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On the Phyllium of Peninsular Malaysia and Sumatra, Indonesia, with range expansions for currently known species, description of the previously unknown Phyllium (Pu.) abdulfatahi Seow-Choen female, and description of the new species Phyllium (Ph.) rubrum n. sp. from Peninsular Malaysia (Phasmida: Phylliidae)

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Abstract. – The Phyllium diversity of Peninsular Malaysia and Sumatra, Indonesia is discussed with records of recent collection and from past literature. The previously unknown female Phyllium (Pulchriphyllium) abdulfatahi Seow-Choen, 2017a. is described along with a discussion of the range in which the species can be found and figures of the female. A range expansion into Sumatra and additional regions of Borneo for Phyllium (Pulchriphyllium) giganteum Hausleithner, 1984 is discussed along with figures of corresponding specimens. Phyllium (Phyllium) hausleithneri Brock, 1999 is also newly recorded from Sumatra and a discussion on intraspecies variation with corresponding figures of females and eggs. From Peninsular Malaysia, a new species Phyllium (Ph.) rubrum Cumming, Le Tirant & Teemsma n. sp. is described and illustrated from Perak, Malaysia. Phyllium (Pu.) mannani Seow-Choen, 2017a. and Phyllium (Pu.) rimiae Seow-Choen, 2017a. ranges are expanded to include Peninsular Malaysia, and Phyllium (Pu.) bioculatum Gray, 1832 is expanded to include Sumatra, with discussion on the intraspecies variation of males of all three species included. To conclude, with Phyllium (Ph.) rubrum Cumming, Le Tirant & Teemsma n. sp. described from a female, a key to females of all currently described species of Phyllium from Sumatra, Singapore, and West Malaysia is included.
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Faunitaxys
Revue de Faunistique, Taxonomie et Systématique
morphologique et moléculaire
Volume 6
Numéro 2 Février 2018 ISSN : 2269 - 6016
Dépôt légal : Février 2018
Faunitaxys
Revue de Faunistique, Taxonomie et Systématique
morphologique et moléculaire
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On the Phyllium of Peninsular Malaysia and Sumatra, Indonesia,
with range expansions for currently known species, description of
the previously unknown Phyllium (Pu.) abdulfatahi Seow-Choen
female, and description of the new species Phyllium (Ph.) rubrum
n. sp. from Peninsular Malaysia (Phasmida: Phylliidae)
ROYCE T. CUMMING (1), STEPHANE LE TIRANT (2) & SIERRA N. TEEMSMA (3)
(1) Entomology Department, San Diego Natural History Museum, POB 121390, Balboa Park, San Diego, California, United States,
92112-1390. Associate researcher for the Montréal Insectarium, Québec, Canada; H1X 2B2 - phylliidae.walkingleaf@gmail.com
- ZooBank : http://zoobank.org/6CA8501F-10BA-4E07-9BF4-65CFCE4E9E92
(2) Collection manager, Montréal Insectarium, 4581 rue Sherbrooke, Montréal, Québec, Canada, H1X 2B2 - sletirant@ville.montreal.qc.ca
- ZooBank : http://zoobank.org/A9391F8A-15D7-4D3B-9E3F-7123BA27EA2E
(3) San Diego, California, United States - sierranteemsma@gmail.com
- ZooBank : http://zoobank.org/1A671E2F-0C27-4750-B1BF-35341BC84644
Abstract.The Phyllium diversity of Peninsular Malaysia and Sumatra, Indonesia is discussed with records
of recent collection and from past literature. The previously unknown female Phyllium (Pulchriphyllium)
abdulfatahi Seow-Choen, 2017a. is described along with a discussion of the range in which the species can be
found and figures of the female. A range expansion into Sumatra and additional regions of Borneo for Phyllium
(Pulchriphyllium) giganteum Hausleithner, 1984 is discussed along with figures of corresponding specimens.
Phyllium (Phyllium) hausleithneri Brock, 1999 is also newly recorded from Sumatra and a discussion on
intraspecies variation with corresponding figures of females and eggs. From Peninsular Malaysia, a new species
Phyllium (Ph.) rubrum Cumming, Le Tirant & Teemsma n. sp. is described and illustrated from Perak,
Malaysia. Phyllium (Pu.) mannani Seow-Choen, 2017a. and Phyllium (Pu.) rimiae Seow-Choen, 2017a. ranges
are expanded to include Peninsular Malaysia, and Phyllium (Pu.) bioculatum Gray, 1832 is expanded to include
Sumatra, with discussion on the intraspecies variation of males of all three species included. To conclude, with
Phyllium (Ph.) rubrum Cumming, Le Tirant & Teemsma n. sp. described from a female, a key to females of all
currently described species of Phyllium from Sumatra, Singapore, and West Malaysia is included.
Cumming R. T., Le Tirant S. & Tee ms ma S. N. , 2018. – On the Phyllium of Peninsular Malaysia and Sumatra, Indonesia,
with range expansions for currently known species, description of the previously unknown Phyllium (Pu.) abdulfatahi
Seow-Choen female, and description of the new species Phyllium (Ph.) rubrum n. sp. from Peninsular Malaysia (Phasmida:
Phylliidae). Faunitaxys, 6(2) : 1 – 21.
Introduction
The large island of Sumatra, Indonesia has historically had
limited phylliid material recovered, and erroneously the few
collected specimens often were misidentified or described
under synonyms (Klante, 1976). To date, no Sumatran specific
work has been published focussing on the Phylliidae diversity
on the island, with all records generally being of single
specimens mentioned in larger works. When this analysis was
first begun it was assumed that there would likely be a new
species described from the island due to the general trend of
many phylliid species being rather island specific with smaller
ranges than what has historically been thought. However, after
series of specimens had been scoured over - males, females,
and eggs included, all materials examined turned out to be
previously known species from nearby localities. These records
are significant as several species ranges are here expanded to
include the island of Sumatra.
Four previously known species were confirmed from the island
of Sumatra based on more recent material, and when coupled
with historic data, help to clarify the ranges of these known
species. The species below discussed and recorded from the
Keywords :
1
Faunitaxys, 6(2), 2018 : 1 – 21.
island are Phyllium (Pulchriphyllium) abdulfatahi Seow-
Choen, 2017a.; Phyllium (Pulchriphyllium) giganteum
Hausleithner, 1984; Phyllium (Phyllium) hausleithneri Brock,
1999 and Phyllium (Pu.) bioculatum Gray, 1832. Of these, Ph.
(Pu.) giganteum has two additional range expansions recorded
from Sabah State, Borneo, Malaysia, and from West
Kalimantan Province, Indonesia. Phyllium (Pu.) abdulfatahi,
besides a range expansion into Sumatra, is also recorded from
Peninsular Malaysia, which the previously undescribed female
is also identified and described for the first time. Phyllium (Ph.)
hausleithneri, from the type locality of Peninsular Malaysia, is
also recorded from Sumatra with the same wide variation of
morphologies recorded.
A new species of leaf insect is also described from West
Malaysia under the name Phyllium (Ph.) rubrum Cumming, Le
Tirant & Teemsma n. sp. With many of the discussed species
coming from Peninsular Malaysia, the authors saw fit to help
fill the current knowledge gaps for this nearby region. With
many of the species from West Malaysia also occuring on
Sumatra, it would not be surprising if Phyllium (Ph.) rubrum
Cumming, Le Tirant & Teemsma n. sp., were also distributed
ZooBank : http://zoobank.org/3F3C9D87-47B8-4CE7-B912-3A3DA1E18E48
Phasmatodea ;
Phasmida ;
Phylliidae ;
Phylliini ;
Phyllium ;
Malaysia ;
Sumatra ;
Indonesia ;
abdulfatahi ;
rubrum ;
description ;
new species.
on the island, although for now it is only confirmed from
Peninsular Malaysia.
With this work expanding several of the currently known
distributions of several species found throughout Sumatra,
Peninsular Malaysia, and Borneo, two species, Phyllium (Pu.)
mannani Seow-Choen, 2017a. and Phyllium (Pu.) rimiae
Seow-Choen, 2017a. previously only known from Bornean
material are here recorded as also occuring in Peninsular
Malaysia. Although these two Bornean species are not currently
recorded from Sumatra or Singapore it would not be surprising
if they eventually were found there as well. Especially due to
the fact that Phyllium (Pu.) abdulfatahi Seow-Choen, 2017a.
and Phyllium (Pu.) giganteum Hausleithner, 1984 are now
recorded from Sumatra, Peninsular Malaysia, and Borneo.
To conclude several species noted from Sumatra in past works
are discussed but not positively identified due to a lack of
material available for observation, and with a new species
described from a female a key to known Phyllium females of
Sumatra, Singapore, and Peninsular Malaysia is included.
Materials and methods
– Measurements of all specimens were made to the nearest 0.1
mm using digital calipers. Morphological examinations were
done with a Leica ZOOM 2000 stereomicroscope. Photos for
figures 1A-D, 2D, 3A-D, 4A,C-E, 6A,C-F, 7A-E, 8A-I, 9A-D,
10A-E were taken at the San Diego Natural History Museum
using a Canon 5D Mark II and a MP-E65mm macro lens and
stacked using Zerene photo stacking software, version 1.04, 64-
bit.
Photos in figure 2A-C were taken by Johnson Chee Min of
Johor, Malaysia, and figure 6B was taken by Alexandre Banko
while visiting the Miyashita collection, Japan.
– Photo for figure 4B was taken by René Limoges of the
Montreal Insectarium using a Nikon D810 DSLR camera with
Nikon Micro-Nikkor 200 mm f/4 lens on Manfrotto 454
micrometric positioning sliding plate. Lighting was provided
by two Nikon SB-25 flash units with Cameron Digital diffusion
photo box. Adobe Photoshop Elements 13 was used as post
processing software.
Photos for figure 1E, 5A-D and 11 were taken by Natural
History Museum United Kingdom (NHMUK) personnel, and
were used under the license CC0-1.0 (NHMUK, 2017).
Eggs from Phyllium (Phyllium) hausleithneri Brock, 1999,
were provided by Bruno Kneubühler and were from a captive
culture originally from Tapah, Perak, Malaysia. Eggs of the
Sumatran population were removed from the abdomens of
females within the first author’s collection. Only eggs with
fully developed frills were observed, any eggs that appeared to
be underdeveloped were discarded.
– Egg orientation terminology follows that presented by Clark,
1978.
Treatment of species
Phyllium (Pulchriphyllium) abdulfatahi Seow-Choen, 2017a.
(Fig. 1A-E, 2A-D, 3A-D, 4A-E)
Type locality. – MALAYSIA, Borneo, Sabah State.
Range expansion. MALAYSIA: Perak and Penang States.
INDONESIA: Sumatra, West Sumatra Province.
Material examined [5 & 5 ]
MALAYSIA:
1 : Perak, Cameron Highlands, Tapah Hills: January, 2017 [Coll RC
17-249]
– 1 : Cameron Highlands, Tapah Hills: April, 2013 [Coll RC 16-039]
1 : Perak, Tapah Hills: February, 2012 [Coll RC 16-040]
1 : Perak, Tapah Hills: May, 2016 [Coll RC 16-222]
1 : Perak, Cameron Highlands: January, 2010 [Coll RC 16-155]
1 : Perak, Tapah Hills: June, 2014 [Coll RC 16-006]
1 & 1 : Perak, Tapah, March, 2014: from the collection of
Stephane Le Tirant
1 : Malaya. Penang, Sungei Pinang Hills, 30.vii.1958, H.T. Pagden
on Nephelium zappaceum. C.I.E.Coll. 16407, Started to lay eggs on
20.vii.58, ceased 24.Xii.58, died 27.xii.58, laid 141 eggs [Catalogue#
NHMUK012497235].
INDONESIA:
1 : Sumatra, West Sumatra Province, Sijunjung Regency, Tanjung
Gadang District, Batang Dikat, Telkom Tower “Bukit Rasam
Tap an gga ng ”: 10 th -1 2th M ay, 20 02 [ Fro m th e c oll ec tio n o f J err i
Larsson, United States, California].
Discussion. This species has for many years been
erroneously referred to as Phyllium bioculatum Gray, 1832,
Phyllium pulchrifolium Audinet-Serville, 1838, and has even
from time to time been in culture within the phasmid breeding
community and called Phyllium bioculatum “morph” or
Phyllium bioculatum “Tapah”. It was not until recently that the
species was described from a single male specimen from
Sabah, Borneo under the name Phyllium (Pulchriphyllium)
abdulfatahi Seow-Choen, 2017a. Seow-Choen did not however
recognize the species as the same one he illustrated and called
Phyllium pulchrifolium Audinet-Serville, 1838 in his book “An
Illustrated Guide to the Stick and Leaf Insects of Peninsular
Malaysia and Singapore” (Seow-Choen, 2000). In the 2000
book, Seow-Choen illustrated both sexes and the eggs of the
species but in the 2017 description of Ph. (Pu.) abdulfatahi
Seow-Choen left the female undescribed. With additional
material examined, the female Phyllium (Pu.) abdulfatahi is
below described and the species is recognized as having a
wider distribution than originally recorded with specimens
from both Peninsular (Perak and Penang) and Bornean
Malaysia (Sabah), and the Indonesian island of Sumatra.
Despite its expansive range, Phyllium (Pu.) abdulfatahi Seow-
Choen, 2017a. appears to be less frequently encountered than
congenerics from the same localities.
Description of previously unknown female
Coloration. – Variable and as observed from dried specimens,
the preservation technique is likely an influence. Overall
coloration green, with several different shades of green
recorded. Occasionally a colored form is found with reddish-
brown markings throughout the body, onto the legs, and even
notable on the vertex of the head (fig. 2C). On the more
commonly colored solid green individuals the antennae are of a
tan color with notable paler spots throughout (fig. 3A-C).
Morphology
Head. – Capsule slightly longer than wide, with a rather smooth
vertex with only fine granulation occasionally along the posterior
margin near the posteromedial tubercle which is broad, but not
prominent. Frontal convexity broad and stout, not exceeding the length
of the first antennomere, and sparsely covered in short, thin,
transparent setae. – Antennae with nine segments, occasionally it appears
as though there are ten segments due to antennomere VIII having a rather
prominent groove around the center, but this groove is superficial and
does not fully split the antennomere (fig. 3A-C). Antennomeres I-VIII
sparsely marked with small transparent setae, leaving the apical
CUMMING, LE TIRANT & TEEMSMA. – Phyllium of Sumatra
2
Faunitaxys, 6(2), 2018 : 1 – 21. 3
Fig. 1. - Phyllium (Pulchriphyllium) abdulfatahi Seow-Choen, 2017; females. - A: Dorsal full body [Coll RC 16-039]. - B: left profemora
[Coll RC 16-039]. - C: left profemora [Coll RC 17-249]. - D: left profemora [Coll RC 16-040]. - E: Dorsal full body (Natural History Museum
United Kingdom collection).
A
B C D
E
antennomere as the only segment covered in dense, stout, brown setae.
Apical antennomere narrower than VIII in which it sits, giving it a
dwarfed appearance (fig. 3A-C). – Compound eyes not exceptionally
large and rather tight against the head capsule, not widely protruding
(fig. 2D). Ocelli absent. Antennal fields wider than the first
antennomere but not exceptionally large.
Thorax. Pronotum with anterior margin only slightly concave and
lateral margins that are slightly convex converging to a narrow,
slightly rounded posterior margin that is only about than half the width
of the anterior rim (fig. 2D). The pronotum is rather smooth, with a
surface that only has fine granulation and a shallow pit in the center.
The pronotum anterior and lateral rims have only a slight rim,
posterior lacks a rim. Pro-, meso-, and metasternum smooth, with
only the slightest amount of fine granulation. – Mesopraescutum only
slightly wider than long, giving it a rectangular appearance, with
lateral margins only slightly converging to the posterior. – Lateral rims
with 11-14 irregularly shaped small granules, with no prominent
tubercles present, which gives the rims a rather smooth, straight
appearance. Mesopraescutum anterior rim smooth, lacking a
tubercle and only has a weak rim. – Mesopraescutum crest with 10-14
fine granules which are made less prominent due to the smooth surface
of the mesopraescutum that rises up to meet it, making the crest little
more than a slight ridge (fig. 2D). – Mesopleurae strongly diverging;
lateral margin with four to five minor tubercles, with one to two small
nodes between each minor tubercle. This small granulation gives the
lateral margins a rather smooth appearance (fig. 2D). Face of the
mesopleurae smooth, with only small granulation near the lateral
margins, and two faint divots, one on the anterior and one closer to the
center. – Tegmina long, extending to the anterior margin of abdominal
segment VIII.
Alae. – Rudimentary (fig. 2B).
Abdomen. – Abdominal segments II through the anterior half of IV
diverging, posterior half of IV through most of VII parallel to
subparallel. – Segment VIII has a single lobe which can be moderate to
strong and curves back towards segment IX. – Segments IX-X with
converging sides with the anal abdominal segment ending in a broad
rounded apex. Anal abdominal segment at its widest is wider than
long giving it a broad appearance. – Subgenital plate starting at the
anterior margin of segment VIII and only extending to the posterior of
segment IX. The subgenital plate is rather stout with a fine pointed tip.
Gonapophyses are exceptionally slender and short, only just slightly
protruding from under the subgenital plate leaving most of the ventral
aspect of the anal abdominal segment exposed (fig. 3D). Cerci short
and rounded, only slightly protruding from under the anal abdominal
segment. Cerci flat not cupped, with a lumpy surface with posterior
most rims that has a row of thin clear setae.
Legs. – Profemora exterior lobes very broad, at least twice the width
of the interior lobe, and with an acute exterior angle. – Posterior edge of
the exterior profemora lobe with three to four fine serrate teeth, lateral
edge is smooth and arcing to the anterior end of the profemora. – Interior
lobe of the profemora with an obtuse angle and marked with three blunted
teeth that are broad and relatively evenly spaced (fig 1B-D). Interior
lobe of mesofemora smoothly arcs from end to end and has nine to ten
small serrate teeth evenly distributed. – Exterior lobe of the mesofemora
arcs end to end but has a slight angle to it with three to four teeth only on
the distal half of the arc. Mesofemora interior and exterior lobes are of
approximately the same width. – Exterior lobe of metafemora thin and
lacking dentition, relatively straight. – Metafemora interior lobe thin near
the base and widening near the end into a rounded lobe that has six to
seven serrate teeth pointing distally, and more prominent on the distal end.
Protibiae exterior lobe narrow, at its widest, near the anterior, is about
the same width as the shaft of the protibiae itself. – Protibiae interior lobe
several times wider than exterior lobe, and shaped as an isosceles triangle
reaching the entire length of the protibiae. Mesotibiae exterior lobe a
rounded scalene triangle reaching end to end, with the wider side,
which is about the width of the shaft of the mesotibiae, located on the
distal end. – Metatibiae exterior lobe reaching end to end, in a large
rounded scalene triangle, with the widest side, which is at least twice
the width of the shaft of the metatibiae, on the distal end.
Measurements of females within the first author’s collection [mm]
length of body (including cerci and head, excluding antennae): 79.1-87.1
length/greatest width of head: 6.8/5.9-7.6/6.9
pronotum: 5.3-5.8
mesonotum: 4.8-5.5
length of tegmina: 46.5-52.6
greatest width of abdomen: 33.6-39.7
profemora: 19.3-19.8
mesofemora: 13.4-14.1
metafemora: 14.5-15.4
protibiae: 8.7-9.2
mesotibiae: 8.0-8.9
metatibiae: 11.6-13.6
antennae: 4.3-4.5.
Variation within sexes
Males. – Only two features were noted as variable within the
males observed, the size of the eyespot on abdominal segment
V and the size of the lobe of abdominal segment VIII. The
eyespots in Ph. (Pu.) abdulfatahi were noted as being relatively
variable ranging in size from 2.5-3.6 mm. The smaller eye spot
variations of Ph. (Pu.) abdulfatahi make them appear similar to
Ph. (Pu.) pulchrifolium (with eye spots that range in size from
1.1-2.4 mm wide), however several features allow quick
separation between the two species. Phyllium (Pu.)
pulchrifolium can be separated from Ph. (Pu.) abdulfatahi by
the lower margin of the exterior lobe of the profemora which is
always with a slightly arcing margin, not straight like Ph. (Pu.)
abdulfatahi. Additionally, the exterior lobe of the protibiae is
always wider than the shaft of the protibiae in Ph. (Pu.)
pulchrifolium and very thin (no more than the width of the
protibiae shaft itself) in Ph. (Pu.) adulfatahi.
The lobes of abdominal segment VIII were also shown to be
rather variable, ranging from small, only slightly protruding
back, (fig. 4D) to a larger lobe, reaching slightly further back
than the apex of the abdomen (fig. 4C). As with other variable
species, this species also has a wide range of intermediate lobe
sizes (fig. 4A, B, E). No stable feature could be detected from
males throughout the range when compared to the holotype Ph.
(Pu.) abdulfatahi and it is therefore determined that all three
locations, Peninsular Malaysia, Borneo, and Sumatra, belong to
the same moderately variable species.
Females. – Only females from Peninsular Malaysia could be
assessed as no known females have been recorded from Borneo
or Sumatra to date. Females observed appear to have a more
stable morphology than their male counterparts. This is a less
common trend of the phylliids as generally the females come in
a variety of forms (lobed, unlobed, intermediate) with males
having a typically more stable morphology.
The only morphological difference that could be noted was a
variation in the profemora. Females with more prominent
features have an exterior lobe of the profemora that can be
elongated, coupled with teeth of the interior lobe that are
generally more pronounced (fig. 1B-C). On the opposite end of
the spectrum, females with less pronounced features generally
tended to have narrower exterior lobes of the profemora and
less pronounced teeth of the interior lobe (fig. 1D).
Phyllium (Pulchriphyllium) giganteum Hausleithner, 1984
(Fig. 5A-D, 6A-F)
Type locality. “Malaysia, Cameron Highlands” (Hausleithner,
1984), presumed to be Perak State.
CUMMING, LE TIRANT & TEEMSMA. – Phyllium of Sumatra
4
Faunitaxys, 6(2), 2018 : 1 – 21. 5
Fig. 2. - Phyllium (Pulchriphyllium) abdulfatahi Seow-Choen, 2017; females. - A: Dorsal full body [Coll RC 17-249]. - B: With tegmina raised,
showing underdeveloped alae [Coll RC 17-249]. - C: Showing front end of live female [Coll RC 17-249]. - D: Head through thorax, dorsal view
[Coll RC 16-040]. Figures 2A-C taken by Johnson Chee Min of Johor, Malaysia of specimen 17-249 right after collecting, while still alive.
A B
C D
Known range. MALAYSIA: (Peninsular) Perak, Pahang, and
Selangor States; (Bornean) Sarawak State. INDONESIA: Sumatra,
Bengkulu Province.
Range expansion. MALAYSIA: (Borneo) Sabah State.
INDONESIA: (Sumatra) “Southern Sumatra”, and (Borneo) West
Kalimantan Province.
Material examined [16 , 8 ]
MALAYSIA [16 , 4 ]:
1 : Perak, Tapah Hills, August, 2014 [Coll RC 16-047]
– 4 : Perak, Tapah Hills, January, 2012 [Coll RC 16-041, 045, 043, 044]
1 : Perak, Tapah Hills, April, 2011 [Coll RC 16-042]
1 : Tapah Hills Reserve, March, 1987 [Coll RC 16-046]
10 : Perak, Tapah Hills, April, 1990, November, 1992, July, 2016
(Collection Stéphane Le Tirant)
1 : Borneo, Sabah, Pensiangan, [91-23] January 9th, 1991, leg.
Cornelio, from the collection of Jerri Larsson, California, United
States
1 : Borneo, Sarawak, Bintulu env., April, 2012 [Coll RC 16-048]
– 1 : Borneo, Sarawak, 1909, J.E.A. Lewis [Catalogue# NHMUK012500164].
INDONESIA [4 ]:
1 : “Southern Sumatra”, from the Miyashita collection
1 : Sumatra, Benkoelen Dist. 1912-1919, C.J. Brooks, 1920-43.
[Catalogue# NHMUK012500165]
2 : West Kalimantan Province, Mount Bawang, elevation 245
meters, January, 2016 [Coll RC 16-220, 16-219].
Known range discussion. – Besides the type locality of Perak
State, Malaysia, two other Peninsular Malaysia states are listed
as having Ph. (Pu.) giganteum records- Pahang: Kuala Tahan
and Selangor: Kuala Lumpur (Brock, 1994). The species is also
known to occur in Sarawak State on the Island of Borneo from
a record within the NHMUK (formerly the British Museum,
fig. 5A-B) and from a male within the first author’s collection
(Coll RC 16-048, fig. 6C). Although Brock, 1994 was not
certain of the Sumatran male he discussed on page 56 being Ph.
(Pu.) giganteum, examination of the specimen clearly confirms
the identity as Ph. (Pu.) giganteum (fig. 5C-D). It is worth
pointing out that the data on the historic label states
“Benkoelen Dist.” which is the historic name for Bengkulu
Province (Harfield, 1995).
Range expansion discussion. Two additional localities from
the island of Borneo are here added. Two males within the first
author’s collection [Coll RC 16-220, 16-219], both from the
Indonesian territory on the island, West Kalimantan Province.
Also a male from the collection of Jerri Larsson, California,
United States, from Borneo, Sabah State, Pensiangan Town,
adds the state of Sabah to the known distribution (fig. 6A).
Regarding Indonesia, besides the additional records from the
island of Borneo, Sumatra is again noted as having Ph. (Pu.)
giganteum, after the first record of Sumatra (Brock, 1994)
appears to have been overlooked by subsequent authors. A male
from the inexact locality of “Southern Sumatra”, Indonesia (fig.
6B) from the Miyashita collection, helps to confirm the
expansion of the range of Ph. (Pu.) giganteum onto the island
of Sumatra.
Variation within sexes
Females. – Widely variable in color and abdomen width.
Overall color of female specimens can range from a pale lime
green, to rich forest green, and in some individuals brown
patches are notable on legs or the tip of the abdomen,
occasionally covering most of the body. Abdomen width and
overall specimen size for all females examined had a wide
range, with overall length 106.7-117.0 mm and abdomen width
51.4-61.8 mm. It is also worth noting that the widest
individuals measured were not always also the largest, for
example the widest female measured was 110.6 mm long, a
very average length individual.
Males. – Male morphology, similar to female, was relatively
stable with only slight ranges of features exhibited. Due to the
wide range of female abdomen widths observed, it was no
surprise that the most variable feature of the males was also the
width of the abdomen. Specimens from all regions of the range
exhibited a spectrum of narrow (fig. 5A) to broad (fig. 6A)
abdomens with intermediate shapes observed (fig. 5C).
Phyllium (Phyllium) hausleithneri Brock, 1999
(Fig. 7A-E, 8A-I)
Type locality. MALAYSIA, Perak.
Range expansion. – INDONESIA: Sumatra, Bengkulu Province.
Material examined [18 , 12 , 62 eggs]:
MALAYSIA [6 , 3 , 35 eggs]
2 : Perak, Cameron Highlands, Tapah Hills: May, 2017 [Coll RC
17-253, 17-254]
1 : Pahang, Cameron Highlands District: June, 2010 [Coll RC
16-085]
1 : Perak, Tapah: June, 2014 [Coll RC 16-087]
2 : Cameron Highlands: May, 2011 [Coll RC 16-084, 16-169]
1 , 2 : Perak, Tapah Hills: April, 2012 [Coll RC 16-086, 16-088,
16-170]
10 Eggs: Origin Perak (from captive culture supplied by Bruno
Kneubühler) [Coll RC 18-002--011]
25 Eggs: Origin Perak (from captive culture supplied by Bruno
Kneubühler).
INDONESIA [12 , 9 , 27 eggs]:
5 : Sumatra Island, Bengkulu Province: January-March, 2017 [Coll
RC 17-064--068]
9 : Sumatra Island, Bengkulu Province: January-March, 2017 [Coll
RC 17-083--090, 17-107]
3 : Sumatra Island: February, 2017 [Coll RC 17-216--218]
4 : Sumatra Island: April, 2017 [Coll RC 17-211, 17-212, 17-213,
17-214]
4 Eggs: Removed from the abdomen of female 17-107 [17-377--380]
5 Eggs: Removed from the abdomen of female 17-089
10 Eggs: Removed from the abdomen of female 17-086
3 Eggs: Removed from the abdomen of female 17-087
5 Eggs: Removed from the abdomen of female 17-085.
Discussion. It was originally thought that the Sumatran records
of this species represented an undescribed species, native to the
island of Sumatra. This thought was fueled by the remarkable
color difference in the eggs as well as a first impression that the
center pattern of short pinnae on the lateral views of the eggs were
consistently different. Due to this first impression of the marked
difference between eggs, an extensive search was begun to
pinpoint morphological differences between adults.
The first impression was that the Sumatran population was smaller
than their Perak counterparts, this was dismissed when actual
measurements were taken to find a recordable difference, and it
was found that both populations had larger and smaller individuals
and a consistent range of sizes in between. Perak females ranged in
size from 74.6-82.8 mm and the Sumatran population ranged in
size from 70.3-79.3 mm, and males ranging in size from 55.8-57.8
mm for the Perak population and 51.4-56.1 mm for the Sumatran
population. Although on average the Sumatran population had
measurements that were slightly smaller than the Perak population,
CUMMING, LE TIRANT & TEEMSMA. – Phyllium of Sumatra
6
Faunitaxys, 6(2), 2018 : 1 – 21. 7
Fig. 3. - Phyllium (Pulchriphyllium) abdulfatahi Seow-Choen, 2017; females. - A: Antennae [Coll RC 16-039]. - B: Antennae [Coll RC
17-249]. - C: Antennae [Coll RC 16-040]. - D: Genitalia, ventral view [Coll RC 16-039].
A B C
D
extremes within each population overlapped significantly. With
this notable overlap in size range and a consistent spectrum of
sizes in each population, this immediate observation was
dismissed.
To adequately differentiate the two populations based on adults,
finer details were then observed. At every point during the search
both populations had individuals that would exhibit extremes of
several features along with the corresponding intermediates
showing mixed forms. Some of the features observed that were
thought to be significant, were later found not to be after a wider
sample of specimens was observed. These features included: the
granulation on the vertex of the head, ratio of width of interior lobe
of profemora versus the exterior lobe, female genitalia (shape and
length of subgenital plate and length of gonapophyses), length of
tegmina, and greatest width of abdomen versus length of abdomen.
Each time a possible unique feature was systematically eliminated,
more and more doubt was placed on their uniqueness.
The last feature to be looked at in fine detail was the female
antennae. Immediately it became clear that all females from the
Sumatran population had nine segmented antennae, a feature that
surprised us as Hennemann et al., 2009 states Ph. (Ph.)
hausleithneri has a ten segmented antennae. The original
description of Ph. (Ph.) hausleithneri however states the type
material as having a nine segmented antennae. After all Sumatran
specimens antennae were examined, the antennae of the Perak
population from the first author’s collections were examined. All
females examined clearly had nine antennae segments (except for
a single aberrant female within the first author’s collection, with
antennomeres VIII and IX fuzed giving it an eight segmented
antennae with a remarkably long terminal segment). This
observation was then sent to Frank Hennemann and Hennemann
reviewed all of his Ph. (Ph.) hausleithneri females, and found that
the illustration that was used in the 2009 work must have been
based on a single aberrant female in his collection with a ten
segmented antennae, as all other specimens in his collection
exhibited the common nine segmented antennae (pers. comm. Jan.
28th, 2018).
The final feature reviewed on the female specimens was the
number of teeth on the pars stridens of antennomere III, a feature
that has proven useful in species differentiation when larger
features seemed to be difficult to keep separated in morphological
keys. Größer in 2008 lists Ph. (Ph.) hausleithneri as having a pars
stridens with 44-46 teeth, and the following year Hennemann et
al., 2009 then listed it as having 44-48 teeth, a moderately wide
range for a species. The number of teeth observed on the Sumatran
populations pars stridens was again eye opening to the wide range
of morphologies of this species with a range of 37-44 teeth. With
this wide range observed from the Sumatran population, it is likely
that the Perak population also has a wider range than has
previously been observed. It is likely to be revised after a larger
sample size is surveyed.
It was thought that possibly the number of teeth of the pars stridens
could be linked to the form of the female (lobed, smooth, or
intermediate), but when compared with abdomen morphology no
clear pattern was present. As illustrated by two females with
extremely different abdominal morphologies (lobed and smooth
abdomens) had the same number of teeth on the pars stridens.
Other more moderate/intermediate forms showed a greater
variability with as few as 37 teeth observed and as many as 44
teeth (the lower threshold previously recorded on the Perak
population).
Eventually eggs were again looked at, under a wider spectrum.
This broader look at the eggs of both the Perak and Sumatran
populations revealed that eggs of the Perak population were not
soley marked with intricate short pinnae designs on the lateral
views. Instead eggs of the Perak population were widely variable,
with eggs that had simple straight markings to ones that were very
complex (fig. 8F-I). Figure 8F-I shows a range of designs, with the
most simple on the left, ranging to a more complex design on the
right which was originally thought to be the average egg for the
Perak population. This thought that the complexly marked eggs
were the most prevalent was what originally led us to assume this
population was a separate species.
When the eggs from the Sumatra population were reassessed and
the sample size was greatly increased, these eggs also showed a
variation in the lateral sort pinnae markings, albeit with a range
slightly less variable than that observed in the Perak population.
The most commonly encountered egg pinnae design was one with
a single marking down the center with relatively straight to wavy
sides, a marking very similar to part of the spectrum that the Perak
eggs, with their greater variety also showed frequently.
The only consistent feature observed between the two populations
from egg morphology, was the color of the eggs, with the Perak
population consistently darker than the Sumatran population.
There is a possibility that because the Sumatran eggs that were
observed in this study were removed from the abdomens of
various females, they could be considered not fully developed due
to this premature removal, and therefore their color could not have
been fully developed. Thanks to discussion with Bruno
Kneubühler, who has received freshly laid and therefore fully
developed eggs from a Sumatran population female, all of his eggs
also exhibited this paler color when compared with the Perak
population.
Another possibility that has not been widely studied as of yet is the
variation of egg color not only within populations but also within a
single females egg color variability. Phyllium (Phyllium)
westwoodii Wood-Mason, 1875 has been recorded as having a
predictable egg color (white or brown) based on if the females
eggs were the product of thelytoky or if they were fertilized (Arai
and Yago, 2015). To date this is the only species to have had a
systematic study done on egg color coupled with egg production
(unfortunately done without replicates however) and it would be
interesting to see the results of a similar study if done with Ph.
(Ph.) hausleithneri. This possibility leaves the authors with the
belief that these two Ph. (Ph.) hausleithneri populations lack any
morphological features that would consistently seperate them to
the grounds of naming them as separate species.
It is likely that future genetic work will help to clarify the
relationship between these Perak and Sumatran populations, but
for now, due to a lack of morphological differences observed
between series of males, females, and eggs, these two populations
are considered the same species.
Variation within sexes
Females. Females of both populations expressed a wide range of
forms with many of their features quite variable. Overall body
length varied with a range of 8-9 mm difference between the
smallest and largest individuals of both populations. Greatest
abdominal width showed moderate variation within both
populations, however this appears to be more closely linked to
overall individual size as larger individuals also frequently had
wider abdomens when compared to their smaller counterparts. The
length of tegmina was only slightly variable within both
populations, either at the anterior edge of abdominal segment VIII
or at most reaching half way into VIII. Considerable variation was
observed in the shape of the abdominal segments, ranging from
smooth continuously converging sides to subparallel sides ending
in abdominal segments VII and VIII widely lobed (fig. 7A-D). The
size of the teeth of the profemora interior lobe was more or less
CUMMING, LE TIRANT & TEEMSMA. – Phyllium of Sumatra
8
Faunitaxys, 6(2), 2018 : 1 – 21. 9
Fig. 4. - Phyllium ( Pulchriphyllium) abdulfatahi S eow- Cho en, 201 7; mal es, dor sal views . - A: Peninsular Malaysia [Coll RC 16-155]. - B: Peninsular
Malaysia, coll. Stephane Le Tirant, photo credit René Limoges. - C: Sumatran record from the collection of Jerri Larsson. - D: Peninsular Malaysia,
reduced lobes, posterior closeup [Coll RC 16-222]. - E: Peninsular Malaysia, moderate lobes, posterior closeup [Coll RC 16-006].
A B C
D E
linked to the general abdominal shape. Individuals with more
pronounced abdominal lobes had larger teeth of the profemora and
those with smaller lobes had smaller finer pointed teeth.
Both populations had the characteristic purple colored coxae and
thorax with the same placement and size range of nodes with
minimal variation. The variation in the number of teeth of the pars
stridens is already discussed above, and with a larger sample size
the variation is expected to be even greater as our small sample
size gave no indication of the average number to expect.
Males. – Unlike females, males of both populations showed very
little variety in morphological features. All features of the
antennae, thorax, legs (including teeth and lobes of legs), and
tegmina were stable features between both populations. The only
detectable variations within both populations were the length of
alae and the maximum width of abdomen. Alae of both
populations ranged from the shortest, reaching the anterior of
abdominal segment IX, to the longest just passed the posterior of
segment IX, with intermediate lengths from both populations
present. Variation of the greatest width of the abdomen was also
notable, although not as extreme as the abdominal variation
expressed within the females. With females expressing a wide
range of abdominal characteristics it is no surprise that males
showed a slight variation as well.
Eggs. Eggs had a steady size range and the arrangement/ length
of the marginal pinnae was consistent. Color was variable, and
from what was assessed it appears as though the color is steady
between the Perak (dark brown) and Sumatra (cream) populations.
Length of the micropylar plate was slightly variable within both
populations showing both longer (fig. 8D) and shorter (fig. 8E)
variations. Inner design of the short pinnae on the lateral surfaces
were more variable in the Perak population (fig. 8F-I), however
the Sumatran also showed some variation, just not as widely as the
Perak population.
Phyllium (Phyllium) rubrum Cumming, Le Tirant &
Teemsma n. sp.
(Fig. 9A-D, 10D-E)
ZooBank : http://zoobank.org/ED5306F6-B1F1-4D46-
AC80-04088F92D764
Holotype, : MALAYSIA, Perak, Tapah Hills, April, 2017 [Coll. RC
17-204]: Deposited in the Montreal Insectarium type collection.
Paratypes:
1 : “Malaysia, Tapah Hills”; bred by staff of the Long Island
Butterfly Exhibit staff, January-March, 2017 [Coll. RC 17-005]:
retained within the collection of the first author.
– 10 Eggs: “Malaysia, Tapah Hills” (from captive culture supplied by
Bruno Kneubühler) [Coll RC 17-339, 17-342, 17-374, 18-012-018]:
retained within the collection of the first author.
Discussion and differentiation. – This species has occasionally
been in culture throughout Europe for a number of years and
has been referred to as Phyllium “red coxae” or Phyllium
“Tapah Hills, red coxae” because of the characteristic red color
of the meso- and metacoxae. This species however seems to be
much less encountered than sympatric species in the same type
locality, with Ph. (Ph.) hausleithneri Brock, 1999 frequently
offered for sale and Ph. (Ph.) rubrum n. sp. rarely offered. The
lack of exterior lobes and the rudimentary alae place this new
species within the siccifolium species-group within the
(Phyllium) subgenus as per Hennemann et al., 2009.
It was originally planned to wait until a male specimen could
be obtained to describe the species, however, due to the lack of
an official name within the phasmid breeding community, a
description of at least the female was warranted to help clear up
current confusion and it is unlikely that a male will be obtained
in the near future due to the species apparent rarity.
The female appears most closely related to Ph. (Ph.)
hausleithneri Brock, 1999, Ph. (Ph.) jacobsoni Rehn and Rehn,
1933, and Ph. (Ph.) palawanense Größer, 2001, as observed by
the abdominal shape and structure of the profemoral lobes and
teeth. From all these species it can be distinguished by its larger
size (~90.0 mm) and the orange-red color of the meso- and
metacoxae. From Ph. (Ph.) palawanense it can more
specifically be distinguished by the singular line of nodes
running along the sagittal plane of the mesopraescutum
(mesopraescutum irregularly granulose in Ph. (Ph. )
palawanense). From Ph. (Ph.) hausleithneri and Ph. (Ph.)
jacobsoni it can further be distinguished by the lateral margins
of the mesopraescutum which have nodes of streadily
increasing size from the anterior to the posterior (in Ph. (Ph.)
hausleithneri and Ph. (Ph.) jacobsoni these nodes are of a
uniform maximum height with the largest on the anterior of a
similar size as those on the posterior).
Males, although not available for this description, from photos
of captive bred stock also have meso- and metacoxae that are
clearly marked with an orange-red color. This clearly colored
coxae separates them from other similar species, Ph. (Ph.)
hausleithneri Brock, 1999, Ph. (Ph.) jacobsoni Rehn and Rehn,
1933, Ph. (Ph.) bradleri Seow-Choen, 2017a., and Ph. (Ph.)
chenqiae Seow-Choen, 2017a. Although Seow-Choen did not
describe the color of the coxae of the two Bornean species that
appear closely related, 6 male Ph. (Ph.) chenqiae Seow-Choen,
2017a. and 23 Ph. (Ph.) bradleri Seow-Choen, 2017a. from
within the first and second authors collections were examined
and all lacked colored coxae. This lack of colored coxae makes
Ph. (Ph.) chenqiae and Ph. (Ph.) bradleri appear more closely
related to Ph. (Ph.) jacobsoni, a congeneric which Seow-Choen
failed to differentiate both species from.
Eggs, morphologically are less similar to Ph. (Ph.)
hausleithneri and are instead more similar in their pinnae
pattern on the lateral view to those of Ph. (Ph.) philippinicum
Hennemann, et al., 2009. Instead of a pinnae pattern on the
lateral view with straight marginal rows and a center row that
varied in pattern from straight sided to intricate, Ph. (Ph.)
rubrum n. sp., has these three rows uniformly marked by
circular bare impressions. All eggs observed of Ph. (Ph.)
rubrum n. sp., showed a stable morphology, very different from
the widely variable eggs of Ph. (Ph.) hausleithneri, which
ranged from straight to intricate irregular shapes. This pattern
of circular bare impressions is a feature that is more similarly
expressed in Ph. (Ph.) philippinicum eggs. Phyllium (Ph.)
rubrum n. sp., eggs however can be differentiated from Ph.
(Ph.) philippinicum eggs by the micropylar plate which is
narrow and covers approximately 5/6th of the dorsal view (in
Ph. (Ph.) philippinicum the micropylar plate only covered
around 2/3rds of the dorsal view and is wider with each side
from the micropylar cup tapering evenly).
Description of female
Coloration. Overall coloration green, with orange antennae
and highlights of the protibiae and profemora. Venation of the
tegmina also with an orange color with a notable red marking
near the branching point of the media anterior from the media
vein (Ragge, 1955). Meso- and metacoxae with red-orange
coloration.
Morphology
Head. – Capsule slightly longer than wide, with a granulose vertex
with 27-30 nodes relatively evenly spaced and of slightly varying sizes
on each side of the sagittal plane. – Posteromedial tubercle is broad,
CUMMING, LE TIRANT & TEEMSMA. – Phyllium of Sumatra
10
Faunitaxys, 6(2), 2018 : 1 – 21. 11
Fig. 5. - Phyllium (Pulchriphyllium) giganteum Hausleithner, 1984, historic males from the Natural History Museum United Kingdom
collection. - A: Record from Sarawak, Borneo, 1909, J.E.A Lewis [Catalogue# NHMUK012500164]. - B: Original data labels for Sarawak
Borneo male. - C: Record from Benkulu Province, Sumatra, 1912-1919, C.J. Brooks [Catalogue# NHMUK012500165]. - D: Original data
label for Sumatra male. Photo credits NHMUK, used under the license CC0-1.0.
A
B
C
D
but not prominent. – Frontal convexity stout, not exceeding the length
of the first antennomere, and with 13-18 tan setae throughout. – Antennae
consisting of nine segments, with I-VII sparsely marked with tan setae of
varying lengths, and segments VIII-IX covered in dense tan setae of
relative uniform length (fig. 9C). – Compound eyes moderately protruding
from head capsule (fig. 9C) and ocelli are absent. – Antennal field wider
than the first antennomere but about as long as the first antennomere is
tall.
Thorax. Pronotum with anterior margin slightly concave with a
distinct rim; lateral margins with moderate rims and relatively straight
sides that are prominently converging to the stout posterior margin
which lacks a rim and that is only about half the length of the anterior
margin (fig. 9C). The pronotum surface is irregularly marked with small
granules and slight pits with the only notable feature being the
prominent pit in the center. – Mesopraescutum slightly wider than long,
with lateral margins running parallel to the posterior. – Lateral rims with
7-8 irregularly shaped nodes, which steadily increase in size from the
anterior to the posterior, with the posterior most being rather prominent
tubercles of a similar size to those fund on the mesopraescutum crest or
the mesopraescutum anterior rim. Mesopraescutum anterior rim
prominent with a lumpy/granulose surface, marked by a prominent
tubercle on the sagittal plane. – Mesopraescutum crest with five evenly
sized turbercles, which are prominent due to the surface of the
mesopraescutum being rather flat with only very small granulation (fig.
9C). Mesopleurae uniformly diverging; lateral margin marked with 14
to 17 nodes of relatively even size and shape all pointed slightly to the
posterior. Face of the mesopleurae mostly smooth, with only small
granulation near the nodes of the lateral margins, and two distinct pits,
one near the anterior and one closer to the center. – Teg min a, nearly
extending to the posterior margin of abdominal segment VII.
Alae. – Rudimentary.
Abdomen. – Abdominal segments II through the anterior twothirds
of IV gradually diverging, posterior third of IV through VII gradually
converging, with segments VIII-X converging more strongly. – Apex
of abdomen appears relatively slender and long likely due to the long
cerci protruding from under the anal abdominal segment which itself is
not particularly broad. Subgenital plate starting at the anterior
margin of segment VIII and extending halfway into segment X. The
subgenital plate is rather broad but near the apex converges to a fine
pointed tip. Gonapophyses long, reaching the apex of the anal
abdominal segment and rather broad, the same width as the cerci
which are flat and with irregular, lumpy surfaces but no prominent
granulation (fig. 9D).
Legs. – Profemora exterior lobe slightly narrower than the interior
lobe, and smoothly arcing end to end. Interior lobe of the profemora
with an angle that is slightly obtuse and marked with three to four
smoothly serrate teeth that are unevenly spaced with notable looping
gaps between each tooth. – Interior lobe of mesofemora narrow, only
as wide as the mesofemora shaft and arcs from end to end. Interior
lobe of mesofemora marked with five small serrate teeth on the distal
half. – Exterior lobe of the mesofemora also only about as wide as the
mesofemora shaft and reaches end to end with a slight angle, marked only
by a single small serrate tooth on the distal third. Exterior lobe of
metafemora thin, lacks dentition, and follows the slight curve of the
metafemora shaft. – Metafemora interior lobe with seven serrate teeth on
the distal half, and only slightly wider than the exterior lobe. – Protibiae
with no exterior lobe, and an interior lobe spanning the length of the
protibiae in a gentle arcing triangle. Meso- and metatibiae simple,
lacking interior and exterior lobes.
Measurements of holotype [mm]
length of body (including cerci and head, excluding antennae): 90.5
length/greatest width of head: 7.1/6.4
pronotum: 5.2
mesonotum: 7.7
length of tegmina: 53.6
greatest width of abdomen: 33.2
profemora: 16.5
mesofemora: 14.5
metafemora: 19.6
protibiae: 10.1
mesotibiae: 9.8
metatibiae: 16.2
antennae: 4.1.
Description of egg (Figs. 10D-E). – The lateral surfaces are
flattened and the dorsal surface slightly convex, which gives
the egg a general kidney shape, which is strengthened by the
long pinnae on the dorsal surface margins. When viewed from
the lateral aspect the anterior width of the capsule is of a
similar size to that of the posterior, with the center width only
slightly wider. When viewed from the lateral view, longitudinal
carinae along the dorsal and posterior margins are each set with
a row of long pinnae which each ends in at least a single split,
occasionally with two larger branches and several smaller ones.
Lateral surfaces with three relatively parallel rows of circular
bare impressions, with the spaces between densely covered
with short, singly branched pinnae. Micropylar plate reaching
almost end to end on the dorsal surface (approximately 5/6th the
length) slightly more situated towards the anterior margin.
Shape of micropylar plate thin and long, only notably wider
around the micropylar cup. Micropylar cup of moderate size
and placed just off of center, on the posterior end of the
midline. Operculum ovular, with the outer margin with a row of
pinnae similar to those along the longitudinal outer edges of the
capsule but slightly longer. Overall color brown with little
variation.
Etymology. – Due to the trend within the phasmid breeding
community to refer to this then undescribed species as Phyllium
“red coxae” the namerubrum was chosen referring to this
unique color.
Phyllium (Pulchriphyllium) rimiae Seow-Choen, 2017a.
(Fig. 10A-C)
Type locality. MALAYSIA, Borneo, Sabah State.
Range expansion. MALAYSIA: “Cameron Highlands” assumed
to be Perak State.
Material examined [5 ]
MALAYSIA:
1 : Sabah, North Borneo, Mt. Trus Madi, [Coll RC 17-281]
1 : Malaysia, Cameron Highlands, June, 2016 [Coll RC 17-223]
1 : Sabah, North Borneo, Mt. Trus Madi, May 14th, 2006 [Coll RC
17-282]
1 : Crocker Range, Sabah, Nord Borneo, Est Malaisie, 15.I.2004
[Within the collection of the second author]
1 : Sabah, North Borneo, Mt. Trus Madi [Within the collection of
the second author].
Variation
Males. Males show variation with robust and slender
individuals with clear intermediate forms between. The size of
the lobes of the abdominal segments VII and VIII varied with
more slender individuals having less pronounced lobes and
robust indivuduals with clear notable lobes. Maximum
abdomen width was somewhat positively linked to the size of
the eyespots in abdominal segment V with abdominal widths
ranging from 17.9-22.1 mm and eye spots ranging from 1.6-3.2
mm. The serration of the exterior lobe of the profemora was
also variable and linked to maximum abdominal width, with
wider, more robust specimens having stronger serration of the
exterior lobe.
The Peninsular Malaysian male examined fell within the range
of the intermediate specimens from Borneo and although
examined in detail, no consistent morphological features could
be identified between the two populations.
CUMMING, LE TIRANT & TEEMSMA. – Phyllium of Sumatra
12
Faunitaxys, 6(2), 2018 : 1 – 21. 13
Fig. 6. - Phyllium (Pulchriphyllium) giganteum Hausleithner, 1984; A-C males, D-F females, dorsal views. - A: Sabah, Borneo record,
Collection of Jerri Larsson. - B: Sumatran record from the Miyashita collection, Japan. - C: Sarawak, Borneo, [Coll RC 16-048]. - D:
Malaysia, Perak, broad abdominal variation [Coll RC 16-042]. - E: Malaysia, Perak, [Coll RC 16-044]. - F: Malaysia, Perak, slender
abdominal variation [Coll RC 16-043].
A B C
D E F
Fig. 7. - Phyllium (Phyllium) hausleithneri Brock, 1999; females, dorsal views. - A: Slender, smooth abdomen [Coll RC 17-083]. - B: Slender,
slightly lobed abdomen [Coll RC 17-085]. - C: Broad, moderately lobed abdomen [Coll RC 17-216]. - D: Broad, strongly lobed abdomen
[Coll RC 17-084]. - E: Left antennae, showing nine segments, dorsal view, from female [Coll RC 17-085].
A B
C D
E
2 : N. Borneo, Sabah, Keningau Dist. Trus Madi Mt., 1,160 meters,
8.XI.2015, A. Klimenko leg. [Coll RC 16-112, 16-131]
1 : N. Borneo, Sabah, Keningau Dist. Trus Madi Mt., 1,160 meters,
23.VI.2014, A. Klimenko leg [Coll RC 16-108]
1 : Borneo, Sabah, Crocker Range, Gunung Alab, 1,700 meters,
3.VI.2014, A. Klimenko leg [Coll RC 16-132]
1 : N. Borneo, Sabah, Keningau Dist. Trus Madi Mt., 1,160 meters,
24.VIII.2012, A. Abramov leg [Coll RC 16-210]
1 : NE Borneo, Sabah, Tambunan reg., Trus Madi Mts. 1,200
meters, Jan. 12-24. 2007, leg. A. Sochivko
1 : Cameron Highlands, June, 2015 [Coll RC 16-231]
1 : Cameron Highlands, June, 2016 [Coll RC 17-219].
Phyllium (Pulchriphyllium) mannani Seow-Choen, 2017a.
Type locality. MALAYSIA, Borneo, Sabah State.
Range expansion. MALAYSIA: “Cameron Highlands” assumed
to be Perak State.
Material examined [12 ]
MALAYSIA:
4 : NE Borneo, Sabah, Tambunan reg., Trus Madi Mts. 1,200
meters, May 13-26, 2007, leg. A Sochivko [Coll RC 16-106,
16-133-134, 16-136]
CUMMING, LE TIRANT & TEEMSMA. – Phyllium of Sumatra
14
Fig. 8. - Phyllium (Phyllium) hausleithneri Brock, 1999; eggs. - A: Lateral, Tapah, Malaysia [Coll RC 18-003]. - B: Lateral, Tapah, Malaysia
[Coll RC 18-004]. - C: Lateral, Sumatra [Coll RC 17-380]. - D: Dorsal, Tapah, Malaysia [Coll RC 18-003]. - E: Dorsal, Sumatra [Coll RC
17-377]. - F-I: Lateral close up views of the variable pattern in the center, Tapah, Malaysia [Coll RC 18-002--18-005].
A B C
D E
F G H I
Faunitaxys, 6(2), 2018 : 1 – 21. 15
Variation
Males. – Males show moderate variation, mostly notable in the
abdomen, with the legs and thorax with a rather uniform
morphology. Abdominal width ranged from 23.0-27.3 mm,
with those with wider abdomens having slightly more
pronounced rounded edges but not by a notable amount. Size of
eye spots on abdominal segment V ranged from 1.1 mm to 2.5
mm, with centers that could be brown or slightly transparent.
Size of eye spots on abdominal segment V did not necessarily
correspond to the width of the abdomen, as moderate width
individuals had a wide range of eye spot sizes. Another feature
that was only weakly variable was the size of the small
serration of the exterior lobe of the profemora, which ranged
from almost not noticeable to small finely pointed teeth. This
feature did not appear to be linked to the robustness of the body
as all moderately size individuals expressed this feature
differently.
The two males examined from Peninsular Malaysia showed an
average range of features with all variable features clearly
falling within the range of those noted from the Bornean
population. With Ph. (Pu.) mannani now also known from
Peninsular Malaysia, it will be interesting to see if Sumatra has
Ph. (Pu.) mannani as well, or has the recently described species
Phyllium (Pu.) shurei Cumming and Le Tirant, 2018, or a yet to
be described adelphotaxon.
Phyllium (Pulchriphyllium) bioculatum Gray, 1832
(Fig. 11)
Material examined [6 ]
MALAYSIA:
2 : Perak, Cameron Highlands, March, 2012 [Coll RC 16-001,002]
1 : Cameron Highlands, June, 2010 [Coll RC 16-149]
1 : Cameron Highlands, August, 2016 [Coll RC 17-224]
1 : Cameron Highlands, April, 2016 [Coll RC 17-225].
INDONESIA:
1 : Sumatra: Lebong Tandai. 14.VII.1921. C.J.Brooks. B.M.
1936-681 [Catalogue# NHMUK012497233].
Variation
Males. Males exhibit moderate variation in the abdomen
(general shape and size of eye spots on abdominal segment V)
and lobes of the pro-, meso-, and metafemora. As expected
from these moderately variable features, not only were both
extremes observed, but also intermediate forms from
throughout the spectrum. Other morphological features were
less variable between individuals, even between individuals
with variable features which were expressed on opposite ends
of the spectrum.
Maximum abdominal width varied little, ranging from
24.1-25.3 mm, instead the variation was observed in the lateral
margins of segments V-VII ranging from distinctly parallel to
subparallel. This feature is only worth mentioning because of
the size of the abdomen and this slight variation quickly
catches the eyes attention, despite the only slight variation that
is measurable. Size of eye spots on abdominal segment V were
more distinct however, and ranged from 1.9 mm to 3.9 mm, all
with coloring that was brown with a slightly transparent center,
which varied in size depending on the overall eye spot
diameter. The pro-, meso-, and metafemora exhibited some of
the most drastic variation of morphologies, with the size of
serration present on the lobes consistently larger or consistently
smaller between the various lobes of individuals. As with the
other variable features, this range in serration size was also
noted to have intermediate individuals with serration of
medium size.
The male examined from Sumatra clearly fell within the
intraspecies variation of the specimens examined from
Peninsular Malaysia and due to the proximity of the two
localities it is not surprising that Ph. (Pu.) bioculatum is also
found on Sumatra. As a species which has shown a moderate
range of morphologies, even within only relatively small
sample sizes, it must be stressed that future authors should
examine series of specimens whenever possible to better
understand intraspecies variation to avoid the naming of
synonyms based solely on common, easily detectable
variations.
Other likely species found on Sumatra as noted from past
authors
Klante, 1976
The female specimen figured by Klante in 1976 (page 75, fig.
11) appears likely to be Ph. (Ph.) hausleithneri but without
seeing the specimen in person this observation cannot be
confirmed. Hennemann et al. 2009 believe this was a separate
species from the limited morphology shown in Klante’s
illustration. However, when a wide range of Ph. (Ph.)
hausleithneri are observed the features shown on the
illustration do not seem out of the range of common features
exhibited in the moderately lobed forms of the population.
Without seeing the specimen in person it is impossible to
positively identify the species, however based on observations
of a wide range of individuals, the specimen illustrated in
Klante is likely the Sumatran population of Ph. (Ph.)
hausleithneri.
Klante also illustrates a female that appears to be a member of
the celebicum species-group (page 75, fig. 5) which he lists as
Phyllium (Ph.) westwoodii Wood-Mason, 1875. This female
instead likely represents a female Phyllium (Ph.) chrisangi
Seow-Choen, 2017b., which at present is confirmed as
occurring in the country of Singapore, a short distance from
Sumatra. Without fresh material from the celebicum species-
group from Sumatra, this identification and presence cannot be
confirmed.
Lucas, 1857
In 1857 Lucas described the species Phyllium magdelainei
Lucas, 1857 which he appears to have named in honor of a Mr.
Magdelaine, marine officer on the ship Duroc. Despite the
ship’s unfortunate sinking, Magdelaine still managed to bring
back Sumatran specimens for Lucas to study and attempt to
rear (Lucas, 1857). This species, was later synomized with
Phyllium (Pulchriphyllium) pulchrifolium by Klante in 1976, a
synonym which has been recognized since. Examination of a
photo of the holotype of Phyllium magdelainei (in Brock et al,
2018) confirms this as a synonym of Phyllium (Pu.)
pulchrifolium [type locality Java, just to the south of Sumatra.]
The synonymy is likely to remain valid, although it would be
useful to examine a series from Sumatra.
Males of Phyllium (Pulchriphyllium) pulchrifolium from Java
show a moderate variation in morphology so it is likely that
Sumatran males will also have a range of morphologies.
Several features that have been noted by the authors as variable
including, overall body length, size of the eyespots on
abdominal segment V, and the size of the lobes of abdominal
segment VIII but only moderately. All other features appear to
be relatively more stable in males.
CUMMING, LE TIRANT & TEEMSMA. – Phyllium of Sumatra
16
Faunitaxys, 6(2), 2018 : 1 – 21. 17
Fig. 9. - Holotype female Phyllium (Phyllium) rubrum Cumming, Le Tirant & Teemsma n. sp. [Coll RC 17-204]. - A: Dorsal view. - B:
Ventral view. - C: Close up of antennae, head, and thorax. - D: Close up of genitalia, ventral view.
A B
C D
Key to known female Phyllium of Peninsular Malaysia, Singapore, and Sumatra.
Localities are given with the following abbreviations following the species and author information: Peninsular Malaysia (PM); Singapore
(SI); Sumatra (SU).
1. Meso- and metatibiae with exterior lobes: [subgenus (Pulchriphyllium)] ...................................................................................... 2.
– Meso- and metatibiae without exterior lobes: [subgenus (Phyllium)] ............................................................................................. 5.
2. Large species (106.7-117.0 mm); gonapophyses long, reaching very near to or to the apex of the anal abdominal segment .........
........................................................................................................................................ Ph. (Pu.) giganteum Hausleithner, 1984 (PM, SU)
- Medium sized species (76.1-89.8 mm); gonapophyses very short, only slightly protruding from under the anal abdominal segment,
leaving most of the ventral aspect of the segment exposed ................................................................................................................ 3.
3. Abdominal segment VIII lobed, not uniformly converging, giving the abdomen a broad rectangular appearance ....................... 4.
– Abdominal segment VIII uniformly converging, not lobed or broad, giving the abdomen a spade shaped appearance ..................
............................................................................................................................................. Ph. (Pu.) bioculatum Gray, 1832 (PM, SI, SU)
4. Posterior margin of exterior lobe of the profemora straight, with only slight serration at most; protibiae exterior lobe thin, generally
thinner than width or at most the same width as the shaft of the protibiae ..................... Ph. (Pu.) abdulfatahi Seow-Choen, 2017a. (PM, SU)
Posterior margin of exterior lobe of the profemora not straight, curved with obvious serration on the margin; protibiae exterior
lobe wide, always wider than the width of the protibiae shaft ................................... Ph. (Pu.) pulchrifolium Audinet-Serville, 1838 (SU)
5. Alae developed: [celebicum species-group] .................................................................... Ph. (Ph.) chrisangi Seow-Choen, 2017b. (SI)
– Alae rudimentary: [siccifolium species-group] ............................................................................................................................... 6.
6. Meso- and metacoxae purple in color; mesopraescutum lateral margins with nodes that are of an even size from the anterior to the
posterior ......................................................................................................................... Ph. (Ph.) hausleithneri Brock, 1999 (PM, SI, SU)
Meso and metacoxae orange to red in color; mesopraescutum lateral margins with nodes that steadily increase in size from the
anterior to the posterior .............................................................................. Ph. (Ph.) rubrum Cumming, Le Tirant & Teemsma n. sp. (PM)
Rehn and Rehn, 1933
The female specimen illustrated in Rehn and Rehn, 1933 (plate
16, fig. 1) is stated as coming from Nias island a small island
off the northwest coast of Sumatra and was listed as Phyllium
(Pu.) pulchrifolium. This female morphologically appears more
similar to Phyllium (Pu.) agathyrsus Gray, 1843 but with
tegmina that are longer than average Phyllium (Pu.) agathyrsus
coming from Sri Lanka. Perhaps fresh material from Nias
island will clarify this.
Conclusional notes on species distribution in Sumatra
With the above species discussed and several distribution
ranges expanded, a clearer picture of the biogeographical
distribution of the phylliids of Sumatra is beginning to form.
The presense of several Peninsular Malaysian species in
Sumatra is not surprising due to the islands close proximity to
the peninsula. The further distribution onto the large island of
Borneo is more surprising however, because of the greater
distance of the islands. It is our hopes that a greater
understanding of the phylliid diversity on Sumatra will prompt
others to conduct additional research into the islands elusive
species. Hopefully future genetic analysis will help reveal the
underlying relationship of these widely distributed species. It
should be noted that throughout this work, and within the
following key, Ph. (Pu.) bioculatum Gray, 1832, Ph. (Pu.)
pulchrifolium Audinet-Serville, 1838, and Ph. (Pu.) agathyrsus
Gray, 1843, are referred to as separate species as per the
original authors descriptions. This is based on a combination of
records of overlap between distributions, and the consistent
morphological characteristics expressed between the different
species, which easily distinguish them. Future genetic analysis
will further reveal their relationships with more clarity, but for
now based upon material examined, they are treated as valid
species.
CUMMING, LE TIRANT & TEEMSMA. – Phyllium of Sumatra
18
Acknowledgments
Paul Brock for sending several of his past works to the authors.
Jim Berrian at the San Diego Natural History Museum for
assistance with the camera equipment and much guidance over
the years. Bruno Kneubühler for providing eggs for
examination as well as lengthy discussion about egg
morphology and variability. Jerri Larsson and Tetsuo Miyashita
for allowing us to use specimens from their personal
collections. Alexandre Banko for taking photos of the
Miyashita collection. René Limoges of the Montreal
Insectarium for photography and many professional courtesies.
Johnson Chee Min for the photographs used in figure 2 as well
as sending the corresponding specimen to the first author.
Works cited
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westwoodii (Phasmatodea: Phylliidae): Preliminary Observations.
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(Orthoptères). Libraire Encyclopédique de Roret, Paris, 18: 1-776.
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giganteum Hausleithner (Insecta: Phasmida, Phylliidae) with a
Description of the Newly Discovered Male. Malayan Nature
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Faunitaxys, 6(2), 2018 : 1 – 21. 19
Fig. 10. - A-C: Male Phyllium (Pulchriphyllium) rimiae Seow-Choen, 2017a. - A: Sabah, North Borneo, Mt. Trus Madi, [Coll RC 17-281]. -
B: Malaysia, Cameron Highlands, [Coll RC 17-223]. - C: Sabah, North Borneo, Mt. Trus Madi, [Coll RC 17-282].
- D-E: Egg of Phyllium (Ph.) rubrum Cumming, Le Tirant & Teemsma n. sp. - D: Lateral view, showing standard design of the pinnae [Coll
RC 1-016]. - E: Dorsal view, showing micropylar plate [Coll RC 18-016].
A B C
D E
Brock P. D., 1999. – Stick and Leaf Insects of Peninsular Malaysia
and Singapore. Malaysian Nature Society, Academe Art &
Printing Sdn. Bhd., Kuala Lumpur, 222 p.
Brock P. D., Büscher T. & Baker E., 2018. Phasmida Species File
Online. Version 5.0/5.0. <http://Phasmida.SpeciesFile.org>.
[Accessed January 1st, 2018].
Clark J. T., 1978. – The eggs of leaf insects (Insecta: Phasmida).
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Phylliidae). Faunitaxys, 6(1): 1-5.
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arranged in conformita with its organisation by the Baron
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Gray G. R., 1843. Description of several species of the Genus
Phyllium. Zoologist, 1(1): 117–123.
Größer, D., 2001. Wan d e l nd e B l ät t e r. E i n K a t al o g a ll e r b is h e r
beschriebenen Phylliinae-Arten und deren Eier mit drei
Neubeschreibungen. Edition Chimaira, Frankfurt am Main, 119 p.
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beschriebenen Phylliinae-Arten und deren Eier mit drei
Neubeschreibungen. 2nd Edition. Edition Chimaira, Frankfurt
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Harfield A, 1995. A History on the Honourable East India
Company's Garrison on the West Coast of Sumatra 1685-1825. A
& J Partnership, Dorset, England, 645 p.
Hausleithner B., 1984. Eine neue Phyllium-Art aus Malaysia
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Insektenbörse, 94(4): 39-43.
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On certain species of the genus Phyllium Illiger, 1798, with
proposals for an intra-generic systematization and the
descriptions of five new species from the Philippines and
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Revision der Gattung Phyllium Ill. (Insecta, Orthoptera,
Phasmatoptera). Zoologische Beiträge, 22(1): 49-76.
Linnaeus C., 1758. Systema Naturae. Vol. 1. 10th Edition.
Holmiae, IV + 824 p.
Lucas M. H., 1857. En faisant passer sous les yeux de la Société
un Orthoptère du genre Phyllium et plusieurs oeufs de cet
insecte, communiqué de la note suivante. Bulletins Trimestriels
de La Societe Entomologique de France, (3)5: 147.
Natural History Museum, United Kingdom (NHMUK), 2017.
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anuary 11th, 2018]. Photographs downloaded from:
http://data.nhm.ac.uk/dataset/56e711e6-c847-4f99-915a-6894bb5c5dea/
resource/05ff2255-c38a-40c9-b657-4ccb55ab2feb/record/8231970
http://data.nhm.ac.uk/dataset/56e711e6-c847-4f99-915a-6894bb5c5dea/
resource/05ff2255-c38a-40c9-b657-4ccb55ab2feb/record/8231912
Ragge D. R., 1955. – The wing-venation of the order Phasmida.
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London, 106 (9): 375-392.
Rehn J. A. G. & Rehn J. W. H., 1933. On certain species of the
genus Phyllium (Orthoptera; Phasmidae). Proceedings of the
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pl. 16-17.
Seow-Choen F., 2000. – An Illustrated Guide to the Stick and
Leaf Insects of Peninsular Malaysia and Singapore. Natural
History Publications (Borneo), Kota Kinabalu, 173 p.
Seow-Choen F., 2017a. A taxonomic guide to the stick insects of
Borneo, Volume II. Natural History Publications (Borneo), Kota
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Seow-Choen F., 2017b. – A taxonomic guide to the stick insects of
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CUMMING, LE TIRANT & TEEMSMA. – Phyllium of Sumatra
20
Résumé
Cumming R. T., Le Tirant S. & Te emsma S. N., 2018. – Note sur les Phyllium de la péninsule Malaise et de Sumatra, Indonésie, avec des
informations sur la répartition géographique des espèces actuellement connues, description de la femelle de Phyllium (Pu.) abdulfatahi Seow-
Choen et description d’une nouvelle espèce Phyllium (Ph.) rubrum n. sp. (Phasmida : Phylliidae). Faunitaxys, 6(2) : 1 – 21.
La diversité des Phyllium de la péninsule Malaise et de Sumatra, en Indonésie, est discutée à partir de mentions récentes et de
références plus anciennes. La femelle de Phyllium (Pulchriphyllium) abdulfatahi Seow-Choen, 2017a., jusqu’alors inconnue,
est décrite et illustrée, et l’aire de répartition de l’espèce est précisée. Une extension de l'aire de répartition de Phyllium
(Pulchriphyllium) giganteum Hausleithner, 1984, à Sumatra et dans d'autres régions de Bornéo, est discutée avec les mentions
correspondantes. Phyllium (Phyllium) hausleithneri Brock, 1999 est également nouvellement mentionnée à Sumatra, avec des
notes sur la variation intraspécifique et les figures correspondantes des femelles et des œufs. Une nouvelle espèce Phyllium
(Ph.) rubrum Cumming, Le Tirant & Teemsma n. sp. est décrite et illustrée en provenance de Perak, en Malaisie. Les
distributions géographiques de Phyllium (Pu.) mannani Seow-Choen, 2017a. et de Phyllium (Pu.) rimiae Seow-Choen, 2017a.
sont élargies pour inclure la péninsule Malaise, celle de Phyllium (Pu.) bioculatum Gray, 1832 pour inclure Sumatra, et des
informations sur la variation intraspécifique des mâles sont incluses. Enfin, la nouvelle espèce Phyllium (Ph.) rubrum
Cumming, Le Tirant & Teemsma n. sp. étant décrite à partir d'une femelle, une clé des femelles des espèces du genre Phyllium
de Sumatra, de Singapour et de la péninsule Malaisie est incluse.
Mots-clés. – Phasmatodea, Phasmida, Phylliinae, Phylliini, Phyllium, Malaisie, Sumatra, Indonésie, abdulfatahi, rubrum, description, espèce
nouvelle.
Faunitaxys, 6(2), 2018 : 1 – 21. 21
Faunitaxys est échangée avec les revues suivantes (« print versions ») :
– Annali del Museo Civico di Storia Naturale G. Doria (Italie)
– Boletín de la Asociación española de Entomología (Espagne)
– Boletín de la Sociedad Andaluza de Entomología (Espagne)
– Bollettino del Museo di Storia Naturale di Venezia (Italie)
– Bulletin de la Société linnéenne de Lyon (France)
– Bulletin of Insectology (Italie)
– Heteropterus Rev. Entomol. (Espagne)
– Israel Journal of Entomology (Israel)
– Klapalekiana (République Tchèque)
– Koleopterologische Rundschau (Allemagne)
– Memorie del Museo Civico di Storia Naturale di Verona (Italie)
– Proceedings of the Entomological Society of Washington (USA)
– Revue suisse de Zoologie (Suisse)
– Spixiana (Allemagne)
– Stuttgarter Beiträge zur Naturkunde A, Biologie (Allemagne)
– Zoosystematica Rossica (Russie)
Fig. 11. - Phyllium (Pu.) bioculatum Gray, 1832, male (Natural History Museum United Kingdom collection).
Faunitaxys
Volume 6, Numéro 2, Février 2018
SOMMAIRE
Note sur les Phyllium de la péninsule Malaise et de Sumatra, Indonésie, avec des informations sur la
répartition géographique des espèces actuellement connues, description de la femelle de Phyllium (Pu.)
abdulfatahi Seow-Choen et description d’une nouvelle espèce Phyllium (Ph.) rubrum n. sp.
(Phasmida : Phylliidae)
Royce T. Cumming, Stéphane Le Tirant & Sierra N. Teemsma ............................................ 1 – 21
CONTENTS
On the Phyllium of Peninsular Malaysia and Sumatra, Indonesia, with range expansions for currently
known species, description of the previously unknown Phyllium (Pu.) abdulfatahi Seow-Choen female,
and description of the new species Phyllium (Ph.) rubrum n. sp. from Peninsular Malaysia (Phasmida:
Phylliidae)
Royce T. Cumming, Stéphane Le Tirant & Sierra N. Teemsma ............................................ 1 – 21
Illustration de la couverture : Forêt tropicale de Sumatra.
Crédit :
Fig. 1A-B-C-D, Fig. 2B-D, Fig. 3A-B-C-D, Fig. 4A-C-D-E, Fig. 6A-C-D-E, Fig. 7A-B-C-D-E, Fig. 8A-B-
C-D-E, Fig. 9A-B-C-D and Fig. 10A-B-C-D-E: © Cumming R. T. (San Diego Natural History Museum)
Fig. 2A-C: © Chee Min J. (Malaysia)
Fig. 4B: © Limoges R. (Montreal Insectarium)
Fig. 1E, Fig. 5A-B-C-D and Fig. 11: © Natural History Museum United Kingdom (NHMUK, used under the
license CC0-1.0)
Fig. 6B: © Banko A.
Publié par l’Association Française de Cartographie de la Faune et de la Flore (AFCFF)
... This record was noted as Phyllium siccifolium in Brock (1999) and treated as a smooth abdominal form of P. hausleithneri in Hennemann et al. (2009), however, without examining the specimen in question it is best to leave this as an unconfirmed record. This is because the morphologically similar Phyllium rubrum Cumming, Le Tirant, & Teemsma, 2018 is also found in Peninsular Malaysia and has in the past been considered a form of P. hausleithneri. Phyllium rubrum can be differentiated from Phyllium hausleithneri by the larger size (90.0 mm or larger in female P. rubrum and 74.6-82.8 ...
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Thirteen species of Phyllium (Phyllium) Illiger, 1798 are studied and (re)described in detail with emphasis on those species which exhibit more or less well developed alae in the females and those occurring in the Philippine Islands and on Palawan. Amongst these five new species are described and illustrated from both sexes and the eggs: Ph. (Ph.) ericoriai Hennemann, Conle, Gottardo & Bresseel n. sp. from the Philippine Islands of Luzon, Marinduque and Batan, Phyllium philippinicum Hennemann, Conle, Gottardo & Bresseel n. sp. from the Philippine Island of Luzon, Phyllium mindorense Hennemann, Conle, Gottardo & Bresseel n. sp. from the Philippine Island of Mindoro, Phyllium mabantai Bresseel, Hennemann, Conle & Gottardo n. sp. from the Philippine Island of Mindanao and Ph. (Ph.) gantungense Hennemann, Conle, Gottardo & Bresseel n. sp. from Palawan. Ph. (Ph.) celebicum de Haan, 1842 is re-described with the male and egg described and illustrated for the first time. It is shown to be restricted to Sulawesi and Ambon with all records from continental Asia based on misidentifications mostly relating to Ph. (Ph.) westwoodii Wood-Mason, 1875. All Philippine records of Ph. (Ph.) celebicum de Haan relate to Ph. (Ph.) ericoriai Hennemann, Conle, Gottardo & Bresseel n. sp.. Both sexes and the eggs of Ph. (Ph.) westwoodii Wood-Mason, 1875 are re-described and illustrated and a survey is provided of its intraspecific variability. This species was misinterpreted by most former authors and is here shown to be widely distributed in southern continental Asia having so far been recorded from the Andamans, Myanmar, Thailand, Laos, Kamputchea, S-China, N-Vietnam, Sumatra and the Riouw Archipelago. The holotype of Phyllium (Ph.) siccifolium (Linné, 1758) is described in detail for the first time with illustrations provided. This, the type-species of the entire family Phylliidae, is shown to have been misinterpreted by almost all previous authors and the distribution to be in fact restricted to the Moluccas (Ambon, Ceram, Halmahera, Sula Islands and Banggai). Ambon is shown to be most likely the type-locality of Ph. siccifolium. Records from Peninsular Malaysia have proven to relate to Ph. (Ph.) hausleithneri Brock, 1999 and Philippine material erroneously referred to as "Ph. siccifolium" by various authors is Ph. (Ph.) philippinicum Hennemann, Conle, Gottardo & Bresseel n. sp.. Ph. (Ph.) tobeloense Größer, 2007 from Halmahera (Moluccas) is shown to represent a junior synonym of Ph. siccifolium (n. syn.). Comparison of the Malayan Ph. (Ph.) hausleithneri Brock, 1999 with Malayan specimens previously referred to as "Ph. siccifolium" has revealed these to be the same species which shows considerable variation concerning to the shape of the abdomen in females. Ph. (Ph.) hausleithneri is characteristic for the conspicuous blue interior marking on the meso- and metacoxae. Both sexes and the eggs as well as the remarkable variation of females are illustrated. Similarly strong variation is recorded and illustrated for females of the Javanese Ph. (Ph.) jacobsoni Rehn & Rehn, 1933. A brief discussion of its variability and distribution as well as a summary of the diagnostic features and illustrations of the females and eggs are presented. The Philippine Ph. (Ph.) bilobatum Gray, 1843 is only known from the unique female holotype and all subsequent records appear to have been based on misidentified material. Subsequent records from Peninsular Malaysia relate to Ph. (Ph.) hausleithneri Brock, 1999 and records from Java have all proven to represent Ph. (Ph.) jacobsoni Rehn & Rehn, 1933. The male allotype of Ph. (Ph.) woodi Rehn & Rehn, 1933 from the Philippine island of Mindanao is specifically distinct from the female holotype from Sibuyan Island and here designated as a paratype of Ph. (Ph.) mabantai Bresseel, Hennemann, Conle & Gottardo n. sp.. The diagnostic features of Ph. (Ph.) woodi, a species so far only known from the island of Sibuyan, are briefly summarized. With emphasis on the Philippine fauna, a checklist and keys are provided for the nine species of Phyllium Illiger, 1798 presently known to occur in the Philippine Islands and Palawan. Critical notes are presented on the current intra-generic systematization of Phyllium Illiger, 1798 along with an extended and more detailed distinction between the two subgenera contained, Phyllium Illiger, 1798 and Pulchriphyllium Griffini, 1898. Based on morphological features of the insects and eggs species-groups are suggested within both subgenus. Phyllium (Phyllium) is proposed to include the siccifolium species-group and celebicum species-group, whereas Phyllium (Pulchriphyllium) subdivides into the bioculatum species-group, schultzei species-group, frondosum species-group and brevipenne species-group. The latter two groups are shown to differ considerably from other members of the subgenus and do not belong in Pulchriphyllium (sensu stricto). Keys are provided for the distinction of the speciesgroups here proposed. The celebicum species-group of Phyllium (Phyllium) is discussed in more detail and provisionally contains all those species in which females have developed alae, a fact overlooked for several species by previous authors. Eight species are here attributed to the celebicum species-group and keys are provided to distinguish these. Five species are transferred from one subgenus to the other. Phyllium drunganum Yang, 1995 and Ph. tibetense Liu, 1993 from S-China are removed from the subgenus Pulchriphyllium and transferred to Phyllium (Phyllium) (n. comb.). Ph. chitoniscoides Größer, 1992 and Ph. frondosum Redtenbacher, 1906 from New Guinea as well as Ph. keyicum Karny, 1914 from they Key-Islands are removed from Phyllium (Phyllium) and transferred to the frondosum species-group of Phyllium (Pulchriphyllium) (n. comb.). Ph. insulanicum Werner, 1922 from the Key Islands is removed from synonymy with the New Guinean Ph. frondosum Redtenbacher, 1906 and synoynmised with Ph. keyicum Karny, 1914; differences between Ph. frondosum and Ph. keyicum are presented. The Philippine Phyllium (Phyllium) pusillulum Rehn & Rehn, 1933 is removed from the genus Phyllium Illiger, 1798 and transferred to Microphyllium Zompro, 2001, hence the valid name now is Microphyllium pusillulum (Rehn & Rehn, 1993 n. comb.). Some taxonomically important features traditionally used for distinguishing the genera and species in the family Phylliidae are critically discussed. The present distinction of Chitoniscus Stål, 1875 and Phyllium Illiger, 1798 is shown to be problematic since research on the length relation of the meso-praescutum (anterior portion of the mesonotum in front of the tegmina) has revealed several species in Phyllium Illiger, 1798 that violate the generic description by having this clearly transverse and actually keying out to Chitoniscus Stål, 1875. The prosternal projection characteristic for Chitoniscus Stål, 1875 is shown to be also present in several members of Phyllium Illiger, 1798. Although the entire family Phylliidae was traditionally diagnosed by females having the antennae with nine segment, six species of Phyllium (Phyllium) Illiger, 1798 are here shown to have in fact ten antennomeres. Another interesting fact are the distinctly pectinate ungues (= claws) seen in Ph. (Ph.) gantungense n. sp. which have so far only been known to occur in the Old World areolate family Aschiphasmatidae.
are of an even size from the anterior to the posterior
  • ............................................................................................................... Si
Meso-and metacoxae purple in color; mesopraescutum lateral margins with nodes that are of an even size from the anterior to the posterior......................................................................................................................... Ph. (Ph.) hausleithneri Brock, 1999 (PM, SI, SU)
mesopraescutum lateral margins with nodes that steadily increase in size from the anterior to the posterior
  • Metacoxae Orange............................................................................ Meso
Meso and metacoxae orange to red in color; mesopraescutum lateral margins with nodes that steadily increase in size from the anterior to the posterior.............................................................................. Ph. (Ph.) rubrum Cumming, Le Tirant & Teemsma n. sp. (PM)
Stick and Leaf Insects of Peninsular Malaysia and Singapore
  • P D Brock
Brock P. D., 1999. -Stick and Leaf Insects of Peninsular Malaysia and Singapore. Malaysian Nature Society, Academe Art & Printing Sdn. Bhd., Kuala Lumpur, 222 p.
-Phasmida Species File Online
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Brock P. D., Büscher T. & Baker E., 2018.-Phasmida Species File Online. Version 5.0/5.0. <http://Phasmida.SpeciesFile.org>. [Accessed January 1st, 2018].
1832.-In Griffith & Pidgeon. The animal kingdom arranged in conformita with its organisation by the Baron Cuvier
  • G R Gray
Gray G. R., 1832.-In Griffith & Pidgeon. The animal kingdom arranged in conformita with its organisation by the Baron Cuvier, 15: 191, pl. 63: 3.
1843. -Description of several species of the
  • G R Gray
Gray G. R., 1843. -Description of several species of the Genus Phyllium. Zoologist, 1(1): 117-123.