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<mods:title id="266E40AE0642CB121A252A9B4FCAC53F">A taxonomic review of the centipede genus Scolopendra Linnaeus, 1758 (Scolopendromorpha, Scolopendridae) in mainland Southeast Asia, with description of a new species from Laos</mods:title>
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<mods:namePart id="235E12C85E93C2425829720FCFE57189">Siriwut, Warut</mods:namePart>
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<mods:namePart id="69BCDB0D7E7D93A8D77AF9A07557FDAC">Edgecombe, Gregory D.</mods:namePart>
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<mods:namePart id="7E6A7F9887EA3E5073F121D1609DAA7D">Sutcharit, Chirasak</mods:namePart>
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<mods:namePart id="F5DB93F50796F2975D8EF6EC31F57ABF">Tongkerd, Piyoros</mods:namePart>
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<subSubSection id="32F8EB61AD72DAE3B6EF158E5D3A693D" pageId="19" pageNumber="20" type="multiple">
<paragraph id="8CB762A3640B44BFF1D533F3CC0C012B" pageId="19" pageNumber="20">Taxon classification Animalia Scolopendromorpha Scolopendridae</paragraph>
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<subSubSection id="178DF298CFD457DED975C87EC976920F" pageId="19" pageNumber="20" type="nomenclature">
<paragraph id="671221B90FEC767F0FFDD9C1210AFDD4" pageId="19" pageNumber="20">
<taxonomicName id="4E977E71BD7D6EBEBFA56A8E077DA65A" ID-CoL="5F3CL" authority="Attems, 1934" authorityName="Attems" authorityYear="1934" class="Chilopoda" family="Scolopendridae" genus="Scolopendra" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Scolopendra subspinipes" order="Scolopendromorpha" pageId="19" pageNumber="20" phylum="Arthropoda" rank="subSpecies" species="subspinipes" subSpecies="piceoflava">Scolopendra subspinipes piceoflava Attems, 1934</taxonomicName>
Figs 14, 15
</paragraph>
</subSubSection>
<subSubSection id="9F31694CBAED21A824F94FFC79AECFEB" pageId="19" pageNumber="20" type="materials_examined">
<paragraph id="8A62B9B798A11B346CF1D3C86CE14EE2" pageId="19" pageNumber="20">Material.</paragraph>
<paragraph id="E21194CB60E6CA46E95A2DCDE9623235" pageId="19" pageNumber="20">Syntypes NMB 391Va spec.1-3, one adult male and two adult females, Central Celebes, don. Z.U.F. Sarasin, 1895 (Figs 14, 15).</paragraph>
<caption id="9ECADBC790BAAB4B86FE6B86C9B936B7" pageId="19" pageNumber="20">
<paragraph id="C6BC2B6CE56B0874364C57915B49D32D" pageId="19" pageNumber="20">
Figure 14.
<taxonomicName id="087046358E2193552C07BB7D5B6A5815" class="Chilopoda" family="Scolopendridae" genus="Scolopendra" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Scolopendra subspinipes" order="Scolopendromorpha" pageId="19" pageNumber="20" phylum="Arthropoda" rank="species" species="subspinipes">Scolopendra subspinipes</taxonomicName>
(Syntypes NMB 391Va of &quot;
<taxonomicName id="90B17944C87ADB4562197B3646B42E25" lsidName="piceoflava" pageId="19" pageNumber="20" rank="species" species="piceoflava">piceoflava</taxonomicName>
Attems, 1934&quot;): A Tooth-plates B Cephalic plate and trunk segments 1-3 C Forcipular segment D Tergites 9-12 E Sternites 8-10.
</paragraph>
</caption>
<caption id="8FB692C611A8704374F93A60025D938E" pageId="19" pageNumber="20">
<paragraph id="1CDBF84BA51A78D88B7808765935517E" pageId="19" pageNumber="20">
Figure 15.
<taxonomicName id="FE12D635B5FD84F7C1075CC60256B26E" class="Chilopoda" family="Scolopendridae" genus="Scolopendra" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Scolopendra subspinipes" order="Scolopendromorpha" pageId="19" pageNumber="20" phylum="Arthropoda" rank="species" species="subspinipes">Scolopendra subspinipes</taxonomicName>
(Syntypes NMB 391Va of &quot;
<taxonomicName id="226E67C041DAA5895B9C3CCC54045882" lsidName="piceoflava" pageId="19" pageNumber="20" rank="species" species="piceoflava">piceoflava</taxonomicName>
Attems, 1934&quot;): A Lateral view of coxopleuron B Tergite of ultimate leg-bearing segment and dorsal view of legs 20 and ultimate leg C Ventral view of Leg 20, coxopleura and ultimate leg D Sternite of ultimate leg-bearing segment, coxopleura and ultimate leg
<normalizedToken id="80964C8DC520D4F643BF6541EB603575" originalValue="EF">E-F</normalizedToken>
Ventro-lateral view of ultimate legs.
</paragraph>
</caption>
<caption id="B6F9589DFDA3741AD547599335191F00" pageId="19" pageNumber="20">
<paragraph id="5BDA20651108850C13D7702836EA6C81" pageId="19" pageNumber="20">
Figure 16.
<taxonomicName id="22DF54F869361F420406C425D86F1E84" class="Chilopoda" family="Scolopendridae" genus="Scolopendra" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Scolopendra subspinipes" order="Scolopendromorpha" pageId="19" pageNumber="20" phylum="Arthropoda" rank="species" species="subspinipes">Scolopendra subspinipes</taxonomicName>
(CUMZ 00315): A Cephalic plate and basal antennal articles B Tooth-plates and trochanteroprefemoral process C Tergites 9-11 D Forcipular segment.
</paragraph>
</caption>
<caption id="8C8475C807BE5C89A4DF422BBB678169" pageId="19" pageNumber="20">
<paragraph id="5233574D8664714252EE6C97239DB3A3" pageId="19" pageNumber="20">
Figure 17.
<taxonomicName id="4F296F0441EE6DFAED4648F9C9A6BA86" class="Chilopoda" family="Scolopendridae" genus="Scolopendra" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Scolopendra subspinipes" order="Scolopendromorpha" pageId="19" pageNumber="20" phylum="Arthropoda" rank="species" species="subspinipes">Scolopendra subspinipes</taxonomicName>
(CUMZ 00315): A Sternites 9-11 B Lateral view of coxopleuron C Tergite of ultimate leg-bearing segment D Sternite of ultimate leg-bearing segment, coxopleura, female genital segment and ultimate leg prefemora E Dorsal view of ultimate leg prefemora.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="021264EE14EE196F10D773F882305209" pageId="19" pageNumber="20" type="type locality">
<paragraph id="8AD0E86718A04AA342D8455FC1BDD339" pageId="19" pageNumber="20">Type locality.</paragraph>
<paragraph id="EF420AC33F628D7F550A9DE427204AB6" pageId="19" pageNumber="20">Tomohon, Sulawesi, Indonesia [Tomohon, North Sulawesi Province, Indonesia].</paragraph>
</subSubSection>
<subSubSection id="67E8D8D31AC26038F8B69DDEA6284FFD" pageId="19" pageNumber="20" type="description">
<paragraph id="7301261A45F0CF91B9C803115D854D31" pageId="19" pageNumber="20">Description.</paragraph>
<paragraph id="6AB8F4F14E869012AE6FC7D276B0F600" pageId="19" pageNumber="20">Body length 16.7 cm in male and 17.1 and 16.5 cm in female syntypes. Preserved male still exhibiting traces of its colouration pattern: cephalic plate and segments dark greenish or brown. Antenna yellowish. Tergites with yellowish or pale colour on posterior margin. All legs blue-greenish, distal part yellow. Cephalic plate without small punctae on anterior part, median sulcus present. Posterior part of cephalic plate without paramedian sulci.</paragraph>
<paragraph id="438863F594B81DDAA1831E07F7B4839F" pageId="19" pageNumber="20">Antenna usually with 17-19 articles, basal 6 subcylindrical and glabrous both on dorsal and ventral sides. Antennae reach segment 4 (Fig. 14B). Forcipular trochanteroprefemoral process bearing denticles in two groups, 2-3 apical and one inner. Tooth-plates wider than long or nearly equivalent, 6-7 teeth (Fig. 14A). Tooth-plate with straight, transverse basal suture. Coxosternite smooth without median suture, with shallow depression in male specimen (Fig. 14C). Article 2 of second maxillary telopodite with spur.</paragraph>
<paragraph id="E7D21DD4E806B003CC8D62F6EBF722D5" pageId="19" pageNumber="20">Anterior margin of T1 underlying cephalic plate (Fig. 14B). Complete paramedian sutures from T4; margination typically starting on TT5-7. Tergite surface (Fig. 14D) smooth, without median sulci. Tergite of ultimate leg-bearing segment (Fig. 15B) curved posteriorly, without median furrow or depression; ratio of width: length of tergite of ultimate leg-bearing segment 0.82:1. Sternites (Fig. 14E) with incomplete paramedian sutures, extending 80% length of sternite on anterior segments. Surface of sternites smooth, without depression. Sternite of ultimate leg-bearing segment (Fig. 15D) with sides converging posteriorly; surface typically without depression (with median depression in one female specimen; NMB391Va sp.1). Pore-field on coxopleuron well developed, with gently curved dorsal margin, reaching nearly to margin of tergite of ultimate leg-bearing segment, anterior part of pore area widest (Fig. 15A).</paragraph>
<paragraph id="E5A58861E5590C987690E8E888F54120" pageId="19" pageNumber="20">
Coxopleural process long (Fig. 15
<normalizedToken id="E5CEA7F7B81A926C60A59021B63255C0" originalValue="CD">C-D</normalizedToken>
) with 1-2 apical spine(s) and absence of lateral and dorsal spines; pore-free area extending 30-50% length from distal part of coxopleural process to margin of sternite of ultimate leg-bearing segment.
</paragraph>
<paragraph id="63C230085B958A210C7FB3A733E8A5E8" pageId="19" pageNumber="20">
All legs without setae and tibial spur. One tarsal spur on legs 1-20. Ultimate legs: slender and long (Fig. 15
<normalizedToken id="7F56E78A1F42261AB6FAA5E0B7E618A3" originalValue="EF">E-F</normalizedToken>
), with ratios of lengths of prefemur and femur 1.2:1, femur and tibia 1.1:1, tibia and tarsus 2 1.3:1, tarsus 1 and tarsus 2 2.3:1. Prefemoral spines: 0-2 VL, 1-2 M, 1-2 DM and prefemoral process with 2-5 spines. Posterior margin of prefemur with acute median groove.
</paragraph>
<paragraph id="92C62C8E3BA0CBC5886B84C58D7084CB" pageId="19" pageNumber="20">Sternite of genital segment 1 round and convex posteriorly, with median suture. In male, sternite of genital segment 2 attached to penis. Tergite of genital segment without small setae. Gonopods present in male.</paragraph>
</subSubSection>
<subSubSection id="0F9A7C27D8E104FC3C63D4C2E821F5FA" pageId="19" pageNumber="20" type="discussion">
<paragraph id="EC0391E67F2DD1D8560695FCCF1381A9" pageId="19" pageNumber="20">Discussion.</paragraph>
<paragraph id="527A9476A327F0DFAFD1DE7C815BAB99" pageId="19" pageNumber="20">
Based on examination of the syntypes, we corroborate the assignment of this nominal subspecies to the
<taxonomicName id="FA280A2E321F6B7CC513D590877FABAD" class="Chilopoda" family="Scolopendridae" genus="Scolopendra" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Scolopendra subspinipes" order="Scolopendromorpha" pageId="19" pageNumber="20" phylum="Arthropoda" rank="species" species="subspinipes">Scolopendra subspinipes</taxonomicName>
group. Some morphological characters that appear, however, not to be identical with
<taxonomicName id="DC5FDFA2A93A3FD1AAC69B79C0C8149A" class="Chilopoda" family="Scolopendridae" genus="Scolopendra" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Scolopendra subspinipes" order="Scolopendromorpha" pageId="19" pageNumber="20" phylum="Arthropoda" rank="species" species="subspinipes">Scolopendra subspinipes</taxonomicName>
are the sharpness and length of the coxopleural process, which bears one or two strong apical spines, the ratio of ultimate leg podomeres, and the colouration pattern on the tergites that is clearly distinct from other geographical populations of
<taxonomicName id="3A59E18331FE0FCF4DFCD2B0B9512A5D" class="Chilopoda" family="Scolopendridae" genus="Scolopendra" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Scolopendra subspinipes" order="Scolopendromorpha" pageId="19" pageNumber="20" phylum="Arthropoda" rank="species" species="subspinipes">Scolopendra subspinipes</taxonomicName>
(the posterior part of the tergites exhibiting a yellowish colouration). On the other hand, the syntypes of
<taxonomicName id="B7DAE47550A8EA7540795F3CC60B26C5" class="Chilopoda" family="Scolopendridae" genus="Scolopendra" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Scolopendra subspinipes" order="Scolopendromorpha" pageId="19" pageNumber="20" phylum="Arthropoda" rank="subSpecies" species="subspinipes" subSpecies="piceoflava">Scolopendra subspinipes piceoflava</taxonomicName>
also display morphological variation between each other with respect to the number of prefemoral spines on the ultimate legs: a male specimen has 4-6 spines on the prefemoral process whereas VL, M and DM spines are absent in one female specimen. The latter is similar to
<taxonomicName id="FD9C47C074DDBC0D1EFDC184D6049C1D" class="Chilopoda" family="Scolopendridae" genus="Scolopendra" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Scolopendra dehaani" order="Scolopendromorpha" pageId="19" pageNumber="20" phylum="Arthropoda" rank="species" species="dehaani">Scolopendra dehaani</taxonomicName>
but it is possible that this absence may be due to regeneration in this individual. However, without additional material and lacking molecular data with which to test relationships among morphological similar species, we tentatively accept
<taxonomicName id="EC62AE6D1CDAB402BE7FA0ABCD260726" class="Chilopoda" family="Scolopendridae" genus="Scolopendra" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Scolopendra subspinipes" order="Scolopendromorpha" pageId="19" pageNumber="20" phylum="Arthropoda" rank="subSpecies" species="subspinipes" subSpecies="piceoflava">Scolopendra subspinipes piceoflava</taxonomicName>
as a junior synonym of
<taxonomicName id="4F6CE1F4E99064A40E17D3660CB7B3B3" class="Chilopoda" family="Scolopendridae" genus="Scolopendra" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Scolopendra subspinipes" order="Scolopendromorpha" pageId="19" pageNumber="20" phylum="Arthropoda" rank="species" species="subspinipes">Scolopendra subspinipes</taxonomicName>
as proposed by
<bibRefCitation id="947C53A69ED42C5C286A9841FBD5C294" author="Kronmueller, C" journalOrPublisher="Spixiana" pageId="48" pageNumber="49" pagination="19 - 27" title="Review of the subspecies of Scolopendrasubspinipes Leach, 1815 with the new description of the South Chinese member of the genus Scolopendra Linnaeus, 1758 named Scolopendrahainanum spec. nov." volume="35" year="2012">
<normalizedToken id="8C83FD12B6C19143E445C35FDEE80717" originalValue="Kronmüller">Kronmueller</normalizedToken>
(2012)
</bibRefCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="5991485C00ABCAFB6D165FF17C8E8898" lastPageId="20" lastPageNumber="21" pageId="19" pageNumber="20" type="distribution">
<paragraph id="4D17CCE3B8F7A8EB93D9A740ADE78723" pageId="19" pageNumber="20">Distribution.</paragraph>
<paragraph id="8BE586E2D1C905131F30E887D58AB39E" lastPageId="20" lastPageNumber="21" pageId="19" pageNumber="20">
Previous studies regarded
<taxonomicName id="2687FDB248DDC196A9D393FBF5CE4D55" class="Chilopoda" family="Scolopendridae" genus="Scolopendra" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Scolopendra subspinipes" order="Scolopendromorpha" pageId="19" pageNumber="20" phylum="Arthropoda" rank="species" species="subspinipes">Scolopendra subspinipes</taxonomicName>
s.l. as a cosmopolitan species in tropical regions (
<bibRefCitation id="99CD7BA0FCDA03EFCA2D32966627B371" author="Schileyko, AA" journalOrPublisher="Arthropoda Selecta" pageId="50" pageNumber="51" pagination="71 - 95" title="The scolopendromorph centipedes (Chilopoda) of Vietnam, with contributions to the faunas of Cambodia and Laos. Part 3." volume="16" year="2007">Schileyko 2007</bibRefCitation>
;
<bibRefCitation id="3D571700D82B217A811653F6793A6342" author="Chao, JL" journalOrPublisher="KG, Saarbruecken, Germany" pageId="46" pageNumber="47" title="Scolopendromorpha (Chilopoda) of Taiwan. VDM Verlag Dr. Mueller Aktiengesellschaft &amp; Co." year="2008">Chao 2008</bibRefCitation>
;
<bibRefCitation id="17C4A1E93273445E4180F823F98D7E94" author="Lewis, JGE" journalOrPublisher="International Journal of Myriapodology" pageId="48" pageNumber="49" pagination="83 - 122" title="A key and annotated list of the Scolopendra species of the Old World with a reappraisal of Arthrorhabdus (Chilopoda: Scolopendromorpha: Scolopendridae)." url="10.1163/187525410X12578602960380" volume="3" year="2010 b">Lewis 2010b</bibRefCitation>
). In this study, most of the sampled specimens were collected on islands. Several old collections in the NHMUK
<pageBreakToken id="AAB0E2CFDF45D5613743A926C891205D" pageId="20" pageNumber="21" start="start">identified</pageBreakToken>
as
<taxonomicName id="C99488919410BD0632EA4AE2704EB5A2" class="Chilopoda" family="Scolopendridae" genus="Scolopendra" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Scolopendra subspinipes" order="Scolopendromorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="subspinipes">Scolopendra subspinipes</taxonomicName>
sensu lato from mainland East and Southeast Asia instead refer to former subspecies of
<taxonomicName id="1E98C7CE02894DEB35CA740933A956F0" class="Chilopoda" family="Scolopendridae" genus="Scolopendra" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Scolopendra subspinipes" order="Scolopendromorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="subspinipes">Scolopendra subspinipes</taxonomicName>
that are now identified as distinct species, including
<taxonomicName id="4FA8E0BA69728F06123E1B0BEB2B7930" class="Chilopoda" family="Scolopendridae" genus="Scolopendra" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Scolopendra dawydoffi" order="Scolopendromorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="dawydoffi">Scolopendra dawydoffi</taxonomicName>
,
<taxonomicName id="C8362B9D174DD9BF4A8EB31681ADDC49" class="Chilopoda" family="Scolopendridae" genus="Scolopendra" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Scolopendra dehaani" order="Scolopendromorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="dehaani">Scolopendra dehaani</taxonomicName>
,
<taxonomicName id="0DD622FFD0176E0AD27E4A5F96924D5F" class="Chilopoda" family="Scolopendridae" genus="Scolopendra" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Scolopendra multidens" order="Scolopendromorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="multidens">Scolopendra multidens</taxonomicName>
and
<taxonomicName id="353A74D18A81F4D8089EEB46F623BB28" class="Chilopoda" family="Scolopendridae" genus="Scolopendra" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Scolopendra japonica" order="Scolopendromorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="japonica">Scolopendra japonica</taxonomicName>
. For this reason, we removed occurrence records of
<taxonomicName id="914CADCD74718BFE982D2A366FC20E2C" class="Chilopoda" family="Scolopendridae" genus="Scolopendra" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Scolopendra subspinipes" order="Scolopendromorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="subspinipes">Scolopendra subspinipes</taxonomicName>
s.l. from Thailand and Laos due to our extensive surveys throughout these two countries, finding that no specimen of
<taxonomicName id="63561E0B75DAEA8DEA925F3720F033FD" class="Chilopoda" family="Scolopendridae" genus="Scolopendra" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Scolopendra subspinipes" order="Scolopendromorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="subspinipes">Scolopendra subspinipes</taxonomicName>
s.str. was found in this area. The updated distribution of this species in Asia (Fig. 18) is as follows: Southeast Asia: Myanmar, Malaysia (Penang and Sarawak), Singapore, Vietnam (fide
<bibRefCitation id="9342D5C8A8EC6CA50F694A328C2C0C68" author="Schileyko, AA" journalOrPublisher="Arthropoda Selecta" pageId="50" pageNumber="51" pagination="71 - 95" title="The scolopendromorph centipedes (Chilopoda) of Vietnam, with contributions to the faunas of Cambodia and Laos. Part 3." volume="16" year="2007">Schileyko 2007</bibRefCitation>
: Lao Cai, Vinh Phuc, HatTay, Hai Phong, Quang Binh, Thua Thien Hue, Da Nang, Dak Lak, Khanh Hoa and Dong Nai Provinces), Indonesia (Surabaya, Java, east coast of Sumatra, Lombok, Sumbawa and Mimika River, New Guinea) and Philippines (Manilla). East Asia: China (Zhouzhan Island, Ningbo and Changsha), Taiwan (Kang-hwa), Japan (Izu and Goto Islands, Yokohama and Kobe) and South Korea.
</paragraph>
<caption id="F4672F887DC54D5C547268FB0F4C4249" pageId="20" pageNumber="21">
<paragraph id="7BFB4D270BAA447015AF4990128BCBF7" pageId="20" pageNumber="21">
Figure 18. Distribution map of
<taxonomicName id="B6E9D95F7E9415B3421F20EC129C2791" class="Chilopoda" family="Scolopendridae" genus="Scolopendra" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Scolopendra dehaani" order="Scolopendromorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="dehaani">Scolopendra dehaani</taxonomicName>
and
<taxonomicName id="EF7BEB7A864ED82A742B45B2A794840B" class="Chilopoda" family="Scolopendridae" genus="Scolopendra" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Scolopendra subspinipes" order="Scolopendromorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="subspinipes">Scolopendra subspinipes</taxonomicName>
in Southeast Asia and China-Japan Sea (small map): Filled and blank colours refer to localities from the present study and in the literature, respectively.
</paragraph>
</caption>
</subSubSection>
</treatment>
</document>