identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
5AB21EA3702159E1AE10A74965A15120.text	5AB21EA3702159E1AE10A74965A15120.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Kronopolites Attems 1914	<div><p>Genus  Kronopolites Attems, 1914</p><p>Kronopolites Attems 1914: 219.</p><p>Kronopolites: Attems 1929: 272; 1931: 113; 1936: 225; 1937: 49; Verhoeff 1939: 274; Takashima 1950: 38; Takakuwa 1954: 30; Hoffman 1962: 579; 1980: 169; Jeekel 1971: 225; 1982: 243; 1988: 98; Chen et al. 2006 a: 252; Golovatch 2009: 121; 2013: 12; Nguyen and Sierwald 2013: 1286; Likhitrakarn et al. 2015: 32; Golovatch 2015: 135; Golovatch 2016: 1; Golovatch 2019: 348; Golovatch and Liu 2020: 170; Golovatch and Semenyuk 2021: 479.</p><p>Kansupus Verhoeff 1934: 17, synonymized by Attems (1936: 233).</p><p>Kansupus: Jeekel 1971: 225; Hoffman 1980: 169.</p><p>Parakansupus Verhoeff 1939: 273, synonymized by Hoffman (1962: 579).</p><p>Parakansupus: Jeekel 1971: 230; Hoffman 1980: 169.</p><p>Type species.</p><p>Kronopolites svenhedini (Verhoeff, 1934) sp. reval., by present designation.</p><p>Diagnosis.</p><p>See Likhitrakarn et al. (2015: 29).</p><p>Remarks.</p><p>Attems (1914) established the genus and designated  Strongylosoma swinhoei Pocock, 1895 as the type species by monotypy. However, due to the misidentification of this species, which was later transferred to  Nedyopus, the correct designation by Attems,  Kronopolites svenhedini (Verhoeff, 1934) sp. reval., has now been reclassified as the type species of the genus.</p></div>	https://treatment.plazi.org/id/5AB21EA3702159E1AE10A74965A15120	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Xiong, Yuan;Chen, Huiming;Jiang, Xuankong;Jiang, Chao	Xiong, Yuan, Chen, Huiming, Jiang, Xuankong, Jiang, Chao (2025): Topotypes of the millipede species Kronopolites swinhoei (Pocock, 1895) reveal a new synonym with revalidation of Kronopolites svenhedini (Verhoeff, 1934) (Diplopoda, Polydesmida, Paradoxosomatidae). ZooKeys 1231: 85-98, DOI: 10.3897/zookeys.1231.137769
1B4CA5D2D0035ACEB00DF66E0D1C04E9.text	1B4CA5D2D0035ACEB00DF66E0D1C04E9.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Kronopolites svenhedini (Verhoeff 1934)	<div><p>Kronopolites svenhedini (Verhoeff, 1934) sp. reval.</p><p>Figs 1 B, 5, 6, 7</p><p>Stronglosoma Swinhoei: Brölemann 1896: 354–357, fig. 9–11; Attems 1898: 304 (misidentified).</p><p>Kansupus svenhedini Verhoeff, 1934: 17, figs. 4–8, synonymized by Hoffman 1962: 581. Type locality: N. O. Szetschuan (= northeastern Sichuan Province), China; ♂ and Süd-Kansu (= Longnan, Gansu Province, China; ♀).</p><p>Kronopolites swinhoei: Attems 1914: 219; Attems 1936: 226, fig. 44; Attems 1937: 50–51, fig. 64; Chamberlin and Wang 1953: 5; Hoffman 1962: 581–583, figs 1, 2; Golovatch 1978: 678; 1982: 298; Wang 1996: 86; Geoffroy and Golovatch 2004: 20; Korsós 2004: 23; Chen et al. 2006 a: 252; Golovatch 2009: 121; 2013: 2, figs 1–4; Nguyen and Sierwald 2013: 1286; Likhitrakarn et al. 2015: 32; Golovatch 2016: 1; 2019: 348, figs 4, 5; 2020: 166; Golovatch and Liu 2020: 170; Golovatch and Semenyuk 2021: 479, figs 28–30 (misidentified).</p><p>Kronopolites svenhedini: Attems 1936: 233; 1937: 53, fig. 66; Zhang and Li 1978: 12; Wang 1996: 86.</p><p>Kansupus svenhedini var. dentiger Verhoeff, 1934: 19, fig. 9; Attems 1936: 233; Attems 1937: 54, synonymized with 
Kronopolites swinhoei
by Hoffman 1962: 581. Type locality: Pei-shui-ho (= Baishuijiang National Nature Reserve), Wen County, Gansu Province, China).</p><p>Material examined.</p><p>China – • Gansu Province: 5 ♂♂ and 40 ♀♀ (20230922044, - 45, - 46, 20230922048–20230922051, 20230922054–20230922057, - 60, - 61, - 63, - 66, - 67, - 68, - 70, - 72, - 73, - 75, - 76, 20230922078–20230922090, 20230922092–20230922098, - 101, - 102, - 104), Lintao County, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=103.8901&amp;materialsCitation.latitude=35.4009" title="Search Plazi for locations around (long 103.8901/lat 35.4009)">Fenghuangshan Forest Park</a> (35.4009 ° N, 103.8901 ° E), 1960 m a. s. l., 22. IX. 2023, Tianyun Chen, Jiabo Fan &amp; Yiying Zhao leg., (CMMI) ; •   2 ♂♂ and 2 ♀♀, Dingxi City, Anding District, Guanying Town,  Yawan Village, 27. VI. 2008, Zhiyong Di leg., (IBGAS)  . •   Henan Province: 1 ♂ and 2 ♀♀, Xinyang City, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=114.0118&amp;materialsCitation.latitude=32.1252" title="Search Plazi for locations around (long 114.0118/lat 32.1252)">Dabieshan station</a> (32.1252 ° N, 114.0118 ° E), 110 m a. s. l., 10. VIII. 2023, Xuankong Jiang &amp; Leilei Shi leg., (IBGAS) ; •   1 ♂ and 1 ♀ 5 J, Xinyang City, Shihe district, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=113.9105&amp;materialsCitation.latitude=31.9987" title="Search Plazi for locations around (long 113.9105/lat 31.9987)">Bailongtan Reservoir</a> (31.9987 ° N, 113.9105 ° E), 230 m a. s. l., 10. VIII. 2023, Xuankong Jiang &amp; Leilei Shi leg., (IBGAS) ; •   17 ♂♂ and 5 ♀♀ 3 J, Xinyang, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=114.0118&amp;materialsCitation.latitude=32.1252" title="Search Plazi for locations around (long 114.0118/lat 32.1252)">Nanwan Reservoir</a> (32.1252 ° N, 114.0118 ° E), 110 m a. s. l., 9. VIII. 2023, Xuankong Jiang &amp; Leilei Shi leg., (IBGAS)  . •   Qinghai Province: 3 ♂♂ and 10 ♀♀,  Huangnan Tibetan Autonomous Prefecture, Jianzha County, Zhiyong Di leg., (IBGAS)  . •   Shaanxi Province: 1 ♂ (20190907029), Mei County, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=107.7836&amp;materialsCitation.latitude=34.0533" title="Search Plazi for locations around (long 107.7836/lat 34.0533)">Honghegu National Forest Park</a> (34.0533 ° N, 107.7836 ° E), 1730 m a. s. l., 7. IX. 2019, Chao Jiang leg., (CMMI) ; •   3 ♂♂ and 4 ♀♀ (20200905117, - 118, 20230802001, - 02), Xi’an, Huyi District, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=108.7163&amp;materialsCitation.latitude=33.9951" title="Search Plazi for locations around (long 108.7163/lat 33.9951)">Taiping National Forest Park</a> (33.9951 ° N, 108.7163 ° E), 540 m a. s. l., 2. VIII. 2023, Tianyun Chen &amp; Yuan Xiong leg., (CMMI)  . •   Zhejiang Province: 3 ♂♂ and 3 ♀♀, Anji County,  Longwangshan Scenic Area, 22. VII. 2018, Rong Fu leg., (IBGAS)  .</p><p>Diagnosis.</p><p>Differs from other species of the genus by the following combination of characters: metazonae have two shapes, either as a transverse band or a median oval spot, and also have two color variations, ranging from pale yellow to orange-red; paraterga relatively poorly developed, set lower (mostly at about 1 / 3 height of segments), caudal corners usually not surpassing rear tergal contours, at most narrowly rounded; ♂ sternal cones present; processes a and b of gonopod on a broad common stem, neither slender nor long (Golovatch 2009).</p><p>Description.</p><p>Length ca 26.0–50.0 mm (♂), 27.0–60.0 mm (♀) with 20 segments. Live color variable (Fig. 1 B). Head, prozonae, anterior part of collum, and metazonae black; posterior part of collum and each metazonite with transverse band or oval spot, pale yellow to orange-red. If the patches not covering ozopores, then the ozopores exhibit the same color of the patches. Telson black with color of tip identical to the patches. Antennae black; legs black to reddish brown (Fig. 5 A – H).</p><p>Head densely setose. Antennae moderately long (Fig. 5 A), extending behind body segment 3 when stretched dorsally. Width gradually widened from collum to 5 th segment, roughly equal in 5 th – 16 th segments, and tapering from 16 th to telson.</p><p>Collum of different specimens with one or two transverse rows of setae, one row with 1 + 1 anterior, two rows with 1 + 1 at both anterior and intermediate. Caudal corner of collum very broadly rounded, declined ventrad, produced behind rear tergal margin (Fig. 5 A, B).</p><p>Cuticle shining, prozonae finely shagreened, metaterga finely rugulose (Fig. 5 A, C, F), surface below paranota finely microgranulate (Fig. 5 B, D). Postcollum metaterga with one transverse row of setae: 4 + 4 or 3 + 3 in anterior (pre-sulcus). Tergal setae long and slender, mostly abraded. Paranota well developed (Fig. 5 A – F), lying rather high (at upper 1 / 2 of body), arched. Ozopores evident, lateral, lying in an ovoid groove at about 1 / 4 in front of posterior edge of metaterga, in segments 5, 7, 9, 10, 12, 13, 15–19. Transverse sulcus usually distinct (Fig. 5 A, C, F), slightly incomplete on segment 18, complete on metaterga 3–18 (♂), narrow, linear, shallow, reaching bases of paranota, faintly ribbed at bottom. Stricture between pro- and metazonae evident, broad and deep, ribbed at bottom down to base of paranota (Fig. 5 A, B, D, G). Pleurosternal carinae with a sharp caudal tooth on segments 2–7, thereafter increasingly strongly reduced until 18 th segment (♂). Epiproct (Fig. 5 G, H) conical, flattened dorsoventrally, with two small apical papillae; tip subtruncate; pre-apical papillae small, lying close to tip. Hypoproct roundly subtriangular, spinnerets at caudal edge small and well separated (Fig. 5 H). Sterna densely setose with a long tongue-shaped process on ♂ coxae 4 (Fig. 5 E). Legs rather long and slender (Fig. 5 A, B, D, G).</p><p>Coxite of gonopods (Fig. 6) thick, pressing inwards on the spermathecal fossa in the prefemur. Prefemur short, with numerous slender setae. Femorite rather stout, with an evident mesal groove and a strong distolateral sulcus demarcating a postfemoral part; the latter well developed, with very prominent bipartite, crescent-shaped, lateral processes: process a rather long and broad; process b short, broad and pointed. Solenophore in the form of a long, tubular branch. Solenomere flagelliform.</p><p>Distribution.</p><p>China: Chongqing, Qinghai, Gansu, Guizhou, Henan (new record), Shaanxi, Sichuan, Yunnan and Zhejiang (Golovatch 2019; Chen et al. 2023).</p><p>Remarks.</p><p>Verhoeff (1934) described a new monotypic genus and species,  Kansupus svenhedini, based on specimens from Gansu, China. Later, Attems (1936) and Hoffman (1962) synonymized the name with  Kronopolites and  K. swinhoei respectively, relying on Brölemann’s (1896) description. However, our research shows that Brölemann (1896) misidentified this species. Thus,  K. svenhedini sp. reval. should be resurrected. To maintain the stability of the taxonomic name  Kronopolites, the type species of  Kronopolites now fixed (under Article 70.3 of the International Code of Zoological Nomenclature 1999) as  Kronopolites svenhedini (Verhoeff, 1934), which was misidentified as  Kronopolites swinhoei (Pocock, 1895) in the original designation by Attems (1914). Furthermore, the publications mentioning this species did not state where the type specimen is preserved, rendering the location of its storage unknown (Verhoeff 1934; Attems 1936, 1937; Hoffman 1962; Zhang and Li 1978; Wang 1996).</p><p>Kronopolites svenhedini (Verhoeff, 1934) sp. reval. is widely distributed in China, with the westernmost occurrence in Qinghai, the easternmost in Zhejiang, the southernmost in Yunnan and the northernmost in Gansu. It shows variation in color, body size, and subtle differences in gonopod shape among different populations (Golovatch 2013). These differences may be attributed to geographic isolation or varying habitats, suggesting the potential presence of cryptic species. This hypothesis could be explored in future research using molecular methods.</p></div>	https://treatment.plazi.org/id/1B4CA5D2D0035ACEB00DF66E0D1C04E9	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Xiong, Yuan;Chen, Huiming;Jiang, Xuankong;Jiang, Chao	Xiong, Yuan, Chen, Huiming, Jiang, Xuankong, Jiang, Chao (2025): Topotypes of the millipede species Kronopolites swinhoei (Pocock, 1895) reveal a new synonym with revalidation of Kronopolites svenhedini (Verhoeff, 1934) (Diplopoda, Polydesmida, Paradoxosomatidae). ZooKeys 1231: 85-98, DOI: 10.3897/zookeys.1231.137769
AACC699E89185D8AA0C17DB1825B24F7.text	AACC699E89185D8AA0C17DB1825B24F7.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Nedyopus Attems 1914	<div><p>Genus  Nedyopus Attems, 1914</p><p>Type species.</p><p>Orthomorpha cingulata Attems, 1898, by original designation.</p></div>	https://treatment.plazi.org/id/AACC699E89185D8AA0C17DB1825B24F7	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Xiong, Yuan;Chen, Huiming;Jiang, Xuankong;Jiang, Chao	Xiong, Yuan, Chen, Huiming, Jiang, Xuankong, Jiang, Chao (2025): Topotypes of the millipede species Kronopolites swinhoei (Pocock, 1895) reveal a new synonym with revalidation of Kronopolites svenhedini (Verhoeff, 1934) (Diplopoda, Polydesmida, Paradoxosomatidae). ZooKeys 1231: 85-98, DOI: 10.3897/zookeys.1231.137769
8C1324A3D28A523B969C713702653B4A.text	8C1324A3D28A523B969C713702653B4A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Nedyopus swinhoei (Pocock 1895) Xiong & Chen & Jiang & Jiang 2025	<div><p>Nedyopus swinhoei (Pocock, 1895) comb. nov.</p><p>Figs 1 A, 2, 3, 4</p><p>Stronglosoma Swinhoei Pocock, 1895: 354–355 . Type specimen: holotype female, collected from Chee Foo (= Zhifu), Yantai, Shandong Province of China, deposited at the British Museum of Natural History (Hoffman 1962), not examined.</p><p>Kronopolites swinhoei: Attems 1914.</p><p>Strongylosoma patrioticum Attems, 1898: 300, figs 12, 13. Type locality: Japan. New synonymy.</p><p>Nedyopus patrioticus: Attems 1914: 201; Attems 1937: 138–139; Takashima 1949: 17; Takakuwa 1954: 47; Takashima and Haga 1956: 332; Miyosi 1959: 49, 71; Wang 1964: 71; Wang 1996: 87; Mikhaljova et al. 2000: 119; Korsós 2004: 23; Chen et al. 2006 b: 3998 –4000; Nguyen and Sierwald 2013: 1231. New synonymy.</p><p>Diagnosis.</p><p>Differs from other species of the genus by the following combination of characters. The metaterga have strong contrasting colors, which are not circularly patterned as in other  Nedyopus species, and the gonopod femur suddenly widens at the base, with l’ and l ” not jagged.</p><p>Material examined.</p><p>China – Anhui Province: • 1 ♂,  Fuyang, 30. I. 2020, Yihao Ge leg. (IBGAS)  . •   Jiangxi Province: 1 ♂ and 1 ♀, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=115.0426&amp;materialsCitation.latitude=27.1439" title="Search Plazi for locations around (long 115.0426/lat 27.1439)">Ji’an</a>, (27.1439 ° N, 115.0426 ° E), 50 m a. s. l., 19. V. 2017, Xuankong Jiang leg. (IBGAS)  . •   Shandong Province: 1 ♂ and 1 ♀ (20231025001), Yantai, Zhifu District, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=121.3716&amp;materialsCitation.latitude=37.61" title="Search Plazi for locations around (long 121.3716/lat 37.61)">Zhifu Island</a>, (37.6100 ° N, 121.3716 ° E), 40 m a. s. l., 25. X. 2023, Xuankong Jiang, Tian Lu and Chongwu Lu leg. (CMMI) ; •  16 ♂♂ and 12 ♀♀, same data, (IBGAS); •   15 ♂♂ and 13 ♀♀, Yantai, Zhifu District, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=121.3972&amp;materialsCitation.latitude=37.5056" title="Search Plazi for locations around (long 121.3972/lat 37.5056)">Tashan Park</a>, (37.5056 ° N, 121.3972 ° E), 290 m a. s. l., 24. X. 2023, Xuankong Jiang, Tian Lu and Chongwu Lu leg. (IBGAS)  .</p><p>Description.</p><p>Length ca 17.5–25.1 mm (♂), 18.2–32.7 mm (♀) with 20 segments. Live color variable (Fig. 1 A). Posterior half of each metazonae with transverse band, pale yellow to orange. Antennomere 1–6 dark brown, antennomere 7 whitish. Legs light yellow (Fig. 2 A – H).</p><p>Clypeolabral region and vertex densely setose. Epicranial suture distinct. Width of body gradually expanded from head to 5 th segment, approximately equal in width from 5 th to 16 th segments, and tapering from 16 th to telson. Caudal corner of collum broadly rounded, declined ventrad, produced behind rear tergal margin (Fig. 2 A, B).</p><p>Cuticle shining (Fig. 2 A, C, F); surface below paranota finely microgranulate (Fig. 2 B, D, G). Paranota strongly developed (Fig. 2 A, B, C, F), slightly upturned, lying rather high (at upper 1 / 3 of body) but below dorsum; anterior edge broadly rounded and narrowly bordered; posterior edge nearly straight. Ozopores evident, lying on paranota at its posterior margin, in segments 5, 7, 9, 10, 12, 13, 15–19. Transverse sulcus usually distinct (Fig. 2 A, C, F), complete on metaterga 5–18 (♂), narrow, linear, shallow, reaching bases of paranota, faintly ribbed at bottom. Stricture between pro- and metazonae evident, broad and deep, ribbed at dorsal side down to base of paranota (Fig. 2 A – F). Pleurosternal carinae with a sharp caudal tooth on segments 3–6. Epiproct (Fig. 2 G, H) conical, flattened dorsoventrally; tip subtruncate; pre-apical papillae small, lying close to tip. Hypoproct roundly subtriangular, spinnerets at caudal edge small and well separated (Fig. 2 H). Sterna densely setose, without modifications, but with two small, rounded, fully separated, setose cones between ♂ coxae 4 (Fig. 2 E).</p><p>Gonopods (Fig. 3) intricate. Coxite elongate, subcylindrical, strongly setose distoventrally. Prefemoral part nearly half femoral length. Femorite short and bulge out at one end like a belly, distal portion carrying two lobes (l’ and l ”). l’ parallel to solenophore. Solenophore lamelliform, twisted distally. Solenomere short and flagelliform.</p><p>Distribution.</p><p>China: Anhui (New record), Jiangsu, Jiangxi (New record), Shandong, Taiwan (Pocock 1895; Chen et al. 2006 b; Zhang et al. 2024); Indonesia, Japan, Korea (Nguyen and Sierwald 2013).</p><p>Remarks.</p><p>The specimens from Zhoushan Island were initially identified as  K. swinhoei by Brölemann in 1896, without providing a justification. However, our investigation reveals a distinct divergence from the original description. For instance, the specimens from Zhoushan are notably larger (47 mm vs 35 mm) and have more vivid in color on the metazonites (orange-red vs yellow).</p><p>During our research in Zhifu, we found a species that closely matches Pocock’s description, leading us to confidently identify it as  K. swinhoei . On examination of the topotypes, we observed significant differences in the gonopods compared to Brölemann’s illustrations (1896). These differences, including the femorite (strongly twisted and expanded in  Nedyopus vs straight in  Kronopolites), the postfemoral sulcus (missing in  Nedyopus vs existed in  Kronopolites) and the solenophore (lamelliform in  Nedyopus vs tubuliform in  Kronopolites), indicate that this species belongs to  Nedyopus rather than  Kronopolites, and is identical to the widespread species  Nedyopus patrioticus (Attems, 1898) . Consequently,  K. swinhoei is formally transferred to  Nedyopus, and  Nedyopus patrioticus is considered a junior synonym of  Nedyopus swinhoei (Pocock, 1895) comb. nov. Additionally,  N. patrioticus consists of two subspecies  Nedyopus patrioticus patrioticus (Attems, 1898) from Japan and  Nedyopus patrioticus unicolor (Carl, 1902) from Indonesia. Therefore, the subspecies unicolor should be treated as  Nedyopus swinhoei unicolor (Carl, 1902) comb. nov.</p><p>Nedyopus swinhoei (Pocock, 1895) comb. nov. has a wide distribution across Asia, from Indonesia to China, Korea, and Japan (Wang 1955; Nguyen and Sierwald 2013). This species is often found near human habitation and may spread through human activities. In China, documented records of this species were limited to Jiangsu, Shandong, and Taiwan provinces (Pocock 1895; Wang 1955; Zhang et al. 2024), possibly due to limited investigation. Our study identified additional distribution locations in Anhui and Jiangxi, suggesting that the species might be found in other regions across China in the future.</p></div>	https://treatment.plazi.org/id/8C1324A3D28A523B969C713702653B4A	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Xiong, Yuan;Chen, Huiming;Jiang, Xuankong;Jiang, Chao	Xiong, Yuan, Chen, Huiming, Jiang, Xuankong, Jiang, Chao (2025): Topotypes of the millipede species Kronopolites swinhoei (Pocock, 1895) reveal a new synonym with revalidation of Kronopolites svenhedini (Verhoeff, 1934) (Diplopoda, Polydesmida, Paradoxosomatidae). ZooKeys 1231: 85-98, DOI: 10.3897/zookeys.1231.137769
