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Article Leaf insects from Luzon, Philippines, with descriptions of four new species, the new genus Pseudomicrophyllium, and redescription of Phyllium (Phyllium) geryon Gray, 1843, (Phasmida: Phylliidae) Associate researcher with the Montreal Insectarium (Quebec-Canada)

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Abstract and Figures

Examination of unidentified Phylliidae specimens revealed a number of undescribed species from the island of Luzon, Philippines. Morphological and molecular study of specimens from the obscure phasmid genus Microphyllium Zompro, 2001, revealed a new species, which we describe as Microphyllium haskelli Cumming sp. nov.. It is here described and differentiated from the two other species in the genus, both currently only known from adults of a single sex. Pseudomi-crophyllium Cumming gen. nov. is described as a new genus within Phylliidae with the type species Pseudomicrophyllium faulkneri Cumming gen. et sp. nov. as the sole known species in the genus. As is unfortunately often the case in the leaf-mimicking family Phylliidae, this new genus and species is only known from a single specimen. In addition to the new genus, two new Phyllium (Phyllium) species from the siccifolium species-group are named and described as Ph. (Ph.) an-tonkozlovi Cumming sp. nov. and Ph. (Ph.) bourquei Cumming & Le Tirant sp. nov.. In addition to the newly described species, Phyllium (Phyllium) geryon Gray, 1843 is redescribed from a nearly perfect specimen, completing some of the morphological knowledge gaps currently missing because of the severely damaged holotype specimen. A key to all known species of Phylliidae from Luzon is included. Holotype specimens for all four new species will be deposited in the National Museum of the Philippines type collection and paratype specimens will be deposited into the San Diego Natural History Museum collection or retained within the first author's collection.
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Accepted by B. Mantovani: 3 Nov. 2017; published: 18 Nov. 2017
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ZOOTAXA
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https://doi.org/10.11646/zootaxa.4365.2.1
http://zoobank.org/urn:lsid:zoobank.org:pub:B166473D-1089-4DD2-866B-9339D152E616
Leaf insects from Luzon, Philippines, with descriptions of four new species,
the new genus Pseudomicrophyllium, and redescription of
Phyllium (Phyllium) geryon Gray, 1843, (Phasmida: Phylliidae)
ROYCE T. CUMMING
1,5,6
, JING V. LEONG
2
& DAVID J. LOHMAN
2,3,4
1
Entomology Department, San Diego Natural History Museum, POB 121390, Balboa Park, San Diego, California, 92112-1390 USA
2
Biology Department, City College of New York, City University of New York, 160 Convent Ave. New York, New York, United States.
10031
3
Biology Ph.D. Program, Graduate Center, City University of New York, 365 Fifth Ave. New York, New York, 10016, United States
4
Entomology Section, National Museum of the Philippines, Manila, 1000, Philippines
5
Associate researcher with the Montreal Insectarium (Quebec-Canada)
6
Corresponding author. E-mail: phylliidae.walkingleaf@gmail.com
Abstract
Examination of unidentified Phylliidae specimens revealed a number of undescribed species from the island of Luzon,
Philippines. Morphological and molecular study of specimens from the obscure phasmid genus Microphyllium Zompro,
2001, revealed a new species, which we describe as Microphyllium haskelli Cumming sp. nov.. It is here described and
differentiated from the two other species in the genus, both currently only known from adults of a single sex. Pseudomi-
crophyllium Cumming gen. nov. is described as a new genus within Phylliidae with the type species Pseudomicrophyllium
faulkneri Cumming gen. et sp. nov. as the sole known species in the genus. As is unfortunately often the case in the leaf-
mimicking family Phylliidae, this new genus and species is only known from a single specimen. In addition to the new
genus, two new Phyllium (Phyllium) species from the siccifolium species-group are named and described as Ph. (Ph.) an-
tonkozlovi Cumming sp. nov. and Ph. (Ph.) bourquei Cumming & Le Tirant sp. nov.. In addition to the newly described
species, Phyllium (Phyllium) geryon Gray, 1843 is redescribed from a nearly perfect specimen, completing some of the
morphological knowledge gaps currently missing because of the severely damaged holotype specimen. A key to all known
species of Phylliidae from Luzon is included. Holotype specimens for all four new species will be deposited in the Na-
tional Museum of the Philippines type collection and paratype specimens will be deposited into the San Diego Natural
History Museum collection or retained within the first author’s collection.
Key words: DNA barcodes, Philippines, Luzon, new species, Phylliinae, holotype, Phyllium, Mountain Province, leaf in-
sect
Introduction
Leaf insects in the family Phylliidae are camouflaged by resembling the color, shape, and venation of
dicotyledonous leaves. This remarkable crypsis presumably reduces predation by visually oriented predators, but
also thwarts attempts to study them in the tropical Asian forests where they live. Many species are known from just
a handful of specimens, and adult specimens of at least one sex are currently unknown for many species.
Specimens in the Philippine-endemic phylliid genus Microphyllium are rarely encountered. The species pusillulum
was described from a single adult female by James A. G. Rehn and John W. H. Rehn (1933), and placed in the
genus Phyllium, making it the smallest species in that taxon. Several decades later, Oliver Zompro (2001) erected
the genus Microphyllium with M. spinithorax as its sole species based on the adult male holotype and two nymph
female paratypes. When Hennemann et al., (2009), reexamined Rehn & Rehn’s small pusillulum holotype, they
transferred the species to the genus Microphyllium based on similarities with the M. spinithorax female nymph
paratypes designated by Zompro in 2001. Hennemann et al., (2009) also hypothesized that M. pusillulum and M.
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spinithorax might be the same, sexually dimorphic species known only from adult female and male specimens,
respectively. Thus, the female nymphs designated by Zompro as M. spinithorax might be M. pusillulum nymphs.
The morphology of an insect changes drastically during its development from egg to adult, and sexual
dimorphism of adult Phylliidae can be pronounced. However, protein-coding mitochondrial DNA sequences are
relatively constant within species but variable enough between species to be a useful tool for associating different
sexes and life stages of a single species (Wells et al., 2001). To clarify the species-level status of Microphyllium
pusillulum and M. spinithorax and to associate immature stages (nymphs) with adults of these taxa, we sequenced
mitochondrial DNA barcodes from six Microphyllium specimens collected on Luzon, Philippines, of varying
developmental stages and sexes, including one confidently identified adult female M. pusillulum.
Abbreviations Used:
ANSP Academy of Natural Sciences, Drexel University, Philadelphia: United States
BMNH Natural History Museum, London: England
SDNHM San Diego Natural History Museum, San Diego, California: United States
PNM National Museum of the Philippines, Manila: Philippines
[Coll. RC] Private collection of Royce Cumming, California: United States
HT Holotype
PT Paratype
M. Microphyllium Zompro, 2001
Ph. (Ph.) Phyllium (Phyllium) Illiger, 1798
Ps. Pseudomicrophyllium Cumming gen. nov.
Materials and methods
Morphological Examination. Morphological evaluations of all specimens in first author’s collection were made
using a Leica ZOOM 2000 microscope. Photos of the material from the first author’s collection were taken by
Danny Burk or by the first author. Photos taken by Danny Burk were done using a Canon 5DS R digital camera
with a 65mm 1–5x macro lens and were focus stacked using photo stacking software. Photos taken by the first
author were taken using a Canon 5D Mark II and a MP-E 65mm macro lens and stacked using Zerene photo
stacking software, version 1.04, 64-bit. Photos of the material from the collection of Stéphane Le Tirant were 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 of the holotype M. pusillulum were taken by Jason Weintraub at ANSP using an Olympus OM-D
E-M10 Micro 4/3 digital camera equipped with an Olympus M.Zuiko Digital ED 60mm f/2.8 macro lens.
Measurements of anatomical figures were made to the nearest 0.1 mm using digital calipers. To identify the
putative M. pusillulum female with confidence, the first author visited the ANSP to compare morphology of the
holotype to the specimen in question. Morphological comparison between the field-collected Microphyllium males
to the M. spinithorax holotype could not be done in person. Instead, the male specimens were compared with the
original description by Zompro (2001) and the genitalia illustration in Wedmann et al. (2007). Morphological
comparison of the Phyllium (Phyllium) geryon holotype could not be done in person, but fortunately detailed
photos of the specimen are available online at the Phasmida Species File http://phasmida.speciesfile.org (author:
Paul D. Brock).
DNA Extraction and Sequencing. We did not attempt to incorporate any type specimens of Microphyllium in
our molecular analysis, as their DNA was probably too degraded for PCR amplification because of their age. Only
specimens from the first author’s collection were used, as they are only around three years old. A single leg was
removed from each dried specimen and stored at room temperature in the dark prior to analysis. DNA was
extracted from each leg using a CTAB phenol-chloroform extraction protocol. A 658 bp fragment of the
mitochondrial gene cytochrome c oxidase subunit I (COI) was amplified from whole genomic extracts using the
universal primers LCO1490 (5’-TAATACGACTCACTATAGGGTCWACWAATCATAAAGATATTGG-3’) and
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HCO2198 (5’-ATTAACCCTCACTAAAGTAAACTTCAGGGTGACCAAAAAATCA-3’). These primer
sequences include the original primers of Folmer et al., (1994) with T7 and T3 tails concatenated on the 5’ end of
LCO1490 and HCO2198 respectively (“tails” indicated in bold; Wahlberg & Wheat, 2008). Addition of these
primer tails increases PCR success for specimens with degraded DNA (Regier & Shi, 2005). Each PCR reaction
consisted of 2.5μl 5x GoTaq® Flexi buffer, 2.5 μl of 25 mM MgCl
2
, 0.5 μl of 10mM dNTPs, 1.25 μl of 10mM for
each primer, 0.125 μl 5U/μl of TaKaRa polymerase, 15.58 μl of H
2
O, and 1.3 μl template DNA for a total volume
of 25 μl per reaction which was amplified under the following thermal cycler conditions: 95° C for 7 min, followed
by 40 cycles at 95° C for 30 sec, 56.5° C for 90 sec and 72° C for 2 min, and 72°C for 10 min. PCR products were
visualized on agarose gels before being sent for sequencing to Macrogen (macrogenusa.com) for EXO-SAP PCR
clean-up and bidirectional sequencing. We sequenced the COI barcode from six leaf insect specimens and included
a COI sequence of Phyllium (Pulchriphyllium) giganteum as an outgroup (GenBank accession AB477461; Kômoto
et al., 2011).
Phylogenetic Analysis. Alignment of nucleotide sequences were performed using MUSCLE (Edgar 2004)
implemented in Seaview 4.2.4 (Gouy et al., 2010) and sequences were assembled and trimmed with Sequencher
5.0 (genecodes.com).
jModelTest 2.1.5 (Darriba et al., 2012) selected the GTR+G model of sequence evolution using the AICc
criterion. Bayesian phylogenetic analysis was performed with MrBayes 3.2.6 on the XSEDE server running
BEAGLE (Ronquist et al., 2012) via the CIPRES Science Gateway (phylo.org; Miller et al., 2010). Four Markov
chains, one cold and three heated, were run simultaneously for 5 million generations. Trees were sampled every
1000 generations, and the first 25% of sampled trees were discarded as burn-in before calculating a consensus tree.
To assess parsimony support for relationships among taxa, TNT 1.1 (Goloboff, et al., 2008) was implemented
to run 1000 bootstrap replicates using symmetric resampling summarized as absolute frequencies (Goloboff et al.,
2003). PAUP* 4.0a152 (Swofford, 2002) was used to calculate uncorrected pairwise distances between COI
sequences.
Results
Microphyllium Zompro, 2001. Morphological examination of the M. pusillulum holotype confirmed the
identification of the unknown specimen [Coll. RC 16-096] as M. pusillulum on morphological grounds, though the
two specimens were slightly different in length (Table 3). Numerous morphological differences were found
between the Microphyllium males in the first author’s collection and descriptions of the M. spinithorax holotype
described in the literature (Table 5).
Analyses of DNA sequence data suggest that five of the six specimens examined are a single species distinct
from M. pusillulum. Moreover, the morphology of the three male specimens differs from M. spinithorax, indicating
that these individuals belong to a new Microphyllium species, which we describe below as M. haskelli. We were
therefore unable to test the hypothesis that M. pusillulum and M. spinithorax are conspecific, nor could we identify
a M. pusillulum male. Indeed, our results raise an additional question: are the two female nymphs designated M.
spinithorax paratypes by Zompro (2001) in fact M. spinithorax, M. haskelli, or—as suggested by Hennemann et
al.—nymphs of M. pusillulum? Microphyllium pusillulum (Rehn & Rehn, 1933) has a wide distribution across the
Cordillera Central mountain range of Luzon, and it would not be surprising if M. spinithorax was synonymous.
However, this decision must be withheld until fresh material of M. spinithorax can be studied.
Uncorrected pairwise distances between a single specimen of Phyllium (Pulchriphyllium) giganteum (Kômoto
et al., 2011) (selected as an outgroup) and five Microphyllium haskelli (Table 1) ranged from 18.8% to 21.3%.
Distances between a single M. pusillulum, and five M. haskelli ranged from 15.5% to 18.8%, while distances
among the five M. haskelli ranged from 0% to 2.2% (Table 1). Phylogenetic trees constructed using parsimony and
Bayesian optimality criteria were completely congruent and highly supported (Fig. 1). The use of mitochondrial
DNA barcodes in taxonomy has been criticized, and there are several situations in which barcode variation or lack
thereof can be misleading, including heteroplasmy, introgression between species, introgression of mitochondrial
genes into the nuclear genome (numts), and infection by Wolbachia or other intracellular bacterial parasites (Hurst
& Jiggins, 2008; Kodandaramaiah et al., 2013; Rubinoff et al., 2006; Song et al., 2008). However, these criticisms
are not particularly relevant here because it is unlikely that all five Microphyllium haskelli and the single M.
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pusillulum samples have numts, since all sequences could be translated into amino acids without stop codons, and
because Wol b a c h i a is unknown within Phasmida (Pérez-Ruiz et al., 2015; Sven Bradler, pers. comm. April, 2017).
TABLE 1. Uncorrected pairwise distances between COI barcodes amplified from three phylliid species show
intraspecific genetic variation of <2% within the M. haskelli sampled, but distances over 15% between different species.
The Phyllium giganteum sequence, referenced by its accession number, was obtained from NCBI (GenBank;
ncbi.nlm.nih.gov).
FIGURE 1. Gene tree of COI barcode sequences from three phylliid species. The inferred relationships are strongly
supported, as indicated by parsimony bootstrap and Bayesian posterior probability (PP) support. The species, gender,
collection code, and collection locality within Luzon, Philippines, are indicated for each Microphyllium specimen.
Pseudomicrophyllium Cumming gen. nov. For several years, specimen RC 16-129 was stored papered in the
Microphyllium tray of the first author’s collection because of its small size. It was not until the specimen was
pinned that its uniqueness became evident. Its antennae, thorax, and abdomen are shaped like a Phyllium
(Phyllium) male, but its size and legs were more akin to Microphyllium. The unique male instead represents an
undescribed genus. Pseudomicrophyllium Cumming gen. nov. is here established with a single species and placed
within the tribe Phylliini due to its similarity to both Phyllium Illiger, 1798 and Microphyllium Zompro, 2001.
Phyllium (Phyllium) Illiger, 1798. With this work focusing on the island of Luzon, three Phyllium (Phyllium)
species are discussed, two of which are new species. The two new species appear closely related to Phyllium
(Phyllium) philippinicum Hennemann et al., 2009 in many aspects. From Isabela Province, Phyllium (Phyllium)
antonkozlovi Cumming sp. nov. males can immediately be differentiated by their more robust antennae. The
female is currently unknown but is assumed to look similar to Phyllium (Ph.) philippinicum because of the size of
the male and the shared morphological similarities between the two species.
Phyllium (Phyllium) bourquei Cumming & Le Tirant sp. nov. is described as a new species from Nueva
Vizcaya province. It was initially thought to be a new population of Phyllium (Ph.) philippinicum because of its
similar size and profemora. However, females can easily be differentiated by their shorter subgenital plate only
(1) (2) (3) (4) (5) (6) (7)
(1) M. haskelli
[Coll. RC 16-097]
-
(2) M. haskelli
[Coll. RC 16-098]
0.00304 -
(3) M. haskelli
[Coll. RC 16-099]
0.0216 0.0183 -
(4) M. haskelli
[Coll. RC 16-128]
0.0216 0.0183 0 -
(5) M. haskelli
[Coll. RC 16-183]
0 0.00304 0.0216 0.0216 -
(6) M. pusillulum
[Coll. RC 16-096]
0.161 0.158 0.155 0.155 0.161 -
(7) P. giganteum
(AB477461)
0.214 0.213 0.213 0.213 0.214 0.188 -
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reaching about half way under the anal abdominal segment (almost reaching the apex of the anal abdominal
segment in Ph. (Ph.) philippinicum) and the margins of the mesopleurae, which are more heavily marked by robust
tubercles (Figs. 18A&B). In Ph. (Ph.) philippinicum the mesopleurae margins are only marked by three prominent
tubercles in the center and a few minor tubercles anteriorly and posteriorly (Fig. 18C). Males can be differentiated
by the mesopraescutum with a distinct crest along the sagittal plane with nodes that steadily decrease in size (Fig.
17B) (vertex of the mesopraescutum almost completely smooth in Ph. (Ph.) philippinicum (Fig. 17C)). These two
new species represent the fifth and sixth described species of Phyllium (Phyllium) recorded from Luzon to date.
In addition, a small Phyllium (Phyllium) specimen from eastern Luzon was analyzed and found to be the
elusive Phyllium (Phyllium) geryon Gray, 1843. This species has not been seen in over a century, and because of its
brief original description and severely damaged holotype, a redescription of the species is presented. The holotype
is missing many morphological features commonly used for identification including antennae, forelegs, and
midlegs; unique features of the thorax were therefore most useful for comparison. These features, coupled with the
size of the specimen, length of tegmina, genitalia, and general body shape, all indicate that the specimen is
Phyllium (Phyllium) geryon.
Treatment of Genera and Species
Microphyllium Zompro, 2001
Type species: Microphyllium spinithorax Zompro, 2001 by original designation
FIGURE 2. Microphyllium pusillulum (Rehn & Rehn, 1933), [Coll. RC 16-096], A: dorsal view; B: ventral view.
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Microphyllium pusillulum (Rehn & Rehn, 1933)
(Figs. 2A&B, 5B, 6A&B, 7A&B)
Material examined [2♀]: PHILIPPINES: 1♀: HOLOTYPE: Philippines, Luzon, Nueva Viscaya Province,
Imugan (W. Boettcher.) [Hebard Collection, Type no. 1255.]; 1♀: Philippines, North Luzon, Mountain Province,
Mt. Barlig, June, 2014. [Coll. RC 16-096].
Microphyllium haskelli Cumming, gen. nov.
(Figs. 3A&B, 4A&B, 5A&C, 6C, 7C, 9A, 10A, 11A&B)
Material examined [3♂, 1♀ adult, & 1♀ L-4 nymph]: HOLOTYPE: 1♂: Philippines, North Luzon, Mountain
Province, Mt. Barlig (June, 2014) [Coll. RC 16-099]. (Deposited in the National Museum of the Philippines type
collection. PNM)
PARATYPES: 1♂: Philippines, North Luzon, Mountain Province, Mt. Polis (July, 2014) [Coll. RC 16-098]
(Deposited in the San Diego Natural History Museum collection. SDNHM)
1♂: Philippines, North Luzon, Mountain Province, Mt. Polis (July, 2014) [Coll. RC 16-097] (Retained within
the Royce Cumming Collection.)
1♀: Philippines, North Luzon, Mountain Province, Mt. Polis (May, 2014) [Coll. RC 16-128]; (Retained within
the Royce Cumming Collection.)
1♀ (L-4 nymph): Philippines, North Luzon, Mountain Province, Mt. Polis (December, 2014) [Coll. RC 16-
183] (Retained within the Royce Cumming Collection.)
Comparison. With the males and females of M. haskelli confirmed through genetic analysis, each gender is
compared against the holotypes of the two currently recognized species. Side by side comparison between the
females of M. pusillulum and M. haskelli can be found in Table 4, and comparisons between males of M. haskelli
and M. spinithorax can be found in Table 5.
Description. Description of the coloration based upon the dried specimens. Both paratype male specimens
match closely to the holotype in morphology and coloration. The female description is based on adult specimen
[Coll. RC 16-128]. Many features characterize this species such as: smallest of the adult female Microphyllium
examined at only 40.5 mm long, and with a rather stout mesopraescutum and mesopleurae; protibiae rather long
and slender; profemora with a smooth triangular lobe with small teeth; abdomen smooth sided and wide (maximum
width 20.6 mm). Also, many features heavily and irregularly granulose (head capsule, pronotum, prosternum,
mesosternum).
♂♂. Coloration. Pale green throughout most of the body but with antennae an off orange and thorax that is
more brown than pale green.
Morphology. Head capsule broad with slightly convex cheeks and rough vertex with ~40 unevenly spaced
granules in no apparent pattern (Fig. 10A). Compound eyes large and bulbous, no ocelli present, instead there is a
notable furrow between the antennal fields. Antennae with 21 segments that are short and with beadlike rounded
segments. Apical antennomere cylindrical with full rounded apex, about 2x longer than wide. Pronotum lateral
sides convex with a slight rim, wider rim on the anterior margin which is concave. Pronotum face irregularly
granulose with ~20 granules, a slight groove along the median plane with a notable pit to the anterior and another
closer to the center. Pro- and mesosternum heavily granulose throughout with evenly sized but irregularly spaced
nodes. Mesopraescutum not cleanly formed, rising above the mesopleurae sharply not allowing the lateral margins
to be distinct, the rudimentary margins of the mesopraescutum are marked with +/-4 tubercles of varying size and
spacing. Overall mesopraescutum marked with 8–10 irregularly spaced and sized tubercles/nodes. Of these, those
on the median plane are; two on the anterior rim, one in the center and a notably larger tubercle on the posterior at
least 2X the size of any other on the mesopraescutum. Mesopleurae only slightly diverging with five spiniform
tubercles unevenly spaced. Surface of the mesopleurae are marked with two pits on the posterior half and an
irregularly granulose surface throughout. Tegmina (length 8.6–9.5 mm, maximum width 2.8–2.9 mm), extending
three-quarters of the way into abdominal segment III. Alae (length 17.3–18.1 mm), well developed oval
configuration, only slightly sclerotized on the exposed fold. Abdominal segment II slightly diverging, segments
III–VIII with approximately parallel margins, IX–X gradually tapering towards the pointed apex. Poculum rather
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long and tapering, completely covering the vomer and projects will into the anal segment (Figs. 11A & B).
Profemora slender with a slight interior lobe on the anterior half with three small teeth and no exterior lobe (Fig.
9A). Protibiae without lobes (Fig. 9A). Exterior and interior lobe of meso- and metafemora gently rounded and of
equal size, interior lobes slightly serrate. Meso- and metatibiae lacking lobes.
♀♀. Coloration. Cream to pale green throughout most of the body and tegimina, thorax closer to pale orange
in color. Compound eyes burnt orange in color. The granulation throughout is of a similar color to the surface upon
which it is found.
TABLE 2. Measurements in mm of Microphyllium haskelli specimens. L = length; W = width.
Morphology. Head capsule approximately as long as wide with an irregularly granulose vertex (some arranged
in anterior to posterior lines, others without a detectable pattern) with a notable posteromedian tubercle. The small
protuberance between the compound eye and antennal base has a notable pit in the center. The frontal convexity
has a slight covering in setae and is approximately a third the size of the compound eyes. Antennae moderately
slender and elongate (1.8 mm), shorter than postocular section of head capsule (2.2 mm), and consisting of nine
segments (Fig. 5A). Antennae covered in setae of varying sizes, with the apical antennomere (IX) with the most
variety in size and the greatest density. Apical antennomere cylindrical with rounded apex, about 2x longer than
wide and >2x as long as VIII. Pronotum roughly rectangular, widest at the anterior, which has a distinct concave
rim. Lateral rims on the pronotum are roughly parallel until the posterior third where the rim begins to weaken and
converge on the posterior which lacks a distinct rim. Face of pronotum irregularly granulose with a distinct furrow
in the center on the median plane, and a weaker furrow perpendicular to it. Prosternum irregularly granulous, rather
small and compact and with a notable protuberance in the center about twice the size of those around it.
Mesopraescutum at its widest point, twice as wide as long (width-length ratio 2.05:1), gradually narrowing towards
the posterior. Lateral margins marked with 5 major and 1–2 minor tubercles of various sizes. Mesopraescutum disk
with a slight rim on the anterior margin, and along the median plane there are two distinct tubercles close together
with the posterior one being about twice the size as the anterior protuberance found on the rim (Fig. 6C). The
anterior protuberance found on the rim is also slightly split with two distinct points. Mesopleurae gradually
diverging with lateral margins armed with 5 unevenly spaced tubercles, the most prominent of which are on the
anterior. Mesopleurae face also marked with several weak granules as well as a clear pit located in the center.
Mesosternum irregularly granulous throughout. Tegmina (length 24.4 mm, maximum width 9.6 mm) extending to
the posterior margin of abdominal segment VII. Alae rudimentary. Abdominal segments II–IV gradually widening
Species
Author
M. haskelli
Cumming sp. nov.
Measurements [mm] [Coll. RC
16-183] (L-4) ♀
[Coll. RC
16-128] ♀
[Coll. RC
16-097] ♂
[Coll. RC
16-098] ♂
[Coll. RC
16-099] ♂
L. body*** 25.1 40.5 26.7 26.0 25.7
L./W. head 2.5/2.6 4.2/4.0 1.8/2.0 1.9/2.0 1.5/2.0
L. antennae 1.1 1.8 9.2 8.4 9.3
L. pronotum 1.8 2.5 1.5 1.6 1.4
L. mesonotum 2.2 2.5 1.8 2.0 1.6
L/greatest W. of tegmina 3.2/1.2 24.4/9.6 9.5/2.8 8.6/2.8 8.7/2.9
L/greatest W. of alae -/- -/- 18.1/10.4 17.3/12.1 17.4/10.1
Greatest W. of abdomen 10.9 20.6 3.9 3.6 2.8
L. profemora 4.7 6.7 4.4 4.8 5.0
L. mesofemora 4.1 6.5 4.4 4.6 5.0
L. metafemora 5.6 8.9 5.8 5.8 5.8
L. protibiae 3.6 5.0 3.4 3.6 3.5
L. mesotibiae 3.4 5.4 3.4 3.6 3.5
L. metatibiae 5.0 7.5 4.3 4.7 4.2
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with the posterior of segment IV the widest segment and V–X gradually tapering towards the apex. Anal segment
at its widest, wider than long (width-length ratio 1.72:1), with a pointed apex. Subgenital plate short, only
extending slightly over the posterior of segment IX, ending in a rounded point (Fig. 7C). Gonapophyses rather
short as well only slightly protruding from under the subgenital plate (Fig. 7C). Profemora with a very narrow
exterior lobe. Interior lobe wider than exterior lobe, obtuse in angle, and only slightly dentate with 3+/- distinct
teeth. Protibiae lacking an exterior lobe and the interior lobe is reduced to a sliver. Exterior and interior lobe of
mesofemora gently rounded, interior lobe with slight dentation and only slightly wider than exterior. Exterior and
interior lobe of metafemora gently rounded with exterior lobe rather thin, interior lobe slightly serrate.
Measurements of the holotype and paratypes can be found in table 2 and can be compared against
measurements of M. pusillulum and M. spinithorax specimens found in table 3.
TABLE 3. Measurements in mm of M. spinithorax and M. pusillulum specimens. L = length; W = width.
* Taken from the original description.
**First author’s measurements.
***Including head and cerci, excludes antennae.
****Missing from the specimen.
TABLE 4. Comparison between species of known adult female Microphyllium Zompro, 2001.
Species
Author
M. spinithorax
Zompro, 2001
M. pusillulum
(Rehn & Rehn, 1933)
Measurements [mm] HT ♂* PT ♀ (L-4)* [Coll. RC
16-096] ♀**
HT: Type no. 1255♀**
L. body*** 24.2 29.2 42.3 47.0
L./W. head 1.8/- 3.2/- 4.6/3.6 4.9/4.4
L. antennae - - 2.0 ****
L. pronotum - - 2.5 3.0
L. mesonotum - - 2.1 3.2
L/greatest W. of tegmina - - 23.4/9.7 25.0/10.5
L/greatest W. of alae - - - -
Greatest W. of abdomen - - 21.4 23.0
L. profemora 4.1 - 6.5 7.2
L. mesofemora 4.0 4.7 6.3 7.3
L. metafemora 4.8 6.2 8.3 8.9
L. protibiae 3.0 - 4.8 5.5
L. mesotibiae 2.9 3.7 4.7 5.5
L. metatibiae 4.0 5.0 7.4 8.2
M. pusillulum
(Rehn & Rehn, 1933)
M. haskelli
Cumming sp. nov.
Head Capsule Vertex w/ ~15 regularly spaced granules Vertex w/ ~40+ small unevenly spaced granules
Pronotum 6 granules (2 X 3 configuration) Irregularly granulose, no pattern
Profemur Interior lobe angle of ~90°, with 2 broad teeth Interior lobe obtuse angle, with 3 small teeth
Protibia Slight exterior lobe and distinct rounded
triangular interior lobe
Without an exterior lobe and interior lobe greatly
reduced
Mesopraescutum Lateral margins marked with 3 tubercles Lateral margins with 4 nubby major tubercles and 1–2
minor tubercles
Subgenital Plate Short, ending in a sharp point Short, ending in a rounded point
Gonapophyses Long and slender reaching apex of abdomen Short, only slightly protruding from under the
subgenital plate
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TABLE 5. Comparison between species of known adult male Microphyllium Zompro, 2001.
*Taken from Zompro, 2001 original description.
**As illustrated by Wedmann, S. et al., 2007.
Etymology. This new species is a patronym named in honor of world famous forensic entomologist Dr. Neal
H. Haskell of Rensselaer, Indiana, United States. Dr. Haskell has been an invaluable mentor to the first author as he
works on his master’s degree.
FIGURE 3. Microphyllium haskelli Cumming sp. nov., PT [Coll. RC 16-128]. A: dorsal view; B: ventral view.
M. haskelli Cumming sp. nov. M. spinithorax Zompro, 2001*
Pedicellus Half or more than height of scapus Short, disk-like, less than half as long as scapus
Antennae segments 21 (count includes scapus and pedicellus) 20 (count includes scapus and pedicellus)
Terminal antennae
segment
A full rounded cone, no excavation or
reduction in size
Anterior part flat, with a spoon-like excavation
Tegmina Reaching posterior of III Reaching posterior of IV
Alae Reaching ½ through IX Reaching posterior margin of VIII
Genitalia Poculum with a pointed apex that covers the
vomer
**Poculum with a rounded apex, exposing the vomer
below
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FIGURE 4. Male Microphyllium haskelli Cumming sp. nov., PT [Coll. RC 16-097], A: dorsal view; B: ventral view.
FIGURE 5. Antennae of female A: Microphyllium haskelli Cumming sp. nov., PT [Coll. RC 16-128] (D. Burk/Indiana, USA);
B: M. pusillulum (Rehn & Rehn, 1933), [Coll. RC 16-096] (D. Burk/Indiana, USA); C: dorsal view female nymph M. haskelli
Cumming sp. nov., PT [Coll. RC 16-183].
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FIGURE 6. Side view of Microphyllium female thorax and head. A: M. pusillulum HT (J.D. Weintraub/ANSP Entomology);
B: M.pusillulum [Coll. RC 16-096]; C: M. haskelli Cumming sp. nov., PT [Coll. RC 16-128].
FIGURE 7. Ventral view of Microphyllium female genital plates A: M. pusillulum HT (J.D. Weintraub/ANSP Entomology); B:
M.pusillulum [Coll. RC 16-096]; C: M. haskelli Cumming sp. nov., PT [Coll. RC 16-128].
Pseudomicrophyllium Cumming gen. nov.
Type Species: Pseudomicrophyllium faulkneri Cumming gen. et sp. nov.
Etymology: The genus is named because it was thought for several years to be a new Microphyllium species
because of its size and it was not until closer examination that it was revealed it to be a new genus evading
detection right under the first author’s view because of those similarities.
Diagnosis. ♂: Appearing closely related to Phyllium (Phyllium) with characteristics such as the antennae,
thorax, but the body size and spination of the legs are reminiscent of Microphyllium. The combination of the below
morphological features characterize males of the new genus and differentiate it from the other known phylliid
genera.
Antennae: Long and filiform with long setae, similar to Phyllium (Phyllium).
Profemora: Completely lacking an exterior lobe, interior lobe greatly reduced, similar to Microphyllium.
Protibia: Lacking both exterior and interior lobes, similar to Microphyllium.
Thorax: Similar to Phyllium (Phyllium) with a distinct mesopraescutum.
The most evident feature that differentiates Pseudomicrophyllium gen. nov. from Microphyllium Zompro,
2001 are the long filiform antennae. Abdominal shape in the walking leaf family is a feature that has proven to be
very variable in female Phyllium and more stable in male Phyllium. However not enough is known about the male
variation of abdominal shape in the small phylliid genera of Luzon to use abdominal shape as a strong
morphological feature in their differentiation and identification. Antennae structure is a feature that has proven
much more stable throughout the family and this emphasis is reflected in the dichotomous key at the end of this
work.
♀: UNKNOWN
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Pseudomicrophyllium faulkneri Cumming sp. nov.
(Figs. 8A&B, 9B, 10B&C, 11C)
HOLOTYPE: ♂: Philippines, North Luzon, Ifugao, Banaue (IX-2014) [Coll. RC 16-129]. (Deposited in the
National Museum of the Philippines type collection. PNM)
Description. Description of the coloration based upon the dried specimen. Body length similar in size to
known Microphyllium males but with antennae just over double the length of the characteristic short bead-like
antennae of Microphyllium.
Coloration. Pale throughout most of the HT likely due to the drying process. The compound eyes are of a
reddish brown coloration. Venation of the tegmina is of a yellow to orange color but this, too, is assumed to have
been a vivid green in life.
FIGURE 8. Male Pseudomicrophyllium faulkneri Cumming gen. et sp. nov., HT [Coll.RC 16-129], A: dorsal view; B: ventral
view.
Morphology. Head capsule broad with convex cheeks and vertex smooth with ~16 evenly spaced granules (Fig.
10B). Compound eyes large and bulbous, no ocelli. Antennae with 23 segments, long and slender with setae length
that at times can be over twice the width of the segment it is protruding from (Fig. 10C). Apical antennomere
cylindrical with rounded apex, about 2x longer than wide. Pronotum with an anterior margin twice as wide as the
posterior margin, and lateral sides rather straight with a slight rim, wider rim on the anterior margin (Fig. 10B).
Pronotum face smooth with 6 evenly spaced nodes, a slight groove along the median plane on the anterior half
along with a slight perpendicular groove. Prosternum with slight uneven granulation throughout. Mesopraescutum
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with slightly narrowing lateral margins marked with 3 evenly spaced tubercles and disk face distinctly raised along
the median plane marked with three tubercles, the one on the anterior rim is the smallest of the three and the
posterior most at least 2X the size of the other two. Just lateral to the tubercles of the median plane are 2 nodes on
each side on the mesopraescutum disk face (Fig. 10B). Mesopleurae only slightly diverging with four tubercles
evenly spaced (Fig. 10B). Mesosternum with slight uneven granulation throughout. Tegmina (length 8.2 mm,
maximum width 1.8 mm) rather small, extending to the middle of abdominal segment II. Alae (length 19.7 mm),
well developed oval configuration, only slightly sclerotized on the exposed fold. Abdominal segment II slightly
converging, segments III–IV gradually diverging with IV the widest segment, V–X gradually tapering towards the
apex. Anal segment as long as wide and with a rounded apex. Poculum projecting well into the anal segment and
covering a majority of the vomer (Fig. 11C). Profemora slender with a slight interior lobe on the anterior 1/5
th
as a
small rounded triangle (Fig. 9B). Protibiae lacking both interior and exterior lobes (Fig. 9B). Exterior and interior
lobe of meso- and metafemora gently rounded with interior lobes slightly serrate.
Measurements [mm]: Length of body (excludes cerci which are lacking in the HT) 27.8, length/width of head
2.4/2.3, length of pronotum 1.7, length of mesonotum 1.7, length of tegmina 8.3, greatest width of tegmina 1.8,
length of alae 19.7, greatest width of abdomen 5.9, length of profemora 5.1, length of mesofemora 4.8, length of
metafemora 5.4, length of protibiae 3.9, length of mesotibiae 3.6, length of metatibiae 4.5, length of protarsi 5.1,
length of antennae 20.1.
Etymology. This new species is a patronym named in honor of David Faulkner of California, USA. Faulkner
has been a mentor to the first author over the years and was the first to introduce him to forensic entomology.
FIGURE 9. Male left foreleg A: Microphyllium haskelli Cumming sp. nov., PT [Coll. RC 16-098] (D. Burk/Indiana, USA); B:
Pseudomicrophyllium faulkneri Cumming gen. et sp. nov., HT (D. Burk/Indiana, USA).
Phyllium Illiger, 1798
Type species: Phyllium (Phyllium) siccifolium (Linnaeus, 1758) by original description.
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FIGURE 10. Male Microphyllium haskelli Cumming sp. nov., PT [Coll. RC 16-098] (D. Burk/Indiana, USA), A: head and
thorax. Male Pseudomicrophyllium faulkneri Cumming gen. et sp. nov., HT (D. Burk/Indiana, USA); B: head and thorax; C:
left antennae.
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FIGURE 11. Ventral view of male genitalia, A: Microphyllium haskelli Cumming sp. nov., PT [Coll. RC 16-098] (genital plate
open away from body); B: M. haskelli Cumming sp. nov., PT [Coll. RC 16-097] (genital plate tucked against the body); C:
male Pseudomicrophyllium faulkneri Cumming gen. et sp. nov., HT.
Phyllium (Phyllium) antonkozlovi Cumming sp. nov.
(Figs. 12A&B & 13A,B,&C)
HOLOTYPE: 1♂ Philippines, Luzon, Isabela Prov., Dindin (VIII-2014) [Coll. RC 16-122]
(Deposited in the National Museum of the Philippines. PNM)
Discussion. With the lack of exterior lobes of the tibiae and simple antennae this new species is placed within
the subgenus Phyllium (Phyllium), species-group placement however is tentative. Hennemann et al., 2009 describe
the males of the siccifolium species-group as having an exterior lobe of the profemora which is always distinctly
narrower than the interior lobe, a feature only weakly accurate for Ph. (Ph.) antonkozlovi sp. nov. (Fig. 13C).
However, the interior lobe leads us to believe that Ph. (Ph.) antonkozlovi sp. nov. is closely related to Phyllium
(Phyllium) philippinicum Hennemann et al., 2009 because of the teeth of the profemora that are small and evenly
spaced. Phyllium (Ph.) antonkozlovi sp. nov. can immediately distinguishable from Ph. (Ph.) philippinicum
Hennemann et al., 2009 by the robust antennae segments and lack of developed ocelli. Because of the male’s
similarity to Phyllium (Phyllium) philippinicum, it is predicted that the female will also be morphologically similar
to Ph. (Ph.) philippinicum females. It is unlikely that this male specimen is the unknown male of the other known
Philippine species, which is known only from females. The large size of the male (57.4 mm) suggests a female
similar in size to Ph. (Ph.) philippinicum females (77.5–88.0 mm, Hennemann et al., 2009). Phyllium (Ph.)
bilobatum Gray, 1843 is known only from the female holotype with the vague locality of “Philippines” is much too
small (65.0 mm) to be the unknown Ph. (Ph.) antonkozlovi sp. nov. female. The holotype Phyllium (Ph.) woodi
Rehn & Rehn, 1933 is closest to the expected size of the unknown Ph. (Ph.) antonkozlovi sp. nov. female (76.0 mm
in length), but Ph. (Ph.) woodi is currently only known from Sibuyan Island, far from the type locality of Ph. (Ph.)
antonkozlovi sp. nov. and is morphologically dissimilar to Ph. (Ph.) philippinicum females. Phyllium (Ph.) geryon
Gray, 1843 is also much too small to be the Ph. (Ph.) antonkozlovi sp. nov. female (Ph. (Ph.) geryon holotype 62.0
mm in length and specimen [Coll RC 17-256] only 65.7 mm in length).
Description. Description is based upon the HT male as no other specimens are known to exist and the female
is currently unknown. Rather average in size for known Phyllium (Phyllium) males at 57.4 mm in length. Antennae
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however are shorter and robust, approximately the length of the extended forelegs (most Phyllium (Phyllium)
antennae are longer than the extended forelegs by 3 to 6 antennal segments).
Coloration. Despite being dried, the lime green coloration has not faded as much as most walking leaves color
often does. Antennae are very deep brown color and are darker than the cherry red compound eyes. Spination is
mostly of a similar color to the surface it is found on.
FIGURE 12. Phyllium (Phyllium) antonkozlovi Cumming sp. nov., HT, [Coll. RC 16-122], A: dorsal view; B: ventral view.
Morphology. Head capsule about as broad as long with rather straight cheeks creating a boxy overall shape
(Fig. 13A). Rather large and distinct antennal fields, vertex smooth with only a slightly noticeable posteromedian
tubercle. Compound eyes of medium size only slightly protruding, ocelli highly reduced to only a slight marking
on the capsule indicating where they were. Antennal rather robust, and consisting of 23 segments. Basal segments
are beadlike and cylindrical (Fig. 13A). Apical antennomere cylindrical with rounded apex, about 2x longer than
wide. Pronotum relatively smooth but with a distinct furrow and slight pit along the median plane and three less
notable pits along the anterior rim. Anterior margin concave, lateral and posterior margins slightly convex. Shape
roughly trapezoidal with the anterior length about twice that of the posterior. Anterior and lateral margins with
slight rims. Prosternum smooth except for slight granulation. Mesopraescutum approximately twice as long than
wide, slightly narrower towards the posterior. Lateral margins with 6–7 tubercles of uniform size but somewhat
unevenly spaced. Mesopraescutum disc only slightly raised along the median plane with a relatively smooth crest
marked by a prominent tubercle on the anterior rim and another in the center, in addition to these two prominent
tubercles there are 4–5 lesser nodes. Mesopleurae only slightly diverging from mesopraescutum lateral margins,
with 3–4 distinct tubercles and 7–9 minor tubercles intermixed. Mesosternum rather smooth without notable
granules. Tegmina (length 21.2 mm, maximum width 6.7 mm), extending about a third the way through abdominal
segment IV. Alae (length 37.9 mm), well developed, oval fan configuration with exposed section slightly
sclerotized. Abdominal segment II slightly tapering, segments III –V gradually widening with VI marking the
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widest segment, VI–X gradually tapering towards the apex. Spiracles distinctly visible on the ventral surface,
located on the anterior margin of segments III–VIII near the median plane. Anal segment not tapering evenly
creating a pentagonal shape with a rounded apex. Poculum rather short with a flat apex, slightly projecting over the
posterior margin of abdominal segment IX and exposing the vomer, which is rather wide (Fig. 13B). Profemora
with a narrow rounded exterior lobe and slightly thicker interior lobe, which is serrate with 5–6 distinct teeth
pointing anteriorly, which are more or less evenly spaced (Fig. 13C). Protibiae lacking exterior lobe, interior lobe
only a smooth arch, not triangular. Exterior and interior lobes of mesofemora gently rounded and slightly serrate
with the exterior lobe slightly narrower. Exterior and interior lobes of metafemora gently rounded with interior lobe
wider and serrate.
FIGURE 13. Phyllium (Phyllium) antonkozlovi Cumming sp. nov., HT [Coll. RC 16-122], A: antennae, head, and thorax; B:
genitalia ventral view; C: right profemora.
Measurements [mm]: Length of body 57.4, length/width of head 3.8/3.5, length of pronotum 3.5, length of
mesonotum 5.0, length of tegmina 21.2, greatest width of tegmina 6.7, length of alae 37.9, greatest width of
abdomen 13.3, length of profemora 13.3, length of mesofemora 11.1, length of metafemora 15.6, length of
protibiae 9.4, length of mesotibiae 7.8, length of metatibiae 10.9, length of protarsi 7.2, length of antennae 27.1.
Etymology. This new species is a patronym named in honor of Anton Olegovich Kozlov, long time friend to
the first author and well-known Russian field entomologist.
Phyllium (Phyllium) bourquei Cumming & Le Tirant sp. nov.
(Figs. 14A,B&C, 15A,B,C,D&E 16A,B&C, 17A&B, 18A,B,&D)
HOLOTYPE: ♀: Philippines, Luzon, Nueva Vizcaya, Kayapa, March, 2017 [Coll. RC 17-203].
(Deposited in the National Museum of the Philippines. PNM)
PARATYPES: 1♀: Philippines, Luzon, Nueva Vizcaya, August, 2007 [Coll. RC 17-255].
1♂: Philippines, Luzon, Nueva Vizcaya, Belanue, May, 2014 [Coll. RC 16-201].
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FIGURE 14. Phyllium (Phyllium) bourquei Cumming & Le Tirant sp. nov. HT female [Coll. RC 17-203], A: dorsal view; B:
ventral view; C: left foreleg.
Coauthored with Stéphane Le Tirant, Montreal, Canada, whose collection originally contained the paratype
female. Discussion. This species was originally thought to be Phyllium (Phyllium) philippinicum Hennemann et
al., 2009 found outside its known range. Upon closer examination, several differences became clear. Like Ph. (Ph.)
philippinicum, this new species falls under the siccifolium species-group of Hennemann, et al., 2009 because the
female lacks developed alae and the male has an exterior lobe of the profemora that is more slender than the
interior lobe. It is possible that future genetic analysis will find that these two populations are not conspecific, but
the marked differences in the female genitalia led us to here describe them with species level status. The
geographic isolation of the two populations, Ph. (Ph.) philippinicum from the Zambales Range, and Ph. (Ph.)
bourquei from the Caraballo Mountains, separated by the Central Luzon Plains, also led to the decision to erect the
new species.
Description. Description is based upon the HT female and the paratype male/female specimens. Coloration is
based upon the dried specimens here illustrated, it is assumed the colors were lighter in life.
♂♂. Coloration. Pale green throughout (except for areas of rot), other areas (antennae and thorax in particular)
paler in color, more straw colored than green. No eyespots noted on the abdomen in the male paratype.
Morphology: Head capsule length and width approximately equal (Fig. 17A). Distinct antennal fields, vertex
mostly smooth, only slight granulation on the posterior end and around a broad but short posteromedian tubercle.
Compound eyes large and protruding, ocelli well developed. Antennal simple, and consisting of 23 segments
(including scapus and pedicelus) and with long thin setae. Apical antennomere cylindrical with rounded apex,
slightly more than 3x longer than wide and covered with short dense setae. Pronotum relatively smooth, with a
distinct furrow and slight pit along the median plane. Anterior margin concave, lateral and posterior margins
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slightly convex. Shape roughly trapezoidal with the anterior length slightly less than twice that of the posterior.
Anterior and lateral margins with distinct rims. Prosternum with moderate granulation. Mesopraescutum
approximately as long as wide, and only slightly narrower towards the posterior. Lateral margins with 5–6 decent
sized tubercles of slightly different sizes and somewhat unevenly spaced. Mesopraescutum disc raised along the
median plane with a crest marked by a prominent tubercle on the anterior rim followed by three others of steadily
decreasing size and even spacing with the smallest ending near the posterior rim (Fig. 17B). The surface of the
mesopraescutum disc on each side of the crest is free of nodes. Mesopleurae slightly diverging from
mesopraescutum lateral margins for the first 1/3 and then gradually widening for the remainder. Mesopleurae
margins with 5 distinct tubercles predominantly on the anterior and 3 minor tubercles intermixed on the posterior
portion. Prosternum fully covered in granules of varying sizes, but none more notable than the others.
Mesosternum rather smooth but with notable granules along the sagittal plane, more heavily marked on the
anterior. Tegmina (length 17.9 mm, maximum width 5.2 mm), extending to the anterior margin of abdominal
segment IV. Alae (length 37.7 mm), well developed, oval fan configuration with exposed section slightly
sclerotized. Abdominal segment II with parallel margins, segments III – first 2/3 of segment IV gradually widening
with IV marking the widest segment, posterior 1/3 of IV–X gradually tapering towards the apex, at first only
slightly then more prominently creating a spade shaped abdomen. Spiracles only just visible on the ventral surface,
located on the anterior margin of segments III–VIII near the median plane. Anal segment posterior half somewhat
evenly rounded apex. Poculum rather stout with a straight posterior margin, slightly projecting over the posterior
margin of abdominal segment IX and exposing the vomer, which is rather wide with a single stout terminal hook
(Fig. 15C). Exterior lobe of profemora narrow, at its widest point only slightly wider than the shaft of the femur.
Exterior lobe of the profemora relatively smooth, only marked by small, barely notable teeth pointing anteriorly.
Interior lobe not starting until a third the way up the femur and arching in a rounded triangle marked with six to
seven small anteriorly pointing teeth that are not quite evenly spaced (Fig. 16C). Protibiae lacking exterior lobe,
interior lobe only a smooth arch, almost triangular in shape. Exterior and interior lobes of mesofemora gently
rounded and approximately equal in width and both lightly serrate. The interior lobe marked more heavily with six
to seven tightly packed teeth, the exterior lobe less serrate, marked with three more widely spaced teeth. Exterior
and interior lobes of metafemora gently rounded; interior lobe notably wider and with eight serrate teeth; exterior
lobe lacking serration. Meso- and metatibiae simple, lacking lobes or serration.
♀♀. Coloration. Most of the holotype is discolored but a pale green is still detectable on the margins of the
abdomen and forelegs. The paratype is mostly a paler color and almost completely discolored.
TABLE 6. Measurements in mm of Phyllium (Phyllium) bourquei Cumming & Le Tirant sp. nov. type specimens. L =
length; W = width.
*Including head and cerci, excludes antennae.
Specimen HT ♀
[Coll. RC 17-203]
PT ♀
[Coll. RC 17-255]
PT ♂
[Coll. RC 16-201]
L. body* 85.2 83.6 51.6
L./W. head 7.6/6.1 7.7/6.1 2.9/2.7
L. antennae 4.7 4.3 28.9
L. pronotum 5.1 4.7 2.5
L. mesonotum 6.9 5.7 3.7
L/greatest W. of tegmina 55.2/18.1 52.6/16.7 17.9/5.2
L/greatest W. of alae 6.6/- -/- 37.7/19.8
Greatest W. of abdomen 38.1 36.0 11.7
L. profemora 18.1 17.6 10.6
L. mesofemora 15.3 15.2 9.0
L. metafemora 19.8 19.4 11.6
L. protibiae 11.1 11.1 7.2
L. mesotibiae 10.6 10.2 6.2
L. metatibiae 14.8 14.7 8.3
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FIGURE 15. Ventral view genitalia of Ph. (Ph.) bourquei Cumming & Le Tirant sp. nov., A: PT female [Coll. RC 17-255] (R.
Limoges/Montreal Insectarium); B: HT female [Coll. RC 17-203]; C: PT male [Coll. RC 16-201]. Female antennae, D: Ph.
(Ph.) bourquei Cumming & Le Tirant sp. nov. PT female [Coll. RC 17-255] (R. Limoges/Montreal Insectarium); E: Ph. (Ph.)
bourquei Cumming & Le Tirant sp. nov. HT female [Coll. RC 17-203]; F: Ph. (Ph.) philippinicum Hennemann et al., 2009
[Coll. RC 16-142].
Morphology. Head capsule longer than wide with the posterior 1/4 sparsely granulose with a notable
posteromedian tubercle at least three times the size of any other node. Antennae moderately slender and elongate,
slightly longer than the postocular section of the head capsule, and consisting of ten segments (Figs. 15D&E).
Antennae mostly lacking setae, the terminal two segments are the only ones with notable setae covering. Apical
antennomere cylindrical with rounded apex, only slightly longer than the preceding segment. Pronotum relatively
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smooth, with a distinct furrow and slight pit along the median plane. Anterior margin concave, lateral and posterior
margins slightly convex. Shape roughly trapezoidal with the anterior width approximately twice that of the
posterior. Anterior margin with distinct rim, lateral margins with weak rims. Mesopraescutum only slightly
narrowing towards the posterior and approximately the same length as the width. Lateral margins marked with ~6
robust major tubercles of relatively uniform size. Mesopraescutum disk with a prominent rim on the anterior
margin marked by a distinct spine followed by four small but clear nodes along the sagittal plane (Figs.
18A,B,&D). Mesopleurae uniformly diverging with lateral margins armed with ~7–8 robust tubercles.
Mesopleurae face relatively smooth but marked with a clear pit located on the anterior third and a slight pit on the
posterior third. Prosternum irregularly granulous throughout. Mesosternum mostly smooth but with notable
granules along the sagittal plane, more heavily marked on the anterior. Tegmina extending slightly past the anterior
margin of abdominal segment VIII. Alae rudimentary. Abdominal segments II- the first 2/3 of IV gradually
widening, with segment IV marking the widest segment. The posterior third of IV–X tapering towards the apex
with segments VII and VIII lobed. Anal segment at its widest, slightly wider than long, with a broad apex.
Subgenital plate with a fine point reaching about half way under the anal abdominal segment (Figs. 15A&B).
Gonapophyses rather long, slightly protruding from under the anal abdominal segment when viewed dorsally.
Profemora with a widely rounded exterior lobe that has a relatively smooth outer margin. Interior lobe, slightly
narrower than exterior lobe and slightly more angled, anterior portion of the margin marked with 5 small saw-like
teeth of equal size and shape, but not with perfectly equal spacing. Protibiae lacking an exterior lobe and the
interior lobe a rounded isosceles triangle. Exterior and interior lobe of mesofemora gently rounded; exterior lobe
slightly wider than interior lobe due to the shape being more angled than the interior lobe, which is gently arching.
Exterior lobe is marked with two to three widely spaced small teeth, interior lobe marked more heavily with serrate
dentition (five to six teeth). Exterior and interior lobe of metafemora gently rounded with exterior lobe rather thin,
interior lobe slightly wider and with seven serrate teeth, exterior lobe lacking dentition. Meso- and metatibiae
simple, lacking lobes or serration.
FIGURE 16. Phyllium (Phyllium) bourquei Cumming & Le Tirant sp. nov. PT male [Coll. RC 16-201], A: dorsal view; B:
ventral view; C: left foreleg.
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FIGURE 17. Male dorsal view of base of antennae, head, and thorax of Phyllium (Phyllium) bourquei Cumming & Le Tirant
sp. nov. PT male [Coll. RC 16-201], A: base of antennae, head, and thorax. Side view of males; B: Ph. (Ph.) bourquei
Cumming & Le Tirant sp. nov. PT male [Coll. RC 16-201], C: Ph. (Ph.) philippinicum Hennemann et al., 2009 [Coll. RC 16-
162].
Measurements of the type material can be found within table 6.
Etymology. This species is dedicated to Mr. Pierre Bourque. He was mayor of the City of Montreal from 1994
to 2001. Mr. Bourque was also one of the most innovative directors of the Montreal Botanical Garden from 1980
to 1994. Among other things, he was responsible for the creation of new greenhouses, the Floralies Internationales
de Montréal, the Japanese Garden, the Chinese Garden, the Tree House, the Montreal Biodome and the Montreal
Insectarium. Thanks to Mr. Bourque, the Montreal Botanical Garden has become the second largest in the world.
Phyllium (Phyllium) geryon Gray, 1843
(Figs. 19A&B & 20A,B,C&D)
Material examined [2♀]: PHILIPPINES: 1♀: HOLOTYPE: 42. 72 Phil. Isl.; Philippine Islands; geryon,
G.R.Gray, Cumings; Phyllium geryon G.R.Gray, Philippine Islds.; BMNH(E) #845232 [examined from detailed
photos available on Phasmida Species File http://phasmida.speciesfile.org (author: Paul D. Brock)]; 1♀:
Philippines, Luzon, Aurora Province, Dingalan Municipality, August, 2017 [Coll. RC 17-256].
Discussion. Originally only known from the inexact type locality of “Inhabits the Philippine Islands” from
Gray’s 1843 description, Phyllium (Phyllium) geryon represents the tenth species of Phylliidae confirmed from
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Luzon. The small Phyllium specimen in the first author’s collection was originally expected to be difficult to
compare morphologically to the damaged holotype, because many of the most frequently used features are missing.
However, several unique features that still remain on the holotype allow us to confidently identify the punitive
specimen. The granulation/spination of the head and thorax of the holotype were the most important features of the
broken specimen to allow confident identification.
FIGURE 18. Female thorax dorsal view, A: Phyllium (Phyllium) bourquei Cumming & Le Tirant sp. nov. HT [Coll. RC 16-
203]; B: Ph. (Ph.) bourquei Cumming & Le Tirant sp. nov. PT [Coll. RC 17-255]; C: Ph. (Ph.) philippinicum Hennemann et
al., 2009 [Coll RC 16-142]. Female thorax side view, D: Ph. (Ph.) bourquei Cumming & Le Tirant sp. nov. HT [Coll. RC 17-
203]; E: Ph. (Ph.) philippinicum [Coll RC 16-142].
The most striking feature on the thorax that identifies Phyllium (Ph.) geryon is the spination along the crest of
the mesopraescutum. The anterior rim lacks a spine, and is instead followed by a notable spine in the center and
ends in a large prominent spine on the posterior that, in profile, is at least twice the size of the anterior rim (Fig.
20D). Also the five well-defined tubercles of the mesopleurae with one to two small nodes placed between each
were perfectly reflected in the specimen and is not something that we have seen in examination of dozens of other
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Phyllium species. The other features of the thorax also match perfectly but are not as unique as the posterior spine
of the mesopraescutum crest.
The only feature that did not match perfectly between specimens was the apex shape of the subgenital plate. In
the holotype the apex is sharply pointed and in the specimen from the first author’s collection the apex is rounded.
The length of the subgenital plate however matches between the two individuals (both short, only reaching the
posterior of segment IX) and the difference in terminal shape is assumed to be the result of being worn down, or a
slight deformity resulting in a dulled point.
Many of the features described below are described for the first time, as it appears the holotype has been
damaged for most of its existence. The profemora that was illustrated in the original description is the only feature
that has drawn notable discussion over the years, as it is unclear if it belonged to the actual holotype or if it was a
recreated illustration from deHaan’s work the year before (de Haan, 1842). The profemora illustration in Gray’s
work certainly appears to be the same as the male nymph illustrated the year before which de Haan listed as
belonging to a Phyllium (Phyllium) siccifolium male nymph. The determination by de Haan is likely erroneous as
he lists the specimen as coming from “Timor, Nova Guinea” a locality not known to have Phyllium (Phyllium)
siccifolium and is instead more likely to belong to one of the many New Guinea native Phylliidae that have since
been described.
FIGURE 19. Female Phyllium (Phyllium) geryon Gray, 1843 [Coll. RC 17-256], A: dorsal view; B: ventral view.
We do find it odd however that while Gray appears to have illustrated a male nymphs profemora and assumed
that it was equivalent to the Phyllium (Ph.) geryon holotype, he did place a great deal of importance on the shape of
the profemora of Phylliidae, dividing his 1843 work into three divisions based solely on the exterior lobe of the
profemora. From the specimen in the first author’s collection it is clear that the exterior lobe of the profemora,
although not very wide (slightly thinner than the interior lobe) is complete and present, not absent as Gray
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describes. This leads us to believe that the holotype, even from the time of Gray’s original observation was missing
its forelegs and Gray looking to use the profemora as a key feature in his work went looking for a specimen with a
similar body shape to the Phyllium (Ph.) geryon holotype and found de Haan’s male nymph adequate. The first
mention that the forelegs were missing from the holotype, and first to suggest that perhaps Gray only illustrated the
nymph from de Haan, was Westwood only sixteen years latter when he observed the specimen and found the
forelegs “wanting” (Westwood, 1859). Rehn and Rehn in 1933 also discussed their suspicion of the missing
exterior lobe of the profemora and surprisingly guessed that Phyllium (Ph.) geryon was native to Luzon, a guess
incorrectly based on their assumption that Phyllium (Ph.) geryon was closely related to Microphyllium pusillulum
which they based on the small size and thin exterior lobe of the profemora.
♀♀. Coloration. Pale to darker green throughout most of the body and tegimina, some areas discolored from
drying. Compound eyes burnt orange. The granulation throughout is of a similar color to the surface upon which it
is found or slightly lighter such as those found on the head capsule.
FIGURE 20. Female Phyllium (Phyllium) geryon Gray, 1843 [Coll. RC 17-256], A: antennae, head, and thorax; B: genitalia
ventral view; C: right profemora; D: lateral view of antennae, head, and thorax.
Morphology. Head capsule slightly longer than wide with a moderately detectable pattern of granules. Lateral
to the prominent posteromedian tubercle there is a node about half the size, followed by two to three other similarly
sized nodes following the margins of the head. Interior to those nodes there are several nodes on the posterior half
of the head capsule pointing from the compound eyes back to the posteromedian tubercle forming a “V” pattern
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(Fig. 20A). The small protuberance between the compound eye and antennal base is marked with a notable pit in
the center. The frontal suture is also well furrowed. The frontal convexity has a slight covering in setae and is only
slightly smaller than the size of the compound eyes, which are also smaller and less bulbous. Antennal fields only
slightly wider than the base of the antennae. Antennae moderately slender and elongate (4.0 mm), approximately
the same length as the postocular section of head capsule, and consisting of nine segments (Fig. 20A). Antennae
covered in setae of varying sizes, most sparse except for antennomeres VIII and IX with the most variety in size
and the greatest quantity of setae. Apical antennomere cylindrical with rounded apex, about 2.5x longer than wide
and about 1.5x as long as segment VIII which is notably longer than any of the other preceding four segments.
Pronotum roughly trapezoidal, widest at the anterior, which is at least twice the length of the anterior rim. Lateral
rims on the pronotum are roughly parallel for the first quarter of the length and then converging to the posterior
margin. The anterior rim is distinct and slightly concave, lateral rims are moderate to weak and the posterior margin
has no rim. Face of pronotum irregularly granulose with a distinct furrow on the median plane on the anterior half.
Mesopraescutum almost a square with a length that equals the width and lateral margins that are only slightly
converging. Lateral margins marked with three large tubercles and slight granulation on the anterior end.
Mesopraescutum disk crest has an anterior rim slightly granulose and lacking a spine, followed by a notable spine
in the center of the disk and ending in a large prominent spine on the posterior that, in profile, is at least twice the
size of the anterior rim (Fig. 20D). Mesopleurae evenly diverging with lateral margins armed with five evenly sized
but slightly unevenly spaced tubercles. Between each tubercle there is one to two small nodes. Mesopleurae face
smooth except for two clear pits, one on the anterior third and one on the posterior third. Prosternum irregularly
granulous throughout, the anterior half is sloped into a point, not flat like the posterior half. Mesosternum
irregularly granulous throughout, those along the sagittal plane slightly larger. Tegmina (length 39.2 mm,
maximum width 12.6 mm) extending almost half way into abdominal segment VII. Alae rudimentary. Abdominal
segments II–IV gradually widening with the posterior of segment IV the widest segment and V–X uniformly
tapering towards the apex. Anal segment at its widest, wider than long (width-length ratio 1.67:1), with a relatively
rounded apex. Subgenital plate short, only reaching the posterior of segment IX, sides slightly convex and ending
in a rounded point (Fig. 20B). Gonapophyses long and reaching the apex of the subgenital plate (Fig. 20B).
Profemora with narrow exterior lobe, slightly thinner than the interior lobe, margin appears smooth without
magnification, but under low magnification, the entire margin is marked with small tight, evenly spaced dentition.
Interior lobe wider than exterior lobe, obtuse in angle, and marked on this specimen with prominent teeth, two teeth
on the left profemora and four teeth on the right profemora. Protibiae lacking an exterior lobe and the interior lobe
is only present on the posterior half of the tibiae as a rounded triangle that drops sharply to the tibiae near the
middle. Meso- and metafemora exterior and interior lobe gently rounded, interior lobe with notable serrate
dentation and approximately twice the width of the exterior lobe. Meso- and metatibiae simple, lacking lobes or
dentition.
Measurements [mm] of specimen [Coll. RC 17-256]: Length of body 65.7, length/width of head 6.5/5.6, length
of pronotum 4.0, length of mesonotum 5.0, length of tegmina 39.2, greatest width of tegmina 12.6, length of alae -
-, greatest width of abdomen 23.9, length of profemora 12.7, length of mesofemora 11.2, length of metafemora
13.8, length of protibiae 9.1, length of mesotibiae 8.0, length of metatibiae 12.2, length of protarsi 8.3, length of
antennae 4.0.
Distribution
Checklist of Phylliidae species currently confirmed from Luzon, Philippines, and their recorded ranges. See figure
21 for an illustrated map of the distributions.
Phylliidae, Phylliinae, Phylliini:
Microphyllium Zompro, 2001
Microphyllium pusillulum (Rehn & Rehn, 1933)
[Central Luzon (Mountain* & Nueva Vizcaya Prov.)]
Microphyllium spinithorax Zompro, 2001
[Luzon (Benguet Prov.**)]
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Microphyllium haskelli Cumming sp. nov.
[Luzon (Mountain Prov.)]
Pseudomicrophyllium Cumming gen. nov.
Pseudomicrophyllium faulkneri Cumming gen. et sp. nov.
[Central Luzon (Ifugao Prov.)]
Phyllium Illiger, 1798
Phyllium (Phyllium) ericoriai Hennemann, et al., 2009
[Luzon (Manila, Aurora*, Quirino*, & Quezon Prov.); Batan Is.; Marinduque Is.; Cantanduanes Is.*]
Phyllium (Phyllium) bonifacioi Lit & Eusebio, 2014
[Northern Luzon (Cagayan Prov.* & Northern Luzon***)]
Phyllium (Phyllium) philippinicum Hennemann, et al., 2009****
[Western Luzon (Bataan Prov.)]
Phyllium (Phyllium) antonkozlovi Cumming sp. nov.
[Eastern Luzon (Isabela Prov.)]
Phyllium (Phyllium) bourquei Cumming & Le Tirant sp. nov.
[Luzon (Nueva Vizcaya Prov.)]
Phyllium (Phyllium) geryon Gray, 1843
[Luzon (Aurora Prov.)*]
*New record for the distribution of the species, see below section.
**The type material of M. spinithorax is only labeled with the location “St. Thomas” which is assumed to be Mt. Santo Tomas
in Benguet Province (Collins & Morris, 1985), located in the Cordillera Central mountain range, the same range that all
Microphyllium collection instances are from, adding validity to this assumption.
***The exact location of the type material was not given by the authors.
****In Grösser’s 2011 work “New Insights and Critical Remarks on certain species of Walking Leaves” he suspected that
Phyllium (Phyllium) philippinicum Hennemann, et al., 2009 was not a valid species and was actually a hybrid. With the
popularity of many Phyllium currently in culture throughout Europe the concern of crossed cultures is valid. However, when
Hennemann et al. described Ph. (Ph.) philippinicum they only used specimens from the original stock collected by Ismael O.
Lumawig and Thierry Heitzmann in June of 2001, not specimens from one of the many untraceable cultures (or possibly
hybridized cultures) currently circulating in the phasmid breeding community.
While the first author was organizing the Phylliidae collection of the California Academy of Sciences
collection, two specimens (a subadult female and an adult male) from the U.S. Naval Base at Subic Bay (a location
near the type locality of Ph. (Ph.) philippinicum) were examined. Morphologically these specimens match with the
original description given by Hennemann et al. and key out to Ph. (Ph.) philippinicum. The pair was collected by
James E. Tobler on the 12
th
of November, 1965, almost 40 years before the species was brought into culture in
Europe, therefore, this historic pair gives validity to the species status of Ph. (Ph.) philippinicum Hennemann et al.,
2009.
Discussion of new distribution records. Microphyllium pusillulum from Mountain Province (Mt. Barlig)
from the single female [Coll RC. 16-096], previously only known from the HT record; Imugan, Nueva Vizcaya
Province.
Phyllium (Phyllium) ericoriai from Aurora Province (Dingalan) and Quirino Province (Nagtipunan) from
several males and females in the first author’s collection; Cantanduanes Island (Ibong Sapa) is a recent record
noted from personal communication with Feliciano Avila de Leon Jr. (June, 2017).
Phyllium (Phyllium) bonifacioi from Cagayan Province (Santa Ana Mts.) from a male/female pair in the first
author’s collection. Note that the type locality in the original description of the species was not specific and could
have been Cagayan Province; communication with the authors of Phyllium (Phyllium) bonifacioi was unsuccessful
Phyllium (Phyllium) geryon from Aurora Province, Luzon is the first precise locality for this little known
species. Gray’s original distribution note of “Inhabits the Philippine Islands”, has led to much confusion over the
years and erroneous records. Several authors over the years have mistakenly noted Ph. (Ph.) geryon from localities
such as Java or New Caledonia (see Hennemann et al., 2009 for a thorough explanation of these erroneous
localities).
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FIGURE 21. Distribution map with all current known records for species confirmed from Luzon, Philippines.
Original image downloaded from: https://commons.wikimedia.org/wiki/File:Relief_Map_Of_The_Philippines.png under the
creative commons license originally created by user: Seventide.
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Average elevation, in meters, of Microphyllium and Pseudomicrophyllium collection sites.*
Mt. Santo Tomas (Benguet Prov.) [~2,200 m]
Mt. Polis (Mountain Prov.): [~1,800 m]
Mt. Barlig (Mountain Prov.): [~1,800 m]
Banaue (Ifugao Prov.): [~1,300 m]
Imugan (Nueva Vizcaya Prov.): [~1,000 m]
*Elevations found using Google Earth (2016 Google sources: Image Landsat/ Copernicus; Data SIO, NOAA, U.S. Navy, NGa,
GEBCO).
Key to known genera, species-groups, and species confirmed from Luzon, Philippines
♂♂*
1. Protibiae lacking both exterior and interior lobes (figs. 9A&B); average size small (~24–28mm) . . . . . . . . . . . . . . . . . . . . . . 2.
- Protibiae with a distinct interior lobe, lacking exterior lobe (fig. 16C); average size medium to large (~49–65mm): Phyllium
(Phyllium) Illiger, 1798: . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.
2. Antennae short (about the length of the outstretched forelegs or less); individual antennae segments short and beadlike: Micro-
phyllium Zompro, 2001. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.
- Antennae filiform and notably longer than forelegs: Pseudomicrophyllium gen. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudomicrophyllium faulkneri Cumming gen. et sp. nov.
3. 21 antennal segments; terminal antennae segment a full rounded cone; pedicellus more than half the length of the scapus, not
short and disk-like; tegmina reaching posterior of abdominal segment III. . . . . . . . . . . . . . . . . . M. haskelli Cumming sp. nov.
- 20 antennal segments; anterior of terminal segment flat with a spoon-like excavation; pedicellus short, disk-like, less than half
as long as the scapus; tegmina reaching posterior of abdominal segment IV. . . . . . . . . . . . . . . . . M. spinithorax Zompro, 2001
4. Exterior lobe of profemora larger than or equal in width to interior lobe; teeth on the interior lobe of varying sizes: (celebicum
species-group) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.
- Exterior lobe of profemora slightly to notably thinner than interior lobe and without a strong angle (figs. 13C & 16C); teeth of
the profemora relatively even in size and shape (figs. 13C & 16C): (siccifolium species-group) . . . . . . . . . . . . . . . . . . . . . . . 6.
5. Abdominal segments V–VII widening; VII and VIII lobed; VII widest segment; interior lobe of mesofemora notably wider
than exterior lobe. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ph. (Ph.) ericoriai Hennemann et al., 2009
- Abdominal segments VI–VII subparallel; VII–X converging, weakly lobed; interior and exterior lobe of mesofemora of a sim-
ilar width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ph. (Ph.) bonifacioi Lit & Eusebio, 2014
6. Antennae thin and filiform, setae frequently longer than twice the width of the segment they are found on; ocelli well devel-
oped (fig. 17A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7.
- Antennae robust, most setae length are less than or equal to the width of the segment they are found on; ocelli severely reduced
(fig. 13A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ph. (Ph.) antonkozlovi Cumming sp. nov.
7. Anterior end of the mesopraescutum width notably shorter than the length of the mesopraescutum giving it a long, slender
appearance; crest along the sagittal plane of the mesopraescutum with a single prominent spine (on the anterior rim), vertex
relatively smooth, at most with small evenly sized nodes (fig. 17C) . . . . . . . . Ph. (Ph.) philippinicum Hennemann et al., 2009
- Anterior end of the mesopraescutum approximately equal in width to the length of the mesopraescutum giving it a stout
appearance; crest along the sagittal plane of the mesopraescutum with a prominent spine (on the anterior rim) followed by
nodes that steadily decrease in size, not nodes of the same size (fig. 17B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ph. (Ph.) bourquei Cumming & Le Tirant sp. nov.
♀♀**
1. Overall body length small (40–47mm); length of apical antennomere approximately equal to or longer than the combined
length of segments IV–VIII (figs. 5A&B): Microphyllium Zompro, 2001. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.
- Overall body length medium to large (~60–100mm); apical antennomere notably shorter than the combined length of segment
IV-final segment before the apical antennomere (figs. 15D,E,&F): Phyllium (Phyllium) Illiger, 1798. . . . . . . . . . . . . . . . . . . 3.
2. Protibiae with distinct interior lobe and slight exterior lobe; subgenital plate with a fine point; gonapophyses long and slender
reaching the apex of the abdomen (figs. 7A&B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. pusillulum (Rehn & Rehn, 1933)
- Protibiae without an exterior lobe and interior lobe greatly reduced; subgenital plate with a smooth rounded point; gonapophy-
ses short, only slightly protruding from under the subgenital plate (fig. 7C) . . . . . . . . . . . . . . . . M. haskelli Cumming sp. nov.
3. Alae well developed: (celebicum species-group) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.
- Alae greatly reduced: (siccifolium species-group) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.
4. Abdominal segment VII lateral margin with distinct lobe; angle of exterior lobe of profemora approaching 90 degrees. . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ph. (Ph.) ericoriai Hennemann et al., 2009
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- Abdominal segment VII lateral margin with indistinct lobe & converging posteriorly; angle of exterior lobe of profemora
obtuse . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ph. (Ph.) bonifacioi Lit & Eusebio, 2014
5. Antennae with ten segments; 5–7 small saw-like teeth on interior lobe of profemora (fig. 14C); interior lobe of the protibiae
extending the full length of the protibiae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.
- Antennae with nine segments; 4 large triangular teeth on the interior lobe of the profemora (fig. 20C); interior lobe of the
protibiae only on the posterior half . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ph. (Ph.) geryon Gray, 1843
6. Subgenital plate almost reaching apex of the anal abdominal segment; mesopleurae margins are only marked by three promi-
nent tubercles in the center and a few minor tubercles anteriorly and posteriorly (fig. 18C). . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ph. (Ph.) philippinicum Hennemann et al., 2009
- Subgenital plate only reaching half way under the anal abdominal segment (figs. 15A&B); mesopleurae margins more heavily
marked by 7–8 robust tubercles (figs. 18A&B) . . . . . . . . . . . . . . . . . . . . . . .Ph. (Ph.) bourquei Cumming & Le Tirant sp. nov.
* Male Microphyllium pusillulum (Rehn & Rehn, 1933) and Ph. (Ph.) geryon Gray, 1843 unknown.
** Female Pseudomicrophyllium Cumming gen. nov. and Ph. (Ph.) antonkozlovi Cumming sp. nov. females unknown, female
M. spinithorax Zompro, 2001 excluded from the key because only known specimens are the immature paratypes.
Acknowledgments
We thank Jason Weintraub, entomology collection manager for the Academy of Natural Sciences of Drexel
University (ANSP), who provided the photos of the holotype M. pusillulum, and for his hospitality while the first
author visited the collection to examine the holotype in person, Jim Berrian at the San Diego Natural History
Museum, for many years of mentoring the first author. We would also like to sincerely thank David Faulkner and
Neal Haskell for their years of guiding he first author in the field of forensic entomology, and Danny Burk for his
photographic expertise for many of the detailed shots used for this manuscript. We want to thank René Limoges,
entomological technician at the Montreal Insectarium for taking several of the photos for this work as well as for
many professional courtesies. Specimen collection was carried out on private land, and holotypes will be
repatriated to the National Museum of the Philippines (PNM).
Literature cited
Brock, P.D., Büscher, T. & Baker, E. (2017) Phasmida Species File Online. Version 5.0/5.0. Available from: http://
Phasmida.SpeciesFile.org (accessed 20 September 2017)
Collins, M. & Morris, M. (1985) Threatened Swallowtail Butterflies of the World, The IUCN Red Data Book. IUCN, Gland,
401 pp.
Darriba, D., Taboada, G.L., Doallo, R. & Posada, D. (2012) jModelTest 2: more models, new heuristics and parallel computing.
Nature Methods, 9 (8), 772.
https://doi.org/10.1038/nmeth.2109
Edgar, R.C. (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research,
32 (5), 1792–1797.
https://doi.org/10.1093/nar/gkh340
Folmer, O., Black, M., Hoch, W., Lutz, R. & Vrijenhock, R. (1994) DNA primers for amplification of mitochondrial
cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3 (5),
294–299.
Goloboff, P., Farris, J.S., Källersjö, M., Oxelmann, B., Ramirez, M. & Szumik, C. (2003) Improvements to resampling
measures of group support. Cladistics, 17, S26–S34.
https://doi.org/10.1111/j.1096-0031.2003.tb00376.x
Goloboff, P.A., Farris, J.S. & Nixon, K.C. (2008) TNT, a free program for phylogenetic analysis. Cladistics, 24 (5), 774–786.
https://doi.org/10.1111/j.1096-0031.2008.00217.x
Gouy, M., Guindon, S. & Gascuel, O. (2010) SeaView version 4: a multiplatform graphical user interface for sequence
alignment and phylogenetic tree building. Molecular Biology and Evolution, 27 (2), 221–224.
https://doi.org/10.1093/molbev/msp259
Gray, G.R. (1843) Description of several species of the Genus Phyllium. Zoologist, 1 (1), 117–123.
Grösser, D. (2011) New Insights and Critical Remarks on certain species of Walking Leaves. ARTHROPODA Generalis, 3, 1–
17.
Haan, W. de (1842) Bijdragen tot de Kennis der Orthoptera. Verhandelingen over de natuurlijke Geschiedenis der
Nederlandsche overzeesche Bezittingen. In: Temminck, C.J. (Ed.), Verhandelingen Zoologie, 2, pp. 95–138.
Hennemann, F.H., Conle, O.V., Gottardo, M. & Bresseel, J. (2009) On certain species of the genus Phyllium Illiger, 1798, with
Zootaxa 4365 (2) © 2017 Magnolia Press
·
131
LUZON PHYL LII DAE
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.
Hurst, G.D.D. & Jiggins, F.M. (2005) Problems with mitochondrial DNA as a marker in population, phylogeographic and
phylogenetic studies: the effects of inherited symbionts. Proceedings of the Royal Society B: Biological Sciences, 272
(1572), 1525–1534.
https://doi.org/10.1098/rspb.2005.3056
Illiger, J.K.W. (1798) Verzeichnis der Käfer Preussens. Johann Jacob Gebauer, Halle, 510 pp.
Kodandaramaiah, U., Simonsen, T.J., Bromilow, S., Wahlberg, N. & Sperling, F. (2013) Deceptive single-locus taxonomy and
phylogeography: Wolbachia-associated divergence in mitochondrial DNA is not reflected in morphology and nuclear
markers in a butterfly species. Ecology and Evolution, 3 (16), 5167–5176.
https://doi.org/10.1002/ece3.886
Kômoto, N., Yukuhiro, K., Ueda, K. & Tomita, S. (2011) Exploring the molecular phylogeny of phasmids with whole
mitochondrial genome sequences. Molecular Phylogenetics and Evolution, 58 (1), 43–52.
https://doi.org/10.1016/j.ympev.2010.10.013
Linnæus, C. (1758) Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus,
differentiis, synonymis, locis. Editio decima, reformata. Laurentii Salvii, Stockholm, 824 pp.
Lit, I. & Eusebio, O. (2014) A new species of leaf insect (Phasmatodea: Phylliidae) from Northern Luzon, Philippines.
Arthropoda Generalis, 5, 1–11.
Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic
trees. Gateway Computing Environments Workshop (GCE). California, La Jolla. Available from: http://www.phylo.org/
sub_sections/portal/sc2010_paper.pdf (accessed 17 September 2017)
Pérez-Ruiz, M., Martínez-Rodríguez, P., Herranz, J. & Bella, J.L. (2015) A survey of Wolbachia, Spiroplasma and other
bacteria in parthenogenetic and non-parthenogenetic phasmid (Phasmatodea) species. European Journal of Entomology,
112 (3), 409–418.
https://doi.org/10.14411/eje.2015.061
Regier, J.C. & Shi, D. (2005) Increased yield of PCR product from degenerate primers with nondegenerate, nonhomologous 5’
tails. BioTechniques, 38 (1), 34–38.
https://doi.org/10.2144/05381BM02
Rehn, J.A.G. & Rehn, J.W.H. (1934[1933]) On certain species of the genus Phyllium (Orthoptera; Phasmidae). Proceedings of
the Academy of Natural Sciences of Philadelphia, 85, 411–427.
Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. &
Huelsenbeck, J.P. (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model
space. Systematic Biology, 61 (3), 539–542.
https://doi.org/10.1093/sysbio/sys029
Rubinoff, D., Cameron, S. & Will, K. (2006) A genomic perspective on the shortcomings of mitochondrial DNA for
“barcoding” identification. Journal of Heredity, 97 (6), 581–594.
https://doi.org/10.1093/jhered/esl036
Song, H., Buhay, J.E., Whiting, M.F. & Crandall, K.A. (2008) Many species in one: DNA barcoding overestimates the number
of species when nuclear mitochondrial pseudogenes are coamplified. Proceedings of the National Academy of Sciences of
the United States of America, 105 (36), 13486–13491.
https://doi.org/10.1073/pnas.0803076105
Swofford, D.L. (2002) PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Version 4. Sinauer Associates,
Sunderland, Massachusetts. [software]
Wahlberg, N. & Wheat, C. (2008) Genomic Outposts Serve the Phylogenomic Pioneers: Designing Novel Nuclear Markers for
Genomic DNA Extractions of Lepidoptera. Systematic Biology, 57 (2), 231–242.
https://doi.org/10.1080/10635150802033006
Wedmann, S., Bradler, S. & Rust, J. (2007) The first fossil leaf insect: 47 million years of specialized cryptic morphology and
behavior. The National Academy of Sciences of the USA, 104 (2), 565–569.
https://doi.org/10.1073/pnas.0606937104
Wells, J.D., Pape, T. & Sperling, F.A. (2001) DNA-based identification and molecular systematics of forensically important
Sarcophagidae (Diptera). Journal of Forensic Science, 46 (5), 1098–1102.
https://doi.org/10.1520/JFS15105J
Westwood, J.O. (1859) Catalogue of Orthopterous insects in the collection of the British Museum. Part 1, Phasmidae. Order of
the Trustees, British Museum, London, 195 pp.
Zompro, O. (2001) Philippine phasmids from the collection of the Staatliches Museums fur Tierkunde, Dresden (Insecta:
Phasmatodea). Reichenbachia, 34 (5), 49–56.
... A similar analysis of pairwise distances of COI sequences in Phasmatodea was performed with leaf insects (Phylliidae) (Cumming et al. 2017), including five specimens of Microphyllium haskelli Cumming in Cumming et al., 2017, one of Pseudomicrophyllium pusillulum (Rehn & Rehn, 1934) and one of Pulchriphyllium giganteum (Hausleithner, 1984). The intraspecific distances of M. haskelli (≤ 2.2%) were very similar to the distances between conspecific specimens of Paraphasma (≤ 2.0%). ...
... A similar analysis of pairwise distances of COI sequences in Phasmatodea was performed with leaf insects (Phylliidae) (Cumming et al. 2017), including five specimens of Microphyllium haskelli Cumming in Cumming et al., 2017, one of Pseudomicrophyllium pusillulum (Rehn & Rehn, 1934) and one of Pulchriphyllium giganteum (Hausleithner, 1984). The intraspecific distances of M. haskelli (≤ 2.2%) were very similar to the distances between conspecific specimens of Paraphasma (≤ 2.0%). ...
... The interspecific distances (15.5-21.4%) were generally higher than those obtained in the present study (6.9-17.5%), in agreement with the fact that in the analysis of Cumming et al. (2017) each species belonged to a different genus whereas our analysis included only congeneric species. ...
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The satyrine butterfly Coenonympha tullia (Nymphalidae: Satyrinae) displays a deep split between two mitochondrial clades, one restricted to northern Alberta, Canada, and the other found throughout Alberta and across North America. We confirm this deep divide and test hypotheses explaining its phylogeographic structure. Neither genitalia morphology nor nuclear gene sequence supports cryptic species as an explanation, instead indicating differences between nuclear and mitochondrial genome histories. Sex-biased dispersal is unlikely to cause such mito-nuclear differences; however, selective sweeps by reproductive parasites could have led to this conflict. About half of the tested samples were infected by Wolbachia bacteria. Using multilocus strain typing for three Wolbachia genes, we show that the divergent mitochondrial clades are associated with two different Wolbachia strains, supporting the hypothesis that the mito-nuclear differences resulted from selection on the mitochondrial genome due to selective sweeps by Wolbachia strains.
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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.
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http://phasmida.speciesfile.org The Phasmida Species File (PSF) is a taxonomic database of the world's Phasmida (stick and leaf insects, known as walking sticks and walking leaves in the U.S.). There is full synonymic and taxonomic information for 3,350 valid species and 5,300 taxonomic names, 37,500 citations to 3,178 references, over 7,600 specimen records and 16,800 images of 75% of valid species, with more being added to on a regular basis. Another future aim of this database is to provide high quality images of living phasmids in the wild. The PSF is annually in fed into the catalogue of life (Species 2000: Naturalis, Leiden, the Netherlands).
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— We studied sequence variation in 16S rDNA in 204 individuals from 37 populations of the land snail Candidula unifasciata (Poiret 1801) across the core species range in France, Switzerland, and Germany. Phylogeographic, nested clade, and coalescence analyses were used to elucidate the species evolutionary history. The study revealed the presence of two major evolutionary lineages that evolved in separate refuges in southeast France as result of previous fragmentation during the Pleistocene. Applying a recent extension of the nested clade analysis (Templeton 2001), we inferred that range expansions along river valleys in independent corridors to the north led eventually to a secondary contact zone of the major clades around the Geneva Basin. There is evidence supporting the idea that the formation of the secondary contact zone and the colonization of Germany might be postglacial events. The phylogeographic history inferred for C. unifasciata differs from general biogeographic patterns of postglacial colonization previously identified for other taxa, and it might represent a common model for species with restricted dispersal.
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The main features of the phylogeny program TNT are discussed. Windows versions have a menu interface, while Macintosh and Linux versions are command-driven. The program can analyze data sets with discrete (additive, non-additive, step-matrix) as well as continuous characters (evaluated with Farris optimization). Effective analysis of large data sets can be carried out in reasonable times, and a number of methods to help identifying wildcard taxa in the case of ambiguous data sets are implemented. A variety of methods for diagnosing trees and exploring character evolution is available in TNT, and publication-quality tree-diagrams can be saved as metafiles. Through the use of a number of native commands and a simple but powerful scripting language, TNT allows the user an enormous flexibility in phylogenetic analyses or simulations. © The Willi Hennig Society 2008.