ArticlePDF Available

A new species of Phyllium Illiger, 1798, from the celebicum species group native to Laos (Phasmida: Phylliidae)

Authors:

Abstract and Figures

Review of leaf-insects collected in Northeastern Laos has revealed a previously unnamed species, which we here describe as Phyllium (Phyllium) oyae species nova. This new species falls within the celebicum species group as described in Hennemann et al. (2009) due to the presence of well developed alae in females and males with a wide exterior profemoral lobe. Upon a review of congenerics, we find that Phyllium (Phyllium) drunganum Yang, 1995 is misplaced and transfer it from the siccifolium species group to the celebicum species group based on the presence of developed alae in females. With so many of the species in this group only known from a single sex, we differentiate Phyllium oyae n. sp. from all species. The species name Phyllium rayongii Thanasinchayakul, 2006 is determined to be a nomen nudum and therefore unavailable according to ICZN Article 16.4.1. To conclude, we present dichotomous keys to males and females known within the celebicum species group.
Content may be subject to copyright.
Faunitaxys
Revue de Faunistique, Taxonomie et Systématique
morphologique et moléculaire
Volume 8
Numéro 4 Février 2020 ISSN : 2269 - 6016
Dépôt légal : Février 2020
Faunitaxys
Revue de Faunistique, Taxonomie et Systématique
morphologique et moléculaire
ZooBank : http://zoobank.org/79A36B2E-F645-4F9A-AE2B-ED32CE6771CC
Directeur de la publication, rédacteur, conception graphique et PAO :
Lionel Delaunay
Cette revue ne peut pas être vendue
Elle est distribuée par échange aux institutions (version papier)
et sur simple demande aux particuliers (format PDF)
à l’adresse suivante :
AFCFF
28, rue Voltaire, F- 42100 Saint Etienne
E-mail : lionel.delaunay@free.fr
Elle est disponible librement au téléchargement à partir du site :
http ://faunitaxys.fr/
La parution de Faunitaxys est apériodique
Imprimée sur les presses de SPEED COPIE
6, rue Tréfilerie, F- 42100 Saint-Etienne
Imprimé le 27 février 2020
A new species of Phyllium Illiger, 1798, from the celebicum species group
native to Laos (Phasmida: Phylliidae)
ROYCE T. CUMMING (1, 2) & STEPHANE LE TIRANT (3)
(1) Associate researcher, Montréal Insectarium, 4581 rue Sherbrooke, Montreal, Quebec, Canada, H1X 2B2 - phylliidae.walkingleaf@gmail.com
- ZooBank : http://zoobank.org/6CA8501F-10BA-4E07-9BF4-65CFCE4E9E92
(2) PhD Student, Richard Gilder Graduate School, American Museum of Natural History, New York City, New York, United States, 10024.
(3) 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
Abstract. Review of leaf-insects collected in Northeastern Laos has revealed a previously
unnamed species, which we here describe as Phyllium (Phyllium) oyae species nova. This new
species falls within the celebicum species group as described in Hennemann et al. (2009) due to the
presence of well developed alae in females and males with a wide exterior profemoral lobe. Upon a
review of congenerics, we find that Phyllium (Phyllium) drunganum Yang, 1995 is misplaced and
transfer it from the siccifolium species group to the celebicum species group based on the presence
of developed alae in females. With so many of the species in this group only known from a single
sex, we differentiate Phyllium oyae n. sp. from all species. The species name Phyllium rayongii
Thanasinchayakul, 2006 is determined to be a nomen nudum and therefore unavailable according
to ICZN Article 16.4.1. To conclude, we present dichotomous keys to males and females known
within the celebicum species group.
Cumming R. T. & Le Tirant S., 2020. A new species of Phyllium Illiger, 1798, from the celebicum species
group native to Laos (Phasmida: Phylliidae). Faunitaxys, 8(4) : 1 – 9.
Introduction
To d ate , on ly in fr eq uent r eco rd s o f P hy ll iida e fr om La os ha ve
surfaced and made their way into publications or into museum
collections, creating a rather insufficient view of the phylliid
diversity of the country. It is not surprising that Phylliidae are only
rarely collected in the wild as their incredible leaf-like crypsis
likely allows them to evade visually oriented predators (and
entomologists alike, Fig. 1). In a preliminary review of numerous
museum collections and more recently fresh specimens sent to the
authors for identification, it became apparent that there were at
least two species of Phyllium Illiger, 1798 present in Laos.
The first noted specimens are those which are morphologically
identical to Phyllium westwoodii Wo od -M as o n, 18 75 w hi ch we re
collected in northern Laos along the same latitude as records of
specimens from northern Thailand. This northern Thailand
population is readily available for sale to collectors in large
quantities each year from breeders in Thailand and is well known.
Currently, molecular analysis of specimens from northern Thailand
and northern Laos are being conducted to confirm/correct this
identification as Phyllium westwoodii.
A second species was also identified but infrequently collected
until larger series of bred specimens became available from Steeve
Collard (Laos, September, 2013 to present) and Johnson Sau
(Malaysia, August, 2018), who sent specimens to the authors.
Upon first review, we thought they could represent a range
expansion for Phyllium drunganum Yang, 1995 which is known
from Drung-Nu Autonomous County, China. After reviewing in
detail the material we received from Laos however, it became
apparent that the structure of the antennae was consistently
different from Phyllium drunganum, and the Laos population
Keywords :
1
Faunitaxys, 8(4), 2020 : 1 – 9.
ZooBank : http://zoobank.org/318DC3DF-A047-4812-AB6C-45D23242310A
Phasmatodea ;
Phasmida ;
Phylliidae ;
Phylliini ;
Phyllium ;
Southeast Asia ;
Laos ;
oyae ;
celebicum ;
Taxonomy ;
description ;
new species.
Fig. 1. Live female Phyllium oyae n. sp. photographed in the wild.
CUMMING & LE TIRANT. – Phyllium oyae n. sp. of Laos
instead represented an undescribed species. This species is
compared below to all members within the celebicum species
group as described by Hennemann, et al., 2009 and is named
Phyllium oyae n. sp. after Steeve Collard’s wife as he first sent us
specimens of this new species and has been incredibly helpful in
supplying fresh material and photographs of the habitat (Fig. 2).
Materials and Methods
Photographs of specimens were taken by René Limoges of the Montreal
Insectarium using a Nikon D810 DSLR camera with Nikon Micro-
Nikkor 200mm f/4 lens on Manfrotto 454 micrometric positioning
sliding plate. Lighting was provided by two Nikon SB-25 flash units
with a Cameron Digital diffusion photo box. Adobe Photoshop
Elements 13 was used as post processing software. Photos of the
holotype and a paratype female were taken by Frank Hennemann
(Germany) using a Nikon D7000 camera equipped with a Nikon DX
AF-S Micro 40 mm lens and a wireless Nikon SU-800 dual speed light
system. Background lihghtning was provided by a 18W 6000K LED
panel light plate. Measurements of the holotype were made to the
nearest 0.1 mm using digital calipers. Egg orientation terminology
follows the terminology presented by Clark (1978). The holotype
specimen is deposited in the Montreal Insectarium type collection.
Abbreviations
IMQC: Insectarium de Montréal, Montréal, Québec / Canada.
Coll RC: Private collection of Royce T. Cumming / U.S.A.
Coll SLT: Private collection of Stéphane Le Tirant / Canada.
Taxonomy
Phyllium (Phyllium) drunganum Yang, 1995
Remarks. – Since the original description, where Phyllium drunganum
was not placed within a particular subgenus (Yang, 1995 & Chen,
1999), there has been significant confusion about the taxonomic
placement of this species. Some authors have placed Phyllium
drunganum within Phyllium (Pulchriphyllium) based on all tibiae with
small exterior lobes (Zompro, 2004; Größer, 2008) and others in the
Phyllium (Phyllium) subgenus (Otte & Brock, 2005; Hennemann, et al.,
2009; Chen & He, 2008).
Most recently with the arrangement of species groups by Hennemann et
al. (2009), Phyllium drunganum was returned to the Phyllium
(Phyllium) subgenus, but unfortunately it was placed within the
siccifolium species group without morphological justification. The
authors were relying on the figure and English summary of the
description provided by Yang (1995) but did not translate the
morphological description in its entirety and assumed that because the
illustration did not show developed alae, that there were none.
Fortunately, we were able to translate the morphological description in its
entirety. To do so we used the most recent version of the phone based ap
“Google Translate” (offere d by Google LLC: version
6.2.0.RC07.268294262) to translate the morphological description of
Phyllium drunganum. We found that the second line on page 19 of the
description translates to “The rear/posterior wing is thin and transparent,
extending only to of the sixth abdominal section” (Yang, 1995). The
presence of developed alae is a feature which helps to define the celebicum
species group as described by Hennemann et al. (2009), therefore we here
note the correct placement of Phyllium drunganum to be within the
celebicum species group, not within the siccifolium species group.
Besides this important morphological feature now known to be present
in Phyllium drunganum, there are other morphological features which
additionally help to justify the transfer. One is the profemoral exterior
lobe which is broad and rounded (which is a feature found in all female
celebicum species group members but also in some Phyllium
(Pulchriphyllium) members as well so it is not an exclusive feature, but
helpful to differentiate the celebicum species group from the siccifolium
species group). Also, all tibiae have small distal exterior lobes, a feature
which is also seen in Phyllium tibetense Liu, 1993. Additionally, the
fourth antennal segment is flattened and disk-like, a feature unique to
and seen in all members of the celebicum species group, except the
Philippine Phyllium bonifacioi Lit & Eusebio, 2014 and Phyllium
ericoriai Hennemann et al., 2009 which lack this disk-like segment.
Based on these morphological features, we here transfer Phyllium
(Phyllium) drunganum to the celebicum species group as described by
Hennemann et al. (2009).
Phyllium (Phyllium) oyae Cumming & Le Tirant n. sp.
(Fig. 1, 3A-C, 4A-D, 5A-E, 6A-D, 7A-C)
ZooBank : http://zoobank.org/0F657962-4BB8-488A-9C75-D35C5AA053B2
Holotype, . – NE-Laos, Mount Phu Phan, 2060 m, IX.2013. Ex:
Frank H. Hennemann private collection / Germany. Deposited in the
Montreal Insectarium type collection. (Fig. 3B).
Paratypes [2 50 ♀♀ , 4 4 ♂♂, 2 5 eggs ]. – LAOS, N.Eastern. Bred locally
in Laos. See below deposition data for individual data.
Depositions.
– 2 ♀♀, 1 [Royce Cumming private collection, California / United States,
Coll RC 18-423 — 18-425: LAOS, N.Eastern, Hua Phan Prov., Xamnuen
District, Ban Saleui, VIII.2018. Local Collector].
14 ♀♀, 6 ♂♂, 2 subadult , 2 subadult ♂♂ [Royce Cumming private
collection, California / United States, Coll RC 19-063 — 19-086: LAOS,
N.Eastern, Hua Phan Prov., Xamnuen District, Ban Saleui, VIII.2018].
– 3 ♀♀ [Royce Cumming private collection, California / United
States19-170 — 19-172: LAOS: Mt Phu Phan 2060m VI.2019 N-E Laos].
– 17 eggs [Royce Cumming private collection, California / United
States, Coll RC 19-183 — 19-199: LAOS, Ban Saleuy, from Bruno
Kneubuhler, Switzerland, December, 2019].
190 , 16 ♂♂, 4 eggs [Stephane Le Tirant private collection, Québec /
Canada]. LAOS, N.Eastern, Hua Phan Prov., Xamnuen District, Ban
Saleui, VIII.2018.
– 10 ♀♀, 10 ♂♂, 1 egg [Insectarium de Montréal collection, Montréal,
Québec / Canada] LAOS, N.Eastern, Hua Phan Prov., Xamnuen
District, Ban Saleui, VIII.2018.
5 ♀♀, 5 ♂♂, Ban Saleuy, Xamnuea, Houaphan Province, NE Laos,
[Oskar V. Conle private collection, Bolsterlang / Germany].
– 4 [Frank H. Hennemann private collection / Germany. NE-Laos,
Mount Phu Phan, 2060 m, IX.2013] [coll. FH, No's 1060-1 1060-4].
2
Fig. 2. Steeve Collard (center) with locals in Hua Phan Province where they collected
the holotype of Phyllium oyae n. sp. and where the paratype series was bred.
Faunitaxys, 8(4), 2020 : 1 – 9. 3
Fig. 3. Dorsal view of Phyllium oyae n. sp. A. Female paratype. B. Female holotype. C. Male paratype.
– 2 ♂♂ [Frank H. Hennemann private collection / Germany. NE-Laos,
Mount Phu Phan, 2060 m, VI.2019] [coll. FH, No's 1060-5 1060-6].
– 10 ♀♀ [Frank H. Hennemann private collection / Germany. LAOS,
N.Eastern, Hua Phan Prov., Xamnuen District, Ban Saleui, VIII.2018.]
[coll. FH, No's 1060-7 1060-16].
3 eggs [Frank H. Hennemann private collection / Germany, ex: Royce
Cumming private collection (Coll RC 19-200—19-202). LAOS, Ban Saleuy,
from Bruno Kneubuhler, Switzerland, December, 2019] [coll. FH, No's 1060-E].
1 [California Academy of Sciences, San Francisco, California /
U.S.A.] LAOS, N.Eastern, Hua Phan Prov., Xamnuen District, Ban
Saleui, VIII.2018. Local Collector.
2 , 1 : [Royal Belgian Institute of Natural Sciences / Belgium, Coll.
I.R.Sc.N.B.]. Laos, NE, Mt Phu Phan, vi.2019, local collectors, I.G.: 34.159.
2 ♀♀: [Royal Belgian Institute of Natural Sciences / Belgium, Coll.
I.R.Sc.N.B.]. Laos, NE, Mt Phu Phan, ix.2019, local collectors, I.G.: 34.159.
1 [Lyman Entomological Museum, McGill University, Québec /
Canada] LAOS, N.Eastern, Hua Phan Prov., Xamnuen District, Ban
Saleui, VIII.2018. Local Collector.
1 [Museum National d’Histoire Naturelle, Paris / France] LAOS, N.Eastern,
Hua Phan Prov., Xamnuen District, Ban Saleui, VIII.2018. Local Collector.
– 1 [The Natural History Museum United Kingdom, London /
United Kingdom] LAOS, N.Eastern, Hua Phan Prov., Xamnuen
District, Ban Saleui, VIII.2018. Local Collector.
– 1 [San Diego Natural History Museum, San Diego / U.S.A]
LAOS, N.Eastern, Hua Phan Prov., Xamnuen District, Ban Saleui,
VIII.2018. Local Collector.
1 [Alexandre Banko private collection, Québec /Canada] LAOS, N.Eastern,
Hua Phan Prov., Xamnuen District, Ban Saleui, VIII.2018. Local Collector.
– 1 [Tetsuo Miyashita private collection / Japan] LAOS, N.Eastern, Hua
Phan Prov., Xamnuen District, Ban Saleui, VIII.2018. Local Collector.
Differentiation. Because the celebicum species group is poorly
known with many species from mainland Asia only known from a
single sex, we discuss all species in the group in order to sufficiently
differentiate and place Phyllium oyae n. sp. taxonomically.
First, we can differentiate Phyllium oyae n. sp. from all species that
have females with mesopleurae margins which do not reach from the
posterior of the mesopraescutum completely to the anterior margin
(Phyllium oyae n. sp. has mesopleurae which span the entire length,
Fig. 4D). This simple and clearly observed feature differentiates
Phyllium oyae n. sp. from: Phyllium athanysus Westwood, 1859;
Phyllium celebicum de Haan, 1842; Phyllium westwoodii Wood-Mason,
1875; and Phyllium chrisangi Seow-Choen, 2017.
Next, two species which can easily be differentiated based on their
antennae are Phyllium bonifacioi Lit and Eusebio, 2014 and Phyllium
ericoriai Hennemann et al., 2009 which have the fourth antennal
segment of a similar length and shape to the following segments (not
short and disk-like as observed in Phyllium oyae n. sp. and all other
species in the celebicum species group, Fig. 4C).
Another species easily differentiated is Phyllium yapicum Cumming &
Tee m sm a, 2 01 8. T hi s sp ec ie s h as t he u ni q ue f e at ur e o f a p ro ti bi al
interior lobe which is only distributed on the proximal half, while the
distal half is thin and almost lacking a lobe. Phyllium oyae n. sp., like
the other species in the group, has a protibial lobe which is fully
spanning the protibial length.
The remaining three species with morphologically similar females are
discussed individually as one of these species is likely the sister species
to Phyllium oyae n. sp. based on morphology and geographic
distribution.
Phyllium tibetense L iu, 19 93. Ma jor form females of Phyllium oyae n. sp.
share the most morphological similarities to Phyllium tibetense such as
A B C
10 mm
more prominent abdominal segment VII lobes and small but developed
tibial exterior lobes on the distal fifth of the tibia, a feature which is
clearly noted and illustrated in the description of Phyllium tibetense Liu,
1993. Also, a shared feature between the two species is the antennal
morphology, with the terminal segment longer than and nearly as wide
as the preceding segment which was uniformly observed throughout the
paratype series of Phyllium oyae n. sp. (Fig. 4C). A simple feature to
differentiate Phyllium tibetense and Phyllium oyae n. sp. is the length of
the female subgenital plate, which at most reaches three quarters through
segment X in Phyllium oyae n. sp., but in Phyllium tibetense is clearly
longer and exceeding the apex of segment X (a feature which helps to
differentiate Phyllium tibetense from all other species group members).
Phyllium rarum Liu, 1993 (only known from the female sex). Features
which are similar between Phyllium rarum and Phyllium oyae n. sp.
are the structure of the genitalia, mesopleurae which span the full length
of the mesopraescutum, and an alae length reaching the posterior margin
of the sixth abdominal segment. Although not the most common form,
there are occasionally female Phyllium oyae n. sp. with a profemoral
exterior lobe having a 90 degree angle (see Fig. 5A for a paratype with
such a shape) as observed in Phyllium rarum. An easy feature to
differentiate the two species is the abdominal shape, with Phyllium
rarum lacking a lobed seventh abdominal segment and having a spade
shaped abdomen instead. Abdominal shape is generally a feature which
we wish to avoid when differentiating between different species.
However, with such a large paratype series, we are confident in the
degree of variation within female Phyllium oyae n. sp. with even the
most slender form having a boxy abdomen with segment VI parallel
sided and not converging as seen in Phyllium rarum.
Based on the female alone, Phyllium oyae n. sp. appears to be most
morphologically similar to Phyllium druganum Yang, 199 5 kn own from
Drung-Nu Aut. Co., China, a distance of nearly 1,000 kilometers from
the Phyllium oyae n. sp. type locality. Features which are shared
between the two species are: mesopleurae which span the full length of
the mesopraescutum (Fig. 4D), small tibial exterior lobes on all legs on
the distal most end, a subgenital plate which does not exceed the apex of
the abdomen (Fig. 4B), and gently lobed abdominal segment VII (Fig. 3A).
The only feature which easily separates these two species is the
morphology of the antennae (which Yang (1995) highlighted as an
important feature to differentiate Ph. drunganum from Ph. tibetense
also) with the terminal segment shorter and narrower than the preceding
segment, not longer and nearly as wide as the preceding segment
observed in Phyllium oyae n. sp. (Fig. 4C). Unfortunately, we do not
yet know male and egg morphology for Phyllium drunganum which
might prove to be important differentiation features, once known.
The two species with only males known which are most
morphologically similar to Phyllium oyae n. sp. are Phyllium
yunnanense Liu, 1993 (described from Mongla, Yunnan Province,
China, just over 400 kilometers from the type locality of Phyllium oyae n. sp.)
and Phyllium parum Liu, 1993 (described from Baisha, Hainan Island,
China, almost 600 kilometers from the type locality of Phyllium oyae n. sp.).
From Phyllium yunnanense, Phyllium oyae n. sp. can be differentiated
by the width of the profemoral exterior lobe which is thinner and arches
smoothly from end to end in Phyllium yunnanense (only about three
times the width of the profemoral shaft itself) versus Phyllium oyae n. sp.
which possesses a wider profemoral exterior lobe with a maximum
width up to five times the width of the profemoral shaft and a distinct
but rounded angle in the center of the span (Fig. 6C).
From Phyllium parum, Phyllium oyae n. sp. can be differentiated by
the length of the tegmina; only reaching the middle of the third
abdominal segment in Phyllium parum and longer in Phyllium oyae n. sp.
reaching the middle of the fourth abdominal segment. The mesopleurae
are also slightly different with the margins in Phyllium oyae n. sp.
almost perfectly straight (Fig. 6A), versus Phyllium parum with a gentle
bend giving the anterior margin a thinner appearance.
CUMMING & LE TIRANT. – Phyllium oyae n. sp. of Laos
4
Ta bl e 1 . Measurements of Phyllium oyae n. sp., holotype female, paratype males and females. All measurements made to the nearest 0.1mm. Measurements for
paratypes are given with a minimum to maximum range.
*Including cerci and head, excluding antennae.
** Only a subset of paratypes had their tegmina opened to measure the alae, the holotype was not measured.
Feature
Holotype
Paratypes ♀♀
Paratypes ♂♂
Length of body*
89
82.1 - 94.2
73.0 - 74.8
Length/greatest width of head
7.9 / 6.4
5.5 - 6.4 / 6.5 - 7.3
3.6 - 3.8 / 3.8 - 3.9
Pronotum
5.2
4.0 - 6.0
3.5 - 3.7
Mesonotum
7.0
6.7 - 7.1
3.8 - 4.0
Length of tegmina
52.3
42.1 - 53.2
27.0 - 28.9
Length of alae
**
17.5 - 18.0
51.0 - 53.2
Greatest width of abdomen
40.0
32.0 - 36.6
22.1 - 22.5
Profemora
18.5
13.1 - 18.5
15.0 - 15.3
Mesofemora
13.9
11.2 - 12.5
11.5 - 12.1
Metafemora
16.2
14.6 - 15.4
13.3 - 13.7
Protibiae
10.2
7.7 - 10.2
8.9 - 9.1
Mesotibiae
9.7
7.3 - 9.8
7.6 - 7.7
Metatibiae
13.8
11.4 - 13.8
9.3 - 9.5
Antennae
4.2
3.2 - 4.1
42.8 - 46.5
Faunitaxys, 8(4), 2020 : 1 – 9. 5
Morphological description of the female.
Coloration. – The coloration description is based on photographs of live
individuals and well preserved specimens. To date only a green color form
has been observed (Fig. 1) with all paratypes nearly identical in coloration
and no variable brown patches of color as are sometimes observed on
other Phyllium species. The base color of the entire individual is a lime
green with several features highlighted with tan to yellow coloration.
These features are the antennae, eyes, posteriomedial tubercle, margins of
the pronotum, spination of the thorax, and the venation of the tegmina.
Morphology
Head. Capsule longer than wide, vertex with slight granulation, the
posteromedial tubercle is not much more prominent than the granulation on the
head capsule (Fig. 4A). Frontal convexity broad and stout; at the longest, it is
half of the length of the first antennomere. Antennae consisting of nine
segments, with the terminal segment slightly longer than the previous two
segments combined and just as wide, not notably narrower (Fig. 4C).
Antennomere IV is notably shorter than V, VI, or VII and is instead compact and
disk-like. Antennomere I through the first half of VIII sparsely marked with
small tan setae; the distal half of VIII and antennomere IX are covered in dense,
stout, dark setae.Compound eyes are not particularly large, only slightly
protruding from the head capsule and taking up about one quarter of the length
of the capsule (Fig. 4A). Ocelli abse nt. Antennal fields only a little wider
than the first antennomere but not protruding back farther than the frontal suture.
Thorax.Pronotum with anterior margin slightly concave and
lateral margins that are relatively straight converging to a narrow,
straight posterior margin that is about half the width of the anterior rim
(Fig. 4D). The pronotum surface is slightly lumpy but lacking any
notable granules or wrinkles, with only a prominent pit in the center,
and weak furrows anterior and lateral to the pit. The pronotum has a
prominent anterior rim which is marked with small granulation,
moderate lateral rims which are relatively smooth, and a posterior
margin that is smooth, lacking a rim (Fig. 4D).Prosternum and the
mesosternum with granulation throughout, and a metasternum which
only has granulation on the lateral margins with the center relatively
smooth.Mesopraescutum approximately as long as wide with the posterior
margin faintly narrower than the anterior margin. Lateral rims with nine to ten
rounded tubercles ranging in size from small to medium, with only minor
granulation between tubercles. Mesopraescutum anterior rim without a
prominent sagittal spine, instead the anterior rim has a granular surface
throughout, none very large. Mesopraescutum crest not prominent,
only slightly raised above the mesopraescutum face and with only
minimal granulation throughout (Fig. 4D). Mesopleurae evenly
diverging with straight margins; lateral margin with five to six major
tubercles and six to seven minor tubercles spread amongst the major
tubercles (Fig. 4D). Face of the mesopleurae wrinkled and with slight
granulation, and the surface has two faint divots, one on the anterior
third and one on the posterior third.Tegmina with slight variation in
Fig. 6. Phyllium oyae n. sp. male paratype detail photos. A. Thorax dorsal.
B. Genitalia ventral. C. Profemoral lobes dorsal. D. Antennae and head capsule.
Fig. 7. Phyllium oyae n. sp. egg. A. Dorsal view showing the micropylar plate.
B. Lateral view. C. Ventral view.
A B C
D
A B C
Fig. 4. Phyllium oyae n. sp. female paratype details. A. Detail of head and antennae, dorsal. B. Genitalia ventral (note that the subgenital
plate is split in half from the gutting process). C. Antennae detail, dorsal. D. Thorax dorsal.
Fig. 5. Phyllium oyae n. sp. female paratypes, profemoral lobes showing the variation observed within a large series of this species.
A B C
D
A B C D E
length, ranging in length from one-third the way through abdominal
segment VII to at most reaching about one-third of the way into
segment VIII.
Alae. Well developed, about 17.5–18.0 mm in length, almost
reaching to the tip of the tegmina.
Abdomen. Abdominal segments II through the anterior one-third of IV
diverging, posterior two-thirds of IV through VI subparallel, only gently
converging. Segment VII is slightly variable, either only gently rounded or can
be moderately rounded, but to date we have not seen a strongly lobed individual.
Segments VIII through X are notably narrower than segment VII and converge
uniformly with the anal abdominal segment ending in a broad rounded apex.
Subgenital plate starts at the anterior margin of segment VIII and extends
about halfway into segment X, ending in a fine point.Gonapophyses r e a c h t h e
posterior margin of segment X and are quite broad, each about as broad as the
subgenital plate projection is wide (Fig. 4B). Cerci o nl y gen tl y cu ppe d, w ith a
lumpy surface and numerous thin brown setae mostly situated along the margins
leaving the center relatively bare.
Legs.Profemoral lobes v a r i a b l e , b u t t h e e x t e r i o r l o b e i s a l w a y s w i d e r t h a n
the interior lobe. Profemoral exterior lobe can be nearly a 90 degree angle at
the smallest (Fig. 5A), or more commonly is an acute angle with the proximal
margin straight and the distal margin slightly convex (Fig. 5E), but this feature is
variable and continuous, not with discrete forms. Edge of the exterior
profemoral lobe with three to six serrate teeth with those at the bend in the angle
presenting as the largest teeth, and those on the margins smaller.Profemoral
interior lobe wide and angular, always with an obtuse angle and marked with
five looping teeth arranged in a 2-1-2 pattern with the central tooth generally
the largest or even in size to the two closest to the protibial shaft (Fig. 5).
Mesofemoral interior lobe arcs from end to end evenly and has five to six
serrate teeth distributed on the distal half. The interior lobe is slightly thinner than
the exterior lobe, and the interior lobe is always a smooth arc without an angle,
while the exterior lobe is slightly wider and has a distinct rounded bend near the
center and can be without small teeth or occasionally has one to two small teeth
on the distal half.Metafemoral interior lobe arcs end to end and has five to six
serrate teeth pointing distally.Metafemoral exterior lobe is thin and smooth,
hugging the metafemoral shaft.Protibial interior lobe spans the entire length
of the shaft, in a scalene triangle with the thicker end on the distal half and is
about three times as wide as the protibial shaft itself. Mesotibiae and
metatibiae lacking interior lobes.. Pro-, meso-, and metatibiae with slender
small exterior lobes on the distal fifth only. These small lobes can be slightly
variable and are more easily observed on broader females.
Morphological description of the male.
Coloration.All coloration descriptions are based on preserved specimens, not
from live individuals. After reviewing photographs of live females compared
with dried females, we expect the males to be a more vibrant green in life like
their female counterparts. Overall coloration pale green to olive green, with
touches of yellowing throughout (likely due to the specimen preservation
process), but particularly notable on the antennae, tegmina, legs, and thorax.
Compound eyes of a rusty brown color. Although uncommon, we did observe a
few males with variable brown coloration on the protibiae (one half to three
quarters colored) and on the same males half or slightly more than half of the
mesofemora were also colored.
Morphology
Head.Capsule about as long as wide, with a vertex that is rather smooth
(Fig. 6D). Frontal convexity stout with sides that evenly converge to the point,
sparsely covered in thin transparent setae. The posteromedial tubercle is not
particularly broad, and is only slightly raised on the posterior of the head capsule.
Antennae consisting of 28 segments (including the scapus and pedicellus), all
segments except the scapus and pedicellus and terminal three are covered in a
moderately dense dark setae that are generally as long as the antennae segment is
wide (Fig. 6D). The terminal three segments have dense dark setae throughout the
surface.Compound eyes l arge and bulb ous , no tabl y pr otr udin g aw ay fr om t he
head capsule..Ocelli are absent.Antennal fields of approximately the same
width as the scapus but not notably broad.
Thorax. Pronotum with anterior margin gently concave and lateral
margins that are nearly straight and converging to the posterior margin which is
less than half of the width of the anterior (Fig. 6A). Anterior and lateral margins
of the pronotum have distinct rims, and the posterior margin lacks a rim. Face of
the pronotum with a slightly wrinkled surface and marked by a distinct medial
and perpendicular furrow and a central pit.Prosternum and mesosternum are
granulose throughout with nodes of somewhat even size and regular spacing.
Metasternum lacking the nodes which are common across the surface of the
pro- and mesosternum and instead has a surface covered in wrinkles throughout.
Mesopraescutum slightly longer than wide, with lateral margins which only
slightly converge to the posterior (Fig. 6A). Lateral rims with eight to ten
medium sized tubercles of slightly varying size, but none notably smaller or
larger than those around them giving the margins a uniform serrate texture. The
surface of the mesopraescutum rises up to slightly along the sagittal plane with a
face that is marked with uniform granules throughout. Mesopraescutum anterior
margin with a granular surface. Mesopleurae wide for a male, starting to
diverge near the anterior margin and uniformly diverging throughout their length
(Fig. 6A). Lateral margin with six to seven medium sized tubercles spread
throughout the length, with six to seven smaller tubercles/nodes intermixed
throughout the more prominent tubercles. Face of the mesopleurae wrinkled and
marked with two faint divots, one on the anterior third and one on the posterior
third.Teg m i n a long, extending half way through abdominal segment IV.
Alae. Well d ev el o pe d in a n o v al f an c on f ig ur at i on , qu i te l en gt h y
reaching half way onto abdominal segment VIII, and occasionally extending
further onto abdominal segment IX.
Abdomen. Abdominal segments II through the anterior half of IV
gradually diverging in a gentle curve, posterior half of IV through X
Fig. 8. Habitat in Hua Phan Province where Steeve Collard frequently
collects various insects.
CUMMING & LE TIRANT. – Phyllium oyae n. sp. of Laos
6
Fig. 9. Leaf samples of the local host plants specimens which Phyllium oyae n. sp.
has been collected feeding on A. Castanopsis sp. B. Quercus sp.
A B
1. Fourth antennal segment approximately as tall as it is wide, not short and disk-like .................................................... 2
Fourth antennal segment short and disk-like, several times wider than tall ................................................................ 3
2(1). Abdominal segment VII lateral margin with distinct lobe; angle of profemora exterior lobe approximately 90 degrees;
Philippines (Luzon Is., Batan Is., Marinduque Is., Catanduanes Is.) ......................... Ph. (Ph.) ericoriai Hennemann et al., 2009
Abdominal segment VII lateral margin with indistinct lobe and converging posteriorly; angle of profemora exterior lobe obtuse;
Philippines (Northern Luzon Is.) .................................................................... Ph. (Ph.) bonifacioi Lit and Eusebio, 2014
3(1). Metatibia with exterior lobe across the full length of the shaft; Sri Lanka ........................ Ph. (Ph.) athanysus Westwood, 1859
Metatibiae simple (lacking an exterior lobe) or in a few species the distal fifth of the metatibia has a small lobe, never with a
full lobe across the entire length ................................................................................................................. 4
4(3). Protibiae interior lobe unevenly distributed, almost entirely on the proximal half, with the distal half greatly reduced;
Micronesia,Yap Is ............................................................................ Ph. (Ph.) yapicum Cumming and Teemsman, 2018
Protibiae interior lobe distributed across the entire length of the shaft, either with even weighting on the distal and proximal
halves, or with the weighting towards the distal half not on the proximal half ........................................................... 5
5(4). Subgenital plate long and thin with the apex exceeding the length of the terminal abdominal segment; China: Tibet,
Xizang ................................................................................................................. Ph. (Ph.) tibetense Liu, 1993
Subgenital not projecting past the apex of the abdominal segment X, only reaching one- quarter to three-quarters of the length
through abdominal segment X .................................................................................................................... 6
6(5). Mesopleurae reaching from the posterior to only about one-half to three-quarters of the way to the anterior margin of the
mesopraescutum, with gently concave margins creating a slender appearance ........................................................... 7
Mesopleurae reaching from the posterior to the anterior of the mesopraescutum, with straight margins, creating a prominent
triangular appearance .............................................................................................................................. 9
7(6). Profemoral exterior lobe recurved, due to a concave proximal margin and convex distal margin; Indonesia
(Sulawesi) ...................................................................................................... Ph. (Ph.) celebicum de Haan, 1842
Profemoral exterior lobe with a right angle or nearly right angle (with straight distal and proximal margins, neither concave), or
with a roundly obtuse angle (with both distal and proximal margins convex) ............................................................ 8
8(7). Alae about half the length of the tegmina, only reaching abdominal segments III or IV; Singapore ....................................
............................................................................................................... Ph. (Ph.) chrisangi Seow-Choen, 2017
Alae nearly as long as the tegmina, reaching abdominal segments VI or VII; Myanmar, Thailand, Laos .................................
.............................................................................................................. Ph. (Ph.) westwoodii Wood-Mason 1875
9(6). Abdominal segments VI and VII distinctly converging, VII without lobes, giving the abdomen a spade shaped appearance;
China: Guangxi-Zhuang ............................................................................................... Ph. (Ph.) rarum Liu, 1993
Abdominal segments VI and VII with parallel or only slightly subparallel margins and segment VII with a smooth but
noteworthy lobe ................................................................................................................................... 10
10(9).Terminal antennal segment shorter and narrower than the preceding segment; China: Yunnan Prov, Gongshan Drury-Nu, Auct. Co.
........................................................................................................................ Ph. (Ph.) drunganum Yang, 19 95
Terminal antennal segment longer and nearly as wide as the preceding segment; Laos ........................... Ph. (Ph.) oyae n. sp.
Key to known females in the Phyllium (Phyllium) celebicum species group
Adapted from key in Cumming & Teemsma (2018). Phyllium (Phyllium) parum and Phyllium (Phyllium) yunnanense females
unknown/ undescribed and are therefore excluded.
Faunitaxys, 8(4), 2020 : 1 – 9. 7
converging to the apex, giving the abdomen a spade shaped appearance.
Poculum broad, and ending in a rounded apex that slightly passes the
anterior margin of segment X (Fig. 6B).Cerci long and slender, protruding
prominently from under the anal abdominal segment, gently cupped, and
covered in numerous stout dark setae and a granular surface throughout.
Vo m e r s t o u t a n d b r o a d w i t h c o n v e x s i d e s c o n v e r g i n g t o t w o a p i c a l h o o k s ( o n e
larger, the other smaller) pointing upwards into the paraproct (Fig. 6B).
Legs. – Profemoral exterior lobe almost as wide as the interior lobe with a
rounded but notable bend instead of smoothly arcing end to end, and marked
with three to four small serrate teeth (Fig. 6C). Profemoral interior lobe
without a strong angle and marked with five to six prominent serrate teeth with a
large looping gap between the third and fourth teeth (Fig. 6C).Mesofemoral
exterior lobe arcs end to end, and lacks a strong angle, but it is slightly
thicker on the distal half which is marked with two to three finely serrate
teeth. Mesofemoral interior lobe is equal in width to the exterior lobe and
similarly shaped but with six to seven fine serrate teeth on the distal half instead.
Metafemoral exterior lobe lacks dentition and hugs the shape of the
metafemoral shaft. Metafemoral interior lobe gently arcs end to end with
seven to eight small serrate teeth on the distal half. Protibial exterior with only
a faint expansion of a slight lobe on the distal fifth, which is occasionally highly
reduced, with the remainder of the protibiae lobeless. Protibial interior lobe
reaching end to end in a rounded scalene triangle, thickest on the distal third.
Meso- and metatibiae simple, lacking lobes completely.
Description of Egg (Fig. 7A-C).
Angular; pentagonal in cross section.Capsule about two times longer than
wide, the lateral and ventral surfaces are flat or only slightly convex,
posterior surface slightly concave. Almost the entire capsule surface is
covered with short moss-like pinnae which are slightly longer along the ribs
of the capsule, and less prominent on the dorsal surface around the
micropylar plate.Capsule lateral and ventral surfaces are set with two
parallel lines of small circular pits running the length of the capsule near the
longitudinal margins of each surface. Each row of circular pits have four to
six pits (the number of which is almost always mirrored on the parallel
running row on the same surface).Micropylar plate long and thin, running
about 80 to 90% of the length of the dorsal surface with the widest portion
around the micropylar cup which is situated around the posterior third of the
capsule with the remainder of the micropylar plate thinner than the
micropylar cup width. Operculum roundly conical, circular in cross-
section with several pits of a similar size and shape to those found along the
longitudinal surfaces of the egg.Coloration caramel to tawny brown, the
moss-like pinnae of a slightly paler brown than the capsule itself.
Measurements [mm].
lateral length (including operculum): 4.2
lateral length (excluding operculum): 3.7
width of dorsal surface: 2.6
width of lateral surface: 3.0
length of micropylar plate: 2.8
Distribution and habitat. – To date this species has only been
confirmed from the Hau Phan Province of Northeastern Laos, although
this species can likely be found elsewhere in nearby mountain ranges or
provinces. This region is classified by the Köppen-Geiger climate
classification index as a temperate region with a dry winter and a warm
to hot summer, with rainfall ranging from only a few mm in the dry
season to 200+ mm or more in the wet season. Average elevation in the
area for this new species was found to range from 850 to 2,060 meters
and is widely lush with thick foliage (Fig. 8, 10).
Steeve Collard was also kind enough to share with us photos of the
local host plant that wild specimens of Phyllium oyae n. sp. have been
found on. Two of which were identified to the genus were
Castanopsis sp. and Quercus sp., both of which are in the beech tree
family Fagaceae (Fig. 9A-B).
Etymology. – Patronym, named after the wife of Steeve Collard, Mme
Oy Houngsavath. Steeve has been a very important collaborator of the
Montreal Insectarium (Canada) for many years. He is now established
permanently in Laos and supplies many important new species to
various specialists. He is also trying to establish an Insectarium in Laos,
as well as helping to create a butterfly farm. His presence in the
contemporary entomology community of Laos is prominent.
Note on the unavailable name Phyllium rayongii
Thanasinchayakul, 2006
The species name Phyllium rayongii Thanasinchayakul, 2006 is
determined to be a nomen nudum and therefore unavailable according to
ICZN Article 16.4.1. (ICZN, 1999). Article 16.4.1 requires the “explicit
fixation of a holotype, or syntypes, for the nominal taxon” (ICZN, 1999), a
simple yet absent requirement from Sorpongpaisal & Thanasinchayakul
(2006). To date, no authors have validated the name Phyllium rayongii by
providing a holotype designation, and therefore the name is unavailable.
Hopefully, future authors working on the Phylliidae of Thailand will
review specimens from throughout Thailand to determine if the
population referenced in Sorpongpaisal & Thanasinchayakul (2006) is in
fact a valid taxon. With the original description of Phyllium rayongii
vague and describing features which apply to nearly all Phylliidae species,
we presently see no reason to believe this population is novel and
speculate it might simply be a southern range expansion of Phyllium
westwoodii.
1. Tegmina long, reaching at least past the anterior margin of abdominal segment IV ..................................................... 2
Tegmina short, only reaching into abdominal segment II or III ............................................................................. 3
2(1). Profemoral exterior lobe that is thinner and smoothly arcing from end to end, only about three times the width of the profemoral
shaft itself; Yunnan Prov ........................................................................................ Ph. (Ph.) yunnanense Liu, 1993
Profemoral exterior lobe that is wider with a maximum width up to five times the width of the profemoral shaft and with a
distinct but smooth angle in the center of the span; Laos ............................................................... Ph. (Ph.) oyae n. sp.
3(1). Profemoral exterior lobe notably wider than the interior lobe; exterior lobe with almost a right angle; Indonesia (Sulawesi)
.................................................................................................................... Ph. (Ph.) celebicum de Haan, 1842
Profemoral exterior lobe the same width as the interior lobe; exterior lobe with a notably obtuse angle ............................. 4
4(3). Mesopraescutum surface black in color; Philippines (Luzon Is., Batan Is., Marinduque Is., Catanduanes Is.) ........................
........................................................................................................... Ph. (Ph.) ericoriai Hennemann et al., 2009
Mesopraescutum surface of a similar color to that of the surrounding area, not black .................................................. 5
5(4). Triangular protibial interior lobe an isosceles triangle, with the widest point in the center of the span; China: Hainan Province,
Daling-Baisha ......................................................................................................... Ph. (Ph.) parum Liu, 1993
Triangular protibial interior lobe a scalene triangle with the widest portion on the distal third, not in the center of the span ................. 6
6(5). Profemoral exterior lobe about two to two and a half times the width of the profemoral shaft; Philippines (Northern Luzon Is.) ........
........................................................................................................... Ph. (Ph.) bonifacioi Lit and Eusebio, 2014
Profemoral exterior lobe three or more times the width of the profemoral shaft ......................................................... 7
7(6). Mesopraescutum surface long and thin, about the length of the pronotum; Singapore ............ Ph. (Ph.) chrisangi Seow-Choen, 2017
Mesopraescutum surface thin but not as long as the pronotum, slightly shorter in length; Myanmar, Thailand, Laos ..............
............................................................................................................. Ph. (Ph.) westwoodii Wood-Mason 1875
Key to known males in the Phyllium (Phyllium) celebicum species group
Phyllium tibetense, Phyllium athanysus, Phyllium rarum, Phyllium drunganum, and Phyllium yapicum unknown/ undescribed and
are therefore excluded.
CUMMING & LE TIRANT. – Phyllium oyae n. sp. of Laos
8
Faunitaxys, 8(4), 2020 : 1 – 9. 9
Résumé
Cumming R. T. & Le Tirant S., 2020. Description d’un nouveau Phyllium Illiger, 1798 du groupe celebicum, nati ve d u Laos (Ph amida : Phylliid ae).
Faunitaxys, 8(4) : 1 – 9.
L’ é tu d e d ’ u n l o t d ’ i n se c t e s f e u i l le s , c o l l ec t é s d a ns l e n o r d - es t d u L a o s , a r é v é l é l a p r é s e nc e d ’ u n e e s p èc e n o u v e ll e , q u e n o u s d é c r iv o n s s o us l e
nom de Phyllium (Phyllium) oyae n. sp. Elle fait partie du groupe de celebicum suivant Hennemann et al. (2009), en raison de la présence d'ailes
bien développées chez les femelles et d’un large lobe profémoral extérieur chez les mâles. Nous constatons ensuite que Phyllium (Phyllium)
drunganum Yang, 1995 est mal placé, et nous le transférons du groupe siccifolium au groupe celebicum, en raison de la présence d'ailes
développées chez les femelles. Puis nous différencions Phyllium oyae n. sp. de toutes les autres espèces. Enfin nous considérons Phyllium
rayongii T h a n a s i n c h a y a k u l , 2 0 0 6 c o m m e é t a n t u n nomen nudum et n 'es t do nc pa s dis po ni ble s elo n l' art ic le 16 .4 .1 de l 'IC ZN . Pou r co ncl ur e, no us
présentons des clés dichotomiques pour les mâles et les femelles des espèces du groupe celebicum connues à ce jour.
Mots-clés. – Phasmatodea, Phasmida, Phylliidae, Phylliini, Phyllium, Asie du Sud-Est, Laos, oyae, celebicum, taxonomie, description, espèce nouvelle.
Acknowledgments
The authors thank René Limoges, entomological technician at the
Montreal Insectarium, Canada for taking many photos for this
work, as well as for many professional courtesies. We thank Steeve
Collard for sending us many Phyllium over the years for study, as
well as the first series of this new species, photos of the host
plants, eggs, habitat and live specimens in the wild. Steeve Collard
wants to thank brothers, Adrien Doré and Gabriel Doré who have
helped him in all of his entomological expeditions in Laos since
2007. Thanks to Johnson Sau (Malaysia) for also sending us a fine
series of quality specimens of this new species. Special thanks to
our two peer reviewers Frank Hennemann (Germany) and Dr
Allan Taylor (Canada) for their quality feedback on this paper.
Additional thanks to Doug Yanega (USA) commissioner for the
International Commision on Zoological Nomenclature (ICZN) and
James Carpenter (USA) who kindly spent time reviewing our
presented nomen nudum case for Phyllium rayongii, for sharing
their knowledgeable opinions in regards to the ICZN code. Thank
you to Connie Cumming (USA) for reviewing the first draft of this
work. Thanks also to Binjie Ge (Shanghai Chenshan Plant
Research Center) and Gilles Vincent Special Advisor of the
Executive President of the Chenshan Botanical Garden-Shanghai,
China for the identification of the plant.
References
Chen S. C., 1999. – Phasmatodea. In: Chen, S. C. [Ed.]: Pictorial Handbook of
Rare and Precious Insects in China. China Forestry Publications House,
Beijing. 332 pp.
Chen S. C. & He Y. H., 2008. – Phasmatodea of China. China Forestry
Publishing House, Beijing, China. 476 pp.
Cumming R. T. & Teemsma S. N., 2018. A new species of Phyllium
(Phyllium) from Micronesia, Yap Island (Phasmida: Phylliidae)
representing a range expansion for the family. Insecta Mundi 0650: 1-9.
Größer D., 2008. – Wandelnde Blätter. Ein Katalog aller bisher beschriebenen
Phylliinae- Arten und deren Eier mit drei Neubeschreibungen. 2nd Edition.
Edition Chimaira, Frankfurt am Main, 175 pp.
Haan W. de, 1842. – Bijdragen tot de Kennis der Orthoptera. In Verhandelingen
over De Natuurlijke Geschiedents der Nederlandsche Overzeesche
Bezittingen, Leiden p. 45-248.
Hennemann F. H., Conle O. V., Gottardo M. & Bresseel J., 2009. 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 Palawan (Phasmatodea: Phylliidae: Phylliinae:
Phylliini). Zootaxa, 2322, 1-83.
Hennemann F. H., Conle O. V. & Zhang W., 2008. – Catalogue of the Stick
and Leaf-insects (Phasmatodea) of China, with a faunistic analysis, review
of recent ecological and biological studies and bibliography (Insecta:
Orthoptera: Phasmatodea). Zootaxa 1735: 1-77.
ICZN, 1999. International Code of Zoological Nomenclature. Fourth
Edition, adopted by the International Union of Biological Sciences.
International Trust for Zoological Nomenclature, London, 306 pp.
Illiger J. C. W., 1798. Ve r s u c h e e i n e r n a t u r l i c h e n O r d n u n g s - u n d G a t t u n g s
Folge der Inseken. In: Ve r z e i c h n i s d e r K ä f e r P r e u s s e n s . Halle, Johann
Jacob Gebauer.: 487-501 [499].
Lit I. & Eusebio O., 2014. – A new species of leaf insect (Phasmatodea:
Phylliidae) from Northern Luzon, Philippines. Arthropoda Generalis:
ISSN 2191-4427.
Liu, S. L., 1993. A preliminary study of the genus Phyllium Illiger of China
(Phasmatodea: Phylliidae). Acta Zootaxonomica Sinica, 18(2), 201-212.
Otte & Brock., 2005. – Phasmida Species File. Catalog of Stick and Leaf
Insects of the world 273
Seow-Choen F., 2017. – A taxonomic guide to the stick insects of
Singapore. Natural History Publications (Borneo); Kota Kinabalu. 175 p.
Sorpongpaisal W. & Thanasinchayakul S., 2006. – Identification of some
stick and leaf insects, (order Phasmida) in Thailand. Kamphaengsean
Academic Journal 4: 18-32.
Wes t wo od , J .O ., 1 85 9. Catalogue of Orthopterous insects in the
collection of the British Museum. Part 1, Phasmidae. British Museum,
London, 195 pp., 40 plates.
Woo d -M as on J. , 18 7 5. – On new or little-known species of Phasmidae,
with a brief preliminary notice of the occurrence of a clasping
apparatus in the males throughout the family. Journal of the Asiatic
Society Bengal, 44(3), 215–220, plates 16-17.
Ya ng J . K. , 1 9 9 5. – Phyllium drunganum Ya ng a ne w s pe c i e s o f L e a f I n s ec t
from Yunnan China (Phasmatodea: Phyllidae). Journal of the Yunnan
Agricultural University, 10(1), 18-22.
Zompro O., 2004. Revision of the genera of the Areolatae, including the
status of Tim em a and Agathemera (Insecta, Phasmatodea). Abhandlungen
des Naturwissenschaftlichen Vereins in Hamburg. Antiquariat Goecke &
Evers, Keltern-Weiler, Germany. (NF) 37. 327 pp.
Fig. 10. Habitat in Hua Phan Province where Steeve Collard frequently
collects various insects.
Faunitaxys
Volume 8, Numéro 4, Février 2020
Illustration de la couverture: Forêt du Laos.
Cover photo: Forest of Laos.
Crédits:
© René Limoges, Insectarium de Montréal, Québec, Canada : Fig. 3C, 4, 5, 6, 7.
© Steeve Collard, Laos : Fig. 1, 2, 8, 9, 10 & couverture.
© Frank Hennemann, Germany : Fig. 3A, 3B.
Publié par l’Association Française de Cartographie de la Faune et de la Flore (AFCFF)
Description d’un nouveau Phyllium Illiger, 1798 du groupe celebicum, native du Laos (Phamida : Phylliidae).
Royce T. Cumming & Stéphane Le Tirant ........................................................................................................ 1 – 9
SOMMAIRE
CONTENTS
A n ew s pecie s of Phyllium Illiger, 1798, from the celebicum species group native to Laos (Phasmida: Phylliidae).
Royce T. Cumming & Stéphane Le Tirant ........................................................................................................ 1 – 9
ResearchGate has not been able to resolve any citations for this publication.
Article
Full-text available
A new species of leaf insect from the celebicum species group, Phyllium (Phyllium) yapicum Cumming and Teemsma, new species (Phasmida: Phylliidae), is described from a female specimen from the California Academy of Sciences collection, United States. This new species is the first recorded species of Phylliidae from the country of Micronesia and represents a notable range expansion for the family. With Phyllium (Phyllium) yapicum Cumming and Teemsma, new species, currently only known from a female holotype; a key to females is included for the celebicum species group.
Article
Full-text available
A complete taxonomic catalogue of the Stick and Leaf-insects (Phasmatodea) recorded or described from the mainland China (excluding Taiwan) is presented. 241 valid species are listed, which are currently attributed to 50 genera, 5 families and 7 subfamilies. Genera and species are listed alphabetically. All available type-data is provided based mainly on literary sources for species described by Chinese workers from 1986 to 2006, including documented depository of type-specimens. The catalogue therefore also provides complete lists of the type-material of Phasmatodea housed in the following Chinese institutions: Administration of Baishuijiang Natural Reserve (ABNR), Beijing Forestry University, Beijing (BFU), China Agricultural University, Beijing (CAU), Geological Museum of China, Beijing (GMC), Inca Science Ltd., Chongqing (INCA), Institute of Zoology, Chinese Academy of Sciences, Beijing (IZCAS), Department of Biology, Nankai University, Tianjin (NKU), Northwest Sci-Tech University of Agriculture and Forestry, Shaanxi (NWSUAF), Institute of Zoology, Shaanxi Normal University, Xi’an (SNU), Institute of Entomology, Sun Yat-sen University (SYSU), Shanghai Institute of Entomology, Academia Sinica, Shanghai (SIES), Tianjin Natural History Museum, Tianjin (TJNHM), Zhejiang Museum of Natural History, Hangzhou (ZJMNH). The known distribution of each species, in means of provinces is provided as well. 14 species are shown to have been recorded from China in error, several of these based on misidentifications. The “Phasmatodea-like” fossil taxa described from the the Late Jurassic Yixian Formation of North Hebei and West Liaoning are listed in a separate section. Two new generic synonyms are recognized: Arthminotus Bi, 1995 synonymised with Lopaphus Westwood, 1859 (n. syn.) and Dianphasma Chen & He, 1997 synonymised with Parasosibia Redtenbacher, 1908 (n. syn.). The genus Linocerus Gray, 1835 (Type-species: Linocerus gracilis Gray, 1835) was erroneously synonymised with the mediterranean Bacillus St. Fargeau & Audinet-Serville, 1825 and is here re-established in Phasmatidae: Pachymorphinae: Gratidiini (rev. stat.). Relationship to Clonaria Stål, 1875 (= Gratidia Stål, 1875, = Paraclonaria Brunner v. Wattenwyl, 1893), Sceptrophasma Brock & Seow-Choen, 2000 and Macellina Uvarov, 1940 is obvious. 13 species are transferred to other genera (new combinations): Asceles dilatatus Chen & He, 2004 and Asceles quadriguttatus Chen & He, 1996 to Pachyscia Redtenbacher, 1908, Arthminotus sinensis Bi, 1995 to Lopaphus Westwood, 1859, Baculum dolichocercatum Bi & Wang, 1998a and Baculum politum Chen & He, 1997 to Medauroidea Zompro, 1999, Dixippus bilippus Chen & He, 1999, Dixippus hainanensis Chen & He, 2002, Dixippus nigroantennatus Chen & He, 2002, Dixippus parvus Chen & He, 2002 and Entoria bobaiensis Chen, 1986 to Lonchodes Gray, 1835, Sipyloidea obvius Chen & He, 1995 to Sinophasma Günther, 1940, and Gratidia bituberculata Redtenbacher, 1889 and Leptynia xinganensis Chen & He, 1993 to Sceptrophasma Brock & Seow-Choen, 2002. Acrophylla sichuanensis Chen & He, 2001 remains of unknown generic assignment, but is shown to be not a member of the Australian genus Acrophylla Gray, 1835. Furthermore, as Baculum Saussure, 1861 is a neotropical genus and most Old World species previously attributed this genus are now listed in Ramulus Saussure, 1861, all Chinese species described in this genus are consequently transferred to Ramulus Saussure. Other changes of specific placements are based on published literature and concern to the following three synonymies not recognized by Chinese workers: Abrosoma Redtenbacher, 1906 (= Prosceles Uvarov, 1940), Necroscia Audinet-Serville, 1838 (= Aruanoidea Redtenbacher, 1908), Lopaphus Westwood, 1859 (= Paramyronides Redtenbacher, 1908). Megalophasma Bi, 1995 is transferred from Necrosciinae to Lonchodinae. Four lectotypes are designated and three new specific synonyms revealed. A lectotype is designated for Rhamphophasma modestus Brunner v. Wattenwyl, 1893, the type-species of Rhamphophasma Brunner v. Wattenwyl, 1893, in order to fix this genus and species. The male paralectotype is shown to be a male of Parapachymorpha nigra Brunner v. Wattenwyl, 1893, the type-species of Parapachymorpha Brunner v. Wattenwyl, 1893. Clitumnus porrectus Brunner v. Wattenwyl, 1907 is synonymised with Bacillus ? artemis Westwood, 1859 with a lectotype designated for the former (n. syn.). A lectotype is designated for Oxyartes lamellatus Kirby, 1904. Paracentema stephanus Redtenbacher, 1908 is shown to have been erroneously synonymised with Neohirasea japonica (de Haan, 1842) and here synonymised with Neohirasea maerens (Brunner v. Wattenwyl, 1907) (n. syn.). In order to fix the new synonymy a lectotype is designated for Paracentema stephanus Redtenbacher, 1908. Finally, a biogeographic analysis of the Chinese phasmid fauna is presented. This includes brief background information on the topography and biogeography of China along with maps showing the seven zoogeographical subregions currently recognized as well as the 4 municipalities, 23 provinces, 5 autonomous regions and 2 special administrative regions of China. A summary of the taxonomic compilation of the fauna is provided and its relationships with neighbouring regions, of both the Palearctic and Oriental realms, are discussed. A study is presented on the distribution of the taxa and species densities of each province / autonomous region.
Article
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.
Pictorial Handbook of Rare and Precious Insects in China. China Forestry Publications House
  • S C Chen
Chen S. C., 1999. -Phasmatodea. In: Chen, S. C. [Ed.]: Pictorial Handbook of Rare and Precious Insects in China. China Forestry Publications House, Beijing. 332 pp.
Wandelnde Blätter. Ein Katalog aller bisher beschriebenen Phylliinae-Arten und deren Eier mit drei Neubeschreibungen
  • D Größer
Größer D., 2008. -Wandelnde Blätter. Ein Katalog aller bisher beschriebenen Phylliinae-Arten und deren Eier mit drei Neubeschreibungen. 2nd Edition. Edition Chimaira, Frankfurt am Main, 175 pp.
Bijdragen tot de Kennis der Orthoptera
  • W Haan
  • De
Haan W. de, 1842. -Bijdragen tot de Kennis der Orthoptera. In Verhandelingen over De Natuurlijke Geschiedents der Nederlandsche Overzeesche Bezittingen, Leiden p. 45-248.
International Code of Zoological Nomenclature. Fourth Edition, adopted by the International Union of
ICZN, 1999. -International Code of Zoological Nomenclature. Fourth Edition, adopted by the International Union of Biological Sciences. International Trust for Zoological Nomenclature, London, 306 pp.
Versuche einer naturlichen Ordnungs-und Gattungs Folge der Inseken
  • J C W Illiger
Illiger J. C. W., 1798. -Versuche einer naturlichen Ordnungs-und Gattungs Folge der Inseken. In: Verzeichnis der Käfer Preussens. Halle, Johann Jacob Gebauer.: 487-501 [499].
A new species of leaf insect (Phasmatodea: Phylliidae) from Northern Luzon
  • I Lit
  • O Eusebio
Lit I. & Eusebio O., 2014. -A new species of leaf insect (Phasmatodea: Phylliidae) from Northern Luzon, Philippines. Arthropoda Generalis: ISSN 2191-4427.