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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.93.1167" ID-GBIF-Dataset="704a30f8-6065-4db2-828f-822d9cfa3082" ID-PMC="PMC3095181" ID-Pensoft-Pub="1313-2970-93-9" ID-PubMed="21594077" ModsDocAuthor="" ModsDocDate="2011" ModsDocID="1313-2970-93-9" ModsDocOrigin="ZooKeys 93" ModsDocTitle="The millipede family Polydesmidae in Taiwan, with descriptions of five new species (Polydesmida, Diplopoda)" checkinTime="1451250468082" checkinUser="pensoft" docAuthor="Golovatch, Sergei I., Mikhaljova, Elena V. &amp; Chang, Hsueh-Wen" docDate="2011" docId="43192597324A3CC494E846D9C06FCDC2" docLanguage="en" docName="ZooKeys 93: 9-42" docOrigin="ZooKeys 93" docSource="http://dx.doi.org/10.3897/zookeys.93.1167" docTitle="Epanerchodus orientalis Attems 1901" docType="treatment" docVersion="4" lastPageNumber="19" masterDocId="FFB1F12DFFDDFFEAFFF43B31FFF6B946" masterDocTitle="The millipede family Polydesmidae in Taiwan, with descriptions of five new species (Polydesmida, Diplopoda)" masterLastPageNumber="42" masterPageNumber="9" pageNumber="15" updateTime="1668168924641" updateUser="ExternalLinkService">
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<mods:titleInfo>
<mods:title>The millipede family Polydesmidae in Taiwan, with descriptions of five new species (Polydesmida, Diplopoda)</mods:title>
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<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Golovatch, Sergei I.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Mikhaljova, Elena V.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Chang, Hsueh-Wen</mods:namePart>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
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<mods:title>ZooKeys</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2011</mods:date>
<mods:detail type="volume">
<mods:number>93</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>9</mods:start>
<mods:end>42</mods:end>
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<mods:url>http://dx.doi.org/10.3897/zookeys.93.1167</mods:url>
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<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.93.1167</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-93-9</mods:identifier>
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<treatment ID-GBIF-Taxon="159364986" LSID="urn:lsid:plazi:treatment:43192597324A3CC494E846D9C06FCDC2" httpUri="http://treatment.plazi.org/id/43192597324A3CC494E846D9C06FCDC2" lastPageId="10" lastPageNumber="19" pageId="6" pageNumber="15">
<subSubSection pageId="6" pageNumber="15" type="nomenclature">
<paragraph pageId="6" pageNumber="15">
<taxonomicName LSID="http://species-id.net/wiki/Epanerchodus_orientalis" authority="Attems, 1901" authorityName="Attems" authorityYear="1901" class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus orientalis" order="Polydesmida" pageId="6" pageNumber="15" phylum="Arthropoda" rank="species" species="orientalis">Epanerchodus orientalis Attems, 1901</taxonomicName>
Figs 2436
</paragraph>
</subSubSection>
<subSubSection pageId="6" pageNumber="15" type="reference_group">
<paragraph pageId="6" pageNumber="15">
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus orientalis subsp. orientalis" order="Polydesmida" pageId="6" pageNumber="15" phylum="Arthropoda" rank="subSpecies" species="orientalis" subSpecies="orientalis">Epanerchodus orientalis orientalis</taxonomicName>
-
<bibRefCitation author="Wang, Y-HM" journalOrPublisher="Quarterly Journal of the Taiwan Museum" pageId="19" pageNumber="28" pagination="155 - 159" title="Serica 1 e: Records of myriapods on Formosa with description of new species (2)." volume="9" year="1956">Wang 1956</bibRefCitation>
: 155 (R);
<bibRefCitation author="Wang, Y-HM" journalOrPublisher="Quarterly Journal of the Taiwan Museum" pageId="19" pageNumber="28" pagination="89 - 96" title="Serica 1 q: Millipedes and centipedes of Quemoy, Fukien Province and Taiwan Island, Botel Tobago (Lan Yu), Taiwan Province and of Singapore." volume="16" year="1963">Wang 1963</bibRefCitation>
: 91 (R)
</paragraph>
<paragraph pageId="6" pageNumber="15">
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus orientalis subsp. takakuwai" order="Polydesmida" pageId="6" pageNumber="15" phylum="Arthropoda" rank="subSpecies" species="orientalis" subSpecies="takakuwai">Epanerchodus orientalis takakuwai</taxonomicName>
-
<bibRefCitation author="Wang, Y-HM" journalOrPublisher="Quarterly Journal of the Taiwan Museum" pageId="19" pageNumber="28" pagination="340 - 344" title="Serica 1 i: On Diplopoda from Taiwan with a new strongylosomids." volume="11" year="1958">Wang 1958</bibRefCitation>
: 341 (R)
</paragraph>
</subSubSection>
<subSubSection pageId="6" pageNumber="15" type="nb">
<paragraph pageId="6" pageNumber="15">NB:</paragraph>
<paragraph pageId="6" pageNumber="15">
Complete catalogue information referring to this species can be found in
<bibRefCitation author="Murakami, Y" journalOrPublisher="Bulletin of the National Science Museum" pageId="19" pageNumber="28" pagination="557 - 582" title="Myriapods found in limestone caves of northern Honshu, Japan." volume="12" year="1969">Murakami (1969)</bibRefCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="6" pageNumber="15" type="material examined">
<paragraph pageId="6" pageNumber="15">Material examined:</paragraph>
<paragraph pageId="6" pageNumber="15">10 ♂ (TFRI), 1 ♂ (NMNS-6557-001), 1 ♀ (NSYSU), 2 ♂ (MNHN JC 333), Taiwan, Taichung County, Shengguang, 25.04.2003; 2 ♀ (TFRI), same locality, 26.03.2003; 1 ♀ (TFRI), same locality, 24.09.2002; 4 ♂ (TFRI), 1 ♂ (NMNS-6557-002), same locality, 24.07.2003, all leg. W.C. Yeh; 2 ♂, 1 ♀ (TFRI), 2 ♂, 1 ♀ (IBSS), Ilan County, Fushan, 23.03.2003; 2 ♀ (NMNS-6557-003), same locality, 18.05.2001; 5 ♂, 1 ♀ (TFRI), same locality, 19.06.2001; 1 ♂ (TFRI), same locality, without date, all leg. J.T. Chao; 1 ♂ (ZMUM), same locality, 20.11.2001, leg. S.S. Lu; 52 ♂, 17 ♀, 4 fragm. (TFRI), 2 ♂ (ZMUC), 2 ♂ (NSYSU), Ilan County, Tatong Township, near Lakes Jialuohu, ca 2,250 m, 4.06.2003; 1 ♀ fragm. (TFRI), same locality, 28.04.2003; 8 ♂, 6 ♀ (TFRI), same locality, 20.06.2002; 1 ♂ (ZMUM), same locality, 25.04.2003; 1 ♂ (TFRI), same locality, 23.07.2002, all leg. Y.M. Chen; 1 ♀ (NSYSU), Taitung County, Yanping Township, Yanping Forest road, ca. 1,250 m, 3.06.2003, leg. M.H. Hsu; 2 ♂ (NSYSU), Hualien County, Fongbin Township, Guangfong Highway, ca 300 m, 6.05.2009; 1 ♀ (NSYSU), same County, Shoufang Township, ca. 230 m a.s.l., 5.05.2009, all leg. M.H. Shu; 2 ♀ (NMNS-6557-004), Pingtung County, Chaozhou Township, Si-Lin, 1.06.1999, leg.?; 1 ♀ (NMNS-6557-005), Kaohsiung County, Taoyuan, Provincial Road No. 20, near 142.5 km road-sign, 3.09.2009, leg. C.Y. Huang.</paragraph>
</subSubSection>
<subSubSection pageId="6" pageNumber="15" type="diagnosis">
<paragraph pageId="6" pageNumber="15">Diagnosis:</paragraph>
<paragraph pageId="6" pageNumber="15">
Differs from the other
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus" order="Polydesmida" pageId="6" pageNumber="15" phylum="Arthropoda" rank="genus">Epanerchodus</taxonomicName>
species known from Taiwan in the variable, mostly medium size, coupled with the caudal corner of most of the paraterga being pointed, and in the gonopod telopodite being relatively slender, complex, highly variable in shape and armature (see also Key below).
</paragraph>
</subSubSection>
<subSubSection lastPageId="8" lastPageNumber="17" pageId="6" pageNumber="15" type="redescription">
<paragraph pageId="6" pageNumber="15">Redescription:</paragraph>
<paragraph lastPageId="7" lastPageNumber="16" pageId="6" pageNumber="15">
Length of both sexes ca 11-19 mm; width of pro- and metazona varying between specimens from 0.8-1.8 to 1.2-3.0 mm, respectively. Usually ♂♂ somewhat smaller than ♀♀. Coloration in alcohol from uniformly pallid (faded?) to light (reddish- to grey-) brown; in the latter case, venter and legs light grey to yellow (Figs 24-31). Body with 20 segments. Tegument mainly dull, at most slightly shining,
<pageBreakToken pageId="7" pageNumber="16" start="start">texture</pageBreakToken>
very delicately alveolate. Labrum, frons and vertex densely pilose; epicranial suture clear but thin; no ridges above antennal sockets; isthmus between antennae considerably broader than diameter of antennal socket (Fig. 29). Antennae long and only slightly clavate (Figs 24, 25, 28), reaching behind end (♂) or midway (♀) of segment 4 dorsally; antennomere 3 longest, 5th highest (Figs 25, 28); antennomeres 5 and 6 each with a small, compact, distodorsal group of bacilliform sensilla; antennomere 7 with a minute dorsoparabasal cone and a distodorsal group of microscopic sensilla.
</paragraph>
<paragraph pageId="7" pageNumber="16">In width, either collum ≤ segment 2 = 3 &lt;4 &lt;head = 5(6)-15(16), or segment 2 = 3 &lt;collum &lt;head = 4 &lt;5-15(16) (both sexes), thereafter body gradually tapering towards telson. Paraterga strongly developed, starting from collum, subhorizontal, set high but always lying slightly below a faintly convex dorsum; front shoulders and caudal edge nearly straight on several postcollum metaterga, caudal corner always pointed. Collum (Fig. 28) nearly regularly elliptical, with an incision near caudal corner and three transverse rows of setae (6+6, 4+4 and 3+3). Starting from segment 5, paraterga extending increasingly beyond rear tergal contour, mostly subspiniform (Figs 25-28). Starting from segment 2, all poreless segments with three, all pore-bearing ones with four, small but evident incisions, each usually bearing a small seta on top at lateral margin. Pore formula normal, ozopores evident, dorsal, located in front of 4th indentation. Metatergal sculpture typical, rather well developed, with three transverse rows of setiferous, polygonal bosses (Figs 25-28). Tergal setae short, mostly retained, a little longer only on collum and in rear row on metatergum 19 (Figs 25, 27). Stricture between pro- and metazona rather obscure, wide, smooth and polished. Limbus very thin, microdenticulate. Pleurosternal carinae absent. Epiproct rather short, conical (Fig. 31), preapical papillae evident. Hypoproct semi-circular; caudal, paramedian, setiferous papillae evident and well-separated.</paragraph>
<paragraph pageId="7" pageNumber="16">Sterna without modifications (Figs 29, 31), densely setose. Epigynal ridge very low, regularly rounded. Legs long and slender (Figs 28, 29), ca 1.8-1.9 (♂) or 1.2-1.3 (♀) times as long as midbody height; ♂ legs evidently enlarged, prefemora only slightly swollen dorsally, femora ventro-parabasally with shorter bifid setae replaced by sphaerotrichomes in remaining parts of femora, as well as on entire postfemora, tibiae and tarsi.</paragraph>
<paragraph lastPageId="8" lastPageNumber="17" pageId="7" pageNumber="16">
Gonopods (Figs 32-36) with large, subquadrate, medially fused coxae carrying a few long setae ventrally. Telopodite stout to rather slender, subfalcate, prefemoral (densely setose) portion one-third to half as long as entire telopodite; seminal groove running mesally over much of its extent, only distally moving frontally to recurve first laterad and then mesad, squeezing neatly between a simple, more or less rudimentary to completely reduced exomere (ex) and a more complex, branching and always well-developed endomere (en), then groove continuing to a considerable extent basad to end into a large accessory seminal chamber lying near base of a prominent excavation, the latter carrying an evident hairy pulvillus; en often but not alwayswith a strong mesal process (s), either slenderer and simple or larger and more complex in shape, as well as usually with a distinct, often again branching, laterobasal outgrowth (p); distal remainder of en elongate, often branching and enlarged, sometimes fringed with short setae or spines
<pageBreakToken pageId="8" pageNumber="17" start="start">.</pageBreakToken>
</paragraph>
<caption pageId="8" pageNumber="17">
<paragraph pageId="8" pageNumber="17">
Figures 24-27.
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus orientalis" order="Polydesmida" pageId="8" pageNumber="17" phylum="Arthropoda" rank="species" species="orientalis">Epanerchodus orientalis</taxonomicName>
Attems, 1901, ♂ from Fushan. 24 habitus, lateral view 25 anterior portion of body, subdorsal view 26 middle portion of body, dorsal view 27 posterior portion of body, dorsal view. Photographed not to scale.
</paragraph>
</caption>
<caption pageId="8" pageNumber="17">
<paragraph pageId="8" pageNumber="17">
Figures 28-31.
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus orientalis" order="Polydesmida" pageId="8" pageNumber="17" phylum="Arthropoda" rank="species" species="orientalis">Epanerchodus orientalis</taxonomicName>
Attems, 1901, ♂ from Jialuohu. 28, 29 anterior portion of body, dorsal and ventral views, respectively 30 middle portion of body, lateral view 31 posterior portion of body, ventral view. Photographed not to scale.
</paragraph>
</caption>
<caption pageId="8" pageNumber="17">
<paragraph pageId="8" pageNumber="17">
Figures 32-36. Right gonopods of
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus orientalis" order="Polydesmida" pageId="8" pageNumber="17" phylum="Arthropoda" rank="species" species="orientalis">Epanerchodus orientalis</taxonomicName>
Attems, 1901, ♂♂ from Shengguang (32), Jialuohu (33-35) and Guangfong Highway (36), mesal, lateral, mesal, sublateral and submesal views, respectively. Scale bar: 0.1 mm.
</paragraph>
</caption>
</subSubSection>
<subSubSection lastPageId="10" lastPageNumber="19" pageId="8" pageNumber="17" type="remarks">
<paragraph pageId="8" pageNumber="17">Remarks.</paragraph>
<paragraph pageId="8" pageNumber="17">
This species seems to be the most widespread and variable among congeners. It has heretofore been accepted as being split into two nominal subspecies:
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus orientalis subsp. orientalis" order="Polydesmida" pageId="8" pageNumber="17" phylum="Arthropoda" rank="subSpecies" species="orientalis" subSpecies="orientalis">Epanerchodus orientalis orientalis</taxonomicName>
and
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus orientalis subsp. takakuwai" order="Polydesmida" pageId="8" pageNumber="17" phylum="Arthropoda" rank="subSpecies" species="orientalis" subSpecies="takakuwai">Epanerchodus orientalis takakuwai</taxonomicName>
Verhoeff, 1913, but, given a similarly profound variation range in body size and shape, and, especially, in gonopod structure as observed in Japan alone, the subspecific status of takakuwai has long been questioned (e.g.
<bibRefCitation author="Miyosi, Y" journalOrPublisher="Arachnological Society of East Asia, Osaka" pageId="19" pageNumber="28" title="Ueber japanische Diplopoden" year="1959">Miyosi 1959</bibRefCitation>
,
<bibRefCitation author="Murakami, Y" journalOrPublisher="Bulletin of the National Science Museum" pageId="19" pageNumber="28" pagination="557 - 582" title="Myriapods found in limestone caves of northern Honshu, Japan." volume="12" year="1969">Murakami 1969</bibRefCitation>
,
<bibRefCitation author="Nishikawa, Y" journalOrPublisher="Faculty of Letters Review of Otemon Gakuin University" pageId="19" pageNumber="28" pagination="261 - 275" title="Distributional records on Japanese millipeds (II)" volume="27" year="1993">Nishikawa and Murakami 1993</bibRefCitation>
).
<bibRefCitation author="Murakami, Y" journalOrPublisher="Shizen-Kankyo-Kenkyu Center. Environmental Agency Japan, Tokyo" pageId="19" pageNumber="28" pagination="95 - 106" title="Diplopoda, Pauropoda, Symphyla. A list of Japanese species. Invertebrates 1" year="1993">Murakami (1993)</bibRefCitation>
, in the latest checklist of the Japanese Diplopoda, treats
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus orientalis" order="Polydesmida" pageId="8" pageNumber="17" phylum="Arthropoda" rank="species" species="orientalis">Epanerchodus orientalis</taxonomicName>
without subspecies. Moreover, numerous samples from Hokkaido, Honshu and Shikoku, Japan, reveal that variation in gonopod conformation appears to be purely individual, failing to demonstrate any meaningful geographical patterns (
<bibRefCitation author="Nishikawa, Y" journalOrPublisher="Faculty of Letters Review of Otemon Gakuin University" pageId="19" pageNumber="28" pagination="261 - 275" title="Distributional records on Japanese millipeds (II)" volume="27" year="1993">Nishikawa and Murakami 1993</bibRefCitation>
). The same concerns the available samples from Taiwan. So we formalize here the long suspected synonymy:
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus orientalis subsp. orientalis" order="Polydesmida" pageId="8" pageNumber="17" phylum="Arthropoda" rank="subSpecies" species="orientalis" subSpecies="orientalis">Epanerchodus orientalis orientalis</taxonomicName>
Attems, 1901 =
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus orientalis subsp. takakuwai" order="Polydesmida" pageId="8" pageNumber="17" phylum="Arthropoda" rank="subSpecies" species="orientalis" subSpecies="takakuwai">Epanerchodus orientalis takakuwai</taxonomicName>
Verhoeff, 1913, syn. n.
</paragraph>
<paragraph pageId="8" pageNumber="17">
It is noteworthy that, formally,
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus orientalis" order="Polydesmida" pageId="8" pageNumber="17" phylum="Arthropoda" rank="species" species="orientalis">Epanerchodus orientalis</taxonomicName>
fails to occur in southernmost Japan, i.e. on Kyushu Island and in the Ryukyus (
<bibRefCitation author="Nishikawa, Y" journalOrPublisher="Faculty of Letters Review of Otemon Gakuin University" pageId="19" pageNumber="28" pagination="261 - 275" title="Distributional records on Japanese millipeds (II)" volume="27" year="1993">Nishikawa and Murakami 1993</bibRefCitation>
,
<bibRefCitation author="Nakamura, Y" journalOrPublisher="Acta Arachnologica" pageId="19" pageNumber="28" pagination="73 - 86" title="Distribution and diversity of millipedes of the Ryukyu Archipelago, with the Senkaku and Daito island groups: A literature review (Arthropoda: Diplopoda)." volume="59" year="2010">
Nakamura and
<normalizedToken originalValue="Korsós">Korsos</normalizedToken>
2010
</bibRefCitation>
), to reappear further south only in Taiwan. In northern Honshu, Japan, it is known to reach 26 mm in length and 3.4 mm in width, with the collum being slightly broader than the head (
<bibRefCitation author="Murakami, Y" journalOrPublisher="Bulletin of the National Science Museum" pageId="19" pageNumber="28" pagination="557 - 582" title="Myriapods found in limestone caves of northern Honshu, Japan." volume="12" year="1969">Murakami 1969</bibRefCitation>
). In Taiwan, as proven by our study of several abundant syntopic samples, the animals tend to be smaller, the collum is invariably narrower than the head, while rather often the gonopods are totally devoid of both an exomere remnant and process s, frequently with the distal, longest part of their endomere being slender, not expanded.
</paragraph>
<paragraph pageId="8" pageNumber="17">
Looking at such a profound variation as observed in a number of peripheral and gonopod characters in
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus orientalis" order="Polydesmida" pageId="8" pageNumber="17" phylum="Arthropoda" rank="species" species="orientalis">Epanerchodus orientalis</taxonomicName>
, one cannot ignore several further nominal congeners described from Japan, including Kyushu, in which the gonopods look especially similar to those of
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus orientalis" order="Polydesmida" pageId="8" pageNumber="17" phylum="Arthropoda" rank="species" species="orientalis">Epanerchodus orientalis</taxonomicName>
:
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus inferus" order="Polydesmida" pageId="8" pageNumber="17" phylum="Arthropoda" rank="species" species="inferus">Epanerchodus inferus</taxonomicName>
Verhoeff, 1941,
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus lobatus" order="Polydesmida" pageId="8" pageNumber="17" phylum="Arthropoda" rank="species" species="lobatus">Epanerchodus lobatus</taxonomicName>
Verhoeff, 1941,
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus satoi" order="Polydesmida" pageId="8" pageNumber="17" phylum="Arthropoda" rank="species" species="satoi">Epanerchodus satoi</taxonomicName>
Takakuwa, 1954,
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus tenuis" order="Polydesmida" pageId="8" pageNumber="17" phylum="Arthropoda" rank="species" species="tenuis">Epanerchodus tenuis</taxonomicName>
Takakuwa, 1954,
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus aculeatus" order="Polydesmida" pageId="8" pageNumber="17" phylum="Arthropoda" rank="species" species="aculeatus">Epanerchodus aculeatus</taxonomicName>
Miyosi, 1954,
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus etoi" order="Polydesmida" pageId="8" pageNumber="17" phylum="Arthropoda" rank="species" species="etoi">Epanerchodus etoi</taxonomicName>
Miyosi, 1955,
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus chichibensis" order="Polydesmida" pageId="8" pageNumber="17" phylum="Arthropoda" rank="species" species="chichibensis">Epanerchodus chichibensis</taxonomicName>
Haga, in
<bibRefCitation author="Takashima, H" journalOrPublisher="Miscellaneous Reports of Yamashina's Institute of Ornithology and Zoology" pageId="19" pageNumber="28" pagination="329 - 343" title="A contribution towards the Japanese cave-dwelling species of the class Diplopoda" volume="1" year="1956">Takashima and Haga 1956</bibRefCitation>
,
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus yoshidai" order="Polydesmida" pageId="8" pageNumber="17" phylum="Arthropoda" rank="species" species="yoshidai">Epanerchodus yoshidai</taxonomicName>
Haga, in
<bibRefCitation author="Takashima, H" journalOrPublisher="Miscellaneous Reports of Yamashina's Institute of Ornithology and Zoology" pageId="19" pageNumber="28" pagination="329 - 343" title="A contribution towards the Japanese cave-dwelling species of the class Diplopoda" volume="1" year="1956">Takashima and Haga 1956</bibRefCitation>
,
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus lacteus" order="Polydesmida" pageId="8" pageNumber="17" phylum="Arthropoda" rank="species" species="lacteus">Epanerchodus lacteus</taxonomicName>
Shinohara, 1958, etc. (e.g.
<bibRefCitation author="Takashima, H" journalOrPublisher="Miscellaneous Reports of Yamashina's Institute of Ornithology and Zoology" pageId="19" pageNumber="28" pagination="329 - 343" title="A contribution towards the Japanese cave-dwelling species of the class Diplopoda" volume="1" year="1956">Takashima and Haga 1956</bibRefCitation>
,
<bibRefCitation author="Shinohara, K" journalOrPublisher="Bulletin of the Chichibu Museum of Natural History" pageId="19" pageNumber="28" pagination="19 - 24" title="Polydesmoidea millipeds from Chichibu" volume="8" year="1958">Shinohara 1958</bibRefCitation>
,
<bibRefCitation author="Miyosi, Y" journalOrPublisher="Arachnological Society of East Asia, Osaka" pageId="19" pageNumber="28" title="Ueber japanische Diplopoden" year="1959">Miyosi 1959</bibRefCitation>
). We suspect that some of them might well prove to represent junior synonyms of
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus orientalis" order="Polydesmida" pageId="8" pageNumber="17" phylum="Arthropoda" rank="species" species="orientalis">Epanerchodus orientalis</taxonomicName>
. Only future in-depth biological observations and such genetic investigations as bar-coding of such particularly similar forms can shed light on their true identities and statuses, because polymorphic variation has long been known in
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus" order="Polydesmida" pageId="8" pageNumber="17" phylum="Arthropoda" rank="genus">Epanerchodus</taxonomicName>
. Thus,
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus polymorphus" order="Polydesmida" pageId="8" pageNumber="17" phylum="Arthropoda" rank="species" species="polymorphus">Epanerchodus polymorphus</taxonomicName>
Mikhaljova &amp; Golovatch, 1981, widespread in the southern part of the Russian Far East and in northern Korea, shows two morphologically distinct morphs both in gonopod and peripheral structure in males, but a complete, overlapping range of the same somatic characters in females. Both male morphs invariably co-occur syntopically and either mates with any female variety (
<bibRefCitation author="Mikhaljova, EV" journalOrPublisher="Zoologicheskiy Zhurnal" pageId="19" pageNumber="28" pagination="1183 - 1189" title="Polymorphism in a new species of Epanerchodus (Diplopoda, Polydesmidae) from the USSR Far East" volume="60" year="1981">Mikhaljova and Golovatch 1981</bibRefCitation>
,
<bibRefCitation author="Mikhaljova, EV" journalOrPublisher="Pensoft, Sofia-Moscow" pageId="19" pageNumber="28" title="The millipedes of the Asian part of Russia" year="2004">Mikhaljova 2004</bibRefCitation>
). A similar situation is found in the nominate species
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus acuticlivus" order="Polydesmida" pageId="8" pageNumber="17" phylum="Arthropoda" rank="species" species="acuticlivus">Epanerchodus acuticlivus</taxonomicName>
Murakami, 1970 and
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus aster" order="Polydesmida" pageId="8" pageNumber="17" phylum="Arthropoda" rank="species" species="aster">Epanerchodus aster</taxonomicName>
Murakami, 1970, both described from the same cave in Shikoku, Japan.
<bibRefCitation author="Murakami, Y" journalOrPublisher="Annotationes Zoologicae Japonenses" pageId="19" pageNumber="28" pagination="151 - 157" title="More new species of Epanerchodus (Diplopoda, Polydesmidae) found in limestone caves in eastern Shikoku, Japan." volume="43" year="1970">Murakami (1970)</bibRefCitation>
explicitly admitted that they were very close, with their females being indistinguishable, while the males showed small but stable differences in gonopod telopodite armature.
</paragraph>
<paragraph pageId="9" pageNumber="18">
<pageBreakToken pageId="9" pageNumber="18" start="start">Based</pageBreakToken>
on the great variation observed in the populations of
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus orientalis" order="Polydesmida" pageId="9" pageNumber="18" phylum="Arthropoda" rank="species" species="orientalis">Epanerchodus orientalis</taxonomicName>
in Japan and, especially, Taiwan, the statuses of
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus polymorphus" order="Polydesmida" pageId="9" pageNumber="18" phylum="Arthropoda" rank="species" species="polymorphus">Epanerchodus polymorphus</taxonomicName>
,
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus acuticlivus" order="Polydesmida" pageId="9" pageNumber="18" phylum="Arthropoda" rank="species" species="acuticlivus">Epanerchodus acuticlivus</taxonomicName>
and
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus aster" order="Polydesmida" pageId="9" pageNumber="18" phylum="Arthropoda" rank="species" species="aster">Epanerchodus aster</taxonomicName>
as further congeners highly similar to
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus orientalis" order="Polydesmida" pageId="9" pageNumber="18" phylum="Arthropoda" rank="species" species="orientalis">Epanerchodus orientalis</taxonomicName>
are likewise to be questioned, as at least some of them might also prove to be the
<normalizedToken originalValue="latter’s">latter's</normalizedToken>
junior synonyms. Yet no formal synonymies are advanced here as obviously being too premature at this purely descriptive taxonomic stage.
</paragraph>
<paragraph pageId="10" pageNumber="19">
<pageBreakToken pageId="10" pageNumber="19" start="start">That</pageBreakToken>
not all of the congeners similar in gonopod structure to
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus orientalis" order="Polydesmida" pageId="10" pageNumber="19" phylum="Arthropoda" rank="species" species="orientalis">Epanerchodus orientalis</taxonomicName>
are synonyms of the latter is proven at least by
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus koreanus" order="Polydesmida" pageId="10" pageNumber="19" phylum="Arthropoda" rank="species" species="koreanus">Epanerchodus koreanus</taxonomicName>
Verhoeff, 1937, a species common throughout Korea, in Kyushu, Japan and in the southern Russian Far East. At least in Russia, it often occurs syntopically together with
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus polymorphus" order="Polydesmida" pageId="10" pageNumber="19" phylum="Arthropoda" rank="species" species="polymorphus">Epanerchodus polymorphus</taxonomicName>
. Yet
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus koreanus" order="Polydesmida" pageId="10" pageNumber="19" phylum="Arthropoda" rank="species" species="koreanus">Epanerchodus koreanus</taxonomicName>
, despite its minor variations in gonopod conformation, is easily recognizable even superficially through its much wider paraterga (3.3-3.8 versus 2.6-3.0 mm); in addition, these species never mate (
<bibRefCitation author="Mikhaljova, EV" journalOrPublisher="Pensoft, Sofia-Moscow" pageId="19" pageNumber="28" title="The millipedes of the Asian part of Russia" year="2004">Mikhaljova 2004</bibRefCitation>
). As another proof to the above statement may also serve
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus pinguis" order="Polydesmida" pageId="10" pageNumber="19" phylum="Arthropoda" rank="species" species="pinguis">Epanerchodus pinguis</taxonomicName>
sp. n., described below and representing still one more congener quite similar in gonopod structure to
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus orientalis" order="Polydesmida" pageId="10" pageNumber="19" phylum="Arthropoda" rank="species" species="orientalis">Epanerchodus orientalis</taxonomicName>
. In Taiwan, both are at least partly sympatric, but
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus pinguis" order="Polydesmida" pageId="10" pageNumber="19" phylum="Arthropoda" rank="species" species="pinguis">Epanerchodus pinguis</taxonomicName>
sp. n. differs markedly enough in a number of peripheral and gonopod characters to warrant recognition of a distinct species (see below).
</paragraph>
<paragraph pageId="10" pageNumber="19">
In other words, several, but definitely not all, of the nominate species of
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus" order="Polydesmida" pageId="10" pageNumber="19" phylum="Arthropoda" rank="genus">Epanerchodus</taxonomicName>
that show their gonopods particularly similar in structure to those of
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus orientalis" order="Polydesmida" pageId="10" pageNumber="19" phylum="Arthropoda" rank="species" species="orientalis">Epanerchodus orientalis</taxonomicName>
are jeopardized as potential junior synonyms of the latter species.
</paragraph>
<paragraph pageId="10" pageNumber="19">
In Taiwan,
<taxonomicName class="Diplopoda" family="Polydesmidae" genus="Epanerchodus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epanerchodus orientalis" order="Polydesmida" pageId="10" pageNumber="19" phylum="Arthropoda" rank="species" species="orientalis">Epanerchodus orientalis</taxonomicName>
occurs in various habitats ranging from lowlands to the mid-montane belt (up to 1,250 m a.s.l.) all over the island (Map 2).
</paragraph>
</subSubSection>
</treatment>
</document>
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