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The new stick insect genus Pterulina gen. nov., a second winged Clitumninae genus from Vietnam with a new combination and a new species (Phasmida, Phasmatidae, Clitumninae, Clitumnini)

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Pterulina gen. nov. is erected to include Sipyloidea distinctissima Redtenbacher, 1908 from the central Annamite mountains and Pterulina simoensi sp. nov. from the southern Annamites. Pterulina distinctissima (Redtenbacher, 1908) comb. nov., the type species of the newly erected genus, is redescribed and figured from both sexes and egg. The species was misplaced in Sipyloidea Brunner von Wattenwyl, 1893, a genus of Necrosciinae (Lonchodidae) and is here transferred to a newly described genus of Clitumninae (Phasmatidae). The new species Pterulina simoensi sp. nov. is described based on male, female and egg. The taxonomic placement and the biogeography of this winged genus in Clitumnini is discussed. A key and distribution map are provided for its species.
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Belgian Journal of Entomology 96: 130 (2020) ISSN: 2295-0214
www.srbe-kbve.be
urn:lsid:zoobank.org:pub:86A63507-90DC-445B-96A7-0BCA112D6C4D
Belgian Journal of Entomology
The new stick insect genus Pterulina gen. nov., a second winged
Clitumninae genus from Vietnam with a new combination and
a new species (Phasmida, Phasmatidae, Clitumninae, Clitumnini)
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, June 29, 2020
BRESSEEL J. & CONSTANT J. Pterulina gen. nov. from Vietnam (Phasmida: Clitumninae: Clitumnini)
2
Citation: BRESSEEL J. & CONSTANT J., 2020. - The new stick insect genus Pterulina gen. nov., a second winged
Clitumninae genus from Vietnam with a new combination and a new species (Phasmida, Phasmatidae,
Clitumninae, Clitumnini). Belgian Journal of Entomology, 96: 130.
ISSN: 1374-5514 (Print Edition)
ISSN: 2295-0214 (Online Edition)
The Belgian Journal of Entomology is published by the Royal
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established on April 9, 1855.
Head office: Vautier street 29, B-1000 Brussels.
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Front cover: Head and thorax lateral view; top: Pterulina distinctissima (Redtenbacher, 1908) comb. nov., male;
bottom: Pterulina simoensi sp. nov., male.
Belgian Journal of Entomology 96: 130 (2020)
3
The new stick insect genus Pterulina gen. nov., a second winged
Clitumninae genus from Vietnam with a new combination and
a new species (Phasmida, Phasmatidae, Clitumninae, Clitumnini)
Joachim BRESSEEL1 & Jérôme CONSTANT2
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
Pterulina gen. nov. is erected to include Sipyloidea distinctissima Redtenbacher, 1908 from the
central Annamite mountains and Pterulina simoensi sp. nov. from the southern Annamites.
Pterulina distinctissima (Redtenbacher, 1908) comb. nov., the type species of the newly erected
genus, is redescribed and figured from both sexes and egg. The species was misplaced in
Sipyloidea Brunner von Wattenwyl, 1893, a genus of Necrosciinae (Lonchodidae) and is here
transferred to a newly described genus of Clitumninae (Phasmatidae). The new species
Pterulina simoensi sp. nov. is described based on male, female and egg. The taxonomic
placement and the biogeography of this winged genus in Clitumnini is discussed. A key and
distribution map are provided for its species.
Keywords: Da Lat Plateau, Global Taxonomy Initiative, Indochina, Kon Tum Plateau,
Lobofemora, Phasmatodea.
Introduction
The identification of specimens of stick insects collected in the framework of the Global
Taxonomy Initiative project “A step further in the entomodiversity of Vietnam” revealed a new
genus including a species previously attributed to Sipyloidea Brunner von Wattenwyl, 1893.
The new genus is described below as Pterulina gen. nov. and can be differentiated from other
genera of Clitumnini by having the median segment about as long as or longer than the
metanotum, the lack of an ovipositor in females and antennae longer than the profemora as well
as the presence of wings in males.
New genera are rarely described in the species rich tribe Clitumnini which currently counts 231
species including 159 currently attributed to Ramulus Saussure, 1862. The stick-like body with
fairly few or subtle diagnostic characters makes species hard to distinguish at generic level.
The new genus Pterulina gen. nov. is only the seventh genus described in over a hundred years
(BROCK et al., 2019), the third one by the authors (BRESSEEL & CONSTANT, 2015; BRESSEEL &
CONSTANT, 2019) and after Lobofemora Bresseel & Constant, 2015, the second winged genus
known within the tribe.
The present paper aims to describe and differentiate Pterulina gen. nov., to propose the new
combination Pterulina distinctissima (Redtenbacher, 1908) comb. nov., with an update on the
distribution of the species and to describe a new species Pterulina simoensi sp. nov. from the
Dac Lac Plateau in the southern Annamites.
BRESSEEL J. & CONSTANT J. Pterulina gen. nov. from Vietnam (Phasmida: Clitumninae: Clitumnini)
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Material & methods
Due to their nocturnal behaviour, the specimens of Pterulina gen. nov. were collected at night.
A light-weight and water-proof Petzl MYO RXP head torch was used during collecting.
Females were kept alive in a mesh pop up cage (exo terra explorarium™) for producing eggs.
The specimens were euthanized by an injection with ethanol, 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.
For each picture of the new species, a number of photographs were taken with a Canon 700D
camera equipped with a Sigma 50 mm Macro lens (for adults), or with a Leica EZ4W
stereomicroscope with integrated camera (for eggs), stacked with CombineZ software and
optimized with Adobe Photoshop CS3. The distribution map was produced with SimpleMappr
(SHORTHOUSE, 2010). Observations were done with a Leica EZ4W stereomicroscope and
measurements were taken with an electronic calliper.
The nomenclature of the morphological characters follows BRAGG (2001); the egg morphology
follows that of CLARCK-SELLICK (1997; 1998). The description of the colouration is based on
live and dried specimens.
Acronyms used for the collections:
MNHN = Muséum National d’Histoire Naturelle, Paris, France.
RBINS = Royal Belgian Institute of Natural Sciences, Brussels, Belgium.
VNMN = Vietnam National Museum of Nature, Hanoi, Vietnam.
Abbreviations:
N.P.: National Park;
N.R.: Nature Reserve;
HT: holotype;
PT: paratype.
Taxonomy
Family Phasmatidae Leach, 1815
Subfamily Clitumninae Brunner von Wattenwyl, 1893
Tribe Clitumnini Brunner von Wattenwyl, 1893
Genus Pterulina gen. nov.
urn:lsid:zoobank.org:act:F36B15B0-CD0C-4084-B77D-3FD3FC4B0FEE
TYPE SPECIES: Sipyloidea distinctissima Redtenbacher; 1908 by present designation.
ETYMOLOGY. The genus name is the combination of pter-, from the Greek word pteron meaning
wing, and -ulina, the ending of Cuniculina Brunner von Wattenwyl, 1907 illustrating the close
relationship between these genera.
DIAGNOSIS AND DIFFERENTIATION. Differing from all other known genera of Clitumnini
Brunner von Wattenwyl, 1893 by the following combination of characters:
1) Presence of tegmina and alae in male.
2) Antennae in males strongly elongated, distinctly projecting over profemora.
3) Females with a distinct praeopercular organ.
4) Median segment about as long as or longer than metanotum.
Belgian Journal of Entomology 96: 130 (2020)
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5) Hemi-tergites of males strongly elongated and tapering into a point.
6) Egg capsule elongated, polar area incised; micropylar plate spear shaped.
As usual for stick insects that have wings in one or both sexes, the median segment in Pterulina
gen. nov. is fairly long in relation to the metanotum and the mesothorax is shorter than twice
the combined length of head and prothorax. Both features are only shared with Lobofemora
Bresseel & Constant, 2015 and distinguish these two genera from all currently known
representatives of Clitumnini. Females are easily distinguished from Lobofemora by the
presence of a distinct praeopercular organ on sternum VII and by lacking a beak-like ovipositor
that is shaped by the elongated subgenital plate and epiproct. Males are distinguished by the
longer antennae, distinctly projecting over profemora; the presence of longer hind wings,
reaching at least to posterior margin of tergum III and by the distinctly elongated and slender
hemi-tergites (very short and broad in Lobofemora).
The eggs somewhat resemble those of Cuniculina Brunner von Wattenwyl, 1907, in having an
elongate minutely granulose capsule with the polar area incised and the operculum inserted in
a distinct negative opercular angle. It can be differentiated by the absence of an elongated
coronal rim and by the spear-shaped micropylar plate.
DESCRIPTION
MALE AND FEMALE
Body: Stick-like with females apterous and males winged. Females vary in colouration to a
certain degree, colouration in males constant.
Head: Longer than wide, slightly narrowing towards the posterior and smooth. Vertex slightly
elongated and flattened, back of head indented medially and submedially creating four
indistinct humps. Eyes circular and projecting hemispherically. Antennae black, males with
antennae projecting over profemora; females with short antennae reaching about halfway
profemora.
Thorax: Pronotum shorter than head, more or less rectangular; anterior margin concave and
slightly thickened; posterior margin slightly rounded. Mesonotum elongated, cylindrical in
cross-section. Metanotum short, about one third of median segment in males and slightly longer
than median segment in females.
Wings: Males with tegmina projecting over posterior margin metanotum, with definite hump
and broadening towards the posterior, apex broadly rounded. Alae projecting at least over
posterior margin of tergum III.
Legs: Profemora laterally flattened and curved basally, all carinae present; females have the
anterodorsal carina serrated and the posterodorsal carina with triangular teeth and have the
mesofemora with both dorsal carina armed with three evenly spaced minute to triangular teeth.
Metafemora as mesofemora.
Abdomen: Elongated and cylindrical in cross-section. Apical portion of abdomen in males with
anal segment distinctly longer than tergum IX, apically split into two elongated and slender
hemi-tergites. Inner portion of hemi-tergite armed with black hook-like teeth. Apical portion of
abdomen in females with anal segment concave apically, lateral margins rounded, angular
posterolaterally. Short triangular epiproct visible, slightly projecting over apex of anal segment
with mediolongitudinal carina. Sternum VII with a distinct praeopercular organ. Subgenital
plate laterally compressed with a mediolongitudinal carina in the posterior portion. Posterior
margin rounded.
BRESSEEL J. & CONSTANT J. Pterulina gen. nov. from Vietnam (Phasmida: Clitumninae: Clitumnini)
6
SPECIES INCLUDED
Pterulina distinctissima (Redtenbacher, 1908) comb. nov.
Pterulina simoensi sp. nov.
Keys to the species of Pterulina gen. nov.
MALES
1. Wings reaching posterior margin of tergum III, head with mediolongitudinal black stripe
dorsally ....................................................................................... Pterulina simoensi sp. nov.
- Wings projecting over posterior margin of tergum IV, head with a distinct round yellow
marking between the eyes .............................................. Pterulina distinctissima comb. nov.
FEMALES
1. Praeopercular organ tapering towards the posterior with posterior margin rounded, not
projecting over anterior margin of subgenital plate .................... Pterulina simoensi sp. nov.
- Praeopercular organ strongly tapering towards the posterior, resulting into a single spine
projecting over anterior margin of subgenital plate ....... Pterulina distinctissima comb. nov.
DISTRIBUTION. Central Vietnam (Fig. 1). The genus is restricted to the Truong Son Range with
Pterulina distinctissima (Redtenbacher, 1908) comb. nov. from the Kon tum Plateau and
Pterulina simoensi sp. nov. from the Dac Lac Plateau.
COMMENT: The Kon Tum and Dac Lac Plateaus of the Truong Son Range are both recognized
as areas of high endemism. (STERLING & HURLEY, 2005).
Fig. 1. Pterulina spp. distribution map.
Belgian Journal of Entomology 96: 130 (2020)
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Pterulina distinctissima (Redtenbacher, 1908) comb. nov.
(Figs 110)
Sipyloidea distinctissima REDTENBACHER, 1908: 546 [described].
Sipyloidea distinctissima OTTE & BROCK, 2005: 317 [catalogued]; DELFOSSE ET AL., 2019:
199 [catalogued].
MATERIAL EXAMINED
TYPE MATERIAL (examined from detailed photographs): Vietnam: Holotype : Cochinchine,
Deyrolle, 1861, MNHN-EO-PHAS355 (MNHN).
Fig. 2. Sipyloidea distinctissima Redtenbacher, 1908, holotype . © MNHN. A, habitus, dorsal view.
B, habitus, ventral view. C, habitus, lateral view. D, labels.
BRESSEEL J. & CONSTANT J. Pterulina gen. nov. from Vietnam (Phasmida: Clitumninae: Clitumnini)
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ADDITIONAL MATERIAL: : Vietnam, Da Nang prov., Ba Na-Nui Chua Nat. Res., 18°09'N
105°55'E, 16-19.vii.2017, GTI Project, Leg. J. Constant & J. Bresseel, I.G.: 33.498 (RBINS);
: C Vietnam, Bach Ma N. P., 16°12’N 107°52’E, 12–17.VII.2011, leg. J. Constant & J.
Bresseel, I.G.: 31.933 (RBINS); 7♂♂, 10♀♀, eggs: Vietnam, Gia Lai prov. Kon Chu Rang
N.R., 6001200m, 13–20.vii.2018, GTI project, 14°28’28”N 108°32’27”E, Leg. J. Constant, J.
Bresseel & X. Vermeersch, I.G.:33.769 (5♂♂, 8♀♀: RBINS; 2♂♂, 2♀♀: VNMN).
COMMENTS: REDTENBACHER (1908) described the species in Sipyloidea Brunner von
Wattenwyl, 1893 based on a single male specimen lacking the abdomen at the time of
description (Fig. 2). Redtenbacher, not having seen the apex of the abdomen, nor the female,
probably placed the species in Sipyloidea because of the presence of wings, the slender
appearance and the relatively long antennae. However, these are not surpassing the front legs
like in Sipyloidea.
DESCRIPTION. The colouration is described based on dried and live specimens. Measurements:
see table 1.
MALE (Figs 25, 6 A, CD).
Colouration: Body colouration constant, head and thorax predominantly black with a distinct
yellowish circular marking between eyes slightly tapering towards the back and yellow
postocular line; genae yellow, mouthparts black. Pro-, meso- and metanotum with lateral
margins with a narrow indistinct yellowish line; prosternum black with distinct yellow marking
anteriorly, profurcasternum black; mesosternum black, mesofurcasternum yellow;
metasternum with broad cream yellow longitudinal band, lateral margins black. Metapleura
black with longitudinal cream-yellow line at lateral margins. Abdominal terga black with
indistinct cream-yellow lateral margins, yellowish margins broader on tergum VIIIIX.
Abdominal sterna IIV with two parallel longitudinal cream-yellow lines submedially, other
portions black; other sterna predominantly black. Profemora black with yellow base. Mid and
hind legs with coxae and trochanter black, femora and tibiae reddish brown (orange brown in
dried specimens). Tegmina with cream-yellow outer margins, otherwise blackish; alae with
costal area black and major veins yellowish; anal area translucent brown, sometimes with few
transparent cells.
Head: Longer than wide, slightly narrowing towards the posterior and smooth. Vertex slightly
elongated and flattened, back of head indented medially and submedially creating four
indistinct humps. Area between eyes slightly raised. Eyes circular and strongly projecting
hemispherically. Antennae black, distinctly projecting over profemora. Antennae with 2325
segments, scapus strongly dorsoventrally flattened, slightly swollen towards the posterior and
only slightly broader than antennomeres. Pedicellus very short, knob-like and cylindrical.
Antennomeres fairly elongated and covered in minute setae, except for apical antennomere.
Thorax: Pronotum shorter than head, more or less rectangular, lateral margins indistinctly
sinuate; anterior margin concave and slightly thickened, followed by a transverse depression;
medially with longitudinal line, not reaching posterior margin; transverse central depression not
reaching lateral margins. Posterior margin slightly concave to more or less straight. Mesonotum
about five times longer than pronotum, first slightly narrowing, but later slightly widening in
the posterior portion; faintly granulose with fine mediolongitudinal carina. Metanotum short,
about one third of median segment.
Wings: Tegmina projecting over posterior margin of metanotum, with definite basal hump and
broadening towards the posterior, apex broadly rounded. Alae projecting over posterior margin
of tergum IV.
Belgian Journal of Entomology 96: 130 (2020)
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Fig. 3. Pterulina distinctissima (Redtenbacher, 1908) comb. nov., . A, habitus, dorsal view.
B. habitus, ventral view. C, head and thorax, dorsal view. D, head and thorax, ventral view. E, habitus, lateral
view. F, head and thorax, lateral view.
BRESSEEL J. & CONSTANT J. Pterulina gen. nov. from Vietnam (Phasmida: Clitumninae: Clitumnini)
10
Legs: Profemora laterally flattened and curved basally; about as long as head and thorax,
including median segment, combined; all carinae present, medioventral carina indistinct. All
carinae armed with minute stiff black setae. Mesofemora about two thirds as long as profemora;
straight basally, medioventral carina absent, other features as in profemora. Metafemora longer
than mesofemora but shorter than profemora, otherwise like mesofemora. Protibiae about one
third longer than profemora, with all carinae armed with minute stiff black setae. Mesotibiae
slightly longer than mesofemora; shaped as protibiae, but carinae with few minute, but acute
spines in posterior portion. Metatibiae about as long as profemora, shaped as mesotibiae, but
with minute spines more numerous, especially towards the posterior.
Fig. 4. Pterulina distinctissima (Redtenbacher, 1908) comb. nov., terminalia. A, dorsal view. B, lateral view.
C, laterodorsal view. D, ventral view.
Abdomen: Abdominal terga smooth with a faint median, longitudinal line. Median segment
longest of all abdominal terga. Abdominal terga IIVI more or less the same length, VIIIX
distinctly decreasing in length. Tergum VIII slightly broadening towards the posterior, IX
shorter than VIII and tectiform. Anal segment distinctly longer than tergum IX, about as long
as VII and with median with longitudinal carina; apically split into two hemi-tergites. Inner
Belgian Journal of Entomology 96: 130 (2020)
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Fig. 5. Pterulina distinctissima (Redtenbacher, 1908) comb. nov., live in situ in Kon Chu Rang N.R.. A, lateral
view. B, dorsal view. C, multiple males with female.
BRESSEEL J. & CONSTANT J. Pterulina gen. nov. from Vietnam (Phasmida: Clitumninae: Clitumnini)
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portion of hemi-tergite armed with black hook-like teeth; hemi-tergites very elongate, slender
and tapered towards a narrow apex; gently curving downwards. Poculum moderately elongated,
gently rounded with a mediolongitudinal carina in the apical half, slightly projecting over apex
of abdominal tergum IX. Apex of poculum narrow, more or less triangular in ventral view.
Cerci short, not reaching apex of anal segment, round in cross-section, indistinctly incurving
with apex rounded.
FEMALE (Figs 5 C, 6 B, 78).
Colouration: Body and head with different shades of brown with darker speckles. Front legs
coloured like body, mid- and hind legs distinctly paler, from pale brown to reddish brown.
Head: Longer than wide, slightly narrowing towards the posterior and slightly granulose.
Vertex slightly elongated and flattened, back of head indented medially and submedially
creating four indistinct humps. Area between eyes slightly raised with transverse black marking.
Eyes circular and strongly projecting hemispherically. Antennae coloured as head; short,
reaching slightly over halfway along profemora. Antennae with 2325 segments, scapus
strongly dorsoventrally flattened, laterally rounded, distinctly broader than pedicellus and
antennomeres. Pedicellus very short, more than three times shorter than scapus, dorsoventrally
Fig. 6. Pterulina distinctissima (Redtenbacher, 1908) comb. nov. A, , head and pronotum dorsal view.
B, , head and pronotum dorsal view. C, wings, dorsal view. D, wings, laterodorsal view. © T. Bollens.
Belgian Journal of Entomology 96: 130 (2020)
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Fig. 7. Pterulina distinctissima (Redtenbacher, 1908) comb. nov., from Kon Chu Rang 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 ventral view. G, terminalia, lateral view. H, head and thorax, lateral view.
BRESSEEL J. & CONSTANT J. Pterulina gen. nov. from Vietnam (Phasmida: Clitumninae: Clitumnini)
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flattened and laterally rounded. Apical antennomeres distinctly shorter than anterior
antennomeres, apical antennomere elongated, club-like.
Thorax: Pronotum sparsely granulose, distinctly shorter and narrower than head. Lateral
margins slightly sinuate; anterior margin concave and slightly thickened, followed by
transverse depression; medially with longitudinal line, not reaching posterior margin; transverse
central depression not reaching lateral margins; posterior margin slightly concave. Mesonotum
about six times longer than pronotum, slightly widening in posterior portion; faintly granulose
with slightly larger, evenly spaced granules laterally and with fine mediolongitudinal line.
Metanotum slightly longer than median segment with fine mediolongitudinal line,
posterolaterally with minute expansion; anterior and posterior margins straight.
Legs: Profemora slightly longer than pro-, meso- and metanotum combined; compressed and
strongly curved basally. All carinae present; anterodorsal carina with small teeth, decreasing
towards the posterior; posterodorsal carina with one to four spaced triangular teeth.
Mesofemora slightly longer than mesonotum, both dorsal carinae armed with three evenly
spaced minute to triangular teeth. Metafemora reaching halfway along tergum V, armed like
mesofemora. Protibiae carinate, distinctly longer than profemora. Dorsal carinae close to each
other, posterodorsal carina with one to five triangular teeth. Posteroventral carina with few
minute, acute teeth; medioventral carina slightly raised and laterally flattened. Mesotibiae about
as long as mesofemora; posterodorsal carina variable, sometimes with few minute spines or
with up to three triangular teeth; anterodorsal carina often with minute spine subapically;
ventral carinae with few minute spines, more numerous towards the posterior. Metatibiae
slightly longer than metafemora; all carinae armed with minute spines; anterodorsal carina
sometimes with few triangular teeth.
Abdomen: Abdominal terga sparsely granulose with faint mediolongitudinal line. Median
segment slightly shorter than metanotum. Abdominal terga II-V slightly gradually increasing
in length; VIX gradually decreasing. Anal segment concave apically, lateral margins rounded,
posterolateral angles about 90°. Short triangular epiproct visible, slightly projecting over apex
of anal segment with mediolongitudinal carina. Sternum VII with distinct praeopercular organ.
Praeopercular organ strongly tapering towards the posterior, terminating in a single spiniform
swelling projecting over anterior margin of subgenital plate. Subgenital plate strongly keeled,
laterally compressed with a mediolongitudinal carina in posterior portion. Posterior margin
rounded.
NYMPH (Fig 9). Newly hatched nymphs are slender and predominantly light green. Head bigger
than pronotum, with black postocular line. Abdomen tipped black. Legs dark with pale
markings. Older nymphs brown or a combination of green and brown.
EGG (Fig 10). Measurements (in mm): length: ~7; width: ~1.2; height: ~2.7.
Capsule predominantly dark brown, distinctly paler dorsally; strongly elongated and laterally
flattened. Ventral margin convex, dorsal margin more or less straight. Capsule minutely and
densely granulose with two more densely granulose, small, pale spots near posteroventral and
posterodorsal margins and two elongate, pale granulose markings on polar area. Anterodorsal
margin longer than anteroventral one in lateral view. Polar area distinctly notched in lateral
view; posterodorsal and posteroventral angles rounded, posteroventral margin slightly longer.
Operculum pale, egg-shaped with a central oval, brown hole from anterior view; surface pale
with distinct granulation; opercular rim distinctly paler than capsule, incurving and granulose.
Micropylar plate distinct, smooth, coloured as capsule with indistinct paler marking anteriorly
of the micropylar cup; elongate spear-shaped and almost completely restricted to dorsal surface;
Belgian Journal of Entomology 96: 130 (2020)
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Fig. 8. Pterulina distinctissima (Redtenbacher, 1908) comb. nov., live in situ. A, dorsal view in Ba Na-Nui Chua
N.R. B, laterodorsal view, head and thorax in Ba Na-Nui Chua N.R. C, nymph in Kon Chu Rang N.R. D, dorsal
view in Kon Chu Rang N.R. E, dorsal view head and thorax in Kon Chu Rang N.R.
BRESSEEL J. & CONSTANT J. Pterulina gen. nov. from Vietnam (Phasmida: Clitumninae: Clitumnini)
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Fig. 9. Pterulina distinctissima (Redtenbacher, 1908) comb. nov., newly hatched nymph. A, dorsolateral view, B-
C, dorsal view. © T. Bollens.
Belgian Journal of Entomology 96: 130 (2020)
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Fig. 10. Pterulina distinctissima (Redtenbacher, 1908) comb. nov. egg. A, lateral view. B, dorsal view. C, latero-
dorsal view. D, ventral view. E, anterolateral view. F, operculum. G, polar view. H, polar area, lateral view.
BRESSEEL J. & CONSTANT J. Pterulina gen. nov. from Vietnam (Phasmida: Clitumninae: Clitumnini)
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margins broad and pale. Micropylar cup small, yellowish, an almost completely circular
swelling with short median line.
BIOLOGY. The species was collected in mountainous tropical evergreen rainforest, at medium
altitude (6001200 m). The specimens were observed on lower vegetation and small trees up
to 3 m and seemed to feed on a variety of plant species. Males are able to fly and fly away when
disturbed during day time. The wings are also used to stridulate by rubbing the outer margins
of the tegmina against the subcostal and radial veins of the alae, which bear numerous tooth-
like structures. The eggs are dropped to the ground. In captivity, the species accepts Rubus spp.,
other Rosaceae and Carpinus betulus L. (Betulaceae) as alternative food-plants.
Table 1. Measurements [mm] of Pterulina distinctissima (Redtenbacher, 1908) comb. nov.
HT*
♂♂
♀♀
75.287.3
109.7112.7
3.44.1
5.56.1
2.52.7
3.73.9
9
13.214.9
23.223.7
8.5
3.54.0
8.38.5
11.011.8
7.37.5
3.94.8
33.738.0
36
32.937.6
37.938.4
22
20.325.4
23.724.9
27
24.630.9
28.328.9
42.150.5
41.244.6
23.827.9
24.126.0
31.338.9
32.434.0
* After REDTENBACHER (1908)
DISTRIBUTION (Fig. 1). Known from Bach Ma N.P., Ba Na-Nui Chua N.P., and Kon Chu Rang
N.R. in the central Annamite Mountains.
Pterulina simoensi sp. nov.
urn:lsid:zoobank.org:act:A45C70AF-8519-4582-AA4B-BEE5BB333560
Figs (1, 1117)
ETYMOLOGY. The species is named after Mr. Rob Simoens (Veurne, Belgium) acknowledging
his help and support with the authors’ study over many years.
TYPE MATERIAL: Holotype : Vietnam, Dak Lak prov., Chu Yang Sin N.P., 650
1000 m,12°27’24”N 108°22’15”E, 9–15.VIII.2019, GTI Project, Leg. J. Constant &
J. Bresseel, I.G.:34.048 (RBINS).
Paratypes (4♂♂, 7♀♀): 1, 3♀♀: same data as holotype. (RBINS); 3♂♂, 3♀♀: same data as
holotype, ex breeding T. Bollens, 2020 (2♂♂, 5♀♀: RBINS; 2♂♂, 2♀♀: VNMN).
ADDITIONAL MATERIAL: Eggs: same data as holotype, ex breeding T. Bollens, 2020 (RBINS).
DIAGNOSIS. The new species much resembles the only other species in the genus: Pterulina
distinctissima (Redtenbacher, 1908) comb. nov., but is distinctly smaller and more slender.
Belgian Journal of Entomology 96: 130 (2020)
19
Males can stridulate but have wings that only reach the posterior margin of tergum III; the
distinct colouration of head and body can easily separate them. Females have the colouration
like Pterulina distinctissima (Redtenbacher,1908) comb. nov. and also have the strongly keeled
and laterally flattened subgenital plate, but can easily be distinguished by the apically rounded
praeopercular organ, not projecting over posterior margin of sternum VII.
DESCRIPTION. The colouration is described from dried specimens and pictures of live
specimens. Measurements: see table 2.
MALE (Figs 1112, 13A, CE, 14A).
Colouration: Body colouration constant. Head with black oval area between eyes continued by
a broad, black mediolongitudinal stripe; postocular line black starting at base of eye; area
between dorsal line and postocular line and beneath postocular line coloured cream-yellow.
Pronotum yellowish brown with broad, black mediolongitudinal stripe; prosternum yellowish
brown. Mesonotum coloured as metanotum, but with black mediolongitudinal stripe distinctly
broader. Mesosternum black, mesofurcasternum yellowish brown; metasternum black with
mediolongitudinal pale line. Abdominal terga IVVIII light brown with outer margins black
and with mediolongitudinal black line, IX darker, hemi-tergites almost completely black. Fore
legs black with yellowish brown base. Mid and hind legs with femora and tibiae reddish brown
(orange brown in dried specimens), apices dark. Tegmina with cream-yellow outer margin,
otherwise blackish; alae with costal area black with major veins yellowish, anal area translucent
brown.
Head: Longer than wide, slightly narrowing towards the posterior and smooth. Dorsally with
faint median line. Vertex slightly elongated and flattened, back of head indented medially and
submedially creating four indistinct humps. Area between eyes slightly raised. Eyes circular
and strongly projecting hemispherically. Antennae black, projecting over profemora. Antennae
with 2023 segments; scapus strongly dorsoventrally flattened, slightly swollen towards the
posterior and broader than antennomeres. Pedicellus very short, knob-like and cylindrical.
Antennomeres elongated and covered in minute setae.
Thorax: Pronotum shorter than head, more or less rectangular, with lateral margins indistinctly
sinuate; anterior margin concave and slightly raised, followed by transverse depression;
medially with longitudinal line, not reaching posterior margin; transverse central depression not
reaching lateral margins. Posterior margin more or less straight. Mesonotum more than 5 times
longer than pronotum, first slightly narrowing, but later slightly widening in posterior portion;
faintly granulose with fine mediolongitudinal carina. Metanotum short, about one third of
median segment.
Wings: Tegmina blackish dorsally and yellowish laterally with definite basal hump; broadening
towards the posterior, apex broadly rounded. Alae reaching posterior margin of tergum III.
Legs: Profemora laterally flattened and curved basally; about as long as head and thorax
(including median segment) combined; all carinae present, medioventral carina indistinct. All
carinae armed with minute stiff black setae. Mesofemora about two thirds as long as profemora;
straight basally, carinae less distinct; medioventral carina absent, other features as in profemora.
Metafemora longer than mesofemora but shorter than profemora, shaped as mesofemora.
Protibiae slightly longer than profemora, with all carinae armed with minute stiff black setae.
Mesotibiae about as long as mesofemora; shaped as protibiae, but carinae with few minute, but
acute spines distally. Metatibiae shorter profemora, shaped as mesotibiae.
Abdomen: Abdominal terga smooth with faint median, longitudinal line. Median segment
longest of all abdominal terga. Abdominal terga IIVI more or less the same length, VIIIX
BRESSEEL J. & CONSTANT J. Pterulina gen. nov. from Vietnam (Phasmida: Clitumninae: Clitumnini)
20
Fig. 11. Pterulina simoensi sp. nov., holotype . A, habitus, dorsal view. B. habitus, ventral view. C, head and
thorax, dorsal view. D, head and thorax, ventral view. E, habitus, lateral view. F, head and thorax, lateral view.
Belgian Journal of Entomology 96: 130 (2020)
21
distinctly getting shorter. Tergum VIII slightly broadening towards the posterior, IX shorter
than VIII and tectiform. Anal segment distinctly longer than tergum IX, slightly shorter than
VII and with median longitudinal carina; apically split into two hemi-tergites. Inner portion of
hemi-tergite armed with black hook-like spines. Hemi-tergites very elongate, slender and
tapered towards pointed apex. Poculum moderately elongated, gently rounded, reaching
posterior margin of tergum IX; apex narrow, more or less triangular from ventral view. Cerci
short, not reaching apex of anal segment, round in cross-section, indistinctly incurving with
apices rounded.
FEMALE (Figs 13B, 14BC, 15).
Colouration: Body and head in different shades of brown, sometimes with darker or paler
markings or speckles. Mid- and hind legs distinctly paler than front legs.
Head: Longer than wide, slightly narrowing towards the posterior and granulose with a pair of
small tubercles between eyes; vertex slightly elongated and flattened; back of head indented
medially and submedially creating four indistinct humps. Eyes circular and strongly projecting
hemispherically. Antennae coloured as head; short, slightly projecting over halfway profemora.
Antennae with 2426 segments; scapus strongly dorsoventrally flattened, laterally rounded,
distinctly broader than pedicellus and antennomeres. Pedicellus very short, more than three
times shorter than scapus; dorsoventrally flattened and laterally rounded. Distal antennomeres
distinctly shorter than proximal ones, apical antennomere elongated, club-like.
Fig. 12. Pterulina simoensi sp. nov., holotype , terminalia. A, dorsal view. B, lateral view. C, ventral view.
BRESSEEL J. & CONSTANT J. Pterulina gen. nov. from Vietnam (Phasmida: Clitumninae: Clitumnini)
22
Thorax: Pronotum sparsely granulose, distinctly shorter and narrower than head. Lateral
margins slightly sinuate; anterior margin concave and slightly thickened, followed by a
transverse depression; medially with longitudinal line, not reaching posterior margin; transverse
central depression not reaching lateral margins; prozona with a raised elongate area on each
side of mediolongitudinal impression. Posterior margin slightly concave. Mesonotum about six
times longer than pronotum, slightly widening in posterior portion; faintly granulose with
slightly larger, evenly spaced granules laterally and with fine mediolongitudinal line.
Metanotum slightly longer than median segment with fine mediolongitudinal line; anterior and
posterior margins straight.
Fig. 13. Pterulina simoensi sp. nov. live specimens. A, , head and pronotum, dorsal view. B, , head and
pronotum, dorsal view. C, , head and pronotum, lateral view. D, , wings, dorsal view. E, wings,
laterodorsal view.
Legs: Profemora slightly shorter than pro-, meso- and metanotum combined; compressed and
strongly curved basally; all carinae present; anterodorsal carina with small teeth, decreasing in
number towards the posterior; posterodorsal carina with one to four spaced saw-like spines.
Mesofemora distinctly shorter than mesonotum, both dorsal carinae armed with three evenly
spaced minute spines. Metafemora projecting over posterior margin of tergum IV, armed as
mesofemora. Protibiae carinate, distinctly longer than profemora. Dorsal carinae close to each
other, posterodorsal carina with four triangular teeth, one subbasally, one medially and two
Belgian Journal of Entomology 96: 130 (2020)
23
Fig. 14. Pterulina simoensi sp. nov., live specimens in situ in Chu Yang Sin N.P., A, , dorsal view, 1000 m.
B, nymph, dorsal view, 1600 m. C, , dorsal view, 1000 m.
BRESSEEL J. & CONSTANT J. Pterulina gen. nov. from Vietnam (Phasmida: Clitumninae: Clitumnini)
24
posteriorly. Posteroventral carina with few minute, acute teeth; medioventral carina slightly
raised and laterally flattened. Mesotibiae about as long as mesofemora; posterodorsal carina
with few minute spines, anterodorsal carina with minute spine subapically; ventral carinae with
few minute spines, more numerous towards the posterior. Metatibiae slightly longer than
metafemora, with all carinae armed with minute spines.
Table 2. Measurements [mm] of Pterulina simoensi sp. nov.
Length of
HT
♂♂
♀♀
Body:
60.8
54.6
82.085.1
Head:
3.0
2.8
4.64.7
Pronotum:
2.1
2.1
3.23.7
Mesonotum:
13.1
11.0
18.719.1
Metanotum:
2.7
5.96.2
Median segment:
7.3
5.05.4
Tegmina:
3.0
2.8
Alae:
18.9
17.0
Profemora:
27.5
26.5
21.325.2
Mesofemora:
16.9
16.8
15.415.5
Metafemora:
21.0
20.6
18.418.8
Protibiae:
31.8
29.8
27.128.3
Mesotibiae:
17.3
17.2
15.615.9
Metatibiae:
24.3
21.722.1
Abdomen: Abdominal terga sparsely granulose with faint median, longitudinal line. Median
segment slightly shorter than metanotum. Abdominal terga IIV indistinctly gradually
increasing in length; VIX gradually getting shorter. Anal segment concave apically, with
lateral margins rounded, posterolateral angles about 90°. Short triangular epiproct visible,
slightly projecting over apex; anal segment with mediolongitudinal carina. Sternum VII with a
distinct praeopercular organ. Praeopercular organ tapering towards the posterior; anterior
portion raised, posterior portion sulcate with posterior margin rounded and not projecting over
anterior margin of subgenital plate. Subgenital plate boat-shaped, laterally compressed with
mediolongitudinal carina in posterior portion; posterior margin rounded.
NYMPH (Fig. 16). Newly hatched nymphs slender and predominantly light green. Head bigger
than pronotum, with black postocular line. Abdomen tipped black. Legs dark with pale
markings. Older nymphs brown or a combination of green and brown.
EGG (Fig. 17). Measurements (in mm): length: ~6.5; width: ~1.1; height: ~1.6.
Capsule predominantly dark brown with indistinct pale network, distinctly paler towards
micropylar plate; strongly elongated and laterally flattened. Ventral margin convex, dorsal
margin more or less straight. Capsule minutely granulose, with two small pale spots near
posterodorsal margin and four pale spots on polar area. Polar area distinctly notched in lateral
view; posterodorsal and posteroventral angles rounded. Operculum pale, oval with central oval,
brown hole in anterior view; surface pale with distinct granulation; opercular rim distinctly
paler than capsule, incurving and granulose. Micropylar plate distinct, smooth, coloured as
capsule with pale oval marking anteriorly of micropylar cup; margins broad and pale; spear-
shaped and almost completely restricted to dorsal surface. Micropylar cup small, yellowish, an
almost completely circular swelling with short median line.
Belgian Journal of Entomology 96: 130 (2020)
25
Fig. 15. Pterulina simoensi sp. nov., . A, habitus, dorsal view. B. habitus, lateral view. C, habitus, ventral view.
D, head and thorax, dorsal view. E, terminalia, anterodorsal view. F, terminalia, dorsal view. G, terminalia ventral
view. H, terminalia, lateral view. I, head and thorax, lateral view.
BRESSEEL J. & CONSTANT J. Pterulina gen. nov. from Vietnam (Phasmida: Clitumninae: Clitumnini)
26
Fig. 16. Pterulina simoensi sp. nov., newly hatched nymph. A, dorsal view. B, laterodorsal view. C, detail of head.
© T. Bollens.
Belgian Journal of Entomology 96: 130 (2020)
27
Fig. 17. Pterulina simoensi sp. nov., egg. A, lateral view. B, dorsal view. C, laterodorsal view. D. Ventral view.
E, anterolateral view. F, operculum. G, polar view. H, polar area, lateral view.
BRESSEEL J. & CONSTANT J. Pterulina gen. nov. from Vietnam (Phasmida: Clitumninae: Clitumnini)
28
BIOLOGY. The species was collected in mountainous tropical evergreen rainforest, at medium
to high altitude (10001600 m). The specimens were observed on lower vegetation and seemed
to feed on a variety of plant species. Males can stridulate. The eggs are dropped to the ground.
In captivity, the species accepts Rubus spp. and other Rosaceae as alternative food-plants.
DISTRIBUTION (Fig. 2). This species is only known from Chu Yang Sin N.P. in the southern
Annamites.
Discussion
PLACEMENT IN THE CLITUMNINI
BRUNNER VON WATTENWYL (1893) first erected the tribe under the name Clitumnidae which he
later corrected to Clitumnini (BRUNNER VON WATTENWYL, 1907). It was in this last publication
that the Clitumnini were described and diagnosed from characters like the absence of wings;
the short antennae not surpassing the profemora; the short and roughly cylindrical cerci; the
medially split anal segment in males and the keeled subgenital plate in females.
HENNEMANN & CONLE (2008) upgraded the tribe to subfamily level which currently includes
Pharnaciini Günther, 1953, Medaurini Hennemann & Conle, 2008 and Clitumnini; the tribe was
re-diagnosed and revised on the generic level.
The traditional definition of Clitumnini was recently adapted (BRESSEEL & CONSTANT, 2015)
to include Lobofemora Bresseel & Constant, 2015 which has a comparatively longer median
segment, and possesses tegmina and vestigial or short alae in the males.
The tribal placement of Pterulina gen. nov. proposed herein is based on a set of characters
which key out to Clitumnini following HENNEMANN & CONLE (2008) and BRESSEEL &
CONSTANT (2015): Very slender and stick-like insects with elongated legs, sexual dimorphism
distinct; head with antennae with less than 28 segments. It has the mesothorax less than 2.5x
the combined length of head and prothorax in females, and has tegmina and short alae in the
males. Cerci are small, distinctly shorter than the anal segment and circular in cross-section.
Males have the anal segment consisting of two movable hemitergites which are minutely
toothed interiorly and lack an external, sclerotized vomer; females have a convex and boat-
shaped subgenital plate which slightly extends over the anal segment. The armature of the legs
is strongly reduced and the medioventral carina of the meso- and metafemora is indistinct and
unarmed.
The eggs are elongated, laterally flattened, more or less parallel-sided and seed-like with a
specialised rim on the outer border of the operculum, a character shared with several other
Clitumnini like Ramulus Saussure, 1862 and Cuniculina Brunner von Wattenwyl, 1907.
The new genus violates a key character for Clitumnini with males having the antennae distinctly
longer than the profemora. The length of antennae in Clitumnini is subject to variation to a
certain degree; in several species of Ramulus saussure, 1862 the antennae reach two thirds along
profemora; but they are almost as long as the profemora in Cuniculina cunicula Brunner von
Wattenwyl, 1907 (HENNEMANN, 2002). The definition of Clitumnini is therefore expanded in
terms of males with antennae at best projecting over profemora, not reaching halfway along the
protibiae.
STRIDULATION
Males in both species of Pterulina gen. nov. are able to stridulate. To produce sound they use
the same stridulation mechanism as Lobofemora Bresseel & Constant, 2015. They rubb the
Belgian Journal of Entomology 96: 130 (2020)
29
outer margins of the tegmina against the subcostal and radial veins of the alae, which bear
numerous very minute tooth-like structures (BRESSEEL & CONSTANT, 2015). The stridulation
occurs when the specimens are disturbed.
BIOGEOGRAPHY
The northernmost distribution data of Pterulina gen. nov. is Bach Ma National Park which is
situated in the central Annamite Mountains and lies on a high mountain ridge that runs west-
east from the Laotian border to the East Sea at the Hai Van pass. It acts as a biogeographical
border between north and south Vietnam. This can also be observed in the distribution pattern
of certain genera of stick insects, e.g. Neohirasea Rehn, 1904, Micadina Redtenbacher, 1908
and Pachyscia Redtenbacher, 1908 have their southern distribution limit at Bach Ma N.P. (HO,
2018a, b; Bresseel & Constant, unpublished data) while Pterulina gen. nov., Lobofemora
Bresseel & Constant, 2015 and Phamartes Bresseel & Constant, 2013 have their northernmost
distribution limit a this same place (BRESSEEL & CONSTANT, 2013, 2015, unpublished data).
The Annamite Range, the region’s dominant upland formation, runs from northwest to
southeast through Laos, Vietnam, and northeastern Cambodia for roughly 1200 km. It
represents an area (or areas) of high endemism and its origin lies with a series of folding events,
the Indosinian Orogeny that took place 400350 Mya, establishing mountains that would later
become parts of the Hoang Lien Son and the northern Annamite ranges. This was followed
350300 Mya by the formation of uplands in Vietnam’s Central and Southern Annamites. The
Southern Annamites consist of a suite of plateaus that are isolated geographically from the
Central Annamites. It is also a younger formation created by different orogenies than those that
uplifted the Central Annamites (BAIN & HURLEY, 1966). Pterulina distinctissima
(Redtenbacher, 1908) comb. nov. and P. simoensi sp. nov. occur in the Central and Southern
Annamites respectively.
It could be hypothesised that populations of the ancestors of Pterulina gen. nov. got isolated
during this period of uplifting, which resulted in the evolution of different species.
Acknowledgments
We thank Dr Hong Thai Pham (VNMN) and his colleagues: Mr Van Dat Nguyen, Mr Thanh Trung Vu, Mrs Thi
Thu Huong Nguyen, Mrs Thi Man Nguyen and Mr Hoang Vu Tru (IEBR), and Mr Xavier Vermeersch
(collaborator, RBINS) for all their help and friendship during the collecting trips; the authorities of the protected
areas where we have sampled, for supporting our studies. We are deeply indebted to Tim Bollens (Herselt,
Belgium) for breeding and photographing both species and providing additional material. Mr Emmanuel Delfosse
and Dr Tony Robillard (MNHN) are thanked for their help during our visit to MNHN; Dr Patrick Grootaert, Dr
Yves Samyn and Dr Marie-Lucie Susini (RBINS) for their permanent support to our projects in Vietnam and Miss
Mado Berthet (RBINS) for adjusting the plates illustrating the paper. This paper is a result of the projects “A step
further in the Entomodiversity of Vietnam” (2010–2019) supported through a grant issued by the capacity building
Programme of the Belgian Global Taxonomy Initiative National Focal Point that runs under the CEBioS
programme with financial support from the Belgian Directorate-General for Development Cooperation (DGD).
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Indochina (Laos, Cambodia, and Vietnam) houses over 600 species of amphibians and reptiles, roughly a quarter of which has been described within the last 15 years. Herein, we undertake the first biogeographic synthesis of the regional herpetofauna since the first half of the 20th century. We review the literature to measure and map species richness and endemism, the contributions of regional faunas, and ecological characteristics of Indochina’s amphibians (Anura, Caudata), and reptiles (Serpentes, Sauria, Testudines, Crocodylia). Dividing Indochina into 19 subregions defined by topography and geology, we estimate the similarity among the regional faunas and appraise the effects of area and survey effort on these comparative analyses. Variation in species composition is broadly correlated with topography, habitat complexity, and proximity to regions outside Indochina. Indochina’s herpetofauna is dominated (in decreasing order) by endemic species, widely distributed species, a South China fauna, and a biota centered in Thailand and Myanmar. Species richness is highest in amphibians and snakes, and peaks in upland forests. Endemism, highest among amphibians and lizards, also peaks in forests of the region’s northern uplands and Annamite Range. Endemic species occupy a narrower range of habitats than nonendemics. Patterns of richness and endemism are partially explained by ecological constraints: amphibians and lizards are more restricted to forests than snakes, turtles, and crocodiles; amphibians are more restricted to uplands, turtles to lowlands. We also assess biogeography in the context of Indochina’s geology, climate, and land cover. In northern Indochina, the Red River either acts as or coincides with an apparent dispersal barrier. Herpetofauna in northeastern upland areas are closely allied with fauna of southeastern China. In southern Indochina there is little evidence that the Mekong River represents a biogeographic barrier to the regional herpetofauna. The Annamite Range is composed of at least three distinct units and its elevated species richness and endemism are also noted in adjacent lowlands. Contribution of subtropical biota to Indochina’s fauna is significantly greater than that of tropical biota and there is little other evidence for intermixing at intermediate latitudes. Our results have implications for biogeography and conservation efforts, although they must be viewed in the context of rapidly evolving systematic knowledge of the region’s amphibians and reptiles. Future survey efforts, and the phylogenetic analyses that come from them, are essential for supporting regional conservation efforts, as they will better resolve the known patterns of amphibian and reptile richness and endemism.
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A new genus and species of stick insect Phamartes coronatus gen. nov. sp. nov. from Bach Ma National Park, central Vietnam is described and illustrated from both sexes and the egg. The genus is most closely related to Oxyartes Stål, 1875, but easily distinguishable by the presence of fully developed alae, the head armature and the split and asymmetrical anal segment, a character previously unknown in Necrosciinae.
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The family Phasmatidae Gray, 1835 is reviewed and the subfamily Phasmatinae shown to be polyphyletic. Based on features of the exosceleton of the insects, egg-morphology and copulation habits a new arrangement of Phasmatidae is proposed. The monophyly of Lanceocercata Bradler, 2001 is confirmed but this name shown to be a synonym of Phasmatidae, hence Lanceocercata is here referred to as Phasmatidae sensu stricto. Six subfamilies belong in Phasmatidae sensu stricto all of which share several common and supposedly apomorphic characters: Phasmatinae, Tropidoderinae, Extatosomatinae (stat. nov.), Xeroderinae, Pachymorphinae and “Platycraninae”. The other two subfamilies contained in Phasmatidae sensu Bradley & Galil, 1977 (Eurycanthinae and Cladomorphinae) are not cosely related and here regarded as subfamilies of Phasmatidae sensu lato. The subfamily Phasmatinae sensu Bradley & Galil, 1977 is shown to be polyphyletic. The two tribes Pharnaciini and Clitumnini (= Baculini Günther, 1953) are removed from Phasmatinae and shown to be closely related to each other. They are transferred to the here established subfamily Clitumninae, a subordinate clade of Phasmatidae sensu lato. The subfamily Lonchodinae is closely related to Clitumninae, hence removed from Diapheromeridae and transferred to Phasmatidae sensu lato. The tribes Achriopterini and Stephanacridini (formerly in Phasmatinae) are shown to be not closely related to either Phasmatinae sensu stricto, Clitumninae or Lonchodinae, and provisionally must be treated as tribes of Phasmatidae sensu lato (incerte sedis). A re-arrangement of Phasmatidae sensu stricto is proposed along with determinating keys to all subfamilies and their tribes. The subfamilies Phasmatinae, Tropidoderinae and Extatosomatinae stat. nov. are re-described and discussed in detail. Full lists of genera are provided for each tribe. Only three of seven tribes formerly in Phasmatinae remain in the subfamily, this is Phasmatini, Acanthomimini and Acanthoxylini. The subfamily Tropidoderinae contains three tribes: Tropidoderini, Monandropterini and Gigantophasmatini trib. nov. The tribe Extatosomatini Clark-Sellick, 1997 is removed from Tropidoderinae and raised to subfamily level (Extatosomatinae stat. nov.). Several genera are transferred to other tribes or subfamilies. Didymuria Kirby, 1904 is removed from Tropidoderini, since it differs by having a closed internal micropylar plate in the eggs (open in all Tropidoderini). It here remains as a genus incerte sedis of Tropidoderinae and its systematic position clearly deserves further clarification. Gigantophasma Sharp, 1898 from the Loyalty Islands is removed from Pharnaciini, and becomes the type genus of the tribe Gigantophasmatini trib. nov.. Anophelepis Westwood, 1859 is removed from “Platycraninae” and shown to belong in Phasmatinae: Acanthomimini. The two Australian genera Arphax Stål, 1875, and Vasilissa Kirby, 1896 are removed from Acanthoxylini and provisionally transferred to Acanthomimini, but their position remains as yet debatable. Echetlus Stål, 1875 is misplaced in “Platycraninae” and shown to be a likely member of Phasmatinae. The two Brazilian species Echetlus evoneobertii Zompro & Adis, 2001 and Echetlus fulgens Zompro, 2004b are obviously misplaced and belong in the New World Diapheromeridae: Diapheromerinae: Diapheromerini. The subfamily Pachymorphinae is briefly discussed and considered polyphyletic. Three genera of Pachymorphinae: Gratidiini Bragg, 1995 (Parapachymorpha Brunner v. Wattenwyl, 1893 and Cnipsomorpha Hennemann et al., 2008) are transferred to Clitumninae: Medaurini trib. nov. The genus Gongylopus Brunner v. Wattenwyl, 1907 is transferred from Pachymorphinae: Gratidiini to Clitumninae: Clitumnini. The subfamily Xeroderinae is briefly discussed and shown likely to be polyphyletic, due to it contains two fundamentally different types of genitalia in the males. Only the genera Xeroderus Gray, 1835 and perhaps Epicharmus Stål, 1875 clearly belong in Phasmatidae sensu stricto. Both, the Pachymorphinae and Xeroderinae certainly deserve more detailed investigation to clarify their systematic positions with confirmation. Two generic groups are recognized within Clitumnini (subfamily Clitumninae). Due to differing by genital features and egg-morphology Medaura Stål, 1875 and Medauroidea Zompro, 2000 are removed from Clitumnini and transferred to the newly described Medaurini trib. nov.. The new tribe furthermore contains two genera formerly included in Pachymorphinae: Gratidiini and transferred here, Cnipsomorpha Hennemann et al., 2008 and Parapachymorpha Brunner v. Wattenwyl, 1893. Phryganistria Stål, 1875 is removed from Clitumnini and transferred to Pharnaciini. Nesiophasma Günther, 1934 is shown to belong in the tribe Stephanacridini. The Australasian subfamily Lonchodinae Brunner v. Wattenwyl, 1893 has formerly been included in Diapheromeridae Zompro, 2001 (= Heteronemiidae by Bradley & Galil, 1977). However, numerous features of the genitalia and egg morphology show close relation to the Oriental subfamily Clitumninae instead. Thus, Lonchodinae is here transferred to the family Phasmatidae (sensu lato). Within Lonchodinae the new tribe Neohiraseini trib. nov. is recognized and contains the five genera formerly placed in the “Neohirasea-complex” of that subfamily, namely Andropromachus Carl, 1913, Neohirasea Rehn, 1904, Pseudocentema Chen, He & Li, 2002, Qiongphasma Chen, He & Li, 2002 and Spinohirasea Zompro, 2001. It differs from all other Lonchodinae (= tribe Lonchodini) by the well developed vomer of males and the lack of a capitulum in the eggs. The genus Cladomimus Carl, 1915 was previously misplaced in Clitumninae: Pharnaciini and is here transferred to Lonchodinae: Lonchodini. It appears to be close to the Australian Hyrtacus Stål, 1875. Leprocaulinus Uvarov, 1940 and Phenacocephalus Werner, 1930 are removed from the subfamily Necrosciinae and transferred to Lonchodinae: Lonchodini. Extensive research on the genera which belong to the tribe Pharnaciini Günther, 1953 and taking features of the genital exosceleton and egg-morphology into account, has shown this tribe to be polyphyletic. Based on such features two generic groups are easily recognized within Pharnaciini sensu Günther, 1953. Males of the first group have a longitudinally split anal segment, which consists of two separate, more or less elongate semi-tergites and forms a clasping apparatus, the vomer is strongly reduced or lacking, the profemora have a prominent, lamellate medioventral carina which is strongly displaced towards the anteroventral carina and the eggs have an open internal micropylar plate with a clear median line. Only the genera falling into this group remain in Pharnaciini. Males of the second group in contrast have an anal segment which is not split, but possess a clearly visible, well sclerotised, triangular or hook-like external vomer, an indistinct medioventral carina on the profemora and eggs with a closed internal micropylar plate. Most of the genera which fall into the second group are here transferred to the tribe Stephanacridini Günther, 1953, this is Diagoras Stål, 1877b, Eucarcharus Brunner v. Wattenwyl, 1907, Phasmotaenia Návas, 1907 and Sadyattes Stål, 1875. A detailed discussion of the differences between Pharnaciini and Stephanacridini is provided along with distinguishing keys, illustrations and maps showing the distinct geographic distributions. The five genera that belong in Pharnaciini are: Baculonistria gen. nov., Pharnacia Stål, 1877a, Phobaeticus Brunner v. Wattenwyl, 1907 (= Baculolonga Hennemann & Conle, 1997a, = Lobophasma Günther, 1934b syn. nov. , = Nearchus Redtenbacher, 1908 syn. nov. ), Tirachoidea Brunner v. Wattenwyl, 1893 stat. rev. and Phryganistria Stål, 1875. Pharnacia annulata Redtenbacher, 1908 and Pharnacia enganensis Redtenbacher, 1908 were misplaced in Pharnacia Stål, 1877 (tribe Pharnaciini) and are transferred to the genus Sadyattes Stål, 1875 (tribe Stephanacridini, comb. nov.). Phobaeticus kuehni Brunner v. Wattenwyl, 1907 is removed from Phobaeticus Brunner v. Wattenwyl, 1907 (Phasmatinae: Pharnaciini) and shown to belong in Nesiophasma Günther, 1934c (tribe Stephanacridini, comb. nov.). Phobaeticus incertus Brunner v. Wattenwyl, 1907 (= Nearchus grubaueri Redtenbacher, 1908 syn. nov.) is unlikely to belong in Pharnaciini and here only retained in the original genus Phobaeticus Brunner v. Wattenwyl, 1907 with doubt, it may belong in Nesiophasma Günther, 1934c (tribe Stephanacridini). Based on a total of almost 700 examined specimens, the Oriental tribe Pharnaciini Günther, 1953 is revised at the species level. The new genus Baculonistria gen. nov. (Type species Baculonistria alba (Chen & He, 1990) comb. nov.), is described to contain three species from Central and Eastern China. Tirachoidea Brunner v. Wattenwyl, 1893 was erroneously synonymised with Pharnacia Stål, 1877 and is here re-established as a valid genus (stat. rev.). All five genera are re-diagnosed and differentiated, their systematic position within Pharnaciini discussed, and complete synonymic and species-listings as well as distribution maps and determination keys to the insects and eggs are provided. Detailed descriptions, diagnoses, synonymic listings, illustrations, material listings, distribution maps and measurements are provided for all 40 valid species. The type material of a further two species appears to be lost. Seven new species are described: Pharnacia borneensis spec. nov. from Borneo; Pharnacia palawanica spec. nov. from Palawan, Phobaeticus mucrospinosus spec. nov. from Sumatra, Phobaeticus palawanensis spec. nov. from Palawan, Tirachoidea herberti spec. nov. from Borneo, Tirachoidea siamensis spec. nov. from Thailand and S-Vietnam and Phobaeticus chani Bragg spec. nov. from Borneo. Phobaeticus chani Bragg spec. nov. is the world’s longest known insect with a maximum body length of 357 mm and an overall length of 567 mm in the female. Twelve new synonymies were discovered: Bactridium grande Rehn, 1920 = Phobaeticus serratipes (Gray, 1835) syn. nov.; Pharnacia rigida Redtenbacher, 1908 = Phobaeticus sumatranus Brunner v. Wattenwyl, 1907, syn. nov.; Clitumnus irregularis Brunner v. Wattenwyl, 1907 = Phibalosoma tirachus Westwood, 1859, syn. nov.; Pharnacia magdiwang Lit & Eusebio, 2008 = Pharnacia ponderosa Stål, 1877 syn. nov.; Pharnacia spectabilis Redtenbacher, 1908 = Phibalosoma hypharpax Westwood, 1859, syn. nov.; Pharnacia semilunaris Redtenbacher, 1908 = Eucarcharus inversus Brunner v. Wattenwyl, 1907, syn. nov.; Pharnacia chiniensis Seow-Choen, 1998c = Pharnacia biceps Redtenbacher, 1908, syn. nov.; Nearchus grubaueri Redtenbacher, 1908 = Phobaeticus incertus Brunner v. Wattenwyl, 1907, syn. nov.; Phibalosoma maximum Bates, 1865 = Cladoxerus serratipes Gray, 1835, syn. nov.; Phobaeticus lambirica Seow-Choen, 1998a = Eucarcharus rex Günther, 1928, syn. nov.; Phobaeticus sichuanensis Cai & Liu, 1993 = Baculum album Chen & He, 1990, syn. nov. and Phobaeticus beccarianus Brunner v. Wattenwyl, 1907 is shown to represent the previously unknown female of Phobaeticus sobrinus Brunner v. Wattenwyl, 1907 (syn. nov.) Lectotypes are designated for: Nearchus redtenbacheri Dohrn, 1910, Pharnacia biceps Redtenbacher, 1908, Pharnacia ingens Redtenbacher, 1908, Pharnacia heros Redtenbacher, 1908, Phibalosoma westwoodi Wood-Mason, 1875, Phobaeticus sinetyi Brunner v. Wattenwyl, 1907, and Phobaeticus sumatranus Brunner v. Wattenwyl, 1907. A neotype is designated for Nearchus maximus Redtenbacher, 1908 and Phobaeticus magnus nom. nov. introduced as a replacement name for Nearchus maximus Redtenbacher, which is a junior homonym of Phibalosoma maximum Bates, 1865. The previously unknown males of Pharnacia heros Redtenbacher, 1908, Phobaeticus ingens (Redtenbacher, 1908), Tirachoidea jianfenglingensis (Bi, 1994), Pharnacia sumatrana (Brunner v. Wattenwyl, 1907), Phryganistria fruhstorferi (Brunner v. Wattenwyl, 1907) and Tirachoidea westwoodii (Wood-Mason, 1875) as well as the females of Pharnacia ponderosa Stål, 1877a and Pharnacia tirachus (Westwood, 1859) are described and illustrated for the first time. A brief description on the basis of colour photos of the so far unknown male of Pharnacia kalag Zompro, 2005 are presented. Detailed descriptions and illustrations are provided for the eggs of 24 species. The eggs of the following 18 species are described and illustrated for the first time: Pharnacia borneensis spec. nov., Pharnacia palawanica spec. nov., Pharnacia ponderosa Stål, 1877a, Pharnacia sumatrana (Brunner v. Wattenwyl, 1907), Pharnacia tirachus (Westwood, 1859), Phobaeticus hypharpax (Westwood, 1859), Phobaeticus chani Bragg spec. nov., Phobaeticus incertus Brunner v. Wattenwyl, 1907, Phobaeticus magnus nom. nov., Phobaeticus philippinicus (Hennemann & Conle, 1997a), Phobaeticus sinetyi Brunner v. Wattenwyl, 1907, Phryganistria grandis Rehn, 1906, Phryganistria virgea (Westwood, 1848), Tirachoidea biceps (Redtenbacher, 1908), Tirachoidea herberti spec. nov., Tirachoidea jianfenglingensis (Bi, 1994) and Tirachoidea siamensis spec. nov.. Several species were originally placed in or subsequently transferred into wrong genera by various authors. Consequently, numerous taxa are here transferred or re-transferred to other genera, which results in 22 new or revised combinations or status of genera and species (comb. nov. / stat. rev. / stat. nov.). A list of the taxonomic changes made in this revision is provided in the summary (see 9.2), which in all lists 70 nomenclatural changes.
Article
Ho, G.W.C. 2018. First report of the genus Micadina Redtenbacher (Phasmatodea: Diapheromeridae) from Vietnam, with description of a new species. Entomotaxonomia, 40(4): 253–256. (http://xbkcflxb.cnjournals.com/xbkcflxb/ch/reader/view_abstract.aspx?file_no=2018026&flag=1)
Article
Eggs of Phasmida are characterized by the presence of a micropylar plate system. The nature of this plate is discussed and the relevance of differences in plate structure to the taxonomy of the order is considered. A survey is made of the range of plate structure throughout the order, covering the external plate structure of 384 species and the internal plate structure of 179 species in forty of the forty-four subgroups of the order.
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An attempt is made to standardize further the descriptive terminology of the phasmid egg capsule by introducing stricter definitions and standard abbreviations. In addition, the various forms of the internal micropylar plate are categorized. Eophasma Sellick is replaced by Eophasmodes nov.n. A key to 131 generic forms of these eggs is provided. Where more than one egg form is associated with a genus, a diagnosis of the subgroups is provided.
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HO G.W.C., 2018a. -The genus Neohirasea Rehn (Phasmatodea: Diapheromeridae: Necrosciinae) from Vietnam. Zoological Systematics. 43(1): 37-51.
Phasmida Species File. Catalog of Stick and Leaf Insects of the world, 2nd Edition. The Insect Diversity Association and the Academy of Natural Sciences
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OTTE D. & BROCK P.D., 2005. -Phasmida Species File. Catalog of Stick and Leaf Insects of the world, 2nd Edition. The Insect Diversity Association and the Academy of Natural Sciences, Philadelphia, CafePress.com.