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https://doi.org/10.11646/zootaxa.5073.1.1
http://zoobank.org/urn:lsid:zoobank.org:pub:AA3269D1-CA2F-4528-BC9D-3A4C75D05BD9
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Accepted by B. Mantovani: 14 Sept. 2021; published: 6 Dec. 2021
5073
ZOOTAXA
Magnolia Press
Auckland, New Zealand
Zootaxa 5073 (1): 001–189
https://www.mapress.com/j/zt/
Copyright © 2021 Magnolia Press Monograph
Stick insects of Sulawesi, Peleng and the Sula Islands, Indonesia—
a review including checklists of species and descriptions of new taxa
(Insecta: Phasmatodea)
FRANK H. HENNEMANN
Associate researcher, Montréal Insectarium, 4581 rue Sherbrooke, Montréal, Québec, Canada, H1X 2B2
�
hennemann@phasmatodea.com; https://orcid.org/0000-0002-6547-4704
HENNEMANN
2 · Zootaxa 5073 (1) © 2021 Magnolia Press
FRANK H. HENNEMANN
Stick insects of Sulawesi, Peleng and the Sula Islands, Indonesia—a review including checklists of spe-
cies and descriptions of new taxa (Insecta: Phasmatodea)
(Zootaxa 5073)
189 pp.; 30 cm.
6 Dec. 2021
ISBN 978-1-77688-418-6 (paperback)
ISBN 978-1-77688-419-3 (Online edition)
FIRST PUBLISHED IN 2021 BY
Magnolia Press
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purpose other than private research use.
ISSN 1175-5326 (Print edition)
ISSN 1175-5334 (Online edition)
STICK INSECTS OF INDONESIA Zootaxa 5073 (1) © 2021 Magnolia Press · 3
Table of Contents
Abstract ...................................................................................................5
Introduction ................................................................................................6
Biogeography ..............................................................................................7
Material and methods .......................................................................................12
Family Aschiphasmatidae Brunner v. Wattenwyl, 1893 .............................................................13
Subfamily Aschiphasmatinae Brunner v. Wattenwyl, 1893 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Tribe Aschiphasmatini Brunner v. Wattenwyl, 1893 ...............................................................13
Genus Orthomeria Kirby, 1904 ...............................................................................13
Orthomeria forstenii (Haan, 1842) ......................................................................14
Orthomeria limogesi Hennemann & Le Tirant n. sp. ........................................................16
Heteropterygidae Kirby, 1896 .................................................................................21
Dataminae Rehn & Rehn, 1939 ...............................................................................21
Datamini Rehn & Rehn, 1939 .................................................................................21
Genus Pylaemenes Stål, 1875 .................................................................................21
Pylaemenes moluccanus (Redtenbacher, 1906) ............................................................22
Pylaemenes pleurospinosus Hennemann & Le Tirant n. sp. ...................................................25
Lonchodidae Brunner v. Wattenwyl, 1893 .......................................................................28
Lonchodinae Brunner v. Wattenwyl, 1893 .......................................................................28
Lonchodini Brunner v. Wattenwyl, 1893 ........................................................................28
Genus Gibbopromachus n. gen. ...............................................................................28
Genus Hermagoras Stål, 1875 ................................................................................30
Hermagoras celebensis Hennemann, 1998 n. stat. ..........................................................30
Genus Leprocaulinus Uvarov, 1940 ............................................................................32
Leprocaulinus digitatus n. sp. ..........................................................................34
Leprocaulinus sulawesiense n. sp. ......................................................................37
Genus Mnesilochus Stål, 1877 ................................................................................41
Mnesilochus bodiense n. sp. ...........................................................................42
Mnesilochus luwuense n. sp. ...........................................................................44
Genus Myronides Stål, 1875. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
Myronides glaucus n. sp. ..............................................................................49
Myronides pussulatus n. sp. ...........................................................................53
Myronides tomohonense n. sp. .........................................................................56
Genus Neomyronides n. gen. .................................................................................59
Genus Paramanduria n. gen. .................................................................................63
Genus Periphetes Stål, 1877 ..................................................................................66
Periphetes borealis n. sp. ..............................................................................66
Family Phasmatidae Leach, 1815 s. l. ...........................................................................69
Subfamily Clitumninae Brunner v. Wattenwyl, 1893 ...............................................................69
Tribe Clitumnini Brunner v. Wattenwyl, 1893 ....................................................................69
Genus Ramulus Saussure, 1862 ...............................................................................69
Ramulus extensus n. sp. ..............................................................................70
Ramulus globosicaput (Brunner v. Wattenwyl, 1907) .......................................................72
Ramulus melanurus n. sp. .............................................................................74
Ramulus pelengense Hennemann, Le Tirant & Purwanto n. sp. ................................................76
Ramulus togianense n. sp. .............................................................................80
Ramulus torajanus n. sp. ..............................................................................84
Family Phasmatidae Leach, 1815 s. str. .........................................................................88
Subfamily Phasmatinae Gray, 1853 ............................................................................88
Tribe Phasmatini Gray, 1853. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88
Genus Anchiale Stål, 1875 ...................................................................................88
Anchiale maculata (Olivier, 1792) ......................................................................89
Genus Paracyphocrania Redtenbacher, 1908 ....................................................................89
Paracyphocrania major Hennemann, Conle & Suzuki, 2015 ..................................................89
Family Phasmatidae Leach, 1815 s. str. .........................................................................94
Subfamily - ...............................................................................................94
Tribe - ..................................................................................................94
Genus Dimorphodes Westwood, 1859 ..........................................................................94
Keys to the Sulawesian species of Dimorphodes ..................................................................95
Dimorphodes celebensis Redtenbacher, 1908 rev. stat. .......................................................95
Dimorphodes elegans n. sp. ............................................................................96
Dimorphodes sarasini Redtenbacher, 1908 rev. stat.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
Family Phasmatidae Leach, 1815 s. l. ..........................................................................102
HENNEMANN
4 · Zootaxa 5073 (1) © 2021 Magnolia Press
Subfamily Platycraninae Brunner v. Wattenwyl, 1893 .............................................................102
Tribe Platycranini Brunner v. Wattenwyl, 1893 ..................................................................102
Genus Platycrana Gray, 1835 ................................................................................102
Platycrana viridana (Olivier, 1792) ....................................................................102
Tribe Stephanacridini Günther, 1953 ..........................................................................103
Genus Nesiophasma Günther, 1934 ...........................................................................103
Keys to the species of Nesiophasma ..........................................................................105
Nesiophasma giganteum Hennemann, Le Tirant & Purwanto n. sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
Nesiophasma kuehni (Brunner v. Wattenwyl, 1907) ........................................................114
Nesiophasma oligarches (Günther, 1935) ................................................................116
Nesiophasma plateni (Dohrn, 1910) ....................................................................116
Nesiophasma sananaense n. sp. ....................................................................... 117
Nesiophasma spinulosum (Brunner v. Wattenwyl, 1907) .................................................... 119
Nesiophasma turbans (Brunner v. Wattenwyl, 1907) .......................................................120
Nesiophasma zanum Hennemann, 1999 .................................................................123
Family Lonchodidae Brunner v. Wattenwyl, 1893 ................................................................124
Subfamily Necrosciinae Brunner v. Wattenwyl, 1893 .............................................................124
Tribe Necrosciini Brunner v. Wattenwyl, 1893 ...................................................................124
Genus Asceles Redtenbacher, 1908 ...........................................................................124
Asceles rufescens (Redtenbacher, 1908) .................................................................126
Genus Calvisia Stål, 1875 ...................................................................................126
Subgenus Conocalvisia Seow-Choen, 2016 .....................................................................126
Calvisia (Conocalvisia) hippolyte (Westwood, 1859) .......................................................126
Genus Hemiplasta Redtenbacher, 1908 ........................................................................127
Keys to the species of Hemiplasta ............................................................................129
Hemiplasta flavifrons n. sp. ...........................................................................130
Hemiplasta mustea (Bates, 1865) n. comb. ...............................................................133
Hemiplasta nigra (Hennemann, 1998) n. comb. ...........................................................137
Hemiplasta parva n. sp. ..............................................................................141
Hemiplasta rostrata (Redtenbacher, 1908) rev. stat. ........................................................143
Hemiplasta styligera (Bates, 1865) .....................................................................145
Genus Necroscia Serville, 1838 ..............................................................................149
Necroscia aruana Westwood, 1859 .....................................................................149
Necroscia malleoformia n. sp. ........................................................................150
Genus Nescicroa Karny, 1923 rev. stat. ........................................................................153
Keys to the Sulawesian species of Nescicroa ....................................................................157
Nescicroa rufescens (Hennemann, 1998) n. stat. ..........................................................157
Nescicroa splendida n. sp. ...........................................................................159
Genus Singaporoidea Seow-Choen, 2017 ......................................................................162
Singaporoidea inconspicua (Redtenbacher, 1908) n. comb. ..................................................164
Singaporoidea normalis (Redtenbacher, 1908) rev. stat., n. comb. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 164
Singaporoidea poeciloptera (Rehn, 1904) n. comb. ........................................................167
Singaporoidea tenella (Günther, 1935) n. comb. ..........................................................168
Checklist of Phasmatodea of Sulawesi .........................................................................169
Erroneous and doubtful Sulawesian records .....................................................................173
Checklist of Phasmatodea of Peleng ...........................................................................174
Summary ................................................................................................175
Conclusion ..............................................................................................180
Acknowledgements ........................................................................................185
References ...............................................................................................185
STICK INSECTS OF INDONESIA Zootaxa 5073 (1) © 2021 Magnolia Press · 5
Abstract
A review of the stick and leaf insects of Sulawesi, Peleng Island and the Sula islands is presented, along with complete
checklists of species. The genera Hemiplasta Redtenbacher, 1908 and Nesiophasma Günther, 1934 are reviewed in detail
and distinguishing keys to all known species are provided. Distinguishing keys are also provided to the Sulawesian
species of Dimorphodes Westwood, 1859 and Nescicroa Karny, 1923. The latter genus is removed from synonymy
with Paranecroscia Redtenbacher, 1908 and re-established as a valid genus (rev. stat.), with a new diagnosis and a
preliminary list of species provided. As a result, 17 species are here transferred back to Nescicroa (rev. comb.). The genus
Cylindomena Günther, 1935 is removed from synonymy with Parasipyloidea Redtenbacher, 1908 and re-established as a
valid genus (rev. stat.). Consequently, the two species originally attributed to Cylindomena are re-transferred to the genus
(rev. comb.). A new diagnosis along with a list of species is also given for the genus Myronides Stål, 1875, which is shown
to be geographically restricted to Wallacea.
Three new genera in Lonchodidae: Lonchodinae are described from Sulawesi. Gibbopromachus n. gen. is established
only for the type-species Pericentrus tripinnatus Redtenbacher, 1908, a species restricted to mossy, mountainous habitats
in North Sulawesi; consequently this species now becomes Gibbopromachus tripinnatus (Redtenbacher, 1908) n. comb..
The new genus Neomyronides n. gen. (Type-species: Pseudostheneboea aberrans emiri Hennemann, 1998) is established
to comprise three species that have previously been misplaced in other genera. Paramanduria n. gen. is erected to
comprise its type-species Neopromachus celebensis Günther, 1935 as well as Prisomera nodosum Günther, 1938; hence
these two species change to Paramanduria celebensis (Günther, 1935) n. comb. and Paramanduria nodosum (Günther,
1938) n. comb..
Twenty-two new species are described, 14 from Sulawesi, one from the Togian islands, five from the island of
Peleng and two from the island of Sanana. Sulawesi: Dimorphodes elegans n. sp. is decribed from the male and female,
Hemiplasta flavifrons n. sp. from the male, Hemiplasta parva n. sp. from the male, Leprocaulinus digitatus n. sp. from
the female, Leprocaulinus sulawesiense n. sp. from the male, female and egg, Mnesilochus bodiense n. sp. from the
male, Mnesilochus luwuense n. sp. from the female, Myronides tomohonense n. sp. from the male and female, Necroscia
malleoformia n. sp. from the male, Nescicroa splendida n. sp. from the male and female, Periphetes borealis n. sp. from
the male, Ramulus extensus n. sp. from the male, Ramulus melanurus n. sp. from the male and Ramulus torajanus n.
sp. from the male, female and egg. Togian Islands: Ramulus togianense n. sp. is described from both sexes and the eggs.
Peleng: Myronides glaucus n. sp., Nesiophasma giganteum n. sp. and Ramulus pelengense n. sp. are described from both
sexes and the eggs, Orthomeria limogesi n. sp. from the male and female and Pylaemenes pleurospinosus n. sp. from the
female only. Sanana Island: Myronides pussulatus n. sp. and Nesiophasma sananaense n. sp. are described only from the
female sex.
Ten species are synonymised (n. syn.): Calvisia (Conocalvisia) octolineata Redtenbacher, 1908 and Calvisia
(Conocalvisia) reductipennis Redtenbacher, 1908 synonymised with Calvisia (Conocalvisia) hippolyte (Westwood,
1859); Lonchodes reductus Brunner v. Wattenwyl, 1907 synonymised with Myronides cristulatus Brunner v. Wattenwyl,
1907; Necroscia albofasciata Redtenbacher, 1908 synonymised with Nescicroa smaragdula (Bates, 1865); Necroscia
densegranulosa Redtenbacher, 1908 and Sipyloidea (Hemiplasta) falcata Redtenbacher, 1908 synonymised with
Hemiplasta mustea (Bates, 1865) n. comb.; Paranecroscia frontalis (Redtenbacher, 1908) synonymised with Nescicroa
viridilineata (Bates, 1865); Paranecroscia graminea (Bates, 1865: 356), here synonymised with Nescicroa smaragdula
(Bates, 1865); Ramulus melanocephalus (Carl, 1913) synonymised with Ramulus supernumerarius (Brunner v. Wattenwyl,
1907); Sipyloidea felderi Redtenbacher, 1908 synonymised with Singaporoidea poeciloptera (Rehn, 1904) n. comb..
Six subspecies are raised (n. stat.) or re-transferred (rev. stat.) to species level: Dimorphodes celebensis Redtenbacher,
1908 rev. stat., Dimorphodes sarasini Redtenbacher, 1908 rev. stat., Hermagoras celebensis (Hennemann, 1998) n. stat.,
Leprocaulinus mammatus (Rehn, 1904) rev. stat., n. comb., Neomyronides emiri (Hennemann, 1998) n. comb., n. stat.,
Nescicroa rufescens (Hennemann, 1998) n. stat.. Hemiplasta rostrata (Redtenbacher, 1908) is removed from synonymy
with Hemiplasta styligera (Bates, 1865) and Sipyloidea normalis Redtenbacher, 1908) is removed from synonymy with
Sipyloidea poeciloptera Rehn, 1904; both are re-established as a valid species (rev. stat.).
37 species are transferred to other genera (n. comb.): Asceles morio (Redtenbacher, 1908), Asceles roseonotata
(Redtenbacher, 1908), Dimorphodes epidicus (Günther, 1935), Gibbopromachus tripinnatus (Redtenbacher, 1908),
Hemiplasta mustea (Bates, 1865), Hemiplasta nigra (Hennemann, 1998), Leprocaulinus heinrichi (Günther, 1935),
Leprocaulinus kaupii (Stål, 1875), Leprocaulinus mammatus (Rehn, 1904), Lopaphus ashmeadi (Rehn, 1904), Lopaphus
baucis (Westwood, 1859), Lopaphus elegans (Brunner v. Wattenwyl, 1907), Lopaphus magnificus (Brunner v. Wattenwyl,
1907), Myronides batesii (Kirby, 1896), Neomyronides aberrans (Günther, 1938), Neomyronides emiri (Hennemann,
1998), Neomyronides obsolefactum (Günther, 1935), Nescicroa ruficeps (Redtenbacher, 1908), Ophicrania bojei (Haan,
1842), Paramanduria celebensis (Günther, 1935), Paramanduria nodosum (Günther, 1938), Phraortes curvithorax
(Brunner v. Wattenwyl, 1907), Ramulus supernumerarius (Brunner v. Wattenwyl, 1907), Rhamphosipyloidea tristis
(Redtenbacher, 1908), Singaporoidea albilateralis (Hennemann, 1998), Singaporoidea dolorosa (Redtenbacher, 1908),
Singaporoidea fruhstorferi (Günther, 1938), Singaporoidea inconspicua (Redtenbacher, 1908), Singaporoidea janus
(Bates, 1865), Singaporoidea lutea (Redtenbacher, 1908), Singaporoidea macra (Redtenbacher, 1908), Singaporoidea
HENNEMANN
6 · Zootaxa 5073 (1) © 2021 Magnolia Press
normalis (Redtenbacher, 1908), Singaporoidea poeciloptera (Rehn, 1904), Singaporoidea pseudosipylus laevis (Günther,
1936), Singaporoidea pseudosipylus pseudosipylus (Redtenbacher, 1908), Singaporoidea pumila (Werner, 1934) and
Singaporoidea tenella (Günther, 1935).
20 species are transferred back to the genera they have been placed in previously (rev. comb.): Cylindomena acuminata
(Redtenbacher, 1908), Cylindomena scalprifera Günther, 1935, Nescicroa angustata (Redtenbacher, 1908), Nescicroa
compacta (Redtenbacher, 1908), Nescicroa contracta (Redtenbacher, 1908), Nescicroa heinrichi (Günther, 1935),
Nescicroa obliterata (Redtenbacher, 1908), Nescicroa papuana (Brancsik, 1898), Nescicroa rammei (Günther, 1935),
Nescicroa redempta (Redtenbacher, 1908), Nescicroa resignata (Redtenbacher, 1908), Nescicroa rivalis (Redtenbacher,
1908), Nescicroa rufescens (Hennemann, 1998), Nescicroa sanguinata (Redtenbacher, 1908), Nescicroa smaragdula
(Bates, 1865), Nescicroa tereticollis (Redtenbacher, 1908), Nescicroa terminalis (Redtenbacher, 1908), Nescicroa
tumescens (Redtenbacher, 1908) and Nescicroa viridilineata (Bates, 1865). The world‘s longest stick insect Phobaeticus
chani Bragg, 2008 is removed from the genus Sadyattes Stål, 1877 (Platycraninae: Stephanacridini) and transferred back
to its original genus Phobaeticus Brunner v. Wattenwyl, 1907 (Clitumninae: Clitumnini) rev. comb..
Lectotypes are designated for 17 species: Aruanoidea inconspicua Redtenbacher, 1908; Calvisia octo-lineata
Redtenbacher, 1908; Calvisia reductipennis Redtenbacher, 1908; Clitumnus redemptus Brunner v. Wattenwyl, 1907;
Cuniculina melanocephala Carl, 1913; Datames moluccanus Redtenbacher, 1906; Dimorphodes bellicosus Redtenbacher,
1908; Dimorphodes celebensis Redtenbacher, 1908; Myronides cristulatus Brunner v. Wattenwyl, 1907; Necroscia
frontalis Redtenbacher, 1908; Necroscia viridilineata Bates, 1865; Phobaeticus kuehni Brunner v. Wattenwyl, 1907;
Phryganistria plateni Dohrn, 1910; Sipyloidea felderi Redtenbacher, 1908; Sipyloidea normalis Redtenbacher, 1908;
Sipyloidea (Hemiplasta) falcata Redtenbacher, 1908; and Sipyloidea (Hemiplasta) rostrata Redtenbacher, 1908.
The male of Hemiplasta styligera (Bates, 1865) and Paracyphocrania major Hennemann, Conle & Suzuki, 2015 are
described and illustrated for the first time and information on the natural habitats, host plants and life-cycle of the latter
species in Peleng are provided. The egg of Dimorphodes sarasini Redtenbacher, 1908 is described and illustrated for the
first time.
Key words: Phasmida, phasmids, Indonesia, Wallacea, Togian Islands, Sanana, new genus, new species, new synonyms,
lectotypes, Hemiplasta, Myronides, Nesiophasma, Nescicroa
Introduction
Phasmatodea are one of the very few insect orders that still lack a robust higher-level phylogeny and apart from the
higher classification also the relationships and delimitations of many genera are still speculative and subject to con-
tinous research. Hence, numerous intergeneric relationships are still very questionable and certainly deserve much
further work to properly delimit genera and to reflect evolutionary history. This should also include re-describing
and defining the known genera and species and uncovering synonymies and misidentifications, to which the present
paper is meant to contribute.
Although Sulawesi is the largest of the Indonesian islands that lie within the Wallacea region, the apparently
very diverse phasmid fauna of Sulawesi and nearby islands is still rather poorly known. Only few papers have up
to now engaged in studying the islands fauna. The first species that have since then been recorded from Sulawesi
and the island of Peleng were described by Linné (1758), Haan (1842), Bates (1865) and Kaup (1871), but some of
them originally from other islands of Wallacea. A considerable number of species from Sulawesi and Peleng were
described by Brunner v. Wattenwyl (1907) and Redtenbacher (1906, 1908) but it was only in 1935 that Günther
published the first paper that exclusively dealt with the Phasmatodea of Sulawesi. Günther (1935) studied the 68
superbly preserved phasmids collected by Gerd Heinrich (1930–1932) from various localities throughout most of
the islands‘ geography and nowadays housed in the collection of the Museum für Naturkunde der Humboldt-Uni-
versität, Berlin (MNHU). These comprised 27 different species belonging to 19 genera, of which Günther (1935)
described 17 species and seven genera as new to science. Only three years later Günther (1938) studied the com-
prehensive collection made by F. Sarasin (1894–1896) and mainly housed in the Naturhistorisches Museum Basel
(NHMB) with some specimens in the Polish Academy of Sciences, Warszawa (ZMPA) and a few in the Naturhisto-
rische Museum, Vienna (NHMW), the latter of which had already mostly been described by Brunner v. Wattenwyl
(1907) and Redtenbacher (1906, 1908). The paper also included specimens collected by H. Fruhstorfer in 1895 and
deposited in the collection of the Staatliches Museum für Naturkunde, Dresden (SMTD). The specimens collected
by F. Sarasin and in the collections of NHMB and ZMPA comprised 27 species and five subspecies, of which Gün-
STICK INSECTS OF INDONESIA Zootaxa 5073 (1) © 2021 Magnolia Press · 7
ther (1938) described seven species and four subspecies as new. This 1938 paper by Günther also included a brief
analysis of the phasmid fauna of Sulawesi as well as a checklist that listed 64 different species and seven subspecies
from 35 different genera. Hennemann (1998) worked on a small number of Phasmatodea collected by himself in
the Central Toraja Highland in 1995 and further fresh material available from the same region as well as the south-
western portion of the island. In total this paper described three new species and three new subspecies, provided
descriptions of previously unknown sexes and eggs of several species and introduced several new synonymies. The
paper furthermore presented a catalogue with information on the known records for all 72 different species and ten
subspecies so far recorded from Sulawesi along with a brief analysis of the islands’ fauna with aspect of the bio-
geographic relationships of the individual taxa. The members of the tribe Phasmatini (subfamily Phasmatinae) and
recorded from Sulawesi very studied in detail by Hennemann, Conle & Suzuki (2015).
The possibility to re-examine most of the historic material in more detail and the availablility of fresh material
for examination has revealed several inaccuracies throughout previous works on the phasmid fauna of Sulawesi.
Along with a good number of new genera and species not only from Sulawesi but also from the nearby island of
Peleng and from the Sula Islands, numerous misidentifications and misinterpretations of taxa, some of which have
resulted in false records for either island, as well as new and erroneous synonymies could be uncovered. Moreover,
several taxa originally described as subspecies were found to represent separate species and were in need of valid
species status. Finally, the much better photographic equipment now available also allows better and comprehen-
sive illustration of several of the lesser known taxa. The numerous taxonomic changes that result from all these
discoveries render it appropriate to provide a new and updated checklist of the Phasmatodea of Sulawesi along with
a checklist for the island of Peleng, which is included in the present paper, intended as a major contribution to the
knowledge of the fauna of these islands, which apparently is more diverse than expected previously.
Biogeography
For better understanding the biogeographic importance of Sulawesi, the former Celebes, and the various influences
on the islands biota a definition of the Wallacea region as well as a summarised explanation of its geography and
geology is provided.
The region of oceanic islands in the centre of the Indo-Australian Archipelago comprising Sulawesi, the Ma-
luku Islands (also called Moluccas), the Kei Islands, Banda Islands and the Lesser Sunda Islands (except Bali) is
termed Wallacea (Dickerson, 1928). This group of several thousand islands of various sizes covers a total land area
of roughly 338.500 km2 located between Sundaland in the west, the Philippines in the north as well as New Guinea
and Australia in the east and southeast are located almost entirely within the borders of Indonesia. The largest and
most western of these islands is Sulawesi, which has an area of approximately 174.600 km². The Lesser Sunda
Islands attributed to Wallacea are located south of Sulawesi and include Lombok, Sumba, Sumbawa, Flores and
Timor, as well as several hundred smaller intervening islands. The Maluku Islands include several hundred islands
in the northern portion of Wallacea, the largest being Halmahera (area 17.780 km²) in the north and Seram (17.100
km²) in the south (Fig. 3). The most northern islands of the Maluku Islands and Wallacea are the Sangihe Islands,
the most southeastern ones are the Kei Islands and Banda Islands, situated west of New Guinea (Fig. 1). In the north,
Wallacea borders the other large region of oceanic islands, the Philippines. While it has been regarded as a distinct
biogeographic entity (e.g. Cox, 2001) because it has a largely unique fauna of its own (e.g. Heaney et al. 2005),
more recent studies have revealed that the Philippines should rather be considered as part of Wallacea (e.g. Vallejo,
2011). Nowadays, most of the Wallacean Islands and Philippines are regarded as true oceanic islands that have had
no terrestrial connection to surrounding areas since they emerged (van Osterzee, 1997). Principally, Wallacea is a
transition zone between the Oriental region and the Australian region with the fauna showing influences from both
these regions (Dickerson, 1928), although the concept of a transition zone was shown to be inadequate in itself
(Simpson, 1977). The largely endemic fauna of this biodiversity hotspot (Myers et al., 2000) is remarkably distinct
from that of the continental islands on the adjacent shelves. Wallacea is regarded to be part of the Papuan region (or
Papuan subregion of the Australian region in the strict zoogeographic sense) as defined by e.g. Beehler et al. (1986)
or Gressitt (1982). The western border of Wallacea is represented by Wallace’s line (named by Huxley in 1868),
which runs between Borneo and Sulawesi in the north and Lombok and Bali in the south (Fig. 2). Wallace’s line is
named after the well known British naturalist Alfred Russel Wallace (1823–1913), who travelled extensively in the
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area between 1854 and 1862 and noticed fairly distinct bird and mammal faunas on either side of the line (Wallace,
1869). The eastern border of Wallacea and the eastern counterpart of Wallace’s line is Lydekker’s line Lydekker
(1896). Lydekker’s line follows the Sahul shelf, thus runs between Seram and the Banda Islands in the west and
New Guinea in the east (Fig. 2). Another well-known and much discussed line is Weber’s line, which runs between
Sulawesi, the Banggai Islands and Sula Islands in the west and the Maluku Islands in the east and represents the
western border of the Papuan region (Gressitt, 1982). Mayr (1944) regarded Weber’s line as a line of faunal balance,
east of which the Papuan faunistic elements have a numerical superiority over Oriental elements. Hence, Sulawesi
as well as the Banggai Islands and Sula Islands belong to Wallacea but are not part of the Papuan region.
Wallacea is regarded as a biodiversity hotspot (Myers et al., 2000) although the subregion contains fewer spe-
cies overall compared to e.g. Sundaland, the Philippines or the Australian region. However, Wallacea contains a
very high percentage of endemic species, i.e. species that are only found in Wallacea or on single islands within
this subregion, which is explained by several factors. First of all the subregion contains a large number of different
islands and geographic isolation is one of the major factors which leads to diversification and formation of endemic
species. Islands are isolated because they are surrounded by areas of sea, which form a barrier for migration and
species inhabiting isolated islands generally tend to evolve into separate species adapted to their local habitats and
ecological niches. Other factors are represented by the geological history of Wallacea. Wallace’s line marks the
position of deep ocean straits and separates shallow waters in the west from deep waters in the east. The sea-level
dropped by estimated 120 metres during the last ice-age and while the shallow waters in the west disappeared and
formed land connections between the Large Sunda Islands (Sumatra and Java), Borneo and the remaining portions
of Sundaland, there were no such connections towards Wallacea. Hence, species were able to disperse throughout
present Sundaland but not into Wallacea, which remained isolated by the deep waters preventing migration from
species of Sundaland and leading to evolution of distinct faunas. This is also the case for islands on the Sahul shelf,
which were connected to the mainland at times of low sea level during the Pleistocene.
FIGURE 1. Map of eastern Indonesia showing the principle islands.
“We now come to the Island of Celebes, in many respects the most remarkable and interesting in the whole
region, or perhaps on the globe, since no other island seems to present so many curious problems for solu-
tion.” Wallace (1876, Vol. 1: 426)
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Sulawesi, the largest of the islands in Wallacea and the world‘s eleventh-largest island is possibly also the oldest
island of Wallacea and apparently has the most diverse fauna of that region (e.g. Whitten et al., 2002). The complex
geological history of Wallacea is mirrored in the geology of Sulawesi, which in contrast to other Wallacea islands
is not truly oceanic, but a composite island that lies at the centre of the Asia-Australia collision zone (Stelbrink et
al., 2012). Today it is estimated, that parts of the island were formerly attached to either the Asian or Australian
continental margin and separated from these areas by vicariant processes (see Stelbrink et al., 2012). In the West,
the opening of the Makassar Strait separated West Sulawesi from Borneo and Sundaland in the Eocene ca. 45 Mya
(Hall, 2011). In the East, it is assumed that extensional fragmentation has taken place following a single collision of
the Sula Spur with West Sulawesi in the Early Miocene ca. 20 Mya (Hall, 2011; Stelbrink et al., 2012).
FIGURE 2. Map of the Wallacea subregion with the Wallace, Weber and Lydekker lines indicated. All islands that biogeo-
graphically belong to Wallacea are marked green.
Basically, Sulawesi consists of four peninsulas, the Minahasa or Northern Peninsula, the East Peninsula, the
Southeast Peninsula and the South Peninsula. Politically, it is divided into the six provinces from north to south:
Sulawesi Utara (North Sulawesi), Gorontalo, Sulawesi Tengah (Central Sulawesi), Sulawesi Barat (West Sulawesi),
Sulawesi Tengah (South Sulawesi) and Sulawesi Tenggara (Southeast Sulawesi). Several nearby islands biogeo-
graphically belong to Sulawesi, the main ones being the Selayar islands that stretch southward into the Flores Sea,
Buton Island and surrounding islands that form an archipelago in the southeast, the Banggai Islands including
Peleng that are positioned between the Sulawesi and the Sula Islands in the east, and the Togian Islands in the Gulf
of Tomini. Sulawesi is largely mountainous with most of the island lying above 500 metres and as much as one-fifth
above the 1000 metres mark. The highest elevations are found in Central and northern South Sulawesi, the highest
peak being Mount Latimojong (local name Gunung Rantemario) in northern Sulawesi Tengah, at 3478 metres. The
lowland forests and mangroves of Sulawesi have largely been lost or degraded with intact forest habitats meanwhile
mostly restricted to mountainous regions, the four terrestrial national parks and various nature reserves. The two
parks with the largest terrestrial area are Bogani Nani Wartabone National Park (2871 km2) on the Minahasa Pen-
insular, Northern Sulawesi, and Lore Lindu National Park (2290 km2) in Central Sulawesi. While there are several
dead volcanoes in South Sulawesi, the most prominent of these is Gunung Lompobatang (“swollen belly”) some 80
kilometres southeast of the capital Ujung Pandang at 2874 metres, the Northern Peninsula has several active volca-
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noes. Sulawesi has 13 lakes of more than 5 km2 surface area, the largest being Lake Towuti in Southeast Sulawesi
and Lake Poso in southern Central Sulawesi. Since the shape of the island precludes the development of any large
rivers, Sulawesi’s longest river, the Lariang is barely 200 kilometres long.
FIGURE 3. Topographic map of Sulawesi, the Banggai and Sula Islands showing the seven provinces and their capitals. (©
http://commons.wikimedia.org; edited)
Sulawesi has a rich fauna of mixed geographic and temporal origin. The geographic origin of Sulawesi’s fauna
and the mode of colonization of the island are still disputed to some extent even almost 150 years after Wallace’s
Line was proposed (Stelbrink et al., 2012). Given Sulawesi’s geographic position, it has been realized that the fauna
is predominantly of Asian origin (e.g. Sarasin & Sarasin, 1901; Whitten et al., 2002), which has gained support by
recent molecular phylogenies that involved taxa from across the Wallace line. These have also identified decisive
STICK INSECTS OF INDONESIA Zootaxa 5073 (1) © 2021 Magnolia Press · 11
shaping of the fauna by recent Miocene to Pleistocene dispersal as the most likely and common mechanism, while
vicariance appears to be true for a considerably lesser number of taxa (see Stelbrink et al., 2012). Moreover, it has
become obvious that the simple and traditional dichotomic approach that has divided the origins of biota into “Asian
origin” and “Australian origin” does not reflect the much more complex true distribution patterns. However, still
considerably more data is needed to gain a comprehensive understanding of Sulawesi’s biogeography and the geo-
graphic and temporal composition of its diverse fauna.
FIGURE 4. Map of Sulawesi showing the localities recorded in this study.
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Off the eastern end of Central Sulawesi is the Banggai Archipelago (Indonesian: Kepulauan Banggai), a group
of islands that politically makes up the Banggai Islands Regency of the Central Sulawesi province (Sulawesi Ten-
gah) and is separated from mainland Sulawesi by the Peleng Strait (Indonesian: Selat Peleng). The main island of
Peleng has an area of about 2340 km2 and is surrounded by more than 300 smaller islands, most of which are situ-
ated south-east ward. Peleng only rises to just over 1000 m and the majority is still covered by tropical rainforest.
Due to its close vicinity to Sulawesi, the fauna and flora generally shows close relation and concordance to that of
mainland Sulawesi. However, although only separated by about 15 km of sea water, Peleng has never been con-
nected to Sulawesi because of a deep sea channel that separates these two islands. Instead, Peleng and some of its
surrounding islands have had Quaternary land connections with each other and have formed one paleoisland. South-
east of the Banggai Islands is the Sula Islands Archipelago, which comprises the islands of Taliabu (Taliabu Island
Regency, Indonesian: Kabupaten Pulau Taliabu) and the islands of Mangole and Sanana (= Sulabesi) in the east
and south-east, which belong to the Sula Islands Regency. Politically the Sula Islands are part of the North Maluku
Province, but biogeographically they show more close affinity to the Banggai Islands rather than to the rest of the
Maluku Islands. Prospecting the avifauna, Rheindt et al. (2020) have supported the close relation between the fau-
nas of these islands and have also shown the great potential of the presence of still undescribed taxa by describing
as many as ten new species and subspecies of birds from mountainous habitats of Peleng and Taliabu.
Material and methods
This study is based on examination of most necessary type-material and a large number of specimens throughout
various museum collections and the collections of the authors. The material used for this study is invariably dried
and pinned. If no live specimens were available, the colouration is described from dried specimens, to which atten-
tion needs to be drawn, as the insects may have changed colour due to preservation. All eggs examined were already
laid and thus fully developed. Examinations of the insects and eggs were carried out using an entomological lens
with 4x magnification and a Russian MBC-10 binocular microscope. Eggs were examined at 10x magnification.
Measurements were taken using a long ruler or a digital calliper and are given to 0.1 mm. The terminology used for
the descriptions of external and internal egg structures follows that of Clark-Sellick (1997). Data of examined speci-
mens are reproduced exactly as on the labels and labels are separated by a semicolon. Photographs of specimens
were taken using a Nikon D7000 camera equipped with a Nikon DX AF-S Micro 40mm lens and a wireless Nikon
SU-800 dual speed light system. Background lightning was provided by a 18w 6000K LED panel light plate. Corel
Photo Paint X3 was used as post processing software. Photos of the holotype of Orthomeria limogesi n. sp. were
taken by René Limoges (IMQC) using a Nikon D810 DSLR camera with a Nikon Micro-Nikkor 200mm f/4 lens on
a Manfrotto 454 micrometric positioning sliding plate. Lightning was provided by two Nikon SB-25 flash units with
a Cameron Digital diffusion photo box. Adobe Photoshop Elements 13 was used as post processing software.
Abbreviatations
ANSP: Academy of Natural Sciences, Philadelphia, Pennsylvania / U.S.A.
CMN: Canadian Museum of Nature, Ontario, Ottawa / Canada.
ETHZ: Eidgenössische Technische Hochschule, Zürich / Switzerland.
IMQC: Insectarium de Montréal, Montréal, Québec / Canada.
IRSN: Institut Royal des Sciences Naturelles de Belgique, Bruxelles / Belgium.
LEMQ: Lyman Entomological Museum, McGill University, Québec / Canada.
MHNG: Museum d’Histoire Naturelle, Geneva / Switzerland.
MNCN: Museo Nacional de Ciencas Naturales, Madrid / Spain.
MNHN: Museum d’Histoire Naturelle, Paris / France.
MNHU: Museum für Naturkunde der Humboldt-Universität, Berlin / Germany.
NHMB: Naturhistorisches Museum, Basel / Switzerland.
NHMUK: Natural History Museum, London / United Kingdom.
NHMW: Naturhistorisches Museum, Vienna / Austria.
NMEG: Naturkundemuseum Erfurt / Germany.
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OXUM: Oxford University Museum, Oxford / England.
RMNH: Rijksmuseum van Natuurlijke Historie, Leiden / Netherlands.
SMTD: Sttatliches Museum für Tierkunde, Dresden / Germany.
ZMAN: Zoologisch Museum, Universiteit van Amsterdam, Amsterdam / Netherlands.
ZMAS: Zoological Museum, Academy of Science, St. Petersburg / Russia
ZMPA: Polish Academy of Sciences, Warszawa / Poland.
ZSMC: State Zoological collections, Munich / Germany.
AB: Personal collection of Alexandre Banko / Canada.
AJH: Personal collection of Allan J. E. Harman / England.
EB: Personal collecton of Edy Bhaskara / Indonesia.
FH: Personal collection of Frank H. Hennemann / Germany.
OC: Personal collection of Oskar V. Conle / Germany.
PEB: Personal collection of Phil E. Bragg / United Kingdom.
RL: Personal collection of René Limoges / Canada.
RTC: Personal collection of Royce T. Cumming / U.S.A.
HT: Holotype
AT: Allotype
PT: Paratype
LT: Lectotype
PLT: Paralectotype
ST: Syntype
Family Aschiphasmatidae Brunner v. Wattenwyl, 1893
Subfamily Aschiphasmatinae Brunner v. Wattenwyl, 1893
Tribe Aschiphasmatini Brunner v. Wattenwyl, 1893
Genus Orthomeria Kirby, 1904
(Figs. 5–8)
Type-species: Phasma (Ascephasma) forstenii Haan, 1842: 114, by original designation.
Comments: A revision of this genus was presented by Bragg (2001: 305) and some of the Bornean species were
illustrated by Seow-Choen (2016 & 2017). Examination of extensive material of the genus, particularly from
throughout the islands of the Philippine but also Wallacea in the authors collection, shows the genus is still in need
of a detailed revision. In particular, the current synonymy of O. pandora (Westwood, 1859) and O. catadromus
(Westwood, 1859) deserves clarification. The first species was originally described upon three ♂♂ originating from
the Philippines and the island of Seram in southern Wallacea, and the latter from three females without confirmed
locality (see comments by Vallotto et al., 2016: 52 and Seow-Choen, 2018: 553).
The validity of Coloratobistus Zompro, 2004 (Type-species: C. dilawitimpakpak Zompro, 2004: 83), a genus
described from the Philippine island of Mindanao, is very questionable and deserves evaluation. The very brief and
insufficient generic diagnostic given by Zompro (2004: 82) provides no characters that would allow distinction from
Orthomeria. Several ♂♂ at hand from the author’s collection are without any doubt congeneric with Orthomeria.
Distribution: Philippines, Wallacea, Borneo and Palawan.
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Orthomeria forstenii (Haan, 1842)
(Fig. 5)
Phasma (Ascephasma) forstenii Haan, 1842: 114. LT, ♂: Forsten, Tondano, Celebes [RMNH]; PLT, ♂: no data [RMNH].
(Not: PLT, ♀: no data [RMNH], this specimen is a different species)
Ascephasma forsteni, Redtenbacher, 1906: 76.
Giglio-Tos, 1910: 5.
Bruner, 1915: 230.
Werner, 1931: 25.
Aschipasma forstenii, Westwood, 1859: 93.
Aschiphasma forsteni, Vanschuytbroeck & Cools, 1981: 23.
Orthomeria forstenii, Kirby, 1904: 420.
Günther, 1938: 57.
Bragg, 1996: 111 (in part—only specimens from Sulawesi)
Hennemann, 1998: 125.
Bragg, 2001: 314, fig. 113 (egg).
Otte & Brock, 2005: 240.
Orthomeria pandora, Günther, 1938: 60.
Material examined: 1 ♂: Celebes, Manado; 60 76; NHMUK 012496930 [NHMUK]; 1 ♂, 1 ♀: Indonesia, N-Su-
lawesi, 1km S Sawangan Flußtal b. River Park Hotel, p1°22’51’’N, 124°56’56’’E, 01.–03.II.2004, 250–300m, leg.
A Weigel UWS/Plant [NMEG]; 14 ♀♀: Indonesien, Zentral Sulawesi, Prov. Sulawesi Tengah, Palolo District, Palu,
VIII.2012 [coll. FH, No’s 0725-1 to 14]; 1 ♂: Nord-Celebes, Toli Toli, 1895, H. Fruhstorfer; det. Redtenb. Asce-
phasma forsteni; 20.729 [NHMW, No. 89]; 2 ♂♂: Nord-Celebes, Toli Toli, Nov.-Dez. 1895, H. Fruhstorfer; det.
Redtenb. Ascephasma forsteni [NHMW, No. 89]; 1 ♀: Nord-Celebes, Toli Toli, Nov.-Dez. 1895, H. Fruhstorfer; Or-
thomeria pandora Gtr. Det. [ZMPA]; 1 ♀: Sararin 6.1894, Tomohon, Nord.-Cel.,; Tomohon; Orthomeria pandora
Westw., K. Günther det. [NHMB]; 2 ♀♀ (nymphs): Sarasin IX.1894, Masarang-Kette, Nord.-Celebes; Orthomeria
pandora Westw., larva K. Günther det. [NHMB], 1 ♂, 1 ♀: Sarasin, 6.II.1895, Lembongpangi (Luwu), elv. 500
m, Cent. Cel.; Orthomeria pandora Westw. K. Günther det. [NHMB]; 3 ♂♂: Indonesien, Zentral Sulawesi, Prov.
Sulawesi Tengah, Sigi Regency, Palolo, Mount Nokilalaki, V.2012 [coll. FH, No’s 0725-15 to 17]; 2 ♂♂: Central
Sulawesi, Prov. Sulawesi Tengah, Sigi Regency, Palolo, Mount Nokilalaki, V.2012 [coll. SLT].
Comments: This species was originally described from two ♂♂ and a ♀ from Tondano, Minahasa in northern
Sulawesi. The specimen which Haan (1842: 114) described as the ♀ of O. forstenii is not the same species, a fact
that was first recognized by Redtenbacher (1906: 76) and subsequently confirmed by Bragg (2001: 316). While
Redtenbacher attributed the specimen to O. pandora (Westwood, 1859), this was rejected by Bragg (2001: 316) who
left the identity as yet unconfirmed. Indeed, the specimen has much longer wings than all other ♀♀ that are here
attributed to O. forstenii and have collecting data. Unfortunately however, Haan’s specimen could only be examined
from a single photograph which is why the exact identity still remains unclarified. Bragg (2001: 314) designated a
lectotype for O. forstenii to guarantee stability of the taxon and provided a description of the ♀♀ based on a speci-
men from Lore Lindu National Park, Sulawesi Tengah. In addition, Bragg (2001: 315) provided a description and
illustrations of the eggs.
With great certainty O. forstenii is an endemic of Sulawesi, hence the records from the Philippines (islands of
Mindanao and Leyte) are here regarded as wrong. Specimens at hand from both these islands in the authors collec-
tion clearly represent a distinct species.
Variability: Females occur in two different colour forms, having the costal region of the alae anterior to the
radial vein either plain orange (Figs. 5A, C) or pale green (Figs. 5B, D); correspondingly the veins posterior to the
radial vein are either orange or green. Green specimens may have the basal portion more or less distinctly orange
although (Fig. 5D). The tegmina are mostly orange but green in specimens that also have the entire anterior portion
of the alae green. The antennae are always black basally and gradually become dark brown towards the tip. Abdomi-
nal tergum VII is pale cream to mid brown and abdominal terga II–VI may have two faint but broad and parallel
longitudinal pale stripes. Body lengths ♀♀: 44.0–51.0 mm, ♂♂ [coll. FH] 36.3–37.5 mm.
Distribution: N-Sulawesi, Sulawesi Utara, Minahasa, Tondano [RMNH]; N-Sulawesi, Minahasa, Tomohon
[NHMB]; N-Sulawesi, Minahasa, S-Sawangan River valley, 250–300 m [NMEG]; N-Sulawesi, Sulawesi Utara,
Minahasa, Gunung Masareng [NHMB]; N-Sulawesi, Sulawesi Utara, Manado [NHMUK]; N-Sulawesi, Sulawesi
Tengah, Tolitoli [NHMW, ZMAS]; Central Sulawesi, Sulawesi Tengah, Lore Lindu National Park, Kamarora [coll.
PEB]; Central Sulawesi, Sulawesi Tengah, Palu [coll. FH]; Central Sulawesi, Sulawesi Tengah, Mount Nokilalaki
[coll. FH]; Central Sulawesi, Sulawesi Tengah, between Palu & Kulavi [IRSN].
STICK INSECTS OF INDONESIA Zootaxa 5073 (1) © 2021 Magnolia Press · 15
FIGURE 5. Orthomeria forstenii (Haan, 1842): A. ♀ (orange colour morp), dorsal view [coll. FH, No. 0725-2]; B. ♀ (green
colour morp), dorsal view [coll. FH, No. 0725-1]; C. ♀ (orange colour morp), dorso-lateral view [coll. FH, No. 0725-6]; D. ♀
(green colour morp), dorso-lateral view [coll. FH, No. 0725-12]; E. ♂ in dorsal view [coll. FH, No. 0725-15]; F. ♀, terminal
abdominal segments in lateral view [coll. FH, No. 0725-6]; G. ♀, terminal abdominal segments in dorsal view [coll. FH, No.
0725-1]; H. ♂, terminal abdominal segments in lateral view [coll. FH, No. 0725-17]; J. ♂, terminal abdominal segments in
ventral view [coll. FH, No. 0725-15]; K. ♂, dorsal view of head, pro- and mesonotum [coll. FH, No. 0725-15].
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Orthomeria limogesi Hennemann & Le Tirant n. sp.
(Figs. 6–8)
HT, ♂: E-Sulawesi, Prov. Sulawesi Tengah, Banggai Ids. E-Peleng Island, Tinangkung Utara District, near Luksagu village, ca.
60 m elev., A. Brata leg., XI–XII.2016 [IMQC].
PT, 27 ♂♂, 24 ♀♀: E-Sulawesi, Prov. Sulawesi Tengah, Banggai Ids., E-Peleng Island, Tinangkung Utara District, near Luk-
sagu village, ca. 60 m elev., A. Brata leg., XI–XII.2016 [IMQC].
PT, 1 ♂, 1 ♀: E-Sulawesi, Prov. Sulawesi Tengah, Banggai Ids., E-Peleng Island, Tinangkung Utara District, near Luksagu vil-
lage, ca. 60 m elev., A. Brata leg., XI–XII.2016 [MNHN].
PT, 1 ♂, 1 ♀: E-Sulawesi, Prov. Sulawesi Tengah, Banggai Ids., E-Peleng Island, Tinangkung Utara District, near Luksagu vil-
lage, ca. 60 m elev., A. Brata leg., XI–XII.2016 [IRSN].
PT, 1 ♂, 1 ♀: E-Sulawesi, Prov. Sulawesi Tengah, Banggai Ids., E-Peleng Island, Tinangkung Utara, District, near Luksagu
village, ca. 60 m elev., A. Brata leg., XI–XII.2016 [NHMB].
PT, 1 ♂, 1 ♀: E-Sulawesi, Prov. Sulawesi Tengah, Banggai Ids., E-Peleng Island, Tinangkung Utara District, near Luksagu vil-
lage, ca. 60 m elev., A. Brata leg., XI–XII.2016 [NHMW].
PT, 1 ♂, 1 ♀: E-Sulawesi, Prov. Sulawesi Tengah, Banggai Ids., E-Peleng Island, Tinangkung Utara District, near Luksagu vil-
lage, a. 60 m elev., A. Brata leg., XI–XII.2016 [SMFM].
PT, 1 ♂, 1 ♀: E-Sulawesi, Prov. Sulawesi Tengah, Banggai Ids., E-Peleng Island, Tinangkung Utara District, near Luksagu vil-
lage, ca. 60 m elev., A. Brata leg., XI–XII.2016 [CMN].
PT, 2 ♂♂, 2 ♀♀: O-Sulawesi, Prov. Sulawesi Tengah, Banggai Ids., W-Peleng Island, Buko District, Tinanasu, VI.2011 [coll.
FH, No’s 0734-1 to 4].
PT, 11 ♂♂, 6 ♀♀: O-Sulawesi, Prov. Sulawesi Tengah, Banggai Ids. E-Peleng Island, Tinangkung Utara District, near Luksagu
village, ca. 60 m elev., E. Brata leg., XI–XII.2016 [coll. FH, No’s 0734-5 to 21].
PT, 5 ♂♂, 5 ♀♀: O-Sulawesi, Prov. Sulawesi Tengah, Banggai-Inseln, W-Peleng Island, Buko District, btw. Tatendeng village
and Eben village 400–550 m, IX.2011 [coll. FH, No’s 0734-22 to 31].
PT, 1 ♂, 1 ♀: E-Sulawesi, Prov. Sulawesi Tengah, Banggai Ids. E-Peleng Island, Tinangkung Utara District, near Luksagu vil-
lage, ca. 60 m elev., E. Brata leg., XI–XII.2016 [coll. RTC].
PT, 1 ♂, 1 ♀: E-Sulawesi, Prov. Sulawesi Tengah, Banggai Ids. E-Peleng Island, Tinangkung Utara District, near Luksagu vil-
lage, ca. 60 m elev., E. Brata leg., XI–XII.2016 [coll. RL].
PT, 2 ♂♂, 2 ♀♀: E-Sulawesi, Prov. Sulawesi Tengah, Banggai Ids. E-Peleng Island, Tinangkung Utara District, near Luksagu
village, ca. 60 m elev., E. Brata leg., XI–XII.2016 [coll. AB].
PT, 3 ♂♂, 3 ♀♀: E-Sulawesi, Prov. Sulawesi Tengah, Banggai-Islands, W-Peleng Island, Bulagi District, Alul village near
Bulagi, VI.2020 [coll. EB].
PT, ♀: Indonesia: Peleng, Tinangkung Utara District, Luksagu, Hutanlomou, Lolou Forest, 1.I.2016, A. Brata leg. [coll. RTC].
PT, ♂: Indonesia, W-Peleng Is., Buko District Tinanasu, 1.2017 [coll. RTC].
PT, ♂: Indonesia, Peleng, 06.2007 [coll. OC, No. 0543-1].
PT, ♂: Indonesia, Peleng, 08.2006 [coll. OC, No. 0543-2].
PT, ♂: Indonesia, Peleng, 04.2007 [coll. OC, No. 0543-3].
Etymology: This new species is dedicated to René Limoges, entomological technician at the Montreal Insectarium
(IMQC), for taking photos of this species for this work as well as for many professional courtesies.
Differential diagnosis: Readily distinguished from the Sulawesian O. forstenii (Haan, 1842) by the larger size,
longer alae, relatively smaller head and dull orange antennae of both sexes (black in forstenii). From the lectotype
of the Philippine O. pandora (Westwood, 1859) ♂♂ of this new species differ by the smaller head, that is notably
narrower than the pronotum, in relation slightly shorter mesonotum, differently shaped cerci (Fig. 6F) and dull or-
ange antennae (except scapus and pedicellus). Since the synonymy of O. catadromus (Westwood, 1859), which is
currently regarded as a synonym of O. pandora (see Vallotto et al., 2016: 52), deserves validation as no confirmed
differentiation of ♀♀ of O. limogesi n. sp. can be presented at this place. It may however be mentioned that ♀♀
of this new species differ from the type-specimens of O. catadromus by the relatively smaller head, that is notably
narrower than the pronotum and the dull orange antennae.
Description: The colouration is described from dried specimens and colour photographs taken of live speci-
mens by local collectors.
♀(Figs.7,8C). Of average size for the genus (body length 46.5–55.5 mm), typical in shape with a compara-
tively small head, that is narrower than the pronotum. Entire body surface covered with fine, cream to dull yellow
setae; setae on legs but tarsi in particular dull orange. General colour mid to dark brown, sometimes with a slight
reddish hue. Abdominal tergum VII sometimes of slightly paler colour than rest of body. Head blackish brown and
somewhat darker than rest of body and legs. Antennae (except scapus and pedicellus) orange to reddish mid brown.
Eyes dark ochre to reddish brown. Tegmina dark orange. Costal region of alae dark brown with veins dark orange;
STICK INSECTS OF INDONESIA Zootaxa 5073 (1) © 2021 Magnolia Press · 17
the basal half of the radial vein and some of the diverging transverse veins sometimes with a green wash (Fig. 8C).
Anal region of alae plain greyish mid brown, slightly transparent with the longitudinal veins dark brown and the
transverse veins transparent (Fig. 7E).
FIGURE 6. Orthomeria limogesi n. sp. ♀♀, paratypes: A. ♀ (PT) from Tinangkung Utara District, dorsal view [coll. FH, No.
0734-1]; B. ♀ (PT) from Tinangkung Utara District, terminal abdominal segments in lateral view [IMQC]; C. ♀ (PT) from
Tinangkung Utara District, subgenital plate in ventral view [IMQC]; D. ♀ (PT) from Tinangkung Utara District, anal segment
(= tergum X) in dorsal view [IMQC]; E. ♀ (PT) from Tinangkung Utara District, left ala in dorsal view [IMQC]; F. ♀ (PT) from
Tinangkung Utara District, head, pro- and mesothorax in lateral view [IMQC]; G. ♀ (PT) from Tinangkung Utara District, head,
pro- and mesothorax in dorsal view [IMQC]. (Figures B–G © René Limoges, IMQC)
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FIGURE 7. Orthomeria limogesi n. sp. ♂, holotype [IMQC]: A. Habitus in dorsal view; B. Left ala in dorsal view; C. Right
protarsus in dorsal view; D. Head, pro- and mesothorax in lateral view; E. Head, pro- and mesothorax in dorsal view; F. Termi-
nal abdominal segments in lateral view. (© René Limoges, IMQC)
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Head: Wider than long, widest behind eyes, prognathous in lateral view, vertex flat (Fig. 7F); shorter and
somewhat narrower than pronotum (Fig. 7G). Genae slightly shorter than diameter of eye. Eyes very large, almost
circular in outline and projecting hemispherically. Two very shallow impressions on frons between bases of anten-
nae. Gula sclerite very narrow and transverse. Antennae cylindrical, reaching to abdominal segment V and consist-
ing of ca. 62–64 antennomeres. Scapus slightly compressed dorsoventrally, rectangular in dorsal aspect and about
1.3x longer than wide. Pedicellus round in cross-section, slightly constricted towards apex and shorter than scapus.
Antennomere III about equal in length to scapus and pedicellus combined. IV much shorter, following ones first
notably increasing then decreasing in length.
Thorax: Pronotum roughly quadrate, the anterior portion with a longitudinal median groove, convex and raised
on both sides of groove and notably higher than posterior portion (Fig. 7F). Glandular field elliptical, the foramen
slightly anteriad directed and at anterior margin. Posterior margin swollen and almost straight. Mesonotum almost
rectangular (Fig. 7G), 1.7x longer than pronotum and approximately 1.6x longer than wide. Mesosternum flat with
a slight longitudinal median furrow in anterior half, furcasternum with demarcated furca. Metanotum transverse,
with central portion convex and strongly setose. Metasternum with a longitudinal median furrow in anterior half.
Tegmina small, spatulate and folded longitudinally (Figs. 7F–G). Alae reaching to posterior portion of abdominal
segment VI. Radial vein very strong and sclerotized; this as well as anterior and posterior median veins simple and
unbranched (Fig. 7E).
Abdomen: Abdomen excluding median segment 1.15x longer than head and complete thorax combined. Seg-
ments II–V very slightly widening, VI –VII progressively narrowing. II–IV slighty increasing and V–VII decreasing
in length with V longest segment; all somewhat longer than wider and almost rectangular. Tergum IX shorter than
VIII. Anal segment strongly tectiform with a very obtuse longitudinal median bulge; posterior margin very slightly
angular (Fig. 7D). Epiproct very small und wholly conealed under anal segment. Cerci small, round in cross-section,
somewhat tapered in the apical half and very gently in-curving; projecting notably over posterior margin of anal
segment (Fig. 7B). Subgenital plate deeply keeled, and very slightly reaching beyond apex of abdomen; posterior
margin rounded (Fig. 7C).
Legs: Short and fairly stocky; all femora with very indistinct and rounded dorsal carinae, the two outer ventral
carinae more defined but unarmed, or at best with a single sub-apical spine weakly indicated. Tibiae slightly shorter
than corresponding femora and almost circular in cross-section with the two outer ventral carinae just very weakly
indicated. Basitarsus of all tarsi a little longer than following two tarsomeres combined. Ungues pectinate (Fig.
6C).
♂(Figs.6,8D).Of average size for the genus (body length 41.5–46.3 mm), shape very typical but with a com-
paratively small head that is notably smaller than the pronotum. Entire body surface, legs and tarsi densely setose,
colour of setae dull yellow (body) to pale orange (tarsi). General colour plain black, tarsomere V and ungues dull
orange. Antennae fairly bright dull orange except for scapus and pedicellus. Tegmina and alae as in ♀♀ but areas
between veins in costal region of alae anterior to radial vein may be increasingly to almost entirely orange (Fig. 6B)
to red (Fig. 8D).
Head: Generally as in ♀♀ but eyes relatively larger and much more strongly projecting. Length of gena cor-
responding to only about ¾ the diametre of eye (Figs. 6-D–E). Antennae as in ♀♀ and reaching to posterior margin
of abdominal segment VI.
Thorax: Pronotum generally as in ♀♀ but the anterior portion somewhat less swollen (Fig. 6D). Mesonotum
very slightly more elongate than in ♀♀, being almost 1.7x longer than wide with the anterior margin strongly swol-
len and gently V-shaped (Fig. 6E). Longitudinal median furrow of meso- and metasternum very shallow. Tegmina
and alae as in ♀♀, but the tegmina notably larger (Figs. 6D–E) and the alae reaching to posterior margin of abdomi-
nal segment VI (Fig. 8D).
Abdomen: Abdomen excluding median segment 1.25x longer than combined length of head and complete
thorax. Segments II–V almost uniform in length and width, VI and VII progressively shorter and slightly widening.
II–V on average 1.6x longer than wide, VII just indistinctly longer than wide. Anal segment slightly longer than
tergum IX, strongly tectiform and without ventral thorn pads at posterolateral angles; the posterior margin broadly
rounded. Epiproct very small, triangular and fully hidden under anal segment. Vomer acutely triangular, strongly
sclerotized and with a pointed, up-curving terminal hook. Cerci large, almost 1.5x length of anal segement, slightly
compressed laterally and generally bone-shaped in lateral aspect with the median portion constricted and the apex
strongly expanded and angular (Fig. 6F). The lower angle much more expanded than the upper one and obtuse, the
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upper angle bearing a sclerotized and shiny, blade-like ridge. Poculum large, scoop-shaped and projecting beyond
apex of abdomen by about 2/3 the length of anal segment (Fig. 6F).
Legs: As in ♀♀ but averaging more slender. Ungues pectinate (Fig. 6C).
Variability: Females do not show any noteworthy variability, except for a rather remarkable range in body
length (see table 1 below) and slight variability in overall colour. Males in contrast show considerable variability in
the colouration of the costal region of the alae anterior to the radial vein. This is mostly coloured more or less like
the posterior portion, being brown with dull orange veins, but a good number of specimens has the areas between the
veins increasingly orange, particularly towards the base of the wing. In a very few specimens the anterior portion is
almost entirely orange. It shall also be mentioned that on average specimens from Tinanasu are slightly larger than
ones from Luksagu.
FIGURE 8. Orthomeria limogesi n. sp.: A. Life ♂ on the host plant (Macaranga sp., Euphorbiaceae) at the type-locality near
Luksagu village, Tinangkung Utara District, Peleng (© Adhi Brata); B. Life ♀ on the host plant (Macaranga sp., Euphorbiaceae)
at the type-locality near Luksagu village, Tinangkung Utara District, Peleng (© Adhi Brata); C. ♀ (PT) from Tatendeng-Eben,
Buko District, Peleng [coll. FH, No. 0734-27]; D. ♂ (PT) from Tatendeng-Eben, Buko District, Peleng [coll. FH, No. 0734-
30].
STICK INSECTS OF INDONESIA Zootaxa 5073 (1) © 2021 Magnolia Press · 21
Comments: This species is fairly common in lowland regions throughout most of Peleng and can frequently be
found on small trees of an unidentified species, that most likely belongs in the genus Macaranga (Euphorbiaceae).
This tree appears to be the main host plant of Orthomeria limogesi n. sp. (Figs. 8A–B). Eggs not available.
Distribution: Apparently endemic to the Island of Peleng, the largest of the Banggai Islands.
TABLE 1. Measurements of Orthomeria limogesi Hennemann & Le Tirant n. sp. [mm]
HT, ♂
[IMQC]
PT, ♂♂ PT, ♀♀
Body 44.4 40.8–45.0 46.5–55.5
Body (incl. poculum) 45.2 41.5–46.3 -
Pronotum 3.5 3.1–3.5 3.9–4.3
Mesonotum 5.7 5.7–6.2 6.0–7.2
Metanotum 2.6 2.5–2.9 2.7–3.1
Median segment 4.8 3.9–4.8 4.0–4.7
Tegmina 1.1 1.0–1.1 0.8–1.1
Alae 24.9 23.5–25.0 19.8–23.0
Profemora 7.5 6.9–7.5 7.3–8.0
Mesofemora 6.7 6.4–6.9 5.8–6.7
Metafemora 10.2 8.5–10.2 8.8–9.6
Protibiae 6.6 5.8–6.7 5.9–6.7
Mesotibiae 6.0 5.2–5.9 5.0–5.8
Metatibiae 10.4 8.5–9.8 8.9–9.7
Antennae 37.5 34.0–38.0 32.5–36.0
Heteropterygidae Kirby, 1896
Dataminae Rehn & Rehn, 1939
Datamini Rehn & Rehn, 1939
Genus Pylaemenes Stål, 1875
(Figs. 9–11)
Type-species: Phasma (Pachymorpha) coronatum Haan, 1842: 137, pl. 14: 4–5, by subgsequent designation of Kirby, 1904:
400.
Comments: Hennemann (1998: 125) correctly synonymised Datames Stål, 1875 with Pylaemenes Stål, 1875 on the
basis that the type species of both genera are clearly conspecific. This was rejected by Bragg (2001: 162) but all sub-
sequent authors have since accepted the synonymy of Datames. Bresseel & Constant (2018: 7) presented additional
diagnostic features for Pylaemenes, differentiated it from the very closely related Orestes Redtenbacher, 1906 and
transferred several species from Pylaemenes to Orestes. In presenting a catalogue of the Phasmatodea of Sulawesi
Hennemann (1998: 125) furthermore provided a synonymic listing for the type-species P. coronatus (Haan, 1842)
and synonymised two further Wallacean species. Recent examination of extensive material of Pylaemenes at hand
from various localities throughout Wallacea however clearly shows some of the synonymies to be wrong and that at
least the Wallacean taxa of the genus are in need of revision.
Distribution: Wallacea, Java, Borneo, Sumatra, Peninsular Malaysia and Singapore. The three species recorded
from Continental Asia (i.e. Vietnam and South China) deserve evaluation and may not belong in Pylaemenes.
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Pylaemenes moluccanus (Redtenbacher, 1906)
(Figs. 9–10)
Datames moluccanus Redtenbacher, 1906: 51. LT (by present designation), ♂: Coll. Br. v. W, Ins. Kalidupa H. Kühne; 24.846;
det. Redtenb. Datames moluccanus; Datames moluccanus Redt. Kalidupa prope Celebes. Type! [NHMW, No. 47]; PLT, 1
♂, 2 ♀♀: Coll. Br. v. W. Ins. Kalidupa, H. Kühne; det. Redtenb. Datames moluccanus [NHMW, No. 47]; PLT, ♂: Coll. Br.
v. W. Ins. Kalidupa bei Celebes, H. Kühne; det. Redtenb. Datames moluccanus; Kalidupa [NHMW, No. 47].
Brock, 1998: 43.
Pylaemenes moluccanus, Zompro, 2004: 226
Otte & Brock, 2005: 297.
Datames oileus, Günther, 1938: 57, 60 [Misidentification; specimens from Sulawesi in NHMB].
Material examined: 1 ♂, 1 ♀ (in copula): Luwu, Lembongpangi, c. 500 m, 6 II. 95; Sarasin, 6.II.1895, Lembong-
pangi (Luwu), ca. 500 m, Centr.- Cel.; Pylaemenes coronatus de Haan. K. Günther det. [NHMB].
Comments: Examination of the two specimens in NHMB recorded by Günther (1938: 60) has shown these to
have been misidentified (Fig. 10). They match best with the five type specimens of P. moluccanus (Redtenbacher,
1906) in NHMW (Fig. 9), that originate from Pulau Kaledupa, an island of the Tukangbesi Islands, Wakatobi Re-
gency, an archipelago to the southeast of Sulawesi (Fig. 3). The two specimens however differ from the typical
examples of P. moluccanus by the smaller size, somewhat less stocky shape and averaging less pronounced body
armature and sculpturing. In comparison, all the spines, tubercles and swellings of the body and legs are mostly
identical but smaller and less acute. In addition to these two specimens, the author has seen pictures of a ♀ from
Mamasa, Central Sulawesi which also matches well with the types of P. moluccanus but in respect of the size and
degree of body armature and sculpturing is intermediate between the specimens in NHMB from Central Sulawesi
and the types of P. moluccanus from Kaledupa. The ♂ syntype which bears a collection number and a handwrit-
ten label by Redtenbacher stating “Type!” is here selected as the lectotype of Datames moluccanus to guarantee
stability of the name (Figs. 9C–D). This is in accordance with Article 74 of the ICZN Code (ICZN, 1999). P. oileus
(Westwood, 1859) differs from these Sulawesian specimens in numerous aspects, e.g. the armature of the head and
shape of the anal segment.
Redtenbacher (1906: 51) cited a body length of 58.0–60.0 mm for ♀♀ and 50.0–52.0 mm for ♂♂ (shortest body
length refers to lectotype). For comparison the measurements of the notably smaller specimens from NHMB are
presented in table 2 below. Illustrations of the type specimens and the couple in copula from NHMB are presented
to illustrate the mentioned differences and support future work on the arrangement of Wallacean representatives of
Pylaemenes.
TABLE 2. Measurements of Pylaemenes moluccanus (Redtenbacher, 1906) from Sulawesi in NHMB [mm]
♀
[NHMB]
♂
[NHMB]
Body 52.8 47.5
Pronotum 3.2 3.5
Mesonotum 12.4 10.8
Metanotum 5.1 4.9
Median segment 1.8 1.5
Profemora 9.1 8.8
Mesofemora 8.2 7.4
Metafemora 8.4 8.9
Protibiae 8.9 8.5
Mesotibiae 6.7 6.8
Metatibiae 9.9 8.9
Antennae > 13.0 ca. 21.0
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FIGURE 9. Pylaemenes moluccanus (Redtenbacher, 1906), type specimens: A. ♀ (PLT) frm Kaledupa, dorsal view [NHMW];
B. ♀ (PLT) from Kaledupa, lateral view [NHMW]; C. ♂ (PLT) from Kaledupa, dorsal view [NHMW, No. 24.846]; D. ♂ (PLT)
from Kaledupa, lateraal view [NHMW, No. 24.846]. © 2019 Natural History Museum Vienna, NOaS Image Collection / H.
Bruckner.
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FIGURE 10. Pylaemenes moluccanus var. (Redtenbacher, 1906), ♂ and ♀ in copula from Luwu, Lembongpangi, Central Su-
lawesi [NHMB]: A. ♂/♀ in dorsal view; B. ♂/♀ in ventral view; C. ♂/♀ in lateral view; D. Terminal abdominal segments of ♀
in lateral view; E. Terminal abdominal segments of ♀ in dorsal view; F. Terminal abdominal segments of ♂ in dorsal view.
STICK INSECTS OF INDONESIA Zootaxa 5073 (1) © 2021 Magnolia Press · 25
Pylaemenes pleurospinosus Hennemann & Le Tirant n. sp.
(Fig. 11)
HT, ♀: Indonesia, Peleng Island, Tinangkung Utara District, near Luksagu village ca. 60 m elev., 1°17’S 123°25.4’ E, VII.2012
[IMQC, ex coll. FH, No. 1147-1].
PT, ♀ (n5): Indonesia, Peleng Island, Tinangkung Utara District, near Luksagu village ca. 60 m elev., 1°17’S 123°25.4’ E,
VII.2012 [coll. FH, No. 1147-2].
Etymology: The name refers to the prominent and long supra-coxal spine of the mesopleurae, which readily distin-
guishes this new species from all other Wallacean representatives of the genus.
Differential diagnosis: Very similar to the type-species P. coronatus (Haan, 1842) but at once differing from
this and all other species of Pylaemenes so far known to occur throughout Wallacea by the presence of a prominent
supracoxal spine on the mesopleurae (Figs. 11F–G). Females, the only sex known furthermore differ from the other
Wallacean species by having the supra-coxal spine of the metapleurae notably more prominent and the mesonotum
relatively more slender with the lateral margins very gently concave and at the anterolateral angle armed with a
prominent, slender laterad directed antero-lateral mesonotal spine (Fig. 11G).
Description: The description is based exclusively on the unique holotype. The nomenclature used for body ar-
mature follows that used for the Heteropteryginae Kirby, 1896 by Hennemann et al. (2016) and that for the cephalic
armature follows the nomenclature used for the genus Orestes Redtenbacher, 1906 by Bresseel & Constant (2018:
6, fig. 1).
♀(Fig.11). Fairly large (body length 59.0 mm) and moderately stocky for the genus with strongly developed
cephalic and body armature and prominent supracoxal spines on the meso- and metapleurae. As typical for the
genus, entire dorsal body surface tectinate longitudinally. General colour of the HT greyish and ochraceous mid
brown, the lateral margins of the thoracic terga and the base of the large supracoxal spines with a slight pale green
wash. All major spines tipped with dull ochre. Abdominal tergum VI with two velvety black, roundly angular central
markings. Eyes dull orange.
Head: Slightly longer than wide with the vertex strongly raised and posteriorly somewhat extending over ante-
rior portion of pronotum, the genae very slightly widening towards the posterior. Frons with a very prominent and
long, somewhat anteriad directed pair of supra-antennals, the anterior and posterior supra-occipitals much smaller
and bluntly rounded; the posterior pair smaller than the anterior pair. Vertex with two converging cristate and un-
evenly tri-tuberculate ridges, each of which terminate in a prominent, conical but blunt anterior coronal spine. The
supra-orbitals large, similar in size to anterior-coronals and positioned close to the aforementioned ridge (Fig. 11F).
The central coronal prominent and bluntly conical; on each side a much smaller, obtuse lateral coronal tubercle. The
two posterior coronals prominent, conical and similar in size to the anterior coronals; the lateral coronals at posterior
margin of head notably smaller than the median pair. Postocular ridge very prominent, unevenly and obtusely crenu-
late and posteriorly terminating in a blunt tubercle. Eyes very small, projecting sub-spherically and their diameter
contained about 3.2x in length of genae (Fig. 11G). Antennae consisting of 20 segments and notably projecting over
apex of protarsi. Scapus longer than wide, somewhat widening towards the base and the outer margin armed with a
fairly acute, lateral directed spine at the base and a more blunt, anteriad directed spine at the apex (Fig. 11F). Anten-
nomere III distinctly elongated and longer than pedicellus and all following antennomeres.
Thorax: Pronotum notably wider and slightly longer than head, roughly quadrate in shape with lateral margins
somewhat concave (Fig. 11G). The transverse median sulcus strongly impressed, straight and short. In front of the
sulcus a very strong pair of bluntly conical pre-median pronotals and just behind it a small but fairly acute pair of
post-medial pronotals. The anterior-mesal pronotals small, the inter-posterior pronotals moderate and rather obtuse.
The antero and postero-lateral pronotals at the outer angles of the pronotum weakly developed and blunt. Sensory
areas of the prosternum and profurcasternum well developed. Mesothorax moderately slender and 3.5x longer than
pronotum, the mesonotum about 2.7x longer than wide and roughly rectangular with the lateral margins very weakly
concave (Fig. 11G). Dorsal surface with a prominent longitudinal median carina, which bears a small pair of closely
placed pre-median tubercles and a very small pair of post-median tubercles (Fig. 11F). The anterior-mesal pronotal
spines at the raised anterior margin very prominent, upright and pointed. The posterior-mesal pronotals notably
smaller than the anterior ones. Surface otherwise very spasely and obtusely tuberclose. Lateral margins distinct
and set with about six spiniform tubercles, which notably decrease in size towards the posterior; the antero-lateral
mesonotal very prominent, long and pointed. Mesopleurae with a longitudinal row of about five blunt tubercles and
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two supra-coxal spines; the posterior spine very prominent, slender, acutely pointed and strongly projecting later-
ally (Fig. 11G). Metanotum less than half the length of mesonotum, about 1.4x longer than wide and with lateral
margins notably concave; sculptured like mesonotum but only with a fairly small pair of post-median tubercles and
the lateral margins with five equally sized blunt tubercles. Metapleurae set with three spiniform lateral tubercles
and the same two supra-coxals seen on the mesopleurae; the posterior supra-coxal however even more prominent,
comparatively stronger and laterally projecting by almost half the width of metanotum (Figs. 11A, C). Meso- and
metasternum with a very faint longutudinal median carina and set with a few indistinct and low paired tubercles
(Fig. 11C).
Abdomen: Median segment transverse with anterior margin widely rounded; the median keel forming an obtuse
swelling anteriorly and with two small tubercles posteriorly. Segments IIII–III widening, IV widest, V–VI narrow-
ing, VII narrowest, VIII widening; all transverse and rectangular in outline, IV about 2.6x wider than long. II–VIII
with a prominent, tectinate median keel, which is most prominent on V and bears a small pair of median tubercles;
near posterior margin with posteromedian pair of spines which is most indistinct on II and most prominent on V.
II–VIII with 3–4 small tubercles along lateral margins and each armed with a prominent, straight, lateral directed
and acutely pointed posterolateral spine. Median carina on tergum IX strongly raised and forming a large, irregu-
larly shaped crista that posteriorly projects over the anterior portion of the anal segment; lateral margins somewhat
deflexed and broadly triangular in outline. Sterna II–VII weakly tectinate longitudinally and with a few scattered,
low tubercles. Preopercular organ on sternum VII formed by an obtuse, transverse, scale-like expansion of the
posterior margin. Anal segment much narrower than all preceding terga, strongly narrowing towards the posterior
and with a blunt longitudinal median carina; the lateral margins with a bluntly triangular expansion in anterior half
and the apex bifid with a distinct triangular median excavation (Fig. 11D). Epiproct small, scale-like and concealed
under anal segment (Fig. 11E). Subgenital plate moderately convex, carinate medio-longitudinally, not reaching
apex of anal segment and the posterior margin broadly rounded (Fig. 11E); the surface unevenly rugulose and with
a single spiniform central tubercle on each lateral surface.
Legs: All moderately long and stocky, all tibiae unarmed. Profemora with a single, blunt sub-apical tooth on
posteroventral carina, otherwise unarmed. Meso- and metafemora with a two sub-apical teeth on the two outer ven-
tral carinae of which the apical one is notably larger; furthermore with two blunt teeth in basal half of posterodorsal
carina.
Comments: Males and eggs unknown. The immature ♀ paratype has a body length of 42.0 mm and well ex-
hibits the characteristic supracoxal spines of this species.
Distribution: Apparently endemic to the Island of Peleng, the largest of the Banggai Islands positioned east of
Sulawesi.
TABLE 3. Measurements of Pylaemenes pleurospinosus Hennemann & Le Tirant n. sp. [mm]
HT, ♀
[IMQC]
Body 59.0
Pronotum 4.4
Mesonotum 12.8
Metanotum 6.5
Median segment 2.1
Profemora 10.0
Mesofemora 8.9
Metafemora 11.0
Protibiae 9.8
Mesotibiae 8.1
Metatibiae 10.8
Antennae 22.5
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FIGURE 11. Pylaemenes pleurospinosus n. sp. ♀, holotype [IMQC]: A. Habitus, dorsal view; B. Habitus lateral view; C. Habi-
tus ventral view; D. Terminal abdominal segments in dorsal view; E. Terminal abdominal segments in ventral view; F. Head,
pro- and mesothorax in dorso-lateral view; G. Head, pro- and mesothorax in dorsal view.
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Lonchodidae Brunner v. Wattenwyl, 1893
Lonchodinae Brunner v. Wattenwyl, 1893
Lonchodini Brunner v. Wattenwyl, 1893
Genus Gibbopromachus n. gen.
(Fig. 12)
Type-species: Pericentrus tripinnatus Redtenbacher, 1998: 352, by present designation.
Pericentrus Redtenbacher, 1908: 352 (in part).
Brock, 1998: 63.
Hennemann, 1998: 119.
Otte & Brock, 2005: 259.
Neopromachus, Günther, 1938: 58, 82 (in part).
Diagnosis: ♀ (Fig.12). Fairly small (body length 78.0 mm) and stocky Lonchodinae with very prominent body
armature, a strongly swollen mesothorax and a bird-beak like ovipositor. General colour brown with a reddish
longitudinal median stripe along most of dorsal body surface. Head and entire body armed with large and strong
but obtuse spines of variable sizes (Figs. 12A–B). Head slightly longer than wide, globose with the vertex roundly
convex and armed with several prominent coronal spines; another strong pair of spines on frons. Antennae slender,
reaching to abdominal segment II and consisting of about 35 antennomeres; scapus compressed dorsoventrally and
somewhat deflexed laterally, pedicellus cylindrical, following increasing in length, the antennomeres in median sec-
tion of antennae extremely elongated, the apical ten or so suddenly much shorter. Antennomere XIV with a shiny,
knob-like sub-basal dorsal swelling; the elongated median antennomeres with apex thickened and gently curved.
Pronotum notably shorter and narrower than head with a prominent, impressed transverse sulcus and at least a pair
of strong spines in posterior half. Mesothorax almost 4x longer than prothorax, gently down-curving, very strongly
swollen medially with the mesonotum very prominently raised and humped (Figs. 12A–B). Mesonotum armed with
numerous very strong spines, mesopleurae with a marginal row of strong spines and a strong supra-coxal spine.
Metanotum roughly 1/3 the length of mesonotum, about 1.6x longer than wide, almost rectangular and almost 2x
longer than median segment; armed with two paired spines. Metapleurae principally like mesopleurae. Meso- and
metasternum with several strong but short, paired spines. Abdomen longer than head and thorax combined. Median
segment somewhat transverse. Segments II–VI decreasing, VII and VIII increasing in length. Terga II and VII with
four spines, III–VII with the lateral margins deflexed into a large, almost semi-circular lobe and armed with six
spines (Fig. 12A). All terga with a fine, longitudinal median carina. Sterna with paired spines, VII with Preopercular
organ formed by two obtusely rounded swellings (Fig. 12E). Terga VIII and IX with an obtuse posteromedian hump.
Anal segment strongly narrowed and declining in lateral aspect (Fig. 12C), the apex strongly elongated into a very
long and straight, lanceolate projection (apex broken in the unique holotype, Fig. 12D). Cerci very small, conical
and gently compressed dorsoventrally. Epiproct very small, triangular and also concealed under apical projection of
anal segment. Subgenital plate convex, strongly keeled longitudinally, bulgy in median portion (Fig. 12 C) and with
apex elongated into a lanceolate projection, that is not reaching the apex of the anal segment; apex obtusely angu-
lar (Fig. 12E). Legs long and moderately slender, profemora a little longer than mesothorax, mesofemora slightly
shorter than mesothorax and metafemora reaching to abdominal segment VI. Profemora strongly compressed and
curved basally. All femora trapezoidal in cross-section, distinctly carinate and on two outer carinae and posterodor-
sal carina with a prominent triangular lobe sub-apically. Medioventral carina obtuse. All tibiae slender and longer
than corresponding femora, unarmed except for a distinct, rounded sub-basal lobe on medioventral carina. Basitarsi
long and slender, somewhat longer than combined length of all remaining tarsomeres.
Egg (Figs. 12F–G): The unique egg available from the ovipositor of the holotype of G. tripinnatus (Redten-
bacher, 1908) n. comb. lacks the operculum. Moderately sized (length 3.5 mm), ovoid about 1.7x longer than wide,
notably higher than wide and oval in cross-section; the polar end slightly narrowed. Capsule surface minutely and
evenly punctured, slightly glossy. General colour flecked with dull ochre and mid brown. Micropylar plate very
elongate with anterior half parallel-sided and posterior half gently widened; covering more than ¾ the length of cap-
sule. Micropylar cup small but well defined and placed some distance off posterior end in a narrow posteromedian
gap of plate. Median line almost obsolete.
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FIGURE 12. Gibbopromachus tripinnatus (Redtenbacher, 1908) n. gen., n. comb., A–E ♀, holotype [NHMW], F–G egg
extracted from ovipositor of holotype (operclum missing) [NHMW]: A. Habitus in dorsal view; B. Habitus in lateral view; C.
Terminal abdominal segments in lateral view; D. Terminal abdominal segments in dorsal view; E. Terminal abdominal segments
in ventral view; F. Egg, dorsal view; G. Egg, lateral view. © 2019 Natural History Museum Vienna, NOaS Image Collection /
H. Bruckner.
Differential diagnosis: Numerous morphological characters suggest close relation to the principally New
Guinean genus Neopromachus Giglio-Tos, 1912, which also has a few representatives on the very eastern islands
of Wallacea (e.g. Aru Islands) but east of Weber’s line; this is mainly the prominent spination of the head and body,
as well as the bird-beak like secondary ovipositor of ♀♀. Females of this new genus, the only sex known, however
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30 · Zootaxa 5073 (1) © 2021 Magnolia Press
differ from those of Neopromachus by the laterally lobed abdominal terga II–VII, strongly swollen and gibbose me-
sonotum, angular apex of the subgenital plate (Fig. 12E), considerably longer basitarsi that are longer than the cor-
responding remaining tarsomeres taken together and extremely elongated median antennomeres. Moreover, the egg
is more ovoid and notably less elongate than all known eggs of Neopromachus. Within the Sulawesian fauna, the
beak-like ovipositor resembles Paramanduria n. gen., but the strong body armature, more globose head, strongly
swollen mesonotum and lateral lobes of the abdominal terga readily distinguish this new genus. The strong body
armature resembles Acanthomenexenus Brock & Hennemann, 2009, but Gibbopromachus n. gen. readily differs
from that genus by the morphology of the terminalia.
Comments: The true relationships of this striking new genus are still not fully known and require knowledge
of the still unknown ♂♂. Redtenbacher (1908: 352) originally placed this species in the genus Pericentrus Redten-
bacher, 1908, which however is not closely related and geographically restricted to northern India and Bhutan.
A discussion of and redescription of Pericentrus was presented by Hennemann, Conle & Zhang (2008: 51), who
already assumed close relation to Neopromachus Giglio-Tos, 1912.
Etymology: The name is a combination of “gibbus” (lat. = hump) and “promachus” from the visually very
similar looking genus Neopromachus Giglio-Tos, 1912. Masculine.
Distribution: Sulawesi (endemic).
Species included:
1. Gibbopromachus tripinnatus (Redtenbacher, 1908: 352) [Pericentrus]. HT, ♀: Coll. Br. v. W., Celebes, Dr.
Sarasin; Lowu Mtne c. 700 m; Pericentrus 3-pinnatus Redt.; det. Redtenb. Pericentrus tripinnatus [NHMW,
No. 713]. n. comb.
Distribution: NE-Sulawesi, Prov. Sulawesi Utara, Mount Lowu 700m [NHMW]; NE-Sulawesi, Minahasa,
Gunung Sudara summit [NHMB].
Genus Hermagoras Stål, 1875
(Figs. 13, 68A)
Type-species: Lonchodes personatus Bates, 1865: 336, pl. 44: 7 [= H. foliopeda (Olivier, 1792)], by subsequent designation of
Kirby, 1904: 322.
Comments: This genus has five known species on Borneo and is represented throughout the southeastern regions of
Wallacea by the type-species H. foliopeda (Olivier, 1792). On Sulawesi it is represented by one species, which was
formerly regarded a subspecies of H. foliopeda but is here raised to species level. The distinction between Herma-
goras and the morphologically very similar Mnesilochus Stål, 1877 presented by Seow-Choen (2016: 279) is partly
based on a misinterpreted character and is clarified in the comments on Mnesilochus below.
Hermagoras celebensis Hennemann, 1998 n. stat.
(Figs. 13, 68A)
Hermagoras foliopeda celebensis Hennemann, 1998: 100, fig. 6, pl. 3: 8–10. HT, ♂: 7916; Holotypus; Hermagoras foliopeda
celebensis Hennemann, 1998, det. F. Hennemann I.1997 ♂; S-Sulawesi, Lembang, Maros, leg. Gunawan 12.1995 [MNHU];
AT, ♀: 7917; Holotypus; Hermagoras foliopeda celebensis Hennemann, 1998, det. F. Hennemann I.1997 ♀; S-Sulawesi,
Lembang, Maros, leg. Gunawan 12.1995 [MNHU]; PT, ♂: Sarasin, 28.I.–1.II.1895, Luwu, Flachland, Boran-Djaladja,
Centr.-Cel.; Lonchodes haematomus K. Günther det. [NHMB, No. VI.D.133].
Hennemann & Conle, 1999: 10. (Catalogued)
Otte & Brock, 2005: 153.
Zompro, 2005: 262. (Catalogued)
Henneman & Conle, 2007: 23.
Comments: Careful re-examination of the type-specimens and comparison with extensive material of the nomi-
nate form of H. foliopeda (Olivier, 1792) from several islands of eastern Wallacea (e.g. Seram, Buru, Ambelau,
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FIGURE 13. Hermagoras celebensis (Hennemann, 1998) n. stat.: A. ♀ (AT), lateral view [MNHU, No. 7917]; B. ♀ (AT),dorsal
view [MNHU, No. 7917]; C. ♂ (HT), lateral view [MNHU, No. 7916]; D. ♂ (HT), lateral view [MNHU, No. 7916]; E. ♂ from
Luwu, dorso-lateral view [NHMB]; F. Apical half of abdomen of ♂ from Luwu, lateral view [NHMB]; G. Terminal abdominal
segments of ♂ from Luwu, dorsal view [NHMB]; H. Terminal abdominal segments of ♂ (HT), lateral view [MNHU, No. 7916];
J. Metanotum and median segment of ♂ from Luwu, dorso-lateral view [NHMB].
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Kei Islands and Waigeo in the Raja Ampat archipelago) at hand from the author‘s collection and evaluation of spe-
cific distinctive characters amongst various species of Hermagoras show the diagnostic characters of this taxon,
along with its isolated western distribution on Sulawesi, to justify raising it to species level. Hence, this Sulawesian
form now becomes H. celebensis Hennemann, 1998 (rev. stat.). Re-examination of the ♀ from Luwu listed along
with the ♂ paratype of H. celebensis in NHMB by Günther (1938: 78) as “Lonchodes haematomus”, clearly shows
the specimen is not conspecific and moreover not a member of Hermagoras. It is here described as a new species
(→ see Mnesilochus luwuense n. sp. below).
A ♂ from Bogani Nani Wartabone National Park, Gorontalo province was examined from a photograph taken
by Pavel Kirillov in December 2012 and is the first record of H. celebensis from northern Sulawesi (Fig. 68A).
Differentiation: Since ♀♀ of H. foliopeda are remarkably polymorphic and show extreme intraspecific vari-
ability (see Hennemann & Conle, 1997), likely differences between ♀♀ are dificult to define with only a unique
♀ of H. celebensis known to date (Figs. 13A–B). Males however show very obvious and constant distinguishing
characters and in contrast to ♀♀ of Hermagoras do not show any noteworthy intraspecific variability. The distinc-
tive black spiniform tubercles near the posterior margin of the metanotum (Fig. 13J), pale granulation of the thorax,
contrasting darker median sections of the meso- and metathorax (colour plain in foliopeda), the pale cream or grey
and black apex of the femora as well as the more slender and apically pointed hemi-tergites of the anal segment (Fig.
13H) readily distinguish ♂♂ from those of H. foliopeda.
Distribution: S-Sulawesi, Prov. Selatan (Lembang, Maros); N-Sulawesi, Prov. Gorontalo (Bogani Nani Warta-
bone N.P.).
Genus Leprocaulinus Uvarov, 1940
(Figs. 14–16)
Type-species: Leprocaulus altecornutus Redtenbacher, 1908: 473, by subsequent designation of Hennemann, 1998: 121.
Comments: This genus appears to be paraphyletic in its present recognition, because it is likely that several species
from Sundaland currently attributed to Carausius Stål, 1875 are congeneric and might prove to be members of Lep-
rocaulinus once a more extensive study of the concerned taxa is conducted. A morphological distinction between
Carausius and Leprocaulinus is difficult. The length relation of the median segment and metanotum traditionally
used to distinguish ♀♀ of the two genera proves not very useful and the three Sulawesian species covered herein,
(two of which are newly described), show that the genus also has apterous ♂♂, a fact not recognised previously.
Furthermore, there is considerable confusion concerning the validity of the species and numerous subspecies cur-
rently attributed to the genus. A re-organisation has been attempted by Günther (1934b: 81) but the author was not
able to examine all necessary type-material in detail and ♀♀ in particular show strong intraspecific variability.
Hence, the genus is in need of a detailed revision at the species level.
Distribution: New Guinea, Thursday Island and Wallacea.
Species recorded from Wallacea:
1. Leprocaulinus digitatus n. sp.
Distribution: Central Sulawesi, Prov. Sulawesi Selatan, Luwu [NHMB].
2. Leprocaulinus heinrichi (Günther, 1935: 6, pl. 1: 4 & 5) [Carausius]. HT, ♂: S. Celebes, Lompobatang, Wawa
Karaeng 2000 m, Sept. 1931 G. Heinrich; Typus [MNHU]; AT, ♀: S. Celebes, Lompobatang, Wawa Karaeng
2000 m, Sept. 1931 G. Heinrich; Typus [MNHU]; PT, 1 ♀, 5 ♂♂: S. Celebes, Lompobatang, Wawa Karaeng
2000 m, Sept. 1931 G. Heinrich; Typus [MNHU]. n. comb.
Comments: Günther (1935: 6) listed six ♂ paratypes, one of which appears lost and could not be traced.
Distribution: S-Sulawesi, Prov. Sulawesi Selatan, Gunung Lompobatang and Gunung Bawakaraeng “Wawa
Karaeng” [MNHU].
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3. Leprocaulinus insularis talaudiensis (Günther, 1934b: 79) [Leprocaulus]. ST, 2 ♀♀: Talaud, Liroeng, Salibaboe,
Erie, V. 1926 [MNHU]; ST, 1 ♂, 2 ♀♀: Talaud, Liroeng, Salibaboe, Erie, V. 1926 [SMTD]; 1 ♂, 5 ♀♀: Talaud,
Liroeng, Salibaboe, Erie, V. 1926 [RMNH].
Comments: The untraced specimens of the type-series (2 ♂♂, 23 ♀♀ in total) were thought to be in the Mu-
seum Zoologicum Bogoriense, Cibinong, Indonesia (MBBJ) by Zompro (2005: 268). The subspecies status of
Günther‘s talaudiensis under L. insularis Kirby, 1896 is very doubtful and requires evaluation. Examination of
the type specimens of both taxa rather suggest talaudiensis as a valid specie, but without presenting a revision
of the entire genus any decisions appear pre-mature at this point.
Distribution: Talaud Islands, Pulau Salebabu, Lirung [MNHU, SMTD, RMNH].
4. Leprocaulinus kaupii (Stål, 1875: 63) [Myronides]. HT, ♂: Coll. Br. v. W., Molukken, Depuiset ded.; det. Br.v.
W. Myronides kaupii Stål; 5023 [NHMW, No. 479]. n. comb.
Comments: The very slender shape, relatively long abdomen and broad hemi-tergites of the anal segment place
this species in Leprocaulinus Uvarov, 1940, hence it is here removed from Myronides Stål, 1875.
Distribution: „Maluku Islands“ [NHMW].
5. Leprocaulinus mammatus (Rehn, 1904: 44) [Carausius]. HT, ♀: Obi Island, Moluccas; Carausius mammatus
Rehn, Type No. 5143; Carausius mammatus Rehn [ANSP]. rev. stat., n. comb.
Comments: Examination of the HT clearly places this species in Leprocaulinus and proves it to be not a syn-
onym of L. vipera bituber (Sharp, 1898), a taxon described from New Britain. Instead, L. mammatus appears to
be closely related to L. vipera vipera (Kaup, 1871) and L. obiensis (Rehn, 1904) and is here re-established as a
valid species. Characters that distinguish it from the latter have been presented by Rehn (1904: 46) and a proper
distinction from the nominate form of L. vipera (Kaup, 1871), originally described from ♂♂ from Sulawesi,
would need material of both sexes.The subspecies and numerous synonymies currently attributed to Kaup’s
vipera are very doubtful and require critical evaluation.
Distribution: Maluku Islands, Obi [ANSP].
6. Leprocaulinus obiensis (Rehn, 1904: 46) [Carausius]. HT, ♀ (penultimate instar): Obi Island, Moluccas;
Carausius obiensis Rehn, Type No. 5142 [ANSP].
= Carausius bilobulatus Brunner v. Wattenwyl, 1907: 273. HT, ♀ (nymph): “Ins. Moluccae” [MNCN, not
traced]. [Synonymised by Günther, 1934b: 84]
= Diardia modesta Redtenbacher, 1908: 485. ST, ♂: Moluques, Obi Major, Waterstadt 1902; type! Diardia
modesta Br.; Sintipo; MNCN Cat. Tipos N° 7623 [MNCN]; ST, ♂: Nouv.-Guinée, Dorey, Raffray & Maindron,
78, 102; Type; Diardia modesta Redt. Type [MNHN]. [Synonymised by Günther, 1934b: 84]
= Carausius stultus Brunner v. Wattenwyl, 1907: 273. HT, ♀: “Ins. Moluccae” [MNCN, not traced].
[Synonymised by Günther, 1934b: 84]
Distribution: Maluku Islands, Obi [ANSP, MNCN, MNHN].
7. Leprocaulinus sulawesiense n. sp.
Distribution: S-Sulawesi, Prov. Sulawesi Selatan, Gunung Latimojong, Rantemario, Uru 800 m [MNHU];
S-Sulawesi, Prov. Sulawesi Selatan, Rantepao Palopo km12, ca. 1100 m [coll. FH]; S-Sulawesi, Prov. Su-
lawesi Selatan, Tana Toraja, 700 m [coll. FH]; SE-Sulawesi, Mengkoka Mountains, Gunung Tanke Salokko
[MNHU].
8. Leprocaulinus vipera vipera (Kaup, 1871: 39) [Necroscia]. LT, ♂: Necroscia vipera Kp. Celebes [HLDH];
PLT, ♂: Necroscia vipera Kp. Celebes [HLDH, not traced].
Comments: Unfortunately, no fresh material of this species has so far become available for properly re-describ-
ing and distinguishing it from closely related taxa. Furthermore, the ♂ LT in HLDH is incomplete and lacks the
apex of the abdomen, which adds to the current difficulties concerning the taxonomy of Leprocaulinus of Wal-
lacea. The three subspecies and six synonyms currently listed under L. vipera are very doubtful. One of these,
C. mammatus (Rehn, 1904), is here removed from synonymy under L. vipera and reinstated as a valid species.
Distribution: “Sulawesi” [HLDH].
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Leprocaulinus digitatus n. sp.
(Fig. 14)
Carausius heinrichi, Günther, 1938: 81 (in part).
Carausius immundus, Günther, 1938: 58, 81 (in part).
HT, ♀: Sarasin—Luwu, Central-Celebes; Carausius immundus Br. K. Günther det.; Carausius sp. cf. K. Günther, Mitt. Zool.
Mus. Berlin XX, 1935, p. 8. K. Günther det. [NHMB].
PT, ♀: Luwu +; Sarasin—Luwu, Central-Celebes; Carausius immundus Br. K. Günther det.; Carausius sp. cf. K. Günther, Mitt.
Zool. Mus. Berlin XX, 1935, p. 8. K. Günther det. [NHMB].
PT, ♀(penultimate instar): Sarasin 28.I.–1.II.1895, Luwu, Flachland, Boran Djaladja, Centr. Cel.; Carausius heinrichi K. Gthr.
K. Günther det. [NHMB].
Etymology: The name (digitatus lat. = with fingers) refers to the long and apically bifid, digitiform projecting pre-
opercular organ of ♀♀ of this new species.
Differential diagnosis: Females (the only sex known) of this new species are very similar to those of L. sulawe-
siense n. sp. but differ by the very prominent Preopercular organ on abdominal sternum VII, which is formed by a
long, digitiform and apically bifid projection (Figs. 14D, F), as well as the somewhat more prominent cephalad pair
of spines.
Description:♀(Figs.14A–C).Medium-sized for the genus (body length 132.5–149.0 mm), form fairly slen-
der with a short median segment and only a pair of low swellings between the eyes. Body surface unevenly tuber-
culose and granulose, sculpturing variable and occasionally with scale-like swellings and excrescences of variable
sizes on abdominal terga I–VII. Colour ranging from buff to greyish brown; occasionally flecked with ochre and in
the adult paratype with some irregularly scattered white dots on the thorax and four basal abdominal segments (Fig.
14C). Large tubercles of the head and thorax pale ochre or black. Antennae dull ochre with a few faint dark brown
annulae. Eyes dark brown.
Head: About 1.25x longer than wide, subcylindrical with the genae indistinctly narrowing towards posterior.
Frons with two small but deep transverse impressions between bases of antennae. Between the eyes with a slightly
swollen transverse area that bears two fairly low and obtusely conical swellings; posteriorly this raised area is
bordered by a V-shaped furrow, that expands almost from one eye to the other. Vertex flat and with a prominently
impressed coronal line, otherwise set with a few scattered nodes and small tubercles, genae usually with 2–3 obtuse
nodes. Posterior margin with a notably enlarged pair of obtuse median swellings. Eyes roughly circular in outline,
moderately projecting and their diametre contained almost 2.5x in length of genae. Antennae reaching to anterior
margin of median segment. Scapus compressed dorsoventrally, somewhat deflexed laterally oval in outline and 1.7x
longer than wide. Pedicellus oval in cross-section and a little more than 1/3 the length of scapus.
Thorax: Pronotum of similar dimensions as head, with the anterior portion somewhat expanded, the lateral
margins notably concave and about 1.4x longer than width of anterior margin. Transverse median sulcus distinct,
weakly curved and almost expanding over entire width of segment; anterior margin with a median pair of blunt
tubercles and some further paired tubercles on dorsal surface. Mesothorax elongate, slender and about 5.3x longer
than prothorax. Mesonotum with a fine but distinct longitudinal median carina and unevenly set with tubercles and
nodes of variable sizes. Mesopleurae with a longitudinal median row of rather small nodes; mesosternum acutely
keeled medio-longitudinally but otherwise smooth except for a few indistinct granules in posterior portion (Fig.
14G). Metanotum half the length of mesonotum and about 4x longer than wide; sculpturing less pronounced than
on mesonotum. Metapleurae and sternum sparsely nodulose.
Abdomen: Median segment about 2x longer than wide, somewhat less than half the length of metanotum and
gently constricted medially; sculpturing alike and occasionally with a transverse scale-like crest or swelling at pos-
terior margin. Abdomen excluding median segment equal in length to head and complete thorax combined; entire
dorsal surface with a fine and acute longitudinal median carina and irregularly granulose to tuberculose. Occasion-
ally each with a posteromedian swelling or scale-like excrescence which is most prominent on tergum VI. Segment
II 1.6x longer than median segment, II–VI uniform in width, II–III and IV–V equal in length, on average some
2.7x longer than wide. Tergum VI usually somewhat swollen post-medially and VII shorter and slightly narrower
than all previous. Sterna sparsely and irregularly tuberculose and slightly rugulose. Preopercular organ formed by
a long, digitiform and apically bifid projection, which is posterior directed and placed in a small distance before
the posterior margin of sternum VII (Figs. 14D, F). Tergum VIII half as long as VII and slightly widening towards
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the posterior, IX shorter and slightly transverse in dorsal aspect. Anal segment about as long as IX, strongly convex
longitudinally and somewhat narrowed posteriorly, with the posterior margin widely triangularly excavated and
the outer angles obtusely angular (Fig. 14E); the lateral margins slightly deflexed in the basal portion (Fig. 14F).
Epiproct distinct, somewhat variable in length and shape, usually wider than long and projecting notably behind
apex of anal segment; roughly triangular in outline (Fig. 14E). Cerci small, compressed laterally and conical with
the apex acute. Subgenital plate strongly keeled medio-longitudinally, convex, bulgy and more or less angular in the
median portion (Fig. 14F); the posterior margin narrowed and projecting slightly beyond apex of abdomen (Figs.
14D–F).
Legs: All moderately long and slender, the profemora about as long as mesothorax, metafemora reaching about
half way along abdominal segment IV and metatibiae reaching to abdominal segment VII. Anterodorsal carina of
profemora moderately deflexed and more or less undulate and wavy in the basal one third; the posteroventral carina
with two obtuse sub-apical teeth. The dorsal carina of the protibiae strongly but almost uniformly deflexed and la-
mellate. Meso- and metafemora slender, laterally compressed and unarmed except for two obtuse sub-apical teeth
on the two outer ventral carinae (Fig. 14G). Medioventral carina moderately developed. Probasitarsus about equal
in length to combined length of following three tarsomeres and with the dorsal carina strongly raised, rounded and
semi-circular in outline (Fig. 14G). Meso- and metabasitarsus slender and about as long as following three tarso-
meres combined.
Variability: The two adult ♀♀ at hand show quite some variability in size (table 4), the paratype being notably
longer than the holotype. Both specimens have been preserved in spirits why only limited observations can be made
on the colouration. The holotype is greyish mid brown (Figs. 14A–B) while the paratype is lighter in colour and
rather buff (Fig. 14C). The latter specimen is also remarkable for having some scattered white dots on the thorax
and four basal abdominal terga (Fig. 14C). The body sculpturing is generally more pronounced in the holotype. The
paratypic penultimate instar nymph (body length 98.0 mm) is remarkable for showing a distinctive body sculpturing
that is not seen in the two adult examples. The meso- and metanotum bear a few irregularly placed black conical
tubercles and there is a scale-like transverse posteromedian excrescence on abdominal terga I–VII, which is small
on II–V and VII (smallest on IV and V) but distinct on the median segment (= tergum I) and VI. The excrescence on
tergum VI is the largest and towards the anterior has a further almost semi-circular medio-longitudinal keel, giving
it a T-shaped outline in dorsal aspect. The typical digitiform Preopercular organ seen in the adult specimens is still
undeveloped in this immature example and the colouration is a yellowish mid brown and particularly with the limbs
irregularly flecked with ochre.
Comments: Males and eggs unknown.
Distribution: Central Sulawesi, Prov. Sulawesi Selatan, Luwu Regency [NHMB].
TABLE 4. Measurements of Leprocaulinus digitatus n. sp. [mm]
HT, ♀
[NHMB]
PT, ♀
[NHMB]
Body 132.5 149.0
Pronotum 5.5 5.6
Mesonotum 32.0 34.0
Metanotum 15.6 16.0
Median segment 6.9 6.2
Profemora 32.7 28.0
Mesofemora 21.0 19.2
Metafemora 24.6 24.0
Protibiae 25.5 27.0
Mesotibiae 19.8 18.5
Metatibiae 23.2 26.5
Antennae ca. 52.0 ca. 61.0
HENNEMANN
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FIGURE 14. Leprocaulinus digitatus n. sp.: A. ♀ (HT), dorsal view [NHMB]; B. ♀ (HT), lateral view [NHMB]; C. ♀ (PT),
dorso-lateral view [NHMB]; D. Terminal abdominal segments of ♀ (PT) in lateral view [NHMB]; E. Terminal abdominal seg-
ments of ♀ (PT) in dorsal view [NHMB]; F. Terminal abdominal segments of ♀ (HT) in lateral view [NHMB]; G. Ventral view
of head, pro- and mesothorax of ♀ (PT) [NHMB].
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FIGURE 15. Leprocaulinus heinrichi (Günther, 1935) n. comb.: A. ♀ (AT), lateral view [MNHU]; B. ♂ (HT), dorso-lateral
view [MNHU]; C. Lateral view of terminal abdominal segments of ♀ (AT) [MNHU]; D. Lateral view of terminal abdominal
segments of ♂ (HT) [MNHU].
Leprocaulinus sulawesiense n. sp.
(Figs. 16)
Carausius sp., Günther, 1935: 8, 10.
Carausius immundus, Günther, 1938: 58, 81 (in part—not the two ♀♀ from Luwu; these are L. digitatus n. sp.).
Carausius insularis (?), Hennemann, 1998: 97, pl. 1: 9–10, pl. 2: 1–4 & 7.
HT, ♀: Celebes, Latimodjong-Geb. Oeroe 800m, G. Heinrich 8.30 [MNHU].
PT, ♂: Celebes, Latimodjong-Geb. Oeroe 1500m, G. Heinrich VI.30 [MNHU].
PT, ♂: Tanke Salokko 1500 m; S-O. Celebes, Mengkoka-Geb., 1.1932, G. Heinrich [MNHU].
PT, 2 ♂♂, 1 ♀, 3 eggs: S-Sulawesi, Straße von Rantepao nach Palopo km12 ca. 1100 m, leg. F. Hennemann 13.–18.VIII.1995
[coll. FH, No’s 0299-1 to 3, E].
PT, ♀: Indonesien, S-Sulawesi, Tanah Toraja, Rantepao 700m, leg. Gunawan X.1995 [coll. FH, No. 0299-4].
PT, 6 ♂♂, 8 ♀♀, 1 ♀ (penultimate instar), 1 ♀ (n4): Indonesien, S-Sulawesi, Tanah Toraja Highland, leg. Tajuddin X.1995–
III.1996 [coll. FH, No’s 0299-5 to 18].
Etymology: The name of this new species refers to the distribution on the island of Sulawesi.
Differential diagnosis: In lacking wings, ♂♂ of this new species resemble the Sulawesian L. heinrichi (Gün-
ther, 1935), which is here transferred from the genus Carausius Stål, 1875, and M. kaupii (Stål, 1875), which is
here transferred from the genus Myronides Stål, 1875. Males of L. heinrichi differ from the new species by the very
distinctive green and brown colouration as well as the much more stocky habitus (Fig. 15B). From L. kaupii the ♂♂
of this new species differ by having just two blunt swellings between the eyes (Fig. 16M). Females are very similar
to those of L. insularis talaudiensis (Günther, 1934) from the Talaud Islands but differ by the somewhat more stocky
shape, more elongate and slender, almost parallel-sided head, much smaller Preopercular organ on abdominal ster-
num VII (Fig. 16H), more deeply excavated posterior margin of the anal segment (Fig. 16G), less convex subgenital
plate and less angular dorsal lobe of the probasitarsi.
Description: The following descriptions are based on all specimens at hand for examination.
♀(Fig.16). Medium-sized for the genus (body length 125.5–147.0 mm), form fairly slender with a short me-
dian segment and only a pair of low swellings between the eyes (Figs. 16A–B). Body surface unevenly tuberculose,
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granulose and rugulose, sculpturing variable and occasionally with scale-like swellings and excrescences on median
segment and abdominal segment VI. Colour variable, ranging from pale ochre over various shades of ochraceous or
greyish brown to very dark brown, more rarely dull green; often with faint darker mottling and occasionally with a
pale ochre longitudinal median streak on abdominal terga VIII–X. Large tubercles of the head and thorax pale ochre
to dull yellow. Genae usually with a small black postocular marking. Antennae dull ochre and sometimes with a
slight reddish wash. Eyes dull ochre.
Head: Longer than wide, subcylindrical with the genae roughly parallel. Frons with two small transverse im-
pressions between bases of antennae (Fig. 16J). Between the eyes with a slightly swollen transverse area that bears
two fairly low and obtuse swellings. Vertex flat and with a fine, impressed coronal line, otherwise set with a few
scattered nodes and small tubercles (Fig. 16J), genae usually with 2–3 obtuse nodes (Fig. 16K). Eyes circular in
outline, moderately projecting and their diametre contained about 2.6x in length of genae. Antennae reaching to
posterior of median segment. Scapus compressed dorsoventrally, somewhat deflexed laterally with both lateral
margins gently rounded and almost 2x longer than wide. Pedicellus oval in cross-section and about 2/5 the length of
scapus.
Thorax: Pronotum of similar dimensions as head, rectangular in outline with the anterior portion very weakly
expanded and about 1.8x longer than wide. Transverse median sulcus moderate, widely V-shaped and almost ex-
panding over entire width of segment; anterior margin with a median pair of tubercles and some further paired
tubercles on dorsal surface (Fig. 16J). Mesothorax elongate, slender and about 5.5x longer than prothorax. Mesono-
tum with a fine longitudinal median carina and unevenly set with tubercles and nodes of variable sizes; those along
the lateral margins more acute and largest. Mesopleurae with a longitudinal median row of nodes; mesosternum
acutely keeled medio-longitudinally but otherwise smooth (Fig. 16L). Metanotum a little less than half the length
of mesonotum, about 3.7x longer than wide; sculpturing like mesonotum. Metapleurae and sternum sparsely nodu-
lose.
Abdomen: Median segment about 1.5x longer than wide and a little less than half the length of metanotum;
sculpturing alike and occasionally with a transverse scale-like crest or swelling at posterior margin. Abdomen ex-
cluding median segment somewhat longer than head and complete thorax combined; entire dorsal surface with a
fine but fairly acute longitudinal median carina and irregularly tuberculose and rugulose. Segment II 1.6x longer
than median segment, II–VI roughly uniform in length and width, on average some 2.8x longer than wide. Tergum
VI usually somewhat swollen medially, sometimes with a pair of low swellings or occasionally with a laterally
compressed, crest-like dorsal lobe (Fig. 16B). VII slightly shorter and narrower than all preceding segments. Sterna
sparsely tuberculose and Preopercular organ formed by a small, transverse, glossy black median tubercle some
distance in front of the posterior margin of sternum VII (Fig. 16H). Tergum VIII somewhat less than half the length
of VII and slightly widening towards the posterior, IX shorter and almost quadrate in dorsal aspect. Anal segment
longer than IX, strongly convex longitudinally and somewhat narrowing towards the posterior, the posterior margin
with a small, roughly semi-circular median excavation and the outer angles obtusely angular (Fig. 16G); the lateral
margins slightly deflexed and angular in the basal portion (Fig. 16F). Epiproct distinct, somewhat variable in length
and shape, usually wider than long and projecting notably behind apex of anal segment (Fig. 16G). Cerci very small,
conical with the apex strongly constricted. Subgenital plate strongly keeled medio-longitudinally, convex, bulgy
and more or less angular in the median portion (Fig. 16F) and the posterior margin narrowed and projecting slightly
beyond apex of abdomen (Fig. 16G).
Legs: All moderately long and slender, the profemora slightly shorter than mesothorax, metafemora reaching
about half way along abdominal segment IV and metatibiae reaching to abdominal segment VII. Anterodorsal carina
of profemora moderately deflexed and more or less undulate and wavy in the basal one third (Figs. 16J, K); the
posteroventral carina with two small, obtuse sub-apical teeth. The dorsal carina of the protibiae strongly but almost
uniformly deflexed and lamellate (occasionally slightly rounded sub-apically; Fig. 16K)). Meso- and metafemora
slender, laterally compressed and unarmed except for two small sub-apical teeth on the two outer ventral carinae.
Medioventral carina moderate. Probasitarsus almost as long as remaining tarsomeres except claw combined and
with the dorsal carina strongly raised and rounded (Fig. 16K). Meso- and metabasitarsus slender and about as long
as following three tarsomeres combined.
♂(Fig.16). Medium-sized for the genus (body length 81.0–105.0 mm), shape slender, apterous with a fairly
short median segment and merely with a pair of low and obtusely conical swellings between the eyes (Fig. 16C).
Body surface very minutely and sparsely granulose. General colour ranging from dull ochre over mid to dark brown,
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occasionally with a slight olive wash. Apical portions of all femora and all coxae may be more or less distinctly red.
Antennae greyish mid brown.
Head: Similar to ♀♀ but widest at the eyes and distinctly narrowing towards the posterior; the coronal line very
distinct in the psterior portion and vertex set only with a very few low tubercles. Pair of swellings between the eyes
more prominent and ranging from obtusely conical to rounded (Fig. 16M). Eyes very large, prominent, projecting
and their diametre contained about 1.6x in length of genae. Antennae reaching to posterior margin of abdominal
segment III. Scapus compressed dorsoventrally with the lateral margins straight but slightly converging towards the
base; almost 2x longer than wide. Pedicellus oval in cross-section.
Thorax: Pronotum roughly of same dimensions as head, the anterolateral angles somewhat expanded and the
lateral margins slightly concave (Fig. 16M). Transverse median sulcus very prominent, very weakly V-shaped and
almost expanding over entire width of segment, a distinct impressed longitudinal median line present and running
along entire length of segment. Tubercles just very weakly defined to obsolete and anterior margin with a small
pair of median tubercles. Mesothorax very elongate and slender and just slightly widened posteriorly; about 6.6x
longer than prothorax. Mesonotum with a very weak and fine longitudinal median carina and mesosternum weakly
tectinate longitudinally. Metanotum about 2/5 the length of metanotum, surface alike.
Abdomen: Median segment a little less than half the length of metanotum, roughly 3x longer than wide and
gently narrowed medially. Segment II 1.3x longer than median segment and very slightly longer than all following
segments, II–VII slightly gradually decreasing in length with VII only about 3/5 the length of II. II–V on average
about 5x longer than wide, VII only about 3.5x longer than wide; II–VII uniform in width and all terga with a very
fine and indistinct longitudinal median line. Tergum VIII roughly half the length of VII and gently widening towards
the posterior, IX shorter and narrowed towards the posterior; medio-longitudinal carina more distinct than on pre-
ceding terga. Anal segment strongly tectiform and split to form two movable hemi-terga (Fig. 16E); these broad and
angular in lateral aspect (Fig. 16D) with the lower angle roughly at 80° and the interior surface of set with several
minute denticles (Fig. 16E). Cerci small, conical, slightly compressed laterally and slightly projecting beyond apex
of anal segment. Poculum fairly small, moderately convex and cup-shaped, bluntly angular in lateral aspect with the
posterior half obtusely carinate medio-longitudinally; the posterior margin declining in lateral aspect, bi-labiate and
weakly indented medially and just not reaching posterior margin of tergum IX.
Legs: All long, slender and unarmed except for 2 small sub-apical denticles on the two outer ventral carinae of
the femora. Profemora a little shorter than mesothorax, metafemora reaching about half way along abdominal seg-
ment IV and metatibiae roughly reaching to apex of abdomen. Basitarsi slender, very elongate and at least equal in
length to remaining tarsomeres combined.
Eggs(Figs16N–O): A description in German and sketches have been presented by Hennemann (1998: 97, fig.
5) but a detailed English description and new illustrations are here provided. The three eggs at hand were laid by
the ♀ from near Rantepao (coll. FH No. 0299-1) and thus are fully developed. Two of these eggs hatched, but the
nymphs refused all alternative food plants offered.
Small, ovoid, 1.4x longer than wide, higher than wide and oval in cross-section. The anterior portion very
slightly narrowed and the polar-area very weakly conical in centre. Entire surface of chorion very minutely granu-
lose and appearing velvety to the naked eye. Micropylar plate about 2/3 the length of capsule, elongate with the
anterior half roughly parallel-sided and the posterior half expanded and club-shaped; polar end with a very slight
median indention. Micropylar cup a small node some distance off the polar end and roughly in centre of the ex-
panded posterior portion. Median line very indistinct and not reaching to polar area. Operculum slightly oval in
outline, convex and with a large, knob-like capitulum on a short stalk in centre; capitulum with a central hollow.
General colour of capsule reddish mid brown irregularly flecked with ochre. Anterior margin and micropylar plate
ochre. Operculum reddish brown and capitulum dark brown. Measurements [mm]: Length including capitulum 3.2,
length 2.7–2.8, width 1.9, height 2.1–2.2, length of micropylar plate 1.8.
Variability: As usual for many members of the subfamily Lonchodinae, much less intraspecific morphological
variability is seen in ♂♂. These merely show variability in size and colouration. The two ♂♂ from near Rantepao
in the first authors collection (coll. FH No’s 0299-2 & 3) have the coxae and apex of all femora distinctively red,
while the femora are plain or just with a very slight reddish wash at the apex in all other specimens at hand. In ad-
dition to size and colouration, ♀♀ in contrast also vary in the degree of body granulation, size of the cephalic pair
of swellings and length of the epiproct and subgenital plate. The subgenital plate either hardly reaches the apex of
the epiproct or it slightly projects beyond the epiproct. The ♀ from near Rantepao in the authors personal collection
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FIGURE 16. Leprocaulinus sulawesiense n. sp.: A. ♀ (PT), lateral view [coll. FH, No. 0299-14]; B. ♀ (PT), lateral view [coll.
FH, No. 0299-1]; C. ♂ (PT), dorso-lateral view [coll. FH, No. 0299-19]; D. Lateral view of terminal abdominal segments of
♂ (PT) [coll. FH, No. 0299-19]; E. Dorsal view of terminal abdominal segments of ♂ (PT) [coll. FH, No. 0299-19]; F. Lateral
view of terminal abdominal segments of ♀ (PT) [coll. FH, No. 0299-5]; G. Dorsal view of terminal abdominal segments of ♀
(PT) [coll. FH, No. 0299-5]; H. Ventral view of terminal abdominal segments of ♀ (PT) [coll. FH, No. 0299-5]; J. Head and
pronotum of ♀ (PT) in dorsal aspect [coll. FH, No. 0299-12]; K. Lateral view of head, prothorax and front legs of ♀ (PT) [coll.
FH, No. 0299-1]; L. Ventro-lateral view of mesothorax of ♀ (PT) showing the keeled mesosternum [coll. FH, No. 0299-7]; M.
Head and pronotum of ♂ (PT) in dorso-lateral aspect [coll. FH, No. 0299-16]; N. Egg (PT), dorsal view [coll. FH, No. 0299-E];
O. Egg (PT), lateral view view [coll. FH, No. 0299-E].
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(coll. FH No. 0299-1, Fig. 16B) has a large, transverse and scale-like swelling at the posterior margin of the median
segment and a prominent, laterally compressed, crest-like lobe on abdominal tergum VI. These have been illustrated
by Hennemann (1998: 99, pl. 2: 1).
Comments: There has previously been some confusion about the identity of this Sulawesian taxon. Günther
(1935: 8) was unsure about the identity of the ♀ and ♂ from Uru in the collection of MNHU and referred to them as
“Carausius spec.”. The three specimens from Bua Karaeng and Luwu in NHMB are conspecific and were referred
to as Carausius immundus Brunner v. Wattenwyl, 1907 by Günther (1938: 81). Also Hennemann (1998: 97) was in
doubt about the identity of these specimens and further conspecific material. Having compared these to the type-
specimens of C. immundus from Sumatra and Java in the collection of NHMW, Hennemann stated these Sulawesian
examples were obviously a distinct species and with doubt listed them as “Carausius insularis (Kirby, 1896) ?”.
Hennemann (1998) provided illustrations of both sexes and also presented a first description and illustration of the
previously unknown eggs (Hennemann, 1998: 102, Fig. 5). Further detailed examination of the Sulawesi material
has now proven this to represent an as yet undescribed species.
Distribution: Apparently widely distributed throughout the southern half of Sulawesi. S-Sulawesi, Prov. Su-
lawesi Selatan, Gunung Latimojong, Rantemario, Uru 800 m [MNHU]; S-Sulawesi, Prov. Sulawesi Selatan, Ran-
tepao Palopo km12, ca. 1100 m [coll. FH]; S-Sulawesi, Prov. Sulawesi Selatan, Tana Toraja, 700 m [coll. FH];
SE-Sulawesi, Mengkoka Mountains, Gunung Tanke Salokko [MNHU].
TABLE 5. Measurements of Leprocaulinus sulawesiense n. sp. [mm]
HT, ♀
[MNHU]
PT, ♂♂
[coll. FH]
PT ♀♀
[coll. FH]
Body 133.0 81.0–105.0 125.5–147.0
Pronotum 4.5 2.8–3.8 5.0–5.4
Mesonotum 30.0 20.4–27.9 27.9–34.5
Metanotum 13.5 9.6–12.0 13.6–15.0
Median segment 6.0 3.9–5.8 6.4–6.9
Profemora 26.0 20.5–27.2 24.8–28.5
Mesofemora 18.0 14.0–18.3 16.7–20.0
Metafemora 22.5 17.7–23.7 21.5–25.0
Protibiae - 19.0–27.0 22.9–25.0
Mesotibiae - 14.3–18.2 15.7–18.6
Metatibiae - 20.2–25.0 23.2–25.7
Antennae - 45.0–67.0 50.0–55.0
Genus Mnesilochus Stål, 1877
(Figs. 17–18)
Type-species: Mnesilochus capreolus Stål, 1877: 39, by subsequent designation of Kirby, 1904: 320.
Comments: This genus is distributed in Borneo, Palawan and the Philippines and the author is aware of an as yet
undescribed species that occurs on Sangihe Island north of Sulawesi. The two species here described are the first
records of the genus from Sulawesi.
Mnesilochus has been removed from synonymy with Carausius Stål, 1875 and reinstated as a valid genus by
Hennemann & Conle (2007: 66). These authors and Seow-Choen (2016: 279) have attempted a clear separation
from the very similar Hermagoras Stål, 1875 but morphological characters in the insects are scarce and merely
include whether the mesosternum is keeled medio-longitudinally (Hermagoras) or not (Mnesilochus) and whether
the eggs are round to ovoid with a rounded polar area (Hermagoras) or more or less laterally compressed and have a
polar mound (Mnesilochus). The distinction presented by Seow-Choen (2016: 279) includes an erroneous character
and thus deserves clarification. In addition to the morphology of the mesosternum, the author stated in the key to
Bornean genera of Lonchodinae: Lonchodinae “Postero-dorsal carina of profemur curved forwards, lamellate and
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may be foliaceous covering totally or partially antero-dorsal carina which is generally straight” for Hermagoras
and “Postero-dorsal carina of profemor nut usually lobes” for Mnesilochus (Seow-Choen, 2016: 279). This distinc-
tion does not hold true because Seow-Choen misinterpreted the anterodorsal carina as the posterodorsal carina and
the anteroventral carina as the anterodorsal carina. Actually, the morphology of the profemur is basically identical
in these two genera and remarkable in that the medioventral carina is very prominently developed, lamellate and
strongly displaced towards the interior/anterior, resulting in that the anteroventral carina is strongly shifted inwards
and upwards and notably approaches the anterodorsal carina. The anterodorsal carina (not the posteroventral carina
as wrongly stated by Seow-Choen) is much expanded, lamellate and curved forwards to partly or completely cover
the anteroventral carina in dorsal aspect. The posterodorsal carina in fact is very indistinct and not raised at all, being
low and merely represented as an obtuse keel or ridge.
Mnesilochus bodiense n. sp.
(Fig. 17)
Phasgania furcata, Günther, 1935: 11, pl. 1: 6.
HT, ♂: Nord-Celebes, Bodi, 30.9.1930, G. Heinrich leg. [MNHU].
Etymology: Named after Bodi, the type-locality and only known record of this small new species.
Differential diagnosis: While fairly typical in general shape for the genus, ♂♂ (the only sex known) of this
small species are easily separated from the ♂♂ of all other known species of the genus by the very broad and in
lateral aspect obtusely rounded hemi-terga of the anal segment (Fig. 17C), which are usually more acuminate and
slender in the other species. The very small size (body length of the holotype 62.0 mm) resembles the two Bornean
M. bushelli (Bragg, 2005) and M. rusticus (Brunner v. Wattenwyl, 1907) but those ♂♂ are much more stocky in
shape and have rather triangular hemi-terga of the anal segment with an acuminate apex.
Description:♂(Fig.17). The unique holotype lacks the right front leg, left protarsus, right mesotibia and meta-
tarsus and the right antenna except for the two basal segments. Thus, no description of the taxonomically important
protarsi can be provided.
Small (body length 62.0 mm) and of average, rather typical shape for the genus. General colour ochraceous
brown, eyes buff, antennae dark reddish brown (except for the two basal segments, which are coloured like the
body). Entire body surface densely granulose, the granules however less in number and notably less pronounced on
abdomen and gradually becoming fewer in number and smaller in size towards apex of abdomen; granules of the
thorax glossy. Abdominal terga with a very fine medio-longitudinal line.
Head: Longer than wide, broadest at the eyes with the genae notably narrowing towards the posterior (Fig.
17A). Vertex flat and between the eyes with a raised area that is densely set with glossy granules and bears two
very obtuse tubercles. Coronal line of vertex indistinct. Eyes large, circular in outline, projecting hemispherically
and their diameter contained 1.7x in length of genae. Antennae slightly projecting over posterior margin of median
segment. Scapus compressed dorsoventrally, almost rectangular and weakly narrowed basally, 2x longer than wide.
Pedicellus cylindrical and about 1/3 the length of scapus.
Thorax: Pronotum roundly rectangular and very weakly narrowed medially, 1.5x longer than wide and narrower
than broadest portion of head. Transverse median sulcus moderately distinct, straight and not reaching to lateral
margins of segment. Mesothorax very elongate, slender, weakly widened at anterior margin and distinctly broad-
ened and slightly swollen in posterior portion; 7.1x longer than pronotum. Metanotum 0.6x the length of mesonotum
with the posterior 1/3 slightly thickened (Fig. 17A). Meso- and metasternum and pleurae without specializations.
Abdomen: Median segment as wide as posterior portion of metanotum, a little less than ¼ the length of meta-
notum, slightly narrowed medially and a about 1.6x longer than wide. Segment II about 1.7x longer than median
segment. II–VI uniform in width and almost uniform in length, on average 3x longer than wide. VII only about ¾
the length of preceding and slightly widened posteriorly. Sterna II–VII simple. Terga VIII and IX acutely keeled me-
dio-longitudinally and with lateral margins weakly concave; VIII moderately widened towards the posterior, widest
segment and slightly shorter than VII; IX transverse and less than half the length of VIII. Anal segment tectiform
and split longitudinally to form two movable hemi-terga, which are connected by a membrane in the basal portion;
in lateral aspect these are rather broad with the apical portion short, obtusely rounded and somewhat downward
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directed (Fig. 17C), the interior surface is irregularly set with small denticles. Cerci short and conical with a fairly
acute apex. Poculum small, cup-shaped and with a rounded posterior margin, that slightly projects beyond tergum
IX (Fig. 17C).
Legs: Long, distinctly carinate with the mesofemora moderately thickened and smooth dorsally; profemora
almost as long as mesothorax, metafemora reaching to posterior margin of abdominal segment IV and metatibiae
reaching to abdominal segment IX. Posteroventral carina of profemora with a one small sub-apical tooth. Two outer
ventral carinae of meso- and metafemora each with three closely placed sub-apical teeth that gradually decrease in
size towards the apex of femur; those of the mesofemora with a common base (Fig. 17B). Meso- and metabasitarsi
about as long as following three tarsomeres combined; slender.
FIGURE 17. Mnesilochus bodiense n. sp. ♂, holotype [MNHU]: A. Habitus dorsal view; B. Habitus, dorso-lateral view; C.
Terminal abdominal segments seen laterally.
TABLE 6. Measurements of Mnesilochus bodiense n. sp. [mm]
HT, ♂
[MNHU]
Body 62.0
Pronotum 2.2
Mesonotum 16.0
Metanotum 10.8
Median segment 2.4
Profemora 15.5
Mesofemora 12.0
Metafemora 13.0
Protibiae 13.5
Mesotibiae 9.7
Metatibiae 12.9
Antennae 34.0
Comments: Females and eggs unknown. Günther (1935: 11) erroneously assigned this specimen to Dixippus
furcatus Brunner v. Wattenwyl, 1907, which is a synonym of Periphetes forcipatus (Bates, 1865), and provided a
description. The possible unknown opposite sex of this small new species may be represented by the ♀ from Luwu,
South Sulawesi in NHMB and erroneously referred to as H. haematomus (Westwood, 1859) by Günther (1938: 78).
This particular specimen is definitely not the opposite sex of the ♂ assigned to H. haematomus by that author in
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NHMB and from the same locality; this is a paratype of Hermagoras celebensis (Hennemann, 1998). If compared
to the holotype of M. bodiense n. sp. the ♀ in NHMB (here described as M. luwuense n. sp.) appears to be too large,
having a body length of 102.0 mm, thus is clearly outside the average size diversity between corresponding sexes
within Mnesilochus. However, since both sexes of species in this genus can show considerably size ranges fresh
material from the same locality will be necessary for any confirmed decision.
Distribution: Central Sulawesi, Prov. Sulawesi Tengah, Buol Regency, Bodi [MNHU].
Mnesilochus luwuense n. sp.
(Fig. 18)
Lonchodes haematomus, Günther, 1938: 78.
HT, ♀: Sarasin, Luwu, Central-Celebes; Lonchodes hosei Kby. K. Günther det. [NHMB, No. VI.D.133].
Etymology: Named after the Luwu Regency, the type-locality and only known record of this new species.
Differential diagnosis: Similar in general shape and appearance to the Philippine M. haedulus Stål, 1877 (as
described and illustrated by Hennemann & Conle, 2007: 73, figs. 107–113) and the Palawanese M. palawanicus
(Carl, 1913) but readily separated from these and from all other known members of the genus by the morphology
of the mesofemora. While these species have the dorsomedian lobe roughly triangular in shape with the posterior
margin dentate, this lobe is distinctively bifid with a median excavation in M. luwuense n. sp. (Fig. 18D).
Description:♀ (Fig. 18). The unique holotype has provisionally been preserved in spirits, what might have
caused discolouration. Moreover, ♀♀ of this genus are known to show considerable morphological and chromatic
variability, why the description here presented on basis of the holotype can in several aspects only be exemplary for
this species.
Of medium size (body length 102.0 mm) and moderately stocky shape for the genus with a large, bifid dorsal
lobe of the mesofemora. Entire body surface unevenly granulose, tuberculose, verruculose and rugulose; abdominal
terga II and V–VIII with posteromedian protrusions. General colour of the holotype greyish mid brown with the
posterolateral portions of the mesothorax, lateral portions of abdominal sterna II–IV buffy. Mesonotum with four
small blackish dorsal markings in anterior half and a narrow black lateral marking some distance before the poste-
rior margin. Metanotum with two black spots sub-anteriorly and a black spot at each posterolateral angle. Median
portion of abdominal tergum II ochre with two elongate and washed blackish median markings and two smaller dark
brown markings posterolaterally. Two similar small blackish markings in anterior portion of tergum III. Tergum V
with lateral surfaces mostly pale ochre and with a blackish streak in anterior half. Terga VII and VIII each with a
dark brown to black central marking. Cerci and epiproct straw. A blackish sub-apical marking also present on inte-
rior surface of protibiae. Mesofemora irregularly dark brown basally and with a buff transverse sub-basal band, the
mesotibiae buff in the median portion and dark brown apically. Metafemora ochre on their interior surface. Anten-
nae greyish mid brown ventrally, ochre dorsally with most antennomeres dark apically; scapus buff. Eyes dark grey.
Head: Elongate, sub-cylindrical, notably longer than wide and very slightly narrowed posteriorly. Between the
eyes with a raised transverse area which has both outer ends protruded into an obtuse, slightly anteriad directed
spine. Vertex flattened and irregularly set with granules, nodes and tubercles of variable sizes, also a few tubercles
present on genae. Posterior margin with a transverse row of four obtusely rounded but enlarged tubercles. Eyes
almost circular in outline, weakly protruding and their diameter contained 2.2x in length of genae. Antennae project-
ing somewhat over posterior margin of mesothorax. Scapus strongly compressed dorsoventrally, narrowed basally,
expanded medially and roughly oval in outline. Pedicellus slightly oval in cross-section and less than half the length
of scapus.
Thorax: Pronotum roughly of same dimensions as head, rectangular and 1.7x longer than wide; the transverse
median sulcus distinctly impressed but narrow, straight and expanding over entire width of segment. Mesothorax
4.2x longer than pronotum and very weakly widened post-anteriorly. Mesonotum with a fine longitudinal median
line and set with a few scattered, enlarged and somewhat wart-like tubercles; a transverse row of four enlarged
tubercles just before posterior margin. Metanotum over 3x longer than wide and about 0.6x length of mesonotum,
the posterior half weakly expanded and the median line like on mesonotum. Mesopleurae with a medio-longitudinal
row of flat, elongate tubercles. Mesosternum densely granulose.
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FIGURE 18. Mnesilochus luwuense n. sp. ♀, holotype [NHMB]: A. Habitus, dorsal view; B. Habitus, dorso-lateral view; C.
Habitus lateral view; D. Right mid leg in postero-lateral aspect; E. Lateral view of terminal abdominal segments; F. Dorsal view
of terminal abdominal segments.
Abdomen: Median segment a little more than half the length of metanotum, 1.8x longer than wide with the
lateral margins weakly concave and a small transverse swelling posteromedially. Abdomen excluding median seg-
ment slightly shorter than pro-, meso- and metathorax combined with segments V–VI notably swollen. Segment II
about equal in length to median segment, II–IV uniform in length and V–VII gradually increasing in length; II par-
allel-sided and 2x longer than wide, III–V gradually widening with V only 1.3x longer than wide. Anterior portion
of VI widest part of abdomen and tergum strongly narrowing towards posterior with a trapezoidal outline; lateral
margins somewhat deflexed. VII parallel-sided, as broad as II and but shorter and hardly longer than wide. Tergum
II with a transverse posterior lobe that bears five rounded tubercles, V and VI each with a similar but more obtuse
transverse posteromedian protuberance. Terga VII and VIII each bear a slightly bilobed posteromedian excrescence,
which posteriorly extends over the anterior portion of the following tergum (Fig. 18E). Preopercular organ formed
by a pair of low and obtuse, verruculose swellings near posterior margin of sternum VII. Tergum IX transverse