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Six new species of the genus Orestes Redtenbacher, 1906, O. bachmaensis sp. nov., O. botot sp. nov., O. diabolicus sp. nov., O. dittmari sp. nov., O. draegeri sp. nov. and O. krijnsi sp. nov. are described and figured from Vietnam. Descriptions are based on both sexes, except for O. diabolicus sp. nov., of which only the male is known. The type species O. mouhotii (Bates, 1865) is redescribed and its distribution range is extended. Dares (Dares) subcylindricus Redtenbacher, 1906 is removed from the synonymy with Orestes mouhotii (Bates,1865), transferred to Orestes and redescribed from both sexes and eggs. The new combination Orestes subcylindricus Redtenbacher, 1906 stat. rev., comb. nov. is proposed. The generic description of Orestes is slightly improved and the delimitation from Pylaemenes Stål, 1875 is precised. A plate of the type species of Pylaemenes, P. coronatus (Haan, 1842) is presented for comparison. Pylaemenes guangxiensis Bi & Li, 1994, P. japonicus Ho, 2016 and P. shirakii Ho & Brock, 2013 are transferred to Orestes and the new combinations O. guangxiensis (Bi & Li, 1994) comb. nov., O. japonicus (Ho, 2016) comb. nov. and O. shirakii (Ho & Brock, 2013) comb. nov. are proposed. The cephalic armature is discussed and figured. A key to the species of Orestes from Vietnam and Cambodia is presented with accompanying plates, as well as a distribution map and live in nature pictures.
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Belgian Journal of Entomology 58: 1–62 (2018) ISSN: 2295-0214
www.srbe-kbve.be
urn:lsid:zoobank.org:pub:38C32CDC-BC98-4094-89C6-42F1035A2229
Belgian Journal of Entomology
The Oriental stick insect genus Orestes Redtenbacher, 1906:
Taxonomical notes and six new species from Vietnam
(Phasmida: Heteropterygidae: Dataminae)
Joachim BRESSEEL¹ & Jérôme CONSTANT²
1,2 Royal Belgian Institute of Natural Sciences, O.D. Phylogeny and Taxonomy, Entomology, Vautier street 29,
B-1000 Brussels, Belgium
1 E-mail:joachimbresseel@gmail.com (corresponding author)
urn:lsid:zoobank.org:author:3C4EF358-9716-46F0-8575-26BE1EDE4349
2 E-mail: jerome.constant@naturalsciences.be
urn:lsid:zoobank.org:author:6E6072A1-9415-4C8D-8E60-2504444DB290
Published: Brussels, January 29, 2018
2
Citation: BRESSEEL J. & CONSTANT J., 2018 - The Oriental stick insect genus Orestes Redtenbacher, 1906: Taxonomical
notes and six new species from Vietnam (Phasmida: Heteropterygidae: Dataminae). Belgian Journal of Entomology, 58:
1–62.
ISSN: 1374-5514 (Print Edition)
ISSN: 2295-0214 (Online Edition)
The Belgian Journal of Entomology is published by the Royal
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Front cover: Orestes bachmaensis sp. nov., male holotype (top) and female paratype (bottom).
Belgian Journal of Entomology, 58: 1–62 (2018)
3
The Oriental stick insect genus Orestes Redtenbacher, 1906:
Taxonomical notes and six new species from Vietnam
(Phasmida: Heteropterygidae: Dataminae)
Joachim BRESSEEL¹ & Jérôme CONSTANT²
1,2 Royal Belgian Institute of Natural Sciences, O.D. Phylogeny and Taxonomy, Entomology, Vautier street 29,
B-1000 Brussels, Belgium.
1 E-mail:joachimbresseel@gmail.com (corresponding author)
urn:lsid:zoobank.org:author:3C4EF358-9716-46F0-8575-26BE1EDE4349
2 E-mail: jerome.constant@naturalsciences.be
urn:lsid:zoobank.org:author:6E6072A1-9415-4C8D-8E60-2504444DB290
Abstract
Six new species of the genus Orestes Redtenbacher, 1906, O. bachmaensis sp. nov., O. botot
sp. nov., O. diabolicus sp. nov., O. dittmari sp. nov., O. draegeri sp. nov. and O. krijnsi sp.
nov. are described and figured from Vietnam. Descriptions are based on both sexes, except for
O. diabolicus sp. nov., of which only the male is known. The type species O. mouhotii (Bates,
1865) is redescribed and its distribution range is extended. Dares (Dares) subcylindricus
Redtenbacher, 1906 is removed from the synonymy with Orestes mouhotii (Bates,1865),
transferred to Orestes and redescribed from both sexes and eggs. The new combination
Orestes subcylindricus Redtenbacher, 1906 stat. rev., comb. nov. is proposed. The generic
description of Orestes is slightly improved and the delimitation from Pylaemenes Stål, 1875 is
precised. A plate of the type species of Pylaemenes, P. coronatus (Haan, 1842) is presented
for comparison. Pylaemenes guangxiensis Bi & Li, 1994, P. japonicus Ho, 2016 and
P. shirakii Ho & Brock, 2013 are transferred to Orestes and the new combinations
O. guangxiensis (Bi & Li, 1994) comb. nov., O. japonicus (Ho, 2016) comb. nov. and
O. shirakii (Ho & Brock, 2013) comb. nov. are proposed. The cephalic armature is discussed
and figured. A key to the species of Orestes from Vietnam and Cambodia is presented with
accompanying plates, as well as a distribution map and live in nature pictures.
Keywords: Global Taxonomic Initiative, ootaxonomy, Indochina, Phasmatodea, Araceae
Introduction
The study of the stick insect material collected in the framework of our Global Taxonomic
Initiative projects in Vietnam and Cambodia (2010–2017) revealed six species of Datamini
(Heteropterygidae: Dataminae) in the genus Orestes Redtenbacher, 1906 which are new to
science. Additionally, specimens of O. mouhotii (Bates, 1865) and O. subcylindricus
(Redtenbacher, 1906) comb. nov. were also collected, allowing the assessment of the
taxonomy of these species and leading to several changes.
The tribe Datamini Rehn & Rehn, 1939 consists of seven genera distributed from Japan,
Taiwan and China to the north over the entire Sundaland as far southeast as Wallacea. The
predominantly ground-dwelling taxa consist of small, cryptic species resembling short pieces
of dead branches. They are considered the most basal tribe in the Heteropterygidae Kirby,
1896; females lack a beak-like ovipositor, have no medio-apical spine on the area apicalis and
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
4
possess paired sensory areas on the prosternum and an additional central sensory area on the
profurcasternum (HENNEMANN et al., 2016).
Several keys to the genera of Datamini have been provided by different authors (BRAGG,
1998, 2001; HO, 2013; SEOW-CHOEN, 2016, 2017) all of which have used the presence or
absence of a crest on the head as a generic diagnostic character.
The genus Orestes was described by REDTENBACHER (1906) based on a female nymph and
was redescribed by ZOMPRO (2004). Zompro’s description is here slightly adapted, the type
species O. mouhotii (Bates, 1865) is redescribed and its distribution range is extended to
Cambodia in the east and to Bangkok in the west. The genus currently only contains one valid
species, O. mouhotii, with three synonyms: Dares fulmeki Werner, 1934, Datames
verruculatus Redtenbacher, 1906 and Dares (Dares) subcylindricus Redtenbacher, 1906.
SHIRAKI (1935) recorded O. mouhotii from Taiwan and Japan but HO & BROCK (2013) have
attributed the Taiwanese material to a new species, Pylaemenes shirakii Ho & Brock, 2013,
while the Japanese material was attributed to another new species, Pylaemenes japonicus Ho,
2016 by HO (2016).
The present paper aims to refine the definition of the genus Orestes, especially as compared to
Pylaemenes Stål, 1875, to describe the six new species, to provide an identification key to the
species from Vietnam and Cambodia and to adjust the generic attribution of some species
currently in Pylaemenes. A standardized nomenclature of the cephalic armature of the
Datamini is proposed, which is adapted from the one proposed for the Heteropteryginae
Kirby, 1896 by HENNEMANN et al. (2016).
Material and methods
Due to their nocturnal behaviour, like most Phasmida, the specimens of Orestes
Redtenbacher, 1906 were collected at night. A light-weight and water-proof head torch: Petzl
MYO RXP was used during collecting. Females were kept alive in a plastic box for
producing eggs. Eggs were collected and hatchlings were reared to adulthood by Mr Rob
Krijns (Netherlands). The wild caught specimens were euthanized in a killing jar with
etylacetate (EtOAc) fumes. The specimens were then stored in airtight plastic “zip”-bags in
wood chips (used in rodent cages) and sprinkled with etylacetate (EtOAc) to prevent rotting,
mould and to keep the specimens flexible. The bags were frozen on arrival and the specimens
mounted later on.
The photographs of the specimens were taken with a Canon EOS 700 D camera with a Sigma
DG Macro lens, those of the eggs with a Leica EZ4W stereomicroscope with an incorporated
camera. They were stacked with CombineZ software and optimized with Adobe Photoshop
CS3. All photographs including those in nature were taken by the second author except when
noted. The distribution map was produced with SimpleMappr (SHORTHOUSE, 2010).
The measurements were done with an electronic calliper. The length of the metanontum
includes the median segment.
The nomenclature of the armature of the head (Fig. 1) is adapted for the Dataminae Rehn &
Rhen, 1939, from that used for the Heteropteryginae Kirby, 1896 by HENNEMANN et al.
(2016). It contains the following additions:
- anterior supra-occipitals = a pair of small spines or granules positioned at anterior margin
of eye, between the supra-antennals and the posterior supra-occipitals.
Belgian Journal of Entomology 58: 1–62 (2018)
5
- posterior supra-occipitals = a pair of, often minute, spines or granules positioned at
posterior margin of eye, between the posterior supra-occipitals and the anterior coronals.
- central coronal = a single granule or spine positioned centrally between anterior- and
posterior coronals.
- postocular carina = a distinct carina starting behind eye, reaching posterolateral margin of
head; posteriorly with a protruding granule or tubercle.
Acronyms used for the collections:
FH = Private collection of Frank Hennemann, Freinsheim, Germany.
MNHN = Museum National d’Histoire Naturelle, Paris, France.
NHMW = Naturhistorisches Museum, Vienna, Austria.
OUMNH = Oxford University Museum, Oxford, United Kingdom.
RBINS = Royal Belgian Institute of Natural Sciences, Brussels, Belgium.
RUPP = Royal University of Phnom Pen, Cambodia.
VNMN = Vietnam National Museum of Nature, Hanoi, Vietnam.
Abbreviations:
N.P.: National Park
N.R.: Nature Reserve
HT: holotype
PT: paratype
Taxonomy
Family Heteropterygidae Kirby, 1869
Subfamily Dataminae Rehn & Rehn, 1939
Tribe Datamini Rehn & Rehn, 1939
Genus Orestes Redtenbacher, 1906
Type species: Orestes verruculatus Redtenbacher, 1906 by original monotopy.
Orestes REDTENBACHER, 1906: 47 [described].
Orestes – REHN & REHN, 1939 [attributed to Datamini]. — BRAGG, 2001 [type specimen
data]. — ZOMPRO, 2004 [redescribed]. — HO, 2013 [recorded from China]. — SEOW-CHOEN,
2017 [recorded from Singapore]. — BROCK et al., 2017 [catalogued].
DIAGNOSIS.
♂♂, ♀♀: moderately sized Datamini (body length ♂♂ 34.0–45.0 mm, ♀♀ 42.6–54.4 mm),
apterous and strongly cryptic. Body robust, surface granulose to tuberculose; colouration with
different shades of brown to black, sometimes whitish. Head armature arranged as in Fig. 1,
from almost indistinct to crest-like. Thorax cylindrical to subcylindrical in cross-section. ♀♀
with mesonotum slightly widening towards the posterior, often with mediolongitudinal line
and two rows of tubercles sublaterally. Legs with carinae often indistinct, most distinct carina
is the anterodorsal carina of the profemora. Abdomen dilated centrally, with posterior “wart”
on terga IV-V; tergum IX with distinct posteromedian crest. Subgenital plate cup-like, not
projecting over anal segment. ♂♂ distinctly smaller and narrower than ♀♀ with mesonotum
and abdomen almost parallel-sided. Poculum relatively large and triangular to cup-like from
lateral view. Legs as in female.
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
6
Fig. 1. Orestes Redtenbacher, 1906, nomenclature of cephalic armature.
Eggs: egg capsule spherical to subspherical. Operculum shape variable elongated to circular
and slightly convex. Capsule and operculum covered with long setae with a distal black two-
to four-hooked grapnel-shaped structure. Micropylar plate trilobate with one anterior
expansion and with two posterior expansions; posterior arms directed laterally and projecting
on sides of capsule. Margin of micropylar plate sometimes indistinct. Micropylar cup small,
black, cup-shaped and protruding.
The genus was redescribed by ZOMPRO (2004) upon both sexes and the egg. ZOMPRO’s
description is based on material of O. mouhotii (Bates, 1865) from Nakhon Ratchasima,
Thailand. In the species description, he stated that the widest abdominal segment is segment
V. However, the posterior margin of tergum IV is slightly wider than V in O. mouhotii and all
species of Orestes. Some species can have terga IV-V almost parallel-sided, whereas other
species have tergum V with distinctly oblique lateral margins.
DIFFERENTIATION.
The genus Orestes differs from all other known genera of Datamini Redtenbacher, 1939 by
the following combination of characters:
1) Mesonotum semicylindrical without distinctly raised mediolongitudinal and lateral carina;
mediolongitudinal line sometimes present, sometimes slightly elevated in males.
2) Females with widest part of body at the posterior margin of tergum IV, sometimes tergum
IV-V almost parallel-sided.
3) Anal segment almost parallel-sided, never triangular or strongly tapering towards the
posterior; posterior margin more or less straight.
4) Anal segment in females dorsally flattened, with two distinct lamellate or tuberculose
oblique carinae reaching posterolateral angles.
5) Females with terga IV-V with a wart-like structure posteriorly and tergum IV being the
highest part of the abdomen.
Belgian Journal of Entomology 58: 1–62 (2018)
7
Fig. 2. Pylaemenes coronatus (Haan, 1842). A, from Indonesia, Molucca, West-Seram, X.2012 (RBINS),
habitus dorsal view. B, from same origin, III.2013 (RBINS), habitus dorsal view.
The most closely related genus is Pylaemenes Stål, 1875. However, Pylaemenes has the
meso- and metanotum flattened or slightly tectiform and with distinct raised
mediolongitudinal and lateral carinae; the anal segment of females is strongly triangular in
dorsal aspect and distinctly notched apically; the males have tergum IX strongly widened
posteriorly and oblique laterally (Fig. 2: P. coronatus (Haan, 1842), the type-species of
Pylaemenes).
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
8
BIOLOGY. All species were collected in tropical evergreen rainforest, at low to moderately
high altitude (50–1500 m). In Vietnam and Cambodia the specimens were observed on low
vegetation, usually very close to the ground (below 50cm) but sometimes up to 1.5–3 m from
the ground. They seemed to be exclusively nocturnal and appeared to feed on a variety of
plant species with a preference for Araceae.
DISTRIBUTION. Far Eastern Palaearctic and Oriental Regions: China, Vietnam, Cambodia,
Thailand, Sumatra, Japan, Taiwan.
Species included
1. O. bachmaensis sp. nov. [Vietnam: Bach Ma N.P., Phong
Dien VNMN Station]
2. O. botot sp. nov. [Vietnam: Bidoup Nui Ba N.P.]
3. O. diabolicus sp. nov. [Vietnam: Bidoup Nui Ba N.P.]
4. O. dittmari sp. nov. [Vietnam: Cat Ba N.P.]
5. O. draegeri sp. nov. [Vietnam: Cat Tien N.P., Dong
Nai Biosphere Reserve]
6. O. guangxiensis (Bi & Li, 1994) comb. nov. [China: Fujian, Guangdong,
Guangxi, Hainan, Hong Kong]
= Pylaemenes hongkongensis Brock & Seow-Choen, 2000
7. O. krijnsi sp. nov. [Vietnam: Nui Chua N.P.]
8. O. japonicus (Ho, 2016) comb. nov. [Japan: Yakushima]
9. O. mouhotii (Bates, 1865) [Cambodia, Tatai, Kirirom N.P.;
Thailand: Bangkok, Chantaburi,
Nakhon Ratchasima; Sumatra:
Medan; Singapore]
= O. verruculatus Redtenbacher, 1906
= Dares fulmeki (Werner, 1934)
10. O. shirakii (Ho & Brock, 2013) comb. nov. [Taiwan: Taipei]
11. O. subcylindricus (Redtenbacher, 1906) comb. nov. [Vietnam: Mt Mauson, Cuc
Phuong N.P., Ngo Luong N.R.,
Tay Ten Tu N.R.]
Keys to the species of Orestes from Cambodia and Vietnam
Figs 3–5
MALES
1. Supra-coxal spine on mesopleuron strongly projecting laterally (Fig. 3 A) ......................... 2
- Mesopleuron without a distinct elongated supra-coxal spine (Fig. 3 B–C) .......................... 3
2. No elongated spines on pro- and mesonotum or on median segment (Fig. 3 D) ....................
....................................................................................................................... O. botot sp. nov.
- A pair of strong, elongated spines on pronotum, two pairs on mesonotum and one on
median segment (Fig. 3 G) ................................................................... O. diabolicus sp. nov.
3. Head without central coronal (Fig. 3 J, M–N) ....................................................................... 4
- Head with central coronal (Fig. 3 K–L, O) ........................................................................... 6
4. Anterior coronals distinctly enlarged; blunt and laterally compressed, creating ear-like
processes; posterior margin of metanotum not raised (Fig. 3 J, N) ....................................... 5
Belgian Journal of Entomology 58: 1–62 (2018)
9
- Anterior coronals spinose, not laterally compressed; posterior margin of metanotum raised
(Fig. 3 M) .................................................................................................... O. krijnsi sp. nov.
5. Posterior margin of mesonotum not distinctly raised, armed with two small tubercles (Fig.
3 E) ................................................................................................ O. mouhotii (Bates, 1865)
- Posterior margin of mesonotum conically raised, sulcate medially (Fig. 3 C, F) ...................
................................................................................................................. O. draegeri sp. nov.
6. Supra-orbitals and anterior coronals lamellate (Fig. 3 L) ........................................................
.............................................................. O. subcylindricus (Redtenbacher, 1906) comb. nov.
- Supra-orbitals and anterior occipitals spinose (Fig. 3 K, O) ................................................. 7
7. Coronals not fused basally; anterior coronals rounded apically (Fig. 3 K) .............................
.................................................................................................................. O. dittmari sp. nov.
- Anterior-, central- and posterior coronals fused basally, together into a raised crest; anterior
coronals acute (Fig. 3 O) .................................................................. O. bachmaensis sp. nov.
FEMALES
1. Head with a lamellate elevation on vertex (Fig. 4 A–B, D) .................................................. 2
- Head without lamellate elevation, vertex granulose to tuberculose (Fig. 4 C) ..................... 5
2. Tergum IX with posteromedian crest distinctly notched posteriorly (Fig. 4 E); tergum V
with posterior margin indistinctly narrower than anterior margin ........................................ 3
- Tergum IX with posteromedian crest strongly tectiform and not distinctly notched
anteriorly, with apex narrow (Fig. 4 F); tergum V with posterior margin distinctly narrower
than anterior margin .............................................................................................................. 4
3. Supra-coxal spine on mesopleuron small but acute and projecting laterally (Fig. 4 I);
posterior margin of mesonotum slightly raised with two short oblique carinae; terga III-IV
without posterolateral elevated tubercle ........................................................ O. botot sp. nov.
- Mesopleuron without a supra-coxa spine, at best present as a small tubercle (Fig. 4 J);
posterior margin of mesonotum not raised; terga III–V with a distinct, raised tubercle
posterolaterally ..................................... O. subcylindricus (Redtenbacher, 1906) comb. nov.
4. Outer ventral carinae of meso- and metafemora with two laterally compressed and rounded
teeth posteriorly (Fig. 4 M); abdominal sternum VII with a conspicuous medial compound
tubercle anteriorly ............................................................................. O. bachmaensis sp. nov.
- Outer ventral carinae of meso- and metafemora unarmed, or with two tubercles posteriorly
(Fig. 4 N); abdominal sternum VII with at best an indistinct pair of tubercles in the anterior
half ............................................................................................................ O. dittmari sp. nov.
5. Tergum IX with posteromedian crest notched posteriorly (Fig. 4 H) .....................................
....................................................................................................... O. mouhotii (Bates, 1865)
- Tergum IX with posteromedian crest elongated, strongly tectiform and not visibly notched
anteriorly, and narrow apically (Fig. 4 G) ............................................................................. 6
6. Meso- and metatibiae unarmed; tergum IV distinctly widening towards the posterior (Fig.
4 L) ............................................................................................................. O. krijnsi sp. nov.
- Meso- and metatibiae with a laterally compressed, central tooth dorsally; tergum IV
indistinctly widening towards the posterior, almost parallel-sided (Fig. 4 K)
................................................................................................................. O. draegeri sp. nov.
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
10
Fig. 3. Orestes spp., diagnostic characters of ♂♂ (not to scale). A–C, mesonotum dorsal view. A, O. botot sp.
nov. B, O. subcylindricus (Redtenbacher, 1906) stat. rev. comb. nov. C, O. draegeri sp. nov. D–I, mesonotum
lateral view. D, O. botot sp. nov. E, O. mouhotii (Bates, 1865). F, O. draegeri sp. nov. G, O. diabolicus sp. nov.
H, O. krijnsi sp. nov. I, O. bachmaensis sp. nov. J–O, head lateral view. J, O. draegeri sp. nov. K, O. dittmari sp.
nov. L, O. subcylindricus (Redtenbacher, 1906) comb. nov. M, O. krijnsi sp. nov. N, O. mouhotii (Bates, 1865).
O, O. bachmaensis sp. nov. Cc = central coronal.
Belgian Journal of Entomology 58: 1–62 (2018)
11
Fig. 4. Orestes spp., diagnostic characters of ♀♀ (not to scale). A–D, head, lateral view. A, O. bachmaensis sp.
nov. B, O. dittmari sp. nov. C, O. draegeri sp. nov. D, O. subcylindricus (Redtenbacher, 1906) comb. nov.
E–H, apex of abdomen, anterolateral view. E, O. botot sp. nov. F, O. dittmari sp. nov. G, O. krijnsi sp. nov.
H, O. mouhotii (Bates, 1865). I–J, mesonotum, lateral view. I, O. botot sp. nov. J, O. subcylindricus
(Redtenbacher, 1906) comb. nov. K–L, abdomen dorsal view. K, O. draegeri sp. nov. L, O. krijnsi sp. nov.
M–N, metafemur, lateral view. M, O. bachmaensis sp. nov. N, O. dittmari sp. nov. C = carina; PmN =
posteromedian notch; PlM = posterolateral margin.
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
12
Fig. 5. Orestes. spp., distribution map.
Belgian Journal of Entomology 58: 1–62 (2018)
13
Orestes bachmaensis sp. nov.
urn:lsid:zoobank.org:act:CDC71921-7274-4C1F-B29F-6E0F0A38C700
Figs 5–7, 8 A–C, 9
ETYMOLOGY. The species epithet refers to the type-locality, Bach Ma National Park in Central
Vietnam.
TYPE MATERIAL. VIETNAM: Holotype : C. Vietnam, Bach Ma N.P., summit 16°12’N
107°52’E 15-16.VII.2011, leg. J. Constant & J. Bresseel , I.G.: 31.933 (RBINS).
Paratypes (41♂♂, 31♀♀, eggs):
2♂♂: same data as HT (1: RBINS; 1: VNMN); 2♂♂, 2♀♀, eggs: same data as HT, ex
breeding D. Dittmar 2017 (1, 1, eggs: RBINS; 1, 1: VNMN); 1, 1: Vietnam, Thua
Thiên-Hué prov., Phong Dien near VNMN station, 16°35’12”N 107°20’31”E, 8-9.IV.2017,
leg. J. Constant & J. Bresseel, I.G.: 33.447 (RBINS). 15♂♂, 15♀♀, eggs: C. Vietnam, Bach
Ma N.P., summit 16°12’N 107°52’E 15-16.VII.2011, leg. J. Constant & J. Bresseel, I.G.:
31.933, ex breeding J. Bresseel 2017 (8♂♂, 8♀♀, eggs: RBINS; 7♂♂, 7♀♀, eggs: VNMN).
11♂♂, 13♀♀, 6 eggs: ex Zucht: F. Hennemann 2013-2015, Herkunft: O-Vietnam: Bach Ma
NP, leg. J. Bresseel & J. Constant VIII.2011 [coll. FH, No’s 0880-1 to 24 & E]. 10♂♂: ex
Zucht: F. Hennemann 2014-2016, Herkunft: O-Vietnam: Bach Ma NP, leg. J. Bresseel & J.
Constant VIII.2011 [coll. FH, No’s 0880-25 to 34].
DIAGNOSIS. Males with indistinct hump on dorsal posterior portion of meso- and metafemur;
abdominal terga IV–V posteromedially raised (Fig. 6 B); all abdominal terga only slightly
varying in length (Fig. 6 A).
Females with an elongated, strongly tapered and tectiform crest posteromedially on tergum IX
(Fig. 7 E–F); abdominal sternum VII with a conspicuous medial compound tubercle
anteriorly, merging with mediolongitudinal carina (Fig. 7 E); outer ventral carinae of meso-
and metafemora with two laterally compressed and rounded teeth posteriorly (Fig. 7 G).
DESCRIPTION.
MALE (Fig. 6).
Measurements: see table 1.
Head: supra-antennals spinose and slightly pointed outwards followed by four smaller supra-
occipitals. Vertex raised, supra-orbitals bluntly spinose, positioned at base of crest. Anterior,
posterior and central coronals raised and fused basally. Central coronal about as long as
anterior coronals; anterior coronals lamellate. Posterior and lateral coronals present as small,
conical tubercles. Behind eye a distinct carina reaching posterior edge of crest, resulting in a
conical tubercle. Eyes relatively small, circular and strongly projecting hemispherically.
Antennae with 23 segments (n = 3), shorter than legs; scapus strongly flattened dorsally,
laterally carinated and with spines laterally and posterolaterally; pedicellus slightly narrowing
towards the posterior. First antennomere about as long as the following three antennomeres
combined; antennomeres gradually getting longer till segment XII; antennal segment XIII
shorter, with minute hump mediolaterally; following segments shiny and distinctly shorter;
apical antennomere elongated, club-shaped and slightly longer than the two preceding
antennomeres combined.
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
14
Fig. 6. Orestes bachmaensis sp. nov., holotype . A, habitus, dorsal view. B, habitus, lateral view. C, habitus,
ventral view. D, head and thorax, dorsal view. E, terminalia, dorsal view. F, terminalia anterodorsal view.
G, terminalia, lateral view. H, terminalia, ventral view. I, head and thorax, lateral view.
Belgian Journal of Entomology 58: 1–62 (2018)
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Thorax: pronotum more or less parallel-sided, quadrate and rugose. Anterior margin slightly
incurved. Prozona with a distinct elevation centrally with two to four tubercles; anterior
tubercles slightly larger, sometimes spinose. Metazona with a pair of granules anteriorly and
posteriorly. Posterior margin straight. Mesonotum rugose with anterior margin thickened,
distinctly concave and with a pair of granules or tubercles anteromedially; about parallel-
sided. Posterior portion slightly raised posteromedially and posterolaterally, posteromedially
sometimes slightly conical. Mesopleura slightly widened above coxae; widened portion
notched medially. Metanotum rugose and unarmed, slightly raised posterolaterally.
Metapleura widened above coxae and slightly notched posterolaterally. Prosternum with
sensory areas elongate oval. Sensory area on profurcasternum small and circular. Ventral
surface of body rugose.
Legs: femora with carinae indistinct except for anterolateral carina of profemora;
posterodorsal margin with triangular apex. Profemora about as long as mesonotum, curved
basally and unarmed. Mesofemora shorter than mesonotum, with outer ventral carina with one
or two minute, spinose tubercles. Metafemora slightly longer than pronotum; ventrally armed
as mesofemora. Tibiae with carinae indistinct, slightly shorter than corresponding femora and
unarmed. Tarsomeres I–III about the same length with a posteromedian rounded hump
dorsally. Euplantulae more or less circular. Claws very small.
Abdomen: median segment unarmed, and rugose, with posterior margin concave. Abdominal
terga rugose, only slightly varying in length. Tergum II transverse, slightly narrowing towards
the posterior. Terga IV–V posteromedially raised and armed with two granules. Terga VI–VII
with two posteromedian granules, sometimes indistinctly raised. Tergum VIII slightly wider
than VII; widening towards the posterior, with posterior margin concave; IX distinctly
transverse, with posterior margin with minute posteromedian ridge. Anal segment
dorsoventrally flattened, almost parallel-sided and indistinctly notched posteromedially.
Posterolateral angles rounded. Anterior margin thickened; merging in oblique carina.
Poculum more or less triangular from lateral view and strongly granulose; posterior rim
dorsoventrally flattened and straight posteriorly; posterolateral angles broadly rounded.
Poculum projecting over base of vomer but not reaching base of apical spine. Cerci short,
strongly flattened and setose, not reaching apex of tergum X, with apices broadly rounded to
almost straight. Vomer well developed, semicircular basally till base of posteromedial spine;
posteromedial spine broad, blackish in posterior portion. Apex curved, not ending medially
but slightly to the right.
FEMALE (Fig. 7).
Measurements: see table 1.
Head: supra-antennals, anterior- and posterior supra-occipitals present as blunt spines
gradually shortening towards the posterior. Vertex elongated and raised, supra-orbitals strong,
slightly flattened laterally and blunt. Anterior coronals laterally compressed, lamellate with
anterior margin reaching apex of supra-orbitals. Central coronal compound and slightly
raised. Posterior and lateral coronals present as triangular tubercles. Between lateral coronal
and supra-orbital, a row of three tubercles; between lateral- and anterior coronal several
tubercles on side of crest. Behind eye, a distinct carina reaching posterior edge of crest,
resulting in triangular tubercle. Eyes relatively small, circular and strongly projecting
hemispherically. Antennae shorter than legs with 25 segments (n = 2); scapus strongly
flattened dorsoventrally, laterally carinated and with a central and subapical spine on outer
lateral carina; pedicellus slightly flattened dorsoventrally and slightly narrowing towards the
posterior. First antennomere about as long as following three antennomeres combined; apical
antennomere elongated, club-shaped and distinctly longer.
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
16
Fig. 7. Orestes bachmaensis sp. nov., paratype , Bach Ma N.P., A, habitus, dorsal view. B, habitus, ventral
view. C, head and thorax, dorsal view. D, terminalia, dorsal view. E, Terminalia anterodorsal view. F, terminalia,
lateral view. G, habitus lateral view. H, terminalia, ventral view. I, head and thorax, lateral view.
Belgian Journal of Entomology 58: 1–62 (2018)
17
Thorax: pronotum trapezoidal, slightly widening towards the posterior; lateral margins with
some granules, anterior margin incurved. Prozona with distinct elevation centrally with four
raised tubercles, anterior pair sometimes more distinct. Metazona with two parallel rows of
three granules; posterior margin straight. Mesonotum rugose with anterior margin thickened
and slightly raised triangularly; lateral margins with several enlarged granules; posterior
margin wider than anterior one; anterior portion with a raised pair of distinct, elongate-oval
dark patches. Posterolateral and posterior margins granulose. Mesopleura slightly widened
above coxae; anterior portion with three spinose tubercles; widened portion notched medially
and with a spinose tubercle anteriorly. Metanotum more or less parallel-sided; sides of median
line granulose. Metapleura widened above coxae, with anterior portion with five small spines;
posterolaterally notched. Prosternum with sensory areas elongate oval, reaching lateral
margin; sensory area on profurcasternum small and circular. Mesosternum with several pairs
of enlarged black granules. Metasternum with few enlarged black granules.
Legs: femora with carinae relatively indistinct except for anterodorsal carina of profemora
which is slightly raised and laterally compressed; dorso-apically ending with a triangular
tubercle. Profemora slightly shorter than mesonotum and curved basally; dorsally with some
minute humps. Mesofemora shorter than profemora; dorsal carinae with flattened and rounded
teeth. Two most distinct teeth on outer dorsal carinae. Ventral carinae posteriorly with a pair
of distinct laterally compressed and blunt teeth; anteriorly with some indistinct granules.
Metafemora about as long as profemora; armed as mesofemora, but armature more distinct.
Tibiae with carinae relatively indistinct, about as long as corresponding femora. Protibiae
with three small humps on posterodorsal carina and two indistinct humps on anterodorsal
carina. Meso- and metatibiae with a small but distinct central tooth, and a smaller posterior,
rounded tooth dorsally. Metatibiae with a second small hump anteriorly. Tarsomeres I–III
about the same length with a posteromedian rounded hump dorsally, IV distinctly shorter.
Claws very small.
Abdomen: median segment strongly transverse, with posterior margin more or less straight
and with indistinct central X-shaped pattern of granules. Abdominal terga rugose; terga II–III
slightly raised mediolongitudinaly; terga III–IV slightly widening and with distinct armature
posterolaterally; posterolateral armature present as enlarged tubercle, mostly triangularly
spinose; tergum IV with two distinct transverse and rounded carinae posteriorly; tergum V
slightly descending obliquely, posterior margin distinctly narrower than anterior one and with
posterior armature as in tergum IV; armature projecting over tergum VI; tergum VI distinctly
descending obliquely, tapering towards the posterior, with an X-shaped mediolongitudinal
carina; tergum VII slightly tapering towards the posterior, with an X-shaped
mediolongitudinal carina and posterolateral spinose tubercle; tergum VIII about as wide as
VII, more or less parallel-sided and slightly ascending; posterolaterally with a spinose
tubercle; IX about as wide as VIII with a distinct elongated and tectiform posteromedial crest;
crest not distinctly notched apically, with apex narrow; sides of tergum IX tuberculose. Anal
segment narrower than tergum IX, obliquely descending and dorsally flattened; anteriorly
with a medial tubercle; oblique carinae distinct, curved and armed with tubercles, lamellate
posteriorly; posterior margin straight with a pair of granules or tubercles. Abdominal sterna
and subgenital plate with a mediolongitudinal carina. Subgenital plate rounded, boat-shaped
with widest part at about mid length; posterior rim dorsoventrally flattened and broadly
rounded apically, almost reaching margin of tergum X. Abdominal sternum VII with a
conspicuous medial compound tubercle anteriorly, merging with mediolongitudinal carina.
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
18
Fig. 8. Orestes spp., eggs. A–C, O. bachmaensis sp. nov. A, lateral view. B, dorsal view. C, operculum. D–F,
O. dittmari sp. nov. D, lateral view. E, dorsal view. F, operculum.
Fig. 9. Orestes bachmaensis sp. nov. in nature, Bach Ma N.P., A–B, adult male, 14.VII.2011. C–D, adult female,
17.VII.2011. E, adult male, IV.2017. F, subadult female nymph, IV.2017.
Belgian Journal of Entomology 58: 1–62 (2018)
19
EGG (Fig. 8 A–C).
Measurements [mm]. Length: 3.6; width: 2.7; height: 2.5.
Egg capsule oval and brown. Operculum suboval, convex and slightly tapering towards the
dorsal side of capsule; ventral margin strongly rounded and distinctly lower than dorsal
margin. Capsule and operculum covered with long setae (about 0.3 mm) with a distal black
three-hooked grapnel-shaped structure. Between long setae some shorter, indistinct hairs
scattered over capsule and operculum. Micropylar plate trilobate with one anterior expansion
and with two posterior expansions. Margins of micropylar plate almost invisible. Anterior
expansion parallel-sided. Apex rounded, reaching edge of capsule. Posterior arms laterally
directed and slightly dilated on lateral surface of capsule, not reaching margins of ventral side;
reaching halfway capsule laterally. Micropylar cup small, black, cup-shaped and protruding.
Table 1. Measurements [mm] of Orestes bachmaensis sp. nov.
Length of HT PT ♂♂ PT ♀♀
Body: 38.2 35.0–43.2 44.0–51.7
Head: 4.8 3.8–5.0 5.0–6.2
Pronotum: 3.1 2.6–3.1 3.3–4.0
Mesonotum: 8.0 6.4–9.0 8.2–11.3
Metanotum: 5.4 5.3–6.2 6.6–7.4
Median segment: 1.5 1.5–1.7 1.8–2.1
Profemora: 7.9 6.3–8.6 7.6–9.3
Mesofemora: 6.4 5.6–6.5 6.9–7.9
Metafemora: 7.4 6.6–8.3 8.7–9.8
Protibiae: 7.4 6.3–7.9 7.0–8.6
Mesotibiae: 5.9 4.7–6.0 5.7–7.0
Metatibiae: 7.3 6.7–7.9 7.7–9.4
DISTRIBUTION: Vietnam, Thua Thien Hue province (Fig. 5).
Orestes botot sp. nov.
urn:lsid:zoobank.org:act:E627C0DC-E48E-4C8D-83A8-24CB0E6239E4
Figs 5, 10–12
ETYMOLOGY. The species epithet is derived from “bò tót”, the Vietnamese name for the gaur
(Mammalia: Bovidae) and refers to the two horns on the head of this species. Gaurs are also
present in Bidoup-Nui Ba National Park, the type-locality of this species of Orestes. It is used
in apposition.
TYPE MATERIAL. VIETNAM. Holotype : Lam Dong prov. Bidoup Nui Ba N.P., 12°26’N
108°30’E, 21-25.VII.2014, night coll. Leg. J. Constant & J. Bresseel, GTI project, I.G.:
32.779 (RBINS)
Paratypes (3♂♂, 2♀♀, 1 nymph, 1 nymph): 2♂♂, 1, 1 nymph, 1 nymph: same data
as HT (2♂♂, 1, 1 nymph, 1 nymph: RBINS; 1: VNMN); 1: same data as HT, ex
breeding Rob Krijns 2015 (VNMN)
DIAGNOSIS. Easy to recognise species. Males with strong supra-coxal spine on mesopleuron
(Fig. 10 D) like the closely related species O. diabolicus sp. nov. (Fig. 13 D), but readily
differentiated by the unarmed median segment and the dark brown colouration with the outer
margins of meso- and metanotum yellowish (Fig. 10 I).
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
20
Fig. 10. Orestes botot sp. nov., holotype . A, habitus, dorsal view. B, habitus, lateral view. C, habitus, ventral
view. D, head and thorax, dorsal view. E, terminalia, dorsal view. F, terminalia anterodorsal view. G, terminalia,
ventral view. H, terminalia, lateral view. I, head and thorax, lateral view.
Belgian Journal of Entomology 58: 1–62 (2018)
21
Females have the posteromedian crest on abdominal tergum IX notched (Fig. 11 F), terga IV–
V almost parallel-sided and with a small but distinct spine on the mesopleura (Fig. 11 A, C);
legs relatively longer when compared to other species (Fig. 11 A).
DESCRIPTION.
MALE (Fig. 10).
Measurements: see table 2.
Head: supra-antennals distinct, conical but blunt and slightly pointed outwards. Posterior
supra-occipitals small, blunt and pointing upwards. Vertex elongated and raised, supra-
orbitals positioned at base of crest, distinctly elongated, slightly incurving and acute apically.
Anterior coronals spinose near apex of crest. Posterior and lateral coronals present as small
granules. Behind eye a distinct carina reaching posterior edge of crest. Eyes relatively small,
circular and strongly projecting hemispherically. Antennae shorter than legs with 23 segments
(n = 4); scapus strongly flattened dorsoventrally and laterally carinated, with a posterolateral
spine. Pedicellus cylindrical, slightly narrowing towards the posterior. First antennomere
longer than following two antennomeres combined. Antennomeres gradually getting longer
till segment XII which is about as long as scapus. Antennal segment XIII shorter, with minute
blunt spine mediolaterally; following segments shiny and distinctly shorter. Apical
antennomere elongated, club-shaped and slightly longer than the two preceding antennomeres
combined.
Thorax: pronotum more or less parallel-sided; lateral margins slightly concave and rugose.
Anterior margin incurved. Prozona with distinct elevation centrally with two to four distinct,
raised tubercles; anterior tubercles slightly larger. Metazona with a pair of granules anteriorly
and posteriorly, and posterior margin straight. Mesonotum rugose with anterior margin
thickened and distinctly concave; lateral margins smooth, slightly concave and posterior
margin slightly wider than anterior one; pre-medial mesonotals spinose and elongated;
posteromedially with two distinct elongated spines, sometimes medially split and bi-apical.
Mesopleura widened above coxae, widened portion notched medially and with an elongated
spine anteriorly. Metanotum unarmed, relatively smooth. Metapleura widened above coxae
and with strong, elongated supra-coxal spine. Prosternum with sensory areas elongate oval.
Sensory area on profurcasternum small and circular. Ventral surface rugose.
Legs: femora with carinae indistinct and posteromedially with a short blunt spine. Profemora
slightly longer than mesonotum and indistinctly curved basally, sometimes with some minute
granules. Mesofemora slightly shorter than mesonotum, posteroventrally with two small,
tuberculose spines laterally. Metafemora about as long as mesonotum and half the pronotum
combined; ventrally armed as mesofemora, dorsally with some minute humps on anterior
carina. Tibiae with carinae indistinct, about as long as corresponding femora and unarmed.
Tarsomeres I–III about the same length with a posteromedian rounded hump dorsally.
Euplantulae more or less circular and of a distinct pale colouration. Claws very small.
Abdomen: median segment unarmed and rugose, with posterior margin concave. Abdominal
terga rugose. Tergum II transverse, slightly narrowing towards the posterior. Terga III–VI
about the same length. Tergum V with posterior margin raised, armed with two granules.
Anterior margin of tergum VI not raised, but with a pair of granules. Tergum VII slightly
shorter than preceding one, with a pair of granules posteriorly. Tergum VIII about as long as
VII, widening towards the posterior; posterior margin concave and slightly raised with a pair
of granules medially; IX shorter than VIII and distinctly transverse, with posterior margin
raised. Anal segment dorsoventrally flattened almost parallel-sided and indistinctly notched
posteromedially. Posterolateral angles rounded. Poculum triangular from lateral view and
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
22
strongly granulose; posterior rim dorsoventrally flattened and slightly notched
posteromedially, projecting over base vomer but not reaching base of apical spine of vomer.
Cerci short, strongly flattened and setose, not reaching apex of tergum X, apices broadly
rounded to almost straight. Vomer well developed, semicircular basally till base of
posteromedial spine; posteromedial spine black in the posterior portion with acute apex; apex
curved, not ending medially but slightly to the right.
FEMALE (Fig. 11).
Measurements: see table 2.
Head: supra-antennals distinct, slightly flattened laterally, blunt and slightly pointed
outwards. Supra-antennals fused basally with anterior supra-occipitals. Anterior supra-
occipitals small, blunt and pointing dorsally. Posterior supra occipitals present as a pair of
granules. Vertex elongated and raised with supra-orbitals strong, laterally compressed and
lamellate. Anterior coronals laterally compressed, lamellate with anterior margin reaching
apex supra-orbitals. Central coronal compound and raised. Posterior and lateral coronals
present as tubercles. Between lateral and anterior coronals, a row of four tubercles on side of
crest. Behind eye, a distinct carina reaching posterior edge of crest. Eyes relatively small,
circular and strongly projecting hemispherically. Antennae shorter than legs with 25
segments (n = 2); scapus strongly flattened dorsoventrally and laterally carinated, with a
posterolateral spine; pedicellus slightly flattened dorsoventrally, slightly narrowing towards
the posterior. First antennomere longer than the following two antennomeres combined.
Antennomeres gradually getting longer till segment XII; segment XIII about as long as XII
and with a granule anterolaterally; following segments shorter except for apical one; apical
antennomere elongated, club-shaped and paler.
Thorax: pronotum trapezoidal, slightly widening towards the posterior; lateral margins
slightly crenulated and anterior margin incurved. Prozona with distinct elevation centrally
with four distinct raised tubercles. Metazona with two parallel rows of three to four granules
and posterior margin straight. Mesonotum rugose with anterior margin thickened, distinctly
concave and strongly granulose; lateral margins more or less smooth; posterior margin wider
than anterior one; posteromedially with two distinct, short elongated carinae pointing
obliquely towards mediolongitudinal line. Mesopleura slightly widened above coxae; anterior
portion with three spinose tubercles; widened portion notched medially and with a spinose
tubercle anteriorly. Metanotum slightly widening towards the posterior, sides of median line
granulose. Metapleura widened above coxae, anteriorly with some small spines followed by a
distinct supra-coxal spine; posterolatelly notched. Prosternum with sensory areas elongate
oval, reaching lateral margin. Sensory area on profurcasternum small and circular.
Mesosternum with several pairs of enlarged black granules dispersed longitudinally.
Metasternum with few enlarged black granules.
Legs: femora with carinae indistinct except for anterodorsal carina of profemora;
posteromedially ending with a short blunt spine. Profemora slightly longer than mesonotum
and curved basally, dorsally with some minute humps. Mesofemora shorter than mesonotum,
posteroventrally with a pair of granules and a pair of conical tubercles posteriorly; dorsally
with several indistinct humps. Metafemora about as long as profemora; ventrally armed as
mesofemora; dorsally with small, lamellate hump subcentrally and subapically. Tibiae with
carinae indistinct, about as long as corresponding femora. Protibiae with three minute humps
on antero- and posterodorsal carina. Mesotibiae with some minute humps dorsally; metatibiae
with a minute hump subanteriorly and centrally. Tarsomeres I–III about the same length with
a posteromedian rounded hump dorsally, IV distinctly shorter. Claws very small.
Belgian Journal of Entomology 58: 1–62 (2018)
23
Fig. 11. Orestes botot sp. nov., paratype . A, habitus, dorsal view. B, habitus, ventral view. C, head and thorax,
dorsal view. D, terminalia, dorsal view. E, terminalia anterodorsal view. F, terminalia, lateral view. G, habitus
lateral view. H, terminalia, ventral view. I, head and thorax, lateral view.
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
24
Fig. 12. Orestes botot sp. nov., in nature. A–B, adult male, Bidoup Nui Ba, 21.VII.2014. C–D, adult female,
same data.
Abdomen: median segment with central X-shaped pattern of granules, strongly transverse and
with posterior margin straight. Abdominal terga rugose; terga II–V with a central X-shaped
pattern of granules; terga II–IV slightly widening; IV–V about parallel-sided and with
posterior armature raised and lamellate. Posterolateral angles of tergum V with a distinct
oblique, short carina. Tergum VI distinctly descending obliquely and strongly narrowing
towards the posterior, medially with a thickened longitudinal carina. Tergum VII more or less
parallel-sided, slightly ascending and with thick X-shaped carina mediolongitudinally.
Tergum VIII about as wide as VII, strongly ascending and with mediolongitudinal carina
slightly diverging at posterior margin; IX about as wide as VIII with two pairs of black
granules in anterior portion and a distinct posteromedial crest; crest notched apically. Anal
segment narrower and shorter than tergum IX, obliquely descending and dorsally flattened;
anteriorly with a median tubercle; oblique carinae present as a row of conical tubercles, only
lamellate posterolaterally; posterior margin straight. Abdominal sterna and subgenital plate
with a mediolongitudinal carina. Subgenital plate rounded, boat-shaped with widest at about
mid length; posterior rim dorsoventrally flattened and slightly tapering towards the posterior.
Apex broadly triangular, almost reaching margin of tergum X.
Table 2. Measurements [mm] of Orestes botot sp. nov.
Length of HT PT ♂♂ PT ♀♀
Body: 44.0 44.2–45 49.8–50.4
Head: 5.0 5.3–5.8 6.8
Pronotum: 2.9 3.3–3.7 4.4–4.5
Mesonotum: 9.0 8.9 9.3–9.6
Metanotum: 5.6 4.7 7.2–7.5
Median segment: 1.5 1.2 2.3
Profemora: 9.4 9.4–10.4 9.4–10.0
Mesofemora: 7.6 7.2–8.7 7.5–7.8
Metafemora: 10.0 10.3–10.7 10.1–11.6
Belgian Journal of Entomology 58: 1–62 (2018)
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Protibiae: 9.6 8.9–9.7 8.5–9.9
Mesotibiae: 7.4 6.8–7.9 6.6–8.7
Metatibiae: 9.7 9.5–10.1 10.6–10.8
DISTRIBUTION. Vietnam, Lam Dong province (Fig. 5).
Orestes diabolicus sp. nov.
urn:lsid:zoobank.org:act:B680C282-DF2B-4593-8FEC-2FBA75D2F34B
Figs 5, 13–14
ETYMOLOGY. The species epithet diabolicus (adj., Latin) means “characteristic of the devil”
and refers to the strongly spinose aspect of the species.
TYPE MATERIAL. VIETNAM. Holotype : Lam Dong prov. Bidoup Nui Ba N.P., 12°26’N
108°30’E, 21-25.VII.2014, night coll. Leg. J. Constant & J. Bresseel, GTI project, I.G.:
32.779 (RBINS).
Paratypes (2♂♂): same data as holotype (1: RBINS; 1: VNMN).
DIAGNOSIS. Easy to recognise species. Males with strong supra-coxal spines on mesopleura,
strong spines on pro-, mesonotum and median segment (Fig. 13 D, I). The reddish brown
colouration, relatively longer legs and body armature (Fig. 13 A) also distinguish this species
from the closely related O. botot sp. nov. Females are unknown.
DESCRIPTION.
MALE (Fig. 13).
Measurements: see table 3.
Head: supra-antennals distinct, conical but blunt and slightly pointed outwards. Supra-
occipitals about as large as supra-antennals, blunt and pointing dorsally. Vertex elongated and
raised; supra-orbitals positioned at base of crest, distinctly elongated, with apices projecting
over sides pronotum, slightly incurving and acute apically. Anterior coronals elongated and
spinose near apex of crest. Central coronal small but acute. Posterior and lateral coronals
present as small granules. Behind eye, a distinct carina reaching posterior edge of crest. Eyes
relatively small, circular and strongly projecting hemispherically. Antennae shorter than legs
with 23 segments (n = 3); scapus strongly flattened dorsoventrally with a posterolateral spine
and a smaller one mediolaterally; pedicellus cylindrical, slightly narrowing towards the
posterior. First antennomere longer than following two ones combined. Antennomeres
gradually getting longer till segment XII which is about as long as scapus. Antennal segment
XIII shorter, with minute blunt spine mediolaterally; following segments shiny and distinctly
shorter. Apical antennomere elongated, slightly longer than the two preceding ones combined
and with apex acute.
Thorax: anterior and posterior margins of pronotum more or less the same width, lateral
margins concave; anterior margin incurved with a small expansion laterally. Prozona with
distinct median line and elevation centrally armed with four distinct raised tubercles.
Metazona with a pair of distinct spinose inter-posterior pronotals. Mesonotum with anterior
margin thickened and concave; lateral margins smooth with posterior margin slightly wider
than anterior one; posteromedially, an inverted V-shaped crest with two distinct spines or
tubercles; anterior margin concave. Mesopleura widened above coxae; widened portion
notched medially with a spine anteriorly. Metanotum rugose with anterior margin convex,
slightly narrowing towards the posterior; suture with median segment indistinct. Metapleura
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
26
Fig. 13. Orestes diabolicus sp. nov., holotype . A, habitus, dorsal view. B, habitus, lateral view C, habitus,
ventral view. D, head and thorax, dorsal view. E, terminalia, dorsal view. F, terminalia anterodorsal view.
G, terminalia, lateral view. H, terminalia, ventral view. I, head and thorax, lateral view.
Belgian Journal of Entomology 58: 1–62 (2018)
27
Fig. 14. Orestes diabolicus sp. nov. in nature. A–B, adult male Bidoup Nui-Ba N.P., 25.VII.2014.
widened above coxae and with strong, elongated supra-coxal spine; anteriorly and posteriorly
of supra-coxal two minute lateral spines. Prosternum with sensory areas almost circular.
Sensory area on profurcasternum small and circular. Mesosternum with four spinose tubercles
and metasternum with two spinose tubercles.
Legs: femora with carinae indistinct and posteromedially with a slightly elongated dorsal
spine. Profemora slightly shorter than pro- and mesonotum combined, indistinctly curved
basally. Mesofemora about as long as mesonotum. Metafemora about as long as profemora.
Tibiae with carinae indistinct, slightly longer than corresponding femora and unarmed.
Tarsomeres I–III about the same length with a posteromedian rounded hump dorsally.
Euplantulae more or less circular and distinctly paler in colour. Claws very small.
Abdomen: median segment with two strong, elongated spines projecting posterodorsally;
posterior margin straight. Abdominal segments relatively smooth. Tergum II distinctly
transverse and slightly wider anteriorly; terga III–IV with a small bituberculate hump
posteromedially and terga V–VIII with a bispinose hump posteromedially; tergum VII
slightly shorter than preceding one; tergum VIII about as long as VII, widening towards the
posterior and distinctly concave posteriorly; IX shorter than VIII and distinctly transverse
with posteromedian and posterolateral humps. Anal segment dorsoventrally flattened and with
a distinct tooth mediolaterally; apex notched, with posterolateral angles rounded. Poculum
triangular from lateral view and strongly granulose; posterior rim dorsoventrally flattened and
deeply notched posteromedially, projecting over base vomer but not reaching base of apical
spine. Cerci short, strongly flattened and setose, not reaching apex of tergum X and with
apices broadly rounded to almost straight. Vomer well developed, semicircular basally till
base of posteromedial spine; posteromedial spine black in the posterior portion with acute
apex; apex curved not ending medially but slightly to the right.
Table 3. Measurements [mm] of Orestes diabolicus sp. nov.
Length of HT PT ♂♂
Body: 40.1 41.0–41.8
Head: 5.3 5.5–5.6
Pronotum: 3.2 3.2–3.4
Mesonotum: 8.0 8.1–8.3
Metanotum: 5.6 5.5–6.4
Median segment: 2.2 2.1–2.7
Profemora: 9.7 10.7–11.2
Mesofemora: 8.0 8.5–8.7
Metafemora: 10.5 10.6–11.4
Protibiae: 10.7 11.1–11.4
Mesotibiae: 8.3 8.7–9.1
Metatibiae: 10.9 12–12.1
DISTRIBUTION. Vietnam, Lam Dong province (Fig. 5).
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
28
Orestes dittmari sp. nov.
urn:lsid:zoobank.org:act:291488DE-BFCA-45BA-A545-F886FB59BB3E
Figs 5, 8 D–F, 15–17
ETYMOLOGY. The species is named after Mr Daniel Dittmar (Germany) in acknowledgement
for breeding several stick insect species that we brought back from Southeast Asia.
TYPE MATERIAL. VIETNAM. Holotype : Vietnam, Cat Ba N.P., 20°48’00”N 107°00’20”E,
12-16.VII.2013, leg. J. Constant & J. Bresseel, I.G.: 32.454 (RBINS).
Paratypes (1, 2 eggs): 1: same data as HT (RBINS).
DIAGNOSIS. Males have the mesonotum distinctly elevated posteriorly (Fig. 15 I); the cephalic
armature is not fused, the anterior coronals are laterally compressed and rounded apically
(Fig. 15 I).
Females have a low crest on the head (Fig. 16 I); the body armature is never acute and the
abdomen is distinctly dilated laterally (Fig. 16 A).
DESCRIPTION.
MALE (Fig. 15).
Measurements: see table 4.
Head: supra-antennals strongly spinose and slightly pointed outwards, about as long as supra-
orbitals; supra-occipitals smaller, the posterior being the smallest. Vertex raised; supra-
orbitals strong, conical and spinose. Anterior coronals strongly flattened laterally and rounded
apically. Central coronal conical, slightly longer than posterior coronals. Posterior and lateral
coronals present as small, conical tubercles. Behind eye, a distinct carina reaching posterior
edge of crest resulting in a conical tubercle. Eyes relatively small, circular and strongly
projecting hemispherically. Antennae shorter than legs, with 23 segments (n = 1); scapus
strongly flattened dorsally, laterally carinated and with a spine laterally and posterolaterally;
pedicellus slightly flattened dorsoventrally, narrowing towards the posterior. First
antennomere about as long as pedicellus. Antennomeres gradually getting longer till segment
XIII. Antennal segment XIII the longest, with minute granule anterolaterally; following
segments shiny and distinctly shorter. Apical antennomere elongated, club-shaped.
Thorax: pronotum rugose, more or less parallel-sided, indistinctly wider posteriorly; almost
quadrate with anterior margin incurved. Prozona with central elevation with four tubercles;
anterior tubercles slightly larger. Metazona slightly lower than prozona, and with three pairs
of granules. Mesonotum rugose with anterior margin thickened, distinctly concave and with a
pair of minute granules anteromedially; posterior portion slightly raised anterolaterally and
distinctly conically raised posteromedially; posteromedial elevation indistinctly V-shaped and
sulcate centrally. Mesopleura widened above coxae; widened portion rounded, slightly
crenulated and indistinctly notched medially. Metanotum more or less parallel-sided, rugose
and slightly raised posterolaterally. Metapleura widened above coxae, rounded and slightly
crenulated. Prosternum with sensory areas elongate oval. Sensory area on profurcasternum
small and circular. Ventral surface rugose.
Belgian Journal of Entomology 58: 1–62 (2018)
29
Fig. 15. Orestes dittmari sp. nov., holotype . A, habitus, dorsal view. B, habitus, lateral view. C, habitus,
ventral view. D, head and thorax, dorsal view. E, terminalia, dorsal view. F, terminalia anterodorsal view. G,
terminalia, lateral view. H, terminalia, ventral view. I, head and thorax, lateral view.
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
30
Fig. 16. Orestes dittmari sp. nov., paratype . A, habitus, dorsal view. B, habitus, ventral view. C, head and
thorax, dorsal view. D, terminalia, dorsal view. E, Terminalia anterodorsal view. F, terminalia, lateral view.
G., habitus lateral view. H, terminalia, ventral view. I, head and thorax, lateral view.
Belgian Journal of Entomology 58: 1–62 (2018)
31
Legs: femora with carinae relatively indistinct except for the anterolateral carina of the
profemora, dorsal posterior margin with indistinct triangular tubercle. Profemora slightly
shorter than mesonotum, curved basally and unarmed. Mesofemora about as long metanotum
and median segment combined with some indistinct granules; outer ventral carina with two
granules subapically. Metafemora about as long as profemora; ventrally armed as
mesofemora; dorsally with a definite blunt tooth centrally and subapically. Tibiae with carinae
relatively indistinct, slightly shorter than corresponding femora; dorsally with some indistinct
granules. Tarsomeres I–III about the same length with a posteromedian rounded hump
dorsally. Claws very small.
Abdomen: median segment unarmed and rugose, indistinctly raised medially. Abdominal
terga rugose; terga II–VI about the same length; tergum II transverse, slightly narrowing
towards the posterior; terga III–VII parallel-sided; II–VIII with a pair of minute granules
posteromedially; tergum VII slightly shorter than VI; tergum VIII slightly wider than VII,
transverse and widening towards the posterior, with posterior margin concave; IX distinctly
transverse, with posterior margin with posteromedian ridge, raised apically. Anal segment
narrower than tergum IX, dorsoventrally flattened, almost parallel-sided and indistinctly
concave posteromedially; posterolateral angles rounded and medially with a transverse row of
three granules and laterally a larger granule in middle. Poculum rounded and strongly
granulose; posterior rim dorsoventrally flattened and broadly rounded anteriorly. Cerci short,
strongly flattened and setose, not reaching apex of tergum X, with apices broadly rounded to
almost straight. Vomer well developed, semicircular basally till base of posteromedial spine;
posteromedial spine broad and blackish in the posterior portion and with apex curved.
FEMALE (Fig. 16).
Measurements: see table 4.
Head: supra-antennals short, conical and blunt. Anterior supra-occipitals present as small
conical humps, posterior supra-occipitals smaller and granulose. Vertex indistinctly raised,
supra-orbitals short, laterally compressed and rounded apically. Anterior coronals fused with
supra orbitals, laterally compressed and lamellate. Central coronal indistinct, only present as
small granule. Posterior and lateral coronals present as conical tubercles. Postocular carina
distinct, apex as triangular tubercle. Eyes relatively small, circular and strongly projecting
hemispherically. Antennae shorter than legs and with 25 segments (n = 1); scapus strongly
flattened dorsally and laterally carinated, with a definite spine posterolaterally and a shorter
spine mediolaterally; pedicellus slightly flattened dorsoventrally, slightly narrowing towards
the posterior. First antennomere longer than the following two antennomeres combined.
Apical antennomere elongated and club-shaped.
Thorax: pronotum trapezoidal, slightly widening towards the posterior; lateral margins
rugose, anterior margin incurved. Prozona with distinct elevation centrally with four granules.
Metazona with some indistinct granules and a pair of tubercles posteriorly and with posterior
margin straight. Mesonotum rugose, indistinctly widening towards the posterior; anterior
margin thickened and distinctly concave; lateral margins posterolaterally with some small
granules. Mesopleura slightly widened above coxae; widened portion notched medially and
slightly crenulated. Metanotum as mesonotum. Metapleura as mesopleura but posterolaterally
notched. Prosternum with sensory areas elongate oval, reaching lateral margins. Sensory area
on profurcasternum small and circular. Mesosternum with few enlarged black granules.
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
32
Fig. 17. Orestes dittmari sp. nov. in nature. A, adult male, Cat Ba N.P., 13.VII.2013. B, adult female, same
location, 7.VII.2013.
Legs: femora with carinae indistinct; posteromedially ending with a short dorsal blunt spine.
Profemora slightly shorter than mesonotum and curved basally. Mesofemora about as long as
metanotum and median segment combined, dorsally with some minute humps;
posteroventrally with two pair of granules. Metafemora about as long as profemora; ventrally
armed as mesofemora, dorsally with a small hump anteriorly, a lamellate tooth centrally and
a rounded tooth subapically. Tibiae with carinae indistinct, slightly shorter than corresponding
femora. Protibiae with three minute humps on posterodorsal carina. Mesotibiae with some
minute humps dorsally, metatibiae with a minute hump subanteriorly and a larger one
centrally. Tarsomeres I–III about the same length with a posteromedian rounded hump
dorsally, IV distinctly shorter. Claws very small.
Abdomen: median segment rugose, unarmed, strongly transverse and with posterior margin
straight. Abdominal terga rugose; terga II–III as median segment but anteriorly with two pairs
of sublateral black granules; terga II–IV distinctly widening towards the posterior; III–IV with
posterolateral tubercle and IV with indistinct granulose mediolongitudinal carina and
posteriorly with twin-crescent-shaped crest; V distinctly narrowing towards the posterior and
slightly descending, with X-shaped carina, and posterior armature raised and projecting over
posterior margin; tergum VI distinctly descending obliquely and strongly narrowing towards
the posterior, medially with a thickened longitudinal carina; tergum VII slightly narrowing
towards the posterior, slightly ascending and with indistinct carina mediolongitudinally;
tergum VIII about as wide as VII, with posterolateral angles with distinct tubercle and
posteromedially with a pair of granules; IX slightly narrower than VIII with central granule
and a distinct posteromedial crest; crest relatively short, not notched apically. Anal segment
narrower and shorter than tergum IX, obliquely descending and dorsally flattened; anteriorly
with a tubercle medially; oblique carinae present as a row of conical tubercles, only lamellate
posterolaterally; posterior margin straight, slightly crenulated. Abdominal sterna and
subgenital plate with an indistinct mediolongitudinal line. Subgenital plate rugose, boat-
shaped with widest part at about mid length; posterior rim dorsoventrally flattened and
slightly tapering towards the posterior. Apex broadly rounded, almost reaching margin of
tergum X.
EGG (Fig. 8 D–F).
Measurements [mm]. Length: 3.5; width: 2.9; height: 3.2.
Belgian Journal of Entomology 58: 1–62 (2018)
33
Egg capsule subspherical and brown. Capsule covered with minute pits. Operculum
subcircular, slightly convex. Capsule and operculum covered with long setae (about 0.3 mm)
with a distal black three-hooked grapnel-shaped structure. Base of long setae surrounded by
very short hairs, creating small circular patches on capsule and operculum. Micropylar plate
trilobate with one anterior expansion and with two posterior expansions. Anterior expansion
almost parallel-sided with apex tapering and rounded, almost reaching edge capsule. Posterior
arms laterally directed and strongly dilated on lateral surface of capsule, reaching margins of
ventral side; almost covering completely lateral side of capsule and almost reaching anterior
margin of capsule. Margin of micropylar plate indistinctly raised and demarcated by more
dense setae. Micropylar cup small, black, cup-shaped and protruding.
Table 4. Measurements [mm] of Orestes dittmari sp. nov.
Length of HT PT
Body: 43.5 50.8
Head: 4.5 6.3
Pronotum: 2.8 4.6
Mesonotum: 8.2 10.8
Metanotum: 6.2 7.6
Median segment: 1.2 1.4
Profemora: 7.8 8.2
Mesofemora: 6.3 7.0
Metafemora: 6.8 8.3
Protibiae: 6.8 7.7
Mesotibiae: 4.8 6.3
Metatibiae: 6.1 8.0
DISTRIBUTION. Vietnam, Ha Long Bay, Cat Ba Island (Fig. 5).
Orestes draegeri sp. nov.
urn:lsid:zoobank.org:act:7A7812DD-F531-4F71-9DA7-B5CBE925FB1B
Figs 5, 18–20
ETYMOLOGY. The species is named after Mr Holger Dräger (Germany) in acknowledgement
for breeding several stick insect species that we brought back from Southeast Asia.
TYPE MATERIAL. VIETNAM. Holotype : Vietnam, Cat tien N.P., 11°20’N 107°20’E, 6-
16.VII.2012, leg. J. Constant & J. Bresseel, I.G.: 32.161 (RBINS).
Paratypes (2♂♂, 12♀♀): 4♀♀: same data as HT (2♀♀: RBINS; 2♀♀: VNMN); 2♂♂, 8♀♀:
Vietnam: Dong Nai Biosphere Res., 11°18’N 107°06’E, 25.VI-6.VII.2012, leg. J. Constant &
J. Bresseel, I.G.: 32.161, ex breeding R. Krijns (1, 6♀♀: RBINS; 1, 2♀♀: VNMN).
DIAGNOSIS. Slender elongate species with short legs (Fig. 18 A, 19 A). Closely related to
Orestes mouhotii Bates, 1865 but distinguishable by the proportionally shorter legs (see
Tables 5 and 7).
Males have the posterior margin of mesonotum conically raised and sulcate medially (Fig. 18
D, I).
Females with cephalic armature indistinct (Fig. 19 I), tergum IV indistinctly widening
towards the posterior, almost parallel-sided (Fig. 19 A). Metatibiae with dorsal, laterally
compressed, teeth anteriorly and centrally. The central tooth being more distinct.
Posteromedial crest on tergum IX distinctly elongated and strongly tectiform (Fig. 19 E–F).
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
34
Fig. 18. Orestes draegeri sp. nov., holotype . A, habitus, dorsal view. B, habitus, lateral view. C, habitus,
ventral view. D, head and thorax, dorsal view. E, terminalia, dorsal view. F, terminalia anterodorsal view.
G, terminalia, ventral view. H, terminalia, lateral view. I, head and thorax, lateral view.
Belgian Journal of Entomology 58: 1–62 (2018)
35
DESCRIPTION
MALE (Fig. 18).
Measurements: see table 5.
Head: supra-antennals spinose and slightly pointed outwards, followed by four smaller,
spinose supra-occipitals. Vertex with anterior coronals distinctly enlarged, blunt and laterally
compressed, creating ear-like processes. Supra-orbitals bluntly spinose. Posterior and lateral
coronals present as small, spinose tubercles forming a corona with the apex of the postocular
carina. Eyes relatively small, circular and strongly projecting hemispherically. Antennae
shorter than legs, with 23 segments (n = 3); scapus strongly flattened dorsoventrally and
laterally carinate with a spine laterally and posterolaterally; pedicellus slightly dorsoventrally
flattened. First antennomere distinctly shorter than scapus. Antennomeres gradually getting
longer till segment XII. Antennal segment XIII the longest, with minute hump anterolaterally;
following segments shiny and distinctly shorter. Apical antennomere elongated, slightly
longer than the two preceding antennomeres combined.
Thorax: pronotum almost parallel-sided, slightly wider posteriorly, longer than wide and
rugose and with anterior margin incurved. Prozona slightly elevated centrally. Posterior
margin indistinctly rounded. Mesonotum with anterior margin concave; slightly bent laterally;
posterior margin wider than anterior one with a distinct conical hump posteromedially. Hump
slightly sulcate medially. Mesopleura slightly widened above coxae; widened portion rounded
and notched medially. Metanotum concave laterally, posteromedially with two minute
granules; posterior margin well marked. Metapleura widened above coxae, rounded and
notched posterolaterally. Prosternum with sensory areas elongate oval. Sensory area on
profurcasternum small and subcircular. Ventral surface of body rugose.
Legs: profemora unarmed, curved basally with carinae more distinct compared to other
femora, except for the medioventral carina; posterior margin with small blunt spine dorsally;
about as long as metanotum and median segment combined. Mesofemora unarmed, only
slightly shorter than profemora and with especially dorsal carinae indistinct. Metafemora
about as long as profemora; unarmed or with one to two humps dorsally. Protibiae with inner
dorsal carina more distinct, slightly shorter than corresponding femora and unarmed. Meso-
and metatibiae with carinae indistinct, slightly shorter than corresponding femora and
unarmed. Tarsomeres I–III about the same length with a posteromedian rounded hump
dorsally. Claws very small.
Abdomen: median segment unarmed, rugose and strongly transverse. Abdominal terga rugose,
terga II–V about the same length; tergum VI slightly shorter; tergum II almost quadrate,
slightly narrowing towards the posterior; terga III–VII parallel-sided posteromedially,
unarmed or sometimes armed with two minute granules posteromedially. Posterior margin of
tergum VII concave; tergum VIII slightly wider than VII, trapezoidal and widening towards
the posterior, with posterior margin concave; IX about as long as VIII, transverse, with
posterior margin slightly raised with minute granules. Anal segment dorsoventrally flattened,
parallel-sided and notched posteromedially. Posterolateral angles broadly rounded. Poculum
more or less rounded and strongly granulose; posterior rim dorsoventrally flattened, thickened
and broadly rounded posteriorly. Poculum projecting over base of vomer but not reaching
base of apical spine. Cerci short, strongly flattened and setose, not reaching apex of tergum X,
with apices broadly rounded to almost straight. Vomer well developed, semicircular basally
till base of posteromedial spine; posteromedial spine broad, strongly upcurving with an angle
of almost 90° and blackish in posterior portion.
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
36
Fig. 19. Orestes draegeri sp. nov., paratype from Dong Nai Biosphere Reserve. A, habitus, dorsal view.
B, habitus, ventral view. C, head and thorax, dorsal view. D, terminalia, dorsal view. E, Terminalia anterodorsal
view. F, terminalia, lateral view. G, habitus lateral view. H, terminalia, ventral view. I, head and thorax, lateral
view.
Belgian Journal of Entomology 58: 1–62 (2018)
37
FEMALE (Fig.19).
Measurements: see table 5.
Head: supra-antennals, supra-occipitals, posterior- and lateral coronals present as minute
granules, posterior coronals more distinct. Postocular carina distinct, apically with minute
granule. Other armature indistinct. Vertex not elongated or raised but slightly rounded.
Between lateral coronal and supra-orbital two equally sized granules. Eyes relatively small,
circular and strongly projecting hemispherically. Antennae slightly shorter than legs, with 25
segments (n = 6); scapus strongly flattened dorsoventrally, laterally carinate and with a central
hump and subapical blunt spine on outer lateral carina; pedicellus slightly flattened
dorsoventrally, almost parallel-sided. First antennomere slightly longer than following two
antennomeres combined; antennal segments IV to XV gradually becoming longer; following
segments shorter except for apical antennomere which is elongated, club-shaped.
Thorax: pronotum trapezoidal, widening towards the posterior; anterior margin strongly
convex. Prozona without elevation centrally and with few minute granules. Metazona with
two to six minute granules and posterior margin slightly convex. Mesonotum rugose with
anterior margin incurved. Anterior portion with four evenly spaced, paired, black granules.
First pair posteriorly, last pair subcentrally. Posterior margin slightly wider than anterior one.
Mesopleura slightly widened above coxae; widened portion slightly crenulated, notched
medially. Metanotum rugose, more or less parallel-sided and concave posteriorly. Metapleura
slightly widened above coxae, anterior portion with few small tubercles, posterolatelly
notched and posteriorly with minute teeth. Prosternum with sensory areas elongate oval, not
reaching lateral margins. Sensory area on profurcasternum small and rounded; anterior margin
more or less straight. Meso- and metasternum relatively smooth with few granules.
Legs: femora with carinae relatively indistinct except for anterodorsal and ventral carinae of
profemora; apex with small granule. Profemora rugose, shorter than mesonotum and curved
basally. Mesofemora about as long as metanotum and median segment combined, dorsally
with some laterally compressed, small granules. Metafemora about as long as profemora, with
two triangular tooth dorsally, one centrally and one subapically. Tibiae with carinae relatively
distinct and distinctly shorter than corresponding femora. Protibiae carinate and unarmed.
Mesotibiae with a central rounded tooth dorsally. Metatibiae with a subanterior and central
rounded tooth dorsally, central one more distinct. Tarsomeres I–III about the same length with
a square posteromedian hump dorsally, IV distinctly shorter. Claws very small.
Abdomen: median segment rugose, strongly transverse and with posterior margin convex.
Abdominal terga rugose; terga II–IV slightly, gradually widening towards the posterior;
tergum IV with posterolateral triangular tubercle; posteromedially with a twin-crescent-
shaped granulose crest; tergum V slightly descending obliquely with central pair of granules,
and posterior margin slightly narrower than anterior one; tergum V with a mediolongitudinal
carina, diverging posteriorly, resulting into flattened triangular tubercles; triangular tubercles
projecting over posterior margin of tergum V; tergum VI distinctly descending obliquely and
with a mediolongitudinal carina, posterior margin distinctly narrower than anterior one and
incurved; tergum VII almost parallel-sided, slightly tapering towards the posterior with a
mediolongitudinal carina and incurved posteriorly; tergum VIII more or less parallel-sided,
and slightly ascending; posterolaterally with a spinose granule and posteromedially with a
pair of minute granules; IX slightly narrower than VIII with a distinct elongated and tectiform
posteromedial crest; crest narrow apically; posterolateral margins of IX rounded and with
several granules. Anal segment narrower than tergum IX and dorsally flattened; anteriorly
with a median tubercle; sublaterally with an oblique row of tubercles reaching posterolateral
angles; posterolateral angles lamellate and raised; posterior margin straight and granulose.
Subgenital plate with a mediolongitudinal carina, more distinct posteriorly; rounded and boat-
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
38
shaped with widest part at about mid length; posterior rim dorsoventrally flattened; apex
rounded, almost reaching margin of tergum X.
Table 5. Measurements [mm] of Orestes draegeri sp. nov.
Length of HT PT ♂♂ PT ♀♀
Body: 40.2 40.0–41.2 45.7–47.1
Head: 3.7 3.9–4.2 4.6–4.8
Pronotum: 2.5 2.8–2.9 3.6–3.9
Mesonotum: 8.4 8.1–8.4 9.6–9.7
Metanotum: 6.0 5.4–6.1 6.4–6.5
Median segment: 1.3 1.2–1.4 1.8
Profemora: 6.3 6.7–6.8 7.2–7.8
Mesofemora: 5.4 5.6–5.9 6.3–6.9
Metafemora: 6.3 5.8–6.0 7.2–7.5
Protibiae: 5.3 5.9 6.3–6.5
Mesotibiae: 4.3 4.6 5.2–5.8
Metatibiae: 5.6 5.5–5.6 6.3–6.5
DISTRIBUTION. Vietnam, Dong Nai and Lam Dong provinces (Fig. 5).
Orestes krijnsi sp. nov.
urn:lsid:zoobank.org:act:69C8EAFB-558E-498B-ADCB-764B2499B5C8
Figs 5, 21–22, 23 A–C, 24
ETYMOLOGY. The species is named after Mr Rob Krijns (Netherlands) in acknowledgement
for breeding several stick insect species that we brought back from Southeast Asia.
TYPE MATERIAL. VIETNAM. Holotype : Vietnam, Ninh Thuan prov., Nui Chua N. P.,
11°42’N 109°09’E, 3-9.VII.2014, night coll. Leg. J. Constant & J. Bresseel, GTI project I.G.:
32.779 (RBINS).
Fig. 20. Orestes draegeri sp. nov. in nature. A, adult male, Dong Nai Biosphere Reserve, 30.VI.2012. B, adul
t
female, same data. C–D, adult female Cat Tien N.P., 8.VII.2012.
Belgian Journal of Entomology 58: 1–62 (2018)
39
Paratypes (3♂♂, 4♀♀, eggs): 3♂♂, 3♀♀, eggs: same data as HT, ex breeding R. Krijns, 2017
(2♂♂, 2♀♀, eggs: RBINS; 1, 1: VNMN); 1: same data as HT, ex breeding D. Dittmar
2017 (RBINS).
DIAGNOSIS. Males have the posterior margin of metanotum distinctly raised (Fig. 21 I). Head
with anterior coronals spinose, not laterally compressed (Fig. 21 I).
Females with cephalic armature except central coronal present as granules (Fig. 22 I), tergum
IV distinctly widening towards the posterior (Fig. 22 A). Metatibiae unarmed. Posteromedial
crest on tergum IX distinctly elongated and strongly tectiform (Fig. 22 E–F).
DESCRIPTION.
MALE (Fig. 21).
Measurements: see table 6.
Head: supra-antennals conically spinose and slightly pointed outwards followed by two
spinose supra-occipitals. Vertex with conically spinose anterior coronals. Supra-orbitals
spinose. Posterior and lateral coronals present as small, conical tubercles forming a corona
with the elongated apex of the postocular carina. Eyes relatively small, circular and strongly
projecting hemispherically. Antennae shorter than legs with 23 segments (n = 1); scapus
strongly flattened dorsoventrally and laterally carinate, with a spine laterally and
posterolaterally; pedicellus dorsoventrally flattened slightly tapering towards the posterior.
First antennomere distinctly longer than pedicellus. Antennomeres gradually getting longer
till segment XII. Antennal segment XIII the longest, with minute hump laterally; following
segments shiny and distinctly shorter. Apical antennomere elongated.
Thorax: pronotum trapezoidal, slightly wider posteriorly, almost quadrate and tuberculose;
anterior margin incurved, prozona slightly elevated centrally with four granules, metazona
with four granules; posterior margin rounded. Mesonotum with anterior margin slightly
elevated with a pair of minute granules; subcentrally, a pair of minute granules; posterior
margin distinctly raised with anterolateral portions tuberculose, posteromedially conically
raised, compound and sulcate centrally. Mesopleura widened above coxae; widened portion
rounded and slightly crenulated laterally. Metanotum more or less parallel-sided, with
posterior margin distinctly raised, posterolaterally with a single tubercle and posteromedially
with a pair of raised tubercles. Metapleura widened above coxae with a single blunt spine and
notched posterolaterally. Prosternum with sensory areas elongate oval. Sensory area on
profurcasternum small and subcircular. Ventral surface of body rugose.
Legs: profemora unarmed, curved basally with anterodorsal carinae slightly more distinct;
posterior margin with triangular tubercle; about as long as metanotum. Mesofemora unarmed
with carinae indistinct; only slightly shorter than profemora, about as long as metanotum and
median segment combined. Metafemora about as long as profemora, shaped like mesofemora.
Protibiae unarmed, slightly shorter than corresponding femora. Meso- and metatibiae with
carinae indistinct, slightly shorter than corresponding femora and unarmed. Tarsomeres I–III
about the same length with a posteromedian rounded hump dorsally. Claws very small.
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
40
Fig. 21. Orestes krijnsi sp. nov., holotype . A, habitus, dorsal view. B. habitus, lateral view. C, habitus, ventral
view. D, head and thorax, dorsal view. E, terminalia, dorsal view. F, terminalia anterodorsal view. G, terminalia,
ventral view. H, terminalia, lateral view. I, head and thorax, lateral view.
Belgian Journal of Entomology 58: 1–62 (2018)
41
Abdomen: median segment rugose, strongly transverse and slightly raised posteriorly.
Abdominal terga rugose; terga II–V about the same length; tergum VI slightly shorter; tergum
II almost quadrate, slightly narrowing towards the posterior; Terga III–VII more or less
parallel-sided, unarmed or sometimes armed with two minute granules posteromedially,
granules more distinct towards the posterior; tergum VII about as wide as long; tergum VIII
wider than VII, trapezoidal and widening towards the posterior, with posterior margin
concave; IX about as long as VIII, with posterior margin with distinct posteromedial raised
carina and posterolateral margin with a tubercle. Anal segment dorsoventrally flattened with a
lateral spine in the anterior half, and notched posteromedially. Posterolateral angles broadly
rounded. Poculum more or less rounded and strongly granulose; posterior rim dorsoventrally
flattened, thickened and broadly rounded posteriorly. Poculum projecting over base of vomer,
reaching base of apical spine. Cerci short, strongly flattened and setose, not reaching apex of
tergum X, with apices broadly rounded to almost straight. Vomer well developed,
semicircular basally till base of posteromedial spine; posteromedial spine broad, strongly
upcurved, with apex black.
FEMALE (Fig. 22).
Measurements: see table 6.
Head: complete armature except central coronal present as granules, supra-antennals slightly
larger and conical. Vertex not elongated or raised but slightly rounded. Between lateral
coronal and supra-orbital, two equally sized granules. Behind eye a distinct carina reaching
posterior edge of crest, resulting in an elongated granule. Eyes relatively small, circular and
strongly projecting hemispherically. Antennae slightly shorter than legs with 25 segments
(n=4); scapus strongly flattened dorsoventrally, laterally carinate and with a central and
subapical, blunt spine on the outer lateral carina; pedicellus slightly flattened dorsoventrally,
narrowing towards the posterior. First antennomere longer than the following two
antennomeres combined; antennal segment IV to XV gradually becoming longer. Segment
XV with a minute spine anterolaterally, following segments shorter except for apical
antennomere which is elongated, club-shaped and distinctly longer.
Thorax: pronotum trapezoidal, widening towards the posterior; anterior margin strongly
convex. Prozona with elevation centrally with four raised granules. Metazona with two to six
granules slightly convex, posterolaterally with a small granule. Mesonotum rugose with
anterior margin thickened, strongly incurved and with two granules anteromedially; lateral
margins with few evenly spaced granules; posterior margin wider than anterior margin;
subcentrally with a raised pair of granules. Posterolateral and posterior margins granulose and
slightly raised. Posterior margin concave. Mesopleura slightly widened above coxae; widened
portion distinctly notched medially and with a granule anteriorly. Metanotum more or less
parallel-sided, posterolateral and posterior margin granulose and indistincly raised.
Posteromedially with a pair of granules Metapleura widened above coxae, anterior portion
with few small tubercles, posterolatelly notched. Prosternum with sensory areas elongate oval,
not reaching lateral margins. Sensory area on profurcasternum small and more or less circular.
Meso- and metasternum relatively smooth with few granules.
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
42
Fig. 22. Orestes krijnsi sp. nov., paratype . A, habitus, dorsal view. B, habitus, ventral view. C, head and
thorax, dorsal view. D, terminalia, dorsal view. E, Terminalia anterodorsal view. F, terminalia, lateral view.
G, habitus lateral view. H, terminalia, ventral view. I, head and thorax, lateral view.
Belgian Journal of Entomology 58: 1–62 (2018)
43
Legs: femora with carinae relatively indistinct except for anterodorsal carina of profemora;
apex with small granule. Profemora rugose slightly shorter than mesonotum and curved
basally. Mesofemora slightly shorter than profemora, dorsally with some small granules.
Metafemora about as long as profemora with three evenly spaced raised granules dorsally,
central one being most prominent. Tibiae with carinae indistinct and unarmed, pro- and
metatibiae about as long as corresponding femora, mesotibiae slightly shorter than
corresponding femora. Tarsomeres I–III about the same length with a square posteromedian
hump dorsally, IV distinctly shorter. Claws very small.
Abdomen: median segment rugose and strongly transverse and with posterior margin more or
less straight. Abdominal terga rugose; terga II–IV distinctly but gradually widening towards
the posterior and with lateral margins oblique; tergum IV with a posterolateral spinose
tubercle, posteromedially with a twin-crescent-shaped granulose crest; tergum V slightly
descending obliquely, with posterior margin slightly narrower than anterior one, with a
mediolongitudinal carina resulting into two flattened triangular tubercles posteromedially, the
triangular tubercles projecting over posterior margin of tergum V, and posterolaterally with a
spinose tubercle; tergum VI distinctly descending obliquely, with posterior magin distinctly
narrower with a posteromedial diverging mediolongitudinal carina; anterolaterally with a
small granule; tergum VII almost parallel-sided, slightly tapering towards the posterior;
tergum VIII about as wide as VII but distinctly longer, more or less parallel-sided and slightly
ascending; posterolaterally with a spinose granule and posteromedially with a pair of
granules; IX about as wide as VIII with a distinct elongated and tectiform posteromedial crest,
the crest not notched apically, apex narrow; posterior margin rounded; sides and crest of
tergum IX granulose. Anal segment narrower than tergum IX and dorsally flattened;
anteriorly with a tubercle medially, sublaterally with an oblique row of two tubercles reaching
posterolateral angles; posterior margin straight but granulose. Subgenital plate with a
mediolongitudinal carina, rounded, boat-shaped with widest part at about mid length;
posterior rim dorsoventrally flattened, narrowing towards the posterior; apex rounded, almost
reaching margin of tergum X. Abdominal sterna with a pair of minute granules
posteromedially.
EGG (Fig. 23 A–C).
Measurements [mm]. Length: 3.4; width: 2.0; height: 2.5.
Egg capsule oval, slightly laterally compressed and brown. Operculum narrow, convex and
strongly tapering towards the dorsal side of capsule; narrow margin visible in dorsal view.
Ventral margin slightly wider and rounded. Capsule and operculum covered with long setae
(about 0.3 mm) with a distal black three-hooked grapnel-shaped structure. Micropylar plate
trilobate with one anterior expansion and with two posterior expansions. Margins of
micropylar plate almost invisible. Anterior expansion parallel-sided, reaching margin of
operculum. Posterior arms laterally projecting but margins obsolete. Micropylar cup small,
black, cup-shaped and protruding.
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
44
Fig. 23. Orestes spp., eggs. A–C, O. krijnsi sp. nov. A, lateral view. B, dorsal view. C, operculum. D–F,
O. mouhotii (Bates, 1865) (photographs by F. Tetaert). D, lateral view. E, dorsal view. F, operculum. G–I,
O. subcylindricus (Redtenbacher, 1906) comb. nov. G, lateral view. H, dorsal view. I, operculum.
Fig. 24. Orestes krijnsi sp. nov., adult pair in nature, Nui Chua N.P., 8.VIII.2014.
Belgian Journal of Entomology 58: 1–62 (2018)
45
Table 6. Measurements [mm] of Orestes krijnsi sp. nov.
Length of HT PT ♂♂ PT ♀♀
Body: 35.4 34.0–37 42.6–44.7
Head: 3.2 3.2–3.6 4.6–4.9
Pronotum: 2.2 2.2–2.3 3.5–3.7
Mesonotum: 6.5 7.0–7.4 7.7–8.1
Metanotum: 5.9 4.9–6.1 6.0–6.3
Median segment: 1.9 1.6–2.0 2.0
Profemora: 6.9 6.4–6.7 8.1–8.5
Mesofemora: 5.3 5.4 6.4–7.7
Metafemora: 6.5 6.3–7.0 7.6–8.5
Protibiae: 6.0 5.3–6.1 7.4–7.7
Mesotibiae: 4.1 3.9–4.6 5.7–6.0
Metatibiae: 5.4 4.9–5.6 7.8–8.1
Orestes mouhotii (Bates, 1865)
Figs 5, 23 D–F, 25–27
Acanthoderus mouhotii BATES, 1865: 342 [described].
Dares fulmeki WERNER, 1934: 81 [described]. Synonymised by BROCK (1998).
Datames verruculatus REDTENBACHER, 1906: 222 [described]. Synonymised by ZOMPRO
(2004).
Datames mouhotii – STÅL, 1875: 93 [transferred to Datames]. — SHIRAKI, 1935: 25 [recorded
in Taiwan and 2Japan]. — REDTENBACHER, 1906: 53 [redescription]. — BROCK, 1999: 138;
fig 93a–c. [recorded in Malaysia, redescription, egg].
Orestes mouhotii – ZOMPRO & FRITZSCHE, 1999: 10 [transferred to Orestes]. — BROCK, 2003:
70 [captive rearing]. — ZOMPRO, 2004: 222; fig. 132 a–b [redescription; description of male;
senior synonym of Dares subcylindricus Redtenbacher, 1906 (error)]. — HO, 2013: 209; fig.
19 [recorded in China]. — SEOW-CHOEN, 2017: 131 [recorded in Singapore].
MATERIAL EXAMINED.
TYPE MATERIAL. THAILAND: HT nymph of Acanthoderus mouhotii Bates, 1865
(examined from photographs): Chantaboun, Mouhot (OUMNH).
INDONESIA, Sumatra: HT of Dares fulmeki Werner, 1934 (examined from photographs):
Medan S.O.K.,I. Fulmek (NHMW).
THAILAND: HT nymph of Orestes verruculatus Redtenbacher, 1906: Bangkok, Harmand,
1885 (MNHN).
ADDITIONAL MATERIAL. (12♂♂, 4♀♀, 1 nymph, 1 nymph, eggs): CAMBODIA: 8♂♂:
Kampong Speu, Kirirom N.P., 11°18’37”N 104°03’04”E, night collecting/light trap, 9-
12.V.2015, leg. J. Constant & V. Sougnez, I.G.: 33.022 (6: RBINS; 2: RUPP); 4♀♀: same
data, ex breeding Rob Krijns, 2017 (2♀♀: RBINS; 2♀♀: RUPP); 1: Koh Kong prov., Tatai,
11°35’13”N 103°05’50”E, leg. J. Constant & J. Bresseel, I.G.: 33.345 (RBINS); 1: Koh
Kong prov., Tatai, 11°35’13”N 103°05’50”E, 7.VIII.2016, G. Chartier. I.G.: 33.345 (RBINS).
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
46
NOTES.
(1) The specimens attributed to O. mouhotii by SHIRAKI (1935) from Japan and Taiwan have
been described as two distinct species, respectively Pylaemenes japonicus Ho, 2016 and
P. shirakii Ho & Brock, 2013 (HO, 2016; HO & BROCK, 2013).
(2) The occurrence of Orestes mouhotii (Bates, 1865) from Malaysia and Singapore is
doubtful as already pointed out by BROCK (1999) who stated that those records might
represent a different species and that material from Cambodia is necessary for comparison.
The synonymy with Dares fulmeki Werner, 1934 from Sumatra (BROCK, 1998) is highly
doubtful as well and needs verification. Therefore, records from Peninsular Malaysia,
Singapore and Sumatra are omitted from the distribution map.
(3) Chantaboun is the former name of Chanthaburi (Chanthaburi Province, 12°3631N,
102°614E), and is located in Thailand near the Cambodian border rather than in Cambodia
as stated by BATES (1865).
(4) ZOMPRO (2004) synonymised Dares subcylindricus Redtenbacher, 1906 and Orestes
verruculatus Redtenbacher, 1906 with O. mouhotii (Bates, 1865). The former here reinstated
as a valid species infra and attributed to Orestes Redtenbacher, 1906.
DIAGNOSIS. Strongly granulose and tuberculose species (Figs 25 A, 26 A).
Males can be separated from the other species of Orestes by the following combination of
characters: antennae with 21 or 22 antennal segments (23 segments in other Orestes species);
posterior margin of mesonotum with a pair of tubercles (Fig. 25 I); first antennomere slightly
longer than pedicellus.
Females can be separated from the other species of Orestes by the following combination of
characters: all cephalic armature except central coronal present as tubercles (Fig. 26 I);
tergum IV indistinctly widening towards the posterior, almost parallel-sided (Fig. 26 A);
metatibiae with dorsal, minute and conical teeth anteriorly and centrally; posteromedial crest
on tergum IX not distinctly elongated but notched (Fig. 26 E–F).
DESCRIPTION
MALE (Fig. 25).
Measurements: see table 7.
Head: supra-antennals spinose and slightly pointed outwards, followed by four spinose supra-
occipitals. Vertex with anterior coronals distinctly enlarged, blunt and laterally compressed,
creating ear-like processes. Supra-orbitals spinose. Posterior and lateral coronals present as
small, elongated tubercles forming a corona with the apex of the postocular carina. Eyes
relatively small, circular and strongly projecting hemispherically. Antennae shorter than legs
with 21–22 segments (n = 6); scapus strongly flattened dorsoventrally and laterally carinate
with a spine laterally and posterolaterally; pedicellus dorsoventrally flattened. First
antennomere slightly longer than pedicellus. Antennomeres gradually getting longer till
segment XII. Antennal segment XIII the longest, with minute hump anterolaterally; following
segments shiny and distinctly shorter. Apical antennomere elongated, longer than the two
preceding antennomeres combined.
Belgian Journal of Entomology 58: 1–62 (2018)
47
Fig. 25. Orestes mouhotii (Bates, 1865), from Cambodia, Kirirom N.P. A, habitus, dorsal view. B, habitus,
lateral view. C, habitus, ventral view. D, head and thorax, dorsal view. E, terminalia, dorsal view. F, terminalia
anterodorsal view. G, terminalia, lateral view. H, terminalia, ventral view. I, head and thorax, lateral view.
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
48
Thorax: pronotum rugose, almost parallel-sided, slightly wider posteriorly, longer than wide.
Anterior margin incurved. Prozona slightly elevated centrally with four granules. Metazona
with four to six granules; posterior margin rounded. Mesonotum more or less parallel-sided
with indistinct mediolongitudinal line, with anterior margin concave with a pair of granules;
posteromedially with a pair of tubercles. Mesopleura slightly widened above coxae; widened
portion rounded and notched medially. Metanotum more or less parallel-sided,
posteromedially with two minute granules; posterior margin concave. Metapleura widened
above coxae, and rounded and notched posterolaterally. Prosternum with sensory areas
elongate oval. Sensory area on profurcasternum small and subcircular. Ventral surface of
body rugose.
Legs: profemora unarmed, curved basally with carinae rather indistinct with anterodorsal
carinae more distinct; posterior margin with small tubercle; almost as long as metanotum,
median segment and tergum II combined. Mesofemora unarmed with carinae indistinct; only
slightly shorter than profemora, slightly longer than metanotum and median segment
combined. Metafemora about as long as profemora, shaped like mesofemora; dorsally with a
small blunt tooth subcentrally. Protibiae unarmed, slightly shorter than profemora. Meso- and
metatibiae with carinae indistinct, slightly shorter than corresponding femora and unarmed.
Tarsomeres I–III about the same length with a posteromedian rounded hump dorsally. Claws
very small.
Abdomen: median segment rugose, strongly transverse and with a pair of minute granules
posteromedially. Abdominal terga rugose; terga II–V about the same length; tergum VI
slightly shorter; tergum II almost quadrate, slightly narrowing towards the posterior; terga III–
VII posteromedially parallel-sided, unarmed or sometimes armed with two minute granules
posteromedially; posterior margin of tergum VII concave; tergum VIII wider than VII,
trapezoidal and widening towards the posterior, with posterior margin concave; IX about as
long as VIII, almost quadrate, with posterior margin with small hump posteromedially. Anal
segment dorsoventrally flattened, parallel-sided and notched posteromedially. Posterolateral
angles broadly rounded. Poculum more or less rounded and strongly granulose; posterior rim
dorsoventrally flattened, thickened and broadly rounded posteriorly. Poculum projecting over
base of vomer and reaching base of apical spine. Cerci short, strongly flattened and setose, not
reaching apex of tergum X, with apices broadly rounded to almost straight. Vomer well
developed, semicircular basally till base of posteromedial spine; posteromedial spine broad,
strongly upcurving with an angle of almost 90°.
FEMALE (Fig. 26).
Measurements: see table 7.
Head: complete armature except central coronal present as tubercles, supra-antennals slightly
larger and conical. Vertex not elongated or raised but slightly rounded. Posterior and lateral
coronals distinct as stalked tubercles. Behind eye, a distinct carina reaching posterior edge of
crest, resulting in an elongated tubercle. Eyes relatively small, circular and strongly projecting
hemispherically. Antennae slightly shorter than legs with 24 segments (n = 4); scapus strongly
flattened dorsoventrally, laterally carinate and with a central tubercle and subapical spine on
the outer lateral carina; pedicellus slightly dorsoventrally flattened, slightly narrowing
towards the posterior. First antennomere about as long as the following three antennomeres
combined; antennal segment IV to XV gradually becoming longer. Segment XV with a
minute spine anterolaterally, following segments shorter than XV except for apical
antennomere which is elongated and club-shaped.
Belgian Journal of Entomology 58: 1–62 (2018)
49
Fig. 26. Orestes mouhotii (Bates, 1865), from Cambodia, Tatai. A, habitus, dorsal view. B, habitus, ventral
view. C, head and thorax, dorsal view. D, terminalia, dorsal view. E, terminalia anterodorsal view. F, terminalia,
lateral view. G, habitus lateral view. H, terminalia, ventral view. I, head and thorax, lateral view.
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
50
Thorax: pronotum trapezoidal, widening towards the posterior; anterior margin strongly
convex; prozona with a pair of granules antero- and posteromedially; metazona with two
longitudinal rows of 3 granules; lateral margin with four evenly spaced granules. Mesonotum
rugose with anterior margin thickened, strongly incurved and with two distinct granules
anteromedially; sublateral margins with few evenly spaced granules; posterior margin wider
than anterior margin. Between lateral margin and median line, a row of evenly spaced
granules almost parallel to lateral margin. Posterior margin with a pair of transversely
elongated granules. Posterior margin slightly concave. Mesopleura slightly widened above
coxae; widened portion with lateral margins crenulated, anterior portion with some raised
granules. Metanotum more or less parallel-sided, armature more or less as in mesonotum.
Metapleura widened above coxae, anterior portion with few small tubercles, posterior portion
rounded with posterolateral notch. Prosternum with sensory areas elongate oval, not reaching
lateral margins. Sensory area on profurcasternum small and rounded, anterior margin more or
less straight. Meso- and metasternum with two longitudinal rows of distinct granules.
Legs: femora with carinae relatively indistinct except for anterodorsal carina of profemora
which is laterally compressed and raised; apex with small granule. Profemora rugose, shorter
than mesonotum and curved basally. Mesofemora about as long as metanotum and median
segment combined and with some indistinct humps dorsally. Metafemora about as long as
profemora with a dorsal conical tooth subcentrally and a granule in the posterior half. Tibiae
shorter than corresponding femora. Protibiae carinate and unarmed. Mesotibiae unarmed.
Metatibiae with some indistinct granules dorsally. Tarsomeres I-III about the same length
with a rounded hump dorsally, IV distinctly shorter. Claws very small.
Abdomen: Median segment rugose, strongly transverse and with posterior margin more or less
straight. Abdominal terga rugose; terga II–IV slightly widening gradually towards the
posterior; II–III with indistinct mediologitudinal carina, distinctly granulose between carina
and lateral margin, medially and posterolaterally with small granule; tergum IV with a
posterolateral triangular tubercle; posteromedially with a twin-crescent-shaped crest, granules
as in III; tergum V slightly descending obliquely with posterior margin slightly narrower than
anterior one, armature as IV but posteriorly with flattened triangular tubercles; triangular
tubercles projecting over posterior margin; tergum VI distinctly descending and granulose,
with posterior margin distinctly narrower than anterior one and incurved; tergum VII almost
parallel-sided, slightly tapering towards the posterior with a mediolongitudinal carina and
incurved posteriorly; tergum VIII more or less parallel-sided, slightly ascending and strongly
granulose; posterolaterally with a triangular granule and posteromedially with a pair of
minute, raised granules; IX about as wide as VIII with a posteromedial crest, crest pointed
more or less upwards and notched apically; posterolateral margins rounded and with several
granules. Anal segment narrower than tergum IX and dorsally flattened; anteriorly with a
tubercle medially, posteromedially with apair of granules; sublaterally with an oblique,
compound and lamellate carina continuing on posterolateral angles; posterolateral angles
lamellate and raised. Subgenital plate with a mediolongitudinal carina; rounded and boat-
shaped with widest part at about mid length; posterior rim dorsoventrally flattened; apex more
or less rounded, almost reaching margin of tergum X.
EGG (Fig. 23 D–F) [ex. breeding O.P.I.E. 22.I.2014, origin: Thailand].
Measurements [mm]. Length: 3.6; width: 2.5; height: 3.1.
Egg capsule subspherical, slightly compressed laterally and brown. Capsule covered with
minute pits. Operculum subcircular, slightly convex. Capsule and operculum covered with
long setae with a distal black two to four-hooked grapnel-shaped structure. Base of long setae
surrounded by very short hairs, creating small circular patches on capsule and operculum.
Belgian Journal of Entomology 58: 1–62 (2018)
51
Micropylar plate trilobate with one anterior expansion and with two posterior expansions.
Anterior expansion distinctly widening towards the capsule. Anterior margin almost straight,
reaching edge of capsule. Posterior arms ax-shaped, laterally directed and projecting on lateral
surface of capsule but not reaching ventral side. Margin of micropylar plate darker and
indistinctly raised. Micropylar cup small, black, cup-shaped and protruding.
Fig. 27. Orestes mouhotii (Bates, 1865), in nature. A–D, Cambodia, Tatai (photographs by G. Chartier). A, adul
t
male, 12.X.2016. B, adult male, 17.V.2015. C, mating pair, 5.II.2017. D, female subadult nymph. E–H,
Cambodia, Kirirom N.P., 11.V.2015. E–F, adult male. G, adult female. H, adult pair.
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
52
Table 7. Measurements [mm] of Orestes mouhotii (Bates, 1865).
Length of ♂♂ ♀♀
Body: 36.0–38.0 45.0–47.0
Head: 3.6–4.1 5.2
Pronotum: 2.6–2.7 3.8–4.0
Mesonotum: 7.1–7.8 9.2–9.7
Metanotum: 5.5–6.0 6.9–7.0
Median segment: 1.6 1.8–1.9
Profemora: 6.5–6.9 7.5–7.8
Mesofemora: 5.7–5.8 6.8
Metafemora: 6.5–7.0 8.1–8.6
Protibiae: 5.6–6.3 6.6
Mesotibiae: 4.5–4.9 4.9–5.3
Metatibiae: 6.4–6.5 6.9–7.3
Orestes subcylindricus (Redtenbacher, 1906) stat. rev., comb. nov.
: urn:lsid:zoobank.org:act:EFD5D814-3F88-4D8E-9718-24EC70AF0DFE
Figs 5, 23 G–I, 28–31
Dares (Dares) subcylindricus REDTENBACHER, 1906: 56 [described].
Dares (Dares) subcylindricus – BRAGG, 1998: 60 [type identified as female nymph]. —
BROCK, 1998: 61 [catalogued]. — BRAGG, 2001: 147 [type identified as female nymph]. —
ZOMPRO, 2004: 222 [synonymised with O. mouhotii (Bates, 1865) in error].
MATERIAL EXAMINED.
TYPE MATERIAL. VIETNAM: holotype: subadult (examined from photographs): Tonkin,
Montes Mauson, April-Mai, 2-3000, H. Fruhstorfer; 24.316 (NMW, No. 55).
ADDITIONAL MATERIAL. (25♂♂, 20♀♀, 2♀♀ nymphs, eggs): VIETNAM: 9♂♂, 10♀♀, 2♀♀
nymphs, eggs: Ninh Binh prov., Cuc Phuong Nat. Park, 20°20’53”N105°35’52”E, 31.VII-
3.VIII.2016, GTI Project, Leg. J. Constant & J. Bresseel, I.G.: 33.282 (7♂♂, 8♀♀, 2♀♀
nymphs, eggs: RBINS; 2♂♂, 2♀♀: VNMN); 6♂♂, 4♀♀: Cuc Phuong N. P.,
20°19’00”N105°36’30”E, 19-23.VII.2011, Leg. J. Constant & J. Bresseel, I.G.: 31.933 (4♂♂,
2♀♀: RBINS; 2♂♂, 2♀♀: VNMN); 3♂♂, 3♀♀: Hoa Binh prov., Ngo Luong Nat. Res.,
20°26’16”N105°20’15”E, 25-30.VII.2016, GTI Project, Leg. J. Constant & J. Bresseel, I.G.:
33.282 (2♂♂, 2♀♀: RBINS; 2♂♂, 1: VNMN); 2♂♂, 1: Tay Yen Tu Nat. Res.,
21°11’10”N 106°43’25”E, 7-11.VII.2013, night collecting, leg. J. Constant & J. Bresseel,
I.G.: 32.454 (1, 1: RBINS; 1: VNMN); 1: Vietnam: Cuc Phuong, 11-18.VII.2010,
I.G.: 31.668, Leg J. Constant & P. Limbourg (RBINS); 1: Ex breeding Rob Krijns 2017,
origin: Cuc Phuong N.P., 20°19’00”N105°36’30”E, VII.2011, J. Constant & J. Bresseel, I.G.:
32.387 (RBINS); 1: Ex breeding Rob Krijns 2012, origin: Cuc Phuong N.P., 20°19’00”N
105°36’30”E, VII.2011, J. Constant & J. Bresseel, I.G.: 32.387 (RBINS); 1: Tonkin, reg. de
Hoa Binh, 1927, A. De Cooman (MNHN).
DIAGNOSIS. Relatively large species with strongly lamellate cephalic crest (Figs 28 I, 29 I, 30
I). Males have tergum IV posteromedially raised and armed with two distinct tubercles (Figs
28 B, 29 B). Females have the leg armature well developed, abdominal terga IV–V almost
parallel-sided (Fig. 30 A) and the posteromedian crest on tergum IX distinctly notched (Fig.
30 D).
Belgian Journal of Entomology 58: 1–62 (2018)
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DESCRIPTION.
MALE (Figs 28–29).
Measurements: see table 8.
Head: supra-antennals spinose and slightly pointed outwards followed by four smaller supra-
occipitals, the anterior being the smallest. Vertex raised, supra-orbitals laterally compressed,
triangular. Anterior coronals strongly compressed laterally and lamellate, about as high as
supra-orbitals. Central coronal conical, about as long as posterior coronals. Posterior and
lateral coronals present as small, conical tubercles. Behind eyes, a distinct carina reaching
posterior edge of crest resulting in a conical tubercle. Eyes relatively small, circular and
strongly projecting hemispherically. Antennae shorter than legs, with 23 segments (n = 10);
scapus strongly flattened dorsoventrally and laterally carinated with a spine laterally and
posterolaterally; pedicellus slightly flattened dorsoventrally, narrowing towards the posterior.
First antennomere distinctly longer than pedicellus. Antennomeres gradually getting longer
till segment XII. Antennal segment XIII about the same length as XII, with minute spine
anterolaterally; following segments shiny and distinctly shorter. Apical antennomere
elongated, club-shaped.
Thorax: pronotum more or less parallel-sided, indistinctly wider posteriorly, almost quadrate
and rugose. Anterior margin incurved. Prozona with slight elevation centrally with four
tubercles; anterior tubercles slightly larger. Metazona with a pair of granules anteriorly and
posteriorly. Mesonotum rugose with anterior margin thickened, distinctly concave and with a
pair of minute granules anteromedially. Posterior portion slightly wider, posteromedially with
a pair of minute granules. Mesopleura widened above coxae; widened portion rounded and
indistincly notched medially. Metanotum more or less parallel-sided, rugose and unarmed.
Metapleura widened above coxae, rounded and slightly crenulated. Prosternum with sensory
areas elongate oval. Sensory area on profurcasternum small and circular.
Legs: femora with carinae relatively indistinct except for the anterolateral carina of the
profemora, with dorsal posterior margin with short spinose tubercle. Profemora about as long
as mesonotum, curved basally and unarmed. Mesofemora about as long as metathorax,
dorsally with a blunt tubercle centrally and subapically; outer ventral carina with one,
sometimes two granules subapically. Metafemora about as long as profemora; ventrally armed
as mesofemora; dorsally with a definite blunt tooth centrally and subapically. Tibiae with
carinae indistinct, slightly than shorter corresponding femora; dorsally with a blunt tooth
centrally, slightly larger on mesotibiae and even more so on metatibiae. Tarsomeres I–III
about the same length with a posteromedian rounded hump dorsally. Claws very small.
Abdomen: median segment unarmed and rugose, with posterior margin slightly concave.
Abdominal terga rugose, only slightly varying in length; tergum II transverse, slightly
narrowing towards the posterior; tergum IV and sometimes V posteromedially raised and
armed with two tubercles; terga VI–VIII with two posteromedian granules; tergum VIII
slightly wider than VII, widening towards the posterior and with posterior margin concave; IX
distinctly transverse, with posterior margin with minute posteromedian ridge. Anal segment
narrower than IX, dorsoventrally flattened, almost parallel-sided and indistinctly notched
posteromedially. Posterolateral angles rounded. Poculum triangular from lateral view and
strongly granulose; posterior rim dorsoventrally flattened and minutely notched posteriorly,
posterolateral angles broadly rounded. Poculum projecting over base of vomer but not
reaching base of apical spine. Cerci short, strongly flattened and setose, not reaching apex of
tergum X, with apices broadly rounded to almost straight. Vomer well developed,
semicircular basally till base of posteromedial spine; posteromedial spine broad and blackish
in posterior portion; apex curved, not ending medially. Ventral surface of body rugose.
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
54
Fig. 28. Orestes subcylindricus (Redtenbacher, 1906) comb. nov., from Vietnam, Cuc Phuong N.P. A, habitus,
dorsal view. B, habitus, lateral view. C, habitus, ventral view. D, head and thorax, dorsal view. E, terminalia,
dorsal view. F, terminalia anterodorsal view. G, terminalia, ventral view. H, terminalia, lateral view. I, head and
thorax, lateral view.
Belgian Journal of Entomology 58: 1–62 (2018)
55
Fig. 29. Orestes subcylindricus (Redtenbacher, 1906) comb. nov., from Vietnam, Tay Yen Tu N.R. A, habitus,
dorsal view. B, habitus, lateral view. C, habitus, ventral view. D, head and thorax, dorsal view. E, terminalia,
dorsal view. F, terminalia anterodorsal view. G, terminalia, ventral view. H, terminalia, lateral view. I, head and
thorax, lateral view.
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
56
Fig. 30. Orestes subcylindricus (Redtenbacher, 1906) comb. nov., from Vietnam, Cuc Phuong N.P. A, habitus,
dorsal view. B, habitus, ventral view. C, head and thorax, dorsal view. D, terminalia, dorsal view. E, terminalia
anterodorsal view. F, terminalia, lateral view. G, habitus, lateral view. H, terminalia, ventral view. I, head and
thorax, lateral view.
Belgian Journal of Entomology 58: 1–62 (2018)
57
FEMALE (Fig. 30).
Measurements: see table 8.
Head: supra-antennals short and conical. Anterior supra-occipitals present as small conical
humps, posterior supra-occipitals smaller. Vertex distinctly raised, supra-orbitals strong,
laterally compressed and lamellate. Anterior coronals laterally compressed and lamellate,
directing towards supra-orbitals. Central coronal only present as small granule. Posterior and
lateral coronals present as conical tubercles. Between lateral coronal and supra-orbital, a row
of tubercles; between lateral and anterior coronal several tubercles on side of crest. Postocular
carina distinct, apex as triangular tubercle. Eyes relatively small, circular and strongly
projecting hemispherically. Antennae shorter than legs, with 25 segments (n = 10); scapus
strongly flattened dorsally and laterally carinated, with a definite spine posterolaterally
mediolaterally; pedicellus slightly flattened dorsoventrally, laterally carinate. First
antennomere longer than the following three antennomeres combined. Apical antennomere
elongated and club-shaped.
Thorax: pronotum trapezoidal, slightly widening towards the posterior; lateral margins rugose
and slightly concave, anterior margin incurved. Prozona with distinct elevation centrally with
four granules; metazona with three pairs of granules, posterior margin straight. Mesonotum
rugose, indistinctly widening towards the posterior; anterior margin thickened with some
granules; sublaterally with two rows of distinct humps. Mesopleura slightly widened above
coxae; widened portion notched medially and armed with few granules. Metanotum as
mesonotum. Metapleura as mesopleura but posterolaterally notched. Prosternum with sensory
areas elongate oval. Sensory area on profurcasternum small and circular. Meso- and
metasternum with several enlarged black granules.
Legs: profemora about as long as mesonotum and curved basally, with some minute humps
on dorsal carina; exteroventral carina with some evenly spaced minute granules. Mesofemora
shorter than mesonotum; externodorsal carina with three flattened and apically rounded teeth,
distal one stronger; inner dorsal carina with three to four minute humps. Medioventral carina
indistinct. Outer ventral carinae with two pairs of granules or small teeth posteriorly;
anteriorly with few indistinct granules. Metafemora about as long as profemora; armed as
mesofemora. Tibiae slightly shorter than corresponding femora. Protibiae with three to four
laterally compressed humps on dorsal carinae, central armature more distinct. Meso- and
metatibiae with a distinct, laterally compressed and rounded lobe dorsally, ventrally unarmed.
Tarsomeres I–III about the same length with a posteromedian rounded hump dorsally; IV
distinctly shorter. Claws very small.
Abdomen: median segment rugose, unarmed, strongly transverse and with posterior margin
almost straight. Abdominal terga rugose; terga II–VIII with X-shaped carina medially; terga
II–IV gradually widening towards the posterior; terga III–V with a distinct triangular process
posterolaterally; terga IV and V almost parallel-sided and slightly notched posteromedially;
posterior margin raised and tuberculose; posterior margin of tergum V projecting over
anterior margin tergum of VI; tergum VI distinctly narrowing towards the posterior and
strongly descending, with posterior margin slightly concave; tergum VII slightly narrowing
towards the posterior, slightly longer than previous one; tergum VIII parallel-sided and
slightly ascending; posterolateral angles with distinct tubercle and posteromedially with at
least one pair of granules; IX slightly triangular, tapering towards the posterior, only slightly
projecting over anal segment; posteromedially tectiform and notched; sublaterally with an
oblique row of granules. Anal segment narrower and shorter than tergum IX, obliquely
descending and dorsally flattened with outer margin lamellate; anteriorly with a tubercle
medially; oblique carinae lamellate; posterior margin relatively straight, slightly crenulated to
indistinctly notched. Abdominal sterna and subgenital plate with a mediolongitudinal carina.
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
58
Fig. 31. Orestes subcylindricus (Redtenbacher, 1906) comb. nov., in nature. A–C, Vietnam, Cuc Phuong N.P.,
20-22.VII.2011. A, adult male. B, adult female. C, adult female. D–E, Vietnam, Ngoc Son-Ngo Luong N.R., 25-
30.VII.2016. D, subadult female. E, mating pair.
Sterna with a pair of granules anteriorly and posteromedially; praeopercular organ knob-like
and slightly flattened laterally. Subgenital plate rugose, boat-shaped with widest part at about
mid length; posterior rim dorsoventrally flattened and rounded posteriorly; laterally with two
rows of granules. Apex broadly rounded, almost reaching margin of tergum X.
EGG (Fig. 23 G–I).
Measurements [mm]. Length: 4.0; width: 3.1; height: 3.5.
Egg capsule subspherical and brown. Capsule covered with minute pits. Operculum
subcircular, slightly convex. Capsule and operculum covered with long setae (about 0.3 mm)
with a distal black two to four-hooked grapnel-shaped structure. Base of long setae
surrounded by very short hairs, creating small circular patches on capsule and operculum.
Micropylar plate trilobate with one anterior expansion and with two posterior expansions.
Anterior expansion slightly widening towards the capsule. Anterior margin almost reaching
edge of capsule. Posterior arms directed laterally and almost completely circling polar area,
Belgian Journal of Entomology 58: 1–62 (2018)
59
but not touching each other on ventral side. Posterior arms dilated laterally on capsule,
projecting over half the capsule length. Margin of micropylar plate indistinctly raised and
demarcated by more dense setae. Micropylar cup small, black, cup-shaped and protruding.
NOTE. Males from Cuc Phuong N.P. and Ngoc Son-Ngo Luong N.R have only tergum IV
posteromedially raised and armed with two distinct tubercles (Fig. 28 B) while males from
Tay Yen Tu N.R. have also tergum V with two distinct tubercles (Fig. 29 B).
Table 8. Measurements [mm] of Orestes subcylindricus (Redtenbacher, 1906)
Length of HT nymph* ♂♂ ♀♀
Body: 42.0 39.5–43.7 47.6–54.4
Head: 4.5–5.0 6.3–7.0
Pronotum: 4.0 3.1–3.4 3.9–4.6
Mesonotum: 8.2 8.2–9.3 9.7–11.6
Metanotum: 6.0 5.9–7.1 7.0–7.6
Median segment: 1.8–2.3** 2.1–2.3
Profemora: 7.4 6.8–7.8 9.7–9.8
Mesofemora: 6.5–6.8 7.4–8.2
Metafemora: 8.3 7.6–8.2 9.1–10.2
Protibiae: 6.9–7.0 8.5–9.2
Mesotibiae: 5.3–5.6 7.0–8.0
Metatibiae: 6.7–7.0 8.4–9.6
* After RETDENBACHER (1906). ** Anterior part of median segment fused with metanotum.
Orestes guangxiensis (Bi & Li, 1994) comb. nov.
urn:lsid:zoobank.org:act:3B71D9C6-E9BE-4986-9233-EA281AA1459A
Datames guangxiensis BI & LI, 1994: 254; figs 1–4 [described and illustrated].
Pylaemenes hongkongensis BROCK & SEOW-CHOEN, 2000: 143; fig. 11 [described,
illustrated]. Synonymised by ZOMPRO (2004).
Dares guangxiensis – ZOMPRO, 2004: 219 [transferred to Dares].
Pylaemenes guangxiensis – OTTE & BROCK, 2005: 296 [transferred to Pylaemenes]. — CHEN
& HE, 2008: 358; fig. 321 A–C; pl. 4 fig. 3 [redescribed (in Chinese), keyed, illustrated].
NOTE. The species is here transferred to Orestes based on the examination of the illustrations
in CHEN & HE (2008) and the photographs of the type specimen of Pylaemenes
hongkongensis Brock & Seow-Choen, 2000 (BROCK et al., 2017). Females have the typical
dorsally flattened and apically straight anal segment. They have the widest part of the body at
the posterior margin of tergum IV. These characters clearly place guanxiensis in Orestes.
Orestes japonicus (Ho, 2016) comb. nov.
urn:lsid:zoobank.org:act:38007A66-B65E-479A-B871-B6C82717C28B
Pylaemenes japonicus HO, 2016: 11; figs 7–10, 15–16 [described and illustrated].
MATERIAL EXAMINED. (4♂♂, 2♀♀, eggs): JAPAN: 2♂♂: Japan, Okinawa, leg. Kazuhisa
Kuribayashi, ex breeding Holger Dräger 2016 [RBINS]; 2♂♂, 2♀♀, eggs: Japan, Okinawa,
ex breeding Illy Klimmert [RBINS].
NOTE. The species is here transferred to Orestes based on the examination of the illustrations
presented by HO (2016) and of cultured specimens from Okinawa Island. Males and females
BRESSEEL J. & CONSTANT J. Orestes from Vietnam and Cambodia (Phasmida, Heteropterygidae)
60
have the typical dorsally flattened and apically straight anal segment. Females have the widest
part of the body at the posterior margin of tergum IV. Those characters place the species in
Orestes and it is very similar to O. dittmari sp. nov. and O. shirakii (Ho & Brock, 2013)
comb. nov.
Orestes shirakii (Ho & Brock, 2013) comb. nov.
urn:lsid:zoobank.org:act:CDF05FCC-6DAD-4EF8-9B41-44E39CC75C9A
Pylaemenes shirakii Ho & Brock, 2013 in HO (2013): 127; figs 29–31 [described and
illustrated].
MATERIAL EXAMINED. 2♀♀: TAIWAN, no precise locality, ex breeding 2014 [RBINS].
NOTE. The species is here transferred to Orestes based on the examination of the illustrations
of the type specimen presented by HO (2013) and of cultured specimens from Taiwan.
Females have the typical dorsally flattened and apically straight anal segment and have the
body with the widest part at the posterior margin of tergum IV. These characters obviously
place shirakii in Orestes.
Discussion
Besides Orestes Redtenbacher, 1906, three other genera of Datamini are recorded from
continental Asia (BROCK et al., 2017): Dares Stål, 1875, Pylaemenes Stål, 1875 and
Hainanphasma Ho, 2013. The genus Dares is represented only by a single species on the
continent, D. ziegleri Zompro & Fritsche, 1999 from Nakhon Ratchasima, Thailand. Only
females are known and all other species of this genus are recorded from Borneo and Palawan,
letting the appearance of a single species in Thailand appear fairly odd. This species warrants
further research and the study of more material including the still unknown male would
resolve the somewhat questionable generic assignment of the species.
The recently described genus Hainanphasma is endemic to Hainan and contains two species.
The genus is obviously closely related to Orestes and HO (2013) separated the genus from the
latter by its more slender antennae, the fin-like lamella on the posterodorsal carina of the
meso- and metafemora, the punctulate occipital crest in females, as well as the elevated
occipital crest and distinctive hump on abdominal terga IV and V in males. Due to the new
species described herein, several of the diagnostic characters provided for Hainanphasma by
HO (2013) appear fairly weak for generic distinction and definitely deserve further assessment
and evaluation. For instance, comparatively slender and thin antennae are also seen in
O. botot sp. nov and O. diabolicus sp. nov., although somewhat smaller the fin-like lamella on
the posterodorsal carina of meso- and metafemora is also present in several species of Orestes
like O. draegeri sp. nov. and O. subcylindricus (Redtenbacher, 1906) comb. nov. and the
distinctive hump on abdominal terga IV and V in males is also found in O. subcylindricus
(Redtenbacher, 1906) comb. nov., and to a lesser degree in O. bachmaensis sp. nov. However,
the egg illustrated by HO (2013) differs considerably from Orestes eggs in being only sparsely
setose, having the operculum medially raised and, in lateral view, showing the dorsal margin
convex and the ventral one oblique.
Six species of the genus Pylaemenes Stål, 1875 were recorded from continental Asia (BROCK
et al., 2017), three of which are here transferred to Orestes: O. guangxiensis (Bi & Li, 1994)
comb. nov., O. japonicus (Ho, 2016) comb. nov. and O. shirakii (Ho & Brock, 2013) comb.
Belgian Journal of Entomology 58: 1–62 (2018)
61
nov. Hence, three continental species are retained in Pylaemenes: P. mitratus (Redtenbacher,
1906), P. pui Ho, 2013 and P. trapezius Ho, 2016.
Currently there are still gaps in the distribution of the subfamily with no records from
Myanmar and Laos and only a few records from Cambodia and Thailand. New collecting
efforts in these areas need to be conducted for any thorough study in due course.
The whole subfamily is in need of a generic and specific revision. New material and a
combined approach including morphology, ootaxonomy, molecular and zoogeographical data
would provide an interesting and more comprehensive insight on the relationships between
the different taxa.
Acknowledgments
We thank Dr Hong Thai Pham (VNMN) and his colleagues and Mr Sophany Phauk (RUPP) and his students as
well as Dr Vincent Sougnez (Lusigny-sur-Barse, France) for all their help and friendship during the collecting
trips; Mr Daniel Dittmar (Germany), Holger Dräger (Germany) and Rob Krijns (The Netherlands) for their help
in captive rearing the species and providing additional study material; Mr Ludovic Nicolas for mounting
numerous specimens we brought back from the field; the authorities of the protected areas where we have
sampled in Vietnam and in Cambodia for their support to our studies; Dr Patrick Grootaert, Dr Yves Samyn and
Dr Marie-Lucie Susini (RBINS) for their permanent support to our projects in Vietnam and Cambodia; Mr
Xavier Vermeersch (scientific collaborator, RBINS) for his help in taking the measurements of many specimens;
Mrs Mado Berthet (RBINS) for the drawing of the cephalic armature; Mr Gerard Chartier (Cambodia) and Mr
François Tetaert (France) for the permission to use their photographs; Mr Emmanuel Delfosse (MNHN) for his
help in the collection of MNHN; Mr Frank Hennemann (Germany) and Mr Oskar Conle (Germany) for
reviewing the manuscript.
This paper is a result of the projects “A step further in the Entomodiversity of Vietnam” (2010–2017) and “A
step further in the Entomodiversity of Cambodia” (2016–2017) supported through a grant issued by the capacity
building Programme of the Belgian Global Taxonomic Initiative National Focal Point that runs under the
CEBioS programme with financial support from the Belgian Directorate-General for Development Cooperation
(DGD), and of an expedition to Cambodia supported by the entomology department of RBINS in 2015.
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... A clade of Oriental stick insects, the Heteropterygidae may be considered to represent a nonadaptive radiation since its members deploy a number of uniform phenotypic and behavioural characteristics associated with living close to the forest floor. Although often generally referred to as ground-dwellers (Bragg, 1998;Hennemann et al., 2016a;Bradler & Buckley, 2018: Bresseel & Constant, 2018, Heteropterygidae can also be found on bark (e.g. Mearnsiana Rehn & Rehn;Hennemann et al., 2016a) and in the vegetation, in particular during nocturnal feeding, whereas during daytime they mostly rest among leaf litter, pieces of bark or between roots of trees (Bragg, 2001). ...
... Sumatra, Java). The monophyly of Orestes could be confirmed and the position of Orestes krijnsi as sister taxon to all remaining species is in fact coinciding with morphological analysis of the egg, which is distinct to other Orestes (Bresseel & Constant, 2018). Heteropteryginae consists of two currently recognized genera Haaniella and Heteropteryx of which the former appears to be paraphyletic with some Haaniella spp. ...
... Several specimens allegedly representing Orestes mouhotii Bates do not cluster together, most probably because of erroneous identifications in previous studies (Kômoto et al., 2011;Goldberg et al., 2015;Robertson et al., 2018). The true O. mouhotii is represented by an individual from the Kirirom National Park (Cambodia; Bresseel & Constant, 2018), which is the sister species of a yet undescribed Orestes species from the Andaman Islands. The erroneous remaining O. mouhotii individuals are in fact members of O. draegeri Bresseel & Constant, as are the Orestes spp. ...
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The new species Pylaemenes scrupeus sp. nov. from Ratchaburi province, Thailand is described and illustrated based on both sexes and the egg. The new species is diagnosed and differentiated from closely related species. The constitution of Pylaemenes Stål, 1875 and the problems arising with generic attribution of species, considering recent phylogenetic studies, are discussed. The genus is for the first time recorded from Thailand and pictures of living specimens and the habitat as well as a distribution map of continental Pylaemenes species are provided. Dares ziegleri Zompro & Fritzsche, 1999 is transferred to Orestes Redtenbacher, 1906 leaving the genus Dares Stål, 1875 restricted to Borneo and Palawan.
... Specimens and structures were photographed under a Leica M205C stereomicroscope with LAS Core software, and measured with Mitutoyo calliper (0.05 mm accuracy) in dorsal view. Terminology of external structures follows Friedemann et al. (2011) for head, Gottardo & Vallotto (2014), Vallotto et al. (2016a, b) and Bresseel & Constant (2018a) for body and terminalia. Terminology of the egg capsule follows Clark Sellick (1997). ...
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Stick insects have received little attention in Brazil, with many taxa lacking taxonomic, biological and morphological information on the literature. This represents a gap of knowledge for inclusive groups in Phasmatodea and for the Neotropical diversity as a whole, including members of poorly known Pygirhynchini (Heteronemiidae). Canuleius similis Redtenbacher belongs to that lineage and is redescribed based on 123 individuals recently collected or raised in captivity. Bacteria ornata Brunner von Wattenwyl is found to be a junior synonym of C. similis. Lectotypes and paralectotypes are designated for Canuleius inermis Redtenbacher, of which part of the syntype series is assigned to C. similis. The male, female, nymphs and eggs were analyzed, illustrated and described. External macromorphology of mouthparts, cerci, tarsi, antennae and internal morphology of genitalia of both sexes, in special the male genitalia were accounted for. The findings on male and female genitalia are discussed considering available information for Phasmatodea, including so far infrequently referenced works on Chilean species, hoping to shed more light in the understanding of genital structures in these insects. The male genitalia have common characters between Heteronemiidae and Pseudophasmatidae, indicating that the former may be a member of Occidophasmata. Female and male internal genitalia vary interespecifically and appears to be conservative in the same population. Traditional taxonomic characters, mostly related to camouflage, are shown to vary whereas detailed morphology is emphasized to be more conservative and encouraged to be included in future analyses. Additional information on habitat, behavior and development are given.
... The nomenclature of the head armature follows Bresseel & Constant (2018). Female terminalia are as in Valloto et al. (2016). ...
Article
The male genitalia have been neglected in Phasmatodea studies over the years. There are few works describing and illustrating this organ and no studies demonstrating its usage. Concerned by this matter, this paper aimed to investigate whether the male genitalia of Phasmatodea is useful for systematics. First, by describing a new species of Creoxylus Audinet-Serville (Creoxylus duckei sp. nov.) and including the description of its male genitalia, along with the differentiation from the male genitalia of the know species Creoxylus spinosus (Fabricius). By comparing these two genitalia, the first case of chirality in Phasmatodea was reported. Secondly, by describing the genitalia of two Prexaspes Stål species (Prexaspes viridipes Stål and another identified to genus level) and making the differentiation between them. Finally, by comparing all four described genitalia. Results showed that the male genitalia are useful to differentiation and identification of species/genera. It is discussed what knowledge can be drawn from these genitalia descriptions in terms of phylogeny of Xerosomatinae and Phasmatodea as a whole, as well as homologies with previous literature.
... The stick insect fauna of Cambodia is extremely poorly known with only two species formally recorded from the country: Orestes mouhotii (Bates, 1865) (BRESSEEL & CONSTANT, 2018) and Ramulus detrectans (Brunner von Wattenwyl, 1907). The records of Calvisia (Calvisia) torquata (Bates, 1865) and Necroscia annulipes (Gray, 1835) were both from Chantaboun, the latter based on the type locality of its junior synonym N. pictipes (Bates, 1865); they were erroneously attributed to Cambodia but the location actually lies in Thailand (BRESSEEL & CONSTANT, 2017b) which was also clearly stated by MOUHOT (1864). ...
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A new species of Medauroidea Zompro, 2000, M. romantica sp. nov., is described from Phnom Tnout mountain, Preah Vihear Province in Cambodia from males, females and eggs, the latter showing a heart-shaped micropylar plate. Male, female, eggs, habitat and colour variation in females are illustrated, and a distribution map is provided. It is the first record of the genus in Cambodia. Ramulus brongniarti (Brunner von Wattenwyl, 1907) is transferred to Medauroidea and the new combination M. brongniarti (Brunner von Wattenwyl, 1907) comb. nov. is subsequently proposed. This represents the first record of the genus in Laos. Ramulus imperialis (Brunner von Wattenwyl, 1907) is proposed as a junior synonym of M. brongniarti, being the female of the latter. A lectotype is designated for R. imperialis. Both latter taxa are illustrated from their type specimens. This paper brings the total number of stick insects taxa formally recorded from Cambodia and Laos to three genera and three species and to two genera and two species, respectively.
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The genus Microrestes gen. nov. is erected to accommodate the new species Microrestes robustus sp. nov. from North Vietnam. Pylaemenes trapezius Ho, 2016 from southern China is transferred to Microrestes and the new combination M. trapezius (Ho, 2016) comb. nov. is proposed. A third species of Microrestes is recorded from northern Thailand based on photographs. The genus is compared with the other genera of Dataminae. Microrestes robustus sp. nov., the type species, is described and figured from both sexes and the egg. The genus is recorded from Vietnam, China and Thailand. A key and distribution map are provided for its species. A standardized nomenclature of the cephalic armature for Microrestes gen. nov. is proposed.
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Rearing and Studying Stick and Leaf Insects OUT OF PRINT
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A detailed listing of this important collection, with notes on Brunner von Wattenwyl and Redtenbacher. 72 pages, paperback A5. OUT OF PRINT but can be downloaded here: https://www.zobodat.at/pdf/kat-nhmw_13_0003-0072.pdf
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Introductory sections on nomenclature and classification, life history, collecting, photography, morphology. Descriptions and identification keys to 103 species, plus species catalogue.
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The areolate Oriental family Heteropterygidae Kirby, 1893 is critically reviewed and the results of the present study contradict the arrangement suggested by Zompro (2004), but in most aspects agree with a molecular study presented by Whiting et al (2003) and a phylogenetic study presented by Bradler (2009). The family is critically discussed and new hypotheses are presented for the phylogeny and intra-familiar relationships, placing the subfamily Dataminae Rehn & Rehn, 1939 as the basalmost clade of Heteropterygidae. The subfamilies Obriminae Brunner v. Wattenwyl, 1893 and Heteropteryginae Kirby, 1893 together represent the sister-group of Dataminae. Arguments and a tree are presented to support this hypothesis. New diagnoses and lists of genera are provided for all three subfamilies contained in Heteropterygidae, along with keys to distinguish between them. The subfamily Obriminae is critically reviewed and the distinction between the three tribes Obrimini Brunner v. Wattenwyl, 1893, Eubulidini Zompro, 2004 and Miroceramiini Zompro, 2004 introduced by Zompro (2004) is shown to be poorly supported. While Obrimini sensu Zompro, 2004 is generally accepted (but now also contains genera that were placed in Eubulidini or Miroceramiini by Zompro (2004)), the tribes Eubulidini and Miroceramiini are not supported. A new arrangement is introduced, which is based on morphological characters neglected or overlooked by Zompro (2004) but were partly discussed by Bradler (2009). The genus Mearnsiana Rehn & Rehn, 1939 is removed from Miroceramiini and transferred to Obrimini. The genera Eubulides Stål, 1877, Heterocopus Redtenbacher, 1906, Theramenes Stål, 1875 and Stenobrimus Redtenbacher, 1906 are removed from Eubulidini and also transferred to Obrimini. Consequently, Eubulidini is synonymised with Obrimini (n. syn.). Miroceramiini is a monotypical tribe and only includes the Wallacean genus Miroceramia Günther, 1934. The new tribe Tisamenini n. trib. is established for the three basal genera Tisamenus Stål, 1875, Ilocano Rehn & Rehn, 1939 and Hoploclonia Stål, 1875 all of which were placed in Eubulidini by Zompro (2004). The latter genus differs from the other two genera by the morphology of the female genitalia, which is unique amongst the entire family. Three generic groups are recognized within Obrimini, the Obrimus-group, Stenobrimus-group and Theramenes-group. Keys are presented to distinguish between the three tribes now contained in the Obriminae, i.e. Obrimini, Tisamenini n. trib. and Miroceramiini. The genus Hennobrimus Conle, 2006 is synonymised with Mearnsiana Rehn & Rehn, 1939, based on the fact that the type-species of both genera are conspecific (n. syn.). Hennobrimus hennemanni Conle, 2006, the type-species of Hennobrimus, and Trachyaretaon manobo Lit & Eusebio, 2005 are synonymised with Mearnsiana bullosa Rehn & Rehn, 1939, the type-species of Mearnsiana (n. syn.). Theramenes dromedarius Stål, 1877 from the Philippines is removed from synonymy with the Wallacean Theramenes olivaceus (Westwood, 1859) and re-established as a valid species (rev. stat.). The subfamily Heteropteryginae Kirby, 1896 is revised at the species-level and a new diagnosis is presented. Keys to the two genera and all 16 known species are provided along with new descriptions, differential diagnoses, lists of examined material, detailed information on the known distributions, measurements and illustrations of the insects and eggs. The intra-subfamiliar and intra-generic relationships are discussed and a cladogram is presented. Heteropteryginae contains two genera: Heteropteryx Gray, 1835 (Type-species: Phasma dilatatum Parkinson, 1798) and Haaniella Kirby, 1896 (Type-species: Phasma (Heteropteryx) muelleri de Haan, 1842). The distribution of this subfamily is restricted to Sundaland with the exception of a single species that is found in Vietnam. All other species are distributed in Borneo, Sumatra, the Mentawai Islands, Singapore, Peninsular Malaysia and Thailand. Heteropteryginae contains the largest and most striking members of the entire family Heteropteryginae, some of which are amongst the heaviest insects known. The subfamily is characterized by apomorphies such as the presence of wings, having a tympanal area (= stridulatory organ) in the basal portion of the alae, straight profemora, strongly shortened tarsi, lack of rough sensory- Areas on the prosternum and typically X-shaped micropylar plate of the eggs. The sister-group of Heteropteryginae is represented by the Obriminae, with which it shares a beak-like secondary ovipositor in the females and presence of a medio- Apical spine on the area apicalis. Both features are synapomorphies of Heteropteryginae + Obriminae. The genus Haaniella Kirby, 1904 contains 16 known species, five of which are newly described herein. The genus Miniopteryx Zompro, 2004 (Type-species: Haaniella parva Günther, 1944) is synonymised with Haaniella on the basis that the distinguishing feature mentioned in the original description is a character that is frequently found throughout the genus (n. syn.). The type-species H. parva Günther, 1944 is automatically retransferred to Haaniella (rev. stat.). Haaniella aculeata n. sp. from western Sumatra is described from the male. Haaniella macroptera n. sp. from Singapore and the Johor state in southern Peninsular Malaysia is described from both sexes and the eggs. Haaniella gintingi n. sp.from Central Sumatra is described from both sexes and the eggs and Haaniella kerincia n. sp. from Western Sumatra is described from the insects only, the eggs being still unknown. One new species, Haaniella gorochovi n. sp., is the only representative of the genus and subfamily Heteropteryginae known from Vietnam and both sexes as well as the eggs are described. Haaniella erringtoniae (Redtenbacher, 1906) is endemic in Peninsular Malaysia, here removed from synonymy with H. muelleri (de Haan, 1842) and re-established as a valid species (rev. stat.). The Sumatran Haaniella glaber (Redtenbacher, 1906) is removed from synonymy with H. muelleri (Haan, 1842) and re-established as a valid species (rev. stat.). Leocrates glaber Redtenbacher, 1906 and Haaniella muelleri simplex Günther, 1944 are removed from synonymy with H. muelleri (Haan, 1842) (rev. stat.) and synonymised with H. glaber. Haaniella mecheli (Redtenbacher, 1906) and H. rosenbergii (Kaup, 1871) are removed from synonymy with H. muelleri (Haan, 1842) and re-established as valid species (rev. stat.). Haaniella erringtoniae novaeguineae Günther, 1934 and Haaniella muelleri var. b. (Haan, 1842) are synonymized with H. rosenbergii (Kaup, 1871) (n. syn.). The type-species Haaniella muelleri(Haan, 1842) is shown to be a fairly rare species that is restricted to Sumatra. All subsequent records of H. muelleri from outside Sumatra and references to captive breeding of stock originating from Peninsular Malaysia in Europe relate to H. erringtoniae (Redtenbacher, 1906). The previously unknown males and eggs of H. rosenbergii (Kaup, 1871) as well as the previously unknown females and eggs of H. parva Günther, 1944 are described and illustrated for the first time. Based on morphological characters of the insects and eggs three distinct species-groups are recognized within Haaniella. The muelleri species-group contains nine species that are distributed throughout Sumatra, the Mentawei Islands, Singapore and Peninsular Malaysia. These are characterized by the smooth ventral surface of the meso- And metafemora and lemon-shaped eggs which entirely lack the setae seen in the two other species-groups. The grayii species-group comprises four species, two of which are endemic in Borneo, one endemic in Sumatra and the fourth species being the only known representative of the subfamily in Vietnam. These species are characteristic for the prominent pair of spines on the abdominal tergites II-IV of males and long apically multidentate epiproct of females. The echinata species-group contains three exceptionally Bornean species, which are characterized by the long and apically pointed subgenital plate of females, which clearly projects beyond the epiproct, as well as the sub-basal lateral tooth of the anal segment of males. The muelleri species-group is sister to the remainder two species-groups. Heteropteryx Gray, 1853 is a monotypical genus and only contains the type-species H. dilatata (Parkinson, 1798), which is found throughout Peninsular Malaysia, Thailand, Sumatra and Northeastern Borneo. This genus differs from Haaniella by the strongly conically elevated head, which posteriorly projects over the anterior margin of the pronotum, females being bright green or yellow in colour with plain and translucent pink alae and having distinct spines on the abdominal tergites, and males having a strongly shortened mesothorax and dull pink alae. Lectotypes are designated for Haaniella parva Günther, 1944, Heteropteryx echinata Redtenbacher, 1906, Heteropteryx saussurei Redtenbacher, 1906 and Heteropteryx scabra Redtenbacher, 1906 to guarantee stability of these names. Information on the habitats, host-plants, biology, life cycle, parasitism and captive breeding of the species of Heteropteryginae is presented and a list summarising all taxonomic changes presented herein.
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The three tribes of Heteropteryginae Kirby, 1896 occurring in Borneo are nocturnal, and ground dwelling species, easily found in both primary and secondary rainforest. The subfamily is reviewed, with keys and redescriptions of all Bornean species; one new species of a predominantly Bornean genus is described from the Philippines. Lectotypes have been selected for a number of species. The eggs of 17 species are described and illustrated. Distribution maps are given for all species. Many of the species have been collected by the author and reared in the United Kingdom; some observations on their natural history and behaviour are included. In the tribe Heteropterygini the synonymy has been re-examined with a revision to the status of several taxa. The five syntypes of Haaniella grayii (Westwood) were found to belong to two different species. In the tribe Datamini new terms are introduced for the spines and tubercules. A new genus, Spinodares, is described with S. jenningsi spec. nov. as the type species. All recorded Bornean specimens of Dares Stål, 1875 have been re-examined and the synonymy revised. Seven new species of Dares are described, six from Borneo: D. kinabaluensis, D. mjobergi, D. multispinosus, D. murudensis, D. navangensis, D. planissimus, and one from Palawan: D. philippinensis; this is the only record of the tribe Datamini from the Philippines. Acanthoderus otys Westwood, 1859 has previously been placed in the genus Dares Stål, 1875, it is found to belong in Datames Stål, 1875 with Pylaemenes infans Redtenbacher, 1906 as a new junior synonym; the female is described for the first time from a specimen in the Nationaal Natuurhistorische Museum, Leiden (RMNH). Two new species of Datames are described, one with three subspecies: D. muluensis, D. borneensis borneensis, D. b. sepilokensis and D. b. waterstradti. Of the twelve new taxa described eight are represented in the RMNH collection.
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http://phasmida.speciesfile.org The Phasmida Species File (PSF) is a taxonomic database of the world's Phasmida (stick and leaf insects, known as walking sticks and walking leaves in the U.S.). There is full synonymic and taxonomic information for 3,350 valid species and 5,300 taxonomic names, 37,500 citations to 3,178 references, over 7,600 specimen records and 16,800 images of 75% of valid species, with more being added to on a regular basis. Another future aim of this database is to provide high quality images of living phasmids in the wild. The PSF is annually in fed into the catalogue of life (Species 2000: Naturalis, Leiden, the Netherlands).
Article
Ho, G.W.C. 2016. The genus Pylaemenes Stål, 1875 (Phasmatodea: Heteropterygidae: Dataminae) of East Asia with descriptions of two new species. Tettigonia: Memoirs of the Orthopterological Society of Japan, 11: 1–14.
Article
This paper deals with four genera and eight species of the subfamily Dataminae Rehn & Rehn, 1939 from China. One new genus and four new species, Hainanphasma cristata Ho gen. nov. spec. nov., H. diaoluoshanensis Ho spec. nov., Pylaemenespui Ho spec. nov. and Pylaemenes shirakii Ho & Brock spec. nov., are described and illustrated. A new combination is proposed: Planispectrum hainanensis (Chen & He, 2008) comb. nov. is transferred from Pylaemenes Stål, 1875 and its male and egg are described for the first time. The occurrence of Orestes mouhotii (Bates, 1865) in China is reconfirmed assessed by an adult specimen collected from Yunnan Province. Pylaemenes guangxiensis (Bi & Li, 1994) is reported for the first time from Vietnam outside the range of China. Keys to the genera and species of the Chinese Dataminae are given.