals who

allowed us to examine type material and other

specimens: Maxwell Barclay, Michael Geiser and

Dmitry Telnov (all BMNH), Shinichi Befu (Kochi,

Japan), Pasquale Ciliberti (RMNH), Alain Drumont

(RBINS), Johannes Bergsten (NHRS), Olaf Jager

(SNSD), Andrea Hastenpflug-Vesmanis (SMF),

Lubos Dembicky (Brno, Czech Republic), Daniel

Heffern (Houston, Texas, U.S.A.), Azadeh Taghavian

(MNHN) and Yaheita Yokoi (Ratingen, Germany).

We are grateful for receiving type photographs

provided by Lubos Dembicky (Brno, Czech

Republic) (Sybra semilunaris), Herbert Schmidt

(Vienna, Austria) (Sybra parabisignatoides) and

Florence Trus (RBINS) (Mimectatina celebica

and M. celebensis). Finally, we sincerely thank D.

Heffern (Houston, Texas, U.S.A.) for English proof of

the manuscript.

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Received: 03.xii.2020.

Accepted: 07.1.2021.

02-Jun-21 21:50:54

Book4.indd 363

Yoxol, Y.: Taxonomic notes on the tribe Callidiopini Lacordaire, 1869 (Coleoptera: Cerambycidae) and some other ...

pp. 363-396

Taxonomic notes on the tribe Callidiopini Lacordaire,

1869 (Coleoptera: Cerambycidae) and some other

taxa of the Papuan Region, mainly from the Solomon

Islands, with descriptions of five new species

urn:lsid:zoobank.org:pub:89A3E229-4149-4C14-96 7/8-60D0/F3132C7

YAHEITA YOKOI

Beerenkothen 33, D-40882, Ratingen, Germany; y.yokoi@kddnet.de

Abstract: Under the auspices of the responsible ministry of the government of Solomon Islands, a research project

on the cerambycid fauna of the country was initiated in February 2019. Specimens of this family were collected

on Guadalcanal Island. As a first step, the taxonomy and biogeography of the Callidiopini was observed. Four new

species of the genus Ceresium Newman, 1842, i.e. C. gracilipenne sp. nov., C. pulekerai sp. nov., C. hurutaraui

sp. nov., and C. perplexum sp. nov. are described. For comparative observation, one new species of the same

genus, C. mirabile sp. nov., from New Guinea, is also described. Of the less-known genus Dendrides Dillon et Dillon,

1952, one new species, D. solomonicus sp. nov., is described. In addition, new distribution records of two known

species, C. diversum (Pascoe, 1866) and Oxymagis horni (Heller, 1912), are recorded. Their genitalia and terminalia

are described and illustrated in detail, in particular their endophalli with manifold ejaculatory duct complexes.

When a male specimen is not available, female genitalia is described. Substantial taxonomic and biogeographical

considerations beyond the description of the individual species are added in the discussion section.

Key words: Cerambycidae, Papuan Region, Ceresium, Oxymagis, Dendrides, genitalia, ejaculatory duct complex.

Introduction

In accordance with the agreement between

the Ministry of Environment, Climate Change,

Disaster Management & Meteorology (MECDM),

Government of Solomon Islands, and the author

of this publication, a research project was

Initiated in February 2019. The objective of the

project is to study the taxonomy, biogeography

and biology of cerambycid beetles in the Country

of Solomon Islands. A number of field studies

were already implemented within the framework.

Cerambycid beetles were collected in Barana area

on Guadalcanal Island by local partners as well

as by the author himself. The new materials were

subsequently classified and carefully examined. In

the course of the examination, interesting facts were

discovered and new observations recorded. These

discoveries and observations will be described ina

series of publications. As a first step, species of the

tribe Callidiopini Lacordaire, 1869 are observed

and described.

For taxonomic and biogeographical

consideration, comparison with the species of

adjacent regions is essential. In this case, the

species of the other parts of the Papuan Region in

sensu lato, as defined by Greke & Telnov (2014),

are most comparable. These subregions include

the Papuan Region in sensu stricto (commonly:

New Guinea), Bismarck Archipelago, Moluccas

and a number of smaller islands. The species from

these subregions are included in the scope. Further,

Vanuatu, Fiji and Micronesia are also taken into

consideration, as their species are often related,

though to a lesser extent.

The tribe Callidiopini in this extensive region,

as defined above, is fairly well explored. Early

discoveries were documented in the classic

literature by Fairmaire (1881; 1883), Schaufuss

(1864) and Pascoe (1869). The knowledge

was subsequently complemented by _ Heller

(1912; 1914), Aurivillius (1927) and Matsushita

(1935). Later, it was greatly supplemented by the

comprehensive publications by both Dillon & Dillon

(1952) and Gressitt (1951; 1956; 1959). In recent

years, Vives, Aberlenc & Sudre (2008) as well as

Waqa-Sakiti & Lingafelter (2009) and Waqa-Sakiti,

Winder & Lingafelter (2015) made substantial

contributions on this subject. Thus, 35 taxa of

the tribe are already recorded from New Guinea,

363

Tan

€

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nearly 22 from Micronesia and 21 from Fiji. For

Ceresium Newman, 1842, the most prolific genus

of the tribe, 12 taxa from New Guinea, 17 from Fiji

and 16 from Micronesia are already known. They

include endemic species and genera, unknown

in the Oriental Region, often with remarkable

characteristics.

In contrast, our knowledge of the tribe in the

Solomon Islands proper is rather limited. Even

though several works on Cerambycid beetles as

a whole were already published, the Callidiopini

was only sporadically referred to. Until recently,

only two widely distributed species, Tethionea

tridentata Pascoe, 1869 and Ceresium unicolor

(Fabricius, 1787) were recorded, both from

Malaita Island (Gressitt 1951). In recent years,

Vives et al. (2008) recorded new distributions of

Ceresium pachymerum (Pascoe, 1869), Examnes

philippensis (Newman, 1842) and Araespor

gazellus Gressitt, 1959 from Vanikoro, one of the

smaller islands remote from the main part of the

archipelago. Given the extensiveness of the region,

composed of many larger islands, and its location

adjacent to New Guinea, a more prolific fauna of

the Callidiopini is to be expected. Obviously, more

extensive exploration is necessary in the major part

of the island group. This research project should be

a contribution to compensate for such a deficiency.

The new material indeed includes several

interesting taxa. Many essential anatomical and

taxonomic aspects have thereby been revealed.

Four hitherto unknown Ceresium species have been

discovered. One new species of the rather enigmatic

genus Dendrides Dillon et Dillon, 1952 has also

been discovered. In addition, new distributions

of two species have been recorded. Of the new

species, C. gracilipenne sp. nov. probably belongs

to a coherent group of atypical, small-sized species

distributed in the Papuan and adjacent regions in

Melanesia. C. pulekerai sp. nov. and C. hurutaraul

sp. nov., especially the former, have genitalia

and terminalia unique in the genus and the tribe.

Likewise, both new species C. perplexum sp. nov.

and C. mirabile sp. nov., resemble each other, and

show peculiar characteristics hitherto unobserved

in the genus. The latter, though collected in New

Guinea, was also included in the observation.

Dendrides solomonicus sp. nov. in turn is the second

member of a poorly known genus whose tribal

position is yet to be clarified. Finally, C. diversum

(Pascoe, 1866) and Oxymagis horni (Heller, 1912),

both newly recorded from Guadalcanal, Solomon

Islands, have revealed remarkable morphologies.

In view of the significance of these species and

it i

364

their newly discovered characteristics, they are

described and illustrated in detail.

Regarding male genitalia and terminalia, their

importance for the taxonomy of the Callidiopini

was already noticed in previous publications

(Yokoi 2019; Yokoi et a/. 2019). In particular, the

endophallus provides an essential character for

classification. The male genitalia and terminalia

were therefore carefully examined. The results are

described and illustrated in particular detail below.

Female genitalia and terminalia are generally

less indicative. However, they often reveal interesting

observations for taxonomy (Yokoi 2019; Yokoi et

al. 2019). When a male specimen is not available,

female genitalia and terminalia are described.

In this publication, 4 new species are compared

in this regard. Differences in the structure of the

8" abdominal segment were observed, as well as

similarities in other regards.

Finally, new observations and questions

on higher taxonomy, beyond the description

of individual species, should be more broadly

discussed. These substantial aspects are added at

the end.

Material and methods

The material for this publication is composed

of specimens collected in Guadalcanal Island of the

Solomon Islands Nation. One specimen from New

Guinea was additionally included in the material.

These specimens in question were either directly

compared to the relevant holotypes, or to their

original descriptions and images.

All labels are cited as in the originals, and are

printed.

For the examination of genitalia, the specimens

were softened first, then submerged in KOH solution

for 12 hours and subsequently cleansed in water.

They were photographed using a digital camera with

a macro lens. Serial photos were stacked onto one

image. Drawings were made using Adobe Illustrator.

Median lobe, tegmen, 8" sternite and tergite were

examined for each male specimen. Endophallus is

observed in natura, i.e. in pre-copulative position in

abdomen. Ejaculatory duct of endophallus, together

with the surrounding manifold sclerites, referred

to as “ejaculatory duct complex”, was observed in

particular detail. In general, the direction, to which

the ejaculatory duct itself is pointed, is referred to

as “apical”, the side nearer to the duct likewise

“dorsal”. Additionally, when relevant, 9" sternites

or tergites were examined. For female, 8" and

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9'" sternites and tergites, soermathecae, median

oviducts and bursae copulatrix were examined. 8"

and 9° abdominal segments are often included

in “terminalia” in publications. However, they

sometimes play significant roles in copulation, as

indicated bytheir often elaborate structures of apical

parts. They are thus often referred to as “external

genitalia”. In this publication, 8" and 9" abdominal

segments are included in the descriptions of genital

organs. For the sake of simplicity and readability,

“terminalia” is nonetheless used as a collective

term. Usually, their external sides are observed and

described, i.e., sternites in ventral view and tergites

in dorsal view. The other sides are described when

relevant. In addition, 7*° abdominal segments are

simultaneously observed when sensible.

The term of the “flying hairs” is used to describe

widely spaced, free-standing, long, neither stout

nor erect nor adpressed dorsal setae, which often

are curved or sinuous, like thin hairs flying in wind.

They are typical for some genera (for instance,

Stenodryas Bates, 1873).

The holotypes and paratypes will be deposited

in the National Museum of Nature and Science

(NMNS), Tsukuba, Tokyo, Japan, where each

type specimen will be labelled with a registration

number. The numbers are given in the descriptions

of the respective species below. Two paratypes of

Ceresium gracilipenne sp. nov. will be temporarily

preserved in the collection of the author for further

comparative observation. The holotype of Ceresium

mirabile sp. nov., from Indonesian New Guinea, will

be preserved in the Indonesian Institute of Sciences

(LIPI), Bogor, Indonesia.

Measurements were taken with = an

ocular micrometer. The abbreviations for the

measurements in the descriptions are as follows:

EL - length of elytra;

EW - humeral width of elytra;

HW - head width across eyes;

M - arithmetic mean;

PA - apical width of pronotum;

PB - basal width of pronotum;

PL - length of pronotum;

PW - maximum width of pronotum;

TL - total length (from apical margin of clypeus to

abdominal apex).

Abbreviations of material stores:

BMNH - The Natural History Museum (formerly British

Museum, Natural History), London, United

Kingdom;

CYY - Collection Yaheita Yokoi, Ratingen, Germany;

pp. 363-396

LIPI - Indonesian Institute of Sciences, Bogor, Indonesia;

MECDM - Ministry of Environment, Climate Change,

Disaster Management, Meteorology, Government

of Solomon Islands, Honiara, Solomon Islands;

NMNS - National Museum of Nature and Science,

Tsukuba, Japan.

Results

Taxonomy

Ceresium gracilipenne sp. nov. (Figs 1-23 & 149

-151)

Holotype @ NSMT-I-C-200337: Solomon _Ils.,

Guadalcanal, Barana Village. 25-28.1I.2019. Y.Yokoi leg.

Paratypes 3 specimens: 19 NSMT-I-C-200338: same

data as holotype; 22 CYY: same data as holotype.

Remark: Antennae of the holo- and paratypes

partly damaged; length estimated.

Derivatio nominis: The name of this species

refers to its slender, evenly punctured elytra with

maculation.

Diagnosis: Small-sized; slender. Head and

pronotum finely reticulate. Pronotal sides evenly

rounded. Elytra evenly punctured; with dark

maculation.

Measurements for holotype male: TL=/.1

mm; EL/EW=3.4; HW/PW=1.04; PL/PW=1.2;

PA/PW=0.81; PB/PW=0.79. Paratype females:

TL=4.8-7.9 mm; EL/EW=3.3-3.4 (M 3.3); HW/

PW=0.96-1.08 (M 1.03); PL/PW=1.04-1.18 (M

1.10); PA/PW=0.84-0.88 (M 0.86); PB/PW=0.79-

0.83 (M 0.8).

Description: Colour yellowish dark brown.

Antennae, tarsi, tibiae, basal halves of femora paler.

Elytral disc pale, with a dark horizontal maculation

near apical 1/3. Setae yellowish to golden.

Head (Figs 2-4): Nearly as broad as pronotum;

universally reticulate; clothed with adpressed,

well aligned hairs. Frons short, reverse-trapezoid;

flattened; steep. Vertex nearly flattened. Eyes large,

separated from one another by 1/4 the width of

occiput or 5/6 of the upper eye-lobes. Antennal

supports flattened. Antennae surpassing elytral

apices by the last two articles in male and the last

one in female. Scape subarcuate and moderately

clavate; dorsum flattened and_ subreticulate.

Antennomeres 3, 4, 5, 6 each 5/4, 9/10, 7/5, 7/5

as long as scape; the remaining articles gradually

reducing in length. Pronotum (Figs 1-2): Longer

than broad; sides evenly rounded. Surface finely

reticulate, clothed with fine, adpressed hairs;

additionally with a few longer flying hairs. Disc well

eee

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rounded, with a diminutive nitid median stripe.

The stripe rudimentary or invisible in two female

paratypes. Scutellum bell-shaped; naked. Elytra:

Slender; subparallel-sided; regularly with dense

setiferous punctures; setae thereof fine, adpressed;

punctures shallower toward apices. Sides slightly

dilated toward apices in one female paratype.

Legs: Rather slender. Femora clavate; moderately

pedunculate in basal 1/4; flattened. Tibiae

Subarcuate. Venter (Fig. 5): Shiny. Prosternum very

convex; with rather sparse setiferous punctures;

setae thereof medium-length, curved, disorderly,

pointing in different directions. Prosternal process

compressed to a narrow strip between procoxae;

emarginate at apex. Mesosternum convex; with

regular large setiferous punctures; setae thereof

stout, adpressed. Mesosternal process broad and

flattened; densely, coarsely punctured; apex widely

emarginated. Metasternum weakly dilated toward

apex; nitid and shiny in middle; regularly and

rather densely punctured elsewhere; setae as of

mesosternum. Male abdomen: Much narrower

than metasternum or elytra; with sparse shallow

setiferous punctures; setae thereof short, curved;

additionally clothed with several longer, flying hairs.

Sternites 4-6 subequal in length; shorter than the

3. Sternite 7 subrectangular; clothed with several

stout hairs near apex. Apex of 7 tergite broadly

arcuate; hairs as on sternite. Female abdomen

(Fig. 8): Sides strongly emarginated. Sternites

subequal in length. 4‘° weakly, 5° moderately, 6"

and 7 strongly flattened. 3' subtrapezoidal; 4

subrectangular; 5 and 6 apically dilated; 7

broadest, subtrapezoidal with rounded sides. Male

genital organs (Figs 6-7 & 12-23): Median

lobe more than 2/5 the length of abdomen; bullet-

Shaped in dorsal view, weakly constricted near

apex; in profile slender and arcuate; dorsal plate

rounded at apex, dehiscent in basal 5/8; ventral

plate longer than the dorsal, pointed at apex,

dehiscent in basal 3/4. Crescent-like sclerites

large and stout. Tegmen 7/10 as long as median

lobe; in profile moderately arcuate. Parameres

about 1/2 the length of tegmen; slender; apical

2/5 bilobed; lobes each clothed with several short

apical setae. Ejaculatory duct complex large, nearly

2/5 as long as median lobe; with a prominent

median cavity in dorsal view; apical part weakly

sclerotized, dehiscent, hooked downward; basal

sclerite slender, elongated. Terminalia (Figs 7 &

21-23): Blade of 8" sternite transverse; narrowed

sublinear toward apex; apex emarginated in a bi-

Sinuate line, clothed with short, stout hairs; sides

fringed with longer hairs; peduncle much shorter

—_—_—

366

than blade. Tergite 8 narrower than the sternite;

more sharply narrowed toward apex; apex simply

emarginated; fringed with hairs of varying lengths.

Female genital organs: (Figs 9-11 & 149-

151). Sternite 8 with peduncle about 2/3 the length

of abdomen; blade elongated-trapezoidal; latero-

basal corners sharply produced; apex clothed with

several stout setae; peduncle nearly three times as

long as blade. Tergite 8 congruent with blade of the

sternite. Aodominal segment 9 moderately long;

coxite nearly as long as paraproct. Vaginal plates

Spatulate, arcuate, flattened. Bursa copulatrix

contiguous to vagina. Spermatheca kidney-shaped;

membrane near base thick, leathery.

Differential diagnosis: With its very small

and slender body, strongly compressed prosternal

process and with several long flying hairs on

pronotum and abdomen, this new species differs

from the type species, Ceresium raripilum Newman,

1842. In fact, it is distinct from all the other

members of Ceresium distributed in the Oriental

Region. It is more similar to a group of smaller

species from the Australian Region. These include

Ceresium gracillipes Fairmaire, 1881, C. decorum

Dillon et Dillon, 1952, and C. promissum Dillon

et Dillon, 1952, all three from Fiji, and Ceresium

huedepohli Yokoi, 2019 from the Philippines.

The new species can be distinguished, however,

above all by the reticulate surface of the head and

pronotum as well as by the distinctive maculation on

elytra. Its procoxae are subcontiguous, while those

of C. decorum, C. promissum, C. huedepohli are

variable, from adjacent to moderately separated.

As for the female abdomen, its strongly emarginated

sides with dilated 6-7" sternites are singular in the

genus. Regarding male genitalia, the new species

is rather ordinary. Ejaculatory duct complex of its

endophallus is similarly sclerotized as in species of

Ceresium, Examnes and Stenodryas (Yokoi 2019),

indicating its fundamental affiliation with them. The

cavity in the middle of the complex is peculiar but

already observed in Ceresium balkei Yokoi, 2019.

In contrast, its male terminalia is rather singular.

The apical part of 8" sternite is distinctly broader

than 8" tergite, with apical margin in an elaborate

bi-sinuate line. The female genitalia manifest few

deviations. Location of bursa copulatrix contiguous

to vagina, basal thickening of spermatheca and

shape of vaginal plates should be noted. So far as

the external characteristics are concerned, it is also

comparable to Examnes dimorpha Gressitt, 1951.

The further taxonomical aspect of this observation

is covered in “Discussion”.

Distribution: Guadalcanal, Solomon Islands.

02-Jun-21 21:50:54

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pp. 363-396

6 * a

Figures 1-7. Ceresium gracilipenne sp. nov., holotype 3. 1 - Habitus; 2 - Head and pronotum, lateral view, angled;

3 - Head, frontal view; 4 - Vertex, surface, enlarged; 5 - Pro- and mesosternum; 6 - Median lobe, lateral view, with

ejaculatory duct complex; 7 - Sternite 8 with tergite in the background, ventral view [not to scale].

367

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

8 10 (spermatheca)

11.

A-— vagina;

B — vaginal plates;

C — bursa copulatrix;

D — spermatheca;

E — median oviduct.

Figures 8-11. Ceresium gracilipenne sp. nov., paratype 9, habitus, ventral view (8) and genitalia (9-11) [not to

scale].

c <=

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pp. 363-396

Figures 12-23. Ceresium gracilipenne sp. nov., holotype 3, genitalia and terminalia. 12 - Median lobe with

endophallus, dorsal view; 13 - ditto, lateral; 14 - ditto, ventral; 15 - Tegmen, dorsal view; 16 - ditto, lateral; 17

- ditto, ventral; 18 - Ejaculatory duct complex; dorsal view; 19 - ditto, lateral; 20 - ditto, ventral; 21 - Sternite 8,

ventral view; 22 - ditto, with 8" tergite in the background; 23 - Tergite 8 with sternite 8 in the background, dorsal

view [scale bars for figures 18-20 - 0.2 mm, scale bars for figures 12-17 & 21-23 - 1 mm].

Ceresium pulekerai sp. nov. (Figs 24-51)

Holotype 4 NSMT-I-C-200339: Guadalcanal Is., Barana

Village. April 2019. Local collector leg.

Derivatio nominis: The name of this species is

dedicated to Mr. Veira Talitotu Pulekera of MECDM,

Solomon Islands.

Diagnosis: Medium-sized; stout. Extensively

black. Pronotum apically inflated; latero-apical

corners minutely punctured. Median lobe extremely

Slender; apex of ventral plate produced in

Sagittiform. Ejaculatory duct complex exceptionally

small.

Measurements: TL=16.5 mm; EL/EW=2.5;

HW/PW=0.72; PL/PW=0.85; PA/PW=0.64; PB/

PE. a> we

PW=0.84.

Description: Colour black; femora, venter and

elytra reddish dark brown. Setae whitish. Head:

Distinctly narrower than pronotum; glossy; sparsely

clothed with short, stout, adpressed hairs; with a

deep median furrow stretching from upper frons to

occiput. Frons not steep, transversely depressed

in middle, with several deep, coarse punctures;

sides emarginated, each with a deep lateral cavity.

Vertex moderately concave; with a row of setiferous

punctures on each side of median furrow. Occiput

with punctures of small to medium size. Eyes

separated from one another by 4/11 the width

of occiput or twice that of the upper eye-lobes.

Antennal supports flattened; each subencircled

aes

369

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by an arcuate row of diminutive setiferous

punctures. Antennae reaching elytral apices by 10%

antennomere. Antennomeres 3, 4, 5, 6 each 1/1,

5/6, 21/20, 21/20 as long as scape; the remaining

articles gradually reducing in length; the last a little

longer than the preceding. Scape subarcuate and

moderately clavate. Pronotum (Fig. 25): Shiny;

transverse; voluminous; strongly inflated in apical

half; strongly constricted just before apex; apex

thus narrower than base; sides unevenly rounded.

Disc convex, well-rounded; with a minute, nitid

median stripe; setiferous punctures dense; hairs

thereof irregular, short, stout, adpressed, pointing

in different directions. Punctures around the latero-

apical corners diminutive; dense; not setiferous

(Fig. 26). Scutellum bell-shaped; well bordered;

with a large puncture in the apical middle; hairs as

of pronotum. Elytra: Shiny; rather short; tapering

toward apices, which are separately rounded.

Punctures dense; setiferous; setae thereof stout,

adpressed, pointing toward apices. Legs: Short

and stout. Femora strongly clavate from base on.

Tibiae arcuate; apically flattened and expanded;

apices produced in spoon-shape, supplemented by

a minute dent (Figs 30-31). Venter (Figs 27-29):

Prosternum convex, well rounded, voluminous; with

a horizontal costa subparallel to apical margin;

densely and coarsely punctured; punctures

laterally reducing in size; hairs denser, shorter than

on pronotum. Prosternal process well bordered;

Shallowly punctured; apically rather steeply

declivous; the apical middle thinned; apex weakly

emarginated. Mesosternum saddle-shaped; steeply

elevated toward the process; the middle with large

punctures; sides clothed with short, adpressed

hairs. Mesosternal process broad; flattened; with

several large punctures; sparsely clothed with

short, curved hairs; apex deeply emarginated.

Metasternum well convex; the middle with large

Sparse punctures; sides subarcuate, clothed with

short, stout hairs. Abdomen: Shiny. Hairs similar to

those on metasternum; sparse in middle; dense on

sides. Sternites 3-6 gradually decreasing in length

and width. 7‘ sternite longer than the two preceding

sternites; apical margin emarginated in a shallow

though sharp bi-sinuate line. 7 tergite projected

forward, surpassing the apex of 8" sternite; apical

margin subspherically rounded, though sharply

emarginated at middle. Male genital organs

(Figs 32-36 & 38-47): Median lobe more than 5/8

the length of abdomen; elongated-bullet-shaped

in dorsal view; extremely slender, the width less

than 1/10 the length; in profile likewise slender,

strongly arcuate; dorsal plate dehiscent in basal

370

1/3; ventral plate distinctly longer than the dorsal,

dehiscent in basal 3/8, with apex strongly produced

in sagittiform (Figs 41-42). Tegmen nearly as long

as median lobe; in profile strongly curved in a bi-

sinuate line; closely embracing the entire apical half

of median lobe. Parameres about 3/5 the length

of tegmen; subparallel-sided; apical 1/3 bilobed;

lobes slender, apices each clothed with several

short setae. Ejaculatory duct complex exceptionally

small, only about 1/20 as long as median lobe;

apical sclerite projected in horn-shape,_ thick,

stout, curved upward; both lateral sclerites thick,

spatulate; basal sclerite simply triangular in dorsal

and ventral view. Membrane of endophallus thick,

widely scaled (Figs 35-36). Terminalia (Figs

37 & 48-51): Blade of 8 sternite subsquarish,

though sides arcuate; thinned in middle; latero-

basal corners produced; apex and sides fringed

with medium-length hairs; peduncle as long as

blade, bifurcated near base. 8" tergite larger than

blade of the sternite; bell-shaped; connected to the

sternite with an extensive membrane structure;

hairs similar to those of the sternite.

Differential diagnosis: The voluminous, apically

inflated pronotum of this new species is peculiar

among Ceresium species from the Australian

Region. It resembles rather those of C. vestigiale

Pascoe, 1866, C. brevipes Pic, 1943, C. grandicolle

Holzschuh, 2015 or C. ikuoyokoii Yokoi, 2019 from

the Oriental Region. Its surface and puncture

are, however, clearly distinct. The diminutive

puncture on the latero-apical corners has not been

recorded before. Further, the apical structure of

tibiae is rather remarkable in the genus. The male

terminalia of the new species, as described above,

is peculiar. Even more remarkable is its genitalia.

The median lobe is not only strongly arcuate but

also exceptionally slender. The sagittiform apical

projection of the ventral plate has not been observed

before in the genus. The elongated, subparallel-

sided parameres of the tegmen are also unusual

among the observed species. The mounting of

tegmen on the median lobe is remarkable. As for

the endophallus, its ejaculatory duct complex is of

a type commonly observed in the genus Ceresium.

It is, however, exceptionally small in relation to the

median lobe. Additionally, the external surface of

the endophallus is heavily scaled, which has not

been observed to this extent in the genus.

Distribution: Guadalcanal, Solomon Islands.

02-Jun-21 21:51:53

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pp. 363-396

Figures 24-31. Ceresium pulekerai sp. nov., holotype <. 24 - Habitus; 25 - Head and pronotum; 26 - Pronotum,

latero-apical corner; 27 - Venter; 28 - Pro-and mesosternum; 29 - Mesosternum, lateral view; 30 - Apical

projection of mid-tibia, frontal view; 31 - ditto, different angle [not to scale].

Ce 371

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TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

3/ 35

Figures 32-37. Ceresium pulekerai sp. nov., holotype 3, genitalia and terminalia. 32 - Median lobe with tegmen,

in natura, lateral view; 33 - ditto, opened, ventral view; 34 - Tegmen, ventral view; 35 - Endophallus; 36 - ditto,

surface, part; 37 - Sternite and tergite 8, lateral view [not to scale].

372 en

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Yoxol, Y.: Taxonomic notes on the tribe Callidiopini Lacordaire, 1869 (Coleoptera: Cerambycidae) and some other ...

pp. 363-396

VIII. ter. au ; ——__—i*Viii. ter.

VII. ter.

VIII, ste. ©

\e = fall

my =

Figures 38-51. Ceresium pulekerai sp. nov., holotype @, genitalia and terminalia. 38 — Median lobe with endophallus,

dorsal view; 39 - ditto, lateral; 40 - ditto, ventral; 41 - Apex of median lobe, dorsal view; 42 - ditto, lateral; 43 -

Tegmen, lateral view; 44 - ditto, dorsal; 45 - Ejaculatory duct complex, dorsal view; 46 - ditto, lateral; 47 - ditto,

ventral; 48 - Tergites 7-8, dorsal view; 49 - Sternites 6-8 and tergites 7-8, ventral view; 50 - Sternite and tergite

8, ventral view; 51 - ditto, latero-ventral, setae removed [scale bars for figures 45-47 - 0.1 mm, for figures 38-44

& 48-51 - 1 mm].

& 373

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TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Ceresium hurutaraui sp. nov. (Figs 52-65)

Holotype 4 NSMT-I-C-200340: Guadalcanal Is., Barana

Village. February-March 2020. Local collector leg.

Derivatio nominis: The name of this species

is dedicated to Mr. Joseph Hurutarau of MECDM,

Solomon Islands.

Diagnosis: Small-sized. Uniform brown in colour.

Setae on pronotum very stout, curved, pointing

in different directions. Elytra short. Punctures on

mesosternum exceptionally large. Outlines of 8"

sternite and tergite congruent. Ejaculatory duct

complex large relative to median lobe.

Measurements: TL=7.0 mm; EL/EW=2.5;

HW/PW=0.88; PL/PW=1.08; PA/PW=0.74; PB/

PW=0.84.

Description: Colour uniformly dark brown; elytra

a little paler. Setae whitish. Head: Narrower

than pronotum, glossy. Frons not steep; coarsely

punctured; sparsely clothed with scattered

Short, stout, adpressed hairs pointing in different

directions; sides deeply emarginated, strongly

declivous. Vertex widely, deeply concave, with

a deep median furrow; with a few large, deep,

setiferous punctures on each side of the furrow.

Occiput with several large punctures near vertex;

fringed with short hairs along the upper eye-lobes.

Eyes separated from one another by 1/4 the width

of occiput or 5/4 of the upper eye-lobes. Antennal

supports flattened; each subencircled by an arcuate

row of diminutive setiferous punctures. Antennae

surpassing elytral apices by 11 antennomere.

Scape elongated, clavate, arcuate; finely and

shallowly punctured. Antennomeres 3, 4, 5, 6, 7

each 7/8, 5/8, 1/1, 1/1, 7/8 as long as scape; the

remaining articles gradually reducing in length; the

last a little longer than the preceding. Pronotum:

Longer than broad; constricted just before apex;

apex narrower than base; sides evenly, weakly

rounded. Surface shiny; with uniform and rather

dense large, setiferous punctures; setae thereof

short, stout, curved, scattered, pointing in different

directions. Disc with an irregular, nitid median stripe

stretching from basal 1/5 to the middle. Scutellum

bell-shaped; thickly clothed with adpressed hairs.

Elytra: Short; tapering toward apices, which

are separately rounded. Shiny. Punctures large,

setiferous; setae thereof stout, curved, pointing

in different directions; apical punctures smaller;

apical hairs finer, more aligned. Legs: Short and

stout; femora clavate from base on. Venter:

Prosternum subparallel-sided; the middle with large

Sparse setiferous punctures; setae thereof stout,

curved, pointing to the process; sides with denser,

374

smaller punctures bearing shorter hairs. Prosternal

process narrow; well bordered; clothed with fine,

adpressed hairs; moderately declivous toward

apex; apex weakly emarginated, thinned in middle.

Mesosternum elevated; the middle with several

extraordinarily large setiferouSs punctures, setae

thereof stout, curved; sides thickly clothed with fine,

adpressed hairs. Mesosternal process broad; with

a few smaller punctures; sides clothed with short

adpressed hairs; apex thickly bordered, deeply

emarginated. Metasternum apically subdilated;

shiny; the basal middle with several large,

setiferous punctures each bearing a stout, curved

seta; punctures elsewhere sparse, smaller; sides

densely clothed with adpressed hairs. Aodomen:

Sides gradually tapering toward apices. Sternites

3-6 shiny; with small sparse setiferous punctures;

setae thereof short, stout, curved. Sternites 4, 5, 6,

7 each 1/2, 1/3, 1/3, 1/2 as long as the 3". The

7 trapezoidal; apex feebly emarginate, fringed with

Short hairs. Male genital organs (Figs 54-63):

Median lobe large in relation, nearly 3/5 as long as

abdomen; fusiform in dorsal view; in profile strongly

arcuate; dorsal plate dehiscent in basal 11/20;

ventral plate distinctly longer, with apex a little

produced to form a tooth; dehiscent in basal 3/4.

Tegmen 8/9 as long as median lobe; moderately

arcuate in profile. Parameres nearly 1/2 the length

of tegmen; weakly tapering toward apices; apical

3/10 bilobed. Lobes slender; apices with several

long hairs; sides and underside clothed with short

setae. Ejaculatory duct complex large, nearly 1/2

as long as median lobe; intricate in structure. Apical

sclerite elongated; subparallel-sided; bilobed.

Both lobes spatulate, apically rounded in spoon-

Shape, inter-connected to each other with a thick

membrane. Ejaculatory duct visible between the

lobes. Both lateral sclerites slender. Basal sclerite

linguate in ventral view; in profile thick and arcuate;

supplemented by a weakly sclerotized ventral

projection. Membrane of endophallus thickened

and scaly in apical 1/2; scales large in apical 1/5,

diminutive from apical 1/5 to 1/2. Terminalia

(Figs 64-65): Blade of 8 sternite subcircularly

rounded in the apical half, though truncated on

apex; the middle narrowly thinned; latero-basal

corners extended; apex and sides fringed with

setae of varying lengths; peduncle 7/10 as long as

blade. Tergite 8 congruent to blade of sternite in

apical 3/4; hairs similar to those of the sternite.

Differential diagnosis: This new species can be

distinguished from the others by the combination of

its universally brown coloration, small though stout

body, short elytra, coarsely punctured pronotum,

02-Jun-21 21:53:47

Yoxol, Y.: Taxonomic notes on the tribe Callidiopini Lacordaire, 1869 (Coleoptera: Cerambycidae) and some other ...

pp. 363-396

1 yv My

Be. ’

Figures 52-55. Ceresium hurutaraui sp. nov., holotype 3. 52 - Habitus, dorsal view; 53 - Venter; 54 - Median lobe,

dorsal view, with ejaculatory duct complex in ventral view; 55 - Endophallus with ejaculatory duct complex, dorsal

view [not to scale].

Gy ee 375

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TeLNnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

disorderly hairs on elytra and large punctures on

mesosternum. In the male genitalia, the ejaculatory

duct complex of the new species, in relation to the

median lobe, is one of the largest in the genus.

It is, for example, relatively 10 times longer than

that of the above described C. pulekerai sp. nov. Its

structure is alSo one of the most intricate among the

hitherto observed species, whereas the ejaculatory

duct itself is openly visible. In addition, the

membrane surface of its endophallus is thickened

and scaly as in the above described C. pulekerai

sp. nov., though to a lesser extent. Regarding male

terminalia, the congruent outlines of the 8" sternite

and tergite overlapping each other is noteworthy.

Distribution: Guadalcanal, Solomon Islands.

Figures 56-65. Ceresium hurutaraui sp. nov., holotype 4, genitalia and terminalia. 56 - Median lobe, dorsal view;

57 - ditto, lateral view; 58 - Tegmen, dorsal view; 59 - ditto, lateral view; 60 - ditto, ventral; 641 - Ejaculatory duct

complex, dorsal view; 62 - ditto, lateral view; 63 - ditto, ventral view; 64 - Sternite 8, ventral view; 65 - Tergite 8,

dorsal view [scale bars for figures 61-63 - 0.5 mm, for figures 56-60 & 64-65 - Imm].

376

02-Jun-21 21:53:50

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Yoxol, Y.: Taxonomic notes on the tribe Callidiopini Lacordaire, 1869 (Coleoptera: Cerambycidae) and some other ...

Ceresium perplexum sp. nov. (Figs 66-71, 146 &

152-153)

Holotype @ NSMT-I-C-200341: Solomon _ls.,

Guadalcanal, Barana Village. 25-28. Il. 2019. Y. Yokoi

leg.

Remark: Antennae of the holotype partly damaged;

length is estimated.

Derivatio nominis: The name of this species

refers to its peculiar appearance.

Diagnosis: Yellowish pale brown; parts of legs

and antennae pitchy black. Pronotum glossy;

subtrapezoidal; sides bi-sinuate. Legs slender.

Measurements: TL=12.4 mm; EL/EW=2.9; HW/

PW=0.9; PL/PW=1.16; PA/PW=0.81; PB/PW=1.0.

Description: Colour yellowish pale brown. Of the

antennal articles, apical 1/2 of the 3% pitchy black;

total of 4-6 pitchy black; apical 1/4 of 7“ blackish;

total of 8-9 blackish. Of the legs, apical 1/4 of

femora, basal 1/5 and apical 1/6 of tibiae pitchy

black; apices of 1%t and 2"? tarsal segments as well

as the whole 3" segment pitchy black. Setae pale

yellowish. Head: Narrower than pronotum; glossy;

sparsely, coarsely punctured. Frons not steep; the

middle transversely depressed; sides emarginated,

each with a deep lateral cavity. Vertex broad,

flattened; with a deep median furrow stretching

to frons. Eyes relatively small, separated from one

another by 2/5 the width of occiput or twice the

width of upper eye-lobes. Antennal supports broad,

markedly flattened; each bordered by an arcuate

row of deep, coarse punctures and, additionally, by

minute, curved hairs. Antennae surpassing elytral

apices by the last article. Antennomeres 3, 4, 5, 6

each 11/10, 4/5, 4/3, 4/3 as long as scape; 7-9

each a little shorter than 6". Scape subarcuate and

moderately clavate; with minute, shallow punctures.

Pronotum (Fig. 68): Elongated-subtrapezoidal

in dorsal view; narrowest at apex and broadest

at base. Sides feebly tapering in a bi-sinuate line

toward apices; constricted just before apices;

uneven with a few protuberances and depressions,

with several coarse punctures. Disc somewhat

elevated, rounded; glossy; with several very shallow

punctures. Scutellum subtriangular; well-bordered;

with several diminutive setiferous punctures.

Elytra: moderate in length; feebly tapering toward

apical 4/5, then arcuate; separately rounded at

apex; disc flattened; regularly punctured; sides and

apices clothed with diminutive, adpressed hairs.

Legs: Rather slender. Femora moderately clavate;

mid and hind femora pedunculate in basal 1/3;

fore femora in about 1/5. Venter (Figs 67, 69-

71): Glossy; more or less clothed with diminutive

pp. 363-396

hairs. Prosternum transverse and subrectangular

in ventral view; convex in the middle; the middle

with sparse setiferous punctures; setae thereof

longer. Prosternal process narrow, well bordered;

with several shallow punctures and diminutive

hairs; apically rather steeply declivous; apex

subtruncated. Mesosternumsaddle-shaped; steeply

elevated toward the process; steeply declivous

on sides; with sporadic large coarse punctures.

Mesosternal process broad, flattened; integument

as of prosternal process; apically bilobed; apex

contiguous at the same level to the basal projection

of metasternum (Fig. 71). Metasternum twice as

longas mesosternum; subparallel-sided; impressed

along median furrow; rugose near the basal

projection; sparsely punctured in middle. Abdomen:

Glossy, impunctate; rather sparsely clothed with

short hairs. Sternites gradually reducing in length

and width toward apex. Sternite 7 strongly narrowed

toward apex, which is widely rounded. Female

genital organs (Figs 146 & 152-153): Blade

of 8 sternite squarish; fringed with apical hairs;

peduncle about 5/2 as long as blade. Base of 8"

tergite widely emarginated. Abdominal segment 9

short; coxite as long as paraproct. Vaginal plates

near median oviduct arcuate; spatulate. Bursa

copulatrix large; oval to subspherical; connected

to the apical vagina by a thick tube. Spermatheca

kidney-shaped; membrane near base and apex

thick, leathery; connected to bursa copulatrix by a

long, winding duct; spermathecal gland visible as a

smaller sack, connected to spermatheca by a short

vessel.

Differential diagnosis: The new species is

peculiar, with its glossy, impunctate, subtrapezoidal

pronotum. The coloration of the legs and antennae

is also unusual for a Ceresium species. The pattern

of coloration is somewhat reminiscent of Ceresium

sibuyanum Yokoi, 2019. However, it obviously

differs in other essential aspects. These peculiar

characteristics distinguish the new species from

other species of Ceresium and related genera.

Distribution: Guadalcanal, Solomon Islands.

Ceresium mirabile sp. nov. (Figs 72-78, 147 &

154-155)

Holotype Q LIPI: Indonesia, W. Papua, Arfak Mts.,

1190 m alt., Duebei Env. 21. 1-8. Il. 2008. 20 km S. of

Warmere, Manokwari distr. Local collector leg.

Derivatio nominis: The name of this species

refers to its remarkable appearance.

Diagnosis: Resembles Ceresium perplexum sp.

nov. at a first glance. Pronotum punctate. Elytral

ort

02-Jun-21 21:53:50

TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

SS OSS

Figures 66-71. Ceresium perplexum sp. nov., holotype 2. 66 - Habitus, dorsal view; 67 - ditto, ventral view; 68 -

Head and pronotum; 69 - Pro- and mesosternum; 70 - Pro-, meso- and metasternum, lateral view; 71 - Mesosternal

process, apex [not to scale].

378 nme

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pp. 363-396

Figures 72-78. Ceresium mirabile sp. nov., holotype 2. 72 - Habitus, dorsal view; 73 - ditto, ventral view; 74 -

Head and pronotum; 75 - ditto, lateral view; 76 - Pro- and mesosternum; 77 - ditto, lateral view; 78 - Elytral apices

[not to scale].

Book4.indd 379 02-Jun-21 21:54:51

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

apices emarginated, bi-dentate. Legs more slender;

partly keeled.

Measurements: TL=13.9 mm; EL/EW=3.1; HW/

PW=0.9; PL/PW=1.16; PA/PW=0.77; PB/PW=0.97.

Description: Colour reddish brown; abdomen and

elytra apically paler. Head black. Apical half of the

5 antennomere and the subsequent articles black.

Apical 1/4—1/3 of femora and basal 1/6—1/5 of

tibiae pitchy black. Setae pale yellowish to golden.

Head: Analogous to that of C. perplexum sp. nov.

Frons narrower; more strongly emarginated on sides.

Vertex narrower, more concave; punctures more

regular. Upper eyes more adjacent to each other,

separated from one another by 1/4 the width of

occiput or 5/4 the width of upper eye-lobes. Occiput

with small to medium-sized punctures near upper

eye-lobes. Antennae more slender, surpassing

elytral apices by the last article. Antennomere three

as long as scape; antennomere 4 about 2/3 as long;

5-8 each 11/10 as long; the subsequent articles

gradually attenuating. Scape a little more slender.

Pronotum: Longer than broad; apex narrower

than base, broadest at middle. Sides uneven with a

few swellings and depressions; with irregular large

punctures; apical half weakly tapering toward apex.

Disc glossy; with a nitid, clavate median elevation

in basal 1/3; with rather irregular small setiferous

punctures around there; setae thereof diminutive.

Scutellum similar to that of C. perplexum sp. nov.

Elytra: A little longer than that of C. perplexum

sp. nov.; subparallel-sided in basal 3/4, then

tapering to apices. Apices each emarginate; bi-

dentate; outer tooth more prominent than the

inner. Punctures on disc smaller and more regular

than in C. perplexum sp. nov. Setae similar to those

of C. perplexum sp. nov. Legs: More slender and

longer than in C. perplexum sp. nov. Fore femora

only feebly clavate. Tibiae and peduncles of femora

keeled. Venter: Similar to that of C. perplexum sp.

nov. Prosternal surface substrigate. Mesosternum

broadly saddle-shaped; coarsely rugose in middle.

Abdomen similar to that of C. perplexum sp. nov.

Female genital organs: (Figs 147 & 154-155).

Similar to those of C. perplexum sp. nov. Blades of

8" sternite elongated-reversed-trapezoidal; apex

fringed with medium-length hairs; peduncle twice

as long as blades. Tergite 8 similarly formed as

blade of the sternite.

Differential diagnosis: This new species

from Papua New Guinea shares many essential

characteristics with the above described C.

perplexum sp. nov. from Solomon Is. However, it

can be easily distinguished, first by the apically

narrower pronotum with punctate surface; second

380

by the emarginate elytral apices and third by the

more slender legs. It additionally differs in such

details as keeled legs or different colouration of

antennae. In the female genitalia, no substantial

difference was observed. In contrast, it differs

obviously from C. perplexum sp. nov. in the female

terminalia, in particular the 8" sternite and tergite.

Distribution: Indonesia, West New Guinea,

Doberai Peninsula.

Ceresium diversum (Pascoe, 1869) (Figs 79-106)

Diatomocephala diversa Pascoe, 1869: 541, type

locality: “Menado”.

Ceresium diversum: Aurivillius (1912: 123)

Ceresium diversum: Yokoi (2015: 198)

Material studied: 2¢ & 39 CYY: Barana Village,

Guadalcanal, Solomon Is. 25-8. Il. 2019. Y.Yokoi leg.;

25 & 69 CYY: same locality but February-March 2020.

Local collector leg.; Holotype Ceresium diversum,

BMNH: Menado / Diatomocephala diversa Pasc. /

Pascoe Coll. 93-60 / Diatomocephala diversa, Type,

Pas. Holotype Ceresium pachymerum, BMNH: Bouru

/ Diatomocephala pachymera, Type, Pasc. / Ceresium

pachymerum Pas. / Type.

Additional description: Venter: Prosternum

convex, well rounded; with sparse, scattered short,

stout, curved hairs pointing in different directions;

surface strigulate near apex; the cotyledonary area

inthe middle coarsely punctured. Prosternal process

narrow; well-bordered; moderately declivous toward

apex; apex weakly arcuate, thinned in middle.

Mesosternum saddle-shaped; though the middle

somewhat impressed; steeply elevated toward

the process; impunctate. Mesosternal process

as In Figs 81-82; broad; clothed with curved,

medium-length hairs; intricate in structure; base

irregularly elevated; apical part widely bilobed;

lobes subrectangular; each lobe with a prominent

protuberance. Metasternum subparallel-sided; well

convex; shallowly punctured around the narrow

median stripe; clothed rather sparsely with fine,

whitish hairs elsewhere. Abdomen: Integument

as of metasternum. Sternites 3-6 gradually

decreasing in length and width. Apex of 7* sternite

thickly bordered; subcircularly emarginated. Apex

of 7" tergite subtriangularly produced; fringed with

medium to long hairs. Male genital organs (Figs

85-93 & 98-103): Median lobe nearly 1/2 the

length of abdomen; fusiform in dorsal view; strongly

arcuate in profile; dorsal plate dehiscent in basal

2/3; ventral plate longer than the dorsal, dehiscent

in basal 7/10, apically produced with a sharp tooth.

Tegmen 19/20 the length of median lobe; in profile

02-Jun-21 21:54:52

Yoxol, Y.: Taxonomic notes on the tribe Callidiopini Lacordaire, 1869 (Coleoptera: Cerambycidae) and some other ...

pp. 363-396

Figures 79-84. Morphology of Ceresium. 79-82 - C. diversum (Pascoe, 1869) 3 from Guadalcanal. 79 - Habitus,

dorsal view; 80 - ditto, ventral view; 81 - Pro- and mesosternal processes; 82 - Mesosternal process, lateral view;

83 - C. diversum, holotype from Sulawesi; 84 - C. pachymerum (Pascoe, 1869), holotype from Buru [not to scale].

& 381

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

PANU |

N AY

| NAA |

Figures 85-97. Ceresium diversum (Pascoe, 1869), 4 from Guadalcanal, genitalia and terminalia. 85 - Median lobe

with endophallus, dorsal view; 86 - ditto, lateral view; 87 - ditto, ventral view; 88 - Tegmen, dorsal view; 89 - ditto,

lateral view; 90 - ditto, ventral view; 91 - Ejaculatory duct complex, dorsal view; 92 - ditto, lateral view; 93 - ditto,

ventral view; 94 - Sternite and tergite 8, ventral view; 95 - Sternites 7-8 and tergite 8, ventral view; 96 - Tergite

8, dorsal view; 97 - Tergites 7-8, dorsal view [scale bars for figures 91-93 - 0.5 mm, for figures 85-90 & 94-97

- Imm].

382

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Yoxol, Y.: Taxonomic notes on the tribe Callidiopini Lacordaire, 1869 (Coleoptera: Cerambycidae) and some other ...

strongly curved near middle. Parameres about 1/2

the length of tegmen; apical 3/5 bilobed; lobes

slender and elongated, each clothed with long

hairs on apex and shorter ones on apical sides.

Ejaculatory duct complex about 1/6 as long as

median lobe. Apical part of apical sclerite dehiscent

in dorsal view; in profile strongly curved downward.

Both lateral sclerites short, wing-shaped. Basal

sclerite thick, stout; horn-shaped in lateral view,

bent upward. Membrane of endophallus scaly

in parts. Terminalia (Figs 94-97 & 104-106):

Blade of 8" sternite bilobed; lobes well rounded;

clothed with several short apical hairs; peduncle

pp. 363-396

distinctly shorter than blade. Tergite 8 as broad

as the sternite; elongated; parallel-sided; apical

margin truncated or weakly emarginated; apical

hairs denser, longer.

Differential diagnosis: Mesosternal process

of the described species is remarkably structured,

with prominent protuberances. This characteristic

was observed in varying degrees among the

examined specimens. In some examples it is

more moderate, in some only rudimentary or not

visible. It is thus not stable in the population. It is

nevertheless peculiar in the genus Ceresium. In the

male genitalia, the tegmen of the new species is

100

105

104

Figures 98-106. Ceresium diversum (Pascoe, 1869), 3 from Guadalcanal, genitalia and terminalia. 98 - Median

lobe with endophallus, dorsal view; 99 - ditto, lateral view; 100 - ditto, ventral view; 101 - Tegmen, dorsal view; 102

- ditto, lateral view; 103 - ditto, ventral view; 104 - Sternite and tergite 8, lateral view; 105 - ditto, latero-ventral

view; 106 - Sternites 7-8 and tergite 8, ventral view [not to scale].

383

02-Jun-21 21:55:37

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384

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

more strongly and abruptly curved in profile than

usual for the genus. In contrast, its ejaculatory

duct complex is of a type commonly observed in

Ceresium species. As to terminalia, apices of its

sternites and tergites 7"—8" are each characteristic

and remarkable. In all, Ceresium diversum shares

essential characteristics with other species of

Ceresium. At the same time, it reveals remarkable

peculiarities hitherto unobserved in the genus.

Distribution: East Java, Bali, Lombok, Sulawesi,

Guadalcanal (first record for the Solomon Islands).

Oxymagis horni (Heller, 1912) (Figs 107-134)

Macroceresium horni Heller, 1912

Oxymagis vitticollis (Fairmaire, 1883): Gressitt (1959:

uD Lie}

Oxymagis horni: Hawkeswood & Dauber (2001: 313)

Oxymagis horni: Hawkeswood & Dauber (2003: 416)

Material studied: 24 CYY: Barana_ Village,

Guadalcanal, Solomon Is. April 2019. Local collector leg.

Additional description: Venter (Figs 108-

113): Prosternum convex; well rounded; irregularly

clothed with short, stout, curved, adpressed hairs;

the extensive cotyledonary area in middle weakly

impressed and densely punctured; punctures near

the process larger. Prosternal process broad; thickly

bordered; surface glossy, uneven, with several large

punctures near base and smaller ones elsewhere;

sporadically clothed with short hairs; apical 1/3

abruptly, subvertically bent downward; apical

margin arcuate. Mesosternum § saddle-shaped;

steeply elevated toward the process; the middle

strigulate on surface. Mesosternal process broad;

flattened, feebly impressed in middle; glossy; with

several punctures and hairs; apex triangularly

emarginated. Mesocoxal cavities closed externally.

Metasternum subparallel-sided; glossy; with sparse

small setiferous punctures; setae thereof short,

Stout, adpressed. Abdomen clothed with short,

adpressed hairs; nitid in middle: sternites 3-6

gradually decreasing in length and width. 7 sternite

longer than the preceding, apically narrowed; apex

truncated. Male genital organs (Figs 115-

121 & 128-133): Median lobe 1/2 the length of

abdomen; elongated-bullet-shaped in dorsal view;

moderately arcuate in profile. Dorsal plate dehiscent

in basal 3/5. Ventral plate longer than the dorsal;

dehiscent in basal 3/4; apical part in ventral

view subcircularly rounded, with a sharp tooth at

the top; impressed twice, first direct at apex and

second a little further; in profile bi-sinuate. Tegmen

9/10 the length of median lobe; narrow; in profile

moderately arcuate. Parameres about 1/2 the

——_

. |

length of tegmen; sides slightly emarginated; apical

2/5 bilobed. Lobes each fringed with a row of long

hairs on sides; apex clothed with several long hairs;

underside with short, stout, curved hairs; dorsum

coarsely punctured. Basal part of endophallus

thick, leathery; in natura multiply folded in close

wrinkles (Figs 128 & 131). Ejaculatory duct complex

about 1/5 the length of median lobe. Apical sclerite

prominent; horn-shaped; supplemented by a large,

weakly sclerotized, pincer-like appendage. Both

lateral sclerites wing-shaped; directly contiguous

to basal sclerite. Basal sclerite thick, monolithic;

Supplemented by a _ subsclerotized membrane

structure. Terminalia (Figs 122-127 & 134):

Blade of 8 sternite transverse; sides and apex

widely rounded; apical half more or less with

setiferous punctures; setae thereof short and stout;

apex fringed with longer hairs; peduncle a little

longer than blade. Tergite 8 similarly broad as blade

of the sternite; longer; apical half well rounded;

apical margin obtusely emarginated, clothed by

hairs of medium length; dorsal side with setiferous

punctures bearing stout hairs; ventral side densely

granulate-punctate near apex. Sternite 9 (spiculum

gastrale) with a pair of prominent protrusions at

the bifurcation, each with several long, stout apical

hairs (Figs 123-127 & 134).

Differential diagnosis: The genitalia of this

species is remarkable. Intrinsic apical structure

of its median lobe, lateral setae on parameres

of tegmen, integument on 8" sternite and tergite

are peculiar in the genus. Above all, its 9% sternite

(Spiculum gastrale) is singular. It has a prominent

projection with setae at the bifurcation. Further, the

endophallus is noteworthy, as its basal part is thick,

leathery, and in natura tightly folded within median

lobe. In contrast, the ejaculatory duct complex of its

endophallus is of a commonly observed type. In all,

the genitalia of Oxymagus horni belongs essentially

to the same type as of the Ceresium and the related

genera in the Callidiopini, so far as ejaculatory

duct complex is concerned. In other aspects it is

obviously distinct.

Distribution: Papua New Guinea (East Sepik,

Northern Solomon Province), Guadalcanal (first

record from the Solomon Islands).

Remarks: Gressitt (1959) erroneously

synonymized this species with Oxymagis vitticollis

(Fairmaire, 1883), which was corrected by

Hawkeswood & Dauber (2001; 2003). It was

originally described from New Britain Island

of Bismarck Archipelago, which now politically

belongs to Papua New Guinea. In other previous

publications, distribution in “Solomon Islands” was

02-Jun-21 21:55:37

Yoxol, Y.: Taxonomic notes on the tribe Callidiopini Lacordaire, 1869 (Coleoptera: Cerambycidae) and some other ...

pp. 363-396

, aay we Ly

YQ Na oe,

{ ENG ES

111 412

Figures 107-114. Oxymagis horni (Heller, 1912), 4 from Guadalcanal. 107 - Habitus, dorsal view; 108 - Venter,

lateral view; 109 - Prosternum; 110 - Mesosternum; 111 - Prosternal process, ventral view; 112 - ditto, vertical

part; 113 - Mesosternal process; 114 - Elytral apices [not to scale].

. <=> oy eoe oe

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

referred to, without the precise location specified.

The distribution in Guadalcanal Is., Solomon Is.,

is hereby confirmed. Oxymagis horni is known as

a pest of cocoa trees in various subtropical and

tropical regions. Aharmful outbreak in the “Northern

Solomon Province in Papua New Guinea” was

already reported 1984 (E. Smith 1985). As cocoa

iS Cultivated on Guadalcanal Island, it cannot be

excluded that this species was artificially imported

there.

Dendrides solomonicus sp. nov. (Figs 135-142,

148 & 156-157)

Holotype 2 NSMIT-l -C-200342: Solomon _Is.,

Guadalcanal, Barana Village. 25-28. Il. 2019. Y.Yokoi.

leg.

Derivatio nominis: The name of this species

refers to its distribution.

Diagnosis: Medium-sized. Robust. Pronotum

distinctly transverse. Antennomeres 3—/ spinose.

Elytral apices broadly emarginated.

Figures 115-127. Oxymagis horni (Heller, 1912), 3 from Guadalcanal, genitalia and terminalia. 115 - Median lobe,

dorsal view; 116 - ditto, lateral view; 117 - Tegmen, lateral view; 118 - ditto, ventral view; 119 - Ejaculatory duct

complex, dorsal view; 120 - ditto, lateral view; 121 - ditto, ventral view; 122 - Sternite 8, ventral view; 123 - ditto,

with tergite 8 and sternite 9 (spiculum gastrale) in the background; 124 - Tergite 8, dorsal view; 125 - ditto, with

spiculum gastrale in the background; 126 - Spiculum gastrale, ventral view; 127 - ditto, apical part, enlarged [scale

bars: Imm].

386

02-Jun-21 21:55:40

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pp. 363-396

133

Figures 128-134. Oxymagis horni (Heller, 1912), 3 from Guadalcanal, genitalia and terminalia. 128 - Median lobe

with endophallus, ventral view; 129 - ditto, apex; 130 - ditto, lateral view; 131 - Endophallus in median lobe; 132

- Tegmen, dorsal view; 133 - ditto, apex; 134 - Sternite 9 (Spiculum gastrale), ventral view [not to scale].

387

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TeLNnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Measurements: TL=20.5 mm; EL/EW=2.1; HW/

PW=0.68; PL/PW=0.74; PA/PW=0.6; PB/PW=0.8.

Description: Colour dark brown to blackish.

Antennae, legs and abdomen reddish brown.

Setae yellowish. Head: Distinctly narrower than

pronotum; sparsely and shallowly punctured: rather

sparsely clothed with short, stout, curved hairs.

Frons transverse; subrectangular; moderately

declivous; transversely, arcuately impressed in

middle; with a pair of deep lateral cavities. Vertex

widely concave; with a rudimentary median furrow.

Eyes deeply emarginate; separated from one

another by 2/5 the width of occiput or 5/2 of the

upper eye-lobes. Antennal supports flattened.

Antennae surpassing elytral apices by 10" article.

Antennomeres 3, 4, 5, 6, 7 each 11/10, 5/6, 6/5,

6/5, 6/5 as long as scape; the remaining articles

gradually reducing in length; Antennomeres 3-6

distinctly spinose; 7 subspinose. Scape moderately

arcuate and clavate; clothed with short, fine,

adpressed hairs. Pronotum (Fig. 137): Distinctly

transverse; strongly constricted just before apex;

apex thus narrower than base. Surface uneven;

irregularly, deeply, coarsely punctured; densely

clothed with minute, adpressed hairs pointing in

different directions. Sides unevenly inflated with

irregular swellings and depressions; the largest

swelling just before the apical constriction. Disc

Subcircularly Surrounded by irregular elevations;

with a narrow, clavate median costa. Scutellum

short and broad, subtriangular; haired as pronotum.

Elytra: Broad; punctures shallow, decreasing in

size toward apices; rather densely clothed with

short, fine hairs. Sides subparallel from basal 1/5

to 1/2, then arcuate to apices. Apices broadly

emarginated; obtusely bi-dentate. Disc rounded;

though subcircularly elevated and flattened just

behind scutellum. Legs: Moderate in length;

hind femora reaching the apex of 5* sternite;

femora moderately clavate from base on. Venter:

(Figs 136 & 138-139). Prosternum convex, well

rounded, voluminous; coarsely punctured; thickly

clothed with short adpressed hairs. Prosternal

process broad; surface uneven; coarsely punctured;

apical 1/3 abruptly subvertically, bent downward;

apex weakly emarginated. Procoxal cavities widely

opened posteriorly. Mesosternum saddle-shaped;

steeply elevated toward the process; with several

large punctures. Mesosternal process broad;

with irregular setiferous punctures; apex broadly

emarginated; apical margin contiguous at the

same level to the basal projection of metasternum.

Mesocoxal cavities closed externally. Metasternum

distinctly transverse; punctate on each side of

———

ty y

388

the median stripe. Abdomen: Shiny; impunctate;

sternites 3-7 gradually decreasing in length and

width; 7th broadly emarginated on apex. Female

genital organs (Figs 148 & 156-157): Blade

of 8'" sternite subsquarish, though the basal half

narrowed posteriorly; widely thinned in middle;

apices fringed with setae of various lengths;

peduncle elongated, about 10/3 as long as blades.

Tergite 8 squarish; apical hairs similar to those of the

sternite. Aodominal segment 9 rather long; coxite

as long as paraproct. Vagina with a pair of arcuate

flagella near median oviduct. Bursa copulatrix

large; contiguous to the apical vagina; with a pair

of small sclerites near base. Spermatheca kidney-

shaped; membrane subsclerotized near base and

apex; connected to bursa copulatrix by a long,

winding tube.

Differential diagnosis: The spinose antennae

of the new species is reminiscent of the tribe

Phoracanthini. In other aspects, however, it does

not resemble any species of the tribe. It is rather

comparable to Dendrides pallidus Dillon et Dillon,

1952 (Figs 143-145). Both species share similar

structures of body, pronotum, legs and elytra,

differing more in details such as punctures and

setae. However, the new species differs obviously

by the spinose antennomeres 3-6. It is the second

member of this rather seldom observed genus. It

should be added that the venter of this new species

is Somewhat similar to that of Oxymagis horni

(Heller, 1912) as described above, particularly

regarding the subvertical apex of prosternal

process. For further taxonomic consideration, see

Discussion.

Distribution: Guadalcanal, Solomon Islands.

Discussion

Ceresium gracilipenne sp. nov. and related species

It is remarkable that Ceresium gracilipenne

Sp. nov. is similar to Examnes dimorpha Gressitt,

1951 from Vanuatu (formerly New Hebrides). In

fact, the body structure as a whole is comparable.

The new species can be distinguished from the

latter by the different coloration and reticulate

surface of the head and pronotum. It indeed shares

the steeply declivous frons with E. dimorpha and

other Examnes species. The question arises if it

should be also classified as Examnes. However, it

differs from Examnes species in several essential

aspects. Compared to Examnes_philippensis

(Newman, 1842), the type species of the genus

02-Jun-21 21:55:41

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pp. 363-396

Figures 135-145. Species of Dendrides. 135-142 - D. solomonicus sp. nov., holotype 9: 135 - Habitus, dorsal

view; 136 - ditto, ventral view; 137 - Pronotum; 138 - Pro-and mesosternal processes; 139 - ditto, lateral view;

140 - Antennomeres 3-7; 141 - Elytral apices; 142 - Third antennomere; 143-145 - D. pallidus Dillon et Dillon,

1952, holotype 3': 143 -Habitus, dorsal view; 144 - Elytral apices; 145 - Antennomere 3 [not to scale].

389

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Examnes, the new species is much smaller and

more slender, its antennae much shorter, its

antennal supports flattened, pronotal surface

reticulate instead of unevenly punctate, and finally,

prosternal process distinctly narrower. Thus, the

differences from Examnes species by far outweigh

the single characteristic of the frons that they have

in common.

At the same time, the slender body of the new

species and its long, very sparse, flying hairs on

pronotum and abdomen are rather reminiscent of

Stenodryas, another genus related to Ceresium.

The new species differs, however, in other essential

aspects such as structure of pronotum and legs.

Regarding genitalia, this new species is

basically similar to those of Ceresium or of related

genera Examnes and Stenodryas. |In contrast, it

has unique male terminalia and female abdomen.

In all, the new species is best placed in

the genus Ceresium, though differing in several

important aspects such as the steep frons, long

flying hairs, peculiar male terminalia and female

abdomen. It is described as Ceresium in this

publication.

The taxonomic inter-relationship of the three

genera Ceresium, Examnes and Stenodryas has

already been discussed in previous publications

(Yokoi 2019; Yokoi et al. 2019). Species embracing

characteristics of these different genera were

already described there. This new species is

another example of such a species.

External characteristics analogous to those of

the new species are also observed for C. gracilipes,

C. decorum, C. promissum and C. huedepohli. They

are small, brownish-yellowish in colour. Their bodies,

antennae and legs are slender. Their pronotal

surfaces are regularly and finely punctured, mostly

even, without lateral callosities. In this respect,

the above compared Examnes dimorpha is also

similar. These smaller species from the Australian

Region probably constitute a coherent group,

though not homogenous. In this connection, it is

noteworthy that these external characteristics are

also commonly shared by species of the subgenus

Ceresiellum, including Ceresium (Ceresiellum)

olidum (Fairmaire, 1881), C. (C.) usingeri Gressitt,

1956 and C. (C.) planatum Gressitt, 1956. Further,

some species from the genus Araespor Thomson,

1878 should also be considered. They also share

some of the above-mentioned characteristics.

However, the tribal position of Araespor must first

be clarified, as it was placed in the Achrysonini

by Aurivillius (1912) but Gressitt (1959) rightly

questioned this, and tentatively included it in the

390

Callidiopini.

The taxonomic positions of these genera and

Subgenera should be carefully considered further.

Genitalia and terminalia of the species concerned,

in particular those of the type specimens, should

be examined.

Remarkable genitalia of C. pulekerai sp. nov.

The genitalia of this new species is unique

within the genus Ceresium. First, the median lobe is

unusually arcuate and slender. It is the slenderest

among the hitherto observed species of the genus,

only the median lobe of Ceresium detanii Yokoi,

2015 is nearly comparable. Second, the peculiar

apical projection of the ventral plate has not been

observed before in Ceresium. Third, the parameres

of tegmen are unusually long and parallel-sided.

It is proportionally by far the longest in the genus.

Finally, the median lobe is much more extensively

and congruently embraced by tegmen than usual.

As to the endophallus of the new species, the

presence of a similarly sclerotized ejaculatory duct

complex indicates a fundamental affiliation with

Ceresium or with the related genera. The complex is,

however, exceptionally small. Usually, the length of

the complex relative to the median lobe varies from

1/4 to 1/3, reducing to 1/8 in an extreme case. It is

1/20 in case of the new species, an extraordinary

small figure. Furthermore, the extensively scaly

surface of endophallus is unique within the genus

Ceresium.

Thus, the male genitalia of the new species are

exceptional in the genus. At the same time, its male

terminalia, as described and illustrated above, is

also unique. Regarding the taxonomy of the new

species, the genitalia and terminalia are almost

distinct enough to justify the introduction of a new

Subgenus or a new genus. In contrast, the external

characteristics of the new species as a whole are

less remarkable. Though the inflated pronotum

with peculiar puncturation and the structure of

mid-tibiae are unusual, other characteristics are

rather commonly observed in Ceresium species.

The difference in external characteristics seems

thus less conclusive. More related species need to

be discovered and observed.

Ceresium hurutataui nov. and Ceresium

pulekeral sp. nov.

Sp.

Endophallus of C. hurutataui sp. nov. is also

scaly or thickened on the surface, though to a

lesser extent than that of C. pulekerai sp. nov.

02-Jun-21 21:56:39

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pp. 363-396

146

A, 9th sternite;

B, vagina;

C, vaginal plates;

D, bursa copulatrix;

E, spermatheca;

F, spermathecal gland;

G, median oviduct;

H, 8th sternite, ventral

view.

147

Far left: 9th sternite.

Clockwise from bottom:

vagina with vaginal plates;

bursa copulatrix;

@ spermatheca;

i Medianoviduct.

148

A, 9th sternite;

B, vagina;

C, vaginal plates;

D, bursa copulatrix;

E, spermatheca;

F, median oviduct.

Figures 146-148. Female copulatory apparatus of Callidiopini. 146 - Ceresium perplexum sp. nov., holotype 9:

A - Sternite 9; B - Vagina; C - Vaginal plates; D - Bursa copulatrix; E - Spermatheca; F - Spermathecal gland;

G - Median oviduct; H - Sternite 8, ventral view; 147 - Ceresium mirabile sp. nov., holotype 9: Far left, 9th

sternite; clockwise from bottom - Vagina with vaginal plates; Bursa copulatrix; soermatheca; Median oviduct; 148

- Dendrides solomonicus sp. nov., holotype 2: A - Sternite 9; B - Vagina; C - Vaginal plates; D - Bursa copulatrix;

E - spermatheca; F - median oviduct [not to scale].

G x

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157

Figures 149-157. Female copulatory apparatus and terminalia of Callidiopini. 149-151 - Ceresium gracilipenne

sp. nov., paratype 2: 149 - Sternite 8, ventral view; 150 - Tergite 8, dorsal view; 151 - Vaginal plate; 152-153 -

Ceresium perplexum sp. nov., paratype 2: 152 - Sternite 8, ventral view; 153 - Tergite 8, dorsal view; 154-155

- Ceresium mirabile sp. nov., paratype 9: 154 - Sternite 8, ventral view; 155 - Tergite 8, dorsal view; 156-157 -

Dendrides solomonicus sp. nov., paratype 9: 156 - Sternite 8, ventral view; 157 - Tergite 8, dorsal view [scale bars

for figures 148-149 & 151-154 - 1 mm, for figure 151 - 0.4 mm].

392

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above. It is noteworthy that the endophalli of these

two sympatric species are similar in this regard.

Their ejaculatory duct complexes are also similar

in design. However, the complex of the latter is

peculiarly small, while that of the former is one of

the largest in the genus. The male terminalia, 8"

sternite and tergite of the two species, differ in

structure, but both are unique within the genus.

Regarding genitalia and terminalia as a whole, both

species differ obviously from the Oriental species of

Ceresium or related genera.

Taxonomical questions regarding Ceresium

perplexum sp. nov. and Ceresium mirabile sp. nov.

These two new species, from Solomon Is. and

from New Guinea respectively, differ in several

significant aspects. They are, however, essentially

similar in body structure. The coloration patterns

are analogous, differing in details. They are also

similar regarding female genitalia, while differing

in terminalia. Both of them reveal, in turn, a few

external characteristics which are not typical of

Ceresium but rather reminiscent of neighbouring

genera. The nitid, impunctate, subtrapezoidal

pronotum of Ceresium perplexum sp. nov. is more

similar to those of Stenodryas species, while that of

Ceresium mirabile sp. nov. is more similar to those

of typical Ceresium species. In contrast, the legs

of the former are unremarkable, while those of the

latter are not typical of Ceresium. The slender, only

feebly clavate fore-femora of the latter are rather

reminiscent of species of Salpinia Pascoe, 1869.

Finally, the emarginated elytral apices of the latter

are seldom observed in Ceresium or related genera.

Only the sympatric Ceresium emarginatum Yokoi,

Makihara et Woro, 2019 shares this characteristic.

Both new species are again examples of taxa

embracing characteristics of different genera. For

further consideration, more comparisons of their

male genital organs are necessary. The two new

species are described here in the genus Ceresium.

Ceresium diversum / Ceresium pachymerum

There seems to be an ambiguity as to the

distinction between Ceresium diversum (Pascoe,

1869), originally described from Sulawesi, and

Ceresium pachymerum (Pascoe, 1869) from Buru

and Seram. According to the original descriptions,

the differences essentially concern coloration and

the form of the pronotum, elytra and legs. However,

these differences are mostly of gradual nature.

Comparison of the holotypes, both female, does

pp. 363-396

not show any clear distinction. Although some

differences can be recognized, such deviations

can also be observed in different geographical

populations of each species. Even within a sympatric

population, similar deviations can be commonly

observed. The changeable nature of their external

characteristics is, for example, underlined by the

observation regarding the peculiar mesosternal

process, described above for Ceresium diversum.

According to the records, C. pachymerum is

widely distributed in the Papuan Region, while C.

diversum is recorded from more western regions

(Yokoi 2015). In this sense, the record of C.

diversum from Solomon Islands is rather isolated.

However, the population from there, as described

above, resembles as a whole rather the holotype

of C. diversum than of C. pachymerum. It is thus

identified as C. diversum in this publication.

The relationship of these two species should

be treated separately in another publication, as it

will involve examination of male genitalia of more

specimens from various regions, and is beyond the

scope of this publication.

Taxonomic position of genus Oxymagis

The endophallus of this species has a strongly

sclerotized ejaculatory duct complex, like that of

Ceresium, Examnes or Stenodryas species. In fact,

Oxymagis is the fourth genus in the Callidiopini

to be observed with this type of endophallus.

In comparison, such a complex is absent in the

nominotypical genus Callidiopsis or other genera of

the Callidiopini such as Salpinia. This observation

indicates that Oxymagis should be affiliated with

the three genera above, Ceresium, Examnes or

Stenodryas of the Callidiopini.

In contrast, other parts of the genitalia differ

obviously from those of the three genera above. Its

9" sternite (spicum gastrale), which has a peculiar

prominent projection, is above all remarkable.

Tribal position of the genus Dendrides

In a formalistic point of view, Dendrides

solomonicus sp. nov. should be classified as a

species of the tribe Phoracanthini, on account

of its spinose antennomeres. However, it does

not resemble any of the species of the tribe in

other essential aspects. In particular, its strongly

transverse pronotum, in combination with broadly

emarginated elytral apices, has no equivalence in

the Phoracanthini.

Instead, the new species shares essential

393

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characteristics with Dendrides pallidus Dillon et

Dillon, 1952 (Figs 143-145). It is noteworthy,

however, that the antennae of D. pallidus are

not so spinose as of the new species, but its 3"

antennomere nevertheless has a tooth or tubercle

(Fig. 145). The essential difference between these

two species is thus relative, which in turn Supports

that the genus Dendrides could indeed include

Species with spinose antennomeres.

The tribal position of the genus Dendrides is

not yet determined. Dillon & Dillon (1952), who

described the genus, noted themselves that it

was impossible to place this genus in a tribe. For

the final classification, male genital organs of the

relevant species must be compared. At present,

the similarity of the pro- and mesosternum, in

particular the processes, with Oxymagis horni

should be noted.

The new species is described here in the genus

Dendrides, temporarily without tribal classification.

Male genitalia and terminalia

It was again observed that the endophallus

of Ceresium species have an intricate, heavily

sclerotized ejaculatory duct complex at the apex,

which is characteristic for the species (Yokoi 2019;

Yokoi et al. 2019). In addition, tt was shown that

a species of the genus Oxymagis has a similar

type of endophallus. Thus, Ceresium, Examnes,

Stenodryas and Oxymagis of the tribe Callidiopini

are similar regarding this important part of genitalia.

It is noteworthy that the endophalli of C.

pulekerai sp. nov. and C. hurutaraui sp. nov., both

from Guadalcanal, are very scaly. It has not been

observed for the Oriental species of the genus

before. A scaly or thickened endophallus is more

commonly observed in Callidiopini in other genera

such as Callidiopsis Thomson, 1864 or Salpinia

Pascoe, 1869.

The terminalia of the genera examined are

diverse and characteristic, as observed in previous

publications (Yokoi 2019; Yokoi et al. 2019). In

addition, it was observed that the apices of the

7" abdominal segments, both sternite and tegrite,

sometimes exhibit interesting structures, as in the

case of C. pulekerai sp. nov. or C. diversum.

The 9" sternite (Spiculum gastrale) of Oxymagis

horni is unique in structure. The prominent

protrusion with stout setae has not been observed

before in the tribe. The function of this peculiar

structure and its hairs is unknown.

Remark: Endophalli of Callidiopsis and Salpinia

are described in a parallel publication (Yokoi, in

it i

394

press).

Female genitalia and terminalia

Female genitalia of the four species examined

are in general not as characteristic as the male

genitalia. Vaginae, bursae copulatrix and median

oviducts are composed of membranes, susceptible

to deformation. Substantial characteristics which

are stable enough were observed solely for

the spermathecae and vaginal plates. Their 9"

abdominal segments are admittedly more stable.

They are, however, often similar, differing mainly in

their relative length. In contrast, the 8" abdominal

segments of these four species show substantial

differences in structure. For example, C. perplexum

sp. nov. and C. mirabile sp. nov., while resembling

each other in general, differ obviously in the

structure of 8" sternite and tergite.

Concluding remark

The above described species have revealed

interesting morphologies of genitalia as well as

of external characteristics, many of which are

observed for the first time in the Callidiopini. Several

new questions must thus be considered. Five new

species of the genus Ceresium, i.e. C. gracilipenne

sp. nov., C. pulekerai sp. nov., C. hurutaraui sp.

nov., C. perplexum sp. nov. and C. mirabile sp.

nov. differ substantially from most species of the

Oriental Region. Their taxonomic positions within

the genus or the tribe must be further examined. In

addition, the tribal position of the genus Dendrides

must be determined. Even the two known species,

C. diversum and Oxymagis horni, have peculiar

characteristics that merit more detailed taxonomic

consideration in future.

Further observations need to include more

comparisons of male genitalia, which could provide

an important character system for classification.

Previous examinations of genitalia and terminalia,

including those of relevant type specimens, have not

been sufficient. Regrettably, many male specimens

of pivotal species are still not available to the

author. In this sense, contributions by colleagues

on genitalia, the male endophallus in particular,

are welcome.

Acknowledgements

| am deeply indebted to Mr. Pulekera, Mr.

Hurutarau and their colleagues of MECDM, the

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Yoxol, Y.: Taxonomic notes on the tribe Callidiopini Lacordaire, 1869 (Coleoptera: Cerambycidae) and some other ...

government of the Solomon Islands (Honiara),

for enabling and supporting the research project.

Thanks to their assistance and collaboration, the

material basis for this publication could be created.

| deeply appreciate my local partners, including Mr.

Peter Besi of Guadalcanal Island, fortheirassistance

during the field research work in their country. |am

grateful to Dr. Tatsuya Niisato (Tokyo, Japan) for

generously allowing me to observe an important

specimen from his collection, to be described as

Ceresium mirabile sp. nov. | am thankful to Prof.

em. Nobuo Ohbayashi (Kanagawa, Japan) for

providing me with many important specimens and

photographs for comparative observations. For the

photograph of holotype of Dendrides pallidus, | also

thank Dr. Junsuke Yamasako (Tsukuba, Japan).

lam grateful to Mr. Maxwell V. L. Barclay (BMNH)

for allowing me to examine relevant type materials.

His proposals and recommendations have also

substantially improved the content and wording

of the manuscript. | would like to thank Dr. Dmitry

Telnov (BMNH), executive editor of this volume,

and his colleagues for their valuable assistance in

the editorial aspect. Further, | would like to thank

Mr. Theodore L. Childers (San Diego, U.S.A.) for

his advice regarding the wording of the text. Last

but not least, | thank my wife, Karin de Wit-Yokoi

(Ratingen, Germany), for proof-reading and other

editorial assistance.

References

Aurivillius C. 1912. Coleoptorum Catalogus. Pars 39:

Cerambycidae: Cerambycinae. W. Junk, Berlin: 574

Pp.

Aurivillius C. 1927. Neue oder wenig bekannte Coleoptera

Longcornia. 22 - Arkiv for Zoologi 19A, No 17:

525-547, 1 pl.

Dillon L., Dillon C. 1952. Cerambycidae of the Fiji Islands.

- Bernice P. Bishop Museum, Bulletin 206: 1-114.

Fabricius J. C. 1787. Mantissa insectorum sistens eorum

species nuper detectas Adiectis Characteribus

genericis, differentiis specificis, emendationibus,

Observationibus. Hafniae, Proft 1: xx + 348 pp.

Fairmaire L. M. H. 1881. Essaisur les Coléopterés des iles

Viti Suite. - Annales de la Sociéte entomologique

de France 6, No 1: 461-492.

Fairmaire L. M. H. 1883. Essai sur les Coléopterés

d’Archipel de la Nouvelle-Bretagne. - Annales de

la Sociéte entomologique de Belgique 27, No 2:

1-58.

Greke K., Telnov D. 2014. Review and assessment of

the literature on recent non-marine molluscs of the

Papuan biogeographical region. In: Telnov D. (ed.)

Biodiversity, Biography and Nature Conservation

pp. 363-396

in Wallacea and New Guinea. Volume Il. The

Entomological Society of Latvia, Riga: 498 pp, 126

pls.

Gressitt J. L. 1951. Longicorn Beetles from New Guinea

and the South Pacific (Coleoptera: Cerambycidae).

Part Il. - Annals of the Entomological Society of

America 44, No 1: 1-30, 1 pl.

Gressitt J. L. 1956. Coleoptera: Cerambycidae. - Insect

of Micronesia 17, No 2: 61-183.

Gressitt J. L. 1959. Longicorn Beetles from New Guinea |

(Cerambycidae). — Pacific Insects 1, No 1: 59-171.

Hawkeswood T., Dauber J. 2001. Biological notes and

host plants of some Papua New Guinean longicorn

beetles (Insecta: Coleoptera: Cerambycidae). -

Entomologische Zeitschrift 111, No 9: 312-315.

Hawkeswood T., Dauber J. 2003. Biological notes and

host plants of some Papua New Guinean longicorn

beetles (Insecta: Coleoptera: Cerambycidae). -

Mauritiana 18, No 3: 415-420.

Heller K. M. 1912. Neuheiten aus Herrn R.v. Benningsen’s

Sammlung von Coleopetreren aus den deutschen

Kolonien. - Entomologische Mitteilungen 1, No 6:

170-176.

Heller K. M. 1914. Coleoptera. Résultat de |l’Expédition

néerlandaise a la Nouvelle Guinée en 1907 et

1909 sous les auspices du Dr. H. A. Lorentz. - Nova

Guinea 9, No 2: 615-666, pl. XVI.

Holzschuh C. 2015. Beschreibung von 50 neuen

Bockkafern aus Sudostasien (Coleoptera,

Cerambycidae). - Les Cahiers Magellanes, nouvelle

série 20: 14-33.

Matsushita M. 1935. Beitrag zur Cerambyciden-Fauna

von Micronesien. - Transactions of the Sapporo

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Newman E. 1842. Cerambycitum Insularum Manillarum

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The Entomologist 15: 243-248.

Pascoe F. P. 1866. Catalogue of Longicorn Coleoptera

collected in the island of Penang by James Lamb,

Esq. Part Il. - The Proceedings of the Scientific

Meetings of the Zoological Society of London for

the year 1866: 504-536, pls XLI-XLIII.

Pascoe F. P. 1869. Longicornia Malayana; or a descriptive

catalogue of the species of the three longicorn

families Lamiidae, Cerambycidae and Prionidae,

collected by Mr. A. R. Wallace in the Malay

Archipelago. - Transaction of the Entomological

Society, Third Series 3: 535-542, pl. 21.

Pic M. 1943. Opuscula martiala X. - L’Echange, Revue

Linnéenne, Numéro spécial 10: 1-16.

Schaufuss L. W. 1864. Neue Coleoptera aus Neu-

Guinea. - Sitzungsberichte und Abhandlungen

der naturwissenschaftlichen Gesellschaft Isis in

Dresden 1864: 19-23.

Smith E. 1985. Outbreak of the longicorn Oxymagis horni

(Heller) (Coleoptera: Cerambycidae) on cocoa in the

Northern Solomon Province of Papua New Guinea.

- Proceedings, 9th International Cocoa Research

Conference, Lomé, Togo, 12-18 Feb. 1984: 607.

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Thomson J. 1864. Systema Cerambycidarum ou exposé

de tous les genres compris dans la famille des

Cerambycides et familles limitrophes. H. Dessain,

Liége: 1-352.

Thomson J. 1865. Systema Cerambycidarum ou exposé

de tous les genres compris dans la famille des

Cerambycides et familles limitrophes. H. Dessain,

Liége: 353-578.

Thomson J. 1864. Systema Cerambycidarum ou exposé

de tous les genres compris dans la famille des

Cerambycides et familles limitrophes. - Mémoires

de la Société Royale des Sciences de Liége 19:

1-540.

Vives E., Aberlenc H.-P., Sudre J. 2008. Entomofauna

de Vanikoro (Islas Salomon). 4a Contribution:

Coleoptera: Cerambycidae. - Heteropterus Revista

de Entomologia 8, No 2: 137-146.

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Waqa-Sakiti H., Lingafelter S. W. 2009. New Fijian

Callidiopini (Coleoptera: Cerambycidae). /n:

Evenhuis N. L., Bickel D. J. (eds) Fiji Arthropods XV.

- Bishop Museum Occasional Papers 106: 3-15.

Waqa-Sakiti H., Winder L., Lingafelter S. W. 2015. Review

of the genus Ceresium Newman, 1842 (Coleoptera,

Cerambycidae) in Fiji. - ZooKeys 532: 15-53.

Yokoi Y. 2015. Notes on the Callidiopini (Coleoptera,

Cerambycidae) across the Lombok Strait. - Elytra,

New Series 5: 185-205.

Yokoi Y. 2019. Callidiopini beetles in the collection of

Zoologische Staatssammlung Munchen (ZSM) Part

|. - Spixiana 42, No 1: 19-46.

Yokoi Y., Makihara H., Woro A. N. 2019. Callidiopini

beetles (Coleoptera, Cerambycidae) in the

collection of Indonesian Institute of Sciences (LIPI).

- Treubia 46, No 1: 1-20.

Received: 11.viii.2020.

Accepted: 30.xii.2020.

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ZIDEK, J.: Revision of Philacelota Heller, 1900 and Engertia Dalla Torre, 1912, with description of a new genus ...

pp. 397-411

Revision of Philacelota Heller, 1900 and Engertia

Dalla Torre, 1912, with description of a new genus

(Coleoptera: Scarabaeidae: Melolonthinae)

urn:lsid:zoobank.org:pub:A82E0/09-212/7-4D1E-B3F1-31F66315A2/B

Jiri ZIDEK

Prague, Czech Republic; jrzidek@gmail.com

Abstract: Philacelota submaculata Heller, 1900 is synonymized with PR maculosa (Brenske, 1896) comb. nov.,

Engertia papuana (Moser, 1913) and E. allolepis Prokofiev, 2019 are synonymized with E. amboinae (Brenske,

1897), and E. germanica Prokofiev, 2016 is synonymized with E. setifera (Moser, 1913). Engertia lii Keith, 2006

possesses characters that do not fit either Engertia Dalla Torre, 1912 or Philacelota Heller, 1900 and is placed in

Philgertia gen. nov.

Key words: Taxonomy, synonymy, new combinations, new genus, distributions, mutual relationships.

Introduction

Lacroix (2010) placed Philacelota Heller, 1910

in the Melolonthini Leach, 1819 and Engertia Dalla

Torre, 1913 in the Leucopholini Burmeister, 1855,

tribes that are maintained as distinct also in other

recent works (e.g. Bezdek 2016). However, Britton

(1978) did not recognize the tribe Leucopholini

and Matsumoto (2010) and Prokofiev (2016a)

questioned the split, as it does not appear to be

Supported by morphological evidence. Since the

question of tribal division is not the topic of this

paper and cannot be resolved by treating merely

two small genera anyway, the tribe Melolonthini is

regarded as an undifferentiated assemblage and

the discussion of the two genera is confined to their

mutual relationship.

Philacelota and Engertia are 14-22 mm long,

vittate melolonthines limited to eastern Indonesia,

New Guinea and the Philippines. They are poorly

represented in collections, with some species

known only from types. The only species currently

known from larger numbers of specimens are

Engertia amboinae (Brenske, 1897) and E. Ili

Keith, 2006, here placed in a new genus, whose

distrioutions are relatively wide. It is hoped that

the diagnoses and illustrations presented below

bring to light additional specimens and improve our

knowledge of these genera.

Material and methods

Generic characters are not repeated in species

diagnoses. Length (L) is given from the tip of the

clypeus to the tip of the elytra, pygidia are excluded.

Of the three antennalsegments (Scape, pedicel,

flagellum) only the flagellum is taxonomically useful.

In its proximal part, the funicle, the antennomeres

are either subequal or differ in length and number

but except for the wedge-shaped distal in the male

remain transversely unexpanded, whereas in the

distal part, the club, they are abbreviated and in

addition to differing numbers are also transversely

extended into thin lamellae. Hence, in this paper

the antennomeres of the club are called ‘lamellae’.

The terms ‘seta’ and ‘scale’ refer to degrees of

hair modification, both stiff and usually pigmented,

that differ in shape and cross-section. Whereas

setae are tubular and either erect or variously

inclined or recumbent, scales are flattened in

cross-section and always recumbent. However,

the distinction between hairs and setae and setae

and scales often becomes arbitrary because of the

presence of transitional forms, which invites terms

such as ‘setiform scales’ (e.g. Prokofiev 2019).

On the abdominal ventrites such forms appear as

‘setiform hairs’, but those that are not undulate are

in this paper called simply ‘setae’.

The presented habitus photos are mostly

of males, as females differ merely in having a

much smaller antennal club and can be assigned

to species without difficulty. Where necessary,

397

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specimens have been dissected and cleaned

aedeagi glued to tips of paper tongues pinned

underneath the specimens.

Collection codens:

BMNH - Natural History Museum (formerly British

Museum, Natural History), London, United

Kingdom;

CDT - Collection Dmitry Telnov, Riga, Latvia;

MFNB - Museum fur Naturkunde, Berlin, Germany;

MNHN - Muséum National d’Histoire Naturelle, Paris,

France;

SMFC - Senckenberg Naturmuseum, Frankfurt am

Main, Germany;

SMTD - Senckenberg Naturhistorische Sammlungen

Dresden, Germany.

Abbreviations:

PNG - Papua New Guinea;

WNG - Western New Guinea, Indonesia.

Results

Taxonomy

Philacelota Heller, 1900

Heller (1900: 11)

Type species: P. submaculata Heller, 1900 (=

Philacelota maculosa (Brenske, 1896) comb. nov.)

subsequent designation by Zidek (2018).

Redescription: Integument brown, reddish

brown to black or bicolored, with black head and

pronotum and brown elytra. Vestiture composed

of grayish-white or brownish oval and lanceolate

scales, grayish-white or yellowish setae and hairs.

Clypeus semicircular, its base formed by complete

frontoclypeal suture. Male antenna composed

of 10 articles with 7-lamellate club and funicle

consisting of a single long antennomere. Female

antenna composed of 10 articles with 5-lamellate

much smaller club and funicle consisting of

3 antennomeres of which proximal is longest.

Pronotum about one-quarter wider than long, widest

at base or at midlength, with margins complete and

sides weakly crenulate, base sinuate, front angles

obtuse, hind angles briefly pointed or obtuse,

and integument punctate except impunctate,

posteriorly wider medial sulcus present throughout

length. Scutellum obliquely triangular with anterior

margin concave and incomplete, micropunctate

and squamose near lateral margins, impunctate

—_—_

if |

398

on disc. Each elytron with 5 low, impunctate,

glabrous ribs separating slightly wider, shallow,

squamose or setose intervals. Sterna densely

covered by long grayish-white to yellowish hairs,

metasternal process short, peg-like, with rounded

tip, metasternal plate with medial furrow present

throughout length. Abdominal ventrites not connate,

covered by white or brownish-yellow scales or short

grayish-white setae. Femora squamose or setose.

Protibia unidentate or incipiently bidentate, with

dorsal and ventral longitudinal carina. Meso- and

metatibiae setose, without transverse carinae.

Distal tarsomere about as long as 2™ through 4%

tarsomeres combined. Tarsal claws large, with

ventral tooth at midlength. All spurs terminal,

attached medial to tarsal base; protibial spur

minute, meso- and metatibial spurs long and

slender. Aedeagus symmetrical, with parameres

about as long as phallobase. Parameres in dorsal

view either gradually tapering toward pointed tips,

or in distal third constricted and ending in weakly

expanded tips; in lateral view either gradually

tapering toward large, upturned shoe-shaped sharp

tips, or in proximal half with large dorsal humps and

in distal third with large, upturned shoe-shaped

round tips.

Philacelota maculosa (Brenske, 1896) comb.

nov. (Figs 1A-E, 2A & 8B)

Schoenherria maculosa Brenske (1896: 201), holotype

2 SMTD from ‘Ceylon’ (seen); Dalla Torre (1912:

181) catalogue; Keith (2006: 94) key, as Engertia.

Known only from the holotype.

= Philacelota submaculata Heller (1900: 12) syn. nov.,

holotype 4 SMTD from ‘Nord-Celebes, Toli-Toli’

(seen).

Other specimens examined: 14 BMNH from

N Sulawesi, Tondano; 14 SMFC from S Sulawesi,

Bantimurung.

Diagnosis: Head and pronotum grayish black,

elytra subtly two-colored, chestnut and dark brown.

Dorsal vestiture composed of spindle-shaped

white scales, on elytra restricted to lighter-colored

parts of intervals. Vestiture of abdominal ventrites

composed of somewhat smaller spindle-shaped

white scales and interspersed reddish-yellow and

white setae. Tips of parameres in dorsal view

weakly expanded but not distinctly sooon-shaped,

in lateral view as in P. sulana.

Remarks: Schoenherria Burmeister, 1855 is

a junior homonym of Schoenherria Blanchard,

1853, the replacement name is Carlschoenherria

Bezdék, 2016 (p. 15). Keith (2006, key) used

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ZIDEK, J.: Revision of Philacelota Heller, 1900 and Engertia Dalla Torre, 1912, with description of a new genus ...

pp. 397-411

His ptt M1) ‘ ais ‘ a eth ay hy ete He * cue RN

iy pe “ah ea hy Vekaa lps Gah iy hak ae Stun

Yael sey rey Miw i | Malet ay : st he BE ey Hi met a f As bith wa

REA A | ae hy ’ x ' uy NOt

Sen OP areca)

whe ARN prin Hy Wi

ikea “hs EATS et aN Hi

‘ a NOX Aug MY

MAY

wit

Figure 1A-G - Philacelota Heller, 1900 species. A - P. maculosa (Brenske, 1896) comb. nov., holotype 9, L = 18.5

mm. A - Dorsal habitus; B - Ventral habitus; C - Original labels; D - RP submaculata Heller, 1900 (= P. maculosa),

male antenna: E-G - Abdominal vestiture in type specimens of: E - P. submaculata, holotype <@; F - P. sulana Heller,

1900, holotype 3; G - P. jakli Zidek, 2018, holotype <' [not to scale].

ee 399

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

the semicircular clypeus to differentiate Engertia

maculosa from other Engertia species, however

it is a feature characteristic of Philacelota. In fact

the male type of Philacelota submaculata (Zidek

2018, fig. 9) and the female type of Schoenherria

maculosa differ from each other only in the size

of the antennal club, which in the latter closely

resembles that in the female of P. jakli (Zidek

2018, fig. 8). The type locality ‘Ceylon’ appears to

be erroneous, as all R submaculata specimens are

from Sulawesi.

Philacelota sulana Heller, 1900 (Figs 1F, 2B & 8C)

Heller (1900: 13), syntypes 1¢ & 12 SMITD from

[Indonesia] ‘Sula Mangoli’ (seen).

Other specimens examined: 20 SMTD from Sula

Mangoli [Mangole Island].

Diagnosis: Vestiture composed of short whitish

setae denser on pronotal disc and abdominal

ventrites, and sparse, confined to lighter-brown

parts of intervals, on elytra. Tips of parameres in

dorsal view expanded and spoon-shaped, in lateral

view as in PR. maculosa.

Remarks: The types are unnaturally red (Fig. LF),

evidently due to the kind of conserving fluid used.

The natural colors are shown in Fig. 2B.

Philacelota jakli Zidek, 2018 (Figs 1G, 2C, 7A &

8A)

Zidek (2018: 2), holotype 3 & allotype 2 SMITD,

paratypes 14’ & 12 BMNH. Known only from the

type series.

Type locality: Indonesia, Flores Island, Mt. Kuwus.

Diagnosis: Dorsal and abdominal vestiture

composed of closely spaced whitish setae.

Pronotum widest at midlength, its hind angles

obtuse. Protibia with an incipient second lateral

tooth, more developed in female. Paramere tips in

dorsal view slender and blunt, in lateral view each

expanded into a rounded ventral keel and a sharp

dorsal peak.

Philacelota (?) leucothea Prokofiev, 2019

Prokofiev (2019: 6, figs. 3-5), holotype 4, paratypes

14 & 22 MNHEN. Known only from the type series.

Type locality: Philippines, Luzon.

Diagnosis: Dorsal and abdominal vestiture

composed of minute grayish-white, spindle-shaped

scales of varying length covering entire elytra

including three feeble, hard-to-discern ribs on each

elytron. Pronotal medial sulcus narrow, present

400

only in anterior half of length. Parameres as in P.

maculosa (Fig. 8B).

Comments: | have not been able to examine this

species, nonetheless the original description and

illustrations are detailed enough to render an

opinion. The male antenna and the parameres

are those of Philacelota, but the female antenna

in Prokofiev’s figure 4E appears to consist of 11

articles (6 in the club, 3 inthe funicle, plus the scape

and pedicel), the scale-covered elytral ribs are said

to be only three on each elytron, and the narrow

medial sulcus is present only in the anterior half of

the pronotum. It appears the author photographed

antennae of both female paratypes, since the one

in his figure 4D consists of only 10 articles. The one

in figure 4E thus may be aberrant, which should be

possible to clarify by comparing it to its opposite

member. This notwithstanding, however, the less

numerous and densely squamose elytral ribs and

the short pronotal sulcus cause me to suspect that

this species belongs to another genus.

Engertia Dalla Torre, 1912

Dalla Torre (1912: 181), nomen nov. pro Phila Brenske.

Type species: Phila amboinae Brenske, 1897 by

monotypy.

= Phila Brenske (1897: 110), nomen praeoccup.

Redescription: Integument creamy — brown,

chestnut brown or black; vestiture composed

of white, grayish-white or brownish-yellow oval,

lenticular, lanceolate, and spindle-shaped scales,

grayish-white or yellowish setae and hairs. Clypeus

obliquely pentagonal, its base formed by faint but

complete frontoclypeal suture. Antenna in both

sexes composed of 10 articles with 5-lamellate

club and funicle consisting of 3 antennomeres

subequal in females and distal shorter, wedge-

Shaped in males; lamellae much shorter in

females. Pronotum about one-quarter wider than

long, widest at base, with margins complete and

sides weakly crenulate, base sinuate, front angles

obtuse, hind angles briefly pointed, integument

punctate except impunctate, posteriorly wider

medial sulcus present throughout length. Scutellum

obliquely triangular, with anterior margin concave

and incomplete, micropunctate and squamose near

lateral margins, impunctate on disc. Each elytron

with 5 low, impunctate glabrous ribs separating

Slightly wider, shallow, densely squamose intervals.

Sterna densely covered by long grayish-white to

yellowish hairs, metasternal process short, peg-

like, with rounded tip, metasternal plate with medial

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pp. 397-411

Figure 2A-C. Dorsal (top) and ventral (bottom) habitus of male Philacelota Heller, 1900. A - R submaculata Heller,

1900 (= P. maculosa (Brenske, 1896)), Sulawesi, L = 15 mm; B - P. sulana Heller, 1900, Sula Islands, Mangole, L

= 14 mm; C - P jakli Zidek, 2018, Lesser Sunda Islands, Flores I|., L = 14 mm [not to scale]. Parameres in Fig. 8A-C.

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TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

furrow present throughout length. Abdominal

ventrites covered by white or brownish-yellow

scales or short grayish-white setae or both, in some

specimens with medial concentration of scales

on 1° through 3” ventrites and 1st + 2™ ventrites

medially connate. Femora squamose or setose, or

both. Protibia unidentate, with dorsal and ventral

longitudinal carina. Meso- and metatibiae setose,

without transverse carinae. Distal tarsomere about

as long as 2™ through 4" tarsomeres combined.

Tarsal claws large, with ventral tooth at midlength.

All spurs terminal, attached medial to tarsal base;

protibial sour minute, meso- and metatibial spurs

long and slender. Aedeagus symmetrical, with

parameres about as long as phallobase. Parameres

in dorsal view gradually tapering, with tips either

pointed or weakly expanded into minute spoon-like

Shape; in lateral view gradually tapering toward

large, upturned shoe-shaped round tips.

Engertia amboinae (Brenske, 1897) (Figs 3-4, 7B

& 8D-E)

Brenske (1897: 111), as Phila; holotype 9 & paratype

13 SMFC (seen).

Type locality: [Indonesia], ‘Uliasser 94’ [Lease Islands].

Other specimens examined: 19 BMNH from Ambon

Island; 19 BMNH without data; 19 CDT from Indonesia,

Raja Ampat, Waigeo Island, 1.5 km W Waisai, Waisai sea

port, 00°25'56"S, 130°48'17’E, 21.11.2012, secondary

coastal vegetation; 34’ & 12 BMNH from WNG, Fakfak

[labelled as E. papuana]; paratypes 1¢ & 29 [labelled

as syntypes] MFNB from ‘Amboina’ [now Ambon]

Island; 22 MGNB from ‘Amboina’ [now Ambon] and

Buru islands; 29 MFNB [labelled as Phila maculosa]

from Finschhafen, PNG; 12 SMFC from Seram Island

[formerly Ceram], Central Maluku.

Engertia papuana Moser (1913: 283) syn. nov., as

Phila; holotype 3 MFNB, WNG, Takar (seen).

Engertia allolepis Prokofiev (2019: 3, figs. 1, 2A-G)

syn. nov.; holotype 3 MNHN from Ambon Island.

Known only from the holotype.

Diagnosis: Dorsal scales of more than one shape,

including on elytra. Abdominal vestiture composed

of oval scales and of setae present either on all

ventrites or only on last ventrite. Parameres in

dorsal view gradually tapering, with tips pointed.

Remarks: Brenske (1897) based the description

of Phila amboinae on a male, a female, and

presumably also on the three MFNB paratypes. The

original ‘n. sp.’ black-frame label is attached to the

female, which apparently resulted in subsequent

labeling of the female as the holotype and the male

as the paratype. Red type labels were obviously

402

added more recently, because the ‘Holo-’ and

‘Para-’ prefixes are written in ballpoint that became

available only much later. The original description

is complete enough to have allowed G. J. Arrow to

identify (‘determined from description’) the BMNH

female specimens as E. amboinae. In contrast to

the holotype (Fig. 3A), the abdominal vestiture of

the paratype (Fig. 3B) consists chiefly of grayish

setae; however, some small scales are present,

leading to the conclusion that the difference is

only a variation expectable in a population. Size,

Shape and color of the scales cannot be used to

discriminate among E. amboinae, E. papuana and

E. allolepis, because they all combine in these taxa,

especially on the elytra. For instance the elytral

scales in the holotype of E. papuana (Fig. 3C) are

markedly smaller and less imbricating than those

in the types of E. amboinae (Fig. 3A-B), but those

In specimens from WNG customarily identified as

FE. papuana (Fig. 4) do not differ in size and shape

from those of E. amboinae. The only detectable

difference in these specimens is the distribution of

setae on abdominal ventrites, which in the types

of E. amboinae are present in variable amounts

on all ventrites, whereas in E. papuana (Figs 3C &

4) and E. allolepis are confined to the 4‘ ventrite.

That could support E. papuana and E. allolepis as

subspecies of E. amboinae if it would not be for the

presence of E. amboinae in the Schouten Islands,

WNG (Prokofiev 2016b) and of E. allolepis in the

Moluccas (Ambon Island, Prokofiev 2019). From

the biogeographic standpoint it reduces these taxa

to infrasubspecific status, and they are therefore

synonymized with E. amboinae.

Engertia setifera (Moser, 1913) (Figs 5 & 8F-G)

Moser (1913: 284), as Phila; holotype 2 MFNB (seen).

Type locality: [Indonesia], Ocliasers, Saparoea [now

Saparua Island, Lease Islands].

Other specimens examined: 14 BMNH from Ceram

Island [now Seram]; 24 & 19 SMTD from ‘Ceram’.

The male, from Seram [Ceram] Island, was described

by Prokofiev (2015); the specimen is said to be later

deposited at the Moscow State University Zoological

Museum.

= Engertia germanica Prokofiev (2016a: 68, figs. 1-5)

syn. nov.; holotype ¢ & paratype 2 MFNB from

PNG, New Berlin.

Diagnosis: Integument usually chestnut brown,

rarely black with purple hue (holotype). Dorsal

vestiture composed of minute brownish-yellow

02-Jun-21 21:57:40

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Zipek, J.: Revision of Philacelota Heller, 1900 and Engertia Dalla Torre, 1912, with description of a new genus

pp. 397-411

aid ae cee =f me

isle: rt

Bele Pad PA ats Hy

A wis

: ace i +‘ te #

AN) | Uy

r ae l, ‘ : : 4 Ui be

, ee h ye ¢

= mas 5 a

{ne -

ns ¢

We RR, as “at a

Ue ay

Y =

“a ay “gh AST

ene ty a Ti ‘

7 (

ryl

{

= =e

= eas, eS

qe ae

Ree

as ye

* ——_

1 te

on. cs -

alas

“ ._- . “a

+h,

oy thet alae

Figure 3A-C. Dorsal habitus (left), base of left elytron (center) and abdominal ventrites (right) of type specimens of

Engertia Dalla Torre, 1912. A - E. amboinae (Brenske, 1897), holotype 2, L = 15.5 mm; B - E. amboinae, paratype

3,L =16 mm; C - E. papuana (Moser, 1913), holotype 3’, L = 16.5 mm [not to scale]. Parameres in Fig. 8D-E

403

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

or rarely white (holotype) spindle-shaped, closely

spaced but non-imbricating scales. Abdominal

vestiture ranging from short yellowish setae to

admixture of grayish-white to yellowish hairs and

sparse oval white scales. Paramere tips in dorsal

view weakly expanded into either brief or elongate

spoon-like shape.

Remarks: Regardless of the color of integument

and vestiture, E. setifera can be distinguished from

EF. amboinae by the slender, spindle-shaped, non-

imbricating dorsal scales (Fig. 5C) and the spoon-

Shaped, variably brief to elongate paramere tips

(Fig. 8F-G). E. germanica is known only from the

types that could not be examined because they

have yet to be returned to MFNB. However, there

is nothing in the original description and figures

that could be deemed to justify a new species, and

the specimens thus are hereby regarded as merely

a geographic extension of E. setifera previously

known only from central Moluccas.

Philgertia gen. nov.

Type species: Engertia lii Keith, 2006 by present

designation.

Derivatio nominis: Feminine, combination of

the names Philacelota and Engertia, alluding to its

relationship.

Description: Integument light to dark brown.

Vestiture composed of chalk white oval and

lanceolate scales, grayish-white and_ yellowish

setae and hairs. Clypeus intermediate between

semicircular and obliquely pentagonal, its base

formed by complete frontoclypeal suture. Male

antenna composed of 10 articles with 6-lamellate

club whose proximal lamella is markedly shorter

than preceding lamellae, and funicle consisting

of 2 antennomeres of which much shorter distal

interlocks with proximal lamella of club. Female

antenna composed of 11 articles with 6-lamellate

much smaller club, and funicle consisting of 3

antennomeres of which proximal spans half of

funicle length. Pronotum about one-quarter wider

than long, widest at base, with margins complete

and sides weakly crenulate, base sinuate, front

angles obtuse, hind angles briefly pointed, and

integument punctate except wide, impunctate

medial sulcus present throughout length. Scutellum

obliquely triangular with anterior margin concave

and incomplete, micropunctate and squamose

near lateral margins, impunctate on disc. Each

elytron with 5 impunctate glabrous ribs separating

slightly wider squamose intervals. Sterna densely

covered by long grayish-white to yellowish hairs,

it i

404

metasternal process short, peg-like, with rounded

tip, metasternal plate with medial furrow present

throughout length. Abdominal ventrites not

connate, covered by evenly distributed white oval

scales that become smaller and denser toward

lateral margins. Femora squamose and setose/

pilose. Protibia unidentate, with dorsal and ventral

longitudinal carina. Meso- and metatibiae setose,

without transverse carinae. Distal tarsomere about

as long as 2" through 4“ tarsomeres combined.

Tarsal claws large, with ventral tooth at midlength.

All spurs terminal, attached medial to tarsal base;

protibial sour minute, meso- and metatibial spurs

long and slender. Aedeagus symmetrical, with

parameres about as long as phallobase. Parameres

in dorsal view gradually tapering to constricted distal

third and ending in weakly expanded elongate tips;

in lateral view proximally with large dorsal humps

and in distal third with large downturned (i.e. convex

margin facing up) shoe-shaped round tips.

Philgertia lii (Keith, 2006) comb. nov. (Figs 6, 7C

& 8H)

Keith (2006: 91), as Engertia; holotype 3 & allotype

2 from Philippines, Leyte Island, Mt. Balocaue [=

Balocawe]; 24 & 29 paratypes from same locality

and date; 14 paratype from Philippines, Kabeng,

May 2000; 2 paratypes from Philippines, Palawan

|., Nov. 2004 [all types in Keith’s collection].

Type locality: Philippines (Leyte, Kabeng, Palawan Is.).

Other specimens examined: 94 & 32 BMNH from

Palawan Island, Salakot Mtn.

Diagnosis: As for genus.

Comments: The male club/funicle connection

consists of a wedge in the distal antennomere of

the funicle that fits in a V-shaped pocket in the

base of the proximal club lamella (Fig. 7C). The

relative length of the proximal lamella is fairly

constant in the specimens examined, but can be

expected to vary with population and/or to differ

in minor/major males. In all the males examined

the overlapping scales completely obscure the

integument of the elytral intervals, partial wear

exposing the integument has been observed only

in the females.

02-Jun-21 21:57:41

Zipek, J.: Revision of Philacelota Heller, 1900 and Engertia Dalla Torre, 1912, with description of a new genus. ..

pp. 397-411

=. = =

oes = =

= =

7 pi fi r,

—_ r 7 a

= m= = =

“

Ns en f

| |

’

‘

(

Figure 4. Engertia amboinae (Brenske, 1897), habitus and enlarged left elytrons of specimens from Fakfak, WNG,

previously identified as E. papuana (Moser, 1913). Top - male, L = 16 mm; Bottom - female, L = 19 mm [not to

scale]. Parameres in Fig. 8E.

405

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02-Jun-21 21:57:41

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

es i me 1) bbigy)

: Re vil pic)

ad a NH ie Wh ey

| aa 1] {a

\ Niwa

ba ye ‘s :

a om ia ee

Figure 5A-E. Engertia setifera (Moser, 1913). A - Male dorsal habitus, L = 16 mm; B - Male right antenna; C -

Enlarged base of left elytron of holotype 9; D-E - Variation in heads and vestiture of abdominal ventrites in two

male specimens from Central Moluccas, Seram (D - BMNH, E - SMTD) [not to scale]. Parameres in Fig. 8F-G.

406

Book4.indd 406 02-Jun-21 21:57:41

ZIDEK, J.: Revision of Philacelota Heller, 1900 and Engertia Dalla Torre, 1912, with description of a new genus ...

pp. 397-411

—— oan

; he di

y %% TA é v4 y YY bi "1 ‘ ‘ '

pas a) byline

1] nu. v vie 2 ny

te ‘

tae 2 ve att, NON rey Ve,

ee a te as re rin,

eu a more 10 ek :

duyars Thy) en 4

ese eee ee

ye. ese poy

hoa

‘i

Nah aa.

NY as mr ie

Figure 6. Philgertia lii (Keith, 2006). Top - male, L = 19 mm, habitus and enlarged left elytron; Bottom - female, L =

20 mm, dorsal habitus and enlarged abdominal ventrites [not to scale]. Antennae in Fig. 7C, parameres in Fig. 8H.

407

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figure 7A-C. Male (left) and female (right) antenna of: A - Philacelota jakli Zidek, 2018; B - Engertia amboinae

(Brenske, 1897); C - Philgertia lii (Keith, 2006) [not to scale].

408

Book4.indd 408 02-Jun-21 21:57:42

ZIDEK, J.: Revision of Philacelota Heller, 1900 and Engertia Dalla Torre, 1912, with description of a new genus ...

Mutual relationships of Engertia, Philacelota and

Philgertia

The habitus of these three genera Is so similar

that Keith (2006, key) placed Philacelota maculosa

(Brenske) and Philgertia Iii (Keith) in Engertia, in

effect regarding the three as one. However, a closer

examination reveals supraspecific differences

that justify maintaining Philacelota Heller and

Engertia Dalla Torre as separate genera and

referring Engertia lii to the new genus, Philgertia.

The supporting characters are shown in Table 1.

Connateness of abdominal ventrites has been

observed in some specimens of Engertia, but is left

out from the table as being too inconsistent. The

most obvious and stable differences can be seen in

the antenna and the aedeagus. The male antenna

of Philgertia is peculiar in possessing a club/

funicle interlocking mechanism, and the female

antenna of Philgertia, which lacks this mechanism,

consists of 11 rather than 10 articles (Fig. 7C).

The parameres of Philgertia Iii (Fig. 8H) differ from

those of Philacelota sulana Heller (Fig. 8C) only in

the downward vs. upward curvature of their tips,

which contrasts with the easily distinguishable

habitus of these species characterized by the

pp. 397-411

densely squamose vs. sparsely setose vestiture

(Figs 2B & 6).

Philacelota and Philgertia share the long

basal antennomere of the funicle (Fig. 7A, C), the

uniformly Squamose or setose vestiture of the

first through fourth abdominal ventrites without

scale/seta combination and without medial

concentration of scales/setae (Figs 1-2 & 6), and

close similarity of the parameres (in Philacelota

two of three species) (Fig. 8B-C, H). In Engertia,

on the other hand, the antennomeres of the funicle

are subequal (Fig. 7B), the vestiture of abdominal

ventrites contains combination of scales and setae

in variable proportions and medial concentration

of scales/setae is to some extent present on at

least the first and second ventrites (Figs 3-5), and

the parameres are more slender, with tips either

Sharp or slightly expanded (Fig. 8D-F). The medial

concentration of scales/setae may have to do with

connateness, which is absent in the other two

genera. Overall, Philacelota and Philgertia appear

to be more derived and more closely related to

each other than either of them is to Engertia.

Table 1. Comparison of generic characters of Engertia, Philacelota and Philgertia gen. nov.

[Characters _—s—Séd[ Ernerrtia Dalla Torre Philacelota Heller Philgertia gen. nov.

Clypeus: margin Obliquely pentagonal

6 — f-lamellate

Q - 5-lamellate

Intermediate semicircular/

pentagonal

S - 6-lamellate, proximal

lamella % as long

2 - 6-lamellate

S - 2, proximal antenno-

mere longer

9 - 3, proximal antenno-

Antenna: club d - 5-lamellate

Q - 5-lamellate

Antenna: funicle S - 3, distal antennomere | - single long antenno-

short mere

9 - 3 subequal antenno- | - 3, proximal antenno-

ryetes mere longest

mere longest

diffuse patches tributed

Paramere tips: lateral view |Shoe-shaped, upturned Shoe-shaped, upturned

Shoe-shaped, downturned

References

Acknowledgements

Bezdék A. 2016. Tribes Leucopholini and Melolonthini:

15, 224-236. In: LObl |., L6bl D. (eds) Catalogue

of Palaearctic Coleoptera. Volume 3. Revised and

updated edition. Brill, London / Boston.

Blanchard C. E. 1853. Description des_ insectes.

In: Hombros J., Jacquinot H. Zoologie, Tome

quatrigme. In: Dumont-d’Urville J. (ed.) Voyage

au pOle Sud et dans |’Océanie sur les corvettes

Astrolabe et la Zélée, executé par ordre du

Roi pendant les années 1837-1838 -1839

/ ay €o

lam grateful to Maxwell V. L. Barclay (BMNH),

Andrea Hastenpflug-Vesmanis (SMFC), Bernd Jager

(MFNB), Olaf Jager (SMTD), and Dmitry Telnov (CDT)

for information and specimen loans. AleS Bezdek

(Czech Academy of Sciences, Ceské Budéjovice),

Denis Keith (Chartres, France) and Richard Sehnal

(Czech University of Life Sciences, Prague) reviewed

and improved the manuscript.

409

Book4.indd 409 02-Jun-21 21:57:42

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figure 8A-H. Dorsal (top) and lateral (bottom) views of parameres in: A - Philacelota jakli Zidek, 2018, holotype;

B - Philacelota submaculata Heller, 1900 (= P. maculosa (Brenske, 1896)); C - Philacelota sulana Heller, 1900; D

— Engertia amboinae (Brenske, 1897), paratype; E - Engertia papuana (Moser, 1913); F - Engertia setifera (Moser,

1913), Seram |., SMTD; G - ditto, Seram |., BMNH; H - Philgertia lii (Keith, 2006) [not to scale].

0 =~ ©

Book4.indd 410 02-Jun-21 21:57:42

Book4.indd 411

ZIDEK, J.: Revision of Philacelota Heller, 1900 and Engertia Dalla Torre, 1912, with description of a new genus ...

-1840, sous le commandement de M. J. Dumont-

d’Urville, capitaine de vaisseau, publié par ordre

du gouvernement, sous la direction supérieure de

M. Jacquinot capitaine de vaisseau commandant

de la Zélée. Gide et Baudry, Paris: 422 pp, 20 pls

[plates issued in 1847].

Brenske E. 1896. Neue Melolonthiden aus Afrika und

Asien. - Stettiner Entomologische Zeitung 57:

178-205.

Brenske E. 1897. Neue Gattungen und Arten der

Melolonthiden aus Afrika und Asien. - Stettiner

Entomologische Zeitung 58: 96-120.

Britton E. B. 1978. A revision of the Australian chafers

(Coleoptera: Scarabaeidae: Melolonthinae).

Volume 2. Tribe Melolonthini. - Australian Journal

of Zoology, Supplement Series 60: 1-150.

Burmeister H. C. C. 1855. Handbuch der Entomologie.

Vierter Band. Besondere Entomologie, Forsetzung,

Zweite Abtheilung. Coleoptera Lamellicornia

Phyllophaga Chaenochela. Enslin, Berlin: x + 569

Pp.

Dalla Torre K. W. 1912. Fam. Scarabaeidae, Subfam.

Melolonthinae Ill, pp. 135-290. In: Junk W.,

Schenkling S. (eds), Coleopterorum Catalogus.

Volumen XX, Pars 49. W. Junk, Berlin.

Heller K. M. 1900. Neue Kafer von Celebes, IV. -

Abhandlungen der Berichte des_ koniglichen

zoologischen und anthropologische-

ethnographischen Museums zu Dresden 9: 1-46.

Keith D. 2006. Réflexions sur le genre Engertia et

description d’une nouvelle espéce des Philippines

(Col. Scarabaeoidea Melolonthidae). - Lambillionea

106: 90-94.

Lacroix M. 2010. Melolonthinae — afrotropicaux

(Scarabaeoidea, Melolonthidae). Genera et

catalogue commenté. Editions Lacroix, Collection

Hannetons, Paris: 277 pp.

pp. 397-411

Matsumoto T. 2010. Taxonomic study on the genus

Psilopholis Brenske (Scarabaeidae, Melolonthinae,

Melolonthini) with consideration of divisions of the

subtribe Melolonthina. - Elytra 38: 239-247.

Moser J. 1913. Beitrag zur Kenntnis der Melolonthiden

(Col.), |. - Deutsche Entomologische Zeitschrift

1913: 271-297.

Prokofiev A. M. 2015. A male of Engertia setifera

(Coleoptera: Scarabaeidae: Melolonthinae) from

Seram Island, Indonesia. - Baltic Journal of

Coleopterology 15: 25-28.

Prokofiev A. M. 2016a. Novye rody i vidy papuanskikh

Leucopholini Melolonthini (Coleoptera:

Scarabaeidae: Melolonthinae). New _ genera

and species of the Papuan Leucopholini and

Melolonthini (Coleoptera, Scarabaeidae:

Melolonthinae). - Euroasian Entomological Journal

15: 68-74. [in Russian, English abstract]

Prokofiev A. M. 2016b. On the phytophagous scarabs

of the subfamilies Dynastinae, Rutelinae, and

Melolonthinae from the Schouten Islands (=

Kepulauan Biak), Indonesian Papua (Coleoptera:

Scarabaeidae). - Species Diversity 21: 71-77.

Prokofiev A. M. 2019. Two new species of Engertia

Dalla Torre, 1913 and Philacelota Heller, 1900

from Indonesia and the Philippines with a revised

identification key (Coleoptera: Scarabaeidae:

Melolonthinae). - European Journal of Taxonomy

561: 1-13.

Zidek J. 2018. A new species of Philacelota Heller

(Scarabaeidae: Melolonthinae: Melolonthini) from

Flores Island, Indonesia. - Insecta Mundi 0679:

1-6.

Received: 05.x.2020.

Accepted: 20.xi.2020.

411

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

412

Book4.indd 412 02-Jun-21 21:57:43

Zi, A., DE Vos, R. & Epwarps, E. D.: The torsilinea species group of the genus Platyja Hubner, [1823] ...

pp. 413-430

The torsilinea species group (Subgenus Mocrendes

Nye, 1975) of the genus Platyja Hubner, [1823]

(Lepidoptera: Erebidae), with descriptions

of four new species

urn:lsid:zoobank.org:pub:EC5D4EC4-3411-4EC8-815F-282BCB6E /47C

ALBERTO ZILLI *?, RoB DE Vos * & Epwarp D. Epwarps °

1 - Corresponding author: Natural History Museum, Life Sciences, Insects Division, Cromwell

Road, SW7 5BD, London, United Kingdom; a.zilli@nhm.ac.uk (ORCID: OO00-0002-3416-8069)

2 - Naturalis Biodiversity Center, Department of Entomology, Darwinweg 2, NL-2333, CR Leiden,

The Netherlands; rob.devos@naturalis.nl

3 - CSIRO Ecosystem Sciences, Australian National Insect Collection, GPO Box 1700, Canberra,

ACT 2601, Australia; ted.edwards@csiro.au (ORCID: O0O00-0002-4637-6391)

Abstract: Following a general introduction to features and putative relationships of the genus Platyja Hubner,

[1823], a review of the Indo-Australian section corresponding to the subgenus Mocrendes Nye, 1975 (= torsilinea

species group) is presented. This group, ranging from the Indian subcontinent to the Australo-Papuan region, is re-

defined on basis of features of the male genitalia (presence of an ampulla-like process on costa termed ‘ampulloid’)

and shown to currently comprise nine species, four of which new to science and herein described, namely P. (M.)

heterogenea sp. nov. (Sulawesi), P. (M.) sukabumensis sp. nov. (Java), P. (M.) lemurella sp. nov. (Java, Flores) and

P. (M.) carpentariella sp. nov. (Queensland). The concepts of P. (M.) torsilinea (Guenée, 1852) and P. (M.) ciacula

Swinhoe, 1893 are found to have been based on the two sexes of the same species and are therefore brought

together, though the latter is ranked as a subspecies (stat. nov.) of the former to account for some singularities

in the nominotypical population from the Andaman Islands. The supposedly long-known females of ‘torsilinea’

are instead shown to be the hitherto unknown ones of P. (M.) exviola Hampson, 1891. The uncertain identity of

Megacephalomana pilosum (Pagenstecher, 1888), nomen dubium, is briefly discussed.

Key words: Review, moths, taxonomy, types, Indo-Australia.

Introduction expected, but instead of running down to the anal

margin, before 1A+2A it turns back upwards so as

Book4.indd 413

By virtue of their greatly atypical pattern, many

authors more or less implicitly recognised the close

relationships of species variously circumscribed

within genera like Platyja HUbner, [1823], lontha

Doubleday, 1842, Facidia Walker, 1865 and

Megacephalomana Strand, 1943 (replacement

name for Megacephalon Saalmuller, 1880) (e.g.,

Tams 1924; Joannis 1929; Sugi 1982; Holloway

2005). Not least, some species have variously

been associated with one or other of these genera,

whose boundaries have therefore remained rather

blurred.

The distinctive feature of these species

occurs in the unusual trend of the postmedial line

of forewing. In fact, it greatly protrudes beyond

the discal cell into the distal field, being actually

positioned where a submarginal line would be

to outline below CuA1 a pedunculate lobe ‘hanging

down’ from the median field. It then reaches the

discal cell from below and proceeds proximally along

Cubitus for a short section up to where it incurves

posteriorly to eventually reach the anal margin. As a

result, the median and distal fields of the forewing

appear as mutually interlobed on the wing disc.

Notably, the pedunculate lobe bubbles as a sort of

drop between CuA2 and 1A+2A perpendicularly to

the anal margin. The filling of this unusual mark is

generally much paler than the ground colour.

There are however exceptions to this

generalised blueprint, such as species in which the

body structure and habitus strictly match those of

typical “Platyja s. |.” but whose crosslines follow

instead another path or are obliterated, and this

most often in only one of the sexes. As a matter

413

Tan

€

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

of fact, in several species the unusual trend of the

postmedial is clearly visible only in the females,

as the pattern of males is either faint or entirely

obscured. Species of this genus group may in fact

show a remarkable sexual dimorphism, with males

highly deviant not just in pattern but also in wing

Shape from the usually more uniform aspect of

females. In fact, sometimes male and female of a

same species have been described as two separate

species. In several species, the males bear tails on

the hindwings or elevated ridges flanked by loosely

scaled interspaces whose most likely function

is that to act as resonators, a feature also seen

in many other erebid genera (e.g., Anisoneura

Guenée, 1852, Delgamma Moore, 1885, Hypospila

Guenée, 1852). Males often also bear voluminous

tufts of scales on legs or abdomen and some of

the species whose hindwings are tailed also show

an exceedingly elongated abdomen. The females

instead are sometimes so similar between species

as to make association of sexes difficult when

more than one species occur sympatrically, while

taxonomic identification of males is usually easier.

Such difficulties long prevented a consistent

taxonomic approach to this assemblage of

genera until the character that species of the

historical generic groups Platyja Hubner, [1823]

and lontha Doubleday, 1842 show tripectinate

antennae in the male sex was highlighted (Zilli,

2001). Such outstanding and rare feature among

the Lepidoptera had long been overlooked

simply because rami of the third row lie precisely

midventrally along the flagellum. Furthermore, in

several species they appear as partially reclinate

lamellae, so that unless the antenna is observed

obliquely it looks bipectinate. A suggestion that

maintaining lontha separate from Platyja could

make the latter paraphyletic (Zilli 2001), was

eventually implemented by Holloway (2005) with

Subsuming the former into the latter.

If the characters of the postmedial line and

tripectinate male antennae are considered, the

majority of species of the historical genera treated

here actually showthem jointly, thoughsome outliers

with one or the other of the two features remain,

that is species with clearly Platyja-like postmedial

but male antennae filiform (or slightly tuberculate)

or vice versa. For the sake of precision though, some

species of this assemblage whose male antennae

look filiform are in reality monopectinate, as they

have lateral rami reduced but the midventral row

remains well-expressed. A more precise statement

of this particular character of the group is thus that

of the male antenna bearing a midventral row of

414

rami or teeth.

Regarding the systematic affinities of the

Platyja genus group, various attempts have been

made to trace its relationships. Holloway (2005;

2011) placed Platyja in unresolved Catocalinae/

Erebinae, but considered Avittonia Hampson,

1926, which, except for the trend of postmedial

(no pedunculated loop being present) strictly

matches in habitus species of Platyja and also

has midventrally serrate male antennae, to be

nearest to “Ugia” serrilinea Hampson, 1926, a

species with normal antennae instead. Zahiri et al.

(2012), in a molecular phylogeny of the Erebidae,

recovered Platyja umminia (Cramer, [1780]) in a

cluster together with Hulodini, Ercheiini and other

genera, namely Ischyja Hubner, [1823], Praxis

Guenée, 1852, Anisoneura Guenée, 1852, Ercheia

Walker, 1858, Hulodes Guenée, 1852 and Ericeia

Walker, 1858, although without support. However,

in the analysis by Homziak et al. (2019) the nearest

neighbour of Platyja umminia is Ischyja with good

support, both clustering with different genera

from the previous analysis except for Anisoneura.

A relationship between Platyja and Ischyja seems

thus to have been assessed, though the exact

topology of the phylogenetic relationships of the

whole group has yet to be reconstructed.

In this respect, it is worth bringing here

important observations by Banziger (1982 and pers.

comm.), who recorded species of Platyja (umminia,

sumatrana [C. & R.] Felder & Rogenhofer, 1874,

suffumata Guenée, 1852) to be particular fruit-

piercing moths without tearing hooks and erectile

barbs on proboscides, typical instead of fruit-

piercers par excellence such as calpine erebids, but

with an extremely sclerotized proboscis ending in

Sharp, beak-like tip, and galeae capable of alternate

motion. These features have also been found,

though with notable modifications, in Sarobacala

albopunctata (Semper, 1901), a species with

habitus profoundly different from the Platyja genus

group and with simple antennae. Banziger (pers.

comm.) has further kindly provided us with other

important details regarding the unique proboscis of

these two genus groups, such as the presence of

blade-like bristles along the dorsal food-entrance

slot. However, these do not work to pierce a hole

through the skin of the fruit but rather slash the

fruit pulp to release its sap (“rasping spines” sensu

Banziger 1980). They also work as a filter to prevent

large pulp clumps from obstructing the food canal,

besides in part to keep the galeae linked together

(Banziger pers. comm.).

Distribution — of

the

group is _ typically

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Zit, A., DE Vos, R. & Epwarps, E. D.: The torsilinea species group of the genus Platyja Hubner, [1823] ...

palaeotropical, with extensions into the SE

Palaearctic and Australasia. Due to the obviously

unsatisfactory assortment of species among

genera of this group and presence of a number of

undescribed ones, it was decided to undertake a

whole review of Platyja and its relatives, of which

this contribution dedicated to the Indo-Australian

torsilinea species group, for which the subgeneric

name Mocrendes Nye, 1975 is available, represents

the first account.

Material and methods

Abbreviations:

ANIC - Australian National Insect Collection, Canberra,

Australia;

BMNH - British Museum (Natural History), London,

United Kingdom (currently NHMUK);

MHNG - Muséum d’histoire naturelle, Geneva,

Switzerland;

NHMUK —- Natural History Museum, London, United

Kingdom;

RMNH — Rijksmuseum voor Natuurlijke Historie, Leiden,

The Netherlands;

SMF - Senckenberg Museum, Frankfurt am Main,

Germany.

Fwl/fwl - forewing length;

N - number of specimens;

X - average.

Results

Taxonomic part

Characterization of the genus-group taxa

Platyja Hubner, [1823]

Platyja Hubner, [1823]. In 1816-[1825], Verzeichniss

bekannter Schmettlinge [sic] (17): 268. Type species:

Phalaena Noctua umminia Cramer, [1780], by

subsequent designation by Moore ([1885], in 1884-

1887).

= lontha Doubleday, 1842. The Entomologist 1:

297. Type species: lontha umbrina Doubleday, 1842, by

monotypy. Synonymised with Platyja by Holloway (2005).

= Cotuza Walker, 1858. List of the specimens of

lepidopterous insects in the collection of the British

Museum 15: 1545 [key], 1551. Type species: Cotuza

drepanoides Walker, 1858, by monotypy. Cotuza

drepanoides is a junior subjective synonym of Phalaena

Noctua umminia.

pp. 413-430

= Ginaea Walker, 1858. List of the specimens of

lepidopterous insects in the collection of the British

Museum 15: 1546 [key], 1637. Type species: Ginaea

removens Walker, 1858, by monotypy. Ginaea removens

is a junior subjective synonym of Phalaena Noctua

umminia.

= Cremnodes [C. & R.] Felder, 1874. See below

under Platyja subg. Mocrendes Nye, 1975.

= Yerongponga T. P. Lucas, 1901. Proceedings of

the royal Society of Queensland 16: 82. Type species:

Yerongponga exequialis T. P. Lucas, 1901, by monotypy.

Synonymised with Platyja by Edwards (1996).

= Mocrendes Nye, 1975. See below under Platyja

subg. Mocrendes Nye, 1975.

Taxonomic remarks: Ginaea was

first synonymised with Cotuza by Moore (1867,

misspelled as Ginea), deemed by this author

to be the correct generic name for PR umminia;

Subsequently, Moore ({[1885] in 1884-1887)

eventually subsumed Cotuza into Platyja. As

noted in the introduction above, lontha was sunk

with Platyja by Holloway (2005), same as _ for

Mocrendes, which in turn had been proposed as a

replacement name for Cremnodes [C. & R.] Felder,

a junior homonym to Foerster’s in the Hymenoptera.

Yerongponga was conditionally synonymised with

Platyja by Kitching (1987), a suggestion brought

forward into a formal way by Edwards (1996).

Among the authors who most recently treated this

group, Holloway (2005) retained all junior generic

names as full synonyms of the senior one, though

he suggested that a number of them, such as lontha

and Mocrendes (as Cremnodes), could actually be

used to circumscribe subgenera. Kononenko &

Pinratana (2013) explicitly recognised subgeneric

Status to Platyja s. str., lontha and Mocrendes.

This approach is preliminarily followed here as

from the screening over the whole genus group so

far undertaken, species of the torsilinea group do

actually show some binding characters that enable

their circumscription under a revised subgeneric

concept of Mocrendes. However, a conclusive

diagnosis and taxonomic arrangement of the

Platyja genus group will become available once the

whole revision of its various lineages is concluded.

Platyja subgenus Mocrendes Nye, 1975

Mocrendes Nye, 1975. The generic names of moths of

the world 1: 319. Replacement name for Cremnodes

[C. & R.] Felder, 1874 (preoccupied by Foerster, 1850).

Type species: Cremnodes lemur [C. & R.] Felder, 1874.

Synonymised with Platyja by Holloway (2005) but used

at subgeneric level by Kononenko & Pinratana (2013) by

virtue of ICZN (1999: art. 61.3.1).

415

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= Cremnodes [C. & R.] Felder, 1874. In: Felder C.

et al., [1865]-1875, Reise der Osterreichischen Fregatte

Novara um die Erde in den Jahren 1857, 1858, 1859

unter den Befehlen des Commodore B. von Wullerstorf-

Urbair (Zoologischer Theil) 2 (2) Lepidoptera (Atlas):

Erklarung der Tafel CXIll, pl. 113, fig. 9. Type species:

Cremnodes lemur [C. & R.] Felder, 1874, by monotypy.

A junior homonym of Cremnodes Foerster, 1850

[Hymenoptera]. The objective replacement name is

Mocrendes Nye, 1975. Synonymised with Platyja by

Holloway (2005).

Diagnostic remarks: Species falling into

this group have usually been considered members

of Platyia whose males have filiform antennae,

elongated forewings andstronglymodifiedhindwings

with either variously lobate termen or loosely scaled

areas with distorted veins (resonator), often with

both. As noted above, this type of antennae should

better be stated as midventrally monopectinate.

This feature, combined with structural characters of

the genitalia, allows a more formal circumscription

of the group than that based on wing features

which are susceptible of great variability due to

phenomena of sexual selection. All species of the

group show in fact a peculiar configuration of the

male genitalia, which bear along the costa a long

Slender, distally clubbed process bearing sparse

microsetae much reminiscent of an ampulla,

if it was not for its position (ampullae typically

branching off from the ventral side of valvae, cf.

Pierce, 1909). In the light of this character, which

will be referred to here as the ‘ampulloid (process)’,

species with less extreme male wing shapes also

need to be combined with Mocrendes.

Some members of the group show a

remarkably similar pattern to African species

currently comprised within Megacephalomana

and Platyia s. |., such as M. rivulosum (Saalmiuller,

1880), and P. semifimbria (Walker, [1858]). The

latter is actually closest to the former but has been

moved to Platyja by Holloway (2005) from Facidina

Hampson, 1926, where it had been placed by

Poole (1989) following its removal from Facidia

Walker, 1865. However, the absence of the costal

ampulloid process, the conspicuously tripectinate

antennae and the normal hindwing configuration in

males of these African taxa indicate that they are

not directly related to Indo-Australian Mocrendes.

Annotated list with descriptions of new species

Platyja (Mocrendes) torsilinea torsilinea (Guenée,

1852) (Figs 1-5, 23-25, 36-40, 51-54 & 64)

Ophisma torsilinea Guenée, 1852. Species général des

416

Lépidoptéres, Noctuélites 3: 240. Locus typicus: Inde

centrale [recte: Silhet]. Holotypus: 2, by monotypy, in

NHMUK (via C. Oberthir collection) [examined].

Note: despite the type locality stated in the

description, the holotype is clearly labelled by

Guenée as from Silhet, with the indication “C’est la

Platyja (Mocrendes) torsilinea ciacula Swinhoe,

1893, stat. nov. (Figs 6, 29, 41, 55 & 65)

Platyja ciacula Swinhoe, 1893. The Annals and

Magazine of natural History, including Zoology, Botany,

and Geology (6) 12 (70): 263. Locus typicus: Port Blair,

Andamans. Holotypus: <@, by monotypy, in NHMUK

[examined].

Taxonomic remarks: The concepts of Platyja

(Mocrendes) torsilinea and P. (M.) ciacula have

hitherto remained separate in consequence of

having been based on specimens of the two sexes,

renowned difficulties in correctly associating males

and females of Platyja and the circumstance that

the holotype (2) of torsilinea either remained

unavailable to authors such as Swinhoe (1893)

and Hampson (1894) or, when it eventually entered

the BMNH holdings in 1923 (via C. Oberthur’s

collection), it was kept in a separate collection.

Accordingly, differences between the holotype

compared to other females from India in the BMNH

that were assumed to be true torsilinea, but which

are the so far unrecognised females of P. (M.)

exviola (See below), were not realized until now. In

fact, the holotype of torsilinea (Fig. 23) matches

females of ciacula where only this species of the

group occurs, while the misidentified females of

‘torsilinea’ originated from places where only males

of exviola are known (SW Ghats), so that the names

torsilinea and ciacula are here brought together.

It is likely that the misunderstanding was driven

by Guenée’s (1852) wrong statement about the

type locality of torsilinea, although correct in the

handwritten holotype’s label, which may have led

to the assumption that this species was present

further south in India. However, when specimens

from the Andaman Islands, type locality for ciacula,

are considered (Figs 6 & 29), the females, and to

a lesser extent also the males, show a particular

pattern feature which is more reminiscent of

exviola than typical torsilinea, namely the inner line

bordering the preapical patch of forewings that more

definitely curves upwards to reach the costa less

obliquely than in typical torsilinea. The outer line

bordering the patch is also feebly but consistently

02-Jun-21 21:57:43

Ziti, A., DE Vos, R. & Epwarps, E. D.: The torsilinea species group of the genus Platyja Hubner, [1823] ...

pp. 413-430

Figures 1-15. Habitus of Platyja (Mocrendes) species (44). 1 - P. (M.) torsilinea torsilinea (Guenée, 1852), Assam,

Cherrapunji; 2 - ditto, Vietnam; 3 - ditto, Vietnam; 4 - ditto, Malaysia, Genting [Sempah]; 5 - ditto, Brunei, Bukit

Retak; 6 - P. (M.) torsilinea ciacula Swinhoe, 1893, holotypus, Andaman; 7 - P. (M.) exviola Hampson, 1891,

holotypus, India, Nilgiris; 8 - R (M.) heterogenea sp. nov., holotypus, Sulawesi, Minahassa; 9-11 - ditto, paratypi,

Bolaang; 12 - P. (M.) sukabumensis sp. nov., holotypus, Java, Priangan; 13 - P. (M.) flavimacula Semper, 1901,

topotypus, Mindanao, Mt. Agtuuganon; 14 - ditto, topotypus, Carmen; 15 - ditto, Luzon, Mt. Polis [scale bar = 1 cm].

& 417

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more sinuous in the Andaman samples.

Besides these features, there are no other similar

traits between the Andaman population and

exviola, which is a species sharply differentiated

in the genitalia of both sexes and the males have

no hindwing resonator. A question arises then as

regards the taxonomic treatment of the Andaman

population. As noted herein, P. (M.) torsilinea is a

broadly distributed species showing quite some

variation. It appears to be very variable in the male

sex, somewhat showing two morphoclines while

proceeding to the Northeast and the Southeast of

its range. In fact, individuals from the NE such as

in North Vietnam and SE China are less extreme in

the elongation of forewings, which are thus broader,

and also with weaker secondary sexual characters

of the hindwings (e.g., Fig. 2). In contrast, males

from the SE (Sundaland) sport particularly narrow

forewings and more strongly modified hindwings

(e.g., Fig. 5). However, males either from other

areas or the same populations are less divergent.

Females instead appear to be more uniform in

external appearance.

This situation prompted us to dissect males from

several geographic locations (Figs 36-41, 51-55,

for a selection), whose outcome was that notable

variation occurs in features such as the slenderness

and elongation of the ampulloids and cuculli, partly

also in the degree of incrassation and very outline

of the saccular lobes, in the thickness of the uncus

and the relative size and orientation of the vesical

lobes. Nevertheless, the observed variation neither

showed geographic congruence nor consistency

among different characters, so that we could notfind

in the genitalia sufficient morphological evidence to

split the concept of torsilinea any further. This does

not exclude the possibility that, with more powerful

insights into the genetic constitution of different

populations, valuable clues to substructure this

morphospecies into more biospecies can be found.

The same morphological considerations basically

apply to individuals from the Andamans, though

their unusually shaped preapical patch of forewings,

peripheral distribution, and insularity, to which also

the somewhat more compact forewing shape of the

females, suggest recognizing their singularity by

ranking the Andaman population at the subspecific

level, as Platyja (Mocrendes) torsilinea ciacula

Swinhoe, 1893 (stat. nov.). It is worth noting that

a male from Peninsular Malaysia (Fig. 4) shows a

forewing patch whose inner edge is not dissimilar

from that of Andaman samples, and that another

pattern feature seen in these, that is the less

oblique forewing postmedial, is occasionally found

—_—_—

418

also in typical torsilinea.

Diagnostic remarks: Subspecies torsilinea

(Figs 1-5, 23-25): Male (fwl = 24-29 mm, x

= 26.17 mm, N = 15) with elongate acuminate

forewing tipped by conspicuous, well-defined, dark

brown subtriangular preapical patch, hindwing with

paired elongated loosely scaled areas alongside

the median vein in relief (resonator) (this feature is

more weakly expressed in populations from the NE

of the range) and smoothly convex distal margin.

Female (fwl = 23-28.5 mm, x = 25.50 mm, N =

16) with elongate albeit broad forewing tipped by

conspicuous, well-defined, dark brown subtriangular

preapical patch, this very elongate and with inner

edge distinctly oblique with respect to costa.

Subsp. ciacula (Figs 6 & 29): As in nominotypical

subspecies (4fwl = 26 mm, x = 26 mm, N = 2; 9fwl

= 24 mm, N = 1) except for inner edge of preapical

patch distinctly arched and ending approximately

perpendicular on costa, particularly in the female,

whose forewing is also less elongated.

In the male genitalia, the saccular lobes are

subtriangular, variably produced, the cucullilongand

Slender, bent ventrally, the ampulloids of variable

thickness, usually outwardly arched, and the juxta

consists of a very broad sub-rhomboid inferior plate,

membranous along the midline, and two very small

apical winglets (Figs 36-41). The phallus vesica

shows a broad central corpus from where two main

anteriorly oriented conical lobes protrude, these of

variable size and relative orientation, among which

another big lobe terminated by an apical lobelet

(this not everted in Fig. 51) and a short blunt apically

bilobed one oriented towards the phallus shaft

lie; a small, serrated plate in continuation of the

Shaft is present at base of the vesica corpus (Figs

51-55). In the female genitalia, the lodix lobes are

subtriangular, slightly constricted at middle, and

terminating into smoothly rounded tips; the ductus

bursae is wide and flat, almost entirely sclerotized,

the sclerotization following up on sternum A8 as a

triangular plate; and the corpus bursae shows a

wrinkled cervical sclerite (Figs 64-65).

Distribution: NE India (See note above regarding

Supposed presence in Central India), Andaman

Islands, Nicobar Islands, Burma, SE _ China,

Thailand, Indochina (Laos, Cambodia, Vietnam),

Malay Peninsula, Sumatra, Borneo, Java, Bali.

The Andaman population corresponds to subsp.

ciacula, to which also that from Nicobar Islands

(Kononenko & Pinratana 2013) should likely be

attributed.

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Ziti, A., DE Vos, R. & Epwarps, E. D.: The torsilinea species group of the genus Platyja Hubner, [1823] ...

Platyja (Mocrendes) exviola Hampson, 1891 (Figs

1, 26, 42, 56 & 66)

Platyja exviola Hampson, 1891. Illustrations of typical

specimens of Lepidoptera Heterocera in the collection

of the British Museum 8: 22, 91, pl. 147, fig. 18 [name

dates from p. 91]. Locus typicus: [India] Nilgiri district,

S. & W. slopes, 3000 feet. Holotypus: 3', by monotypy, in

NHMUK [examined].

Taxonomic remarks: A_ generally poorly

known species whose females have mostly been

misidentified as those of Platyja (Mocrendes)

torsilinea (See above).

Diagnostic remarks: Compared to other

relatives of the torsilinea group, P. (M.) exviola

(Sfwl = 24-26 mm, x = 24.67 mm, N = 3; °fwl

= 23-25 mm, x = 24 mm, N = 2) is the species

with the least marked secondary sexual characters

of the male (Fig. 7). In fact, its forewing is more

compact than in its allies, and its hindwing is

without signs of distorted veins and is devoid of a

resonator; a preapical patch is present but greatly

blurred into the overall dark brown ground colour

of the forewing. The female looks like one of

torsilinea albeit with less elongated wings (Fig. 26);

in consequence of this, the outer half of the median

field of the forewing is less projected outwardly

and the dark brown preapical patch shorter, with

its inner edge, if not fully perpendicular, much less

oblique with respect to costa than in its congener

(notably, these differences do not apply in relation

to torsilinea subspecies ciacula, see above and Fig.

29). The median field is also overall concolorous,

warm brown, with the exception of the paler

coloured pedunculate lobe, which also appears to

be narrower, less globular than in nominotypical

torsilinea. In the male genitalia, the superior angle

of the sacculus is nearly a right angle and very

feebly produced, so that the saccular lobes are

practically non-existant; the ampulloids are very

long, slender and outwardly arched, the cuculli

are also very long and slender, arched dorsally, so

as to form a U-configuration with the ampulloids;

the juxta is similar to that of torsilinea, but the

inferior plate is more broadly rounded and there

is a longer and thinner constriction between this

and the apical winglets (Fig. 42). The phallus is

almost abruptly bent at middle, with a voluminous

vesica whose main corpus projects posteriorly into

a conical, heavily scobinate lobe and, beyond a

mesial constriction, anteriorly into a large trilobate

sack, the largest lobe subquadrate and distally

scobinate, the smallest bearing a stout apical

cornutus, and the intermediate in size conical in

pp. 413-430

Shape (Fig. 56). In the female genitalia the lodix is

massive, with greatly expanded, obliquely truncated

lobes; the ductus bursae is long and comparatively

narrow, sclerotized only in its anterior half, the

sternum A8 being thus wholly membranous; in the

bursa copulatrix there are two cervical sclerites and

another one in correspondence with small conical

projection (Fig. 66).

Distribution: All known records are from the SW

Ghats in Peninsular India, from where the species

is likely to be endemic.

Platyja (Mocrendes) heterogenea sp. nov. (Figs

8-11, 27-28, 43, 57 & 67)

Holotypus 4 NHMUK: [Indonesia: Celebes (=

Sulawesi)] Minahassa, 1920, Coll. P. J. v. d. Bergh Lzn.,

NHMUK010918195, slide NHMUKO10315357.

Paratypi 446 &29: 2 NHMUK: Menado, Nord Celebes,

[coll.] W. Niepelt, Zirlau, Joicey Bequest 1934-120,

NHMUK 010918198, slide NHMUKO10316309; 4,

2 RMNH: 3, Manado, Tanggarie, 410 m, VI.1936, J. P.

A. Kalis leg., coll. J. M. A. v. Groenendael; 4, Bolaang

[= Bolaang-Mongondow]; 26), idem, 1919; 2, Dumoga-

Bone N.P. [= Bogani Nani Wartabone National Park],

Gunong Mogogonipa, 0°27’N 123°57’E, 900-1008 m,

20-23.V.1983, multistrate evergreen forest/moss forest,

R. de Jong leg.

Additional material: 3 MHNG: Sud Sulawesi, R[ou]

te de Rantepao a Palopo, 1700 m, G. Lecourt leg.

Derivatio nominis: The species’ name, derived

from the Greek-derived Latin word “heterogeneus”,

stresses the ‘heterogeneous’ appearance of the

moth regarding several respects, namely the

discrepancy in wing shape between the fore and

hindwing of the male, the combination of features

otherwise appearing disjunct in other species

(torsilinea, flavimacula) and the dimorphism of the

two sexes. The name is an adjective in feminine

gender in agreement with the generic name.

Male (Figs 8-11): Fwl = 26-27 mm, x= 26.60 mm,

N = 5; head large, frons flat, eye globular, antenna

midventrally uniserrate, labial palpus stout, second

joint thick and compactly scaled, blackish brown,

slightly incrassate distally, third joint short rod-

like, distally clubbed. Patagium flap-like, compactly

scaled, tegula long and slender. Ground colour of

head and notum medium brown, pectus blackish

brown tufted. Forewing oblong-elongated, with costa

strongly bowed before apex, oblique termen and

smoothly convex tornus; ground colour medium to

pale brown, darkest in basal field, paler with some

minute white sprinkles in outer half; transverse lines

medium brown, subbasal indistinct, antemedial

very oblique from costa to anal margin, lined pale

419

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TeLNov, D. et al. (eds) 2021:

Figures 16-35. Habitus of Platyja (Mocrendes) species (42). 16 - P. (M.) lemur ([C. & R.] Felder, 1874), 3, Seram;

17 - P. (M.) lemurella sp. nov., 4, holotypus, Java, Trettes; 18 - <, ditto, Flores, Bea Nio; 19 - P. (M.) carpentariella

sp. nov., 4, holotypus, Australia, Queensland, Cooks Hut; 20 - P. (M.) cyanopasta (A. J. Turner, 1908), 4, New

Guinea, Nomnagihé; 21 - 4, ditto, Snow Mts.; 22 - <, ditto, topotypus, Australia, Kuranda; 23 - P. (M.) torsilinea

torsilinea (Guenée, 1852), 2, holotypus, Silhet; 24 - 9, ditto, Assam, Digboi; 25 - 9, ditto, Sumatra, Lebong Tandai;

26 - P. (M.) exviola Hampson, 1891, 9, India, Malabar; 27 - P. (M.) heterogenea sp. nov., 2, paratypus, Sulawesi,

Menado; 28 - 9, ditto, paratypus, Gunung Mogogonipa; 29 - P. (M.) torsilinea ciacula Swinhoe, 1893, °, topotypus,

Andamans; 30 - P. (M.) flavimacula Semper, 1901, 9, holotypus, Mindanao; 31 - P. (M.) lemurella sp. nov., 9,

paratypus, Java; 32 - Q, ditto, paratypus, Java, Neglirip; 33 - P. (M.) carpentariella sp. nov., 29, paratypus, Australia,

Queensland; 34 - P. (M.) cyanopasta (A. J. Turner, 1908), 2, Papua, Humboldt Bay; 35 - Q, ditto, topotypus, Australia,

Kuranda [scale bar = 1 cm].

420

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Ziti, A., DE Vos, R. & Epwarps, E. D.: The torsilinea species group of the genus Platyja Hubner, [1823] ...

pp. 413-430

Figures 36-50. Male genitalia of Platyja (Wocrendes) species (phalli removed). 36 - P. (M.) torsilinea torsilinea

(Guenée, 1852), Khasi; 37 - ditto, Assam, Cherrapunji; 38 - ditto, Hong Kong; 39 - ditto, Malaysia, Genting

[Sempah]; 40 - ditto, Brunei, Bukit Retak; 41 - P (M.) torsilinea ciacula Swinhoe, 1893, topotypus, Andaman; 42

- P. (M.) exviola Hampson, 1891, topotypus, India, Nilgiris; 43 - P (M.) heterogenea sp. nov., holotypus, Sulawesi,

Minahassa; 44 - P. (M.) sukabumensis sp. nov., holotypus, Java, Priangan; 45 - P. (M.) flavimacula Semper, 1901,

topotypus, Mindanao; 46 - P. (M.) lemur ([C. & R.] Felder, 1874), Seram; 47 - P. (M.) lemurella sp. nov., holotypus,

Java, Trettes; 48 - P. (M.) carpentariella sp. nov., holotypus, Australia, Queensland, Cooks Hut; 49 - P. (M.) cyanopasta

(A. J. Turner, 1908), New Guinea; 50 - ditto, topotypus, Australia, Kuranda [scale bar = 1 mm].

€ 421

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yellowish brown proximally, subparallel with section

of postmedial line below cubitus, orbicular stigma

large, round, reniform kidney-shaped, both darker

brown than ground colour, outlined pale yellowish

brown, pedunculate lobe beige, one to three faint

beige or white dots beyond reniform, albeit not

always present, postmedial lined pale yellowish or

white externally, distinctly edged white near costa;

preapical patch present, large and conspicuous,

dark chocolate brown, regularly rounded alongside

edge facing median field; veins lined chocolate

brown, except for 1A+2A on basal field, lined beige;

terminal line thin, medium brown, fringe beige

basally, medium brown distally. Hindwing rhomboid

Sub-pentagonal, greatly bowed costa, with

prominent apex and broad vannus, the wing margin

between which is smoothly concave; veins greatly

distorted and forming a large semi-hyaline yellowish

resonator, affecting almost whole anterior half of

wing, crossed by conspicuously ridged median vein;

remaining surface of wing middle brown, almost

pattern-less, with dark brown section of postmedial

line occurring only from below cubitus to before

anal margin, externally lined pinkish white and with

white sprinkles around; terminal line and fringe as

in forewing. Underside of wings medium brown,

semi-hyaline in correspondence of resonator. Fore-

and midleg conspicuously tufted blackish brown,

hindleg medium brown, with elongated crest of

scales. Abdomen slender, medium brown.

Male genitalia (Figs 43 & 57): Tegumen and

vinculum slender, V-shaped. Valva broad at base,

with greatly expanded, elongated obovate sacculus,

saccular lobe lying approximately at middle of valva,

subrectangular and feebly bilobed; costa expanded,

prolonged into thick, comparatively wide and short,

ventrally bent cucullus; ampulloid long and slender,

outwardly arched, distally clubbed, followed along

costa by short spine at base of cucullus. Uncus

long, slightly bent at base and feebly incrassate

distally before apical spine. Juxta stout and strongly

sclerotized, actually in shape of half of a vertically

bisected ‘carafe’. Phallus long, coecum thin and

relatively long, slightly curved, shaft beyond junction

with ductus ejaculatorius broadly arched, ending

into pointed tip; vesica with compact corpus slightly

scobinate posteriorly, three equally sized anterior

subconical lobes, and two sclerotized platelets,

subbasal and distal, respectively (Fig. 57).

Female (Figs 27-28): Fwl = 26 mm, x = 26

mm, N = 3; habitus as in nominotypical P. (M.)

torsilinea (Figs 23-25), with the exception of more

bowed forewing costa before apex, more oblique

antemedial and postmedial lines, larger and more

422

rounded dark brown preapical patch, dots beyond

reniform stigma, if present, not filled dark brown,

and generally unicolorous paler brown ground

colour. Underside of wings medium brown, with

pinkish white irroration on hindwing; faint pale

dotting on veins in correspondence of postmedial

lines.

Female genitalia (Fig. 67): Sternum A/ and

apparatus as in P. (M.) torsilinea (Figs 64-65),

with broader lobes of lodix not constricted at

middle, ductus bursae narrower and slightly funnel-

Shaped anteriorly, stronger sclerotized all through,

midventral sclerotization on A8 longer and wider,

and bursa more saccate with broad cervical sclerite

to the left.

Diagnostic remarks: This’ species _ is

unmistakable in the male (Figs 8-11) as it combines

the pattern features of male P. (M.) torsilinea with

the wing shape of male P. (M.) flavimacula Semper,

1901 (Figs 1-6, 13-15). The female instead

(Figs 27-28) shows the greatest similarity to that

of torsilinea from the nominotypical subspecies

of which (Figs 23-25) it can be distinguished by

the differences outlined in the description above.

As concerns P. (M.) torsilinea subsp. ciacula (Fig.

29), and partly also the fairly similar P. (M.) exviola

(Fig. 26), female heterogenea is larger, slate-

brown-coloured, with more conspicuous and darker

preapical patch of forewing, and with a less oblique

antemedial line of forewing. In the male genitalia,

the new species looks closest to P. (M.) flavimacula

(Figs 43 & 45), with which it shares the same

basic configuration of valvae and juxta. However, it

can easily be distinguished from the latter by the

narrower sacculus with smaller and more rugulose

saccular lobes, longer cuculli, more prominent

superior ‘beak’ of the juxta, longer and more evenly

curved distal section of the phallus, and larger lobes

of the vesica. The female genitalia of flavimacula

have not been studied, as the only female known

to us and whose identity was not in doubt is the

holotype. To distinguish the new species from its

close relative P. (M.) sukabumensis sp. nov. see the

description of this latter.

Distribution: So far known from Sulawesi.

Platyja (Mocrendes) sukabumensis sp. nov. (Figs

12,44 & 58)

Holotypus @ RMNH: [Nederlands Indié (= Indonesia)]

W. Java, Priangan, 19.1.1940, J. M. A. v. Groenendael

[leg. et coll.], slide RMNH.INS1108703.

Taxonomic remarks: Due to the scanty

material available, important differences in the

02-Jun-21 21:57:54

Zi, A., DE Vos, R. & Epwarps, E. D.: The torsilinea species group of the genus Platyja Hubner, [1823] ...

pp. 413-430

Figures 51-63. Phalli of Platyja (Mocrendes) species. 51 - P. (M.) torsilinea torsilinea (Guenée, 1852), same as

Fig. 37 (backslash points to non-everted apical lobelet); 52 - ditto, same as fig. 38 (two views); 53 - ditto, same as

Fig. 39 (two views); 54 - ditto, same as Fig. 40; 55 - P. (M.) torsilinea ciacula Swinhoe, 1893, same as Fig. 41 (two

views); 56 - P. (M.) exviola Hampson, 1891, same as Fig. 42 (two views); 57 - P. (M.) heterogenea sp. nov., Same as

Fig. 43; 58 - P. (M.) sukabumensis sp. nov., same as Fig. 44 (vesica partly everted); 59 - P. (M.) flavimacula Semper,

1901, same as Fig. 45; 60 - P. (M.) lemur ([C. & R.] Felder, 1874), same as Fig. 46; 61 - P. (M.) lemurella sp. nov.,

same as Fig. 47 (two views); 62 - P. (M.) cyanopasta (A. J. Turner, 1908), same as Fig. 49 (two views); 63 - P. (M.)

carpentariella sp. nov., same as Fig. 48 (vesica not everted) [scale bar = 1 mm].

@qg ke 423

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male genitalia shown by a singleton examined from

Java compared to populations of its Sulawesian

relative P (M.) heterogenea sp. nov., coupled

with notably sharper elongation of forewings and

allopatry (c. 1500 km across the Java sea from

nearest locality Known in South Sulawesi), leads

us to propose herein the erection of a new species

reluctantly.

Derivatio nominis: The species is named after

the city and district of Sukabumi in Parahyangan

(= Priangan) region of West Java. The name is a

feminine adjective agreeing in gender with the

generic name.

Male (Fig. 12): Fwl = 26 mm, N = 1; similar in all

respects to P. (M.) heterogenea sp. nov., with the

exception of the forewing which is more elongate

and more acutely tipped.

Male genitalia (Figs 44 & 58): Apparatus as in

P. (M.) heterogenea sp. nov., excepted for narrower

valva, more slender clavus, stout cucullus, straight

(aligned with remainder of valva) and apically blunt,

thicker and erect ampulloid, feeble hump in place

of costal spine, and superior ‘beak’ of juxta smaller.

Phallus shaft as in heterogenea [but vesica not

compared because it was only partially everted

during preparation].

Female: unknown.

Diagnostic remarks: As noted above, the

habitus of the holotype closely matches that of P. (M.)

heterogenea sp. nov. (Figs 8-11) but its forewings

are distinctly more apically elongated, so that the

termen is even more oblique than in the Wallacean

taxon. In the male genitalia, the new species can

easily be distinguished from heterogenea (Fig. 43)

by the straight, apically truncated cuculli (incurved

ventrally and apically rounded in heterogenea), the

thick erect ampulloids reaching in length the apex of

cuculli and forming a V-configuration with the latter

(thin and smoothly arched into U-configuration in

heterogenea), the different degree of development

between the juxta sections, the inferior plate being

longer, and the superior ‘beak’-like protrusion

smaller.

Distribution: So far known from West Java and

to be preliminarily regarded as a vicariant sister

species of Sulawesian P. (M.) heterogenea, the two

thus indicating a Javanese-Celebian connection

which excludes Borneo, from where neither of them

is recorded.

Platyja (Mocrendes) flavimacula Semper, 1901

(Figs 13-15, 30, 45 & 59)

Platyja flavimacula Semper, 1901. Die Schmetterlinge

der Philippinischen Inseln 2: 572, pl. 61, fig. 3. Locus

424

typicus: [Philippinen] Ost-Mindanao. Holotypus: 9°, by

monotypy, in SMF [examined].

Taxonomic remarks: A poorly known species

whose identity could only be solved after the

rediscovery of its type during the present work. Only

a few specimens appear to be known.

Diagnostic remarks: Male (Figs 13-15) (fwl

= 27-29 mm, x = 27.88, N = 4), with very narrow

elongated forewing devoid of distinct preapical

patch, rather with blurred black irroration ending

well before apex, crosslines distinctly crenulated,

ground colour brownish black irrorated with

ash grey; orbicular filled with grey and outlined

black, reniform deep black with indistinct edges,

pedunculate lobe with stem concolorous with

median field, only with globular end paler coloured,

dull beige-yellow; hindwing shaped as in male P.

(M.) heterogenea sp. nov., darker coloured and with

sinuous median ridged vein crossing the resonator.

Female (fwl = 27.8 mm, N = 1) (Fig. 30) patterned

as in male, with shorter and broader forewing and

hindwing normal, that is devoid of secondary sexual

modifications. Despite the remarkable differences

in pattern, in the male genitalia this species looks

closest to heterogenea (for differences see above

under this).

Distribution: Philippines (Luzon, Mindanao).

Platyja (Mocrendes) femur ([C. et R.] Felder, 1874)

(Figs 16, 46 & 60)

Cremnodes lemur ([C. & R.] Felder, 1874). In: Felder C.

et al., [1865]-1875, Reise der Osterreichischen Fregatte

Novara um die Erde in den Jahren 1857, 1858, 1859

unter den Befehlen des Commodore B. von Wullerstorf-

Urbair (Zoologischer Theil) 2 (2) Lepidoptera (Atlas):

Erklarung der Tafel CXIll, pl. 113, fig. 9. Locus typicus:

[in] Ins[ulis] Mollucc[is]. Holotypus: <, by likely monotypy,

in NHMUK (via L. W. Rothschild collection) [examined].

Taxonomic remarks: The holotype of Platyja

(Mocrendes) lemur is in very poor condition, with

worn and shrunken wings and even with unnaturally

darkened parts which seem to have been mouldy or

to have been fumigated. The unnatural narrowness

of the forewing and approximation in the drawing

of the pattern in the original illustration, indicate

that it was already damaged at the time of the

description. Such a poor illustration evidently

misled other authors to place in Cremnodes totally

unrelated species, including Snellen (1880) with

his Cremnodes macrocera. Nevertheless, surviving

details of the pattern, hindwing lobes and venation

show a full match to a male singleton from Seram

in the then Joicey collection, now in NHMUK, which

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Ziti, A., DE Vos, R. & Epwarps, E. D.: The torsilinea species group of the genus Platyja Hubner, [1823] ...

enables us to outline the features of this apparently

most rare species.

Diagnostic remarks: Male (fwl = 30 mm, N

= 1) with broader, less elongated forewing than

that of P. (M.) flavimacula (Figs 13-15) and with

a Straight, not markedly convex anal margin, albeit

with essentially similar forewing pattern to the

latter, although differing by the warm brown ground

colour, complete absence of any patch or darker

irroration before apex, presence of white or golden

dotting wherever the crosslines of both wings

(terminal line included) intersect with veins and a

well-defined reniform stigma. The pedunculate lobe

appears to be very slender, slightly hooked inwards,

filled with pale orange brown. In the hindwing, the

margin between apex and tornus is not concave but

protrudes at the middle, thus conferring to the whole

outline a lobed silhouette that recalls that of P. (M.)

cyanopasta (A. J. Turner, 1908), and the resonator

iS present but less strongly developed than in the

Philippine species, particularly below median vein.

In the male genitalia, the closest similarities are

with P. (M.) torsilinea, P. (M.) lemurella sp. nov., and

P. (M.) cyanopasta. From the first (Figs 36-41) it

can be distinguished by the basally broader valva,

that is also slightly angular along the ventral margin,

less curved cuculli and slightly more expanded and

more rugulose apical winglets of the juxta; from the

last (Figs 49-50) by the less asymmetrical saccular

lobes, more slender and more ventrally incurved

cuculli and less developed and widely opened

winglets of juxta; for differences from /emurella see

below under this species. Differences in the vesical

configuration with the three species are notable (cf.

Figs 51-55, 60-62). Female unknown.

Distribution: Seram, and likely in other islands

of the Moluccas.

Platyja (Mocrendes) lemurella sp. nov. (Figs 17-

18, 31-32, 47, 61 & 68)

Holotypus 6 NHMUK: [Indonesia] Efast}Java,

Trettes, 3000 [ft], May 1932, J. P A. Kalis [leg.],

NHMUK010918832, slide NHMUK010315429.

Paratypi 32 RMNH: 9, Gedanganvak [= Gedanganak],

20 m, 17.”II” [= February vel November].[19]27; 9,

Nglirip, 1936, Meyr. Walsh [leg.]; 2, [illegible, Javanese

site inferred from collector’s biography], Fr. A. Th. H.

Verbeek [leg.]; all via W. K. J. Roepke collection.

Additional material: 4 RMNH: Flores, Bea Nio,

16.X1.1952, J. M. A. v. Groenendael [leg. et coll.]; 4, sine

data [probably Java], W.K.J. Roepke coll.

Derivatio nominis: The species is named in

analogy to its bigger-sized relative, P (M.) lemur,

with a term of endearment based on its specific

pp. 413-430

name, adjectivized into feminine gender agreeing

with the generic name.

Male (Figs 17-18): Fwl = 22 mm, x = 22 mm, N =

2; habitus, pattern and colour fully corresponding to

P. (M.) lemur except for the smaller size, presence

of a faint preapical patch at forewing, slightly

darker than ground colour and with sinuous outer

edge, thicker and duller brown pedunculate lobe,

and much weaker expression of Secondary sexual

characters. This leads the new species to have

neither distorted veins nor loosely scaled resonator

on the hindwing, whose distal margin is furthermore

not distinctly lobed but almost regularly convex.

Underside of wings dark chocolate brown, with

straw-coloured dots on veins in correspondence of

postmedial lines. Fore- and midleg conspicuously

tufted blackish brown, hindleg medium brown, with

elongated crest of scales.

Male genitalia (Figs 47 & 61): Tegumen and

vinculum slender, V-shaped. Valva broadest at base,

slightly angular along ventral margin of saccular

part, sacculus moderately developed, projected

into conspicuous subtriangular lobe pointing

towards articulation between valva and tegumen,

cucullus long and slender, with ventrally incurved

tip, ampulloid slender and outwardly arched. Uncus

bent at very base, then straight, of uniform width,

bearing recurved hook apically. Juxta membranous

along midline, consisting of broad inferior plate in

Shape of narrow circle sector and two large apical

winglets. Phallus with fairly elongated and curved

coecum, straight middle section, and curved distal

third; vesica with compact saccate corpus projected

anteriorly into large subcylindrical lobe, densely

scobinate at apex, a smaller bilobed pedunculate

diverticulum with opposed lobes towards the shaft,

and some minor lobes (as in Fig. 61).

Female (Figs 31-32): Fwl = 19-22 mm, x =

20.67, N = 3; as in male, with filiform antenna,

broader wings and less oblique, more convex

forewing termen.

Female genitalia (Fig. 68): Sternum A7 and

apparatus as in P. (M.) torsilinea (Figs 64-65),

with lodix lobes more acute and not mesially

constricted, rather with angle at middle of inner

margin, ductus bursae narrower, well sclerotized

only in its posterior two thirds, emitting midventral

sclerotization onto A8 into long and narrow strip of

uniform width; cervical sclerite in shape of narrow

sinuous belt.

Diagnostic remarks: Closest to both P. (M.)

lemur (Fig. 16), of which it looks like a dwarf

version without conspicuous modifications in wing

Shape and venation, and P. (M.) carpentariella

oes

425

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Figures 64-70. Female genitalia of Platyja (Wocrendes) species (64-66 & 70: lodices removed and placed to left).

64 - P. (M.) torsilinea torsilinea (Guenée, 1852), Assam, Digboi; 65 - P. (M.) torsilinea ciacula Swinhoe, 1893,

topotypus, Andaman; 66 - P. (M.) exviola Hampson, 1891, Travancore; 67 - P. (M.) heterogenea sp. nov., paratypus,

Sulawesi, Menado; 68 - P. (M.) lemurella sp. nov., paratypus, Java, Nglirip; 69 - P. (M.) carpentariella sp. nov.,

paratypus, Australia, Queensland; 70 - P. (M.) cyanopasta (A. J. Turner, 1908), topotypus, Australia, Queensland,

Kuranda [scale bar = 1 mm].

sp. nov. described below. From the former, it can

also be distinguished by the wider and duller-

coloured pedunculate lobe and presence of a

preapical patch on forewing, that however is not

as Sharply defined and contrastingly dark against

the ground colour as in P. (M.) torsilinea and other

outstandingly patched species of the group. In the

male genitalia, the most evident differences from

its Moluccan relative occur in the less incrassate

uncus, the shape of the juxta with much broader

apical winglets and the configuration of vesica (cf.

Figs 60-61). To distinguish it from carpentariella

see below.

Distribution: So far known from Java to the

western Lesser Sunda Islands (Flores).

Platyja (Mocrendes) carpentariella sp. nov. (Figs

19, 33, 48, 63 & 69)

143.18E, Cooks Hut, Iron Ra. Q. GPS, 9 July 1998,

J. C. Cardale [leg.], Barcode of Life DNA voucher

specimen, S[a]mple ID: LOANIC-O6605, BOLD Proc. ID:

ANICJ608-10, Gen. prep. ANIC 18649.

426

Paratypus 19 ANIC: [Australia, Queensland] 11.41S

142.42E, 14 km ENE of Heathlands RF Qld, at light, 25-

28 Feb 1993, P. Zborowski [leg.], Barcode of Life DNA

voucher specimen, S[a]mple ID: LOANIC-O6606, BOLD

Proc. ID: ANICJ609-10, Gen. prep. ANIC 18650.

Derivatio nominis: The species’ name is a

term of endearment based on “Carpentaria”, an old

toponym for the Cape York Peninsula, from where

the species is known, adjectivized into feminine

gender agreeing with the generic name.

Male (Fig. 19): Fwl = 23.5 mm, N = 1; habitus

and pattern as in male P. (M.) lemurella sp. nov.

but with darker brownish black colour and without

pale spots at the junction between wing veins

and transverse lines, wings finely irrorated with

whitish grey sprinkles, forewing with transverse

lines thicker and black, antemedial more vertical,

postmedial distinctly crenulated in correspondence

of preapical patch and with pedunculate lobe

concolorous with median field and feebly projected

outwardly; hindwing with more distorted venation

and looser scaled areas configuring weak albeit still

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present resonator, and showing slightly different

trend of postmedial line whose widest pale lining,

orangeish brown, is external. Underside of wings

dark chocolate brown, with sparse pinkish white

irroration on basal and median areas of hindwing.

Fore- and midleg conspicuously tufted blackish

brown, hindleg medium brown, with elongated

crest of scales.

Male genitalia (Figs 48 & 63): General

configuration as in P. (M.) lemurella sp. nov. but

with cucullus straight and of almost uniform width

till apex, this blunt and slightly oblique, shorter and

stouter ampulloid, and juxta with sub-pentagonal

inferior plate and smaller apical winglets. Phallus

Shaft as in lemurella [vesica not studied as it was

not everted].

Female (Fig. 33): Fwl = 21.5 mm, N = 4;

habitus and pattern as in male, secondary sexual

characters excepted and with some blurred pale

dotting at veins, and similar also to female P. (M.)

lemurella sp. nov.; ground colour warm brown,

with sparse whitish grey irroration, especially in

basal field of forewing and submarginal fields of

both wings; transverse lines thick black, orangeish

brown-lined; conspicuous milky white filling affects

section between ante- and postmedial lines below

cubitus and, with a pink hue, whole pedunculate

lobe of forewing, white lining extending also

along anterior sections of both transverse lines.

Underside of wings chocolate brown, with diffuse

pinkish white irroration; postmedial lines visible

and distinctly crenulate, dark brown, lined pinkish

white externally.

Female genitalia (Fig. 69): Sternum A7 and

apparatus as in P. (M.) lemurella sp. nov. but with

lodix lobes broadly obtuse triangular with straight

inner margin and midventral strip on sternum A&

wider.

Diagnostic remarks: Evidently a close relative

of PR (M.) lemurella sp. nov., whose male (Figs

17-18) can easily be distinguished from that of

carpentariella by the brown colour, presence of pale

dotting where veins intersect transverse lines and

at termen, thinner black lines, forewing postmedial

smoother in preapical area, hindwing without

resonator and with postmedial line that is more

conspicuously lined pale internally, not externally. In

the male genitalia the most remarkable differences

of Jemurella with respect to carpentariella consist

of the slender, apically tapered and ventrally curved

cuculli compared to the stout straight ones of the

latter, the thinner and curved ampulloids and the

much larger apical winglets of the juxta. Females

of the two species are strikingly different in

pp. 413-430

appearance (cf. Figs 31-33), though it is unknown

whether the white markings seen in the unique

female known of carpentariella are a constant

feature of this species. In the pattern elements, the

unique female carpentariella known shows thicker,

more crenulate and darker black transverse lines

than its ally, this difference is most evident where

the postmedial line of forewing faces the preapical

area.

Distribution: Hitherto known from the Cape

York Peninsula in northern Queensland.

Platyja (Mocrendes) cyanopasta (A. J. Turner,

1908) (Figs 20-22, 34-35, 49-50, 62 & 70)

Ischyja cyanopasta A. J. Turner, 1908. Transactions and

Proceedings and Report of the royal Society of South

Australia 32: 64. Locus typicus: [Australia] N[orth]

Q[ueensland], Kuranda. Holotypus: 6, by original

designation, in ANIC (via Council for Scientific and

Industrial Research, Canberra) [examined].

= Plateja [sic] plagosa Rothschild, 1915. In

Rothschild W. & Durrant J. H., Lepidoptera of the British

Ornithologists’ Union and Wollaston expeditions in the

Snow Mountains, Southern Dutch New Guinea: 60.

Locus typicus: Snow Mountains, Southern Dutch New

Guinea [...] Base Camp; Oetakwa River, 3000 ft. Syntypi:

1¢, 19 in NHMUK (via L. W. Rothschild collection) [9

syntypus examined, 3 not traced].

Taxonomic remarks: Dissections _ of

specimens from New Guinea and _ Australia

revealed slight differences in the saccular lobes

which could however be simply ascribable to

infraspecific geographic variation, so that pending

upon a more extensive screening over a broader

sample from different geographic locations the

synonymy between Plateja [sic] plagosa and Ischyja

cyanopasta, originally proposed by Edwards (1996),

is here maintained. In the NHMUK collection there

is a watercolour of the series commissioned by G.

F. Hampson on valuable specimens preserved in

other collections for use in his “Catalogue of the

Lepidoptera Phalaenae in the British Museum”

that portrays a male of this species, labelled as

“Platyja trogopera Queensland Kuranda Turner”. It

is not known whether such painting antedates the

description by Turner (1908) or not, but trogoptera

must be regarded as an unavailable manuscript

name.

Diagnostic remarks: An unmistakable

species which, besides several unique features,

strangely enough combines a series of features

seen in other species of the group, as Summarised

here for the male sex (Figs 20-22): fwl = 26-31

mm, x = 28.97, N = 15; shiny appearance and

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Sharp forewing apex (torsilinea), Conspicuous

preapical patch (torsilinea, heterogenea, and

sukabumensis, albeit shorter, less elongated),

trend and waving of transverse lines of forewing

(flavimacula, lemur, lemurella, and carpentariella),

lobate outline of hindwing (/emur), strong resonator

(torsilinea, heterogenea, sukabumensis and

flavimacula), and trend of hindwing postmedial

(torsilinea, heterogenea, and sukabumensis).

Besides the broader wings and, obviously, absence

of male secondary sexual modifications, which lead

males of this species to show also the heaviest leg

tufting among its allies, the female (fwl = 27-31.5

mm, x = 29.50, N = 8) (Figs 34-35) retains the

same pattern features seen in the male. It is also

the species with the most regularly present and

conspicuous dots in median field of the forewing

beyond the reniform stigma. In the male genitalia,

the valval outline is remarkably similar to that of the

lemur-lemurella-carpentariella lineage, with cuculli

that are however more tapered than in P. (M.)

carpentariella sp. nov., but not as much as in P. (M.)

femur and P. (M.) lemurella sp. nov., and only feebly

curved ventrally (cf. Figs 46-50). Notable is the

great asymmetry in the saccular lobes, the left one

being more elongate and digitiform, while the right

one Is shorter and broader based. Differences have

been noted in the relative degree of development

of such lobes between dissected specimens from

New Guinea and Australia (Figs 49-50). The juxta

bears small apical winglets. The phallus vesica

has a unique configuration (Fig. 62). In the female

genitalia, the lobes of the lodix are stout, prominent

but constricted at the middle, with the apex rather

blunt; the ductus bursae sclerotized all through,

broadest anteriorly, and the sclerotized belt on

sternum A8 long and of uniform width (Fig. 70).

Distribution: New Guinea, Australia (northern

Queensland), New Britain.

Nomen dubium

Megacephalomana pilosum (Pagenstecher, 1888)

Megacephalon pilosum Pagenstecher, 1888. Jahrbucher

des Nassauischen Vereins fur Naturkunde 41: 156.

Locus typicus: Amboina. Holotypus: <, by monotypy, in

coll. Staudinger [not traced].

Taxonomic remarks: Thisspecies was implicitly

transferred to Megacephalomana by Strand (1943)

following the replacement of Megacephalon

Saalmuller, 1880 (preoccupied by Temminck,

1844 [Aves]). The absence of an illustration in

the original description by Pagenstecher (1888)

428

and unsuccessful attempts to locate the holotype

either in the collection Staudinger (Museum

fur Naturkunde, Berlin, Germany) or the coll.

Pagenstecher (Museum Wiesbaden, Germany)

compel us to provisionally treat this nominal taxon

as of doubtful identity. We mention it here as it

is formally a member of the Platyja group and its

provenance from the Moluccas (Ambon) leaves

open the possibility that it is either conspecific or

related to one of the species treated in the present

work. Unfortunately, the original description is

unclear because of several conflicting statements.

On one hand, Pagenstecher (1888) compares his

pilosum to the Madagascan Megacephalomana

stygium (Saalmuller, 1881) (as Megacephalon),

which is actually a very differently looking species

from Megacephalomana_ rivulosum (Saalmiuller,

1880), which superficially resembles the

Moluccan P. (M.) lemur. On the other, he recalls its

Subquadrate hindwing which would seem a feature

of some Mocrendes species but he mentions that

Its base is shiny translucent, not the anterior half

where a resonator is placed. He records also a very

small size (wingspan 42 mm) for any Wallacean

species with a resonator and distribution of colours

and pattern elements which does not appear to

match any known species of Platyja. The chestnut

colour would seem compatible with the much

bigger, currently described P (M.) heterogenea

Sp. nov. (well over 50 mm), but pilosum is said to

have pectinated antennae, a straight transverse

line in the basal field of the forewing, to be much

darker beyond this, with a very short termen and

metatibiae thickly clothed with blackish scales, and

no dark preapical patch is mentioned. Interestingly,

Pagenstecher’s description looks partly compatible

with males of some species of the genus Crithote

Walker, 1864, but conclusive evidence is still

lacking.

Discussion

Subgenus Mocrendes of Platyja currently

consists of nine species, four of which are

described herein, distributed in the Indo-Australian

Region from the Indian subcontinent eastwards to

the Bismarck Archipelago (New Britain). Peripheral

areas or definite groups of islands often host

endemic taxa, such as P. (M.) exviola in the SW

Ghats, P. (M.) flavimacula in the Philippines and P.

(M.) carpentariella in northern Queensland.

Despite the often-striking differences in

external appearance, members of the group

02-Jun-21 21:57:59

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Zit, A., DE Vos, R. & Epwarps, E. D.: The torsilinea species group of the genus Platyja Hubner, [1823] ...

show quite limited male genital variation, with the

most western species (exviola) sporting possibly

the most extreme development of valval shape.

Broadly speaking, configuration of the valva and

juxta allows us to recognise two distinct groups,

a broadly distributed one with elongated cuculli

and broad-based juxta bearing paired small apical

winglets, and another group with shorter, stouter

cuculli with a small process (Spine or hump along

costa at their base), and the juxta in the shape of a

bisected ‘carafe’ with a prominent superior ‘beak’.

The second group is settled in the central area of

the range of the subgenus, with P. (M.) flavimacula

(Philippines), P. (M.) heterogenea (Sulawesi) and P.

(M.) sukabumensis (Java). Vicariance events have

evidently led to formation of allopatric species

pairs, such as aforementioned heterogenea and

sukabumensis or P. (M.) lemurella (Java and W

Lesser Sundas) and P. (M.) carpentariella (Cape

York Peninsula). The latter thus clearly represents

a south-eastern lineage which is further closely

related to the Moluccan P. (M.) lemur, which in

turn shares with eastern P (M.) cyanopasta an

essentially similar hindwing configuration, with a

protruding lobe along the wing margin between the

apex and tornus.

Further research will be necessary to

detail the phylogenetical relationships among

species of the group, a task which will benefit

from resolution of other sympatrically occurring

groups of Platyja, and to assess whether broadly

distributed, geographically varying taxa (e.g.,

torsilinea, cyanopasta) do actually comprise more

evolutionary significant entities.

List of species

Platyja (Mocrendes) exviola Hampson, 1891

Platyja (Mocrendes) torsilinea (Guenée, 1852)

Platyja (Mocrendes) torsilinea torsilinea (Guenée,

1852)

Platyja (Mocrendes) torsilinea ciacula Swinhoe,

1893 stat. nov.

Platyja (Mocrendes) lemur ([C. et R.] Felder, 1874)

Platyja (Mocrendes) lemurella sp. nov.

Platyja (Mocrendes) carpentariella sp. nov.

Platyja (Mocrendes) cyanopasta (A. J. Turner, 1908)

Platyja (Mocrendes) flavimacula Semper, 1901

Platyja (Mocrendes) heterogenea sp. nov.

Platyja (Mocrendes) sukabumensis sp. nov.

LS ON ON ms am

pp. 413-430

Acknowledgements

The authors are deeply indebted for the

realisation of this work to Hans Banziger (Chiang

Mai University, Thailand) for providing original

information and discussion on _ functional

morphology of proboscides, Wolfgang Nassig and

Massimo Terragni for their incessant assistance

during AZ’s visitto SMF, Fritz Geller-Grimm for having

kindly allowed AZ to examine the Pagenstecher

collection at Museum Wiesbaden, Théo Léger,

Viola Richter and Konrad Ebert (Museum fur

Naturkunde, Berlin, Germany) for their extensive

search for the type of Megacephalon pilosum,

Bernard Landry (MHNG) for supplying collection

data, Mark Sterling (London, United Kingdom) for

providing important comparison material from

Hong Kong, and You Ning Su for detailed photos

of specimens and preparations at ANIC. Finally,

Bernard Landry, and Martin R. Honey (London,

United Kingdom) are gratefully acknowledged for

their thorough reviewing of the manuscript which

led to its substantial improvement.

This research was largely possible thanks to a

NHMUK grant to AZ (DIF 2019: SCR13057).

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général des Lépidoptéres 7. Noctuélites 3. Roret,

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Hampson G. F. 1894. The fauna of British India, including

Ceylon and Burma. Moths 2. Taylor and Francis,

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Holloway J. D. 2005. The Moths of Borneo [Parts 15 &

16]: Family Noctuidae, subfamily Catocalinae. -

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Holloway J. D. 2011. The Moths of Borneo [Part

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Zygaenidae; revised and annotated checklist. - The

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Pierce F. N. 1909. The genitalia of the group Noctuidae of

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species. - Journal of the natural History Society of

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Received: 02.iv.2021

Accepted: 15.iv.2021

02-Jun-21 21:58:00

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BIENKOWSKI, LARA S., STEPHAN, ROBIN S., KIRSCHEY, TOM & SARYANTHI, Ria: Population density of Tarsius supriatnai ...

pp. 431-442

Population density of Tarsius supriatnai along a forest

degradation gradient in Popayato-Paguat landscape

(Gorontalo Province, Sulawesi)

urn:lsid:zoobank.org:pub:5F18F501-F6B8-4940-82 7C-A69F /BC3EDD9

LARA SuiRIN BiENKOWSKI *, RoBIN SIMON STEPHAN *, Tom KIRSCHEY * & RIA SARYANTHI °

1 - Eberswalde University for Sustainable Development, Schicklerstrasse 5, D-16255,

Eberswalde, Germany; lara.bienkowski@hnee.de and robin.stephan@hnee.de

2 —- NABU Headquarters, International Department, Charitéstrasse 3, D-10117, Berlin, Germany;

tom.kirschey@nabu.de (ORCID: O0O00-0002-66 /6-4981)

3 - Burung Indonesia, Bogor, Jalan Dadali 32, RT.03/ RW.05, Tanah Sareal, 16161, Kota Bogor,

Jawa Barat, Indonesia; rsyanthi@burung.org

Abstract: Jatna's Tarsier Tarsius supriatnai Shekelle, Groves, Maryanto et Mittermeier, 2017 is a species of primitive

primates described very recently from the western part of the Gorontalo Province in North Sulawesi (Shekelle et al.

2017). The main aim of the present study is to identify correlations between forest degradation and conversion and

population densities of Jatna’s Tarsier in a typical forest edge to agricultural land transition area in Popayato Paguat

landscape and provide ecological data about the species and its spatial distribution. The results show that Jarsius

supriatnai is not entirely restricted to the secondary forest. As long as small particular remnants of natural vegetation

remain in the landscape, tarsiers are found be able to persist, but their population densities are considerably lower.

Average population densities on agricultural-agroforest area (1.22 individuals/ha) differ significantly from those

within the secondary rainforest (5.37 individuals/ha). Moreover, average population densities at the study plots

undergoing forest conversion (1.05 individuals/ha) differ significantly from those unaffected (5.65 individuals/ha)

by rainforest degradation (4.07 individuals/ha). Regarding to the IUCN Red List assessment guidelines, we confirm

the criterion "VU Vulnerable" for Tarsius supriatnai based on the results of the present study.

Key words: Jarsius supriatnai, population biology, Popayato Paguat landscape, Sulawesi.

Introduction

Deforestation and rainforest degradation rates

in Indonesia are still the highest in Southeast Asia

and even globally rank among the top three among

tropical countries. Deforestation, forest degradation

and conversion of natural forests into agricultural

land are the main terrestrial threats to many

species in the global biodiversity hotspot Wallacea,

including its most enigmatic island of Sulawesi -

a place of extraordinary endemism in mammals.

In Popayato Paguat landscape, a designated Key

Biodiversity Area (KBA), the non-governmental

organization Burung Indonesia, with support of

the NABU and KfW (German Development Bank)

aim to improve forest management and maintain

landscape connectivity between Nantu Forest

Reserve and Panua Protected Area. Our research

aims to test the indicator function of an endemic

species towards forest disturbances along a

gradient of forest use and conversion in this

landscape and provide some additional biological

data on this elusive and insufficiently studied

small primate species. Tarsiers are remarkable,

fully insectivorous primates. Originally thought to

be only a single species (7Jarsius tarsier (Erxleben,

1777), also widely known under the synonymized

name Tarsius spectrum (Pallas, 1778)), tarsiers

from Sulawesi and surrounding islands represent a

complex comprised of 12 species as is considered

today. Especially in recent years tarsier taxonomy

has been improved by clarification of the taxonomic

status of taxa from the northern arm of Sulawesi

Island, including the recent descriptions of Tarsius

spectrumgurskyae Shekelle, Groves, Maryanto et

Mittermeier, 2017 and Tarsius supriatnai Shekelle,

Groves, Maryanto et Mittermeier, 2017 (Shekelle

et al. 2017) (Fig. 1). While the taxonomic status of

South and South-Central Sulawesi populations is

still unclear, with the description of Tarsius niemitzi

431

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% 7

Figure 1. Jatna's Tarsier Tarsius supriatnai Shekelle, Groves, Maryanto et Mittermeier 2017, specimen from the

plot No 108 (photograph: Tom Kirschey, 17.i.2020).

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BIENKOWSKI, LARA S., STEPHAN, ROBIN S., KIRSCHEY, TOM & SARYANTHI, Ria: Population density of Tarsius supriatnai ...

Shekelle, Groves, Maryanto, Mittermeier, Salim et

Springer, 2019 from the Togean Islands in the Gulf

of Tomini the unclear situation has been solved

for the northern part of Sulawesi and adjacent

islands (Shekelle et al. 2019). Tarsius supriatnai

is distributed West of the Gorontalo (geotectonic)

fault between the Popalo Bay in the North and

Gorontalo City (including Limboto) in the South,

whereas populations East of the Gorontalo fault

are represented by Tarsius spectrumgurskyae

(Shekelle et al. 2017).

Material and methods

The field research was conducted for a two

months period in December 2017 and January

2018 around a small village next to the rainforest

edge in the Popayato-Paguat landscape, Gorontalo

province, Sulawesi. During this period on a daily

basis excursions along a transect between the

village and the forest area have been conducted

during the maximum activity period of the tarsiers.

The forest, agricultural area and different transition

stages have been classified in order to verify

the habitat suitability for tarsiers. For the study,

29 study plots (all subject to human-induced

disturbance), have been arrayed along river bank

oriented transects (Fig. 2). Fourteen of these trial

plots were set up within a tropical old-secondary

lowland rainforest on mineral soil, which served as

reference site by providing ideal habitat conditions.

Another 15 study plots were delineated on diversely

used agricultural land, which has undergone

severe forest degradation and conversion. The data

collection was conducted by recording the duet

vocalizations at dawn and locating the direction of

sound. Defined cultivation types, highest reached

vegetation layer, density of riparian vegetation,

river bank connecting vegetation (closed tree

canopy across rivers), forest cover loss since 2001,

occurrence of secondary forest fragments, height

differences within each study plot and the potential

sleeping trees were considered in order to answer

the goals of this study.

Investigation area

Popayato Paguat landscape is an area in the

West of the Gorontalo Province in the regencies of

Pohuwato and Boalemo in the Northern part of the

island of Sulawesi. The area has been classified as

a Key Biodiversity area (KBA) (Burung Indonesia

2014; Wood et al. 2015). The landscape includes

pp. 431-442

the lowlands and southern slopes of the mountain

ridge between 48 and 980 m altitude and edges of

villages as well as agricultural land. Some authors

refer to this landscape as the Polahi-Marissa Forest

Complex (Cannon et al. 2007). The settlement today

known as Makarti Jaya village area (Fig. 3) was

established in 1994, when the first families were

brought to our study area under the Indonesian

transmigration settlement program. The settlement

was then known under the name “Unit Pemukiman

Transmigrasi (UPT) Marisa V”. Because of the

Figure 2. Arrangement of trial plots (Scale 1 : 35 OOO,

QGIS 3.4, OSM Standard).

433

02-Jun-21 21:58:05

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

-- "Makarti Jaya

es Village !

Figure 3. Satellite image showing the forest coverage around the study site in Makarti Jaya village, Sulawesi.

ot = 3 Forest fragments,

a & " 5 Pala nantu trees

fo F: fe oh

aa — v

eet peli : J 3

<etttene Oe SG :

% Banana trees,

T coconut palm,

bamboo,

oil palm

Cocoa, shrubs, orange

1,5

0 Corn, grass, rice,

bare land

Figure 4. Agricultural crops and agroforestry composition in Makarti Jaya village, Sulawesi, based on a secondary

forest stratification model.

434

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BIENKOWSKI, LARA S., STEPHAN, ROBIN S., KIRSCHEY, TOM & SARYANTHI, Ria: Population density of Tarsius supriatnai ...

compatibility of cocoa trees with soil and climate in

this region, their dowry provided by the state were

cocoa seedlings to establish a livelinood. Since

2002 the former UPT counts as official village

under the name Makarti Jaya (Makarti = work, Jaya

= success; Makarti Jaya = success through work).

The cultivation of cocoa has proved successful for

the villagers. The community today has several

cocoa nurseries and most of the livelinood of the

village is generated by farming cocoa and corn. The

forest itself is classified as production forest ("Hutan

Produksi") and comprises of a secondary forest as

a result of natural succession after this area was

logged out more than three decades ago (Fig. 4). Itis

a seasonally dry lowland to mid-montane Fagaceae-

Myrtaceae forest with Lithocarpus spp., Ficus spp.,

and Calamus spp. When activities to establish an

Ecosystem Restoration Concession (ERC) in the

Production Forest block have been started, Makarti

Jaya was selected to be a key village in a forest

management program by Burung Indonesia and its

partners (Kirschey & Schell 2016).

Results

The numbers of tarsiers per trial plot in the

reference area were higher than the numbers on

the study plots within the agricultural land. With a

total number of 238 individuals on 44.33 ha in the

pp. 431-442

secondary forest, there is an average density of 5.37

Jatna’s Tarsier per ha and 17 tarsiers per trial plot.

On the research plots within the cultivated land, 58

tarsiers were detected on an area of 47.5 ha. This

indicates an average density of 1.22 Jatna’s Tarsier

per ha and 3.8/7 tarsiers for each trial plot. In the

reference area tarsier numbers vary between six to

30 individuals per plot, whereas the numbers in the

cultivated land show a lower variance with counts

between three and nine individuals.

Inside the agricultural used land 15 cultivation

types could be identified. In addition, there are four

uncultivated vegetation types, which fall under the

term ‘cultivation type’ for this research. These are

bare land, shrub land, forest fragments and river

bank vegetation. The only monoculture where

tarsiers have been found was cocoa (n = 6) and the

only mixtures where tarsier have been observed

were a mixture of cocoa and pala nantu (n = 1) and

a mixture of cocoa and corn (n = 3). However, most

tarsiers were found in the cultivation types which

are currently not in agricultural use. Hereby, most

tarsiers have been found in forest fragments (n =

21), followed by river bank vegetation (n = 20) and

Shrub land (n = 6).

Riverside vegetation has been classified into

four different categories, ranging from O to 3 (Table

1). The parameter contains information about the

intensity of riparian vegetation along the river and

its horizontal connections.

Table 1. Classes of riverside vegetation in Popayatpo Paguat area.

eo No river bank vegetation at all

Poor river bank vegetation, not connected

Moderate river bank vegetation, partly connected

Rampant river bank vegetation, well connected

The conducted generalized linear module

(GLM) shows that a significant relation exists for the

occurrence of Tarsius supriatnail (p-value of 2./76e-

15). Same is true when considering the agricultural

used plots only (p-value of 0.0019).

The evaluation for forest degradation and

conversion was done in accordance with the degree

of tree cover loss for each trial plot (Hansen et al.

2013). Here, the thresholds of forest degradation

(<90%) and forest conversion (>90%) were

consulted according to their definition by the FAO

(FAO 2011). The outcome shows that the tarsier

occurrence on the plots that have been affected by

forest conversion vary significantly from those which

are unaffected or experienced forest degradation

(p-value of 2e-16).

The area where tarsiers have been detected

and where actual cultivation and forest conversion

occur included only plantations on which cocoa

trees represented the most dominant crop. Tarsius

supriatnai is able to adapt towards changing habitat

conditions, including even forest conversion, to a

certain extent. However, it was found that Tarsius

supriatnai is restricted to areas providing dense

vegetation structures like bamboo thickets or

rattan/duruh palm stands (Fig. 5). Using this

presence - absence - abundance data showed that

Tarsius supriatnai has a limited indicating function

435

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TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

Figure 5. Bamboo as a crucial habitat structure for Tarsius supriatnai Shekelle, Groves, Maryanto et Mittermeier

2017 (photograph: Tom Kirschey, 1.7.i.2020).

to forest degradation or conversion, although

population densities decrease significantly with

forest conversion. However, Tarsius supriatnai could

contribute as indicator for (Sustainable) agroforest

practices including cocoa cultivation that allow

well-connected, dense vegetation structures.

Discussion

Already prior to its formal scientific description,

Tarsius Supriatnai was studied and referred to as

the "Gorontalo form“ of Spectral Tarsier T. tarsier

(MacKinnon & MacKinnon 1980; Gursky et al.

2008) or the "Libuo form“ (Shekelle et al. 1997;

Shekelle 2003; 2008). The terra typica of Tarsius

Supriatnai is Sulawesi, Nantu Forest Reserve,

adjoining Popayato Paguat landscape in the

Northeast (Shekelle et a/. 2017). Its characteristic

duet call was described as a ~2 to 5-note female

phrase accompanied by male calls - a distinct

character different from other representatives of

the genus (Fig. 6). Population ecology considered

similar as in TJarsius spectrumgurskyae (e.g.,

Gursky 2007). Some aspects like site fidelity may be

similar in other Tarsius Storr, 1780 species, but our

436

investigations allow only preliminary conclusions.

Gursky (2007) reported 33 sleeping trees within

an area of 100 hectares in Tangkoko Dua Saudara

Nature Reserve, North Sulawesi. Twenty trees were

still used by Tarsius spectrumgurskyae five years

later, indicating suitable microhabitats in a dynamic

landscape are traditionally used by tarsiers over

long time periods.

Our results demonstrate no significant

differences between unaffected and degraded

plots. The density of riparian vegetation showed

a significant relation towards the occurrence of

Tarsius Supriatnai, while the presence of closed

tree canopy across the river banks did not correlate

with projected population numbers. Out of the 19

identified habitat conversion types, tarsiers were

detected in only six of those, whereas the three

types showing the highest tarsier occurrence

consisted of remnants of natural vegetation.

The results of the present study also

demonstrated that the population densities of

Tarsius Supriatnai in the agriculturally used land

differ significantly from those inside the secondary

rainforest. With a value of 5.37 individuals per ha

within the rainforest, the population density is 4.4

times higher as inside the cultivated/converted land

02-Jun-21 21:58:11

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BIENKOWSKI, LARA S., STEPHAN, ROBIN S., KIRSCHEY, TOM & SARYANTHI, Ria: Population density of Tarsius supriatnai ...

pp. 431-442

at " hy , "y

i Na

. 1b | { ) baal

A se iW } ‘ :

iy ~~]

hah

Figure 6. Tarsius supriatnai Shekelle, Groves, Maryanto et Mittermeier 2017 spectrograms (created with Praat

version 6.0.30): A - A typical duet call; B - A separate female call.

(1.22 tarsiers per ha). Compared to other studies

on population densities of species belonging to

the genus Tarsius, this value is comparatively high

(Merker 2003; Gursky 2007; Saroyo et al. 2017).

This may be due to the proximity of open water

bodies, which support rich populations of insects,

and because home ranges are not used linearly.

Slightly disturbed forest, as it is the case in the study

area, is the habitat where tarsier densities were

expected to be the highest. According to Gursky

(2007), the density of Tarsius soectrumgurskyae (a

species very similar to 7. supriatnal) inthe secondary

forest is even more than twice as high, compared

to their actual natural habitat, the undisturbed

primary forest. This can be explained by increased

light incidence in secondary forests, causing shrub

layer to develop stronger, and thus amenable

sleeping sites to the benefit of tarsier populations.

This demonstrates that forest disturbances may

promote tarsier densities to a certain degree. Due

to the fact that the secondary rainforest provides

dense vegetation, aS well as a high number of

suitable sleeping trees, tarsiers might not have the

need to travel far for their nightly forage activities,

which would mean that they require smaller home

range sizes. This would confirm the observations

by Merker (2003) and Merker et al. (2005), who

found out that Jarsius dentatus Miller et Hollister,

1921 adapts its ranging behaviour according to the

degree of habitat disturbance.

Analysis of forest disturbances shows that

tarsier densities significantly respond to forest

conversion but not to forest degradation. The

average population density within the area of

conversion (1.05 individuals/ha) is significantly

smaller than the density of the other groups (5.65

and 4.07 individuals/ha), whereas the unaffected

forest land has the highest population density.

This stepwise decrease shows similarities with

research on Tarsius dentatus by Merker (2003),

who stated that population densities are lower

in more disturbed habitats, and that the tarsiers

do not seem to substantially distinguish between

unaffected and slightly disturbed habitats. The

reason for the comparatively low population density

within conversion plots could be that these rarely

provide the vegetation structures as required by

tarsiers. It follows that the habitat must at least

provide some vegetation which can be used for

foraging and as sleeping sites. This corresponds

well with the research by Leksono et al. (1997) and

Yustian et al. (2008). Moreover, vegetation height

might matter as well. High and dense tree stands

provide more shade during the day when tarsiers

are sleeping and prevent the inner space from

overheating. This kind of microclimate could also be

a reason for the high population density within the

forested land, compared with the agricultural land.

437

02-Jun-21 21:58:11

Book4.indd 438

TeLnov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

In conclusion, it can be said that forest degradation

and also forest conversion do not amount to total

habitat loss for this species, but create new habitat

types which are changed to an extent that tarsiers

occur in significantly lower numbers. This implies

that forest conversion does automatically entail

habitat degradation for Tarsius supriatnai.

Considering the agricultural used land, only in

six out of 19 identified types of cultivation tarsiers

have been detected at all. Out of these, three of

the types do not actually count as cultivated land,

while showing the highest numbers of tarsiers.

Specifically, secondary rainforest fragments show

the highest number of tarsiers (n = 21), followed

by river bank vegetation (n = 20) and shrub land (n

= 6). The vegetation types under cultivation were a

cocoa monoculture (n = 6), a mixture of cocoa and

corn (n= 3) anda mixture of cocoa and nutmeg trees

(pala nantu) (n = 1). Since it has been observed

that all three cultivation types include cocoa as the

most dominant crop, this suggests that cocoa is,

compared to the other cultivation types within the

agricultural land, a crop which Tarsius supriatnal

does use (Fig. 7). However, there are three more

cultivation types which include cocoa but did also

not show tarsiers.

Moreover, it iS important to stress that

compared to the total area of cocoa plantations, the

identified number of tarsiers is still relatively low.

Additionally, tarsiers have never been observed to

actually sleep within these plantations. Whenever

tarsiers have been detected in cocoa plantations,

a Suitable sleeping location was close-by. In these

cases, observed sleeping structures were dense

thickets inside forest fragments, bamboo bushes,

rattan and salak palms. This circumstance confirms

the observations by Merker & Yustian (2008) and

Leksono et al. (1997) on Tarsius dentatus and

other Tarsius spp. showing that a limiting factor

for tarsier occurrence Is the availability of suitable

sleeping sites. It seems that the combination of

cocoa trees (monoculture and mixture) and any

dense thicket for resting, does provide sufficient

habitat conditions for tarsiers to survive. In these

cases, the question remains whether tarsiers, even

though they can survive in this habitat, are able

to reproduce in numbers sufficient for long-term

species survival (Fig. 8).

The results demonstrate that particular

riverside vegetation features, especially dense

structures containing shrubs as well as trees

of various heights and widths play a key role in

tarsiers’ habitat e.g. by provision of sleeping trees.

This could also be one reason why the intensity of

———

ty y

438

riverside vegetation correlates significantly with

tarsier occurrence for both the totality of plots and

those used agriculturally only (Fig. 7). Riverside

vegetation is providing sufficient shade and some

kind of microclimate. Cooling effects provided by

forests due to evapotranspiration and a closed

canopy is reduced to a higher extent on agricultural

land. This could also be a reason why most tarsiers

have been observed within forest fragments,

shrubland and dense riverside vegetation.

Concludingly, it can be said that TJarsius

Supriatnai can persist within agricultural land and

does use certain cultivation types and discriminate

others. The primates were mostly found on

uncultivated land, but were also identified on

plantations where cocoa is the most dominant

crop. All tarsier-occurrence sites share a dense

vegetation structure or close proximity to shrub-

like vegetation or bamboo thickets. Moreover, the

intensity of river bank vegetation seems to be an

important factor conditioning tarsier occurrence,

as well as left over secondary forest fragments

which have been spared from conversion.

Since it was described in 2017, Tarsius

Supriatnai has been evaluated for IUCN Red List

very recently (Shekelle 2020) and included in the

category “VU Vulnerable”. This corresponds with

earlier informal assessments proposed by Gursky

et al. (2008), but not with “DD Data Deficient“ as of

Supriatna (2019).

With the present study we aim to contribute to

extend the knowledge about this elusive species.

With an Extent of Occurrence (EOO) of about 11

000 km? and a remaining habitat (potential Area

of Occupancy, AOO) of less than 5 OOO km? area

limitations are not as prominent as those for Tarsius

sangirensis Meyer, 1897 (limited to Sangir Island),

T. tumpara Shekelle, Groves, Merker et Supriatna,

2008 (limited to Siau Island), T. pelengensis Sody,

1949 (limited to Peleng), 7. niemitzi Shekelle,

Groves, Maryanto, Mittermeier, Salim et Springer,

2019 (limited to the Togians) or 7. tarsier sensu

Stricto (limited to Selayar). Habitat loss due to

deforestation is an ongoing threat on Sulawesi,

which progresses at an alarming rate (Cannon et

al. 2007; Supriatna et al. 2020). In Taluditi area of

North Sulawesi, deforestation rates increased by

8.6% since year 2000 according to Global Forest

Watch (2021) (Fig. 9). Human population increased

by 22.4% in Gorontalo Province and by 19.4% in

Central Sulawesi Province during 2000-2010,

indicating an increasing pressure to the ecosystems

(Burung Indonesia 2014).

With continuous habitat loss, decline in its

02-Jun-21 21:58:11

BIENKOWSKI, LARA S., STEPHAN, ROBIN S., KIRSCHEY, TOM & SARYANTHI, Ria: Population density of Tarsius supriatnai ...

pp. 431-442

swiss, Meer ide op SS

ih 7 the, ) Ae -

| ee

ME Cocoa [I Rice ME oil Palm ME Bare Land

[SS Coconut [SS Rumput Gajah HM Banana GS Bamboo

I) corn (Mix Cocoa Coconut WE Mix Cocoa Banana WY) Shrub Land

[2 Mix GocoaCom =) Mix Cooma Gorn Coconut (Fa Mix BananaOrange $l Forest fragment

Figure 7. Cultivation types in the agricultural-agroforestry area near Makarti Jaya village, Sulawesi, with tarsier

occupation [not to scale].

@q ke 439

Book4.indd 439 02-Jun-21 21:58:12

TeLNov, D. et al. (eds) 2021: Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, IV

15 20 25 30

No, of tarsiers (n) per plot

Figure 8. Habitat capacity and tarsier densities in a comparison of 29 trial plots.

Tree cover

2010

Tree cover loss

2001-2018

a Tree cover gain

2001-2012

Figure 9. Recent forest degradation and deforestation development in Popayato Pauat landscape

in Makarti Jaya area, Sulawesi (Source: Global Forest Watch 2021).

sac es V ®& “<a”

Book4.indd 440 02-Jun-21 21:58:13

Book4.indd 441

BIENKOWSKI, LARA S., STEPHAN, ROBIN S., KIRSCHEY, TOM & SARYANTHI, Ria: Population density of Tarsius supriatnai ...

extend and quality and other not ceased drivers, we

independently support the proposed assessment

of Tarsius supriatnai as “VU Vulnerable” as of

Shekelle (2020) made with accordance with the

Guidelines for using the IUCN Red List Categories

and Criteria version 14 (2019).

Acknowledgements

We like to thank Hans-Christian Mittag and

the NABU International Foundation for financial

support of the field survey for the first two authors

as part of their bachelor thesis. Prof. Dr. Siegfried

Rieger and the third author jointly supervised the

thesis. Burung Indonesia Gorontalo Programme

facilitated data collection in Popayato Paguat

landscape in close collaboration with Prof. Evi

Hulukati from Universitas Negeri Gorontalo (UNG).

We also want to thank the Ministry of Research

Technology and Higher Education (RISTEK) of

Indonesia for issuance the Research Permit No

416/SIP/FRP/E5/Dit.KI/XI/2017; 417/SIP/FRP/

E5/Dit.KI/XI/2017). Moreover, field research was

Supported by Makarti Jaya farmers Anmad Fauzan

and Djoko Danang Santoso (Indonesia), who

attended the whole data collection and shared their

knowledge on the local agroforest ecosystem. Tom

Kirschey and Ria Saryanthi like to thank Prof. Jatna

Supriatna, Myron Shekelle, Hanom Bashari, and

Andrea Schell for valuable discussions on tarsier

taxonomy and ecology.

On behalf of NABU and Burung Indonesia,

third and fourth author are grateful to KfW German

Development Bank for supporting sustainable

forest management in Popayato Paguat landscape

with help of the project grant “Nature conservation

concessions to protect tropical rainforest in

Indonesia” by the International Climate Initiative

(IKI) of the German Federal Ministry for the

Environment, Nature Conservation and Nuclear

Safety (BMU).

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Book4.indd 443

Index to new taxa described in this volume

two new genera

one new subgenus

61 new species

amnicola, Litoria

bacchusi, Plateros

biaka, Taheitia

bicolorinotus, Sulabanus

bifurca, Taheitia

brevispina, Phrictaetypus

brunneifasciatus, Sulabanus

caelestis, Endelus (Hexagonodelus)

camarinensis, Lalahonia

camarinensis, Manilla

candalagaensis, Manilla

candalagensis, Baeocera

capitatus, Perignamptus

carpentariella, Platyja (Mocrendes)

cenderawasih, Palaina (s. |.)

cyclops, Tanycricos

doberaiensis, Plateros

evanescens, Palaina (s. |.)

gebeensis, Taheitia

gigantea, Taheitia

glabra, Baeocera

gracilipenne, Ceresium

halmaheraensis, Plateros

heterogenea, Platyja (Mocrendes)

Hexagonodelus subgen. nov.

hurutaraui, Ceresium

inarmata, Huona

jodiae, Taheitia

kagainisi, Palaina (s. |.)

kerleyi, Perignamptus

Lalahonia gen. nov.

lata, Scaphobaeocera

latimojongensis, Xylobanus

lemurella, Platyja (Mocrendes)

leuseriana, Manilla

longeolivaceus, Metriorrhynchus

longepilosus, Perignamptus

longpela, Taheitia

malagan, Taheitia

marazziorum, Catypnes

mirabile, Ceresium

mundaensis, Perignamptus

nitidifasciatus, Sulabanus

onerosa, Baeocera

pallgergelyi, Palaina (s. |.)

pangoensis, Xylobanus

perplexum, Ceresium

Philgertia gen. nov.

philippinus, Erymus

pronotata, Lalahonia

pulekerai, Ceresium

rendovaensis, Perignamptus

sarmi, Palaina (s. |.)

saxatilis, Palaina (s. |.)

sinajiensis, Xylobanus

solomonicus, Dendrides

speciosa, Lalahonia

striatifrons, Mossula

subcostulatus, Acanthiulus

sukabumensis, Platyja (Mocrendes)

telnovi, Taheitia

torajaensis, Sulabanus

uluwayensis, Mangkutanus

wau, Sohenomorphus

Index to new taxa

145

oe igre

258

268

116

261

re Ware

404

369

116

118

261

386

208

107

422

147

259

200

443

02-Jun-21 21:58:13