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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.1018.61033" ID-GBIF-Dataset="6b752569-7f6e-4e32-8e0b-f7dc8b3711d4" ID-PMC="PMC7907054" ID-Pensoft-Pub="1313-2970-1018-1" ID-Pensoft-UUID="84B0D9BEE71D5171B80C3F4CBFDC7366" ID-PubMed="33664609" ID-ZooBank="7E9360A5A359437A91C004C74B1FE9D6" ModsDocID="1313-2970-1018-1" checkinTime="1613747250157" checkinUser="pensoft" docAuthor="Cumming, Royce T., Bank, Sarah, Bresseel, Joachim, Constant, Je ́ ro ̂ me, Tirant, Stephane Le, Dong, Zhiwei, Sonet, Gontran &amp; Bradler, Sven" docDate="2021" docId="B7FA223E233555279DF4863A17B08329" docLanguage="en" docName="ZooKeys 1018: 1-179" docOrigin="ZooKeys 1018" docPubDate="2021-02-18" docSource="http://dx.doi.org/10.3897/zookeys.1018.61033" docTitle="Cryptophyllium echidna gen. et 2021, sp. nov." docType="treatment" docUuid="076CABC1-2772-4E56-9918-F3804B62356E" docUuidSource="ZooBank" docVersion="5" id="84B0D9BEE71D5171B80C3F4CBFDC7366" lastPageNumber="1" masterDocId="84B0D9BEE71D5171B80C3F4CBFDC7366" masterDocTitle="Cryptophyllium, the hidden leaf insects - descriptions of a new leaf insect genus and thirteen species from the former celebicum species group (Phasmatodea, Phylliidae)" masterLastPageNumber="179" masterPageNumber="1" pageNumber="1" updateTime="1668149808794" updateUser="ExternalLinkService">
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<mods:titleInfo>
<mods:title>Cryptophyllium, the hidden leaf insects - descriptions of a new leaf insect genus and thirteen species from the former celebicum species group (Phasmatodea, Phylliidae)</mods:title>
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<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Cumming, Royce T.</mods:namePart>
<mods:nameIdentifier type="ORCID">https://orcid.org/0000-0001-7930-1292</mods:nameIdentifier>
<mods:affiliation>Montreal Insectarium, 4581 rue Sherbrooke est, Montreal, Quebec, Canada, H 1 X 2 B 2 &amp; Richard Gilder Graduate School, American Museum of Natural History, New York, NY 10024, USA &amp; Biology, Graduate Center, City University of New York, NY, USA</mods:affiliation>
<mods:nameIdentifier type="email">roycecumming@gmail.com</mods:nameIdentifier>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Bank, Sarah</mods:namePart>
<mods:nameIdentifier type="ORCID">https://orcid.org/0000-0001-6952-1590</mods:nameIdentifier>
<mods:affiliation>Department of Animal Evolution and Biodiversity, Johann-Friedrich-Blumenbach Institute for Zoology and Anthropology, University of Go ̈ ttingen, Untere Karspu ̈ le 2, 37073, Go ̈ ttingen, Germany</mods:affiliation>
<mods:nameIdentifier type="email">sarah.bank@uni-goettingen.de</mods:nameIdentifier>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Bresseel, Joachim</mods:namePart>
<mods:affiliation>Royal Belgian Institute of Natural Sciences, O. D. Taxonomy and Phylogeny and JEMU, rue Vautier 29, B- 1000, Brussels, Belgium</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Constant, Je ́ ro ̂ me</mods:namePart>
<mods:affiliation>Royal Belgian Institute of Natural Sciences, O. D. Taxonomy and Phylogeny and JEMU, rue Vautier 29, B- 1000, Brussels, Belgium</mods:affiliation>
</mods:name>
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<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Tirant, Stephane Le</mods:namePart>
<mods:affiliation>Montreal Insectarium, 4581 rue Sherbrooke est, Montreal, Quebec, Canada, H 1 X 2 B 2</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Dong, Zhiwei</mods:namePart>
<mods:affiliation>State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, Yunnan, 650223, China</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Sonet, Gontran</mods:namePart>
<mods:affiliation>Royal Belgian Institute of Natural Sciences, O. D. Taxonomy and Phylogeny and JEMU, rue Vautier 29, B- 1000, Brussels, Belgium</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Bradler, Sven</mods:namePart>
<mods:nameIdentifier type="ORCID">https://orcid.org/0000-0001-9307-1032</mods:nameIdentifier>
<mods:affiliation>Department of Animal Evolution and Biodiversity, Johann-Friedrich-Blumenbach Institute for Zoology and Anthropology, University of Go ̈ ttingen, Untere Karspu ̈ le 2, 37073, Go ̈ ttingen, Germany</mods:affiliation>
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<mods:date>2021</mods:date>
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<mods:number>2021-02-18</mods:number>
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<mods:number>1018</mods:number>
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<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.1018.61033</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-1018-1</mods:identifier>
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<treatment ID-GBIF-Taxon="177429200" LSID="urn:lsid:zoobank.org:act:076CABC1-2772-4E56-9918-F3804B62356E" httpUri="http://treatment.plazi.org/id/B7FA223E233555279DF4863A17B08329" lastPageNumber="1" pageId="0" pageNumber="1">
<subSubSection pageId="0" pageNumber="1" type="nomenclature">
<paragraph pageId="0" pageNumber="1">
<taxonomicName LSID="http://zoobank.org/076CABC1-2772-4E56-9918-F3804B62356E" authority="gen. et" authorityName="gen. et" authorityYear="2021" class="Insecta" family="Phylliidae" genus="Cryptophyllium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cryptophyllium echidna" order="Phasmida" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="echidna" status="sp. nov.">Cryptophyllium echidna gen. et</taxonomicName>
<taxonomicNameLabel pageId="0" pageNumber="1">sp. nov.</taxonomicNameLabel>
<figureCitation captionStart="Figure 30" captionStartId="F30" captionText="Figure 30. Cryptophyllium echidna gen. et sp. nov. holotype female, photographs by Rene ́ 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." figureDoi="10.3897/zookeys.1018.61033.figure30" httpUri="https://binary.pensoft.net/fig/510494" pageId="0" pageNumber="1">Figure 30</figureCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="materials_examined">
<paragraph pageId="0" pageNumber="1">Material examined.</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" italics="true" pageId="0" pageNumber="1">Holotype</emphasis>
♀: INDONESIA: Wangi-wangi Island. Collected prior to 2020, but no exact date given. Deposited in the Montreal Insectarium (IMQC).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="remarks">
<paragraph pageId="0" pageNumber="1">Remarks.</paragraph>
<paragraph pageId="0" pageNumber="1">
This is the first phylliid record we have seen from the small island of Wangi-wangi in the Wakatobi Regency in Southeast Sulawesi Province. This small island appears to be rather unique biogeographically and unexplored as a yet to be described bird species presently known as the &quot;Wangi-wangi White-eye&quot; (
<taxonomicName class="Insecta" family="Zosteropidae" genus="Zosterops" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zosterops" order="Phasmida" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="1">Zosterops</emphasis>
</taxonomicName>
sp. nov.) has recently been identified as well, suggesting this island may hold many endemic undescribed species (
<bibRefCitation DOI="https://doi.org/10.1093/zoolinnean/zlz022" author="O'Connell, DP" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="0" pageNumber="1" pagination="701 - 724" refId="B48" refString="O'Connell, DP, Kelly, DJ, Lawless, N, O'Brien, K, O Marcaigh, F, Karya, A, Analuddin, K, Marples, NM, 2019. A sympatric pair of undescribed white-eye species (Aves: Zosteropidae: Zosterops) with different origins. Zoological Journal of the Linnean Society 186: 701 - 724, DOI: https://doi.org/10.1093/zoolinnean/zlz022" title="A sympatric pair of undescribed white-eye species (Aves: Zosteropidae: Zosterops) with different origins." url="https://doi.org/10.1093/zoolinnean/zlz022" volume="186" year="2019">
<normalizedToken originalValue="OConnell">O'Connell</normalizedToken>
et al. 2019
</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="differentiation">
<paragraph pageId="0" pageNumber="1">Differentiation.</paragraph>
<paragraph pageId="0" pageNumber="1">
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
<taxonomicName authorityName="Cumming &amp; Bank &amp; Bresseel &amp; Constant &amp; Tirant &amp; Dong &amp; Sonet &amp; Bradler" authorityYear="2021" baseAuthorityName="de Haan" baseAuthorityYear="1842" class="Insecta" family="Phylliidae" genus="Cryptophyllium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cryptophyllium celebicum" order="Phasmida" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="celebicum">
<emphasis italics="true" pageId="0" pageNumber="1">Cryptophyllium celebicum</emphasis>
</taxonomicName>
comb. nov. which has a much wider distribution to the north on the islands of Buton, Sulawesi, and Peleng (Fig.
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="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 / differentiate (indicated by dashed lines). Note that the line for Cryptophyllium westwoodii comb. nov. is pointing to the neotype locality and the type locality for Cryptophyllium athanysus comb. nov. is simply &quot; Ceylon &quot; 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 fit within the main map. The colors in this map are noted to the left of the names within the phylogenetic tree in Fig. 4. Newly herein described species have names noted in bold. Note that with Cryptophyllium 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 Thai offshore islands of Ko Phangan and Ko Samui which could not be differentiated 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 specimens and links to observational records&gt;." figureDoi="10.3897/zookeys.1018.61033.figure2" httpUri="https://binary.pensoft.net/fig/510466" pageId="0" pageNumber="1">2</figureCitation>
).
</paragraph>
<paragraph pageId="0" pageNumber="1">
Female
<taxonomicName authorityName="gen. et" authorityYear="2021" class="Insecta" family="Phylliidae" genus="Cryptophyllium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cryptophyllium echidna" order="Phasmida" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="echidna">
<emphasis italics="true" pageId="0" pageNumber="1">Cryptophyllium echidna</emphasis>
</taxonomicName>
sp. nov. can be differentiated from
<taxonomicName authorityName="Cumming &amp; Bank &amp; Bresseel &amp; Constant &amp; Tirant &amp; Dong &amp; Sonet &amp; Bradler" authorityYear="2021" baseAuthorityName="de Haan" baseAuthorityYear="1842" class="Insecta" family="Phylliidae" genus="Cryptophyllium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cryptophyllium celebicum" order="Phasmida" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="celebicum">
<emphasis italics="true" pageId="0" pageNumber="1">Cryptophyllium celebicum</emphasis>
</taxonomicName>
comb. nov. by only subtle differences in the thorax and profemoral exterior lobes. In
<taxonomicName authorityName="gen. et" authorityYear="2021" class="Insecta" family="Phylliidae" genus="Cryptophyllium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cryptophyllium echidna" order="Phasmida" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="echidna">
<emphasis italics="true" pageId="0" pageNumber="1">Cryptophyllium echidna</emphasis>
</taxonomicName>
sp. nov. the prescutum is slightly broader and with a weaker sagittal crest (Fig.
<figureCitation captionStart="Figure 30" captionStartId="F30" captionText="Figure 30. Cryptophyllium echidna gen. et sp. nov. holotype female, photographs by Rene ́ 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." figureDoi="10.3897/zookeys.1018.61033.figure30" httpUri="https://binary.pensoft.net/fig/510494" pageId="0" pageNumber="1">30D</figureCitation>
) than in
<taxonomicName authorityName="Cumming &amp; Bank &amp; Bresseel &amp; Constant &amp; Tirant &amp; Dong &amp; Sonet &amp; Bradler" authorityYear="2021" baseAuthorityName="de Haan" baseAuthorityYear="1842" class="Insecta" family="Phylliidae" genus="Cryptophyllium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cryptophyllium celebicum" order="Phasmida" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="celebicum">
<emphasis italics="true" pageId="0" pageNumber="1">Cryptophyllium celebicum</emphasis>
</taxonomicName>
comb. nov. (Fig.
<figureCitation captionStart="Figure 22" captionStartId="F22" captionText="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." figureDoi="10.3897/zookeys.1018.61033.figure22" httpUri="https://binary.pensoft.net/fig/510486" pageId="0" pageNumber="1">22E</figureCitation>
). Additionally the profemoral exterior lobe of
<taxonomicName authorityName="gen. et" authorityYear="2021" class="Insecta" family="Phylliidae" genus="Cryptophyllium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cryptophyllium echidna" order="Phasmida" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="echidna">
<emphasis italics="true" pageId="0" pageNumber="1">Cryptophyllium echidna</emphasis>
</taxonomicName>
sp. nov. has a right exterior angle (Fig.
<figureCitation captionStart="Figure 30" captionStartId="F30" captionText="Figure 30. Cryptophyllium echidna gen. et sp. nov. holotype female, photographs by Rene ́ 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." figureDoi="10.3897/zookeys.1018.61033.figure30" httpUri="https://binary.pensoft.net/fig/510494" pageId="0" pageNumber="1">30A</figureCitation>
), not acute like in
<taxonomicName authorityName="Cumming &amp; Bank &amp; Bresseel &amp; Constant &amp; Tirant &amp; Dong &amp; Sonet &amp; Bradler" authorityYear="2021" baseAuthorityName="de Haan" baseAuthorityYear="1842" class="Insecta" family="Phylliidae" genus="Cryptophyllium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cryptophyllium celebicum" order="Phasmida" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="celebicum">
<emphasis italics="true" pageId="0" pageNumber="1">Cryptophyllium celebicum</emphasis>
</taxonomicName>
comb. nov. (Fig.
<figureCitation captionStart="Figure 22" captionStartId="F22" captionText="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." figureDoi="10.3897/zookeys.1018.61033.figure22" httpUri="https://binary.pensoft.net/fig/510486" pageId="0" pageNumber="1">22D</figureCitation>
).
</paragraph>
<paragraph pageId="0" pageNumber="1">
Males are presently unknown, but as the sister species to
<taxonomicName authorityName="Cumming &amp; Bank &amp; Bresseel &amp; Constant &amp; Tirant &amp; Dong &amp; Sonet &amp; Bradler" authorityYear="2021" baseAuthorityName="de Haan" baseAuthorityYear="1842" class="Insecta" family="Phylliidae" genus="Cryptophyllium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cryptophyllium celebicum" order="Phasmida" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="celebicum">
<emphasis italics="true" pageId="0" pageNumber="1">Cryptophyllium celebicum</emphasis>
</taxonomicName>
comb. nov. the males likely have a similar morphology.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="distribution">
<paragraph pageId="0" pageNumber="1">Distribution.</paragraph>
<paragraph pageId="0" pageNumber="1">At present only known from the small Indonesian island of Wangi-wangi off the east coast of Buton Island.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="description">
<paragraph pageId="0" pageNumber="1">Description.</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">
Female.
<emphasis italics="true" pageId="0" pageNumber="1">Coloration</emphasis>
.
</emphasis>
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. The rotten areas are brown but are assumed to have been green in life. The remainder of the female is lime-green in color throughout, with no indication of natural brown patches (which even on somewhat rotten specimens can generally be identified) but this female appears to have been uniform green in life.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" italics="true" pageId="0" pageNumber="1">Morphology.</emphasis>
<emphasis italics="true" pageId="0" pageNumber="1">Head.</emphasis>
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. The 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 first 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.
<normalizedToken originalValue="¼">1/4</normalizedToken>
of the length of the lateral head capsule margins (Fig.
<figureCitation captionStart="Figure 30" captionStartId="F30" captionText="Figure 30. Cryptophyllium echidna gen. et sp. nov. holotype female, photographs by Rene ́ 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." figureDoi="10.3897/zookeys.1018.61033.figure30" httpUri="https://binary.pensoft.net/fig/510494" pageId="0" pageNumber="1">30A</figureCitation>
). Ocelli absent. Antennal fields slightly wider than and about as long as the length of the first antennomere.
<emphasis italics="true" pageId="0" pageNumber="1">Antennae.</emphasis>
Antennae consist of nine segments, with the terminal segment approximately the same length as the preceding two
<normalizedToken originalValue="segments">segments'</normalizedToken>
lengths combined (Fig.
<figureCitation captionStart="Figure 30" captionStartId="F30" captionText="Figure 30. Cryptophyllium echidna gen. et sp. nov. holotype female, photographs by Rene ́ 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." figureDoi="10.3897/zookeys.1018.61033.figure30" httpUri="https://binary.pensoft.net/fig/510494" pageId="0" pageNumber="1">30B</figureCitation>
). Antennomeres I-III sparsely marked with small transparent setae (with the longest on the first segment), the terminal two antennomeres are densely covered in stout, brown setae.
<emphasis italics="true" pageId="0" pageNumber="1">Thorax.</emphasis>
Pronotum with a 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.
<figureCitation captionStart="Figure 30" captionStartId="F30" captionText="Figure 30. Cryptophyllium echidna gen. et sp. nov. holotype female, photographs by Rene ́ 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." figureDoi="10.3897/zookeys.1018.61033.figure30" httpUri="https://binary.pensoft.net/fig/510494" pageId="0" pageNumber="1">30D</figureCitation>
). The 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.
<figureCitation captionStart="Figure 30" captionStartId="F30" captionText="Figure 30. Cryptophyllium echidna gen. et sp. nov. holotype female, photographs by Rene ́ 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." figureDoi="10.3897/zookeys.1018.61033.figure30" httpUri="https://binary.pensoft.net/fig/510494" pageId="0" pageNumber="1">30D</figureCitation>
). The pronotum has a prominent anterior rim and moderate lateral and posterior rims (Fig.
<figureCitation captionStart="Figure 30" captionStartId="F30" captionText="Figure 30. Cryptophyllium echidna gen. et sp. nov. holotype female, photographs by Rene ́ 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." figureDoi="10.3897/zookeys.1018.61033.figure30" httpUri="https://binary.pensoft.net/fig/510494" pageId="0" pageNumber="1">30D</figureCitation>
). 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.
<figureCitation captionStart="Figure 30" captionStartId="F30" captionText="Figure 30. Cryptophyllium echidna gen. et sp. nov. holotype female, photographs by Rene ́ 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." figureDoi="10.3897/zookeys.1018.61033.figure30" httpUri="https://binary.pensoft.net/fig/510494" pageId="0" pageNumber="1">30D</figureCitation>
). Lateral rims with 6-8 prominent tubercles with various small, lumpy granules interspersed throughout the margins (Fig.
<figureCitation captionStart="Figure 30" captionStartId="F30" captionText="Figure 30. Cryptophyllium echidna gen. et sp. nov. holotype female, photographs by Rene ́ 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." figureDoi="10.3897/zookeys.1018.61033.figure30" httpUri="https://binary.pensoft.net/fig/510494" pageId="0" pageNumber="1">30D</figureCitation>
). Prescutum anterior rim prominent but not strongly protruding, surface is granular and lacks a prominent sagittal spine (Fig.
<figureCitation captionStart="Figure 30" captionStartId="F30" captionText="Figure 30. Cryptophyllium echidna gen. et sp. nov. holotype female, photographs by Rene ́ 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." figureDoi="10.3897/zookeys.1018.61033.figure30" httpUri="https://binary.pensoft.net/fig/510494" pageId="0" pageNumber="1">30D</figureCitation>
). Prescutum surface without a strongly raised sagittal crest, instead the surface is only slightly raised along the sagittal plane. The prescutum surface has moderate granulation throughout ranging in size from small to medium with irregular spacing (Fig.
<figureCitation captionStart="Figure 30" captionStartId="F30" captionText="Figure 30. Cryptophyllium echidna gen. et sp. nov. holotype female, photographs by Rene ́ 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." figureDoi="10.3897/zookeys.1018.61033.figure30" httpUri="https://binary.pensoft.net/fig/510494" pageId="0" pageNumber="1">30D</figureCitation>
). 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 five or six larger, sharp tipped tubercles with an additional five or six smaller nodes interspersed throughout (Fig.
<figureCitation captionStart="Figure 30" captionStartId="F30" captionText="Figure 30. Cryptophyllium echidna gen. et sp. nov. holotype female, photographs by Rene ́ 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." figureDoi="10.3897/zookeys.1018.61033.figure30" httpUri="https://binary.pensoft.net/fig/510494" pageId="0" pageNumber="1">30D</figureCitation>
). Face of the mesopleura lacking granulation, but instead highly wrinkled and with two notable pits, one on the anterior ⅓ and one nearer the posterior ⅓ (Fig.
<figureCitation captionStart="Figure 30" captionStartId="F30" captionText="Figure 30. Cryptophyllium echidna gen. et sp. nov. holotype female, photographs by Rene ́ 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." figureDoi="10.3897/zookeys.1018.61033.figure30" httpUri="https://binary.pensoft.net/fig/510494" pageId="0" pageNumber="1">30D</figureCitation>
).
<emphasis italics="true" pageId="0" pageNumber="1">Wings.</emphasis>
Tegmina long, reaching nearly to the posterior margin of abdominal segment VII. The subcosta (Sc) is the first vein in the forewing and runs subparallel with the wing for the first half of its length, and then bends towards the wing margin for the second half. The radius (R) spans the central portion of the tegmina with two subparallel branched veins. The first radius (R1) branches ca. ⅗ of the way through the radius length and terminates ca. ⅓ of the way through the wing length. The 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. There 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). The media (M) is simply bifurcate with both the media anterior (MA) and media posterior (MP) terminating close to the posterior
<normalizedToken originalValue="¼">1/4</normalizedToken>
of the wing. The 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. The first 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.
<emphasis italics="true" pageId="0" pageNumber="1">Abdomen.</emphasis>
Abdominal segments II through the anterior
<normalizedToken originalValue="½">1/2</normalizedToken>
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.
<emphasis italics="true" pageId="0" pageNumber="1">Genitalia.</emphasis>
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 fine point (Fig.
<figureCitation captionStart="Figure 30" captionStartId="F30" captionText="Figure 30. Cryptophyllium echidna gen. et sp. nov. holotype female, photographs by Rene ́ 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." figureDoi="10.3897/zookeys.1018.61033.figure30" httpUri="https://binary.pensoft.net/fig/510494" pageId="0" pageNumber="1">30E</figureCitation>
). Gonapophyses VIII are long (exceeding the tip of 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.
<figureCitation captionStart="Figure 30" captionStartId="F30" captionText="Figure 30. Cryptophyllium echidna gen. et sp. nov. holotype female, photographs by Rene ́ 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." figureDoi="10.3897/zookeys.1018.61033.figure30" httpUri="https://binary.pensoft.net/fig/510494" pageId="0" pageNumber="1">30E</figureCitation>
). Gonapophyses IX are smaller and shorter, and mostly covered from view by the notably larger gonapophyses VIII. Cerci flat, with a granular surface throughout and few setae on the ventral surface, the dorsal surface has thin, transparent setae throughout the surface (Fig.
<figureCitation captionStart="Figure 30" captionStartId="F30" captionText="Figure 30. Cryptophyllium echidna gen. et sp. nov. holotype female, photographs by Rene ́ 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." figureDoi="10.3897/zookeys.1018.61033.figure30" httpUri="https://binary.pensoft.net/fig/510494" pageId="0" pageNumber="1">30E</figureCitation>
).
<emphasis italics="true" pageId="0" pageNumber="1">Legs.</emphasis>
Profemoral exterior lobes broad (ca. 2
<normalizedToken originalValue="×">x</normalizedToken>
width of the interior lobe) and approximately right angled (Fig.
<figureCitation captionStart="Figure 30" captionStartId="F30" captionText="Figure 30. Cryptophyllium echidna gen. et sp. nov. holotype female, photographs by Rene ́ 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." figureDoi="10.3897/zookeys.1018.61033.figure30" httpUri="https://binary.pensoft.net/fig/510494" pageId="0" pageNumber="1">30A</figureCitation>
). 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.
<figureCitation captionStart="Figure 30" captionStartId="F30" captionText="Figure 30. Cryptophyllium echidna gen. et sp. nov. holotype female, photographs by Rene ́ 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." figureDoi="10.3897/zookeys.1018.61033.figure30" httpUri="https://binary.pensoft.net/fig/510494" pageId="0" pageNumber="1">30A</figureCitation>
). Profemoral interior lobe with a slightly obtuse angle and marked with five large, serrate teeth with looping gaps between them. These 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 off 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.
<normalizedToken originalValue="¼">1/4</normalizedToken>
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 notably 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.
</paragraph>
<caption doi="10.3897/zookeys.1018.61033.figure30" httpUri="https://binary.pensoft.net/fig/510494" pageId="0" pageNumber="1" start="Figure 30" startId="F30">
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Figure 30.</emphasis>
<taxonomicName authorityName="gen. et" authorityYear="2021" class="Insecta" family="Phylliidae" genus="Cryptophyllium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cryptophyllium echidna" order="Phasmida" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="echidna">
<emphasis italics="true" pageId="0" pageNumber="1">Cryptophyllium echidna</emphasis>
</taxonomicName>
gen. et sp. nov. holotype female, photographs by René Limoges (IMQC)
<emphasis bold="true" pageId="0" pageNumber="1">A</emphasis>
habitus, dorsal
<emphasis bold="true" pageId="0" pageNumber="1">B</emphasis>
dorsal details of the antennae
<emphasis bold="true" pageId="0" pageNumber="1">C</emphasis>
habitus, ventral
<emphasis bold="true" pageId="0" pageNumber="1">D</emphasis>
dorsal details of the thorax
<emphasis bold="true" pageId="0" pageNumber="1">E</emphasis>
ventral details of the genitalia.
</paragraph>
</caption>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">
<emphasis italics="true" pageId="0" pageNumber="1">Measurements of holotype female</emphasis>
[mm].
</emphasis>
Length of body (including cerci and head, excluding antennae) 94.5, length/width of head 7.8/7.5, antennae 5.0, pronotum 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.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="etymology">
<paragraph pageId="0" pageNumber="1">Etymology.</paragraph>
<paragraph pageId="0" pageNumber="1">
Noun, Greek in origin. Relating to the tenth labor of Heracles (apparently a favorite story of
<bibRefCitation author="Gray, GR" journalOrPublisher="Zoologist" pageId="0" pageNumber="1" pagination="117 - 123" refId="B25" refString="Gray, GR, 1843. Description of several species of the Genus Phyllium. Zoologist 1: 117 - 123" title="Description of several species of the Genus Phyllium." volume="1" year="1843">Gray (1843)</bibRefCitation>
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
<taxonomicName class="Insecta" family="Phylliidae" genus="Phyllium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phyllium geryon" order="Orthoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="geryon">
<emphasis italics="true" pageId="0" pageNumber="1">Phyllium geryon</emphasis>
</taxonomicName>
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. Those three sons were Agathyrsus, Gelonus, and Scythes (for which Gray named
<taxonomicName class="Insecta" family="Phylliidae" genus="Phyllium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phyllium agathyrsus" order="Orthoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="agathyrsus">
<emphasis italics="true" pageId="0" pageNumber="1">Phyllium agathyrsus</emphasis>
</taxonomicName>
Gray, 1843;
<taxonomicName class="Insecta" family="Phylliidae" genus="Phyllium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phyllium gelonus" order="Orthoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="gelonus">
<emphasis italics="true" pageId="0" pageNumber="1">Phyllium gelonus</emphasis>
</taxonomicName>
Gray, 1843; and
<taxonomicName class="Insecta" family="Phylliidae" genus="Phyllium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phyllium scythe" order="Orthoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="scythe">
<emphasis italics="true" pageId="0" pageNumber="1">Phyllium scythe</emphasis>
</taxonomicName>
Gray, 1843).
</paragraph>
<paragraph pageId="0" pageNumber="1">
We felt that this species could help to finish telling the story which Gray was so fond of. With
<taxonomicName class="Insecta" family="Phylliidae" genus="Phyllium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phyllium geryon" order="Orthoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="geryon">
<emphasis italics="true" pageId="0" pageNumber="1">Phyllium geryon</emphasis>
</taxonomicName>
being a species from the Philippines, one biogeographical bridge for species to the Philippines is from Sulawesi through the Sangihe Islands (
<bibRefCitation DOI="https://doi.org/10.1080/10635150390251063" author="Evans, BJ" journalOrPublisher="Systematic Biology" pageId="0" pageNumber="1" pagination="794 - 819" refId="B23" refString="Evans, BJ, Brown, RM, McGuire, JA, Supriatna, J, Andayani, N, Diesmos, A, Iskandar, D, Melnick, D, Cannatella, D, 2003. Phylogenetics of fanged frogs: testing biogeographical hypotheses at the interface of the Asian and Australian faunal zones. Systematic Biology 52: 794 - 819, DOI: https://doi.org/10.1080/10635150390251063" title="Phylogenetics of fanged frogs: testing biogeographical hypotheses at the interface of the Asian and Australian faunal zones." url="https://doi.org/10.1080/10635150390251063" volume="52" year="2003">Evans et al. 2003</bibRefCitation>
), therefore as a steppingstone to the Philippines we felt there should be an Echidna along the route to the Philippines where
<taxonomicName class="Insecta" family="Phylliidae" genus="Phyllium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phyllium geryon" order="Orthoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="geryon">
<emphasis italics="true" pageId="0" pageNumber="1">Phyllium geryon</emphasis>
</taxonomicName>
can be found.
</paragraph>
</subSubSection>
</treatment>
</document>