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Cryptophyllium, the hidden leaf insects –
descriptions of a new leaf insect genus
and thirteen species from the former celebicum
species group (Phasmatodea, Phylliidae)
Royce T. Cumming1,2,3, Sarah Bank4, Joachim Bresseel5, Jérôme Constant5,
Stéphane Le Tirant1, Zhiwei Dong6, Gontran Sonet5, Sven Bradler4
1Montreal Insectarium, 4581 rue Sherbrooke est, Montréal, Québec, Canada, H1X 2B2 2Richard Gilder
Graduate School, American Museum of Natural History, New York, NY 10024, USA 3Biology, Graduate
Center, City University of New York, NY, USA 4Department of Animal Evolution and Biodiversity, Johann-
Friedrich-Blumenbach Institute for Zoology and Anthropology, University of Göttingen, Untere Karspüle 2,
37073, Göttingen, Germany 5Royal Belgian Institute of Natural Sciences, O.D. Taxonomy and Phylogeny and
JEMU, rue Vautier 29, B-1000, Brussels, Belgium 6State Key Laboratory of Genetic Resources and Evolution,
Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, Yunnan, 650223, China
Corresponding author: Royce T. Cumming (phylliidae.walkingleaf@gmail.com);
Sarah Bank (sbank.bio@gmail.com)
Academic editor: M. Gottardo|Received 23 November 2020|Accepted 16 January 2021|Published 18 February 2021
http://zoobank.org/7E9360A5-A359-437A-91C0-04C74B1FE9D6
Citation: Cumming RT, Bank S, Bresseel J, Constant J, Le Tirant S, Dong Z, Sonet G, Bradler S (2021) Cryptophyllium,
the hidden leaf insects – descriptions of a new leaf insect genus and thirteen species from the former celebicum species
group (Phasmatodea, Phylliidae). ZooKeys 1018: 1–179. https://doi.org/10.3897/zookeys.1018.61033
Abstract
While the leaf insects (Phylliidae) are a well-supported group within Phasmatodea, the genus Phyllium
Illiger, 1798 has repeatedly been recovered as paraphyletic. Here, the Phyllium (Phyllium) celebicum species
group is reviewed and its distinctiveness from the remaining Phylliini genera and subgenera in a phyloge-
netic context based on morphological review and a phylogenetic analysis of three genes (nuclear gene 28S
and mitochondrial genes COI and 16S) from most known and multiple undescribed species is shown.
A new genus, Cryptophyllium gen. nov., is erected to partially accommodate the former members of the
celebicum species group. Two species, Phyllium ericoriai Hennemann et al., 2009 and Phyllium bonifacioi
Lit & Eusebio, 2014 morphologically and molecularly do not fall within this clade and are therefore left
within Phyllium (Phyllium). e transfer of the remaining celebicum group members from Phyllium Illiger,
1798 to this new genus creates the following new combinations; Cryptophyllium athanysus (Westwood,
ZooKeys 1018: 1–179 (2021)
doi: 10.3897/zookeys.1018.61033
https://zookeys.pensoft.net
Copyright Royce T. Cumming et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC
BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
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Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
2
1859), comb. nov.; Cryptophyllium celebicum (de Haan, 1842), comb. nov.; Cryptophyllium chrisangi
(Seow-Choen, 2017), comb. nov.; Cryptophyllium drunganum (Yang, 1995), comb. nov.; Cryptophyl-
lium oyae (Cumming & Le Tirant, 2020), comb. nov.; Cryptophyllium parum (Liu, 1993), comb.nov.;
Cryptophyllium rarum (Liu, 1993), comb. nov.; Cryptophyllium tibetense (Liu, 1993), comb. nov.; Crypto-
phyllium westwoodii (Wood-Mason, 1875), comb. nov.; Cryptophyllium yapicum (Cumming & Teemsma,
2018), comb. nov.; and Cryptophyllium yunnanense (Liu, 1993), comb. nov.
e review of specimens belonging to this clade also revealed 13 undescribed species, which are
described within as: Cryptophyllium animatum gen. et sp. nov. from Vietnam: Quang Nam Province;
Cryptophyllium bankoi gen. et sp. nov. from Vietnam: Quang Ngai, ua ien Hue, Da Nang, Gia Lai,
Quang Nam, and Dak Nong Provinces; Cryptophyllium bollensi gen. et sp. nov. from Vietnam: Ninh
uan Province; Cryptophyllium daparo gen. et sp. nov. from China: Yunnan Province; Cryptophyllium
echidna gen. et sp. nov. from Indonesia: Wangi-wangi Island; Cryptophyllium faulkneri gen. et sp. nov.
from Vietnam: Quang Ngai and Lam Dong Provinces; Cryptophyllium icarus gen. et sp. nov. from Vi-
etnam: Lam Dong and Dak Lak Provinces; Cryptophyllium khmer gen. et sp. nov. from Cambodia: Koh
Kong and Siem Reap Provinces; Cryptophyllium limogesi gen. et sp. nov. from Vietnam: Lam Dong, Dak
Lak, and Dak Nong Provinces; Cryptophyllium liyananae gen. et sp. nov. from China: Guangxi Province;
Cryptophyllium nuichuaense gen. et sp. nov. from Vietnam: Ninh uan Province; Cryptophyllium phami
gen. et sp. nov. from Vietnam: Dong Nai and Ninh uan Provinces; and Cryptophyllium wennae gen.
et sp. nov. from China: Yunnan Province. All newly described species are morphologically described, il-
lustrated, and molecularly compared to congenerics.
With the molecular results revealing cryptic taxa, it was found necessary for Cryptophyllium westwoo-
dii (Wood-Mason, 1875), comb. nov. to have a neotype specimen designated to allow accurate dieren-
tiation from congenerics. To conclude, male and female dichotomous keys to species for the Cryptophyl-
lium gen. nov. are presented.
Keywords
Description, Greek Mythology, new species, Phasmida, Phylliini, Phyllium, Southeast Asia, Vietnam
Table of contents
Introduction ............................................................................................................. 4
Materials and methods ............................................................................................. 6
Specimens .......................................................................................................... 6
Photography ...................................................................................................... 8
Literature reviewed ............................................................................................. 8
Rules of zoological nomenclature ....................................................................... 8
Molecular laboratory work and phylogenetic analysis ......................................... 8
Global Taxonomy Initiative (GTI) projects in Vietnam and Cambodia .............. 9
Results .................................................................................................................... 11
Phylogenetic analysis ........................................................................................ 11
Taxonomy ........................................................................................................ 11
Phylliidae Brunner von Wattenwyl, 1893 ......................................................... 11
Species boundary interpretations ...................................................................... 12
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 3
Cryptophyllium gen. nov. ............................................................................ 14
Key to females of the genera of the Phylliini Brunner von Wattenwyl, 1893..... 16
Key to males of the genera of the Phylliini Brunner von Wattenwyl, 1893 ....... 16
Key to eggs of the genera of the Phylliini Brunner von Wattenwyl, 1893 ......... 17
Generic characteristics ...................................................................................... 18
Species recorded per country ............................................................................ 23
Cryptophyllium animatum gen. et sp. nov.................................................... 25
Cryptophyllium athanysus (Westwood, 1859), comb. nov. ........................... 30
Cryptophyllium bankoi gen. et sp. nov. ........................................................ 31
Cryptophyllium bollensi gen. et sp. nov. ....................................................... 38
Cryptophyllium celebicum (de Haan, 1842), comb. nov. .............................. 51
Cryptophyllium chrisangi (Seow-Choen, 2017), comb. nov. ........................ 54
Cryptophyllium daparo gen. et sp. nov. ........................................................ 58
Cryptophyllium drunganum (Yang, 1995), comb. nov. ................................ 62
Cryptophyllium echidna gen. et sp. nov. ....................................................... 65
Cryptophyllium faulkneri gen. et sp. nov. ..................................................... 69
Cryptophyllium icarus gen. et sp. nov. ......................................................... 73
Cryptophyllium khmer gen. et sp. nov. ......................................................... 85
Cryptophyllium limogesi gen. et sp. nov. ...................................................... 97
Cryptophyllium liyananae gen. et sp. nov. .................................................. 106
Cryptophyllium nuichuaense gen. et sp. nov. .............................................. 111
Cryptophyllium oyae (Cumming & Le Tirant, 2020), comb. nov. .............. 116
Cryptophyllium parum (Liu, 1993), comb. nov. ........................................ 120
Cryptophyllium phami gen. et sp. nov. ....................................................... 121
Cryptophyllium rarum (Liu, 1993), comb. nov. ......................................... 132
Cryptophyllium tibetense (Liu, 1993), comb. nov. ...................................... 137
Cryptophyllium wennae gen. et sp. nov. ..................................................... 142
Cryptophyllium westwoodii (Wood-Mason, 1875), comb. nov. .................. 145
Cryptophyllium yapicum (Cumming & Teemsma, 2018), comb. nov. ....... 155
Cryptophyllium yunnanense (Liu, 1993), comb. nov. ................................. 157
Cryptophyllium spp. which are not yet described or identied ................... 162
Key to known males in the Cryptophyllium gen. nov. ...................................... 164
Key to known females in the Cryptophyllium gen. nov. ................................... 166
Phylliidae conservation in Vietnam ................................................................ 168
Discussion ............................................................................................................ 169
Acknowledgements ............................................................................................... 172
References ............................................................................................................ 173
Supplementary material 1 ..................................................................................... 178
Supplementary material 2 ..................................................................................... 178
Supplementary material 3 ..................................................................................... 178
Supplementary material 4 ..................................................................................... 179
Supplementary material 5 ..................................................................................... 179
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
4
Introduction
Phylliidae comprise the true leaf insects, a subordinated clade within the plant mim-
icking lineage of Phasmatodea (Bradler and Buckley 2018; Simon et al. 2019). While
most phasmatodeans exhibit an elongated and slender body form to camouage per-
fectly in the foliage, among branches or on bark (Bedford 1978), leaf insects mastered
the imitation of angiosperm leaves (Fig. 1).
Of the ve phylliid genera, the majority of species are attributed to Phyllium, which
comprises 65 of the 89 currently described and valid Phylliidae species (Brock et al.
2020). In previous studies on Phasmatodea, the Phylliidae were always recovered as
monophyletic whereas Phyllium itself appeared to be paraphyletic (Buckley etal. 2009;
Bradler et al. 2015; Robertson et al. 2018). us, the Phyllium are in need of a thorough
phylogenetic analysis. Taxonomically, Phyllium is further divided into four subgenera:
Phyllium, Pulchriphyllium, and the recently described Comptaphyllium and Walaphyl-
lium (Cumming et al. 2019; Cumming et al. 2020b). For the two traditional subgenera
Phyllium and Pulchriphyllium, an intra-generic systematization had been proposed in
order to facilitate dierentiation in a taxonomical context (Hennemann et al. 2009).
e subdivision of the most diverse subgenus Phyllium (Phyllium) into the siccifolium
and celebicum species groups was mainly based on the presence of developed alae in
females of the latter, a problematic character in a group with strong sexual dimorphism
and several species only known from single sexes (e.g., Liu 1993; Yang 1995).
One method that has allowed condent matching of opposite sexes is through cap-
tive cultures as was demonstrated for the Nanophyllium (described only from males)
whose opposite sex was found to be already described in the Phyllium (Cumming et
al. 2020c). Although laboratory rearing is substantial to ascertain the useful taxonomic
trio of female, male, and egg morphology, and therefore an important factor for taxo-
nomic classication, it has its limitations. With the aid of molecular data, in addition
to matching up sexes and identifying new species, it is also possible to uncover poten-
tial cryptic species (Cumming et al. 2020a).
Since 2010, sta from the Royal Belgian Institute of Natural Sciences (RBINS),
led by Jérôme Constant, have participated in expeditions to document the phasmid
fauna of Vietnam and Cambodia in the framework of the Global Taxonomy Initiative
(GTI) projects “A step further in the entomodiversity of Vietnam” (ten expeditions,
2010–2019) and “A step further in the entomodiversity of Cambodia” (three expe-
ditions, 2016–2018) (Constant et al. 2018). e expeditions were jointly organized
with the Institute of Ecology and Biological Resources (2010–2014) and the Vietnam
National Museum of Nature (2015–2019), which are both part of the Vietnam Acad-
emy of Science and Technology, in Vietnam, and with the Royal University of Phnom
Penh in Cambodia. For each expedition, the local institutions organized the collecting
permits and ocial paperwork. GTI was created in the framework of the Convention
on Biological Diversity to remove the ‘taxonomic impediment’ (www.cbd.int/gti). e
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 5
Figure 1. Paratype Cryptophyllium phami sp. nov. where it was found in Cat Tien N.P. (Vietnam) feed-
ing on a Guava tree (Psidium guajava) in July 2012 by Jérôme Constant (RBINS) and Joachim Bresseel
(RBINS). e photograph was taken without ash to show how they appear naturally from underneath
with the light passing through their thin bodies like the light passing through a leaf.
goals of these GTI projects organized by RBINS and partners are to document and
describe the biodiversity, to provide taxonomy capacity building (in the eld and in
the lab) for Vietnamese and Cambodian students, and to assist and advise about the
management of insect collections of the participating institutions. rough the eorts
of the many participants of these expeditions, many scientically signicant specimens
from the celebicum species group were discovered and utilized herein.
e celebicum species group currently contains 13 of the 42 species of the Phyl-
lium (Phyllium) with a distribution spanning from Sri Lanka over southern China
and mainland Southeast Asia to the Philippines, Sulawesi, and Micronesia (Fig. 2).
Here, we aim to review the Phyllium (Phyllium) celebicum species group by extensive
investigation of morphology as well as analysis of molecular data to test whether the
taxon represents a monophyletic group, to conrm matching of sexes, to identify un-
described species, and reveal potential hidden (cryptic) diversity within the lineage. We
also formally describe any new species for which species status appears warranted based
on the obtained morphological and/or molecular data.
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
6
Figure 2. Distribution map for the 24 Cryptophyllium gen. nov. species presently known (with solid lines
from their name pointing to the type locality) as well as additional Cryptophyllium gen. nov. species which
we could not herein describe/dierentiate (indicated by dashed lines). Note that the line for Cryptophyl-
lium westwoodii comb. nov. is pointing to the neotype locality and the type locality for Cryptophyllium
athanysus comb. nov. is simply “Ceylon” therefore the line is pointing to the present-day localities we are
aware of. Inset is of southern Vietnam showing the distributions of three additional species which could
not t within the main map. e colors in this map are noted to the left of the names within the phyloge-
netic tree in Fig. 4. Newly herein described species have names noted in bold. Note that with Cryptophyl-
lium khmer sp. nov. not easily distinguishable from Cryptophyllium westwoodii comb. nov. from photos
alone, only the locations for these two species where they were genetically sampled are solid colored, all
observational images without genetic sampling have bicolored circles and could represent either of these
species. Additionally, one symbol is split three ways for Cryptophyllium gen. nov. specimens from the ai
oshore islands of Ko Phangan and Ko Samui which could not be dierentiated from C. chrisangi comb.
nov., C. westwoodii comb. nov., and C. khmer sp. nov. from images alone. See Suppl. material 4 for a full
list of the specimens/observations utilized to make the distribution map with deposition data for speci-
mens and links to observational records.
Materials and methods
Specimens
We examined and sequenced numerous specimens from both institutional and private
collections for this review. Many of the specimens, which were sequenced for this work,
are illustrated but not all specimens that were sequenced are included as gures. To ensure
reproducibility for future investigators, images and institution accession numbers of ad-
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 7
ditional specimens which were sequenced but not illustrated within the main body of this
work can be found in Suppl. material 1. For each species discussed within we explicitly
state the number of specimens reviewed, their collection data, and the collection they
are contained within to allow the specimens utilized for this work to be located by other
interested researchers to ensure reproducibility of our work. In multiple cases we were pre-
sented with photographs of individuals which we based morphological comparisons upon,
in these instances many of these images were reproduced and referenced herein or their
source is stated to allow the reviewed images to be located. Additionally, we utilized the
extensive collection of high-quality images of captive reared specimens available publicly
on Phasmatodea.com “e World of Stick Insects” (http://www.phasmatodea.com/taxon-
omy/term/1788) as well as the many type material images available on “Phasmida Species
Files Online” (http://phasmida.speciesle.org/HomePage/Phasmida/HomePage.aspx).
Measurements of specimens were made to the nearest 0.1 mm using digital calipers.
Measurements for paratype specimens are given with a minimum to maximum range.
Holotype and paratype specimens are deposited within several institutions which are
explicitly listed within the type material information of the new species descriptions.
e following collection acronyms are used. Specimen data listed within quotations is
verbatim and is therefore in a variety of non-standard formats, but is presented as is to
ensure traceability of specimens utilized.
CAS California Academy of Sciences, San Francisco, California, USA;
BPBM Bishop Museum, Honolulu, Hawaii, USA;
IMQC Insectarium de Montréal, Montréal, Québec, Canada;
KIZ Kunming Institute of Zoology, Yunnan, China;
LKCNHM Lee Kong Chian Natural History Museum, Singapore;
MNHN Muséum national d’Histoire naturelle, Paris, France;
NHMUK Natural History Museum United Kingdom, London, United Kingdom;
NZSI Zoological Survey of India, National Zoological Collection, Calcutta, India;
OUMNH University Museum of Natural History, Oxford, United Kingdom;
RBINS Royal Belgian Institute of Natural Sciences, Brussels, Belgium;
RMNH Naturalis Biodiversity Centre, Leiden, Netherlands;
RUPP Royal University of Phnom Penh, Phnom Penh, Cambodia;
SDEI Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany;
UCR University of California Riverside, California, USA;
VNMN Vietnam National Museum of Nature, Hanoi, Vietnam;
GTI Global Taxonomy Initiative (https://www.cbd.int/gti/);
Coll FH Private collection of Frank H. Hennemann, Germany;
Coll MO Private collection of Maxime Ortiz, France;
Coll OC Private collection of Oskar V. Conle, Duisburg, Germany;
Coll RC Private collection of Royce T. Cumming, California, USA;
Coll SLT Private collection of Stéphane Le Tirant, Québec, Canada;
Coll TB Private collection of ies Büscher, Kiel, Germany;
Coll ZD Private collection of Zhiwei Dong, Yunnan, China.
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
8
Photography
Photographs of specimens deposited within the IMQC collection were taken by René
Limoges using a Nikon D850 DSLR camera (Nikon Corporation, Tokyo, Japan) with
Nikon Micro-Nikkor 200mm f/4 lens on Manfrotto 454 micrometric positioning
sliding plate (Manfrotto, Casolla, Italy). Lighting was provided by two Nikon SB-25
ash units with a Cameron Digital diusion photo box (Henry’s, Vancouver, Canada).
Adobe Photoshop Elements 13 (Adobe Inc., San Jose, USA) was used as post-process-
ing software.
Photographs of specimens within the RBINS collection were taken by Jérôme
Constant (RBINS). For each specimen a number of photographs were taken with a
Canon 700D camera (Canon Inc., Ota City, Tokyo, Japan) equipped with a Sigma
50 mm Macro lens (Sigma Corporation, Kawasaki, Japan) for adults, or with a Leica
EZ4W stereomicroscope (Leica Microsystems Ltd., Wetzlar, Germany) with integrat-
ed camera for eggs and vomers, and stacked with CombineZ software (https://combin-
ezp.software.informer.com) and optimized with Adobe Photoshop software.
Additional photographs used which are not from these two above institutions are
explicitly listed within the gure captions with citation to the photographers.
Literature reviewed
All original species descriptions for relevant species were reviewed, and in several cases
when not originally published in English the works were translated. To translate the
relevant works of Liu (1993), Yang (1995), and Sorpongpaisal and anasinchayakul
(2006), we used the most recent version of a phone-based application “Google Trans-
late’’ (oered by Google LLC: version 6.2.0.RC07.268294262).
Rules of zoological nomenclature
We here follow the International Code of Zoological Nomenclature (ICZN 1999) as
a means of describing new phylliid taxa and designating the neotype Cryptophyllium
westwoodii (Wood-Mason, 1875), comb. nov.
Molecular laboratory work and phylogenetic analysis
Tissue samples from 56 specimens of the Phyllium (Phyllium) celebicum species group
as well as two outgroup samples (Microphyllium haskelli and Pseudomicrophyllium
faulkneri) were obtained to generate molecular data for phylogenetic analysis. DNA
extraction, PCR amplication of cytochrome oxidase subunit I (COI), 16S rRNA
(16S) and parts of 28S rRNA (28S), and subsequent Sanger-sequencing was carried out
following the protocol given by Cumming et al. (2020a). For two samples (RBINS13
and RBINS14), DNA was extracted using the NucleoSpin Tissue kit (MACHEREY-
NAGEL, Düren, Germany) and eluted in 100 µl elution buer and Sanger-sequencing
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 9
was run on an ABI3130xl Genetic Analyzer (Applied Biosystems, Foster City, USA;
Laboratory of Molecular Systematics of the RBINS, Brussels, Belgium). We decided
that for most samples, especially possible duplicates, it is sucient to sequence only
one gene. DNA sequences were checked for quality and edited in Geneious Prime ver-
sion 2020.0.4 (www.geneious.com) prior to deposition in GenBank (accession num-
bers MW161173–MW161229, MW165169–MW165218, see Suppl. material 5).
For the phylogenetic inference, we furthermore included sequence data of nine
additional phylliid species and the three outgroup species published by Cumming
et al. (2020a). Nucleotide sequences of a total of 70 specimens were aligned sepa-
rately for each gene using the G-INS-I method of MAFFT version 7.453 (Katoh and
Standley 2013). e alignments were manually trimmed at the beginning and end
before concatenation with FASconCAT (Kück and Meusemann 2010). e phyloge-
netic inference was conducted in IQ-TREE version 2.1.1 (Minh et al. 2020) with a
random starting tree and the best-t substitution model selected by ModelFinder un-
der AICc (GTR+F+R5, Kalyaanamoorthy et al. 2017). Node support was estimated
using the Ultrafast Bootstrap approximation and 10,000 pseudo-replicates (UFBoot,
Hoang et al. 2018). e resulting tree was visualized in FigTree v. 1.4.4 (https://
github.com/rambaut/gtree) and rooted with Aschiphasmatinae (Orthomeria kangi
Vallotto, Bresseel, Heitzmann & Gottardo, 2016), which has been repeatedly re-
covered as sister group to the remaining Euphasmatodea (= Neophasmatodea) (e.g.,
Robertson et al. 2018; Simon et al. 2019).
Global Taxonomy Initiative (GTI) projects in Vietnam and Cambodia
Since 2010 a total of 206 days was spent in the eld in Vietnam, with 32 locations sam-
pled for stick/leaf insects leading to eleven collecting events (place and time where col-
lecting occurred) involving leaf insects; in Cambodia 36 days of eldwork in eight loca-
tions provided a single collecting event (Fig. 3). In total, eight species of leaf insect were
collected, including seven yet to be described, and an additional young nymph that was
unidentiable, which did not survive. e collecting of stick and leaf insects is always
better at night as these insects are mostly nocturnal. However, it has also been possible
to nd them during the day when keeping in mind important aspects of their biology,
for example that guava trees (Psidium guajava L., Myrtaceae) are an attractive food plant
for the leaf insects which can be located by searching for their telltale feeding damage
amongst the leaves (Fig. 1). Additionally, knowing the local name of the guava tree (cây
ổi in Vietnamese; ដ ើមត្របែក “derm trobaek / derm trorbek” in Khmer) and of the
leaf insects (bọ lá in Vietnamese; កណ្ដូបឈើ or កណ្ដូបស្លឹក “kandob chheu”
or “kandob sloek” in Khmer) (H.T. Pham, P. Heang, and B. Doeurk pers. comm.) and
interacting with local people proved a useful way to locate potential leaf insect habitats
(Fig. 1). Being canopy dwellers, leaf insects are usually out of reach in the forest and have
proven easier to nd in anthropized habitats such as along roads (Fig. 32), on regrowth
in recently burnt or logged areas (Figs 16, 48), or even on cultivated trees (mostly guava
trees) near houses (Fig. 37). Adult male specimens, the only sex able to y, were found
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
10
Figure 3. Map of Vietnam and Cambodia where GTI joint expeditions between the teams from RBINS,
the Institute of Ecology and Biological Resources (Vietnam), the VNMN, and the RUPP in Cambodia
have conducted eldwork between 2010 and 2019. All sampled locations are marked, with the green stars
indicating locations where leaf insect specimens were located and the red circles denoting sampled sites
where leaf insects were not encountered.
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 11
attracted to lights a night. During the expeditions, males of Cryptophyllium phami gen.
et sp. nov., Cryptophyllium khmer gen. et sp. nov., Cryptophyllium rarum comb. nov.,
and Cryptophyllium westwoodii comb. nov. were sampled using a light trap with mercury
vapor lamps. One male of Cryptophyllium rarum comb. nov. was collected when it was
attracted to the streetlights in Cuc Phuong N.P. Field experience has shown that it is
important to check the surroundings of the light trap as it seems that males are attracted
to the lights, but not necessarily likely to alight on the white sheet like other nocturnal
insects. Due to the somewhat feeble nature of males’ ight, we expect that only males
in the vicinity of the light trap are attracted, not ying from great distances to reach it.
Whenever possible the female specimens were kept alive for several days to allow
them to lay eggs which provide supplemental morphological characters to dierentiate
species. Additionally, these eggs were shared with enthusiastic phasmid breeders in Bel-
gium and Switzerland who were instrumental in documenting the nymphal stages and
in obtaining important specimens such as males of species of which only the female
was found in the wild, rearing additional specimens to study intraspecic variation, or
when species were only collected as nymphs from the wild these breeders helped by
rearing them to adulthood.
Results
Phylogenetic analysis
e morphological review of all examined specimens revealed 20 species of which eleven
could be dierentiated from already described species. In order to conrm our assump-
tions, we conducted a phylogenetic analysis based on a molecular dataset comprising
1804 characters resulting from the concatenation of 69 sequences of the COI, 39 of the
16S, and 33 of the 28S gene. e phylogenetic inference under Maximum Likelihood
(Figure 4) recovered Phylliidae as monophyletic with maximum support. e Phyllium
was recovered paraphyletic with Microphyllium + Pseudomicrophyllium as sister group to
the Phyllium (Phyllium) celebicum species group. e specimens assigned to this group,
here described as the Cryptophyllium gen. nov. based on morphology, formed a strongly
supported monophyletic group. Species were recovered as distinct clades as initially pro-
posed by morphology. In two cases, the molecular phylogeny revealed cryptic species.
Taxonomy
Phylliidae Brunner von Wattenwyl, 1893
Phylliidae can be dierentiated from other phasmids by the broad, dorsolaterally at-
tened abdomen and attened femoral lobes (and for many taxa lobes on the tibia as
well), giving them a distinct leaf-like appearance; a protuberance on the attachment
site in the head capsule from the dorsal tentorial arms; and dorsoventrally attened
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
12
maxillary- and labial palps (Bedford 1978; and see other notable works for additional
discussion on phylliid apomorphies: Bradler 2009; Klug and Bradler 2006; Friede-
mann et al. 2012). Additionally, to add believability to the leaf shape, the phylliids
come in leaf-like colors such as green, brown, yellow, and a wide variety of mixed
color forms and even sway as they walk like leaves in the wind. Currently, there are
two divisions of leaf insects within the singular Phylliinae, the Phylliini Brunner von
Wattenwyl, 1893 (which contains the bulk of genera and species) and the Nanophyl-
liini Zompro & Groesser, 2003 (which only contains the Nanophyllium Redtenbacher,
1906) (Brock et al. 2020). Our molecular and morphological review has warranted the
description of a novel genus of leaf insect to accommodate the former celebicum species
group (sensu Hennemann et al. 2009), which is erected herein.
Species boundary interpretations
We determined our species boundaries based upon molecular, morphological, and
geographic data. Although this was not done objectively through the use of species
delimitation programs, we felt that due to the gaps in our sampling, a case-by-case
delimitation was necessary based on a thorough understanding of the taxa in question.
In some instances, species boundaries were easy to interpret as molecular and mor-
phological data clearly agreed with little room for abstract interpretation. For example,
note the clade in Fig. 4 containing Cryptophyllium limogesi gen. et sp. nov., Cryptophyl-
lium icarus gen. et sp. nov., and Cryptophyllium bankoi gen. et sp. nov. where these
three species are 1) molecularly unique, 2) morphologically easily distinguishable, and
3) geographically sympatric (see inset within Fig. 2).
However, not all species boundary interpretations were as clearly observed, and
our boundary lines are less distinctly dened for some clades (e.g., see Fig. 4 with
our treatment of Cryptophyllium westwoodii comb. nov., Cryptophyllium rarum comb.
nov., or the clade formed by Cryptophyllium bollensi gen. et sp. nov., Cryptophyllium
nuichuaense gen. et sp. nov., and Cryptophyllium phami gen. et sp. nov.). In instances
where our molecular results gave ambiguous delimitations, we turned to morphology
and biogeography to help clarify our species boundaries for this work.
Within the instances of Cryptophyllium westwoodii comb. nov. and Cryptophyllium
rarum comb. nov. we found these to be geographically widespread populations with
notable molecular variability (which one would expect from a widespread popula-
tion of not particularly mobile insects). When reviewing the morphology of these two
groups we were unable to identify a consistent and reliable morphological feature for
dierentiation to warrant splitting of these clades into multiple species. Additionally,
we maintain these somewhat molecularly variable clades due to our present lack of ex-
tensive molecular sampling from throughout their large geographic distributions. For
example, in Figure 4 note the Cryptophyllium westwoodii comb. nov. sample 18-029
from Karathuri, Myanmar. Although this sample has a signicant molecular distance
from the northern ailand samples to which it is most closely related, we lack ad-
ditional molecular data from the over 800 kilometers between this location and the
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 13
Figure 4. Phylogenetic relationships among 35 phylliid species (67 specimens). e species of the Cryp-
tophyllium gen. nov. are highlighted with colors and symbols corresponding to the distribution map
(Fig.2). Support values based on maximum likelihood (UFBoot, IQ-TREE) are indicated at each node.
samples from the north. Considering the lack of morphological features for dierentia-
tion and the possibility that within this signicant 800 kilometer range is a continuous
population lling in the molecular distances between these north/south extremes, we
felt a conservative approach was best. erefore, even though these expansive clades
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
14
may in fact represent more than one species in reality, we felt it premature to split these
clades apart and hope that future molecular analyses from these missing geographic
areas can eventually bring greater clarity to these populations.
In contrast, similar molecular distances were observed within the clade formed by
Cryptophyllium bollensi gen. et sp. nov., Cryptophyllium nuichuaense gen. et sp. nov.,
and Cryptophyllium phami gen. et sp. nov. as were observed within the diverse Cryp-
tophyllium westwoodii comb. nov. or Cryptophyllium rarum comb. nov. but with the
important caveats that 1) these three species had notable molecular distance between
them, 2) these three could reliably and consistently be morphologically separated, and
3) due to their geographic proximity and extensive sampling within this area we do
not have large areas of unsampled uncertainty (with the distance between these three
species ranging from only 60–100 kilometers apart).
erefore, although our interpretations at the present may not perfectly represent
biological reality, we feel that due to limited data a conservative and individual ap-
proach was best for species delimitation within this work. For additional details on our
species boundary determinations please see the individual species sections below for
individual discussions where warranted.
Cryptophyllium gen. nov.
http://zoobank.org/35EF6752-C08C-4583-8516-A7AA3167B704
Type species. Phyllium celebicum de Haan, 1842: 111, herein designated.
Dierentiation. e only single feature that alone can distinguish Cryptophyllium
gen. nov. from others phylliids is the vomer of the males, which has two apical hooks
(Fig. 5). Größer (2011) was the rst to review the vomer morphology across a wide
sampling of species and noted that the vomer of males in this group is two-hooked,
whereas all other known phylliid males have a singularly hooked vomer. Besides
this distinct feature, it is a combination of other features, not a singular feature,
which allows dierentiation of Cryptophyllium gen. nov. from the other phylliid
genera. Morphologically, Cryptophyllium gen. nov. falls within the Phylliini due to
the posteromedial tubercle being singular, not split into two lobes (a feature which
dierentiates Phylliini and Nanophylliini; Cumming et al. 2020c).
Genera and subgenera included within the Phylliini Brunner von Wattenwyl,
1893 are: Chitoniscus Stål, 1875; Microphyllium Zompro, 2001; Pseudomicrophyllium
Cumming, 2017; Phyllium (Phyllium) Illiger, 1798; Phyllium (Pulchriphyllium) Gri-
ni, 1898; Phyllium (Comptaphyllium) Cumming, Le Tirant, & Hennemann, 2019;
Phyllium (Walaphyllium) Cumming, urman, Youngdale & Le Tirant, 2020, and the
herein described Cryptophyllium gen. nov. To dierentiate Cryptophyllium gen. nov.
from other Phylliini and place it taxonomically we present a key to genera for each sex
individually as well as a key to egg morphology.
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 15
Figure 5. Details of the Cryptophyllium gen. nov. male vomer. All from the RBINS collection, prepared
and photographed by Jérôme Constant (RBINS). Views 1 ventral 2 right lateral of the ventrally oriented
vomer 3 left lateral of the ventrally oriented vomer A Cryptophyllium westwoodii comb. nov., ailand,
Na Haeo B Cryptophyllium khmer gen. et sp. nov. Cambodia, Kbal Spean C Cryptophyllium phami gen. et
sp. nov., Vietnam, Cat Tien D Cryptophyllium khmer gen. et sp. nov., Cambodia, Tatai E Cryptophyllium
bollensi gen. et sp. nov., Vietnam, Phuoc Binh F Cryptophyllium oyae comb. nov., Laos, Mt. Phu Phan
GCryptophyllium bankoi gen. et sp. nov., Vietnam, Ba Na-Nui Chua H Cryptophyllium icarus gen. et
sp.nov., Vietnam, Bidoup-Nui Ba I Cryptophyllium rarum comb. nov., Vietnam, Cuc Phuong J Crypto-
phyllium limogesi gen. et sp. nov., Vietnam, Ta Dung. Scale bars, 1.0 mm.
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
16
Key to females of the genera of the Phylliini Brunner von Wattenwyl, 1893
Unknown for Pseudomicrophyllium Cumming, 2017
1 Tegmina venation with the posterior cubitus split into an anterior cubitus (CuA),
rst posterior cubitus (CuP1), and second posterior cubitus (CuP2) .....................
................................................Phyllium (Walaphyllium) Cumming et al. 2020
– Tegmina cubitus venation simple (unsplit) or bifurcate (into an anterior cubitus
(CuA) and posterior cubitus (CuP1) only) ..........................................................2
2 Terminal antennomere as long as the preceding four or ve segments combined ....3
– Terminal antennomere as long as the preceding one or two segments combined ....4
3 Prescutum sagittal plane with two prominent spines, one on the anterior rim and
another along the sagittal plane in the middle of the length; small species (42.0–
47.0 mm long); restricted to the Philippines ........ Microphyllium Zompro, 2001
– Prescutum anterior rim with a prominent singular spine, the remainder of the sag-
ittal plane of the prescutum with small nodes; small to large species (56.0–102.0
mm long) ............................ Phyllium (Comptaphyllium) Cumming et al. 2019
4 Prescutum which is wider than long (ca. 2× wider) ...........Chitoniscus Stål, 1875
– Prescutum which is the same width as length, or notably longer than wide ......... 5
5 Tegmina with media and cubitus veins running side by side and touching through-
out the majority of their lengths ....... Phyllium (Pulchriphyllium) Grini, 1898
– Tegmina with media and cubitus veins distinctly separated with several vein widths
distance between them throughout the length, not touching ............................... 6
6 Antennae typically with the fourth segment short and disk-like at least 3× wider
than long and notably shorter than any of the following three segments, or rarely a
similar length to the following segment, but still at least 2× as wide as long ..........
....................................................................................Cryptophyllium gen. nov.
– Antennae with the fourth antennal segment as tall as wide and of a similar height
to each of the following three segments length, not notably shorter ......................
.......................................................................Phyllium (Phyllium) Illiger, 1798
Key to males of the genera of the Phylliini Brunner von Wattenwyl, 1893
1 Small (< 30.0 mm in length); protibiae lacking an interior lobe; restricted to the
Philippines .......................................................................................................... 2
– Medium to large (35.0 mm to > 80.0 mm); protibiae almost always with a half to
fully developed interior lobe, or rarely highly reduced to a sliver on the proximal
half only .............................................................................................................. 3
2 Antennae short (only ca. the length of the outstretched front legs), with bead-like
antennomeres that are no more than 2× longer than they are wide ........................
............................................................................ Microphyllium Zompro, 2001
– Antennae notably longer than the outstretched front legs, with antennomeres 4–5×
longer than wide ....................................Pseudomicrophyllium Cumming, 2017
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 17
3 Prescutum stout, ca. 2× as wide as long .............................Chitoniscus Stål, 1875
– Prescutum as long as wide or notably longer than wide ....................................... 4
4 Vomer with two apical hooks .......................................Cryptophyllium gen. nov.
– Vomer with a single apical hook .......................................................................... 5
5 Alae radial sector, media anterior, and media posterior veins fusing to the cubitus
at dierent locations along the vein and running together to the wing margin ......
........................................................ Phyllium (Pulchriphyllium) Grini, 1898
– Alae radial sector, media anterior, and media posterior not fusing with the cubitus ....6
6 Tegmina media vein splits into the anterior media vein (MA) and posterior media
vein (MP) very early on, immediately or at most ¼ of the way through the wing
length and they run unbranched and subparallel through the wing length; protibial
interior lobe not reaching from end to end of the shaft, only restricted to the proxi-
mal ½ to ⅔ but never more; a head capsule with clearly dened nodes arranged in
evenly spaced patterns ......... Phyllium (Comptaphyllium) Cumming et al. 2019
– Tegmina media vein running unbranched for the rst ⅓ to ⅖ of the wing length,
and then branching with either a single short media posterior running to the wing
margin or two short media posteriors branching from the notably longer media
anterior and running to the margin; protibial interior lobe variable, either fully
spanning the full length or only ½ of the length; head capsule at most with random
granulation but frequently bare ........................................................................... 7
7 Abdominal shape rectangular, with segments V and VI with fully parallel-sided
margins (segments IV and VII with only half parallel-sided and the remainder
curved) .....................................Phyllium (Walaphyllium) Cumming et al. 2020
– Abdominal shape variable, either spade-shaped (with the margins of V parallel or
strongly converging and segment VI strongly converging), ovular (with margins
expanding and then contracting, no segments parallel-sided), thin and slender with
converging margins, bell-shaped (with margins expanding until segment VI then
strongly converging) or boxy with only segment V parallel-sided (segments IV and
VI only partially parallel-sided, the remainder rounded .........................................
.......................................................................Phyllium (Phyllium) Illiger, 1798
Key to eggs of the genera of the Phylliini Brunner von Wattenwyl, 1893
Unknown for Microphyllium Zompro, 2001 and Pseudomicrophyllium Cumming, 2017
1 Surface lacking pinnae, instead porous and stiy spongy .................................... 2
– Surface with pinnae (moss-, rope- or feather-like) ...............................................4
2 Lateral margins fanned out into distinct ns with an operculum which is typically
longer than wide (but not always), or if the capsule ns are reduced (not promi-
nently protruding), the egg in cross-section is distinctly triangular (not pentagonal
or rectangular), with the dorsal surface notable broader than the other surfaces ....
........................................................ Phyllium (Pulchriphyllium) Grini, 1898
– Capsule boxy and rectangular without distinct ns and the operculum is always
notable wider than long (length ca. ½ the greatest width) ...................................3
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18
3 Eggs medium to large, 5.0–7.0 mm long ..............................................................
................................................Phyllium (Walaphyllium) Cumming et al. 2020
– Eggs small, ca. 3.0 mm long .............................................. Chitoniscus Stål, 1875
4 Pinnae short and moss-like over the entire capsule............Cryptophyllium gen. nov.
– Pinnae long and feather- or rope-like ..................................................................5
5 Operculum with a row of pinnae along the sagittal plane, not pinnae encircling the
margin of the operculum ..... Phyllium (Comptaphyllium) Cumming et al. 2019
– Operculum with pinnae encircling the margin, not along the sagittal plane ...........
.......................................................................Phyllium (Phyllium) Illiger, 1798
Generic characteristics
In general, the Cryptophyllium gen. nov. are medium to large, with females ranging
from 77.0 mm (in the smallest recorded Cryptophyllium athanysus comb. nov.; Henne-
mann et al. 2009) to 107.0 mm long (in the only recorded Cryptophyllium daparo
sp.nov.), and males from 55.4 mm (in the smallest recorded Cryptophyllium bollensi sp.
nov.) to 89.4 mm (in the only recorded Cryptophyllium animatum sp. nov.). Both sexes
only have interior tibial lobes on the protibiae and some species either have full exterior
meso-, and metatibial exterior lobes; lobes on the pro-, meso-, and metatibial exterior,
which are small and reduced to the distal end only; or have meso-, and metatibiae,
which are completely devoid of exterior lobes. Females have antennae with nine seg-
ments of which the fourth antennal segment is short and disk-like (Fig. 6) and males
have antennae which range between 24 and 29 segments. Exterior profemoral lobes
on both sexes can be rounded and obtuse, roundly angled, distinctly right angled, or
even acutely angled with recurved exterior angles. Interior profemoral lobes can have
uniformly sized and spaced teeth or doubly serrate and triangular teeth in a variety of
arrangements, but generally there are ve teeth arranged in a two-one-two pattern.
e thorax in both sexes is variable with mesopleura that can be narrow on the ante-
rior and only fanning out halfway through the length, or on the other extreme, broad
and uniformly diverging throughout the entire length, with some species exhibiting a
somewhat intermediate range of shapes. Female tegmina are always long and have a
subcoastal vein, radial vein split into the rst radial and radial sector and ending in a
small radial to medial crossvein, a bifurcate medial vein, a bifurcate cubitus vein, and
a rst anal which fuses with the cubitus early on (Fig. 7A). Females can have highly
reduced alae (rare) or more commonly decently developed alae which can be half to al-
most the same length as the tegmina. Male tegmina range in length slightly with most
species having tegmina, which reach onto abdominal segment III or less commonly
longer onto abdominal segment IV. Male tegminal venation has two media posterior
veins and the cubitus is unbranched. Male alae are always long and fully developed
with a set of coastal and subcoastal veins on the anterior margin, a radial split into a
rst radius and radial sector which do not reconnect before terminating on the wing
margin, a media which splits into the media anterior and media posterior but then
reconnects and runs fused to the wing margin, and a typical set of a cubitus and anal
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 19
Figure 6. Details of female phylliid antennae illustrated from specimens within the RBINS collection, il-
lustrations by Julien Caudron (RBINS). Note that the gures start with the scapus, therefore the diagnostic
fourth antennomere is the third one shown; to ensure clarity A has the segment number listed to the right
of the image. Note that the fourth antennomere of Phyllium (A) is longer than the fth antennomere in
comparison to the Cryptophyllium gen. nov. representatives (B–H), which have the fourth antennomere
short and disk-like A Phyllium ericoriai, Philippines B Cryptophyllium bollensi gen. et sp. nov., Vietnam,
Phuoc Binh C Cryptophyllium celebicum comb. nov., Indonesia, Sulawesi D Cryptophyllium khmer gen. et
sp. nov., Cambodia, Siem Reap Province E Cryptophyllium nuichuaense gen. et sp. nov., Vietnam, Nui Chua
F Cryptophyllium phami gen. et sp. nov., Vietnam, Cat Tien G Cryptophyllium westwoodii comb. nov., Laos,
Bokeo Province H Cryptophyllium westwoodii comb. nov., ailand, Chiang Mai. Drawn to relative scale.
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
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Figure 7. Typical tegmina and alae venation present in the Cryptophyllium gen. nov., photographed by
RTC A female: Cryptophyllium celebicum comb. nov. (Coll RC 16-075) B male: Cryptophyllium yunnan-
ense comb. nov. (Coll RC 16-120). Abbreviations: C (costa); Sc (subcosta); R (radius); R1 (radius 1); R2
(radius 2); Rs (radial sector); M (media); MA (media anterior); MP (media posterior); MP1 (rst media
posterior); MP2 (second media posterior); MA+MP (media anterior fused with media posterior); Cu
(cubitus); CuA (cubitus anterior); CuP (cubitus posterior); Cu+1AA (cubitus and rst anterior anal); 1A
(rst anal); 1AA–7AA (rst–seventh anterior anal); 1PA–7PA (rst–seventh posterior anal).
veins lling the remainder of the alae as a fused cubitus and rst anterior anal which
splits before the wing margin, seven total anterior anals and typically ve to seven
posterior anals (Fig. 7B). Abdominal shape is variable in both sexes and can be boxy,
spade-shaped, ovoid, narrow and long, or strongly tapered. Female subgenital plates
can be short and stout or very long and thin with the apex reaching the tip of the
abdomen. Males have a unique vomer which is two-pronged, with the hooks either
the same size and shape or with one more prominent hook and a second smaller hook
adjacent to the rst (Fig. 5).
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 21
Egg morphology is variable in regard to color and pitting size/arrangement, but
the general features are relatively uniform. All known eggs are relatively small with-
out strongly formed lateral ns. Instead, the eggs are rather boxy in appearance with
relatively straight margins and short moss-like pinnae on most surfaces. e egg oper-
culum is conically raised and shares a similar surface texture to the overall egg with
short moss-like pinnae throughout, but typically with slightly longer pinnae along the
margins of the operculum (Fig. 8).
Freshly hatched nymphs have a base coloration of brown to red, and many of the
known species have abdominal segments II and III with distinct green patches which
stand out from the rest of the general nymph coloration. All freshly hatched nymphs
have partial white bands on their meso- and metafemoral lobes, and some have these
white bands weakly to fully formed on their profemoral lobes as well (Fig. 9). e
nymphs with bright red coloration such as Cryptophyllium icarus sp. nov. (Fig. 9I)
bear a striking resemblance to nymphs of Hymenopus coronatus Olivier, 1792 (Man-
todea: Hymenopodidae) and Paradasynus spp. China, 1934 (Hemiptera: Coreidae)
Figure 8. Known eggs for the Cryptophyllium gen. nov. set in couplets of lateral and then dorsal views. All
images except for K and L were taken by Bruno Kneubühler (Switzerland), K and L images taken by the
rst author, Q and R images taken by René Limoges (IMQC) A, B Cryptophyllium westwoodii comb. nov.
C , D Cryptophyllium tibetense comb. nov. E, F Cryptophyllium bollensi gen. et sp. nov. G, HCryptophyl-
lium oyae comb. nov. I , J Cryptophyllium icarus gen. et sp. nov. K , L Cryptophyllium liyananae gen. et sp.
nov. M , N Cryptophyllium chrisangi comb. nov. O, P Cryptophyllium celebicum comb. nov. Q, R Crypto-
phyllium limogesi gen. et sp. nov. Scale bars 2.0 mm long.
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
22
with their two-toned aposematic coloration (Hawkeswood and Sommung 2019). It
appears as though Hymenopus coronatus and Cryptophyllium icarus sp. nov. freshly
emerged nymphs are benetting in this aposematic mimicry which is common in
the Pentatomorpha true bugs warning of their distastefulness (Burdeld-Steel and
Shuker 2014).
Distribution. Sri Lanka to the west, Tibet to the north, Micronesia to the east, and
Wangi-wangi Island, Indonesia to the south (Fig. 2). To date we have recorded Crypto-
phyllium gen. nov. from 14 countries with China and Vietnam the current hotspots of
diversity with nine and eleven species presently known from each, respectively (Fig. 2).
Etymology. Adjectival, meaning the hidden leaves. From Greek kryptós = hidden +
phyllium = leaf. is genus is neuter in gender like Phyllium. We chose this name with
a double meaning in mind. First, this clade of leaf insects has been “hidden” within the
true Phyllium, and only recently have they been noted as unique and placed within the
celebicum species group by Hennemann et al. (2009) of Phyllium, but still not complete-
ly recognized as separate. Secondly, these insects are masters of disguise and well-hidden
within their canopy habitat (Fig. 1) which means only rarely are individuals found.
Species included within Cryptophyllium gen. nov. (listed and treated below
alphabetically).
Cryptophyllium animatum gen. et sp. nov.
Cryptophyllium athanysus (Westwood, 1859), comb. nov.
Cryptophyllium bankoi gen. et sp. nov.
Cryptophyllium bollensi 1gen. et sp. nov.
Cryptophyllium celebicum (de Haan, 1842), comb. nov.
Cryptophyllium chrisangi (Seow-Choen, 2017), comb. nov.
Cryptophyllium daparo gen. et sp. nov.
Cryptophyllium drunganum (Yang, 1995), comb. nov.
Cryptophyllium echidna gen. et sp. nov.
Cryptophyllium faulkneri gen. et sp. nov.
Cryptophyllium icarus gen. et sp. nov.
Cryptophyllium khmer gen. et sp. nov.
Cryptophyllium limogesi gen. et sp. nov.
Cryptophyllium liyananae gen. et sp. nov.
Cryptophyllium nuichuaense gen. et sp. nov.
Cryptophyllium oyae (Cumming & Le Tirant, 2020), comb. nov.
Cryptophyllium parum (Liu, 1993), comb. nov.
Cryptophyllium phami gen. et sp. nov.
Cryptophyllium rarum (Liu, 1993), comb. nov.
Cryptophyllium tibetense (Liu, 1993), comb. nov.
Cryptophyllium wennae gen. et sp. nov.
Cryptophyllium westwoodii (Wood-Mason, 1875), comb. nov.
Cryptophyllium yapicum (Cumming & Teemsma, 2018), comb. nov.
Cryptophyllium yunnanense (Liu, 1993), comb. nov.
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 23
Figure 9. Freshly hatched nymphs known for the Cryptophyllium gen. nov., all except for C bred and
photographed by Bruno Kneubühler (Switzerland) A Cryptophyllium celebicum comb. nov. B Cryptophyl-
lium chrisangi comb. nov. C Cryptophyllium khmer gen. et sp. nov. bred and photographed by Tim Bollens
(Belgium) D Cryptophyllium tibetense comb. nov. E Cryptophyllium oyae comb. nov. F Cryptophyllium
westwoodii comb. nov. G Cryptophyllium phami gen. et sp. nov. H Cryptophyllium bollensi gen. et sp. nov.
I Cryptophyllium icarus gen. et sp. nov.
Species recorded per country
Nepal
Cryptophyllium tibetense: Gandaki Pradesh (Tanahun District); Province No. 1 (Ilam)
India
Cryptophyllium tibetense: West Bengal (Kalimpong); Sikkim; Assam (Digboi)
Cryptophyllium sp.: Andaman Islands [Wood-Mason 1875]
Bhutan
None yet known, but biogeographically assumed Cryptophyllium gen. nov. should
be present.
Bangladesh
None yet known, at this time we are only aware of Phyllium (Pulchriphyllium) bio-
culatum scythe Gray, 1843 from Bangladesh, but biogeographically assumed
Cryptophyllium gen. nov. should be present as well.
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
24
China
Cryptophyllium tibetense: Tibet (Xizang Autonomous Region), Mêdog County
(Motuo); Southeast Tibet (Tenga Valley, Anjaw District, Lohit District, East
Siang district);
Cryptophyllium drunganum: Yunnan Province (Nùjiāng Lisu Autonomous Prefecture)
Cryptophyllium yunnanense: Yunnan Province (Xishuangbanna Dai Autonomous
Prefecture, Pu’er prefecture-level city, Honghe Hani and Yi Autonomous Pre-
fecture, Yuxi prefecture-level city)
Cryptophyllium wennae: Yunnan Province (Xishuangbanna Dai Autonomous Pre-
fecture, Pu’er prefecture-level city)
Cryptophyllium daparo: Yunnan Province (Xishuangbanna Dai Autonomous
Prefecture)
Cryptophyllium rarum: Guangxi Zhuang Autonomous Region (Wuzhou prefec-
ture-level city, Baise prefecture-level city, Nanning City)
Cryptophyllium parum: Hainan Island (Baisha Li Autonomous County, Ledong Li
Autonomous County)
Cryptophyllium liyananae: Guangxi Zhuang Autonomous Region (Wuzhou pre-
fecture-level city)
Cryptophyllium oyae: Yunnan Province (Maguan County)
Myanmar
Cryptophyllium westwoodii: Kayin Province, Tanintharyi Province (Karathuri)
Cryptophyllium sp.: Mandalay Region (Myingyan District)
Vietnam
Cryptophyllium oyae: Ha Giang Province (Tung Ba Commune)
Cryptophyllium yunnanense: Lai Chau Province (Mount Fansipan), Yen Bai Prov-
ince (Nghia Lo)
Cryptophyllium rarum: Vinh Phuc Province (Tam Dao), Quang-Ninh Province
(Tay Yen Tu), Ninh Binh Province (Cuc Phuong), Da Nang Province (Ba Na),
Kon Tum Province (Ngoc Linh), Hoa Binh Province
Cryptophyllium animatum: Quang Nam Province (Tay Giang)
Cryptophyllium bankoi: ua ien Hue Province (Bach Ma), Da Nang Province
(Ba Na Nui Chua), Quang Nam Province (Axan)
Cryptophyllium bollensi: Ninh uan Province (Phuoc Binh N.P.)
Cryptophyllium phami: Dong Nai Province (Cat Tien N.P.), Binh uan Province
(Dong Tien)
Cryptophyllium nuichuaense: Ninh uan Province (Nui Chua N.P.)
Cryptophyllium icarus: Lâm Dông Province (Bidoup Nui Ba N.P., Bao Loc), Dak
Lak Province
Cryptophyllium limogesi: Dak Lak Province (Chu Yang Sin), Dak Nong Province
(Ta Dung), Lam Dong Province (Dambri)
Cryptophyllium faulkneri: Quang Ngai Province (Bato), Lam Dong Province (Bao Lam)
Laos
Cryptophyllium oyae: Houaphan Province (Xamnuen District)
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 25
Cryptophyllium westwoodii: Bokeo Province (Ban Muang Kan), Louang Prabang
Province, Vientiane Province (Muang Feuang District)
ailand
Cryptophyllium westwoodii: Chiang Mai Province (Fand District), Lamphun Prov-
ince (Mae a District), Loei Province (Na Haeo)
Cryptophyllium chrisangi: Nakhon Si ammarat Province (ung Song District),
Satun Province (Tarutao Island)
Cryptophyllium oyae: Nan Province (Bo Kluea Tai District), Phetchabun Province,
Chiang Mai Province
Cambodia
Cryptophyllium khmer: Koh Kong Province (Tatai), Siem Reap Province (Kbal Spean)
Sri Lanka
Cryptophyllium athanysus: Southern Province (Sinharaja Forest Reserve)
Philippines
Cryptophyllium sp. “Cebu”: Cebu Island
Cryptophyllium sp. “Zamboanga”: Mindanao (Zamboanga Peninsula)
Malaysia
Cryptophyllium chrisangi: Perak (Tapah), Sarawak State (Kuching)
Singapore
Cryptophyllium chrisangi: mainland Singapore and St. John’s Island
Indonesia
Cryptophyllium chrisangi: Pulau Weh Island (north coast of Sumatra)
Cryptophyllium sp. “Bangka”: Bangka Island (o the southeastern Sumatra coast)
Cryptophyllium celebicum: Sulawesi; Peleng Island; and Buton Island
Cryptophyllium echidna: Wangi-wangi Island
Micronesia
Cryptophyllium yapicum: Yap Island
Cryptophyllium animatum gen. et sp. nov.
http://zoobank.org/0EA5C7BF-DD05-4889-AEBA-8203220B57CE
Figure 10
Material examined. Holotype ♂: “VIETNAM: Quang Nam, Tay Giang, Axan Mt,
1,300 meters: July 2017 (Coll RC 17-261)”. Deposited in the Montreal Insec-
tarium (IMQC).
Remarks. is large species is presently only known from the singular holotype
male (89.4 mm in length) from Central Vietnam. With such a large male we look
forward to when the female is located as the female must also be signicantly large
and possibly larger than any of the presently known Cryptophyllium gen. nov. females.
Dierentiation. Females unknown. Males can be dierentiated by the profemoral
interior lobe which is marked by teeth which are uniform in size and spacing, a feature which
no other known congenerics possess (Fig. 10B). Molecularly Cryptophylliumanimatum
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
26
sp. nov. is most closely related to Cryptophyllium rarum comb. nov. and shares several
morphological similarities such as the large size, serrated profemoral exterior lobe, similar
tegmina lengths, and a similar shape of the mesothorax and similar arrangement/sizing of
the teeth (Fig. 10C). ese species can be dierentiated, as Cryptophyllium rarum comb.
nov. have profemoral interior lobe teeth, which are unevenly sized and spaced (Fig. 61E),
and a broad rounded abdomen and not a thin spade-like abdomen as in Cryptophyllium
animatum sp. nov.
An additional species morphologically similar to Cryptophyllium animatum sp.
nov. is Cryptophyllium faulkneri sp. nov., which can also be found in Central Vietnam.
Morphological similarities between these species are the thin spade-shaped abdomens,
large sizes, and similar tegmina lengths. Although supercially they look similar with
their large thin appearance, it is easy to dierentiate the species when the details of
the thorax are observed, as Cryptophyllium animatum sp. nov. has small numerous
teeth throughout the mesopleura length (Fig. 10C) vs. Cryptophyllium faulkneri sp.
nov., which has four or ve large tubercle-like teeth and smaller interspersed teeth
between (Fig. 31A). Additionally, the exterior profemoral lobes dier in their serration
as Cryptophyllium animatum sp. nov. has eight or nine sharply serrate teeth (Fig. 10B)
vs. Cryptophyllium faulkneri sp. nov. which at most has four rounded, less prominent
teeth (Fig. 31B).
Distribution. Currently only known from the type locality in Central Vietnam:
Quang Nam Province, Axan Mountain.
Male. Coloration. Coloration description based on the single dried holotype
specimen, not on living individuals which are likely a more vibrant green. Overall col-
oration is pale green with variable patches of tan throughout due to the drying of the
specimen (Fig. 10A). Compound eyes are cherry red (Fig. 10C). e antennae, head,
and thorax are tan (Fig. 10C). Abdominal segment V has a set of eye spots which are
clearer and situated closer to the midline than the margin.
Morphology. Head. Head capsule slightly longer than wide, with a vertex that is
weakly granular in a slightly longitudinal arrangement; posteromedial tubercle small
but notable and slightly raised above the head capsule (Fig. 10C). Frontal convexity
stout and nely pointed with two or three short setae near the apex. Compound eyes
large but not overly bulbous, occupying > ⅓ of the head capsule lateral margins
(Fig.10C). ree ocelli are well-developed and located between and slightly posterior
to the compound eyes. Antennal elds are slightly wider than, and approximately as
long as the scapus. Antennae. Antennae (including the scapus and pedicellus) consists
of 27 segments. e scapus and pedicellus are bare, all other segments are covered in
dense, thin, pale setae that are as long as or longer than the antennae segment is wide,
and the terminal three segments have shorter setae which are denser than the other
segments. orax. Pronotum with anterior margin gently concave and lateral margins
that are nearly straight and converge to a straight posterior margin that is approxi-
mately half the width of the anterior rim (Fig. 10C). Anterior and lateral margins of
the pronotum with distinct rims and the posterior margin lacks a rim (Fig. 10C). Face
of the pronotum is marked by a smooth surface, distinct sagittal furrow, pit just pos-
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 27
Figure 10. Holotype Cryptophyllium animatum gen. et sp. nov., photographs by RTC A habitus, dorsal
B details of the protibial and profemoral lobes C details of the head and thorax, dorsal D genitalia, ventral
E head–thorax, lateral.
terior to the center, a moderate perpendicular furrow just anterior to the central pit,
and two distinct pits along the anterior margin (Fig. 10C). Prosternum is nearly
smooth, with only a slightly wrinkled surface. Mesosternum surface marked densely
with prominent nodes, with the largest along the sagittal plane and more strongly on
the anterior margin, posterior margin with less prominent and slightly smaller nodes.
Prescutum longer than wide, with lateral margins slightly converging to the posterior
which is slightly narrower than the anterior rim width (Fig. 10C). Lateral rims with
ten or eleven small tubercles of varying sizes, none large or prominent, most approxi-
mately even in size and one or two not much more prominent than nodes (Fig. 10C).
e surface of the prescutum is heavily granulose and rises slightly up to meet the
prescutum crest (Fig. 10E). Prescutum crest along the sagittal plane with seven or
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
28
eight nodes which are denser on the anterior and slightly wider spaced on the poste-
rior (Fig. 10E). In addition to the granular surface there is shallow pitting along the
lateral margins, but none very prominent (Fig. 10C). Prescutum anterior rim with a
heavily granulose surface and no distinct central tubercle (Fig. 10E). Mesopleura nar-
row, gradually diverging from the anterior to the posterior (Fig. 10C). Lateral margin
with four or ve moderately sized tubercles throughout the length, and between six or
seven smaller minor tubercles interspersed throughout (Fig. 10C). Face of the meso-
pleura slightly wrinkled and with two distinct pits, one on the anterior ⅓ and one on
the posterior ⅓. Wings. Tegmina moderate length, extending slightly > ½ through
abdominal segment III. Tegmina wing venation (see Fig. 7B for general venation
found in the Cryptophyllium gen. nov.): the subcosta (Sc) is the rst vein and termi-
nates the earliest, slightly > ca. ⅓ of the way through the overall tegmina length. e
radius (R) spans nearly the entire length of the tegmina with the rst radius (R1)
branching just anterior to the middle and terminating just posterior to the middle of
the wing, the second radius (R2) branches near the distal ⅓ of the wing, and then the
radial sector (Rs) runs parallel and side by side with the media anterior until the pos-
terior ⅕ of the tegmina, at which point the radial sector bends away from the media
anterior and then terminates just shy of the tegmina apex. e media (M) spans the
entire length of the tegmina, terminating at the wing apex as the media anterior (MA).
e rst media posterior (MP1) begins and terminates near the tegmina mid length
followed by the second media posterior (MP2) which begins ca. ⅔ of the way through
the tegmina length and terminates near the posterior quarter of the wing. e cubitus
(Cu) runs through the wing surface angled towards the margin which it meets ca. ⅓
of the way through the tegmina length and then runs along the margin as the two
media posterior veins then meet it and the cubitus eventually dissipates ca. ⅔ of the
way through the tegmina length. e rst anal (1A) vein runs subparallel to the cubi-
tus until it meets it slightly > ⅓ of the way through the tegmina length. Alae well
developed in an oval fan conguration, reaching to the anterior margin of abdominal
segment IX. Alae wing venation (see Fig. 7B for general venation found in the species
of this genus): the costa (C) is present along the entire foremargin giving stability to
the wing. e subcosta (Sc) spans ca. ⅔ of the wing length and is mostly fused with
the radius in the beginning but terminates when it meets the costa. e radius (R)
spans the entire wing and branches slightly > ⅖ of the way through into the radius 1
(R1) and radial sector (Rs) which run nearly parallel through most of their length
until they terminate at the wing apex near each other but not touching. e media
(M) branches early, ca. ⅐ of the way through the wing into the media anterior (MA)
and the media posterior (MP) which run parallel with each other until the distal ⅐ of
the wing where the media posterior fuses with the media anterior which then run
fused together to the wing margin. e cubitus (Cu) runs unbranched and terminates
at the wing apex. Of the anterior anal veins, the rst anterior anal (1AA) fuses with
the cubitus near the point where the media branches into the media anterior and
media posterior and then the rst anterior anal branches from the cubitus ⅔ of the
way through the wing length where it uniformly diverges from the cubitus until it
terminates at the wing margin. e anterior anal veins two–seven (2AA–7AA) have a
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 29
common origin and run unbranched in a folding fan pattern of relatively uniform
spacing to the wing margin. e posterior anal veins (1PA–7PA) share a common
origin separate from the anterior anal veins and run unbranched to the wing margin
with slightly thinner spacing than the anterior anal veins. Abdomen. Abdomen gen-
eral shape is a narrow spade. Abdominal segments II through the anterior half of seg-
ment IV slightly diverging to the widest point with the remainder of segment IV
parallel sided. Segments V–IX are uniformly converging slowly with straight margins
to the terminal abdominal segment X which at rst also converges slowly but then
quickly tapers o to a blunt rounded apex. Genitalia. Poculum broad and rounded,
ending in a straight margined apex that passes beyond the anterior margin of segment
X (Fig. 10D). Cerci long and slender, with ca. ½ of their length extending out from
under the anal abdominal segment. e cerci are slightly cupped, covered in a granu-
lose surface and numerous short setae throughout (Fig. 10D). Vomer broad and stout
with straight sides evenly converging to the apex, which is armed with two upwards
turning hooks, one ca. 2× as wide and ⅓ longer than the other smaller hook adjacent
to it. Legs. Profemoral exterior lobe a rounded arc, only slightly wider than the inte-
rior lobe (ca. 3× the width of the profemoral shaft at its widest), and with the anterior
half marked by eight or nine small but sharp anteriorly pointing teeth (Fig. 10B).
Profemoral interior lobe rounded near the midline and then straight to the anterior
margin, ca. 2½× as wide as the profemoral shaft at its widest. e profemoral interior
lobe is marked with eight or nine small, serrate, anteriorly pointing teeth which are
nearly uniform in size and spacing (Fig. 10B). Mesofemoral exterior lobe arcs end to
end but with the widest portion distal to the midline, and the widest point about as
wide as the interior lobe or the mesofemoral shaft at its widest. e mesofemoral ex-
terior lobe is marked with three or four small serrate teeth distal to the widest point,
with the proximal portion of the lobe smooth. e mesofemoral interior lobe is ap-
proximately the same width as the exterior lobe, but more evenly weighted from end
to end with the widest portion near the midline. e distal half is marked with nine
small serrate teeth with those most distal slightly closer together than those near the
middle. Metafemoral exterior lobe lacks dentation, and has a straight margin along
the metafemoral shaft. Metafemoral interior lobe smoothly arcs end to end with ten
or eleven small serrate teeth on the distal half only. Protibiae lacking exterior lobe,
interior lobe reaching end to end in a smooth triangle slightly > 2× as wide as the
protibial shaft, with the widest point just distal to the midline (Fig. 10B). Mesotibiae
and metatibiae simple, lacking lobes completely.
Measurements of holotype male [mm]. Length of body (including cerci and head,
excluding antennae) 89.4, length/width of head 5.5/4.5, antennae 55.8, pronotum
4.5, mesonotum 5.8, length of tegmina 27.1, length of alae 63.7, greatest width of
abdomen 21.6, profemora 19.6, mesofemora 15.6, metafemora 17.6, protibiae 12.4,
mesotibiae 9.7, metatibiae 13.4.
Etymology. Noun, Latin for “endowed with spirit”. We felt that these incredible
insects are so leaf-like in texture, color, and movement, they simply appear to be a leaf
on a tree come to life and walking right out of a fantasy novel and therefore the full
binomial translating to “hidden leaf endowed with spirit” was tting.
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
30
Cryptophyllium athanysus (Westwood, 1859), comb. nov.
Figures 11, 12
Material examined. Very few specimens are known. We examined the holotype female
in the NHMUK “Holo-type. Ceylon. Athanysus, Westw. Phyllium Athanysus Westw,
Ceylon. BMNH(E) #845230” (Fig. 11A) and a male in the OUMNH collection “P.
athanysus, Westw. Ceylon. E Coll. (1830–73), W. W. Saunders., Purchased and, pres.
‘73 by Mrs. F. W. Hope.” Greene (1906: g. 1b, plate between page 220 and 221)
illustrated a single female specimen as well as an egg (g. 2b, page 221). We were un-
able to trace these specimens for proper examination. Besides these few specimens, we
reviewed images of one live female (Fig. 12A, B) and one live male (Fig. 12C) from the
Sinharaja Forest Reserve, but no other records are known to us.
Remarks. is rare species has only had a number of instances over the years where
preserved specimens were illustrated. Here we present the rst known photographs of
live male and female individuals observed on two dierent occasions, both within the
Sinharaja Forest Reserve, Sri Lanka (Fig. 12).
Dierentiation. Females can be dierentiated by the smaller size (the smallest
known species of Cryptophyllium gen. nov., ca. 77.0 mm) and the fully developed exte-
rior metatibial lobes, which span the entire metatibial shaft, and the weak but notable
full mesotibial exterior lobes, which also span the entire length of the shaft (when there
are exterior lobes present in the other Cryptophyllium gen. nov. species, they are small
and restricted to the distal end of the shaft only). Cryptophyllium athanysus comb. nov.
females are most morphologically similar to Cryptophyllium bollensi sp. nov. and to
Cryptophyllium chrisangi comb. nov. due to the anteriorly narrow mesopleura and the
broad rounded profemoral exterior lobes. Additionally, Cryptophyllium chrisangi comb.
nov. also has a tapered abdomen like Cryptophyllium athanysus comb. nov. without dis-
tinct abdominal lobes on segment VII, which many Cryptophyllium species have. Both
of these morphologically similar species can immediately be dierentiated however by
the lack of metatibial exterior lobes.
Males can also be dierentiated from all other species by the presence of fully
developed exterior metatibial lobes which span the entire metatibial shaft, as
no other Cryptophyllium gen. nov. species are known to have such a feature. Male
Cryptophylliumathanysus comb. nov. are morphologically very similar to Cryptophyllium
westwoodii comb. nov. as they both have similar narrow abdominal shapes, similar
profemoral morphology, and similar thorax shape and serration. Cryptophyllium
athanysus comb. nov. can be dierentiated easily however by the presence of metatibial
exterior lobes, which Cryptophyllium westwoodii comb. nov. lacks.
Distribution. Sri Lanka. All old records simply state “Ceylon’’ as the location, but
the two modern records we have come across were both from within the Sinharaja For-
est Reserve on the southern end of the island (Fig. 12). Even Greene (1906) regarded
this species as rarely encountered with most records belonging to the other phylliid
species present on the island.
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 31
Figure 11. Cryptophyllium athanysus comb. nov. holotype female, photographs by Paul Brock (NHMUK)
A habitus, dorsal view B details of the thorax, dorsal C genitalia details, ventral D illustration of a Cryp-
tophyllium athanysus comb. nov. egg from Greene (1906).
Cryptophyllium bankoi gen. et sp. nov.
http://zoobank.org/A99E5AA5-D9A3-405B-92F7-DE8B83A57384
Figures 5G, 13, 14, 15
Material examined. Holotype ♂: “VIETNAM: Quang Nam, Tay Giang, Axan
Mt, 1,300 meters: June 2017, Coll RC 17-228”. Deposited in the Montreal Insec-
tarium (IMQC).
Paratypes: (4 ♂♂, 1 ♂ nymph, 1 ♀ nymph) • 1 ♂; “Vietnam; Daknong, June,
2017, Coll RC 17-338” (Coll RC) • 1 ♂; “Vietnam; Quang Ngai Province, Bato Mt.
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
32
Figure 12. Live Cryptophyllium athanysus comb. nov. observed in the Sinharaja Forest Reserve, Sri Lanka
A, B female photographed by Alberto Carrera (A photograph purchased from Alamy photo stock website
B photograph purchased from Adobe Stock) C rst record of a live male observed and photographed by
Angela Christine Chua (Singapore) in January 2020.
900 m. elv: May 2015, Coll RC 16-141” (Coll RC) • 1 ♂: “Coll. I.R.Sc.N.B., Da Nang
prov., Ba Na-Nui Chua Nat. Res., 18°09’N 105°55’E, 16-19.vii.2017, GTI Project,
Leg. J. Constant and J. Bresseel, I.G.: 33.498” [vomer dissected] (RBINS) • 1 ♂: “Ngoc
Linh, Kon Tum prov., Vietnam, 1700 m, VI.2016, leg. Luong coll. TB-05-134” (Coll
TB) • 1 subadult ♀: “C. Vietnam, Bach Ma N.P., 16°12’N 107°52’E, 12-17.vii.2011,
Leg J. Constant and J. Bresseel, I.G.: 31.933” [RBINS- Phyllium-DNA sample 0006]
(RBINS) • 1 subadult ♂: “Vietnam, Gia Lai prov. Kon Chu Rang N.R., 600-1200 m,
13-20.vii.2018, GTI project, 14°28’28”N 108°32’27”E, Leg. J. Constant, J. Bresseel
and X. Vermeersch, I.G.:33.769” [RBINS- Phyllium-DNA sample 0008] (RBINS).
Remarks. is species, in true “cryptic leaf” fashion, was only recognized as unique
when the molecular results were reviewed as morphologically it was hidden within the
Cryptophyllium rarum comb. nov. males. eir morphological resemblance is uncanny
and even with a series of males it is dicult to adequately dierentiate these two species
based on male morphology. Only the male Cryptophyllium bankoi sp. nov. is known
at present and therefore it is unknown if the female also morphologically resembles
Cryptophyllium rarum comb. nov. or is morphologically more unique. Despite several
expeditions to southern Vietnam by the RBINS expedition members, only one nymph
female Cryptophyllium bankoi sp. nov. has been collected to date, unfortunately she did
not survive to adulthood (Fig. 13). With this species recorded from three Vietnamese
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 33
provinces, we hope that as a widely ranging species the female can one day be identied
and morphologically described.
Dierentiation. Females unknown. Males are morphologically very similar to
Cryptophyllium rarum comb. nov. in their antennae length, profemoral interior lobe
serration, protibial interior lobe shape, tegmina length, and their thorax shape and
spination. e molecular results revealed however that this species was in fact not iden-
tical to Cryptophyllium rarum comb. nov. with which it shares a sympatric geographic
range in central Vietnam and a striking morphological resemblance. ankfully with
the molecular analysis we were able to separate out these two species to allow us to
see the subtle dierences between them, which we originally thought to be simple
morphological variation. e only consistent morphological dierences we were able
to locate between these species is that Cryptophyllium rarum comb. nov. tends to have
an abdomen which is slightly more rounded (Fig. 59A), vs. Cryptophyllium bankoi sp.
nov. which has the abdomen ever so slightly spade-shaped (Fig. 14). Additionally, it
appears as though Cryptophyllium rarum comb. nov. tend to be larger (79.8–89.0 mm)
vs. Cryptophyllium bankoi sp. nov. (61.9–69.5 mm), and the profemoral exterior lobes
have slightly diering serration, with Cryptophyllium bankoi sp. nov. having no teeth or
up to ve small teeth, vs. Cryptophyllium rarum comb. nov., which tend to have seven
or eight teeth.
Distribution. Central and Southern Vietnam. Known from Quang Ngai, ua
ien Hue, Da Nang, Gia Lai, Quang Nam, and Dak Nong Provinces.
Male. Coloration. Living individuals are a more vibrant green, and our descrip-
tions are only based on preserved specimens. Overall coloration pale green throughout
Figure 13. Cryptophyllium bankoi gen. et sp. nov. paratype female nymph collected in Bach Ma N.P. on
Rubus sp. (Rosaceae), Vietnam, photographed by Jérôme Constant (RBINS) on 11th July 2011. Molecular
sample RBINS06 within this study A dorsal view with abdomen curled B dorsal view with abdomen
held at.
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
34
Figure 14. Cryptophyllium bankoi gen. et sp. nov. holotype male, photographs by RTC A front leg de-
tails, dorsal B details of the base of antennae, head, thorax C habitus, dorsal D details of genitalia, ventral.
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 35
with variable patches of yellow and tan coloration (Figs 14, 15). Compound eyes are
slightly reddish to pink (Figs 14B, 15D). e antennae are yellow to orange in color.
In all specimens examined there were no variable brown patches on the lobes of the legs
as is sometimes common in male phylliids.
Morphology. Head. Head capsule approximately as long as wide, with a vertex that
is irregularly lumpy and with small granulation throughout, and a posteromedial tu-
bercle which is larger than any of the nodes on the head (Fig. 14B). Frontal convexity
stout with sparse thin setae throughout the surface. Compound eyes large and bulbous,
occupying ca. ⅖ of the head capsule lateral margins (Fig. 14B). Between the com-
pound eyes are three well-developed ocelli (Fig. 14B). Antennal elds approximately as
wide and as long as the scapus. Antennae. Antennae (including the scapus and pedicel-
lus) consists of 25 or 26 segments, all segments except the scapus and pedicellus and
terminal four segments are covered in dense setae that are as long as or longer than the
antennae segment is wide. e terminal four segments are covered in dense short setae
and the scapus and pedicellus are nearly bare, lacking setae. orax. Pronotum with
anterior margin slightly concave and lateral margins that are straight and converging to
a straight posterior margin that is ca. ½ as wide as the anterior margin (Fig. 14B). An-
terior and lateral margins of the pronotum have distinct rims, but the posterior margin
has a weakly formed rim (Fig. 14B). Face of the pronotum is marked by a distinct pit
in the center with prominent anterior and lateral furrows from this central pit, and the
pronotum surface is marked with wrinkles throughout, but not prominent nodes (Fig.
14B). Prosternum surface is moderately granulose throughout, the mesosternum sur-
face is mostly smooth with only small granulation along the sagittal plane which then
continues onto the metasternum surface which has moderate granulation throughout
(Fig. 15B). Prescutum ca. ⅓ longer than the anterior margin width, with lateral mar-
gins slightly converging to the posterior (Fig. 14B). Lateral margins lacking distinct
tubercles, instead with granulation throughout giving them a rough appearance (Fig.
14B). Prescutum surface throughout with granulation, with smaller and tighter packed
granulation along the lateral margins, and slightly larger nodes along the sagittal plane
(Fig. 14B). Prescutum anterior margin distinct but not signicantly raised, with a sur-
face that is marked with minimal granulation. Mesopleura not notably wide, narrow
near the anterior margin and then only gradually diverge for the remainder of their
length (Fig. 14B). Lateral margin with ve or six small tubercles throughout the length
and several smaller minor tubercles interspersed throughout. Face of the mesopleura
slightly wrinkled and with two faint divots, one near the anterior margin and one
on the posterior ⅓ (Fig. 15E). Wings. Tegmina moderate length, extending ⅓ to ½
through abdominal segment III. Tegmina wing venation: the subcosta (Sc) is the rst
vein, is simple, and terminates the earliest ca. ⅓ of the way through the overall tegmina
length. e radius (R) spans the entire length of the tegmina with the rst radius (R1)
branching ca. ⅖ of the way through the wing length and terminating near the midline,
followed by the branching and termination of the second radius (R2) near the distal
⅓ of the wing, and the radial sector runs to the wing apex. e media (M) also spans
the entire length of the tegmina with the rst media posterior (MP1) branching o ca.
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
36
Figure 15. Cryptophyllium bankoi gen. et sp. nov. paratype male from the Ba Na-Nui Chua Nature
Reserve (RBINS), photographs by Jérôme Constant (RBINS) A habitus, dorsal B habitus, ventral C pro-
femoral and tibial, dorsal D details of the base of antennae, head, and thorax, dorsal E details of the base
of antennae, head, and thorax, lateral F terminalia, dorsal G genitalia, ventral.
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 37
⅓ of the way through the wing length, the second media posterior (MP2) branching
near the midline, and the media anterior (MA) running to the wing apex. e cubitus
(Cu) runs along the edge of the wing as the two media posterior veins fuse with it and
as the cubitus reaches the apex it fades. e rst anal (1A) vein terminates upon reach-
ing the cubitus ca. ⅓ of the way through the wing length. Alae well developed in an
oval fan conguration, long, reaching to the middle of abdominal segments VIII. Alae
wing venation: the costa (C) is present along the entire foremargin giving stability to
the wing. e subcosta (Sc) is long, spanning ca. ⅔ of the wing length and is mostly
fused with the radius in the beginning but terminates when it meets the costa. e
radius (R) spans the entire wing and branches ca. ⅓ of the way through into the rst
radius (R1) and radial sector (Rs) which run gently diverging for most of their length
and then converge at the apex of the wing where they terminate near each other but
not touching. e media (M) branches early, ca. ⅙ of the way through the wing into
the media anterior (MA) and the media posterior (MP) which run parallel with each
other throughout the wing until the distal ⅕ of the wing when the media posterior
fuses with the media anterior which then run fused together to the wing apex where
they terminate near the radial sector. e cubitus (Cu) runs unbranched and termi-
nates at the wing apex. Of the anterior anal veins, the rst anterior anal (1AA) fuses
with the cubitus near the point where the media branches into the media anterior
and media posterior and then the rst anterior anal branches from the cubitus ⅔ of
the way through the wing length where it uniformly diverges from the cubitus until
it terminates at the wing margin. e anterior anal veins two–seven (2AA–7AA) have
a common origin and run unbranched in a folding fan pattern of relatively uniform
spacing to the wing margin. e posterior anal veins (1PA–6PA) share a common ori-
gin separate from the anterior anal veins and run unbranched to the wing margin with
slightly thinner spacing than the anterior anal veins. Abdomen. Abdominal segment II
with parallel margins, III through the anterior ⅓ of segment IV gradually diverging
to the widest portion. e posterior of IV–V are parallel or slightly subparallel. Seg-
ments VI–X uniformly converging, giving the abdomen a spade-shaped appearance
(Fig. 14C). Abdominal segment X distinctly longer than wide (Fig. 15F). Genitalia.
Poculum broad, and ending in a rounded apex that slightly passes the anterior mar-
gin of segment X (Fig. 14D). Cerci long and slender, extending from under the anal
abdominal segment by slightly > half of their length, surface slightly cupped and with
a granulose surface with numerous short setae throughout (Fig. 14D). Vomer broad
and stout with straight sides evenly converging to the apex, which is marked by two
apical hooks, one slightly larger than the other (Fig. 5G). Legs. Profemoral exterior
lobe slightly thinner than the interior lobe, arcing end to end in a smoothly bending
lobe which has a granular margin throughout and can range from lacking dentation
to having four or ve small serrate teeth on the distal half of the lobe (Figs 14A, 15C).
Profemoral interior lobe roundly triangular and marked with ve teeth arranged in a
two-one-two pattern with large looping gaps between these groups (Fig. 15A). Mes-
ofemoral exterior lobe arcs end to end, but is more heavily weighted toward the distal
half which is slightly broader than the mesofemoral shaft width, and which is marked
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
38
with 1–3 serrate teeth on the distal half only, and the proximal half is rather thin and
lacks dentation. Mesofemoral interior lobe is about the same width as the mesofemoral
shaft and arcs end to end, is slightly broader on the distal end and is marked with four
or ve small serrate teeth. Metafemoral exterior lobe lacks dentation, and has a straight
margin hugging the metafemoral shaft. Metafemoral interior lobe smoothly arcs end to
end with seven or eight serrate teeth on the distal half only. Protibiae lacking exterior
lobe, interior lobe reaching end to end in a smoothly rounded triangle which is slightly
weighted to the distal half and at the widest point slightly < 2× the protibial shaft width
(Fig. 15A). Meso- and metatibiae simple, lacking lobes completely.
Measurements of holotype male [mm]. Length of body (including cerci and head,
excluding antennae) 64.3, length/width of head 3.8/3.6, antennae 34.5, pronotum
2.9, mesonotum 3.7, length of tegmina 17.4, length of alae 47.0, greatest width of
abdomen 20.6, profemora 13.7, mesofemora 10.6, metafemora 12.5, protibiae 8.2,
mesotibiae 7.0, metatibiae 9.4.
Measurements of paratype males [mm]. Length of body (including cerci and
head, excluding antennae) 61.9–69.5, length/width of head 3.2–3.6/3.1–3.7, anten-
nae 32.8–41.2, pronotum 2.7–3.6, mesonotum 4.0–5.1, length of tegmina 19.7–19.9,
length of alae 48.3–50.1, greatest width of abdomen 18.4–20.6, profemora 12.5–15.7,
mesofemora 9.9–12.1, metafemora 10.8–14.2, protibiae 7.6–9.2, mesotibiae 7.8–8.2,
metatibiae 8.0–10.4.
Etymology. Patronym. is species is dedicated to our friend and colleague Alex-
andre Banko for his extensive eorts to discover new species and his long collaboration
with Team Phyllies to present us with fresh material to sequence and study.
Cryptophyllium bollensi gen. et sp. nov.
http://zoobank.org/8023142D-4BAA-4FBF-A26F-B280DC7C5E02
Figures 5E, 6B, 8E, 8F, 9H, 16, 17, 18, 19, 20, 21
Material examined. Holotype ♂: “Coll. I.R.Sc.N.B., Ex breeding Tim Bollens, 2018,
Coll. I.R.Sc.N.B., Vietnam, Ninh uan prov., Phuoc Binh N.P., 12°04’N, 108°45’E,
26.vii.2014, night coll., Leg.: J. Constant & J. Bresseel, GTI project, IG: 32.779”.
Deposited in the Royal Belgian Institute of Natural Sciences (RBINS).
Paratypes: (16 ♀♀, 25 ♂♂, 46 eggs) • 1 ♀: “Coll. I.R.Sc.N.B., Vietnam, Ninh
uan prov., Phuoc Binh N.P., 12°04’N, 108°45’E, 26.vii.2014, night coll., Leg.: J.
Constant and J. Bresseel, GTI project, IG: 32.779” (RBINS) • 6 ♂♂: same data as HT
[3 with vomer dissected] (RBINS) • 2 ♂♂, 3 ♀♀, 1 subadult ♂: “Coll. I.R.Sc.N.B.,
Ex breeding Tim Bollens, 2019, Vietnam, Ninh uan prov., Phuoc Binh N.P.,
12°04’N, 108°45’E, 26.vii.2014, night coll., Leg.: J. Constant and J. Bresseel, GTI
project, IG: 32.779” (RBINS) • 1 ♂: “Coll. I.R.Sc.N.B., Ex breeding B. Kneubuh-
ler, Vietnam, Ninh uan prov., Phuoc Binh N.P., 12°04’N, 108°45’E, 26.vii.2014,
night coll., Leg.: J. Constant and J. Bresseel, GTI project, IG: 32.779” (RBINS) • 1 ♂,
1♀: “Coll. I.R.Sc.N.B., Ex breeding Bruno Kneubühler, 2017” “Coll. I.R.Sc.N.B.,
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 39
Figure 16. Live Cryptophyllium bollensi gen. et sp. nov. paratype where she was found in Phuoc Binh
N.P., Vietnam in July 2014 by Joachim Bresseel (RBINS) and Jérôme Constant (RBINS). Photograph by
Jérôme Constant (RBINS).
Vietnam, Ninh uan prov., Phuoc Binh N.P., 12°04’N, 108°45’E, 26.vii.2014,
night coll., Leg.: J. Constant and J. Bresseel, GTI project, IG: 32.779” (VNMN)
• 1 ♀; “VIETNAM: Ninh uan prov. Phuoc Binh N.P., 12°04’N 108°45’E, Bred
from eggs supplied by Bruno Kneubühler (Switzerland), May, 2018, Coll RC 18-
414” (Coll RC) • 2 subadult ♀♀; “VIETNAM: Ninh uan prov. Phuoc Binh N.P.,
12°04’N 108°45’E, Bred from eggs supplied by Bruno Kneubühler (Switzerland),
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
40
May, 2018” Coll RC 18-412 and 18-413 (Coll RC) • 1 ♂; “VIETNAM: Ninh -
uan prov. Phuoc Binh N.P., 12°04’N 108°45’E, Bred from eggs supplied by Bruno
Kneubhler (Switzerland), April, 2018, Coll RC 18-2017” (Coll RC) • 19 eggs; “VI-
ETNAM: Ninh uan prov. Phuoc Binh N.P., 12°04’N 108°45’E, eggs supplied by
Bruno Kneubühler (Switzerland), 2018”, Coll RC 17-375, 17-376, 18-282–18-298
(Coll RC) • 1 ♀, 3 ♂♂, 27 eggs; “ex Zucht T. Bollens 2018, Herkunft: Vietnam,
Prov. Nin uan, Bác Ái Distr., Phuoc Binh N.P., leg. Bresseel and Constant 2014”
[coll. FH, No’s 1061-1 to 4, E], (Coll FH) • 1 ♀, 1 ♂ (nymph n3); “ex Zucht F.
Hennemann 2019, Herkunft: Vietnam, Prov. Nin uan, Bác Ái Distr., Phuoc Binh
NP, leg. Bresseel and Constant 2014” [coll. FH, No’s 1061-5 and 6], (Coll FH) • 2
♀♀, 3 ♂♂; “VIETNAM: Ninh uan prov. Phuoc Binh N.P., 12°04’N 108°45’E,
Bred by Maxime Ortiz, France, circa 2020” (Coll MO) • 4 ♀♀, 6 ♂♂; “VIETNAM:
Ninh uan prov. Phuoc Binh N.P., bred by Bruno Kneubühler (Switzerland), circa
2017-2018” (Coll OC).
Remarks. is species was rst collected by Jérôme Constant (RBINS) and Joachim
Bresseel (RBINS) during a GTI research expedition. Only a single adult female was
found (Fig. 16) while collecting at night in Phuoc Binh N.P.. e female was found on
a shrub in a eld that had been cleared of trees and planted with crops. e clearing
of the forest had been done fairly recently as tree stumps were still present. In this area
there was no adjacent old growth forest, only secondary forest nearby. e female was
kept alive long enough to lay a series of fertilized eggs which were shared with expert
phylliid breeder Tim Bollens (Belgium) who was able to rear a nice series of specimens
and eventually share this species with other leaf insect breeder enthusiasts (Fig. 17).
Dierentiation. Females are morphologically most similar to Cryptophyllium
phami sp. nov., Cryptophyllium chrisangi comb. nov., and Cryptophyllium nuichuaense
sp. nov. based on the general femoral lobe shapes, the broad rounded exterior pro-
femoral lobe, and the thorax shape and spination. Cryptophyllium bollensi sp. nov. have
moderately long alae reaching onto abdominal segment IV that can be dierentiated
from Cryptophyllium phami sp. nov., which have shorter alae only reaching the anterior
margin of abdominal segment III. e ventral surface of the antennae dierentiates
Cryptophyllium bollensi sp. nov. from the other similar species as Cryptophyllium bol-
lensi sp. nov. have the ventral surface of segments VI, VII, and VIII ush (Fig. 6B) vs.
Cryptophyllium chrisangi comb. nov. and Cryptophyllium nuichuaense sp. nov. which
have the ventral surface of antennal segments VI and VII projecting beyond segment
VIII, giving the antennae a slight lamellate appearance (Fig. 6E).
Males are morphologically most similar to Cryptophyllium westwoodii comb. nov.,
Cryptophyllium chrisangi comb. nov., Cryptophyllium phami sp. nov., and Cryptophyl-
lium khmer sp. nov. due to the femoral shape and spination, the length of antennae
and alae, and thorax shape and spination. Cryptophyllium westwoodii comb. nov. and
Cryptophyllium chrisangi comb. nov. can be dierentiated by their narrower abdominal
shape with a maximum width only 30–34% of the abdominal length, vs. the oth-
ers which have an abdominal shape that is broadly elliptical or broadly spade-shaped
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 41
Figure 17. Live Cryptophyllium bollensi gen. et sp. nov. bred and photographed by Bruno Kneubühler
(Switzerland) A adult female, dorsal B adult male, dorsal C genitalia, female, ventral.
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
42
with a maximum width ca. 38–45% of the abdominal length. Due to similarities in
morphology and the intraspecic variation within Cryptophyllium bollensi sp. nov.,
Cryptophyllium phami sp. nov., and Cryptophyllium khmer sp. nov., we could not iden-
tify a reliable morphological feature for dierentiation within the males. e female
morphology does allow dierentiation of these species, and of course molecular analy-
sis (Fig. 4) allows reliable dierentiation even between these variable and dicult to
distinguish species.
Distribution. At present only known from the type locality of Phuoc Binh N.P.,
Ninh uan Province, southern Vietnam.
Description. Female. Coloration. Coloration description is based upon living in-
dividuals (Figs 16, 17A). Overall coloration is pale green throughout, with variable ar-
eas highlighted with burnt red or brown coloration. ese areas tend to be the margins
on the lobes of the legs, some striping on the lobes of the legs, the thorax, abdominal
margins, and the venation in the tegmina (Fig. 16).
Morphology. Head. Head capsule about as long as wide, vertex relatively smooth
with the only notable feature being the posteromedial tubercle which is nely pointed
(Fig. 18E). Frontal convexity broad and blunt, with a slightly granular surface. Com-
pound eyes slightly protruding from the head capsule, and are not particularly large,
taking up ca. ¼ of the head capsule lateral margins (Fig. 18E). Ocelli absent. Anten-
nal elds slightly wider than the width of the rst antennomere. Antennae. Antennae
consist of nine segments, with the terminal segment about the same length as the
preceding 2½ segments’ lengths combined (Fig. 6B). Antennomeres I–VIII sparse-
ly marked with small transparent setae, the terminal antennomere and the anterior
margin of antennomere VIII are covered in stout, brown setae (Fig. 18C). orax.
Pronotum with gently concave anterior margin and slightly convex lateral margins,
which converge to a straight posterior margin that is half the width of the anterior
margin (Fig. 18E). e pronotum surface is smooth, with only a prominent pit in the
center, and slight furrows anterior and lateral to the pit (Fig. 18E). e pronotum has
moderately formed anterior and lateral rims and a weakly formed posterior rim, all of
which are relatively smooth (Fig. 18E). Prosternum and the mesosternum are covered
with numerous broad nodes, but the metasternum has a somewhat wrinkled surface.
Prescutum longer than wide, lateral rims with 9–11 small to medium tubercles, simi-
lar in size giving the margin a rough appearance (Fig. 18E). Prescutum anterior rim
prominent but not strongly protruding, rim surface is granular, lacking a large sagittal
spine (Fig. 18F). Prescutum surface heavily granular, with those along the sagittal plane
slightly larger than the rest (Fig. 18E). Mesopleura begin ca. ¼ of the way through the
prescutum length and evenly diverge; lateral margin with nine or ten small tubercles
with about half of those slightly larger than the rest, with the smaller ones interspersed
throughout (Fig. 18E). Face of the mesopleura smooth or slightly wrinkled, with two
notable divots, one on the anterior margin and one near the middle (Fig. 18F). Wings.
Tegmina long, reaching ½ through abdominal segment VII. Tegmina venation; the
subcosta (Sc) is the rst vein in the forewing, running parallel with the margin for the
rst half, and then bending and running towards the margin. e radius (R) spans
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 43
Figure 18. Cryptophyllium bollensi gen. et sp. nov. paratype female, photographs by Jérôme Constant
(RBINS) A habitus, dorsal B habitus, ventral C details of the antennae, dorsal D front leg details, dorsal
E details of the antennae, head, and thorax, dorsal F details of the antennae, head, and thorax, lateral
Gterminalia, dorsal H genitalia, ventral.
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
44
the central portion of the forewing with two subparallel branched veins; the rst ra-
dius (R1) branches ca. ¼ of the way through the wing length and terminates slightly
proximal to the midline, and the radial sector (Rs) branches ca. ⅖ of the way through
the wing length and terminates near the distal ⅓ of the wing length. ere is a weak
continuation of the radius following the prominent Rs branching which continues
on as a short and thin R–M crossvein that weakly connects the two veins. e media
(M) is simply bifurcate with both the media anterior (MA) and media posterior (MP)
terminating near to the posterior ¼ of the wing. e cubitus (Cu) is also bifurcate,
branching near the posterior ⅕ of the wing into the cubitus anterior (CuA) and cubi-
tus posterior (CuP) which both terminate at or very near the wing posterior apex. e
rst anal vein (1A) is simple and fuses with the cubitus early on, at the length about
midway between the splitting of the R1 and Rs. Alae short, with their apex only just
passing the anterior margin of abdominal segment IV. Abdomen. Abdominal segments
II through the anterior half of IV uniformly diverging. e posterior half of segment
IV through the anterior of segment VII are parallel, giving the abdomen a boxy ap-
pearance. e posterior half of segment VII ends in a slightly rounded lobe. Segments
VIII–X are notably narrower than the previous segments, and have converging margins
to the broad rounded apex (Fig. 18G). Genitalia. Subgenital plate starts at the anterior
margin of tergum VIII, is moderately broad, and extends halfway onto tergum X with
straight margins ending in a ne point (Fig. 18H). Gonapophyses VIII are long and
moderately broad, slightly exceeding the apex of abdominal tergum X; gonapophyses
IX are shorter and narrower, hidden below (Figs 17C, 18H). Cerci at, not strongly
cupped, with a granular surface and few detectable setae (Fig. 18H). Legs. Profemoral
exterior lobe broad, rounded, and obtusely angled, smoothly arcing from end to end,
ca. ⅓ wider than the width of the interior lobe (Fig. 18D). Edge of the profemoral
exterior lobe granular, or with a slightly serrate surface of eight or nine small teeth (Fig.
18D). Profemoral interior lobe ca. 2× as wide as the greatest width of the profemoral
shaft, obtusely angled, and marked with ve teeth arranged in a two-one-two pattern
with looping gaps between them (Fig. 18D). Mesofemoral exterior lobe arcs from end
to end but is slightly bent in the center, weighted towards the distal half, and marked
with three or four small serrate teeth distributed on the distal half only. Interior lobe is
about the same width as the mesofemoral shaft, and the exterior lobe is slightly wider.
Mesofemoral interior lobe arcs smoothly end to end with 6–8 small serrate teeth only
on the distal half of the arc which is slightly wider than the proximal half of the arc.
Metafemoral interior lobe arcs end to end, with the distal half slightly wider than
the proximal half and marked with 7–10 serrate teeth on the distal half of the lobe.
Metafemoral exterior lobe is thin and smooth, hugging the metafemoral shaft and
lacks dentation. Protibiae lacking an exterior lobe (Fig. 18D). Protibiae interior lobe
spans the entire length of the protibiae and is ca. 2× the width of the protibiae shaft
itself. e lobe is roundly triangular with the widest portion on the distal half. Mesoti-
biae and metatibiae lacking exterior and interior lobes.
Measurements of paratype female [mm] (wild caught). Length of body (includ-
ing cerci and head, excluding antennae) 75.6, length/width of head 8.0/6.1, anten-
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 45
nae 4.3, pronotum 4.8, mesonotum 7.2, length of tegmina 45.3, length of alae 23.1,
greatest width of abdomen 28.6, profemora 16.7, mesofemora 13.6, metafemora 16.3,
protibiae 10.8, mesotibiae 9.4, metatibiae 12.6.
Measurements of paratype females [mm] (ex culture). Length of body (includ-
ing cerci and head, excluding antennae) 74.6–80.1, length/width of head 7.6–8.5/5.9–
6.2, antennae 3.9–4.2, pronotum 4.5–5.4, mesonotum 6.7–6.8, length of tegmina
43.2–45.1, length of alae (not measurable as they were hidden by the tegmina), great-
est width of abdomen 29.7, profemora 17.2–18.2, mesofemora 13.7–14.4, metafemo-
ra 16.5–16.6, protibiae 10.9–11.0, mesotibiae 9.1–9.6, metatibiae 12.5–12.8.
Male. Coloration. Coloration based upon live bred specimens in captivity
(Fig.17B). Overall coloration pale green throughout with variable patches of tan to
reddish coloration (Fig. 17B). ese tan to reddish areas are primarily around the mar-
gins of the lobes of the legs, the margins of the thorax, the tips of the antennae, and
the margins of the abdomen. In darker colored specimens the mesofemoral lobes can
also have coloration, not just along the margins. Abdominal segment V has a pair of
slightly transparent eye spots.
Morphology. Head. Head capsule about as long as wide, with a vertex that is
relatively smooth with only light granulation throughout. Frontal convexity stout
with sparse thin setae. e posteromedial tubercle is not broad but is distinctly raised
from the head capsule. Compound eyes large and bulbous, taking up ca. ⅖ of the
head capsule lateral margins (Fig. 19E). ere are three well-developed ocelli lo-
cated between and slightly posterior to the compound eyes. Antennae. Antennae (in-
cluding the scapus and pedicellus) consist of 23–26 segments, all segments except
the scapus and pedicellus and terminal three segments are covered in dense setae
that are as long as or longer than the antennae segment is wide. e terminal three
segments are covered in dense short setae and the scapus and pedicellus are nearly
completely bare. orax. Pronotum with anterior margin slightly concave and lateral
margins that are slightly convex and converging to a straight posterior margin that
is ca. ½ the width of the anterior rim (Fig. 19E). Anterior and lateral margins of the
pronotum have moderately formed rims and the posterior margin lacks a rim (Fig.
19D). Face of the pronotum is marked by a distinct furrow and pit in the center and
a relatively smooth lumpy surface with weak granulation (Fig. 19D). Prosternum
surface is weakly granular with small nodes of even size and spacing. Mesosternum
surface marked with slightly more prominent nodes, with the largest along the sagit-
tal plane and denser on the anterior margin, posterior margin with less prominent
and smaller nodes. Prescutum slightly longer than wide, with lateral margins that
are only slightly converging to the posterior (Fig. 19E). Lateral rims with eight or
nine node-like tubercles, giving the lateral margins a rough textured appearance.
Prescutum surface with minimal nodes throughout, with those along the sagittal
plane slightly larger than the others. Prescutum anterior rim moderately formed but
not strongly raised, with a granular surface and lacking a prominent sagittal tubercle.
Mesopleura begin on the anterior prescutum margin but are narrow throughout the
anterior ⅓ of their length, only diverging gently for the posterior ⅔ (Fig. 19E). Lat-
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
46
Figure 19. Cryptophyllium bollensi gen. et sp. nov. holotype male, photographs by Jérôme Constant
(RBINS) A habitus, dorsal B habitus, ventral C pro- tibial and femoral lobes, dorsal D details of the base
of the antennae, head, and thorax, lateral E details of the antennae, head, and thorax, dorsal F terminalia,
dorsal G genitalia, ventral.
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 47
eral margin with eight or nine minor tubercles throughout the length except for the
posterior ⅓ which is relatively smooth. Face of the mesopleura mostly smooth, with
slight wrinkling throughout. Wings. Tegmina moderate length, extending ⅓ of the
way onto abdominal segment III. Tegmina wing venation: the subcosta (Sc) is the
rst vein, is simple, and terminates the earliest ca. ⅓ of the way through the overall
tegmina length. e radius (R) spans the entire length of the tegmina with the rst
radius (R1) branching just proximal to the midline and terminating just distal to the
midline, followed by the branching and termination of the second radius (R2) near
the distal ⅓ of the wing, and then the radial sector runs to the wing apex. e media
(M) also spans the entire length of the tegmina with the rst media posterior (MP1)
branching o slightly more than ⅓ of the way through the wing length, and then
the second media posterior (MP2) branches just distal to the midline, and the media
anterior (MA) runs to the wing apex. e cubitus (Cu) runs along the edge of the
wing as the two media posterior veins fuse with it and as the cubitus reaches the apex
it fades. e rst anal (1A) vein terminates upon reaching the cubitus ca. ⅓ of the
way through the wing length. Alae well developed in an oval fan conguration, long,
reaching to the middle or posterior of abdominal segments IX. Alae wing venation:
the costa (C) is present along the entire foremargin giving stability to the wing. e
subcosta (Sc) is long, spanning ca. ⅔ of the wing length and is mostly fused with the
radius in the beginning but terminates when it meets the costa. e radius (R) spans
the entire wing and branches slightly proximal to the midline into the rst radius
(R1) and radial sector (Rs) which run gently diverging for most of their length and
then converge at the apex of the wing where they terminate near each other but not
touching. e media (M) branches early, ca. ⅙ of the way through the wing into the
media anterior (MA) and the media posterior (MP) which run parallel with each
other throughout the wing until the distal ⅕ of the wing where the media posterior
fuses with the media anterior which then run fused to the wing apex where they
terminate near the radial sector. e cubitus (Cu) runs unbranched and terminates
at the wing apex. Of the anterior anal veins, the rst anterior anal (1AA) fuses with
the cubitus near the point where the media branches into the media anterior and
media posterior and then the rst anterior anal branches from the cubitus ⅔ of the
way through the wing length where it uniformly diverges from the cubitus until it
terminates at the wing margin. e anterior anal veins two–seven (2AA–7AA) have
a common origin and run unbranched in a folding fan pattern of relatively uniform
spacing to the wing margin. e posterior anal veins (1PA–6PA) share a common
origin separate from the anterior anal veins and run unbranched to the wing margin
with slightly thinner spacing than the anterior anal veins. Abdomen. Lateral margins
of abdominal segment II parallel, III through the anterior ⅔ of segment IV gradually
diverging, the remainder of IV and segment V are parallel-sided, segment VI starts
parallel-sided but then gently starts to converge and the remaining segments con-
verge uniformly to the rounded apex of the abdomen. Genitalia. Poculum broad and
ends in a rounded apex that slightly passes the anterior margin of segment X (Fig.
19G). Cerci long and slender, extending from under the anal abdominal segment,
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
48
nearly at, not strongly cupped, covered in a granulose surface and numerous short
setae (Fig. 19G). Vomer broad and stout with straight sides evenly converging and
ending in a thick apical hook with a smaller second hook adjacent to it (Fig. 5E).
Interestingly while examining the vomers of type material we found several aberrant
vomers with some bearing only a singular hook (Fig. 20A) or even three hooks (Fig.
20B), based on other specimens examined and the trend within the Cryptophyllium
gen. nov. males we expect that a typical male of this species has a two hooked vomer.
Legs. Profemoral exterior lobe slightly broader than the interior lobe, ca. 2½× the
greatest width of the profemoral shaft, roundly arcing end to end in a broad obtuse
angle that is not distinctly bent, with the proximal margin slightly granulose, and
the distal margin with four or ve small serrate teeth (Fig.20C). Profemoral interior
lobe roundly triangular and marked with ve sharp teeth arranged in a two-one-two
pattern with looping gaps between them, and the central tooth slightly larger than
the others (Fig. 20C). Mesofemoral exterior lobe arcs end to end but is slightly wider
on the distal ⅓ which is marked with three or four serrate teeth, and a proximal half
that is rather thin. Mesofemoral interior lobe is about the same width as the exterior,
is broader on the distal end and is marked with 6–8 small serrate teeth. Metafemoral
exterior lobe lacks dentation and has a straight margin along the metafemoral shaft.
Metafemoral interior lobe smoothly arcs end to end with eight or nine small serrate
teeth on the distal ⅔, which is slightly wider than the proximal ⅓. Protibiae lacking
exterior lobe, interior lobe reaching end to end in a smooth triangle which is slightly
weighted to the distal half and at its widest is ca. 2½ as wide as the protibial shaft
(Fig. 20C). Meso- and metatibiae simple, lacking lobes completely.
Measurements of holotype male [mm]. Length of body (including cerci and head,
excluding antennae) 57.6, length/width of head 4.⅓.4, antennae 41.4, pronotum 3.0,
mesonotum 4.3, length of tegmina 18.3, length of alae 43.4, greatest width of abdo-
men 16.0, profemora 12.8, mesofemora 11.2, metafemora 13.0, protibiae 8.9, mesoti-
biae 7.7, metatibiae 9.5.
Measurements of paratype males [mm] (ex culture). Length of body (including
cerci and head, excluding antennae) 55.8–65.5, length/width of head 4.1–4.9/3.2–
3.5, antennae 39.9–41.5, pronotum 3.0–3.4, mesonotum 4.0–4.8, length of tegmina
18.2–20.1, length of alae 41.5–47.2, greatest width of abdomen 13.6–16.3, profemora
12.4–13.6, mesofemora 10.4–12.3, metafemora 12.7–14.4, protibiae 8.7–10.2, mes-
otibiae 7.5–8.4, metatibiae 9.0–10.2.
Eggs. (Fig. 21). e overall color is muted dark brown, with the moss-like pinnae
lighter in color, generally tan or light brown. e lateral surfaces are at or slightly con-
vex, with eggs either the same width anterior to posterior or with the posterior of the egg
slightly wider. e lateral surfaces are marked with 40–50 small to medium sized pits,
unevenly spaced in no detectable pattern, with sparse tufts of moss-like pinnae between
these pits (Fig. 21A). e dorsal surface has the micropylar plate spanning a majority of
the length but not quite reaching end to end. On either side of the micropylar plate is
variable pitting (generally eight or so pits) with those one the anterior and posterior ends
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 49
Figure 20. Abnormal male vomer found in Cryptophyllium bollensi gen. et sp. nov. from Vietnam, Phuoc
Binh, observed in the RBINS collection bred from the original limited stock. Prepared and photographed
by Jérôme Constant (RBINS). Scale bars, 1.0 mm. Views 1 ventral 2 right lateral of the ventrally oriented
vomer 3 left lateral of the ventrally oriented vomer A abnormal singularly hooked vomer B abnormal
three hooked vomer.
slightly larger than the central pits (Fig. 21C). e micropylar plate is symmetrical with
the anterior and posterior thin and the middle the widest point. e micropylar cup is not
located at this widest midpoint but is instead located on the posterior ⅓ of the micropy-
lar plate. e micropylar plate margin is lined with short moss-like pinnae. Operculum
slightly ovular, outer margin with a distinct row of short moss-like pinnae and in from
the outer margin is a singular semi-circle of small to medium pits on the dorsal and lateral
aspects (not fully surrounding the apex of the operculum as the ventral portion lacks these
pits; Fig. 21D). Operculum is roundly raised with the height ca. ½ the operculum width
and the apex of the raised operculum has a tuft of moss-like pinnae. e ventral surface
of the egg capsule has a slightly raised sagittal crest marked sparsely with short moss-like
pinnae on the anterior ⅔, and the posterior ⅓ has longer moss-like pinnae. On either side
of this raised sagittal crest is pitting, near the posterior ⅓ on each side of the longer moss-
like pinnae of the sagittal crest is a large circular pit, and anterior to the lowest large pit are
around ten small to medium pits arranged in no detectable pattern (Fig. 21F).
Measurements including the extended pinnae [mm]. Length (including opercu-
lum): 4.1–4.4; maximum width of capsule when viewed from lateral aspect 2.6–2.8;
length of micropylar plate 3.3–3.4
Newly hatched nymphs. (Fig. 9H). e general color throughout the body is dark
brown with slightly lighter brown on the legs. e basitarsi are yellow and remaining
tarsal segments are dark brown. All tibiae lack exterior lobes but do have extremely
thin smoothly arcing interior lobes which have several tan to brown stripes throughout
their length. All femoral lobes are similar in width and have distinct serration on their
distal halves. e interior profemoral lobe lacks a white spot, but the exterior lobe
has a distinct white patch on the proximal ⅓ with an additional small white patch at
the proximal most margin. e meso- and metafemoral interior lobes have two white
patches, one on the proximal most edge, and a larger white patch ⅓ of the way through
the length. e meso- and metafemoral exterior lobes also have a large white patch on
the proximal ⅓, but lack a smaller white patch on the proximal most margin. e dis-
tal ends of the meso- and metafemora also have minimal white edges. e abdomen is
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
50
Figure 21. Cryptophyllium bollensi gen. et sp. nov. egg (RBINS), photographs by Jérôme Constant A lateral
view B dorso-lateral view C dorsal view D opercular (anterior) view E posterior view F ventral view.
mostly brown, but abdominal segments II and III have distinct green patches on their
lateral surfaces (the centerline of the abdomen is uniform brown throughout). e
terminal three abdominal segments also have a little bit of green on their margins. e
widest point of the abdomen is abdominal segment IV.
Etymology. Patronym. Named after Tim Bollens (Belgium) who has been instru-
mental in bringing many new phylliid species into the phasmid breeding community
over the years. With his expertise in breeding these dicult phasmids he has allowed
us to compare the informative sets of male, female, freshly hatched nymph, and egg
morphology instead of only comparing singular dead specimens collected in the wild.
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 51
Cryptophyllium celebicum (de Haan, 1842), comb. nov.
Figures 6C, 8P, 8P, 9A, 22, 23
Material examined. (11 ♀♀, 3 ♂♂, 5 eggs): 6 ♀♀: “Indonesia: Sulawesi” (Coll RC
16-069, 16-070, 16-075, 16-238, (nymph) 16-074, (nymph) 16-072); 2 ♀♀: “In-
donesia: Sulawesi, Palolo, Palu, 2.2008” (Coll RC 16-071, (nymph) 16-073); 1 ♀:
“Indonesia: Peleng, Tattendeng, Sept. 2019” (Coll RC 19-181); 2 ♂♂: “Indonesia:
Sulawesi” (Coll RC 16-146, 16-076); 1 ♂: “Sulawesi, Central Sulawesi Province,
Palu Palolo: February, 2008” (Coll RC 16-145); 1 ♀: “Coll. I.R.Sc.N.B., Indone-
sia, Sulawesi, Puncak BEI, Palopo, VI.2001” (RBINS); 1 ♀: “Indonesia: Bugadidi,
ex culture T. Bollens” (RBINS); 2 eggs: “Indonesia: S-Sulawesi, Tiulapolu leg. Jasmin
III.2008, F-1 Generation, Cultured F.Hennemann 2009 Ex. Coll. Frank Hennemann
(Germany)” (Coll RC 18-250, 18-251); 3 eggs: “Indonesia: Sulawesi; removed from
specimen Coll RC 16-075” (Coll RC 17-345, 17-346, 17-347).
Remarks. is was the rst species described within the newly erected Cryptophyl-
lium gen. nov. and we herein designate it as the type species for this new genus. is
species is now well-known and little confusion surrounds this species’ true identity.
is has not always been the case however as for years it was the subject of repeat-
ed misidentications by many authors (see Hennemann et al. (2009) for a thorough
list of misidentications which instead represented species such as Phyllium ericoriai
Hennemann et al., 2009 from the Philippines and the closely related Cryptophyllium
westwoodii comb. nov. from mainland Asia). Gray (1843) appears to be the rst to er-
roneously state that ‘Phyllium celebicum’ occurs in the Philippines and Wood-Mason
(1875) disrupted the mainland Asia identications when he claimed that ‘Phyllium
celebicum’ could be found in Myanmar. ese two works snowballed for decades as
nearly all specimens from Northern ailand (a major commercial breeding site for
Cryptophyllium westwoodii comb. nov.) were sold as ‘Phyllium celebicum’ therefore con-
fusing collectors and researchers. Additional confusion likely occurred due to the fact
that the last publication explicitly recording the holotype ‘Phyllium celebicum’ appears
to have been by Willemse (1947 [1945]) when he illustrated it and then it subsequent-
ly went missing despite several attempts to locate it by other authors (for example by
Hennemann et al. in April of 2006 in their review of the RMNH collection). ank-
fully the holotype specimen was located by the authors of this work while reviewing
photographs of the RMNH collection and appears to have been overlooked as it was
misplaced and labeled with “Type, Phyllium crurifolium Serv., 1938 (sic!)”, a simple
mistake but one that shows how important proper labeling can be. Now the holotype
is properly labelled, and we here present the rst photographs of this important speci-
men (Fig. 22).
Interestingly, this species is commonly collected and sold from the forest of Sulawesi
solely as green color form specimens, but nearly all captive bred individuals are yellow
to orange in coloration (Fig. 23B, D) with green individuals captive reared quite rare
(Fig. 23A). Molecularly, we unfortunately do not have a wide sampling from through-
out Sulawesi or the surrounding islands so we do not yet know the average intraspecic
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
52
Figure 22. Holotype, Cryptophyllium celebicum (de Haan, 1842), comb. nov. the type species for the
Cryptophyllium gen. nov. Photographs by Luc Willemse, Naturalis Biodiversity Center (RMNH) A dorsal,
habitus B genitalia, ventral C ventral, habitus, and original collection label inset to left D details of the
front legs, head, and thorax, dorsal E details of the antennae and thorax, dorsal.
variation on Sulawesi. We were able to obtain a molecular sample from Peleng Island
o the northeast Sulawesi coast which shows a notable molecular distance from our
single mainland sample (Fig. 4). is Peleng specimen did not have signicant mor-
phological dierences to dierentiate it from the mainland series we examined, and
due to our lack of sampling from throughout Sulawesi within this review we treat this
oshore population as identical. Perhaps additional molecular sampling will reveal
the true intraspecic variation of this species and warrant the Peleng population to be
described as a sister species one day.
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 53
Dierentiation. Females can be dierentiated by the following combination of fea-
tures: mesopleura which are narrow on the anterior half, alae which are ca. ½ the length
of the tegmina, and profemoral exterior lobes which are broad and slightly recurved
which gives them an acute angle at the bend. Two species which are morphologically
very similar are Cryptophyllium echidna sp. nov. and Cryptophyllium limogesi sp. nov. due
to the abdominal and femoral lobe shapes. Cryptophyllium echidna sp. nov. is the molec-
ular sister species to Cryptophyllium celebicum comb. nov. and morphologically very sim-
ilar with the only easy to dierentiate feature being the profemoral exterior lobe which
in Cryptophyllium echidna sp. nov. is nearly right angled, not slightly recurved with an
acute angle. e male and egg morphology are not known for Cryptophylliumechidna
sp. nov. but hopefully once that is observed, additional features can be identied. Cryp-
tophyllium limogesi sp. nov. has a very similarly shaped abdomen and exterior profemoral
lobes, but can immediately be dierentiated by the mesopleura, which are prominent
and reach nearly to the anterior rim (Fig. 42E) vs. Cryptophyllium celebicum comb. nov.
which has the mesopleura narrowed on the anterior rim (Fig. 22E).
Figure 23. Live Cryptophyllium celebicum comb. nov. A green form female, dorsal, bred and photo-
graphed by omas Stijnts (Belgium; Flanders) B–D bred and photographed by Bruno Kneubühler
(Switzerland) B orange form adult female, dorsal C lateral view of the female genitalia holding an egg
ready to be icked away. Note the large gonapophyses VIII and the smaller gonapophyses IX holding the
egg. D orange form adult male, dorsal.
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54
Males are rather morphologically unique as they have profemoral exterior lobes, which
are broad and strongly angled almost to a right angle. No other known males of Crypto-
phyllium gen. nov. have such a prominent profemoral exterior lobe as they either have a
narrow rounded lobe (like in Cryptophyllium yunnanense comb. nov.; Fig. 75B) or an ex-
terior profemoral lobe, which at most is broad with a distinct bend, but still clearly obtuse
(like in Cryptophyllium limogesi sp. nov. or Cryptophyllium oyae comb. nov.; Fig. 50B).
Distribution. Known from throughout the island of Sulawesi and from the nearby
oshore islands of Peleng to the east and Buton to the south.
Cryptophyllium chrisangi (Seow-Choen, 2017), comb. nov.
Figures 8M, 8N, 9B, 24, 25, 26
Material examined. (7 ♀♀, 5 ♂♂, 10 eggs): 2 ♂♂, 2 ♀♀: “Singapore, ex breeding”
(RBINS); 1 ♂: “Singapore (Pulau Ubin), Collected at light, by Francis Seow-Choen,
January 5th, 2018 (0-75 m elv)” (Coll RC 18-279); 1 ♂: “ailand: Satun Province,
Tarutao Island” (UCR); 1 ♀: “Sarawak: Kuching, 12.12, J.M. Bryan., B.M.1931-150.”
(NHMUK); 1 ♀: “P.siccifolium, 4.93 Tapah, Dr. Yeh” (LKCNHM); 10 eggs: “Singapore”
(Coll RC 18-332–18-341); 1 ♀: “Presented by Dr. Brooke, St. John’s I. 26.4.09, Coll
Freie Universitat Berlin Sammlung Exotische Insekten, DEI Hemimetabola #100117”.
Photographic records: 2 ♀♀: ailand: Nakhon Si ammarat Province, ung
Song District, August 2020 (photographed by Tatsatorn Dharithai, ailand);
1 ♂: Indonesia, Pulau Weh Island o the coast of Sumatra (photographed by
Mathieu MJP Van Goethem, South Africa).
Remarks. is species is one which was brought into the breeding community
(Fig. 24A) and has therefore allowed thorough observation of the adults, nymph, and
egg morphology. Molecularly, Cryptophyllium chrisangi comb. nov. is sister species to
Cryptophyllium westwoodii comb. nov. with these two species biogeographically sepa-
rated by the Isthmus of Kra. e Isthmus of Kra is a notable line of biogeography for
several organisms (e.g., Li and Li 2018; Parnell 2013) which separates the Ranong and
Chumphon Provinces of ailand (Fig. 2). is isthmus appears to be signicant for the
Phyllium (Phyllium) as no species are presently known north of this line, but insigni-
cant for the Phyllium (Pulchriphyllium) as this group is found on both sides of this line.
With our herein designation of a neotype for Cryptophyllium westwoodii comb.
nov. we can help to clear up possible signicant confusion which surrounds this spe-
cies. When Wood-Mason described this species in 1875 he did so with a male and
female pair of syntypes from two dierent localities. e female was from “South
Andaman” and the male was from “near Pahpoon, ca. 150 miles north of Moulmein,
in the Salween country” (Wood-Mason 1875). It is almost certain biogeographically
that these syntypes represent two dierent phylliid species, not a singular Cryptophyl-
lium westwoodii comb. nov. as he intended. Morphologically, Wood-Mason’s descrip-
tion and wonderful illustration of the female syntype interestingly appears very similar
morphologically to Cryptophyllium chrisangi comb. nov. with the correct femoral and
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 55
Figure 24. Live Cryptophyllium chrisangi comb. nov. A female bred and photographed by Bruno
Kneubühler (Switzerland) B male genitalia, ventral view, bred by Bruno Kneubühler (Switzerland)
Cmale observed and photographed by Mathieu MJP Van Goethem (South Africa) on Weh Island o
the north shore of Sumatra D female observed and photographed in ailand: Nakhon Si ammarat
Province, ung Song District, by Tatsatorn Dharithai (ailand) in August 2020.
abdominal shapes, and alae length (Fig. 25). With our designation of a neotype Cryp-
tophyllium westwoodii comb. nov. from a male from the mainland, this leaves the Anda-
man female somewhat mysterious as few records of phylliids have been noted from the
Andaman Islands, all of which are Phyllium (Pulchriphyllium) bioculatum-like species,
none representing a Cryptophyllium gen. nov. species. Hopefully one day a fresh speci-
men of this Cryptophyllium gen. nov. species from the Andamans can be located and
molecularly compared with congenerics to identify if it is an additional Cryptophyllium
chrisangi comb. nov. range expansion or an undescribed species.
Dierentiation. Females can be dierentiated by the following combination of
features: spade-shaped abdomen (as segment VII lacks lobes), mesopleura which are
distinctly narrower on the anterior half, and alae which are only ca. ½ as long as the teg-
mina. Morphologically, this species is similar to Cryptophyllium westwoodii comb.nov.
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
56
Figure 25. Wood-Mason’s 1875 plate XVII with the beautifully illustrated ‘Phyllium westwoodii’ female
syntype from South Andaman. Note the morphological similarity to Cryptophyllium chrisangi comb. nov.
and given the wide range of Cryptophyllium chrisangi comb. nov. outside of the type locality of Singapore,
this record may represent a range expansion. Public domain work downloaded from the Biodiversity Her-
itage Library (https://www.biodiversitylibrary.org/item/114410#page/227/mode/1up)
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 57
Figure 26. Tentative female Cryptophyllium chrisangi comb. nov. from Kuching, Sarawak, Malaysia
which represents a range expansion for this species, photographs by Jérôme Constant (RBINS) A habitus,
dorsal B details of the anterior of the head, dorsal C habitus, ventral D front leg details, dorsal E details of
the head and thorax, dorsal F details of the head and thorax, lateral G genitalia details, dorsal H genitalia
details, ventral I data label.
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
58
due to the femoral and mesopleura shape, but can be dierentiated by the shorter alae
(only half of the tegmina length) as Cryptophyllium westwoodii comb. nov. has alae
which are nearly the same length as the tegmina (Fig. 68D). Additionally, Cryptophyl-
lium athanysus comb. nov. is morphologically similar due to the femoral, mesopleura,
and abdominal shape, but can immediately be dierentiated by the metatibial exterior
lobes which are lacking in Cryptophyllium chrisangi comb. nov.
Males are morphologically very similar to Cryptophyllium westwoodii comb. nov.
and we have yet to nd a reliable morphological feature to dierentiate these two spe-
cies. Both are morphologically variable and can even have a wide range of sizes which
does not allow for condent dierentiation when molecular markers and locality are
unknown. One of the more consistent features however is the abdominal shape as
Cryptophyllium westwoodii comb. nov. tends to have a slightly more spade-shaped ab-
domen with segments V–IX converging, and Cryptophyllium chrisangi comb. nov. hav-
ing a slightly more ovoid abdomen with segments V–VI parallel or subparallel, but we
have seen morphological intermediates which do not allow this as a diagnostic feature.
Distribution. e type locality for Cryptophyllium chrisangi comb. nov. is mainland
Singapore and it has additionally been recorded from St. John’s Island from a record in
the SDEI collections. Additionally, we have observed specimens and photographs from
several areas, including several from ailand: Nakhon Si ammarat Province, ung
Song District (Fig. 24D) and Satun Province, Tarutao Island (UCR coll.); one tenta-
tive old record of a large female from Malaysia, Sarawak, Kuching, from the NHMUK
collection (Fig. 26); an adult female from Tapah, Malaysia (LKCNHM); and a record
from Indonesia, Pulau Weh Island o the coast of Sumatra (Fig.24C). Hopefully mo-
lecular samples from these far-reaching areas can one day be obtained to conrm the
identication as Cryptophyllium chrisangi comb. nov. but for now morphologically this
is what these records appear to represent.
Cryptophyllium daparo gen. et sp. nov.
http://zoobank.org/92D078EF-D482-4F03-AB57-E63EEFE742A5
Figure 27
Material examined. Holotype ♀: “CHINA: Yunnan, Wangtianshu, Mengla County,
Xishuangbanna Prefecture, VII-2016, Legit: Xiao-Yu Zhu”. Deposited in the Kun-
ming Institute of Zoology (KIZ), Yunnan, China.
Remarks. is large species is at present only known from the single holotype
female, which has a unique set of morphological features which do not link it to the
molecularly recovered closely related species of Cryptophyllium drunganum comb. nov.
and Cryptophyllium tibetense comb. nov. Geographically, these species do represent
some of the highest latitude species and interestingly Cryptophyllium daparo sp. nov.
and Cryptophyllium tibetense comb. nov. are the largest species in this genus. Hopefully
further collection eorts in this region will reveal the unknown male and the presently
unknown egg morphology.
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 59
Dierentiation. Females are morphologically most similar to Cryptophyllium ath-
anysus comb. nov. and Cryptophyllium rarum comb. nov. due to the tapered, spade-like
abdomen and the anteriorly narrower mesopleura. From both species Cryptophyllium
daparo sp. nov. can be dierentiated by the exterior profemoral lobe shape which is
distinctly obtuse and nearly rounded in its shape, not smoothly right-angled like in the
other species. Cryptophyllium daparo sp. nov. is also notably larger than Cryptophyllium
athanysus comb. nov. and Cryptophyllium rarum comb. nov. which are only 77 and 88
mm long respectively (Hennemann et al. 2009; Liu 1993).
Figure 27. Holotype female Cryptophyllium daparo gen. et sp. nov., photographs by Zhiwei Dong (KIZ)
A habitus, dorsal B details of the face and antennae, rostral view C profemoral lobes, dorsal D details of
the head and thorax, dorsal E terminalia, dorsal F details of the head and thorax, lateral G genitalia, ventral.
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
60
Males are presently unknown, and the unique female morphology means we can-
not predict much about the male morphology. e only feature which can be guessed
is that the male must also be rather large due to the female size.
Distribution. At present only known from the unique holotype collected in Chi-
na, Yunnan, Wangtianshu, Mengla County, Xishuangbanna Prefecture.
Description. Female. Coloration. Coloration description is based upon the dried
holotype which is somewhat discolored (Fig. 27). Leaf insects are a more vibrant lime
green in life and we expect that this specimen was likely originally this color. e holo-
type female is mostly tan to pale green throughout and appears to not have any natural
brown patches of color as are sometimes present on the lobes or thorax.
Morphology. Head. Head capsule about as long as wide, vertex with moderately
spaced small granulation, and two notable tubercles, one on each side of the sagittal
plane near the midline of the head capsule which are larger than the rest, but not as
prominent as the posteromedial tubercle (Fig 27D). e posteromedial tubercle is
about as broad as the two tubercles on the capsule, but the posteromedial tubercle is
notably taller and pointed (easily seen from the lateral aspect; Fig. 27F). Frontal con-
vexity broad but not very large, with an irregularly lumpy surface and sparse transpar-
ent setae throughout. Compound eyes only slightly protruding from the head capsule
and only occupying ca. ⅕ of the head capsule length (Fig. 27D). Ocelli absent. Anten-
nae. Antennae consisting of nine segments, with the terminal segment slightly shorter
than the length of the preceding two segments’ lengths combined (Fig. 27B). Anten-
nomeres I–VIII sparsely marked with small transparent setae, the terminal antenno-
mere is covered in stout, brown setae. orax. Pronotum with a distinctly concave
anterior margin and nearly straight lateral margins, which converge to a straight pos-
terior margin that is half the width of the anterior margin (Fig. 27D). e pronotum
surface is marked with slight granulation, with a prominent pit in the center, and dis-
tinct furrows anterior and posterior to the pit and slight furrows lateral to the central
pit (Fig. 27D). e pronotum has a prominent anterior rim which is marked through-
out by minute setae and moderately formed lateral rims, and a posterior rim which
is weakly formed (Fig. 27D). Prosternum with moderate nodes, irregularly sized and
spaced. Mesosternum with similar nodes as on the pronotum, but only along the mar-
gins and on the anterior half of the sagittal plane, the remainder of the surface is rather
smooth. Metasternum with an irregularly lumpy surface, no strong nodes. Prescutum
slightly longer than wide and with nearly parallel lateral margins (Fig.27D). Lateral
rims with eight or nine irregularly shaped but short tubercles with various small nodes
mixed throughout, giving the margins a rough textured appearance (Fig.27D). Pres-
cutum anterior rim distinct, but not strongly raised above the prescutum surface, the
rim has a granular surface, no distinct sagittal tubercle larger than the rest (Fig.27F).
Prescutum surface with irregular nodes throughout, with those along the sagittal plane
slightly more prominent (Fig. 27D). Mesopleura beginning slightly posterior to the
anterior margin of the prescutum and evenly diverging; lateral margin with three or
four larger tubercles throughout the length, and ten or eleven smaller node-like tu-
bercles interspersed (Fig. 27D). Face of the mesopleura with granulation throughout
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 61
and slight wrinkling on areas where the nodes are less prominent (Fig. 27F). Wings.
Tegmina long, extending three quarters of the way through abdominal segment VII.
e subcosta (Sc) is the rst vein in the forewing and runs parallel with the wing for
the rst half and then distinctly bends towards the distal margin where it terminates
ca. ¼ of the way through the wing length. e radius (R) lls approximately the ante-
rior ⅓ of the wing as two subparallel branched veins; radius 1 (R1) terminates ca. ⅓
of the way through the wing length, and the radial sector (Rs) terminates posterior to
the widest portion of the tegmina, just past the midline. ere is a thinner continua-
tion of the radius following the prominent Rs branching which continues on as a short
R–M crossvein that does not appear to solidly connect the two veins. e media (M)
is bifurcate with both the media anterior (MA) and media posterior (MP) terminat-
ing close to the posterior ⅓ of the wing. e cubitus (Cu) is also bifurcate, branching
near the posterior ¼ of the wing into the cubitus anterior (CuA) and cubitus posterior
(CuP) which both terminate at or very near the wing posterior apex. e rst anal
vein (1A) is simple and fuses with the cubitus early on, only slightly past the branch-
ing distance of the R1 from the R. Alae well developed, 58.5 mm long, only a little
shorter than the tegmina. Abdomen. Segments II through the anterior ⅓ of IV gradu-
ally diverging, with the posterior ⅔ of segments IV and V parallel-sided. Segments
VI–X are gradually converging to the broadly rounded apex, giving the abdomen a
smooth spade-shaped appearance (Fig. 27A). Genitalia. Subgenital plate starts at the
anterior margin of segment VIII, is broad, and extends ca. ⅔ of the way onto segment
X, ending in a ne point (Fig.27G). Gonapophyses VIII are long and moderately
broad, slightly exceeding the apex of abdominal segment X (Fig. 27G); gonapophyses
IX are thinner and shorter and are concealed below the larger gonapophyses VIII.
Cerci at, not strongly cupped, with a granular surface and rough granular lateral
margins (Fig. 27E). Legs. Profemoral exterior lobes slightly broader than the interior
lobe, roundly arcing from end to end in a broad obtuse angle (Fig. 27C). Edge of
the profemoral exterior lobe with a highly granular surface on the proximal margin,
and the distal margin has less granulation, but does have ve or six small but notable
teeth (Fig. 27C). Profemoral interior lobe ca. 3× as wide as the greatest width of the
profemoral shaft, and with a distinct obtuse angle and a distal margin marked by four
or ve prominent serrate teeth, with a larger gap between the middle teeth (Fig. 27C).
Mesofemoral exterior lobe arcs from end to end with a distinct bend near the center
slightly weighted towards the distal half and marked with two or three dulled serrate
teeth on the distal half only. Interior mesofemoral lobe is slightly narrower than the
exterior lobe. Mesofemoral interior lobe arcs end to end with ve or six serrate teeth
on the distal half of the arc only. Metafemoral interior lobe narrow, arcing end to end,
and marked with ve or six serrate teeth and slight granulation on the distal half of
the lobe only. Metafemoral exterior lobe is thin and smooth, hugging the metafemoral
shaft and lacks teeth. Pro-, meso-, and meta- tibiae lacking exterior lobes. Protibial
interior lobe spans the entire length, is ca. 2× the width of the shaft, and is roundly
triangular with the widest portion on the distal half (Fig. 27A). Meso- and metatibiae
lacking interior lobes.
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62
Measurements of holotype female [mm]. Length of body (including cerci and
head, excluding antennae) 107.0, length/width of head 8.5/7.0, antennae 5.0, prono-
tum 6.3, mesonotum 7.0, length of tegmina 67.5, length of alae 58.5, greatest width of
abdomen 32.0, profemora 27.0, mesofemora 19.2, metafemora 20.0, protibiae 17.4,
mesotibiae 18.0, metatibiae 18.5.
Etymology. Noun, named for the artistic company “DAPARO”, owned by Dap-
aro-Yeung which is well-known for their beautiful natural history themed brooches.
Several years ago, DAPARO even produced a beautifully crafted leaf insect themed
brooch which helped to shed light on these beautiful creatures and bring them into
the public eye.
Cryptophyllium drunganum (Yang, 1995), comb. nov.
Figures 28, 29
Material examined. We examined the holotype female from within the Beijing Agri-
cultural University from detailed photographs taken by Yu-Chen Zheng (China Ag-
ricultural University, China). Additionally, we examined a tentatively identied male
from “Yunnan China: Qinglangdang, Dulongjiang Township, Gongshan County, Nu-
jiang Prefecture, II-2016, Local” (Coll ZD).A tentative male specimen collected very
near the type locality was used in our molecular analysis, which cannot be condently
conrmed as Cryptophyllium drunganum comb. nov. due to extreme sexual dimor-
phism of the phylliids and lack of a fresh tissue sample from a true Cryptophyllium
drunganum comb. nov. female (Fig. 29).
Remarks. is species is only known at present from the morphologically unique
holotype female from northern Yunnan Province (Fig. 28). However, this area is not
known for a high diversity of species, so we are fairly condent that this male repre-
sents the undescribed Cryptophyllium drunganum comb. nov. male. Also, this male
specimen was molecularly recovered as distinct to the other species described from
China as we have successfully sampled almost all species of Cryptophyllium gen. nov.
and included them in our molecular phylogeny (Fig. 4).
Dierentiation. Females are morphologically similar to Cryptophyllium tibetense
comb. nov. and Cryptophyllium liyananae sp. nov. due to the long alae, rounded exte-
rior profemoral lobe, mesopleura which are distinctly reaching the anterior margin but
slightly curved on the anterior end (not perfectly straight margined), boxy abdomen
with a notable bend on abdominal segment VII, and the presence of small exterior
lobes on all tibiae. Both of these species can be dierentiated by the length of the
subgenital plate as in Cryptophyllium drunganum comb. nov. it is short, just passing
the anterior margin on the tenth abdominal segment, and in the other species it is at
least three quarters of the length of the tenth abdominal segment (in Cryptophyllium
liyananae sp. nov.) or even longer and exceeding the tip of the abdomen (in Cryptophyl-
lium tibetense comb. nov.).
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 63
Figure 28. Cryptophyllium drunganum comb. nov. holotype female within the Beijing Agricultural Uni-
versity, photographed by Yu-Chen Zheng (China Agricultural University, China) A habitus, dorsal, inset
specimen data labels B details of antennae, head, and anterior of the thorax.
Our male specimen is morphologically similar to Cryptophyllium tibetense comb.
nov. and Cryptophyllium yunnanense comb. nov. due to the shape of the profemoral
exterior lobe which smoothly arcs end to end without a distinct bend, the exterior
profemoral lobe that is the same width or slightly thinner than the interior lobe (not
wider as is common in many of the Cryptophyllium gen. nov. species), tegmina which
are long reaching the anterior margin of abdominal segment IV or slightly passing it, a
similar spade-shaped abdomen, and prominent tubercles on the mesopleura.
Cryptophyllium tibetense comb. nov. males additionally have small exterior tibial
lobes on the distal ends like are present in our Cryptophyllium drunganum comb. nov.
male. Our male Cryptophyllium drunganum comb. nov. can however be dierentiated
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
64
Figure 29. Presumed male Cryptophyllium drunganum comb. nov. male used within our molecular analy-
sis (sample DZW05), from Du Longjiang Township, Nujian Lisu Autonomous Prefecture, photographs
by Zhiwei Dong (KIZ) A habitus, dorsal B details of the legs and thorax, ventral.
from Cryptophyllium tibetense comb. nov. by the presence of eight or nine small serrate
teeth present throughout the full length of the profemoral exterior lobe vs. Cryptophyl-
lium tibetense comb. nov. males which only have two or three small teeth on the distal
end only. Additionally, the mesofemoral exterior lobe also can dierentiate these spe-
cies as it is distinctly angled in our Cryptophyllium drunganum comb. nov. male and
smoothly arcing without a distinct bend in Cryptophyllium tibetense comb. nov. males.
Cryptophyllium yunnanense comb. nov. can be dierentiated by the absence of ex-
terior tibial lobes and the lack of a distinctly serrate exterior profemoral lobe margin,
with Cryptophyllium yunnanense comb. nov. only having two or three small teeth vs.
our male Cryptophyllium drunganum comb. nov. which has eight or nine small serrate
teeth present throughout the full length of the profemoral exterior lobe.
Distribution. At present only known from northern Yunnan Province, from the
type locality of Nujian Lisu Autonomous Prefecture, Gongshan County (Drung-Nu),
and our tentative male Cryptophyllium drunganum comb. nov. from Du Longjiang
Township, Qing Lang Dang in the same prefecture.
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 65
Cryptophyllium echidna gen. et sp. nov.
http://zoobank.org/076CABC1-2772-4E56-9918-F3804B62356E
Figure 30
Material examined. Holotype ♀: INDONESIA: Wangi-wangi Island. Collected prior
to 2020, but no exact date given. Deposited in the Montreal Insectarium (IMQC).
Remarks. is is the rst phylliid record we have seen from the small island of
Wangi-wangi in the Wakatobi Regency in Southeast Sulawesi Province. is small
island appears to be rather unique biogeographically and unexplored as a yet to be
described bird species presently known as the “Wangi-wangi White-eye” (Zosterops
sp. nov.) has recently been identied as well, suggesting this island may hold many
endemic undescribed species (O’Connell et al. 2019).
Dierentiation. Presently we only know of a single female specimen of this new
species which we here designate as the holotype. Morphologically and molecularly this
species is closely related to Cryptophyllium celebicum comb. nov. which has a much
wider distribution to the north on the islands of Buton, Sulawesi, and Peleng (Fig. 2).
Female Cryptophyllium echidna sp. nov. can be dierentiated from Cryptophyllium
celebicum comb. nov. by only subtle dierences in the thorax and profemoral exte-
rior lobes. In Cryptophyllium echidna sp. nov. the prescutum is slightly broader and
with a weaker sagittal crest (Fig. 30D) than in Cryptophyllium celebicum comb. nov.
(Fig.22E). Additionally the profemoral exterior lobe of Cryptophyllium echidna sp.
nov. has a right exterior angle (Fig. 30A), not acute like in Cryptophyllium celebicum
comb. nov. (Fig. 22D).
Males are presently unknown, but as the sister species to Cryptophyllium celebicum
comb. nov. the males likely have a similar morphology.
Distribution. At present only known from the small Indonesian island of Wangi-
wangi o the east coast of Buton Island.
Description. Female. Coloration. At present we only have the dried holotype
female to describe the color from which has a bit of rot through the legs, head, thorax,
and the central area of the abdomen. e rotten areas are brown but are assumed to
have been green in life. e remainder of the female is lime-green in color through-
out, with no indication of natural brown patches (which even on somewhat rotten
specimens can generally be identied) but this female appears to have been uniform
green in life.
Morphology. Head. Head capsule slightly longer than wide, vertex with minimal
granulation throughout the surface, all relatively well-spaced with no areas on the
head tightly packed. e posteromedial tubercle is the most prominent feature on the
vertex of the head capsule. Frontal convexity broad and stout, shorter than the length
of the rst antennomere, and with a lumpy surface marked by few short transparent
setae. Compound eyes not particularly large, only slightly protruding from the head
capsule, taking up ca. ¼ of the length of the lateral head capsule margins (Fig. 30A).
Ocelli absent. Antennal elds slightly wider than and about as long as the length of
the rst antennomere. Antennae. Antennae consist of nine segments, with the terminal
segment approximately the same length as the preceding two segments’ lengths com-
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
66
Figure 30. Cryptophyllium echidna gen. et sp. nov. holotype female, photographs by René Limoges
(IMQC) A habitus, dorsal B dorsal details of the antennae C habitus, ventral D dorsal details of the
thorax E ventral details of the genitalia.
bined (Fig. 30B). Antennomeres I–III sparsely marked with small transparent setae
(with the longest on the rst segment), the terminal two antennomeres are densely
covered in stout, brown setae. orax. Pronotum with a gently concave anterior mar-
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 67
gin and slightly convex lateral margins, which converge to a straight posterior mar-
gin that is half the width of the anterior margin (Fig. 30D). e pronotum surface
lacks granulation but is slightly lumpy, with only a prominent pit in the center, and
slight furrows anterior and lateral to the pit (Fig. 30D). e pronotum has a promi-
nent anterior rim and moderate lateral and posterior rims (Fig. 30D). Prosternum
and mesosternum with numerous nodes throughout the surface, all about the same
size and spacing throughout. Metasternum with slightly less granulation but they are
slightly larger than those on the pro- and mesosternum. Prescutum longer than wide,
with a slightly broader anterior margin (Fig. 30D). Lateral rims with 6–8 prominent
tubercles with various small, lumpy granules interspersed throughout the margins
(Fig. 30D). Prescutum anterior rim prominent but not strongly protruding, surface is
granular and lacks a prominent sagittal spine (Fig. 30D). Prescutum surface without a
strongly raised sagittal crest, instead the surface is only slightly raised along the sagittal
plane. e prescutum surface has moderate granulation throughout ranging in size
from small to medium with irregular spacing (Fig. 30D). Mesopleura not spanning
the entire length, instead with the anterior ⅓ narrow and only starting to fan out
near the midline of the prescutum length. Mesopleura lateral margins with ve or six
larger, sharp tipped tubercles with an additional ve or six smaller nodes interspersed
throughout (Fig. 30D). Face of the mesopleura lacking granulation, but instead highly
wrinkled and with two notable pits, one on the anterior ⅓ and one nearer the pos-
terior ⅓ (Fig. 30D). Wings. Tegmina long, reaching nearly to the posterior margin
of abdominal segment VII. e subcosta (Sc) is the rst vein in the forewing and
runs subparallel with the wing for the rst half of its length, and then bends towards
the wing margin for the second half. e radius (R) spans the central portion of the
tegmina with two subparallel branched veins. e rst radius (R1) branches ca. ⅗ of
the way through the radius length and terminates ca. ⅓ of the way through the wing
length. e radial sector (Rs) branches from the end of the radius and runs angled to
the wing margin where it terminates just posterior to the wing midline length. ere
is a weak continuation of the radius following the prominent radial sector branching
which continues on as a short and thin radius to media crossvein (R–M). e media
(M) is simply bifurcate with both the media anterior (MA) and media posterior (MP)
terminating close to the posterior ¼ of the wing. e cubitus (Cu) runs throughout
the entire wing length simply, and then near the posterior ⅕ of the wing becomes
bifurcate into the cubitus anterior (CuA) and cubitus posterior (CuP) which both ter-
minate at or very near the wing posterior apex. e rst anal vein (1A) is simple and
fuses with the cubitus early on, near where the radial sector branches from the radial.
Alae of moderate length, reaching abdominal segment IV. Abdomen. Abdominal seg-
ments II through the anterior ½ of IV diverging, with the middle of segment IV the
widest segment. Segments V–VII gently converging, with segment VII ending with
a distinct lobe which bends inward to a notably narrower segment VIII. Segments
VIII–X converging to a broad apex. Genitalia. Subgenital plate starts at the anterior
margin of segment VIII, is broad, and only extends ⅓ of the way under segment X,
ending in a ne point (Fig. 30E). Gonapophyses VIII are long (exceeding the tip of
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
68
the abdomen but not as long as the tips of the cerci) and moderately broad (together
side by side are about as broad as the subgenital plate; Fig.30E). Gonapophyses IX are
smaller and shorter, and mostly covered from view by the notably larger gonapophyses
VIII. Cerci at, with a granular surface throughout and few setae on the ventral sur-
face, the dorsal surface has thin, transparent setae throughout the surface (Fig. 30E).
Legs. Profemoral exterior lobes broad (ca. 2× width of the interior lobe) and approxi-
mately right angled (Fig. 30A). Proximal edge of the exterior profemoral lobe gently
undulates giving this margin a slightly wavy appearance, whereas the distal margin is
nearly straight, both margins have notable serration throughout their lengths (Fig.
30A). Profemoral interior lobe with a slightly obtuse angle and marked with ve large,
serrate teeth with looping gaps between them. ese teeth are arranged somewhat into
a two-one-two pattern with the gaps between these sets larger. Mesofemoral exterior
lobe arcs from end to end and is almost evenly weighted on both sides, but with the
broadest point just o center to the distal side of the midline, and on the distal half
only marked with three small serrate teeth. Interior mesofemoral lobe arcs end to end,
is ca. ¼ narrower than the exterior lobe, not as strongly angled, and marked with six
or seven teeth on the distal half. Metafemoral interior lobe arcs end to end but is no-
tably wider on the distal ⅔ of the lobe and this wider portion is marked by 11 or 12
serrate teeth. Metafemoral exterior lobe is thin and smooth, hugging the metafemoral
shaft and marked with one or two small, rounded teeth on the distal edge. Protibiae
lacking an exterior lobe. Protibiae interior lobe spans the entire length of the protibiae
as a rounded scalene triangle with the widest portion on the distal ⅓. Mesotibiae and
metatibiae simple, lacking exterior and interior lobes.
Measurements of holotype female [mm]. Length of body (including cerci and
head, excluding antennae) 94.5, length/width of head 7.8/7.5, antennae 5.0, prono-
tum 6.6, mesonotum 7.9, length of tegmina 59.1, length of alae 31.6, greatest width of
abdomen 41.3, profemora 20.7, mesofemora 16.5, metafemora 21.2, protibiae 14.0,
mesotibiae 12.1, metatibiae 16.4.
Etymology. Noun, Greek in origin. Relating to the tenth labor of Heracles (ap-
parently a favorite story of Gray (1843) as four of his therein described species came
from this myth) in which Heracles was tasked with capturing the red cattle from the
monster Geryon (for which Gray named Phyllium geryon Gray, 1843). Before Heracles
sailed out to the island Erythia where Geryon and his red cattle lived, he rested on the
mainland and forgot to tie up his horses. When he eventually found them, they were
in a cave with an Echidna (a monster which is described as women from the waist up
and snake on her lower half). She refused to give back his horses unless he lay with her,
which being a classic Greek hero, he did. Before he left the Echidna, she told him that
she was pregnant with three of his sons and asked him which of the three should rule
her lands one day. Heracles then left her with his bow and a girdle, and told her that
whichever of his three sons could draw the bow and wear the girdle best would inherit
her land and the other two should be banished. ose three sons were Agathyrsus,
Gelonus, and Scythes (for which Gray named Phyllium agathyrsus Gray, 1843; Phyl-
lium gelonus Gray, 1843; and Phyllium scythe Gray, 1843).
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 69
We felt that this species could help to nish telling the story which Gray was so
fond of. With Phyllium geryon being a species from the Philippines, one biogeographi-
cal bridge for species to the Philippines is from Sulawesi through the Sangihe Islands
(Evans et al. 2003), therefore as a steppingstone to the Philippines we felt there should
be an Echidna along the route to the Philippines where Phyllium geryon can be found.
Cryptophyllium faulkneri gen. et sp. nov.
http://zoobank.org/30DBFBC7-7AF0-46ED-9A3F-E50D0F17E457
Figure 31
Material examined. Holotype ♂: “VIETNAM: Quang Ngai Province, Bato Mt. 900
m. elv: May 2015 (Coll RC 16-114)”. Molecularly sampled within our analysis. De-
posited in the Montreal Insectarium (IMQC).
Paratypes (2♂): 1 ♂: “Ngoc Linh, Kon Tum Prov. Vietnam, 1700 m, VI.2016,
leg. Luong coll. TB-05-134’ (Coll TB) • ♂ nymph: “VIETNAM: Lam Dong, Bao
Lam, Da Tom: March 2016 (Coll RC 16-236)”, molecularly sampled within our anal-
ysis (Coll RC).
Remarks. is large species was immediately identied as distinct by the size and
additionally by the large, prominent tubercles of the mesopleura, which are not as
distinct in males of other Cryptophyllium gen. nov. species. Despite several expeditions
to southern Vietnam by the RBINS expedition members, no possible female for this
species has been located to date, despite the fact that the female is likely a very large
phylliid. Hopefully future expeditions reveal the female so the morphology to conge-
nerics can be compared.
Dierentiation. Females are presently unknown. Male Cryptophyllium faulkneri
sp. nov. are most morphologically similar to Cryptophyllium limogesi sp. nov. due to the
large size and thorax spination and Cryptophyllium animatum sp. nov. due to the super-
cially similar appearance of a slender abdomen and large overall size. Cryptophyllium
faulkneri sp. nov. can be dierentiated from Cryptophyllium animatum sp. nov. by the
mesopleura spination (as Cryptophyllium faulkneri sp. nov. has meseopleurae with four
large protruding tubercles and ve smaller interspersed nodes; Fig. 31A) vs. Cryptophyl-
lium animatum sp. nov. which has six small sized tubercles and six or seven small nodes
interspersed (Fig. 10C). Additionally, Cryptophyllium faulkneri sp. nov. has a rather
average interior profemoral lobe serration pattern with irregular sized and spaced teeth
(Fig. 31B), vs. Cryptophyllium animatum sp. nov. which has teeth all evenly spaced and
sized (Fig. 10B). Cryptophyllium faulkneri sp. nov. can be dierentiated from Crypto-
phyllium limogesi sp. nov. by the narrower abdomen (rather broad and spade-shaped
in Cryptophyllium limogesi sp. nov.), a lack of small exterior tibial lobes (as Cryptophyl-
lium limogesi sp. nov. has distinct small exterior tibial lobes), and the interior protibial
lobe shape which in Cryptophyllium limogesi sp. nov. is weighted towards the anterior
⅓ in a scalene triangle, vs. Cryptophyllium faulkneri sp. nov. which has the lobe evenly
weighted with the widest portion in the middle as an isosceles triangle.
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
70
Figure 31. Holotype Cryptophyllium faulkneri gen. et sp. nov., photographs by RTC A details of the
base of antennae, head, and thorax, dorsal B habitus, dorsal C details of thorax, ventral D details of head
and thorax, lateral.
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 71
Distribution. Presently only known from central and southern Vietnam, no oth-
er specimens are presently known. e two Vietnamese provinces that this species is
known from are Quang Ngai and Lam Dong Provinces.
Male. Coloration. Coloration description based on the dried holotype specimen,
not on living individuals which are likely a more vibrant green. Overall coloration is
pale green with variable patches of yellow throughout due to the drying of the specimen
(Fig. 31B). Compound eyes are rust red (Fig. 31A). ere are no hints of natural brown
coloration on the holotype and there are no eye spots present on the abdomen.
Morphology. Head. Head capsule slightly longer than wide, with a vertex that
lacks granulation; posteromedial tubercle notable as the only feature on the poste-
rior half of the head capsule (Fig. 31A). Frontal convexity stout, with a somewhat
broad point, and with numerous short setae throughout the surface. Compound eyes
large but not overly bulbous, occupying slightly > ⅓ of the head capsule lateral mar-
gins (Fig.31A). Between the compound eyes are three well-developed ocelli. Anten-
nal elds are slightly wider than, and approximately as long as the scapus. Antennae.
Antennae (including the scapus and pedicellus) consists of 28 segments. e scapus
and pedicellus are mostly bare but with few very short setae throughout, segments III–
XXV are covered in dense, thin, pale setae that are as long as or longer than the anten-
nae segment is wide. e terminal three segments have dense short setae throughout
that are notably shorter than the segments are wide. orax. Pronotum with anterior
margin gently concave and lateral margins that are slightly convex and converge to a
straight posterior margin that is ca. ½ the width of the anterior rim (Fig. 31A). Ante-
rior and lateral margins of the pronotum have distinct rims and the posterior margin
lacks a rim (Fig.31A). Face of the pronotum is marked by a smooth surface, distinct
sagittal furrow on the anterior half, a pit just posterior to the center, and a moderate
perpendicular furrow just anterior to the central pit (Fig. 31A). Prosternum only has
minimal small granulation on a mostly smooth surface, the mesosternum surface on
the anterior half is heavily granulose and the posterior half is wrinkled to smooth (Fig.
31C). e metasternum lacks granulation but has a smooth wrinkled surface through-
out. Prescutum narrow, longer than wide, with lateral margins that are straight and
marked with four or ve large smooth tubercles of almost equal size with few smaller
nodes interspersed between them, spaces between the prominent tubercles relatively
smooth (Fig. 31A).e surface of the prescutum is slightly raised along the sagittal
plane which is marked with nine or ten, smooth nodes of slightly varying size, with
the remainder of the prescutum surface lacking nodes, but with a slightly wrinkled
surface instead (Fig.31A). Prescutum anterior rim with a slightly lumpy surface and a
moderately formed central tubercle that rises above the prescutum slightly (Fig. 31A).
Mesopleura narrow on their anterior ⅓, then gradually diverging with straight mar-
gins on the posterior ⅔ (Fig. 31A). Lateral margin with ve or six large protruding
tubercles and ve or six smaller nodes interspersed throughout (Fig. 31A). Face of
the mesopleura smooth but slightly wrinkled and with two distinct pits, one near the
anterior and one on the posterior ⅓. Wings. Tegmina moderate length, extending ½
through abdominal segment III. Tegmina venation: the subcosta (Sc) is the rst vein,
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
72
is simple, and terminates the earliest ca. ⅖ of the way through the overall tegmina
length. e radius (R) spans the entire length of the tegmina with the rst radius (R1)
branching just proximal to the midline and terminating just distal to the midline,
followed by the branching and termination of the second radius (R2) near the distal
⅓ of the wing, and then the radial sector runs to the wing apex. e media (M) also
spans the entire length of the tegmina with the rst media posterior (MP1) branching
o just proximal to the midline, and then the second media posterior (MP2) branches
just posterior to the midline, and the media anterior (MA) runs to the wing apex. e
cubitus (Cu) runs along the edge of the wing as the two media posterior veins fuse with
it and as the cubitus reaches the apex it fades. e rst anal (1A) vein terminates upon
reaching the cubitus ca. ⅓ of the way through the wing length. Alae well-developed
in an oval fan conguration, long, reaching ⅔ of the way through abdominal segment
VIII. Alae venation: the costa (C) is present along the entire foremargin giving stability
to the wing. e subcosta (Sc) is long, spanning slightly < ⅔ of the wing length and
is fused with the radius in the beginning but terminates when it meets the costa. e
radius (R) spans the entire wing and branches ca. ⅖ of the way through into the rst
radius (R1) and radial sector (Rs) which run gently diverging for most of their length
and then converge at the apex of the wing where they terminate near each other but
not touching. e media (M) branches early, ca. ⅙ of the way through the wing into
the media anterior (MA) and the media posterior (MP) which run parallel with each
other throughout the wing until the distal quarter of the wing where the media pos-
terior fuses with the media anterior which then run fused together to the wing apex
where they terminate near the radial sector. e cubitus (Cu) runs unbranched and
terminates at the wing apex. Of the anterior anal veins, the rst anterior anal (1AA)
fuses with the cubitus near the point where the media branches into the media anterior
and media posterior and then the rst anterior anal branches from the cubitus ⅔ of
the way through the wing length where it uniformly diverges from the cubitus until
it terminates at the wing margin. e anterior anal veins two–seven (2AA–7AA) have
a common origin and run unbranched in a folding fan pattern of relatively uniform
spacing to the wing margin. e posterior anal veins (1PA–6PA) share a common ori-
gin separate from the anterior anal veins and run unbranched to the wing margin with
slightly thinner spacing than the anterior anal veins. Abdomen. Abdomen is a narrow
dagger-like shape. Abdominal segment II with parallel-sided margins, III with gen-
tly diverging margins, the anterior half of segment IV gently diverging to the widest
point of the abdomen. e remainder of segment IV through the apex are converging
uniformly with straight margins to the apex. Genitalia. Poculum broad and rounded,
ending in a straight margined apex that passes onto segment X. Cerci not exceptionally
long, with only ca. ½ of their length protruding from under the terminal abdominal
segment. e cerci are relatively at, not strongly cupped, covered in a granulose sur-
face and numerous short setae throughout. Vomer broad and stout with straight sides
evenly converging to the apex which is armed with two upwards turning hooks. Legs.
Profemoral exterior lobe a rounded arc, about as wide as the interior lobe (ca. 3× the
width of the profemoral shaft at its widest), and with the anterior half marked by ve
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 73
small, rounded teeth (Fig.31B). Profemoral interior lobe an obtusely rounded triangle
with six, serrate, anteriorly pointing teeth arranged in pairs with looping gaps between
them (in the holotype the middle pair of teeth are uneven in size, one notably larger
than the other; Fig. 31B). Mesofemoral exterior lobe arcs end to end but with the wid-
est portion slightly distal to the midline, and the widest point slightly wider than the
interior lobe. e mesofemoral exterior lobe is marked with six or seven small serrate
teeth distal to the widest point, with the proximal portion of the lobe smooth. e
mesofemoral interior lobe is approximately the same width as the mesofemoral shaft
with ve or six small serrate teeth on the distal half only. Metafemoral exterior lobe
thin, lacks dentation, and has a straight margin along the metafemoral shaft. Metafem-
oral interior lobe smoothly arcs end to end with nine or ten small serrate teeth on the
distal half only. Protibiae lacking exterior lobe, interior lobe reaching end to end in
a smooth evenly weighted triangle two and a half times as wide as the protibial shaft
(Fig.31B). Meso- and metatibiae simple, lacking lobes.
Measurements of holotype male [mm]. Length of body (including cerci and head,
excluding antennae) 87.0, length/width of head 5.4/4.9, antennae 49.4, pronotum
4.4, mesonotum 5.9, length of tegmina 27.3, length of alae 60.8, greatest width of
abdomen 19.7, profemora 19.5, mesofemora 17.4, metafemora 20.0, protibiae 12.9,
mesotibiae 10.7, metatibiae 14.0.
Etymology. Patronym. Dedicated to David Faulkner, California, United States.
Forensic entomology mentor to RTC and dear friend.
Cryptophyllium icarus gen. et sp. nov.
http://zoobank.org/BFC89435-7C71-4751-B404-CACE95B79616
Figures 5H, 8I, 8J, 9I, 32, 33, 34, 35, 36
Material examined. Holotype ♂: “Coll. I.R.Sc.N.B., Ex breeding Tim Bollens,
2018, Coll. I.R.Sc.N.B., Vietnam, Lam Dong prov., Bidoup-Nui Ba N.P., 12°26’N,
108°30’E, 21-25.vii.2014, Leg. J. Constant and J. Bresseel, GTI Project, I.G.: 32.779”.
Deposited in the Royal Belgian Institute of Natural Sciences (RBINS).
Paratypes: (26 ♀♀, 49 ♂♂, 78 eggs) • 1 ♂: “Coll. I.R.Sc.N.B., Ex breeding
Tim Bollens, 2018, Coll. I.R.Sc.N.B., Vietnam, Lam Dong prov., Bidoup-Nui Ba
N.P., 12°26’N, 108°30’E, 21-25.vii.2014, Leg. J. Constant and J. Bresseel, GTI Pro-
ject, I.G.: 32.779” [vomer dissected] (RBINS) • 5 ♀♀, 4 ♂♂: “Coll. I.R.Sc.N.B., Ex
breeding Tim Bollens, 2018, Coll. I.R.Sc.N.B., Vietnam, Lam Dong prov., Bidoup-
Nui Ba N.P., 12°26’N, 108°30’E, 21-25.vii.2014, Leg. J. Constant and J. Bresseel,
GTI Project, I.G.: 32.779” (RBINS) • 1 ♀, 1 ♂: “Coll. I.R.Sc.N.B., Ex breeding Tim
Bollens, 2018, Coll. I.R.Sc.N.B., Vietnam, Lam Dong prov., Bidoup-Nui Ba N.P.,
12°26’N, 108°30’E, 21-25.vii.2014, Leg. J. Constant and J. Bresseel, GTI Project,
I.G.: 32.779” (VNMN) • 1 ♀, 5 ♂♂: “Coll. I.R.Sc.N.B., Ex breeding Tim Bollens,
2019, Coll. I.R.Sc.N.B., Vietnam, Lam Dong prov., Bidoup-Nui Ba N.P., 12°26’N,
108°30’E, 21-25.vii.2014, Leg. J. Constant and J. Bresseel, GTI Project, I.G.: 32.779”
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
74
Figure 32. Live Cryptophyllium icarus found in July 2014 by Joachim Bresseel (RBINS) and Jérome
Constant (RBINS) in Bidoup Nui Ba N.P., Vietnam. Photographs by Jérome Constant (RBINS) A para-
type female, dorsal closeup B male nymph, paratype C same female as in A but showing the host tree she
was found on D disturbed area along the road where the original stock was found with arrows pointing
out host trees.
(RBINS) • 1 ♀, 1 ♂: “Coll. I.R.Sc.N.B., Ex breeding Bruno Kneubühler, 2017, Coll.
I.R.Sc.N.B., Vietnam, Lam Dong prov., Bidoup-Nui Ba N.P., 12°26’N, 108°30’E,
21-25.vii.2014, Leg. J. Constant and J. Bresseel, GTI Project, I.G.: 32.779” (RBINS)
• 6 ♀♀, subadult ♀, ♂: “Coll. I.R.Sc.N.B., Vietnam, Lam Dong prov., Bidoup-Nui
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 75
Ba N.P., 12°26’N, 108°30’E, 21-25.vii.2014, Leg. J. Constant and J. Bresseel, GTI
Project, I.G.: 32.779” (RBINS) • 1 ♂; “Vietnam, Lam Dong, Bao Loc” (molecular
sample SLT005 within our analysis) (Coll SLT) • 3 ♂♂; “ex Zucht F. Hennemann
2019, Herkunft: Z-Vietnam, Lâm Dông Prov., Dam Rông, Bidoup Nui Ba NP, leg.
Bresseel and Constant VII.2014 [ex coll. FH]” (Coll SLT) • 1 ♀; “VIETNAM, Lam
Dong Prov., Dam Rong, Bidoup-Nui Ba NP, Leg. Bresseel and Constant VII.2014. Ex
breeding F. Hennemann 2015-2017. Ex coll. F. H. Hennemann (Germany); Coll RC
18-226” (Coll RC) • 4 ♂♂; “ex Zucht F. Hennemann 2019, Herkunft: Z-Vietnam,
Lâm Dông Prov., Dam Rông, Bidoup Nui Ba NP, leg. Bresseel and Constant VII.2014
[ex coll. FH]”, Coll RC 20-099–20-102 (Coll RC) • 1 ♂; “VIETNAM, Dak Lak
Prov., May 2018. Coll RC 18-408” (Coll RC) • 1 ♂; “VIETNAM, Lam Dong Prov.,
Dam Rong, Bidoup-Nui Ba NP, Leg. Bresseel and Constant VII.2014. Ex breeding
F. Hennemann 2015-2017. Ex coll. F. H. Hennemann, Germany; Coll RC 18-225”
(Coll RC) • 1 ♂; “VIETNAM, Lam Dong Prov., Bao Loc, October 2016, Coll RC
16-248” (Coll RC) • 1 ♂; “VIETNAM, Lam Dong Prov., Bao Loc, May 2016, Coll
RC 17-264” (Coll RC) • 3 ♂♂; “VIETNAM, Lam Dong Prov., Bao Loc, July 2017”,
Coll RC 17-266, 17-267, 17-268 (Coll RC) • 1 ♂; “VIETNAM, Lam Dong Prov.,
Bao Loc, June 2017. Coll RC 17-265” (Coll RC) • 2 eggs: “VIETNAM, Lam Dong
Prov., Dam Rong, Bidoup-Nui Ba NP, Leg. Bresseel and Constant VII.2014. Ex breed-
ing F. Hennemann 2015-2017. Ex coll. F. H. Hennemann, Germany”; Coll RC 18-
238 and 18-239 (Coll RC) • 20 eggs: “VIETNAM: Lam Dong Prov., Bidoup Nui Ba
N.P., culture from Maxime Ortiz (France), 2018”. Coll RC 18-343–18-362 (Coll RC)
• 7 ♀♀, 1 ♂; “VIETNAM: Lam Dong Prov., Bidoup Nui Ba N.P., bred by Bruno
Kneubhler (Switzerland), circa 2016” (Coll OC) • 3 ♀♀, 7 ♂♂, 28 eggs: “ex Zucht
F. Hennemann 2018, Herkunft: Z-Vietnam, Lâm Dông Prov., Dam Rông, Bidoup
Nui Ba NP, leg. Bresseel and Constant VII.2014” [coll. FH, No’s 0896-1 to 10, E1]
(Coll FH) • 1 ♀, 7 ♂♂, 1 ♂ nymph n5, 28 eggs; “ex Zucht F. Hennemann 2019,
Herkunft: Z-Vietnam, Lâm Dông Prov., Dam Rông, Bidoup Nui Ba NP, leg. Bres-
seel and Constant VII.2014” [coll. FH, No’s 0896-11 to 19, E2], (Coll FH) • 1 ♀:
“ex Zucht F. Hennemann 2020, Herkunft: Z-Vietnam, Lâm Dông Prov., Dam Rông,
Bidoup Nui Ba NP, leg. Bresseel and Constant VII.2014” [coll. FH, No’s 0896-20],
(Coll FH) • 3♀♀, 4♂♂; “Vietnam, Lâm Dông Prov., Dam Rông, Bidoup Nui Ba,
bred by Maxime Ortiz, France, circa 2020” (Coll MO).
Remarks. Cryptophyllium icarus sp. nov. was rst collected in 2014 by Joachim
Bresseel (RBINS) and Jérome Constant (RBINS) from Bidoup Nui Ba N.P., Vietnam.
Several adult females and a subadult male nymph were found on small trees between
the side of the road and next to a high cli with primary forest on top (Fig.32C,D).
is species was successfully brought into the phasmid breeding community which
allowed observation of morphological variation in the females and description of the
adult male, egg, and freshly hatched nymph morphology (Fig. 33). Interestingly, a
signicant change in abdominal shape occurred when this species was reared in cap-
tivity vs. the wild collected females. e wild collected paratype females (Fig. 32A)
has a slenderer abdomen, with segments VI and VII notably converging towards the
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
76
Figure 33. Live adult Cryptophyllium icarus bred and photographed by Bruno Kneubühler (Switzerland)
A female B male C lateral view of female with tegmina held up to expose the dwarfed alae underneath.
posterior with only a slight bend near the posterior of segment VII, vs. individuals
reared in captivity (Fig. 33A) which instead develop a rather boxy abdominal shape,
with segments VI and the majority of VII parallel-sided and ending in a distinct lobe
on the posterior of VII. Interestingly all wild caught females were slender and all bred
females have been boxy; at this time we are unsure what mechanism is promoting this
polymorphism within this species. Hopefully additional eld work in the region will
reveal additional observations to review.
An additional unique feature for this species is the coloration of the freshly hatched
nymphs, which do not match well with the coloration of other known congenerics.
is could be due to our lack of knowledge as freshly hatched nymph coloration is
known only for nine species (Fig. 9). Molecularly, we found Cryptophyllium icarus
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 77
sp. nov. to be sister species to Cryptophyllium limogesi sp. nov. whose freshly hatched
nymph morphology we do not know. It is possible that these may share similar colora-
tion, but at the present Cryptophyllium icarus sp. nov. with their solid red abdomen and
dark extremities (Fig. 9I) appear quite unique within the Cryptophyllium gen. nov. as
presently known.
Dierentiation. Female morphology is supercially similar to Cryptophyllium
daparo sp. nov. and Cryptophyllium chrisangi comb. nov. due to the tapered abdominal
shape, obtusely rounded profemoral exterior lobes, and similarly shaped and textured
thorax. Cryptophyllium icarus sp. nov. can be dierentiated from both species by the
highly reduced alae not reaching abdominal segment II (Fig. 33C), vs. the other species
which both have well-developed alae.
Males with their thorax shape and spination, the spade-shaped abdomen, shorter
tegmina, and exterior profemoral lobe which is thinner than the interior, are morpho-
logically similar to Cryptophyllium bankoi sp. nov. and Cryptophyllium rarum comb.
nov. males. From both species, Cryptophyllium icarus sp. nov. can be dierentiated by
the width of the profemoral lobe which in Cryptophyllium icarus sp. nov. is only ca.
2× as wide as the profemoral shaft, vs. the other species which are at least 2½ or 3× as
wide as the profemoral shaft. Additionally, males of Cryptophyllium rarum comb. nov.
are notably larger than the largest Cryptophyllium icarus sp. nov. males.
Distribution. Only presently known from southern Vietnam, from the provinces
of Lam Dong and Dak Lak.
Description. Female. Coloration. Coloration is variable, as when it was bred in
captivity females were almost uniformly pale green, with only slightly orange/tan areas
along the profemoral lobes, antennae, eyes, thorax, and terminal abdominal margins
(Fig. 33A). In contrast, the paratype female when she was found in the wild was quite
colorful with a vibrant green body with highlights of red throughout the antennae,
legs, thorax, tegmina, and abdomen (Fig. 32A), much darker and more plentiful than
on captive bred specimens.
Morphology. Head. Head capsule longer than wide, vertex relatively smooth, with
the only notable feature the posteromedial tubercle which is not notably broad but
is signicantly raised above the head capsule (Fig. 34E). Frontal convexity broad and
stout, with a lumpy surface, and with several setae throughout (Fig. 34E). Compound
eyes slightly protruding from the head capsule, not notably large, only taking up slight-
ly < 1/4 of the length of the lateral head capsule margins (Fig. 34E). Ocelli absent. An-
tennal elds slightly wider than the rst antennomere (Fig. 34C). Antennae. Antennae
consisting of nine segments, with the terminal segment about as long as 2½× the pre-
ceding segments lengths (Fig. 34C). Antennomeres I–VIII sparsely marked with small
transparent setae, the terminal antennomere and the distal margin of antennomere
VIII has darker, shorter, and denser setae than the other segments (Fig. 34C). orax.
Pronotum with a slightly concave anterior margin and slightly convex lateral margins,
which converge to a straight posterior margin that is half the width of the anterior mar-
gin (Fig. 34E). e pronotum surface and moderately formed pronotum rims are only
slightly lumpy, lacking signicant granulation, with only a prominent pit in the center,
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
78
and slight furrows anterior, posterior, and lateral to the pit (Fig.34E). Prosternum with
moderate granulation, mesosternum anterior half and lateral margins with moderate
granulation (Fig. 34B). Metasternum relatively smooth, lacking notable nodes. Pres-
cutum slightly longer than wide, lateral rims with four larger nodes and four or ve
nodes interspersed throughout (Fig. 34E). Prescutum anterior rim prominent but not
strongly protruding, surface marked throughout with irregular granulation, no promi-
nent singular sagittal spine present (Fig. 34F). Prescutum surface covered densely by
small tubercles and numerous nodes, with those along the sagittal plane the largest
(Fig. 34E). Mesopleura beginning slightly posterior to the anterior margin of the pres-
cutum and evenly diverging; lateral margin with ve larger tubercles, and ve or six
smaller tubercles interspersed unevenly throughout (Fig. 34E). Face of the mesopleura
slightly wrinkled, with a distinct pit near the anterior margin and one near the center
(Fig. 34E). Wings. Tegmina long, reaching ½ way through abdominal segment VII.
e subcosta (Sc) is the rst vein in the forewing and runs parallel with the tegmina
lateral margin for the rst half of the vein, then bends gently and runs to the to the
lateral margin of the wing where it terminates ca. ⅓ through the length. e radius (R)
spans the central portion of the forewing with two subparallel branched veins; radius 1
(R1) terminates slightly proximal to the midline, and the radial sector (Rs) terminates
ca. ⅔ of the way through the wing length. ere is a weak continuation of the radius
following the prominent Rs branching which continues on as a short and thinner
R–M crossvein that does not solidly connect the two veins as it reaches the media. e
media (M) is simply bifurcate with both the media anterior (MA) and media posterior
(MP) terminating on the posterior ¼ of the wing. e cubitus (Cu) is also bifurcate,
branching near the posterior ⅕ of the wing into the cubitus anterior (CuA) and cu-
bitus posterior (CuP) which both terminate at or very near the wing posterior apex.
e rst anal vein (1A) is simple and fuses with the cubitus early on, halfway between
the branching of the R1 and the Rs. Alae are highly reduced, not reaching the anterior
margin of abdominal segment II. Abdomen. Abdominal segments II through the an-
terior half of IV diverge uniformly towards the posterior. e posterior half of IV and
segment V are parallel-sided and are the widest portion of the abdomen. Segment VI
and VII are variable depending on the environmental conditions, with wild collected
females (such as in the paratype; Fig. 32A) having segments VI and VII strongly con-
verging and VII ending with a small bulge on the posterior margin with a nal width
about the same as the anterior margin of segment VIII, vs. captive bred individuals
generally having VI and VII subparallel and ending with VII having a distinct lobe
which is notably wider than the anterior margin of segment VIII (Fig. 33A). Segments
VIII–X converge uniformly to the rounded apex. Genitalia. Subgenital plate starts at
the anterior margin of segment VIII, is long and narrow reaching ca. ½ onto segment
X (Fig. 34H). Gonapophyses VIII are long and signicantly broad, each about as wide
as the subgenital plate projection, with their tips reaching the apex of segment X, gona-
pophyses IX are smaller and slender, hidden below the gonapophyses VIII (Fig. 34H).
Cerci at, not strongly cupped, with a heavily granular surface and a setae throughout
(Fig. 34H). Legs. Profemoral exterior lobes notably wider than the interior lobe with
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 79
a rounded obtuse angle (Fig. 34D). Exterior lobe margin is not marked by teeth and
is instead rather smooth or at most slightly granular (Fig. 34D). Profemoral interior
lobe ca. 2× as wide as the greatest width of the profemoral shaft, with an obtuse angle,
and marked with ve prominent teeth arranged in a two-one-two pattern with large
looping gaps between the teeth (Fig. 34D). Mesofemoral exterior lobe arcs from end
to end in a slightly bent lobe slightly weighted to the distal ½ and marked with two or
three small serrate teeth on the distal ½ only. Interior and exterior lobes are of a similar
width, or the exterior is slightly wider. Mesofemoral interior lobe arcs smoothly end to
end, is marked with six or seven serrate teeth only on the distal half of the arc, and is
about as wide as the mesofemoral shaft. Metafemoral interior lobe arcs end to end, but
is slightly wider on the distal half, and has six or seven serrate teeth on the distal half of
the lobe only. Metafemoral exterior lobe is thin and smooth, hugging the metafemoral
shaft and lacks dentation. Protibiae interior lobe spans the entire length of the proti-
biae and is at ca. 2× as wide as the protibial shaft. e lobe is distinctly triangular with
the broadest point distal to the midline (Fig. 34D). Pro-, meso-, and meta- tibiae lack-
ing exterior lobes; meso-, and meta- tibiae lack interior lobes as well.
Measurements of paratype females [mm]. Length of body (including cerci and
head, excluding antennae) 81.8–94.1, length/width of head 8.0–9.7/6.4–7.4, anten-
nae 3.3–4.2, pronotum 4.6–5.7, mesonotum 6.7–9.1, length of tegmina 45.4–57.5,
length of alae 6.0–7.2, greatest width of abdomen 30.7–34.4, profemora 17.9–21.6,
mesofemora 13.8–15.9, metafemora 16.2–18.6, protibiae 11.1–11.8, mesotibiae
10.1–10.7, metatibiae 13.6–13.7.
Male. Coloration. Coloration description based on live captive reared males
(Fig.33B) and the male nymphs found in the wild (Fig. 32B). Captive males are gen-
erally mostly pale green throughout with the margins of the legs, thorax, and abdomen
a reddish brown, and slightly transparent eye spots on abdominal segment V. e wild
caught nymphs had these areas typically darker, more dark red than reddish brown and
dark eyespots on abdominal segment V.
Morphology. Head. Head capsule about as long as wide, with a vertex that is weak-
ly granular; posteromedial tubercle small but notable and slightly raised above the head
capsule (Fig. 35D). Frontal convexity stout with a few short setae near the apex. Com-
pound eyes large and bulbous, occupying ca. ⅖ of the head capsule lateral margins and
signicantly protruding from the head capsule (Fig. 35D). ere are three well-devel-
oped ocelli between and slightly posterior to the compound eyes. Antennae (including
the scapus and pedicellus) consists of 23 segments. e scapus and pedicellus are bare,
all other segments are covered in dense, thin, pale setae that are as long as or longer than
the antennae segment is wide. e terminal three segments have shorter darker setae.
orax. Pronotum anterior margin is distinctly concave and lateral margins are slightly
convex and converge to a straight posterior margin that is slightly > ½ width of the
anterior rim (Fig. 35D). Anterior and lateral margins of the pronotum have moderate
rims and the posterior margin lacks a rim (Fig. 35D). Face of the pronotum is slightly
lumpy, has a distinct sagittal furrow, a pit just posterior to the center, a moderate per-
pendicular furrow just anterior to the central pit, and has a distinct pit on each side near
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80
Figure 34. Cryptophyllium icarus gen. et sp. nov. female paratype, photographs by Jérôme Constant
(RBINS) A habitus, dorsal B habitus, ventral C antennae and anterior of head capsule D profemoral
and protibial lobes E head and thorax details dorsal F head and thorax details lateral G genitalia dorsal
H genitalia ventral.
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 81
Figure 35. Cryptophyllium icarus gen. et sp. nov. male holotype, photographs by Jérôme Constant
(RBINS) A habitus, dorsal B habitus, ventral C profemoral and protibial lobes D head and thorax details
dorsal E head and thorax details lateral F genitalia dorsal G genitalia ventral.
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
82
the anterior margin (Fig. 35D). e prosternum surface is slightly granular. e mesos-
ternum surface is marked densely with prominent nodes, with the largest along the
sagittal plane and more prominent on the anterior margin, posterior margin with less
prominent and slightly smaller nodes. Prescutum slightly longer than wide, with lateral
margins slightly converging to the posterior which is ca. ¾ the width of the anterior rim
(Fig. 35D). Lateral rims with nine or ten nodes of slightly varying size, none very large
or prominent, but each marked with a single sti seta protruding from the tip. e
surface of the prescutum is notably granulose along the sagittal plane with lateral sur-
faces rather smooth (Fig. 35D). Prescutum anterior rim slightly granulose with no dis-
tinct central tubercle (Fig. 35E). Mesopleura narrow, only gradually diverging from the
anterior to the posterior (Fig. 35D). Lateral margin granulose throughout, with only
four or ve slightly larger than the rest, but not signicantly larger. e largest nodes
along the mesopleura have a singular seta protruding from them like those on the pres-
cutum margins. Face of the mesopleura smooth but slightly wrinkled and with two
faint pits, one on the anterior margin and one near the middle of the mesopleura.
Wings. Tegmina short, only reaching the anterior margin of abdominal segment III.
Tegmina wing venation: the subcosta (Sc) is the rst vein and terminates the earliest, ca.
⅖ of the way through the overall tegmina length. e radius (R) spans the entire length
of the tegmina with the rst radius (R1) branching ca. ⅓ of the way through the wing
length and terminating just posterior to the middle of the wing, the second radius (R2)
branches near the distal ⅓ of the wing, and then the radial sector (Rs) runs straight to
the tegmina apex and terminates. e media (M) spans the entire length of the tegmina,
terminating at the wing apex as the media anterior (MA) with the rst media posterior
(MP1) beginning and terminating near the tegmina mid length followed by the second
media posterior (MP2) which begins ca. ⅔ of the way through the tegmina length and
terminates near the posterior quarter of the wing. e cubitus (Cu) runs through the
wing surface angled until it meets the margin ca. ⅓ of the way through the tegmina
length and then runs along the margin as the two media posterior veins then meet it
and fuse and the cubitus continues to run nearly to the wing apex. e rst anal (1A)
vein runs subparallel to the cubitus until it meets it slightly > ⅓ of the way through the
tegmina length and fuses with it. Alae well-developed in an oval fan conguration,
reaching to the anterior margin of abdominal segment IX or halfway through it. Alae
wing venation: the costa (C) is present along the entire foremargin giving stability to the
wing. e subcosta (Sc) spans ca. ⅔ of the wing length and is mostly fused with the
radius in the beginning but terminates when it meets the costa. e radius (R) spans the
entire wing and branches ca. ⅖ of the way through into the radius 1 (R1) and radial
sector (Rs) which run slightly diverging for the rst ⅓ of their length, parallel for the
central portion until the terminal quarter where they converge and terminate on the
wing margin near each other but not touching. e media (M) branches early, ca. ⅙ of
the way through the wing into the media anterior (MA) and the media posterior (MP)
which run parallel with each other until the distal ⅙ of the wing where the media pos-
terior fuses with the media anterior which then run fused together to the wing margin.
e cubitus (Cu) runs unbranched and terminates at the wing apex. Of the anterior
anal veins, the rst anterior anal (1AA) fuses with the cubitus near the point where the
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 83
media branches into the media anterior and media posterior and then the rst anterior
anal branches from the cubitus ⅔ of the way through the wing length where it uni-
formly diverges from the cubitus until it terminates at the wing margin. e anterior
anal veins two–seven (2AA–7AA) have a common origin and run unbranched in a fold-
ing fan pattern of relatively uniform spacing to the wing margin. e posterior anal
veins (1PA–7PA) share a common origin separate from the anterior anal veins and run
unbranched to the wing margin with slightly thinner spacing than the anterior anal
veins. Abdomen. Segment II, parallel-sided, segment III and the anterior half of IV di-
verging, the posterior ½ of IV through segment V either parallel-sided or slightly di-
verging, VI through the apex converging to a blunt rounded apex. e margins of seg-
ments VIII–X have a line of setae along them (Fig. 35F). Genitalia. Poculum broad and
rounded, ending in a rounded apex that passes beyond the anterior margin of segment
X (Fig. 35G). Cerci long and slender, with ca. ½ of their length extending out from
under the anal abdominal segment. e cerci are slightly cupped, with a granulose sur-
face and numerous short setae throughout (Fig. 35F). Vomer broad and stout with
rounded sides converging to the apex which is armed with two upwards turning hooks,
one at the apex which is larger and one lateral to it which is slightly smaller (Fig. 5H).
Legs. Profemoral exterior lobe a rounded arc without a distinct angle, slightly thinner
than the interior lobe (at its widest slightly > 2× the greatest width of the profemoral
shaft), and with the distal half marked by three or four small but sharp anteriorly point-
ing teeth (Fig. 35). Profemoral interior lobe roundly triangular, at its widest ca. 2½× as
wide as the profemoral shaft at its widest. e profemoral interior lobe is generally
marked with ve, serrate, anteriorly pointing teeth arranged in a two-one-two pattern,
with shallow looping gaps between them, and occasionally marked with an extra tooth
within the set (like as can be seen in Fig. 35C). Mesofemoral exterior lobe arcs end to
end but with a more prominent bend near the distal ⅓ of the lobe which is marked with
two or three small serrate teeth on the distal ⅓ only, with the proximal portion of the
lobe smooth. e mesofemoral interior lobe at its widest is approximately the same
width as the exterior lobe, but the proximal half is slightly wider than the proximal half
of the exterior lobe. e distal half of the mesofemoral interior lobe is marked with six
or seven small serrate teeth and the proximal half is rather smooth. Metafemoral exte-
rior lobe lacks dentation and has a straight margin along the metafemoral shaft.
Metafemoral interior lobe smoothly arcs end to end with six or seven small serrate teeth
on the distal ½ only. Protibiae lacking exterior lobe, interior lobe reaching end to end
in a smooth triangle ca. 3× as wide as the protibial shaft, with the widest point just
distal to the midline, and all margins notably marked with short setae throughout
(Fig.35C). Mesotibiae and metatibiae simple, lacking lobes completely.
Measurements of holotype male [mm]. Length of body (including cerci and head,
excluding antennae) 58.6, length/width of head 3.7/3.3, antennae 32.5, pronotum
2.8, mesonotum 4.5, length of tegmina 16.1, length of alae 42.0, greatest width of
abdomen 15.8, profemora 12.0, mesofemora 9.5, metafemora 11.4, protibiae 7.5,
mesotibiae 6.3, metatibiae 8.2.
Measurements of paratype males [mm]. Length of body (including cerci and
head, excluding antennae) 56.0–69.4, length/width of head 2.9–4.6/2.9–4.0, anten-
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
84
nae 34.5–43.9, pronotum 2.5–3.0, mesonotum 3.8–4.8, length of tegmina 16.4–19.8,
length of alae 40.5–49.9, greatest width of abdomen 14.4–18.3, profemora 11.7–14.8,
mesofemora 9.5–12.3, metafemora 11.0–13.6, protibiae 7.7–9.9, mesotibiae 6.5–8.0,
metatibiae 8.5–10.4.
Description of egg (Fig. 36). e lateral surfaces are at but with the posterior
half slightly wider than the anterior half. e dorsal surface is slightly convex, which
gives the margin a slight undulating appearance when viewed from the lateral aspect as
the middle is thinner than either end of the egg. When viewed from the lateral aspect;
the ventral margin is also not straight but is instead with the posterior slightly protrud-
ing more than the anterior. All surfaces have numerous small sized pits throughout
with short moss-like pinnae interspersed throughout the capsules surfaces with those
on the margins and those on the dorsal surface slightly more prominent. Dorsal sur-
face with irregular medium sized pitting and moss-like pinnae around the micropylar
plate. Micropylar plate long, ca. 7/8 of the overall dorsal surface length, with the wid-
est portion around the micropylar cup. Micropylar plate nearly symmetrical with the
anterior and posterior thin and the area around the micropylar cup the widest point.
Micropylar cup of moderate size and placed just slightly posterior to the micropylar
plate midline. Operculum slightly ovular, with the outer margin with a distinct row of
moss-like pinnae surrounding the operculum. Operculum is roundly raised with the
height ca. ½ the operculum width. e overall color is light brown, with the moss-like
pinnae sometimes slightly lighter in color.
Measurements including the extended pinnae [mm]. Length (including opercu-
lum) 2.2, maximum width of capsule when viewed from lateral aspect 1.3 mm, length
of micropylar plate 1.1 mm.
Newly hatched nymphs. (Fig. 9I). e body is made up of two general blocks of
color, the legs, head, pronotum, and mesonotum are primarily chocolate brown and the
mesonotum and abdomen are burnt red in color. Basitarsi are white and the remaining
tarsal segments are burnt reddish. e tibiae only have interior lobes which smoothly
span the full length and lack exterior lobes. e tibiae interior lobes are brown with two
white patches on the proximal half only, the proximal most is notably larger. Interior
and exterior femoral lobes are all about the same width, are smoothly arcing, and all have
minimal serration. On the profemoral interior lobe there is a notable whitish patch, the
exterior lobe and profemoral shaft itself are devoid of prominent white markings. e
meso- and metafemoral interior lobes are similar in that they have a small white patch
right at the proximal end, and then another white marking ca. ⅖ of the way through
the length. e exterior meso- and metafemoral lobes only have one white patch located
on the proximal ⅓, but notably wider than the interior lobe white patch. e meso-
and metafemoral shafts lack white coloration. e abdomen is slender with segment II
through the anterior ½ of IV diverging and the posterior ½ of IV–X converging.
Etymology. Noun, from Greek mythology. Named for the tragic story of Icarus,
son of Daedalus. During their escape from the island of Crete, Icarus ew too close
to the sun and melted the wax wings his father built. We felt it was tting that this
mythological name is shared with this species that lacks the hindwings within the for-
mer celebicum species group (characterized by females with well-developed alae).
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 85
Figure 36. Cryptophyllium icarus gen. et sp. nov. egg, RBINS collection, photographs by Jérôme Con-
stant A lateral view B dorso-lateral view C dorsal view D opercular (anterior) view E posterior view
Fventral view.
Cryptophyllium khmer gen. et sp. nov.
http://zoobank.org/118A6B83-0408-4373-8C02-3705528BB026
Figures 5B, 5D, 6D, 9C, 37, 38, 39, 40, 41
Material examined. Holotype ♂: “Coll. I.R.Sc.N.B., collected as nymph, Coll.
I.R.Sc.N.B., Cambodia, Koh Kong prov., Tatai, 11°35’13”N 103°05’50”E,
9-19.x.2016, day collecting, GTI Project, Leg. J. Constant and J. Bresseel, I.G.: 33.345
(RBINS-PHYLLIUM DNA sample 0002)” [vomer dissected], deposited in RBINS.
Paratypes (9 ♀♀, 2 ♂♂): • “Coll. I.R.Sc.N.B., collected as nymph, Coll.
I.R.Sc.N.B., Cambodia, Koh Kong prov., Tatai, 11°35’13”N 103°05’50”E,
9-19.x.2016, day collecting, GTI Project, Leg. J. Constant and J. Bresseel, I.G.:
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
86
33.345, RBINS-PHYLLIUM DNA sample 0001” (RBINS) • 3 ♀♀, 1 ♂: “Coll.
I.R.Sc.N.B., Ex breeding Tim Bollens, 2018, Coll. I.R.Sc.N.B., Cambodia, Koh Kong
prov., Tatai, 11°35’13”N 103°05’50”E, 9-19.x.2016, day collecting, GTI Project, Leg.
J. Constant and J. Bresseel, I.G.: 33.345” • 2 ♂♂: “Coll. I.R.Sc.N.B., Ex breeding
Tim Bollens, 2018, Coll. I.R.Sc.N.B., Cambodia, Koh Kong prov., Tatai, 11°35’13”N
103°05’50”E, 9-19.x.2016, day collecting, GTI Project, Leg. J. Constant and J.
Bresseel, I.G.: 33.345; ex breeding Tim Bollens, 2018” • 2 ♀♀: “Coll. I.R.Sc.N.B.,
CAMBODIA, Siem Reap Prov., Phnom Kulen N.P., Forest near Preah om, 26-27-
VII-2006, Leg K. Smets, Y. Oul and D. Jump.” (1 ♀: RBINS; 1 ♀: RUPP) • 1 ♂:
“Cambodia, Siem Reap; Kbal Spean, 13°40.858’N 104°01.111’E, 122 m, 6-jul-2015,
Hap, Sour, Phauk, Khearn, Chhum, Ly, Lom, Heang, Hok, CA0028, Lighttrap in
the forest with canopy cover.” (RUPP) • 3 ♀, 1 ♂: “Coll. I.R.Sc.N.B., Ex breeding
Tim Bollens, 2019, Coll. I.R.Sc.N.B., Cambodia, Koh Kong prov., Tatai, 11°35’13”N
103°05’50”E, 9-19.x.2016, day collecting, GTI Project, Leg. J. Constant and J.
Bresseel, I.G.: 33.345; ex breeding Tim Bollens, 2019” (RBINS) • 1 ♀, 1 ♂: “Coll.
I.R.Sc.N.B., Ex breeding Tim Bollens, 2019, Coll. I.R.Sc.N.B., Cambodia, Koh Kong
prov., Tatai, 11°35’13”N 103°05’50”E, 9-19.x.2016, day collecting, GTI Project, Leg.
J. Constant and J. Bresseel, I.G.: 33.345; ex breeding Tim Bollens, 2018” (Coll RC).
Remarks. When this species was rst reviewed morphologically, it was assumed to
be an additional distribution record of Cryptophyllium westwoodii comb. nov., which
is known from a relatively expansive range (Fig. 2). However, our molecular analysis
revealed that the Tatai and Siem Reap (Cambodia) populations instead represented an
undescribed species distinct from Cryptophyllium westwoodii comb. nov. e recogni-
tion of this cryptic species from Cambodia, leaves many observational records from
Laos, ailand, and Cambodia without conrmed identication (represented by the
bi-colored circles in our distribution map noting a record which, due to the lack of
molecular conrmation, could represent either species; Fig. 2).
During GTI expeditions several nymphs ranging from L1 to subadult were col-
lected on multiple closely situated guava trees behind a house near the start of the trail
leading to Tatai falls. Nymphs were successfully reared to adulthood by Tim Bollens
(Belgium). Strangely locals had never noticed the insects before due to their excellent
camouage (Fig. 37).
Interestingly, in 2006, an attempt was made to describe a Cryptophyllium west-
woodii comb. nov. like species from Rayong, ailand as ‘Phyllium rayongii’ (Sor-
pongpaisal and anasinchayakul 2006). Cumming and Le Tirant (2020) note how-
ever that this name is a nomen nudum and therefore unavailable according to ICZN
Article 16.4.1. (ICZN 1999). With this population rather geographically close to
the type locality of Cryptophyllium khmer sp. nov. it is entirely possible that ‘Phyllium
rayongii’ may have been intended to represent a valid population, but with the lack
of a holotype specimen to dene this species it was never conrmed and is now a
moot point.
Due to the cryptic nature of this new species, we hope that eorts will be under-
taken in the future to molecularly sample from throughout ailand, Myanmar, Laos,
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 87
Figure 37. Live Cryptophyllium khmer gen. et sp. nov. type material found by the joint eort of the
RBINS, RUPP, and VNMN teams in Tatai, Cambodia in October 2016, all found as nymphs. Indi-
viduals in photographs A–H were found feeding on Guava trees (Psidium guajava) and the specimen in
photograph I was found feeding on an unidentied species. Photographs by Jérôme Constant (RBINS)
A female B Hong ai Pham (left) and Sisonila Kangsun (right), participants of the GTI project, photo-
graphing the leaf insects, which are very popular in Asia C male nymph D female nymph E male nymph
F several nymphs G nymph H nymph I female nymph on unidentied host species.
and Cambodia to determine with more clarity the geographic distributions where
Cryptophyllium khmer sp. nov. and Cryptophyllium westwoodii comb. nov. occur.
Dierentiation. Morphological dierentiation of this species has proven to be
dicult, with the only clear and consistent dierences being ascertained through mo-
lecular analysis (Fig. 4).
Females are most morphologically similar to Cryptophyllium westwoodii
comb. nov., Cryptophyllium bollensi sp. nov., Cryptophyllium phami sp. nov., and
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
88
Cryptophyllium nuichuaense sp. nov. females based on the general shape of the
abdomen, lobes of the legs, and the thorax. e later three species can be dierentiated
by their shorter alae reaching to abdominal segments II or III vs. Cryptophyllium khmer
sp. nov. which has long alae reaching onto abdominal segment VI. We have yet to
identify a reliable morphological feature between Cryptophyllium khmer sp. nov. and
Cryptophyllium westwoodii comb. nov. as both species have long alae and at least for
Cryptophyllium westwoodii comb. nov. there is signicant intraspecic variation which
often encompasses the range of Cryptophyllium khmer sp. nov. female variation.
Males are most morphologically similar to Cryptophyllium westwoodii comb. nov.,
Cryptophyllium chrisangi comb. nov., Cryptophyllium bollensi sp. nov., and Cryptophyl-
lium phami sp. nov. based on features of the thorax, tegmina, and lobes of the legs.
Cryptophyllium khmer sp. nov. males can be dierentiated from the rst two species by
the general shape of the abdomen as it is thinly elliptical with a maximum width only
30–34% the abdominal length in Cryptophyllium westwoodii comb. nov. and Cryp-
tophyllium chrisangi comb. nov. vs. broadly elliptical or broadly spade-shaped with a
maximum width ca. 38–45% the abdominal length in Cryptophyllium khmer sp. nov.,
Cryptophyllium bollensi sp. nov., and Cryptophyllium phami sp. nov. males. Unfortu-
nately, due to intraspecic variation within Cryptophyllium khmer sp. nov., Crypto-
phyllium bollensi sp. nov., and Cryptophyllium phami sp. nov. we could not identify a
reliable morphological feature for dierentiation males based on morphology alone.
is diculty for dierentiating a single sex alone emphasizes the importance of
captive rearing of specimens to reveal the informative set of female, male, and egg
morphology, and of course the importance of molecular comparison.
Distribution. At present only conrmed from two Cambodian provinces, Koh
Kong Province (Tatai) and Siem Reap Province (Kbai Spean and Phnom Kulen N.P.,
Forest Near Prean om). It is likely that other nearby localities may also represent
this species, but due to a lack of molecular data we cannot at this time conrm them.
Description. Female. Coloration. Coloration description is based upon photo-
graphs of living individuals (Fig. 38A, B) reared by Tim Bollens (Belgium). Overall
coloration pale mint green with variable slight highlighting of orange or tan coloration
throughout. Compound eyes are slightly more yellow with tan highlights. Antennae
are tan. e prescutum and mesopleura are reddish tan with pale cream granulation
throughout. roughout the head, legs, and body there is slight speckling as granula-
tion is slightly paler in color than the surface it is found on. In lighter individuals, the
venation of the tegmina is pale yellow to pale mint green (Fig. 38A) and in darker
individuals the venation is yellow with highlights of orange interspersed throughout
(Fig. 38B). Darker individuals also have variable reddish patches throughout the lobes
of the legs and slightly darker coloration on the abdomen.
Morphology. Head. Head capsule slightly longer than wide, vertex with granula-
tion throughout the surface, none as prominent as the posteromedial tubercle which is
not notably wide but is distinctly taller than any other nodes on the head (Fig. 39E).
Frontal convexity stout, marked throughout with slight granulation and several short
setae. Compound eyes slightly protruding from the head capsule, but are signicantly
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 89
Figure 38. Live Cryptophyllium khmer gen. et sp. nov. type material bred and photographed by Tim Bolllens
(Belgium), dorsal views A paler form female (paratype) B darker form female (paratype) C male (paratype).
large, taking up slightly < ⅓ of the head capsule margins (Fig. 39E). Ocelli absent.
Antennal elds slightly wider than the rst antennomere. Antennae. Antennae consist-
ing of nine segments, with the terminal segment slightly longer than the preceding
two segments’ lengths combined (Fig. 39C). Antennomeres I–VIII sparsely marked
with small transparent setae, the terminal antennomere is covered densely in slightly
shorter setae. orax. Pronotum slightly wider than long, with gently concave anterior
margin and slightly convex lateral margins, which converge to a slightly convex pos-
terior margin that is half the width of the anterior margin (Fig. 39E). e pronotum
surface is marked with granulation throughout, a prominent pit in the center, and
slight furrows anterior and lateral to the pit (Fig. 39E). e pronotum has a prominent
anterior rim and weakly formed lateral and posterior rims (Fig. 39E). Prosternum and
the anterior half of the mesosternum are marked with stout and numerous nodes, with
the remainder of the mesosternum and the metasternum lacking prominent nodes
(Fig.39B). Prescutum about as long as wide with lateral rims with 11 or 12 lumpy tu-
bercles ranging in size from small to medium with granulation present throughout the
length giving the margins a tough textured appearance (Fig. 39E). Prescutum anterior
rim not strongly protruding and marked with a granular surface (Fig. 39F). Prescutum
surface with 14 or 15 distinct nodes predominantly along the sagittal plane, with those
on the anterior half slightly larger than the rest (Fig. 39E). Mesopleura are narrow and
parallel on the anterior ⅓, and then bend distinctly and diverge uniformly throughout
their length; lateral margin with 13–16 small to medium lumpy tubercles, of which
three or four are slightly larger than the rest, but most are small and variable in shape,
giving the margin a rough textured appearance (Fig. 39E). Face of the mesopleura with
granulation along the margin, with the remainder of the surface relatively smooth or
with slight wrinkles. e surface of the mesopleura also has two distinct pits, one near
the anterior ⅓ where the mesopleura bend, and one near the posterior ⅓ (Fig. 39E).
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
90
Figure 39. Cryptophyllium khmer gen. et sp. nov. paratype female, molecular sample RBINS01 in our anal-
ysis, from Tatai, Cambodia, photographs by Jérôme Constant (RBINS) A habitus, dorsal B habitus, ventral
C details of the antennae, dorsal D pro- tibial and femoral lobes, dorsal E details of the antennae, head, and
thorax, dorsal F details of the antennae, head, and thorax, lateral G terminalia, dorsal H genitalia, ventral.
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 91
Wings. Tegmina long, reaching onto abdominal segment VIII. e subcosta (Sc) is the
rst vein in the forewing and runs parallel with the wing for the rst half of its length,
and then bends towards the wing margin for the second half, terminating ca. ⅓ of the
way through the wing length. e radius (R) spans the central portion of the tegmina
with two subparallel branched veins. e rst radius (R1) branches ca. ½ through the
radius length and terminates ca. ⅖ of the way through the wing length. e radial sec-
tor (Rs) branches from the end of the radius and runs angled to the wing margin where
it terminates near the posterior ⅓ of the wing length. ere is a weak continuation
of the radius following the prominent radial sector branching which continues on as
a short and thin radius to media crossvein (R–M). e media (M) is simply bifurcate
with both the media anterior (MA) and media posterior (MP) terminating close to
the posterior ⅕ of the wing. e cubitus (Cu) runs throughout the entire wing length
simply, and then near the posterior ⅕ of the wing splits into the cubitus anterior (CuA)
and cubitus posterior (CuP) which both terminate at or very near the wing posterior
apex. e rst anal vein (1A) is simple and fuses with the cubitus early on, at around
the midline between the rst radial branching and the radial sector branching. Alae
well-developed, reaching abdominal segment VI. Abdomen. Abdominal segments II
through the anterior half of IV diverging, the posterior half of IV through the anterior
half of VII parallel-sided (giving the abdomen a boxy appearance), the remainder of
VII smoothly rounded and converging to the apex with segments VIII–X. Genitalia.
Subgenital plate starts at the anterior margin of segment VIII, is moderately broad, and
extends ½ to ¾ of the way onto segment X, ending in a ne point (Fig. 39H). Gona-
pophyses VIII are long and moderately broad, exceeding the apex of the abdomen with
the tips slightly longer than the cerci, gonapophyses IX are thinner and shorter, hidden
below gonapophyses VIII (Fig. 39H). Cerci at, not strongly cupped, with a nely
granular surface and moderately marked with a few short setae. Legs. Profemoral exte-
rior lobe broad and smoothly rounded, ca. 1½ to ca. 2× wider than the interior lobe
(Fig. 39D). Margin of the profemoral exterior lobe with 10–12 small weakly formed
teeth throughout the length (Fig. 39D). Profemoral interior lobe obtusely angled and
typically marked with ve teeth arranged in a two-one-two pattern with looping gaps
between them, but occasionally individuals can have doubly serrate teeth or an extra
small tooth between sets (Fig. 39D). Mesofemoral exterior lobe arcs from end to end
but is weighted towards the distal half with a detectable bend and marked with four
or ve rounded teeth distributed on the distal half only. Interior and exterior lobes of
a similar width. Mesofemoral interior lobe arcs end to end smoothly with ve or six
small serrate teeth only on the distal half of the arc which is slightly wider than the
proximal half of the arc. Metafemoral interior lobe arcs end to end and has ve or six
serrate teeth on the distal half of the lobe which is slightly wider than the proximal half.
Metafemoral exterior lobe is thin and smooth, hugging the metafemoral shaft and lacks
notable teeth but the distal ⅓ can be slightly granular. Protibial interior lobe spans the
entire length of the protibiae and is ca. 2½ the width of the protibiae shaft itself. e
lobe is roundly triangular and is slightly wider on the distal half. Protibiae lacking a
distinct exterior lobe. Mesotibiae and metatibiae lacking exterior and interior lobes.
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
92
Measurements of paratype females [mm] (from Tatai, Cambodia). Length of
body (including cerci and head, excluding antennae) 83.3–90.0, length/width of head
8.4–8.7/6.6–7.1, antennae 4.1–4.6, pronotum 5.6–6.0, mesonotum 7.6–7.8, length
of tegmina 52.8-53.6, length of alae 42.6 (only measured on one specimen, the others
have the alae covered by the tegmina), greatest width of abdomen 31.3–36.2, profem-
ora 19.1–21.4, mesofemora 15.1–15.4, metafemora 18.7–19.6, protibiae 12.5–12.6,
mesotibiae 11.4–11.6, metatibiae 14.7–15.0.
Male. Coloration. Coloration description based on images of live males bred by
Tim Bollens (Belgium). Overall coloration pale mint green throughout with highlight-
ing of tan to orange (Fig. 39C). e areas most often with the orange highlighting are
the tips of the antennae, margins of the lobes of the legs, the thorax, and the margins
of the abdomen. Additionally, on more prominently colored individuals the base of
the antennae and the posteromedial tubercle of the head capsule can also be colored.
Compound eyes are a muddled tan to reddish.
Morphology. Head. Head capsule about as long as wide, with a vertex that has
moderate granulation throughout and a prominent but not broad posteromedial tu-
bercle which is larger than any of the granules on the head capsule (Fig. 40E). Frontal
convexity not particularly long but ending in a ne point and covered with sparse thin
setae. Compound eyes large and bulbous, taking up ca. ⅖ of the head capsule lateral
margins (Fig. 40E). ere are three moderately developed ocelli located between and
slightly posterior to the compound eyes. Antennal elds about as wide as the scapus.
Antennae. Antennae (including the scapus and pedicellus) consists of 25 segments, all
segments except the scapus and pedicellus and terminal three segments are covered in
dense setae that are as long as or longer than the antennae segment is wide. e termi-
nal three segments are covered in dense short setae and the scapus and pedicellus are
nearly completely bare. orax. Pronotum with anterior margin slightly concave and
lateral margins that are straight or slightly convex and converging to a straight posterior
margin that is half the width of the anterior rim (Fig. 40E). Anterior margin of the
pronotum has a distinct rim, lateral margins have moderate rims, and the posterior
margin lacks a rim (Fig. 40E). Face of the pronotum is marked by a distinct sagittal fur-
row and pit in the center, a granular surface, and a slight perpendicular furrow from the
central pit. Prosternum is granulose throughout with small nodes of nearly even size.
Mesosternum anterior half with nodes of a similar size to the prosternum and those on
the posterior half slightly less prominent. e metasternum has a slightly wrinkled sur-
face and sparse granulation. Prescutum longer than wide, with lateral margins slightly
converging to the posterior (Fig. 40E). Lateral rims with small granulation throughout
giving them a rough textured appearance, only three or four are slightly larger than the
rest. Prescutum surface with granulation throughout with those along the sagittal plane
slightly larger than the others. Prescutum anterior rim weakly formed but marked with
a surface which is granular. Mesopleura narrow, almost parallel for the anterior quarter,
and then only gradually diverge for the remainder of the length (Fig.40E). Lateral mar-
gin lacking prominent tubercles, instead marked with sharp granulation throughout
with only two or three slightly larger than the rest, giving the margins a rough textured
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 93
Figure 40. Cryptophyllium khmer gen. et sp. nov. holotype male, from Tatai, Cambodia, photographs by
Jérôme Constant (RBINS) A habitus, dorsal B habitus, ventral C pro- tibial and femoral lobes, dorsal
D details of the antennae, head, and thorax, lateral E details of the antennae, head, and thorax, dorsal
Fterminalia, dorsal G genitalia, ventral.
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
94
appearance. Face of the mesopleura slightly wrinkled and with two faint divots, one
near the anterior margin and one half-way through the length (Fig.40D). Wings. Teg-
mina moderate length, reaching ⅓ to ½ onto abdominal segment III. Tegmina wing
venation: the subcosta (Sc) is the rst vein, is simple, and terminates ca. ½ through the
overall tegmina length. e radius (R) spans the entire length of the tegmina with the
rst radius (R1) branching < ca. ½ through the wing length and terminating just distal
to the midline, followed by the branching and termination of the second radius (R2)
near the distal ⅓ of the wing, and then the radial sector runs to the wing apex. e
media (M) also spans the entire length of the tegmina with the rst media posterior
(MP1) branching o ca. ⅖ of the way through the wing length, and then the second
media posterior (MP2) branches near the midline, and the media anterior (MA) runs
to the wing apex. e cubitus (Cu) runs along the edge of the wing as the two media
posterior veins fuse with it and as the cubitus reaches the apex it fades. e rst anal
(1A) vein terminates upon reaching the cubitus slightly < ⅓ of the way through the
wing length. Alae well-developed in an oval fan conguration, long, reaching onto ab-
dominal segments IX or X. Alae wing venation: the costa (C) is present along the entire
foremargin giving stability to the wing. e subcosta (Sc) is long, spanning slightly >
⅔ of the wing length and is mostly fused with the radius in the beginning but termi-
nates when it meets the costa. e radius (R) spans the entire wing and branches ca.
⅓ of the way through into the rst radius (R1) and radial sector (Rs) which run gently
diverging for most of their length and then converge at the apex of the wing where
they terminate near each other but not touching. e media (M) branches early, ca.
⅙ of the way through the wing into the media anterior (MA) and the media posterior
(MP) which run parallel with each other throughout the wing until the distal ⅙ of the
wing where the media posterior fuses with the media anterior which then run fused
together to the wing apex where they terminate near the radial sector. e cubitus (Cu)
runs unbranched and terminates at the wing apex. Of the anterior anal veins, the rst
anterior anal (1AA) fuses with the cubitus near the point where the media branches into
the media anterior and media posterior and then the rst anterior anal branches from
the cubitus ⅔ of the way through the wing length where it uniformly diverges from
the cubitus until it terminates at the wing margin. e anterior anal veins two–seven
(2AA–7AA) have a common origin and run unbranched in a folding fan pattern of rela-
tively uniform spacing to the wing margin. e posterior anal veins (1PA–6PA) share a
common origin separate from the anterior anal veins and run unbranched to the wing
margin with slightly thinner spacing than the anterior anal veins. Abdomen. Margins of
abdominal segment II either slightly converging or parallel-sided. Abdominal segments
III through the anterior ⅔ of IV diverging. Segment V with parallel margins and VI–X
converging slowly at rst then more prominently for the terminal three segments, giv-
ing the abdomen a spade-shaped appearance. Genitalia. Poculum broad, posteriorly
rounded and with a shallow notch medioapically; slightly passes the anterior margin of
segment X (Fig. 40G). Cerci long and slender, extending from under the anal abdomi-
nal segment, slightly cupped with a granular surface and numerous short setae through-
out (Fig. 40F). Vomer broad and stout with sides evenly converging and terminating
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 95
in an upward hooking apical spine with a smaller hook next to the base of the primary
spine (Fig. 5D). Legs. Profemoral exterior lobe about the same width as the interior
lobe or slightly wider, smoothly arcing from end to end and marked with a granular
margin and ve or six small serrate teeth on the distal half only (Fig. 40C). Profemoral
interior lobe roundly triangular and marked with ve teeth arranged in a two-one-two
pattern with prominent looping gaps between the sets and the middle tooth larger than
the others (Fig. 40C). Mesofemoral exterior lobe arcs end to end, but is slightly more
bent than the interior lobe and is broader on the distal half which can either be lacking
dentation or have three or four dulled teeth, and the proximal half that is rather thin
and lacking teeth. Mesofemoral interior lobe of a similar width to the exterior lobe, is
broader on the distal end and is marked with ve or six serrate teeth mostly situated
on the distal ⅓ to ½ of the lobe. Metafemoral exterior lobe lacks dentation, and has a
straight margin along the metafemoral shaft. Metafemoral interior lobe smoothly arcs
end to end with eight or nine serrate teeth on the slightly wider distal half. Protibiae
lacking exterior lobe, interior lobe reaching end to end in a smoothly rounded triangle
with the widest portion ca. 3–3½× as wide as the protibial shaft and situated just distal
to the midline. Meso- and metatibiae simple, lacking lobes completely.
Measurements of holotype male [mm]. Length of body (including cerci and head,
excluding antennae) 61.9, length/width of head 4.5/3.9, antennae 37.3*, pronotum
3.6, mesonotum 4.3, length of tegmina 19.0, length of alae 49.3, greatest width of
abdomen 17.3, profemora 13.5, mesofemora 11.3, metafemora 13.4, protibiae 9.7,
mesotibiae 7.8, metatibiae 10.1.
Measurements of paratype males [mm]. Length of body (including cerci and head,
excluding antennae) 63.8–70.2, length/width of head 5.0–5.5/4.1–4.3, antennae 38.8–
39.5, pronotum 3.6–4.1, mesonotum 5.0–6.2, length of tegmina 20.0–20.4, length of
alae 50.0–52.1, greatest width of abdomen 17.1–17.9, profemora 15.7**, mesofemora
12.1, metafemora 13.7–14.1, protibiae 9.8**, mesotibiae 8.7–8.9, metatibiae 11.2–11.4.
Eggs. (Fig. 41). e lateral surfaces are at with a length ca. 1½× the width with
parallel margins, giving the capsule a rectangular appearance. All surfaces have nu-
merous small to medium sized pits throughout, the lateral surface has around 35 pits
(mostly on the smaller end of the spectrum) arranged in no detectable order, some
more closely spaced than others. In addition, between the pits the surfaces are covered
with short moss-like pinnae with the pinnae along the margins slightly longer than the
pinnae on the other surfaces. e dorsal surface is marked with six or seven slightly
irregular medium sized pits on each half running the length of the capsule with short
moss-like pinnae around the micropylar plate and between the pits. e micropylar
plate is not overly long, occupying ca. ½ of the dorsal surface length but not perfectly
centered, with ca. ⅓ of the unoccupied space below and ⅔ above the micropylar plate.
e micropylar cup is the widest portion of the micropylar plate and is located ca. ⅓
of the dorsal surface length from the posterior. e micropylar plate is approximately
* Antennae of the holotype broken, measurement is only until segment 23.
** Only measured on the largest specimen, the smallest are missing their front legs.
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
96
Figure 41. Cryptophyllium khmer gen. et sp. nov. egg, RBINS collection, photographs by Jérôme Con-
stant (RBINS) A lateral view B dorso-lateral view C dorsal view D opercular (anterior) view E posterior
view F ventral view.
teardrop-shaped with the anterior portion longer and thinner than the posterior after
the micropylar cup. Operculum slightly ovular, with the outer margin encircled with
short moss-like pinnae surrounding the operculum and four or ve medium pits sur-
rounding the dorsal and lateral margins. e operculum is roundly raised with a height
slightly > ½ operculum width. is rounded raised cap is marked with a sagittal raised
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 97
row of pinnae similar in length to those along the capsule margins. e rounded raised
cap is not perfectly centered and instead the rounded projection is shifted slightly to-
wards the ventral surface. e overall egg color is tan to light brown, with the moss-like
pinnae sometimes slightly lighter in color.
Measurements including the extended pinnae [mm]. Length (including opercu-
lum): 5.6; maximum width of capsule when viewed from lateral aspect 3.2; length of
micropylar plate 3.0.
Etymology. Noun. e species epithet is the Hindi word khmer, meaning Cambo-
dia, referring to the country of origin for this species.
Cryptophyllium limogesi gen. et sp. nov.
http://zoobank.org/411639C9-07AD-4BE6-AFAD-F8D5D4EA1152
Figures 5J, 8Q, 8R, 42, 43, 44, 45
Material examined. Holotype ♀: “VIETNAM, Lam Dong Province, Bao Lam, Dam-
bri, V.2018”. Deposited within the Montreal Insectarium (IMQC).
Paratypes: (13 ♀♀, 2 ♂, 7 eggs) • 1 ♂: “Coll. I.R.Sc.N.B., VIETNAM, Dak Nong
prov., Ta Dung N.P., 11°52’22”N 107°58’40”E, 5-8.viii.2019, GTI Project, Leg. J.
Constant and J. Bresseel, I.G.: 34.048 (Coll. I.R.Sc.N.B.)”, vomer dissected (RBINS)
(SB0531 molecular sample within our analysis) • 1 ♂: “Vietnam: Dak Lak Province,
Local collector, September 2020” (IMQC) • 5 ♀♀; “Vietnam: Dak Lak Province, Lo-
cal collector, September 2020” (Coll RC 20-127, 20-128, 20-129) • 5♀♀; “Vietnam:
Dak Lak Province, Local collector, September 2020” (Coll SLT) • 3 ♀♀; “Vietnam:
Dak Lak Province, Local collector, September 2020” (IMQC) • 1 egg: “Vietnam: Dak
Lak Province, Local collector, September 2020” (Coll SLT) • 1 egg: “Vietnam: Dak
Lak Province, Local collector, September 2020” (Coll FH) • 3 egg: “Vietnam: Dak
Lak Province, Local collector, September 2020” (IMQC) • 2 egg: “Vietnam: Dak Lak
Province, Local collector, September 2020” (Coll RC).
Remarks. is large and morphologically unique species has been located in sev-
eral provinces of southern Vietnam over the last few years and molecularly we found
it to be sister species to Cryptophyllium icarus sp. nov. (Fig. 4) a species also only at the
present known from southern Vietnam (Fig. 2). Interestingly, these molecular sister
species are morphologically drastically dierent, perhaps due to their geographic sym-
patry. During a GTI joint expeditions two small nymphs were collected in Ta Dung
N.P. on the forest edge adjacent to a coee plantation. Tim Bollens (Belgium) reared
one nymph to adulthood, thus revealing the male morphology (Fig. 42).
Dierentiation. Female Cryptophyllium limogesi sp. nov. are most morphologically
similar to Cryptophyllium celebicum comb. nov. and Cryptophyllium tibetense comb.
nov. based on the wide profemoral exterior lobe with an acute angle and broad boxy
abdomen. From Cryptophyllium celebicum comb. nov., the easiest observed dierence
is the structure of the thorax, with the mesopleura of Cryptophyllium celebicum comb.
nov. notably narrower on the anterior half vs. the mesopleura of Cryptophylliumlimogesi
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
98
Figure 42. Holotype female Cryptophyllium limogesi, photographs by René Limoges (IMQC) A habitus,
dorsal B habitus, ventral C head and antennae dorsal D antennae detail dorsal E thorax dorsal F front leg,
dorsal G terminalia, dorsal H genitalia, ventral.
sp. nov. reaching nearly to the anterior margin of the prescutum with straight margins
(Fig. 42E). From Cryptophyllium tibetense comb. nov. the female subgenital plates easily
dierentiate these species as Cryptophyllium tibetense comb. nov. has a long subgenital
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 99
plate with a point which exceeds the apex of the terminal abdominal segment (Fig.
63B) vs. Cryptophyllium limogesi sp. nov. where the subgenital plate is short, reaching
no more than ½ through the terminal abdominal segment (Fig. 42H).
Male Cryptophyllium limogesi sp. nov. are similar morphologically to Cryptophyl-
lium oyae comb. nov. due to the broad spade-shaped abdomen and the profemoral
lobe shapes. ese species can be dierentiated by the length of their tegmina (only
reaching onto abdominal segment III in Cryptophyllium limogesi sp. nov. but reaching
halfway onto segment IV in Cryptophyllium oyae comb. nov.); and they can be dieren-
tiated by the shape of the mesopleura as they are broad and nearly straight margined in
Cryptophyllium oyae comb. nov. but slightly narrower on the anterior in Cryptophyllium
limogesi sp. nov. males (Fig. 44C).
Distribution. Southern Vietnam: presently known from three provinces: the type
locality of Lam Dong Province, Bao Lam, Dambri; the male paratype record from Dak
Nong Province, Ta Dung N.P.; an observational record from Dak Lak Province, Chu
Yang Sin N.P.; and paratype records from Dak Lak Province.
Description. Female. Coloration. Coloration description is based on photos of
the holotype female shortly after being preserved. Nearly the entire body was of a uni-
form lime-green without diering colored markings (all legs and even wing venation
a similar color to the rest of the body). Only the compound eyes were slightly yellow
and not the same shade of green as the rest of the body.
Morphology. Head. Head capsule longer than wide, vertex with a moderately
granular surface, and the posteromedial tubercle which is three or four times larger
than the most prominent granules on the capsule. Frontal convexity broad and stout,
notably shorter than the length of the rst antennomere, and with several long, thin,
clear setae across the surface. Compound eyes are not large, only slightly protruding
from the head capsule and with a width of ca. ¼ the head capsule length (Fig. 42C).
Ocelli absent. Antennal elds wider than the rst antennomere but not protruding
back farther than the frontal suture. Antennae. Antennae consist of nine segments,
with the terminal segment approximately the same length as the previously three seg-
ments combined (Fig. 42D). e eighth antennal segment has a distinct furrow around
the middle which makes the segment appear to be two separate segments (giving the
antennae a ten segmented appearance), but this furrow appears to only be supercial
(Fig.42D). Antennomeres I-III sparsely marked with thin transparent setae, similar to
those found on the frontal convexity, but slightly shorter in length. Antennomere IV
is short, disk-like, and wider than the following segments, and interestingly has a base
which is narrow and somewhat longer than other congeneric antennal segments IV,
giving it a raised appearance (Fig. 42D). e terminal antennomere and the distal half
of segment VIII (distal to the midline furrow) are covered in dense, stout, setae. orax.
Pronotum with anterior margin slightly concave and lateral margins that are relatively
straight, converging to a narrow, straight posterior margin that is ca. ½ the width of the
anterior rim (Fig. 42E). e pronotum surface is smooth, with only a prominent pit
in the center, and slight furrows anterior and lateral to the pit, no prominent wrinkles
or granulation. e pronotum has a prominent anterior rim and moderate lateral rims,
the posterior is lacking a rim. Prosternum with notable nodes throughout the surface,
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
100
relatively evenly spaced. Mesosternum with prominent nodes on the anterior margin,
followed by moderate nodes on the anterior ⅓ of the surface, with the remainder
with dispersed, weak granulation which continues onto the metasternum. Prescutum
notably longer than wide, with lateral margins running parallel to the posterior mar-
gin giving it a distinctly rectangular appearance. Lateral rims with ve or six medium
sized tubercles situated on the anterior ⅔, with only small granulation on the remain-
der. Prescutum anterior rim prominent but not strongly protruding, with the surface
granular and lacking a prominent sagittal tubercle. Prescutum surface with granulation
throughout with those along the sagittal plane slightly larger (Fig. 42E). Mesopleura
start ca. ⅓ down the prescutum and evenly diverge with straight lateral margins. Lat-
eral margin with six or seven major and distinctly pointed tubercles and six or seven
smaller tubercles intermixed amongst them. is mix of tubercles is only prominent
along the anterior ⅔ of the length with the remaining ⅓ lacking notable tubercles and
instead marked with consistent granulation (Fig. 42E). Face of the mesopleura with
slight wrinkles throughout most of the surface and a few irregular nodes along the
lateral margin, as well as two faint divots, one on the anterior margin and one closer
to the center. Wings. Tegmina reaching slightly past the anterior margin of abdominal
segment VII. Tegmina venation is rather typical for the Cryptophyllium gen. nov. e
subcosta (Sc) is the rst vein in the forewing and runs parallel with the wing for the
rst half of its length, and then bends towards the wing margin for the second half.
e radius (R) spans the central portion of the tegmina with two subparallel branched
veins. e rst radius (R1) branches ca. ½ through the radius length and terminates
ca. ⅓ of the way through the wing length. e radial sector (Rs) branches from the end
of the radius and runs angled to the wing margin where it terminates near the wing
midline length. ere is a weak continuation of the radius following the prominent ra-
dial sector branching which continues on as a short and thin radius to media crossvein
(R–M). e media (M) is simply bifurcate with both the media anterior (MA) and
media posterior (MP) terminating close to the posterior ¼ of the wing. e cubitus
(Cu) runs throughout the entire wing length simply, and then near the posterior ⅕
of the wing becomes bifurcate into the cubitus anterior (CuA) and cubitus posterior
(CuP) which both terminate at or very near the wing posterior apex. e rst anal vein
(1A) is simple and fuses with the cubitus early on, at around the midline between the
rst radial branching and the radial sector branching. Alae short, only 21.8 mm long.
Abdomen. Abdominal segments II through the anterior half of IV diverging, posterior
half of IV through the anterior ⅓ of VII parallel. Abdominal segment VII with a dis-
tinct looping lobe which meets abdominal segment VIII which is notably narrower.
Segments VIII–X uniformly converge to a broad rounded apex. Genitalia. Projecting
portion of the subgenital plate stout, beginning at the anterior margin of abdominal
segment IX and projecting with nearly straight sides to just under the anterior mar-
gin of the terminal abdominal segment (Fig. 42H). Gonapophyses VIII are long and
broad with a dagger-like shape (parallel-sided at rst and then after ca. ½ of the length
uniformly converging to the point) with the points just projecting from under the
terminal abdominal segment (Fig. 42H). Gonapophyses IX are smaller and not visible
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 101
from under the large gonapophyses VIII. Cerci strongly pointed and relatively at (not
strongly cupped) with weakly crenate margins, and the dorsal surface heavily granular
and marked by thin transparent setae throughout (Fig. 42G, H). Legs. Profemoral
exterior lobes broad, at its broadest ca. 2× wider than the interior lobe, and distinctly
serrate throughout the entire length (with 13–17 small, pointed teeth). e proximal
edge is slightly concave and the distal edge is smoothly convex, therefore giving the
lobe a distinct recurved appearance and an acute exterior angle (Fig. 42F). Interior
profemoral lobe ca. 2½× the width of the profemoral shaft at its widest and with dou-
bly serrate, large, triangular teeth. e largest teeth are grouped into a two-one-two
pattern with large looping gaps between these groupings, with these large gaps also
nely serrate, not smooth (Fig. 42F). Interior mesofemoral lobe arcs evenly weighted
from end to end, and at its widest is approximately the same width as the mesofemoral
shaft itself. e interior mesofemoral lobe is nely serrate for ca. ¾ of its distal length
with seven or eight teeth. Mesofemoral exterior lobe is also approximately at its widest
as wide as the mesofemoral shaft, but the exterior lobe is distinctly bent in the center
with straight margins, not smoothly arcing from end to end. On the distal half of the
lobe only there are six or seven small serrate teeth. Metafemoral interior lobe arcs end
to end and has ve or six dull teeth pointing distally. Metafemoral exterior lobe is thin
and smooth, hugging the metafemoral shaft and lacks dentation. Protibiae with a thin
but notable exterior lobe on the distal ⅕ only. Protibial interior lobe spans the entire
length as a broad scalene triangle with the broad end on the distal half of the protibiae.
Mesotibiae simple, completely lacking lobes. Metatibiae lacking interior lobes, exterior
is marked by a very slender lobe which only occupies the distal ¼ of the shaft.
Measurements of holotype female [mm]. Length of body (including cerci and
head, excluding antennae) 99.0, length/width of head 8.9/6.8, antennae 5.1, prono-
tum 5.9, mesonotum 6.5, length of tegmina 55.7, length of alae 21.8, greatest width of
abdomen 41.5, profemora 22.8, mesofemora 16.0, metafemora 19.1, protibiae 11.2,
mesotibiae 11.8, metatibiae 15.0.
Male. Coloration. Coloration description based on the captive reared paratype
male when it was alive (Fig. 43). Overall coloration mint-green throughout with high-
lights of tan coloration on the distal half of the protibiae, along all femoral lobe mar-
gins, the frontal convexity, the margins of the thorax, the base of the antennae and the
distal tips of each longer antennomere, and intermittently along the tegmina and alae
sclerotized veins. Abdominal segment V has a slightly darker patch and a transparent
eye spot on each side of the midline. Compound eyes are pale yellow with slight orange
marbling throughout.
Morphology. Head. Head capsule approximately as long as wide, with a vertex that
is only slightly granular with no discernable pattern (Fig. 44C). e posteromedial
tubercle is not broad and is only weakly raised from the head capsule. Compound eyes
are large and bulbous, taking up ca. ⅖ of the head capsule lateral margins (Fig. 44C).
ere are three well-developed ocelli between and slightly posterior to the compound
eyes (Fig. 44C). Antennal elds are about as wide as the scapus. Antennae. Antennae
(including the scapus and pedicellus) consist of 29 segments. e scapus and pedicel-
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102
Figure 43. Paratype male Cryptophyllium limogesi gen. et sp. nov. from Ta Dung N.P. reared to adult-
hood and photographed by Tim Bollens (Belgium) A habitus, dorsal B detail of the front legs and head
C genitalia details, ventral.
lus are nearly completely bare, lacking long setae. e following segments except the
terminal three are covered in dense setae that are as long as or longer than the anten-
nae segment is wide, and the terminal three segments are covered in dense short setae.
orax. Pronotum with anterior margin slightly concave and lateral margins that are
straight and converging to a slightly curved posterior margin that is ca. ½ the width
of the anterior rim (Fig. 44C). Anterior and lateral margins have moderate rims and
the posterior margin lacks a rim (Fig. 44C). Face of the pronotum is marked by a
distinct pit in the center with a furrow anterior to the pit along the sagittal plane,
weakly formed furrows lateral to the central pit, and a smooth surface with only slight
granulation in no detectable pattern (Fig. 44C). e prosternum is slightly granu-
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 103
lose throughout and the anterior ⅓ of the mesosternum surface is marked with more
prominent nodes, with the remainder of the surface with fewer and smaller nodes
(Fig.44B). Prescutum longer than wide, with lateral margins slightly converging to
the posterior (Fig. 44C). Lateral rims with eight or nine small tubercles (Fig. 44C).
Prescutum surface slightly raised along the sagittal plane with six or seven nodes of
varying size, with the remainder of the surface with only slight granulation through-
out. Prescutum anterior margin weakly formed and with a granular surface, lacking a
prominent central tubercle. Mesopleura narrow on the anterior quarter of the length
but then gently diverging with nearly straight margins to the posterior (Fig. 44C).
Mesopleura lateral margin with three large conical tubercles, three or ve small tuber-
cles, and four or ve nodes throughout the length (Fig. 44C). Face of the mesopleura
slightly wrinkled, with slight granulation throughout, and with two faint divots, one
on the anterior margin and one near the midline. Wings. Tegmina moderate length,
extending halfway through abdominal segment III. Tegmina wing venation: the sub-
costa (Sc) is the rst vein, is simple, and terminates the earliest slightly < ½ through
the overall tegmina length. e radius (R) spans the entire length of the tegmina with
the rst radius (R1) branching ca. ⅓ of the way through the wing length and terminat-
ing near the midline, followed by the branching and termination of the second radius
(R2) slightly distal to the midline, and then the radial sector runs to the wing apex.
e media (M) also spans the entire length of the tegmina with the rst media poste-
rior (MP1) branching o ca. ⅓ of the way through the wing length, then the second
media posterior (MP2) branching near the midline, and the media anterior (MA) runs
to the wing apex. e cubitus (Cu) runs along the edge of the wing as the two media
posterior veins fuse with it and as the cubitus reaches the apex it fades. e rst anal
(1A) vein terminates upon reaching the cubitus ca. ⅓ of the way through the wing
length. Alae well-developed in an oval fan conguration, long, reaching the middle
of abdominal segment VIII. Abdomen. Abdominal segment II gently diverging, III
through the anterior half of segment IV diverging to the widest portion of the abdo-
men. e posterior of IV–V parallel-sided giving the abdomen a broad spade-shaped
appearance. Segments VI–X uniformly converging with slightly undulating margins
(Fig. 44B). Genitalia. Poculum broad, and ends in a broadly rounded apex that slightly
passes the anterior margin of segment X (Fig. 44G). Cerci long and slender, with > ½
their length extending from underneath the terminal abdominal segment (Fig. 44F),
relatively at, covered in a heavily granulose surface and with equally granular margins,
and surface with numerous short setae (Fig. 44G). Vomer stout with slightly rounded
sides converging to an apex with two side by side thick apical hooks which are the same
size and hook upwards into the paraproct (Fig. 43C). Legs. Profemoral exterior lobe
broader than the interior lobe, arcing end to end with a distinct rounded bend in the
center, with the proximal half margin with a distinctly granular surface, and the distal
half with six small serrate teeth (Fig. 44E). Profemoral interior lobe roundly triangular,
at least 3× wider than the profemoral shaft, and marked with ve large, serrate teeth
arranged in a two-one-two pattern with looping gaps between them (Fig. 44E). Mes-
ofemoral exterior lobe arcs end to end, is slightly wider than the mesofemoral shaft,
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104
Figure 44. Paratype male Cryptophyllium limogesi gen. et sp. nov. same individual as in Figure 43 photo-
graphs by Jérôme Constant (RBINS) Ahabitus, dorsal B habitus, ventral C details of the base of anten-
nae, head, and thorax, dorsal D details of the base of antennae, head, and thorax, lateral E front leg details,
dorsal F genitalia details, dorsal G genitalia details, ventral.
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 105
but with the widest portion on the distal ⅓ which is marked with granulation or with
one or two weakly formed teeth. Mesofemoral interior lobe is slightly thinner than the
exterior lobe and is slightly broader on the distal end which is marked with six serrate
teeth. Metafemoral exterior lobe lacks dentation, and has a straight margin hugging
the metafemoral shaft. Metafemoral interior lobe smoothly arcs end to end, with the
distal half wider than the proximal half, and the distal half is marked with seven or
eight serrate teeth on the distal half. Protibiae with a small but notable exterior lobe on
the anterior ⅕ which is no wider than the width of the protibial shaft (Fig. 44E). Proti-
bial interior lobe reaching end to end in a smoothly rounded triangle with the widest
portion on the distal third ca. 3× as wide as the protibial shaft (Fig. 44E). Meso- and
metatibiae simple, lacking lobes.
Measurements of reared paratype male [mm]. Length of body (including cerci
and head, excluding antennae) 79.7, length/width of head 4.9/3.9, antennae 41.8,
pronotum 3.7, mesonotum 5.7, length of tegmina 24.4, length of alae 57.5, greatest
width of abdomen 26.5, profemora 16.4, mesofemora 11.4, metafemora 13.7, proti-
biae 9.1, mesotibiae 7.7, metatibiae 10.9.
Eggs. (Fig. 45). When viewed from the anterior the egg capsule cross section is
rounded pentagonal, therefore the lateral surfaces are raised into dorsolateral and ven-
trolateral surfaces. All surfaces and margins slightly undulate giving the egg an overall
lumpy appearance. e ventrolateral surface is marked by four large, evenly spaced pits
from the anterior to the posterior in a singular line. e dorsolateral surface has ve
large pits, arranged with one on the anterior, one on the posterior, and three in the
middle spaced out in a two-one pattern. Both lateral surfaces are primarily bare but do
have sparse and small moss-like pinnae between the pits. e dorsal surface is marked
with seven medium sized pits total (one on the anterior end along the sagittal plane at
the apex of the micropylar plate, followed by three on each slide of the plate with broad
spacing between them ending with the posterior most near the base of the capsule).
e dorsal surface is also marked throughout with short moss-like pinnae around the
micropylar plate, with the area immediately around each pit bare. e micropylar plate
is long, ca. 5/7 of the overall dorsal surface length with the micropylar cup situated on
the posterior ⅓ of the length. e micropylar plate is thin with the widest portion the
area around the micropylar cup. e operculum is slightly ovular with a surface that
is roundly raised and a height slightly < ½ the operculum width. e operculum is
marked intermittently with moss-like pinnae similar in shape but slightly smaller than
those found on the rest of the capsule, as well as marked with two medium sized pits
(one on each side of the sagittal plane). e overall egg color is dark brown, with the
moss-like pinnae light brown in color so they stand out clearly on the surface.
Measurements including the extended pinnae [mm]. Length (including opercu-
lum): 5.2; maximum width of capsule when viewed from lateral aspect 3.8; length of
micropylar plate 3.0.
Etymology. Patronym. Named after René Limoges (Canada) from the Montreal
Insectarium to thank him for his many years of assisting Team Phyllies with countless
publication worthy photographs.
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
106
Figure 45. Paratype Cryptophyllium limogesi gen. et sp. nov. egg, laid by paratype females from Dak Lak
Province, Vietnam. Photographs by René Limoges (IMQC) A lateral B dorsal C ventral D opercular
(anterior) view.
Cryptophyllium liyananae gen. et sp. nov.
http://zoobank.org/87CFC18E-3479-47CD-B052-37AEB4A07BDB
Figures 46, 47
Material examined. Holotype ♀: “CHINA: Guangxi Prov., Jinxiu County, Liuzhou
City, Dayaoshan Mountain, 875-1,500 m., 18-19.IX.2019. (Coll RC 20-002)”. De-
posited in the Montreal Insectarium (IMQC).
Paratypes: (6 ♀♀, 4 eggs) • 1 ♀: “CHINA: Guangxi Prov., Jinxiu County, Liu-
zhou City, Dayaoshan Mountain, 875-1,500 m., 19.IX.2019. Coll RC 19-182” (Coll
RC) • 1 ♀: “CHINA: Guangxi Prov., Jinxiu County, Liuzhou City, Dayaoshan Moun-
tain, 875-1,500 m., 18-19.IX.2019. Coll RC 20-003” (Coll RC) • 3 ♀♀: Same data
as the holotype (Coll SLT) • 1 ♀: Same data as the holotype (Coll FH)• 4 eggs: Re-
moved from the abdomen of paratype female 19-182. “CHINA: Guangxi Prov., Jinxiu
County, Liuzhou City, Dayaoshan Mountain, 875-1,500 m., 19.IX.2019”; Coll RC
20-069–20-072: 20-069, 20-072 (Coll RC); 20-070, 20-071 (IMQC).
Photographic record: (1 ♀) In addition to the type material examined, images
of a live female observed by Dr. Lu Qiu in Guangxi, Huaping Nature Reserve, 900–
1000m, in August 2020 (Fig. 46B) were shared with us and compared to type material.
Remarks. is species appears to be the same which was illustrated by Liu (1993)
and called Phyllium (Phyllium) celebicum within the work. Not only do the illustra-
tions clearly show similar morphology, but one of the specimens is also from the lo-
cality of Dayao Mountain, Jinxiu, Guangxi. e illustrated female (gures 10–12;
Liu 1993) clearly shows gently rounded lobes on abdominal segment VII, the same
tegmina length, a small but notable metatibial exterior lobe on the distal end, the same
shape/length features of the genitalia (Fig. 47E), mesothorax shape (Fig. 47D), similar
rounded profemoral exterior lobes (Fig. 47C), and the VIII antennal segment which is
notably longer than the other preceding individual segments (Fig. 47B).
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 107
Figure 46. Female Cryptophyllium liyananae gen. et sp. nov. A live holotype female, photograph by
Chengzhi Bian (China) B live female observed and photographed by Dr. Lu Qiu in Guangxi, Huaping
Nature Reserve, 900–1000 m, August 2020 C paratype female Coll RC 19-182 shortly after being col-
lected before the color faded, photograph by Chengzhi Bian.
Dierentiation. Females are morphologically similar to Cryptophyllium drunga-
num comb. nov. due to the tibial and femoral shapes (including small exterior pro- and
metatibial anteriorly situated lobes), alae long (reaching abdominal segment VI), a
seventh abdominal segment which is slightly lobed, and similar genitalia shapes and
lengths of features. ese species can be dierentiated by the mesopleura as they are
broader on the anterior end in Cryptophyllium liyananae sp. nov. and slightly narrower
in Cryptophyllium drunganum comb. nov. and the overall size as Cryptophyllium liya-
nanae sp. nov. are 88.0–92.0 mm long and the holotype Cryptophyllium drunganum
comb. nov. is 75.0 mm long. Males are presently unknown.
Distribution. Presently only conrmed from two localities in Guangxi Province,
Liuzhou prefecture-level city (Fig. 46A, C) and Guilin prefecture-level city (Fig. 46B).
e other specimens within Liu (1993) with varying locality data which may represent
additional localities have not been reviewed so it is possible that these may represent
other species or range expansions.
Description. Female. Coloration. Coloration descriptions are based upon photos
of the live individuals which became the type material herein described (Fig. 46A, C).
e overall coloration is pale mint green, with highlights of yellow, orange, and red.
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
108
e antennae, interior profemoral lobe margins, and the margins of the terminal ab-
dominal segments are red. e mesopleura margins, veins of the tegmina, and exterior
profemoral lobe margins are orange to yellow.
Morphology. Head. Head capsule about as long as wide, vertex smooth (Fig. 47D).
e posteromedial tubercle is the only notable feature on the head capsule. Frontal
convexity broad for the posterior half then narrowing on the anterior half, about as
long as the rst antennomere, and with slight granulation on the dorsal surface and
several setae present which are longer than any setae on the antennae. Compound eyes
slightly protruding from the head capsule, taking up ca. ¼ of the length of the lateral
head capsule margins (Fig. 47D). Ocelli absent. Antennal elds slightly wider than the
rst antennomere and slightly shallower than the rst antennomere is tall (Fig. 47B).
Antennae. Antennae consisting of nine segments, with the terminal segment slightly
shorter than the preceding two segment lengths combined (Fig. 47B). Antennomeres
I–VIII sparsely marked with small transparent setae, the terminal antennomere is more
densely covered in stout, brown setae. e pars stridens of antennomere III has 46 or
47 teeth. orax. Pronotum with gently concave anterior margin and straight lateral
margins, which converge to a convex posterior margin that is ½ the width of the an-
terior margin (Fig. 47D). e pronotum surface is smooth, with only a prominent pit
in the center, and slight furrows anterior, posterior, and lateral to the pit (Fig. 47D).
e pronotum has a prominent anterior rim and weakly formed lateral and posterior
rims, all of which are relatively smooth (Fig. 47D). Prosternum and the mesosternum
with stout and numerous nodes, with the central area of the mesosternum with less
nodes and relatively smooth. Metasternum with granulation reduced and only mini-
mal. Prescutum anterior margin as wide as the presuctum is long, with margins slightly
narrowing on the anterior ⅓ and then running parallel to the posterior margin which
is slightly narrower than the anterior margin (Fig. 47D). Lateral rims with 7–9 short
tubercles which are all about the same size (Fig. 47D). Prescutum anterior rim promi-
nent but not strongly protruding, with a surface that is granular, lacking a singular
prominent sagittal tubercle (Fig. 47D). Prescutum surface with minimal granulation
throughout, with those along the sagittal plane only slightly larger (Fig. 47D). Meso-
pleura beginning near the anterior margin of the prescutum and evenly diverging;
lateral margin with 9–13 tubercles which are variable in size, with three larger than the
rest (Fig. 47D). Face of the mesopleura relatively smooth, and with two notable divots,
one near the anterior margin and another ca. ⅗ of the way through the length (Fig.
47D). Wings. Tegmina reaching ca. ½ through abdominal segment VII. e subcosta
(Sc) is the rst vein in the forewing, is distinctly bent in the center, and terminates ca.
¼ of the way through the wing length. e radius (R) spans the anterior half of the
forewing with two subparallel branched veins; radius 1 (R1) terminates ca. ⅓ of the
way through the wing length, and the radial sector (Rs) terminates in the center of the
wing at the widest portion. ere is a weak continuation of the radius following the
prominent Rs branching which continues on as a short and thinner R–M crossvein
that does not appear to solidly connect the two veins fading as it reaches the media.
e media (M) is bifurcate with both the media anterior (MA) and media posterior
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 109
Figure 47. Holotype Cryptophyllium liyananae gen. et sp. nov., photographs by René Limoges (IMQC)
A habitus, dorsal B details of the antennae, dorsal C profemoral lobes, dorsal D details of the antennae,
head, and thorax, dorsal E genitalia, ventral.
(MP) terminating on the posterior ⅓ of the wing. In some individuals there is a weak
continuation of the media as a thin crossvein to the cubitus, but this was not present
in all individuals observed. e cubitus (Cu) is also bifurcate, branching near the pos-
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
110
terior ⅓ to ¼ of the wing into the cubitus anterior (CuA) and cubitus posterior (CuP)
which both terminate at or very near the wing posterior apex. e rst anal vein (1A)
is simple and fuses with the cubitus early on, only slightly past the branching distance
of the rst radius from the radius. Alae well-developed, reaching abdominal segment
VI (41.0–43.0 mm long). Abdomen. Abdominal shape relatively stable between all
females observed. Segments II through the anterior ⅓ of IV evenly diverging, with the
posterior ⅔ of segment IV the widest segment. Segments V–VI are slightly subparallel,
converging gently to the posterior, giving the abdomen a slight narrowing appearance.
Segment VII is distinctly rounded to the terminal three segments which are notably
narrower than the previous segments (segment VIII on the anterior is slightly > ½
width of the widest portion of the abdomen). Segments VIII–X converge to the apex
which is broad and rounded. Genitalia. Subgenital plate starts at the anterior margin
of segment VIII, is moderately broad, and extends ca. ⅔ of the way onto segment X,
ending in a ne point (Fig. 47E). Gonapophyses VIII are long and moderately broad,
with their tips notably exceeding the apex of the abdomen, and slightly shorter than
the tips of the cerci; gonapophyses IX are slender and long, extending ca. ¾ of the
way onto segment X (Fig. 47E). Cerci at, not strongly cupped, with a heavily granu-
lar surface (Fig. 47E). Legs. Profemoral exterior lobe broad (broader than the interior
lobe), approximately right angled, and with a rounded exterior angle. e margin is
marked by 7–9 small serrate teeth throughout the length, none prominent (Fig. 47C).
Profemoral interior lobe narrower than the exterior lobe (only ca. 3× the greatest width
of the profemoral shaft) and with a slightly obtuse angle giving the interior lobe a tri-
angular appearance marked by ve teeth on the distal margin (Fig. 47C). ese teeth
are arranged in a two-one-two pattern with the exterior pairs closer together and with
a shallow gap between them, and the gap to the center tooth is deeper and wider than
these exterior pairs (Fig. 47C). Mesofemoral exterior lobe arcs from end to end as a
rounded triangle, but is slightly weighted towards the distal ½ and marked with two
or three small serrate teeth distributed on the distal half only. Mesofemoral exterior
lobe is slightly wider than the interior lobe. Mesofemoral interior lobe arcs end to end
more evenly than the exterior lobe, is marked with ve or six small serrate teeth only
on the distal half of the arc. Metafemoral interior lobe arcs end to end but is signi-
cantly more heavily weighted on the distal half and has six or seven serrate teeth on the
broader distal half of the lobe. Metafemoral exterior lobe is thin and smooth, hugging
the metafemoral shaft. Protibiae interior lobe spans the entire length of the protibiae
and is at its widest ca. 2× the width of the protibiae shaft itself. e lobe is distinctly
triangular and widest on the distal half. ere is a small but notable exterior protibial
lobe on the distal quarter of the length, but this is only ca. ½ as wide as the protibial
shaft itself. Mesotibiae simple, lacking lobes completely. Metatibiae lacks an interior
lobe, but does have a small but notable anteriorly situated exterior lobe.
Measurements of holotype female [mm]. Length of body (including cerci and
head, excluding antennae) 88.7, length/width of head 7.2/7.1, antennae 4.6, prono-
tum 5.3, mesonotum 7.5, length of tegmina 52.3, length of alae 41.5, greatest width of
Cryptophyllium, the hidden leaf insects; description of a new leaf insect genus 111
abdomen 37.9, profemora 19.7, mesofemora 15.0, metafemora 17.2, protibiae 12.6,
mesotibiae 11.4, metatibiae 15.2.
Measurements of paratype females [mm]. Length of body (including cerci and
head, excluding antennae) 90.8–92.0, length/width of head 7.5–7.7/6.6–7.3, anten-
nae 4.9–5.3, pronotum 5.5–5.6, mesonotum 8.0–8.4, length of tegmina 50.5–53.8,
length of alae 41.0–43.5, greatest width of abdomen 35.9–39.5, profemora 19.9–21.1,
mesofemora 15.1–16.0, metafemora 17.6–18.0, protibiae 12.0–12.6, mesotibiae
10.8–11.0, metatibiae 15.6–15.9.
Description of egg (Fig. 8K, L). e entire capsule is covered in short moss-like
pinnae and pitting of various size, depth, and spacing. e dorsal, ventral, and lateral
surfaces are attened, giving the egg a rectangular appearance. When viewed from the
lateral aspect, the egg has an almost uniform width throughout. e lateral margins
have slightly longer pinnae than the faces, but not drastically. Lateral surfaces slightly
raised along the center of the length of the egg, and the entire surface has various shallow
pitting in no detectable pattern. Micropylar plate spans the entire dorsal surface, with
the thickest portion near the posterior ⅓ around the micropylar cup. e remainder of
the micropylar plate is narrower, but not thin, at the thinnest still ca. ½ of the width of
the widest portion. Operculum ovular and shallowly raised in the center and a singular
circle of eight or nine shallow pits around the margin. Overall color light tan to brown.
Measurements including the extended pinnae [mm]. Length (including oper-
culum) 4.8–5.0, maximum width of capsule when viewed from lateral aspect 3.5–
3.6mm, length of micropylar plate 3.9–4.0 mm.
Etymology. Patronym. e type specimens for this species were discovered by
Chengzhi Bian (China) who recognized the scientic importance of the specimens
and shared them with the authors to review. Chengzhi Bian has decided to name this
species after his mother Liyanan to thank her for her amazing support of his passion
for entomology.
Cryptophyllium nuichuaense gen. et sp. nov.
http://zoobank.org/A801EC6F-5F97-4C17-A338-002FA9C9F112
Figures 6E, 48, 49
Material examined. Holotype ♀: “Coll. I.R.Sc.N.B., Vietnam, Ninh uan prov.,
Nui Chua N. P., 11°42’N 109°09’E, 3-9.VII.2014, night coll. Leg. J. Constant and
J. Bresseel, GTI project I.G.:32.779, DNA PH006”. Deposited in the Royal Belgian
Institute of Natural Sciences (RBINS).
Paratype: 1 ♀, “Coll. I.R.Sc.N.B., Vietnam, Ninh uan prov., Nui Chua N. P.,
11°42’N 109°09’E, 3-9.VII.2014, night coll. Leg. J. Constant and J. Bresseel, GTI
project I.G.:32.779” (VNMN).
Remarks. is species was collected on a night walk in 2014 within Nui Chua
National Park by Jérôme Constant (RBINS) and Joachim Bresseel (RBINS) (Fig. 48).
Royce T. Cumming et al. / ZooKeys 1018: 1–179 (2021)
112
Unfortunately, only two females were found, and this species could not be brought
into breeding to reveal details about the egg, male, or freshly hatched nymph morphol-
ogy. is is one of the many apparently highly endemic species within southern Viet-
nam, and we hope that future expeditions to this region reveal the male morphology
and the range of this species with more clarity.
Dierentiation. Females are morphologically most similar to Cryptophyllium pha-
mi sp. nov. and Cryptophyllium bollensi sp. nov. due to their general femoral lobe shape,
abdominal shape, femoral lobe spination, and thorax spination. When ner details are
observed however these species can easily be dierentiated. Cryptophyllium nuichuaense
sp. nov. have slightly shorter alae which only reach onto the anterior of abdominal seg-
ment III, whereas the other species have longer alae reaching to the middle of segment
III or even to the anterior margin of segment IV. Additionally, the antennae readily
dierentiate these three species as Cryptophyllium nuichuaense sp. nov. has antennal
segments V, VI, and VII with ventral margins which project past the margin of seg-
ment VIII, giving the antennae a slightly lamellate appearance (Fig. 6E), vs. the other
two species which have these segments ventral margin ush with the ventral margin of
segment VIII (Fig. 6B, F).
Males are presently unknown, but due to the adult morphology we expect that
they likely look similar to Cryptophyllium phami sp. nov. and Cryptophyllium bollensi
sp. nov. males.
Distribution. At present only known from Nui Chua N.P., in Ninh uan Prov-
ince, Vietnam.
Description. Female. Coloration. Coloration description is based upon the liv-
ing type material (Fig. 48). Overall coloration is pale green throughout. e antennae,
compound eyes, interior profemoral lobe margin, small patches along the protibial
interior lobe, and the anterior of the prescutum margins are orange to red, but these
areas are only sparsely marked.
Morphology. Head. Head capsule about as long as wide, vertex relatively smooth
with the only notable feature being the posteromedial tubercle which is nely point-
ed (Fig. 49E). Frontal convexity broad and blunt, with a slightly granular surface.
Compound eyes slightly protruding from the head capsule, and are not particularly
large, taking up slightly < ¼ of the head capsule lateral margins (Fig. 49E). Ocelli
absent. Antennal elds slightly wider than the width of the rst antennomere. An-
tennae. Antennae consisting of nine segments, with the terminal segment about the
same length as the preceding two segments’ lengths combined (Fig. 49C). Anten-
nomeres I–VII sparsely marked with small transparent setae, the terminal anten-
nomere and antennomere VIII are covered in stout, brown setae (Fig.49C). Anten-
nomeres V–VII ventral margins project farther than the ventral margin of segment
VIII, therefore giving the antennae a slight lamelatte appearance (Fig. 6E). orax.
P