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The millipedes of Yintiaoling National Natural Reserve, Southwest China, were investigated. In total, thirteen species from five orders (Sphaerotheriida, Spirobolida, Callipodida, Chordeumatida and Polydesmida) and eight families were discovered. Among them, six species are new to science: Zephronia linkouzi sp. nov., Paracortina inflata sp. nov., Orthomorpha laminata sp. nov., Polylobosoma corollifera sp. nov., Epanerchodus wuxi sp. nov. and Riukiaria spina sp. nov. As usual, the polydesmidan family Paradoxosomatidae is the most diverse group, containing five species. All new species are described and illustrated, and photographs of the habitus and the gonopods of all species are also provided.

Zephronia linkouzi sp. nov. Figs 1–4 Type materials. Holotype male: China, Chongqing, Wuxi County, Yintiaoling National Nature Reserve, Linkouzi, 31°28’19.47” N, 109°52’58.34” E, alt. 1680 m, 17 August 2022, X.K. Jiang & H.M. Chen leg. Paratypes: 6 males and 2 females, same data as holotype; 1 female, same locality, 18 August 2022, X.K. Jiang & H.M. Chen leg. Diagnosis. This species is very similar to Zephronia hui Liu & Wesener, 2022 from Jiangkou County, Guizhou Province, China (Zhao et al. 2022), but it differs from the latter by the antennae slender (Fig. 2A) (short and stout in Z. hui), the process of telopoditomere 2 of the anterior telopod obviously shorter than the length of telopoditomeres 3 and 4 (Figs 3C, 4C) (subequal in Z. hui). Etymology. This species is named after the type locality, Linkouzi, noun in apposition. Description. Male body length ca. 19–31 mm. Width of thoracic shield = 8.0–11.0 mm, of tergite 8 = 8.8–12.0 mm. Height of thoracic shield = 4.3–6.0 mm, of tergite 8 = 6.0–8.2. Holotype 31 mm long, 11.0 mm (thoracic shield) wide, 6.0 mm (thoracic shield) high, 12.0 (tergite 8) wide, 8.2 (tergite 8) high. Female body length ca. 32–34 mm long. Width, of thoracic shield = 10.8–11.0 mm, of tergite 8 = 11.9–12.0 mm. Height, of thoracic shield = 6.4–6.9 mm, of tergite 8 = 8.3–8.9 mm. Body generally brown with irregular dark marks scattered on body surface (Fig. 1). Eyes with ca. 55 ocelli. Antennae short and thick. Last antennomere obviously longer than other antennomeres, lengths of antennomeres: 1=2=3=4=5< Male gonopore covered with a small, inconspicuous plate. Telopoditomere 1 of anterior telopods (Figs 3A, B; 4A, B) stout, slightly wider than long. Telopoditomere 2 as long as telopoditomere 3 in anterior view. Process of telopoditomere 2 originated posteriorly, broader than telopoditomere 3, tapering apically, curved, and protruding as high as basal part of telopoditomere 4. Inner margin towards movable finger with one large, membranous lobe. Posterior surface with a row of 5 crenulate and sclerotized teeth (Figs 3C, 4C). Telopoditomere 4 short, with at inner margin two long spines (Figs 3A–C, 4A–C). Mesial and lateral parts of telopoditomere 1 and anterior and lateral parts of telopoditomere 2 covered by setae. Other parts glabrous. Telopoditomere 1 of posterior telopods cylindrical, twice as long as wide. telopoditomere 2 stout (Figs 3D, E; 4D, E). Immovable finger (process of telopoditomere 2) slightly shorter than movable finger, consisting of telopoditomeres 3 and 4. Margin towards movable finger with two large, membranous lobes. Tip of immovable finger with a white spot in posterior view. Telopoditomere 3 elongated, 3 times longer than telopoditomere 4. Margin near immovable finger with a large membranous lobe and two slender spine, posterior surface with a row of 15 crenulate and sclerotized teeth. Telopoditomere 4 with at inner margin two long spines and a single membranous lobe. Entire telopod with few setae, mostly at anterior margins of telopoditomere 1 and inner margins of telopoditomere 2 (Figs 3D, E; 4D, E). Female vulva (Fig. 2E) large, covering 2/3 of coxa, located at mesal margin, extending mesally to basal third of prefemur length. Operculum rounded. Subanal plate large and wide, not subdivided. Distribution. Known only from the type locality.

Zephronia linkouzi sp. nov. Figs 1–4 Type materials. Holotype male: China, Chongqing, Wuxi County, Yintiaoling National Nature Reserve, Linkouzi, 31°28’19.47” N, 109°52’58.34” E, alt. 1680 m, 17 August 2022, X.K. Jiang & H.M. Chen leg. Paratypes: 6 males and 2 females, same data as holotype; 1 female, same locality, 18 August 2022, X.K. Jiang & H.M. Chen leg. Diagnosis. This species is very similar to Zephronia hui Liu & Wesener, 2022 from Jiangkou County, Guizhou Province, China (Zhao et al. 2022), but it differs from the latter by the antennae slender (Fig. 2A) (short and stout in Z. hui), the process of telopoditomere 2 of the anterior telopod obviously shorter than the length of telopoditomeres 3 and 4 (Figs 3C, 4C) (subequal in Z. hui). Etymology. This species is named after the type locality, Linkouzi, noun in apposition. Description. Male body length ca. 19–31 mm. Width of thoracic shield = 8.0–11.0 mm, of tergite 8 = 8.8–12.0 mm. Height of thoracic shield = 4.3–6.0 mm, of tergite 8 = 6.0–8.2. Holotype 31 mm long, 11.0 mm (thoracic shield) wide, 6.0 mm (thoracic shield) high, 12.0 (tergite 8) wide, 8.2 (tergite 8) high. Female body length ca. 32–34 mm long. Width, of thoracic shield = 10.8–11.0 mm, of tergite 8 = 11.9–12.0 mm. Height, of thoracic shield = 6.4–6.9 mm, of tergite 8 = 8.3–8.9 mm. Body generally brown with irregular dark marks scattered on body surface (Fig. 1). Eyes with ca. 55 ocelli. Antennae short and thick. Last antennomere obviously longer than other antennomeres, lengths of antennomeres: 1=2=3=4=5

Zephronia golovatchi Srisonchai, Sutcharit & Likhitrakarn, 2021 New locality data: 1 &male;, ZFMK MYR6262, Nakhon Ratchasima Province, Pak Ching District, near Highway 3052, 14°30'59.2''N, 101°23'52.9''E, 10.VI.2017; 1 &male;, 2 &female;&female;, ZFMK MYR6779, same data as previous; 1 &male;, ZFMK MYR7236, same data as previous; 2 &female;&female;, NHMD K53, Nakhon Nayok Province, Khao Yai National Park, 700 m, leg. 29.IX–6.X.1984, leg. Karsholt, Lomholdt & Nielsen. Remarks. Both localities lie close to the type locality (Fig. 2A). However, specimens from both localities differ significantly genetically and might represent two different species. Updated key to confirmed species of Zephronia in Thailand 1: Tarsi of leg pair 4–21 with single apical spine, third leg pair without apical spine. Male antennae, antennomere 6 swollen, not axe-shaped........................................ Z. viridisoma Rosenmejer & Wesener in Rosenmejer et al. 2021 - Tarsi of leg pair 4–21 with 3 apical spines, third leg pair with apical spine. Male antennae, antennomere 6 axe-shaped...... 2 2: Conspicuous colour pattern with a vivid green, glabrous or dull, not golden setae. Palpi of gnathochilarium, sensilla in clusters............................................................................................. 3 - Brown, grey, or brown-greenish color pattern with dense coat of 'golden' setae. Palpi of gnathochilarium, sensilla in single field................................................................................................ 4 3: Small bodied, around 20 mm. Unique color pattern, green-whitish with red on the anal shield. Tergites setose, dull. Anterior telopod, podomere 3 with up to 7 crenulated teeth. Antennae, antennomere 6 with 50–60 cones.... Z. siamensis Hirst, 1907 - Large, around 40 mm. Unique color pattern, with a vivid green-lightbrownish stripes. Tergite glabrous, shiny. Anterior telopod variable, without or only very few sclerotized teeth. Antennae, antennomere 6 with 70–90 cones........................................................................ Z. phrain Likhitrakarn & Golovatch in Likhitrakarn et al. 2021 4: Male second leg, mesal margin extended in short membranous process........................................... 5 - Male second leg, mesal margin not extended................................................................ 7 5: Endotergum, two rows of marginal bristles which project beyond tergite margin..................................... 6 - Endotergum, two rows of marginal bristles which end clearly before the tergite margin................ Z. erawani sp. nov. 6: Female vulval operculum margin straight and wide.Antennae, antennomere 6 with 90–100 cones. Male anal shield bell-shaped.......................................................................... Z. golovatchi Srisonchai et al., 2021 - Female vulval operculum narrow in posterior view. Antennae, antennomere 6 with around 75 cones. Male anal shield rectangular................................................................. Z. enghoffi Srisonchai et al. 2021 7: Brown. Inner area of endotergum without setae......... Z. lannaensis Likhitrakarn & Golovatch in Likhitrakarn et al. 2021 - Greyish. Inner area of endotergum with setae............................................................... 8 8: Small, 20–22 mm long. Legs with 7–9 ventral spines. Male antennae, antennomere 6 with ca. 50 cones. Pleurite 1 extended into long, sharp tip. Endotergum with two rows of marginal bristles........................ Z. panhai Srisonchai et al. 2021 - Medium, 40 mm long. Legs with 10–12 ventral spines. Male antennae, antennomere 6 with 78–88 apical cones. Pleurite 1 weakly extended, well-rounded. Endotergum with three rows of marginal bristles............... Z. chrysomallos sp. nov., Published as part of Bhansali, Sneha & Wesener, Thomas, 2022, New Thai giant pill-millipede species, with new genetic barcoding data (Diplopoda Sphaerotheriida, Zephroniidae), pp. 357-380 in Zootaxa 5105 (3) on pages 376-377, DOI: 10.11646/zootaxa.5105.3.2, http://zenodo.org/record/6332731, {"references":["Srisonchai, R., Sutcharit, C. & Likhitrakarn, N. (2021) The giant pill-millipede genus Zephronia Gray, 1832 from Thailand, with a redescription of Z. siamensis Hirst, 1907 and descriptions of three new species (Diplopoda, Sphaerotheriida, Zephroniidae). ZooKeys, 1067, 19 - 56. https: // doi. org / 10.3897 / zookeys. 1067.72369","Rosenmejer, T., Enghoff, H., Moritz, L. & Wesener, T. (2021) Integrative description of new giant pill-millipedes from southern Thailand (Diplopoda, Sphaerotheriida, Zephroniidae). European Journal of Taxonomy, 762, 208 - 232. https: // doi. org / 10.5852 / ejt. 2021.762.1457","Hirst, A. S. (1907) On four new pill-millipedes from the Malay Peninsula and Siam. The Annals and Magazine of Natural History, Series 7, 20 (117), 215 - 219. https: // doi. org / 10.1080 / 00222930709487327","Likhitrakarn, N., Golovatch, S. I., Srisonchai, R. & Sutcharit, C. (2021) Two New Species of the Giant Pill-Millipede Genus Zephronia Gray, 1832 from Thailand (Diplopoda: Sphaerotheriida: Zephroniidae). Tropical Natural History, 21, 12 - 26."]}

Zephronia erawani Bhansali &Wesener, new species Figures 2A, 12, 13, 14. Diagnosis. The position of the organ of Tömösváry at the brim and not in the antennal groove identifies this species as a member of the Zephronia senso-stricto group with which it also aligns weakly genetically. The sensory cones on the palpi of the gnathochilarium are arranged in a single field, not in separate clusters and the body is covered in a dense coat of golden setae like in several other Thai Zephronia. The greyish-brown color resembles those of Z. chrysomallos sp. nov. and Z. panhai. The male second leg mesal coxal margin is extended into a membranous process (Fig. 13B arrow), a character shared only with Z. golovatchi and Z. enghoffi. Z. erawani sp. nov. differs from both Z. golovatchi and Z. enghoffi in the shorter marginal bristles of the endotergum which end well before the margin, while the bristles are projecting above the two in both other species. Z. erawani sp. nov. differs genetically in the COI barcoding fragment from Z. panhai by a p-distance of 11.6%, from Z. chrysomallos sp. nov. by a p-distance of 11.9% and from Z. lannaensis by 12.6–13.3% (Sup. file 1). Derivatio nominis. Named after the type locality, the Erawan waterfalls. Erawan is a mythological elephant, similar to the Hindi Airavata. Noun in apposition. Material examined (total: 1 &male;) Holotype: THAILAND: 1 &male;, NHMD K56-9, Kanchanaburi Province, Si Sawat District, 50 km W of Kanchanaburi, Erawan Waterfall, [probably: 14°21'31.32"N, 99° 8'26.23"E, 377 m], 6.VI.1990, leg. A. R. Rasmussen & B. Helwigh. Description (based on holotype) Size. Length: 31.4 mm. Width of thoracic shield 14.8 mm, of widest segment (6) 16.5 mm. Height of thoracic shield 9.3 mm, of highest segment (6) 9.8 mm. Colour. Faded after 30 years in 75% ethanol. Collum medium brown, head and legs with traces of green. Antenna green. Tergites and anal shield pale greyish-brown, darker brown towards posterior margin. Thoracic shield-like tergite with darker groove and margins. Anal shield colour like tergite. Head. Number of ommatidia 95. Tömösváry organ placed on the rim of antennal groove. Antennae short, not reaching to center of head (Fig. 12A–D). Antennomere lengths: 1>2=3=4=5< Epipharynx. Not dissected. Gnathochilarium (Fig. 12D). Gnathochilarium palpi with the sensory cones arranged in single field. Mandible. Not dissected. Collum. Numerous setae, distributed towards margins, few setae in central region. Thoracic shield. Thoracic shield grooves shallow and wide with 2 or 3 weak ledges (Fig. 12A, B). First stigmatic plate. Smaller than coxa, rounded apex. Tergites. Tergites covered in short, dense setae visible at higher magnification, with dull orange skin-like surface. Paratergite tips weakly projecting backwards (Fig. 12A, B). Pleurites (Fig. 12C). Pleurite 1 projecting, tip well-rounded. Pleurite 2 well-rounded, weakly projecting. Endotergum. With a regular flat margin. Outer zone with three dense rows of irregular marginal setae, not extending beyond posterior margin, but reaching 4/5 of the outer area. Middle section with cuticular impressions. Inner area without setae. Legs. Ventral spines on leg 1 2/2, at 2 4 /4, at 3 7/7. Leg 3 with single apical spine. Mid-body legs with 2 or 3 apical and 8–10 ventral spines (Fig. 13A). Inner margin of femur with 14 small triangular teeth, concentrated at apical half, but not excavated. Femur 1.7, tarsus 3.6 times longer than wide. Anal shield. Bell-shaped and like tergites covered with short dense setae visible at higher magnification. Underside with single black locking carina of medium length, located close to pleurite (Fig. 12C). Male gonopore. Located directly on mesal margin of coxa 2, covered posteriorly by a single semicircular sclerotized plate. Membrane rising mesally into a projection (Fig. 13B). Anterior telopods (Fig. 13C–E). Syncoxite with few setae. Podomere 1 rectangular, as long as wide. Numerous setae, mainly at median margins. Podomere 2 slightly narrower than podomere 1. Process massive, well-rounded, apically slightly tapering, relatively short and wide (Fig. 13D, E). Process visible protruding and exceeding podomere 3 in anterior view (Fig. 13D). Podomere 3 cylindrical, with a black spot at the anterior margin visible in posterior view. Podomere 4 partially fused to podomere 3, suture can be seen in anterior view (Fig. 13C), while it is completely fused in posterior, lateral and mesal views. Podomere 4 small, narrowing at the apex. Meso-anterior margin with one spine and a black spot at apex visible in posterior and lateral view. Posterior telopods (Fig. 14). Syncoxite inner horns (not drawn): apex well rounded, apically diverging. Podomere 1 with short setae distributed in median and lateral margin, median margin glabrous. Podomere 2 with long and wide immovable finger, 2.5 times longer than wide. Podomere 2 with numerous setae visible in anterior view, almost entirely glabrous in posterior view. Immovable finger slightly narrow anteriorly and straight. Apical process with sclerotized round spots in anterior view (Fig. 14A, arrow). Membrane toward podomere 3 with a two-tipped membranous lobe. Podomere 3 setose in anterior view, with more setae concentrated medially. With row of 15 large, crenulated teeth on posterior side (Fig. 14B, arrow). Large membranous ledge on podomere 3 with single spine present basal region. Podomere 4 glabrous on both sides, slightly curved towards immovable finger. Membranous lobe with 2 long spines. Immovable finger shorter than movable finger. Distribution Z. erawani sp. nov. is currently only known from the type locality (Fig. 2A) and might be microendemic, as are other, closely related species in nearby forests.

Zephronia phrain Likhitrakarn & Golovatch in Likhitrakarn et al. 2021 New locality data: 1 &male;, SMF, Chiang Mai Province, Chiang Dao District 20 km NNE of Chiang Dao, Tham Houay Luk, 19°32'18''N, 99°2'44''E, 623 m, outside cave, by hand, leg. 26.VI.2014, P. Jäger & E. Grall; 1 &male;, SMF, Chiang Mai Province, Chiang Dao District, Doi Chiang Dao, Ma Lee's Resort, 19°24'12''N, 98°55'34''E, 470 m, garden, by hand, by day and night, leg. 24–30.VI.2014, P. Jäger & E. Grall; 1 &male;, 1 &female;, SMF, Chiang Mai Province, Chai Prakan District, near Si Dong Yen, ThamNgam [=pretty cave], 19°39'19.46''N, 99°6'11.10''E, 660 m, inside cave and in front of cave, by day, by hand, leg. 29.VI.2014, P. Jäger & E. Grall; 1 &male;, ZFMK MYR4907, Chiang Mai, Chiang Dao, 19°23'54" N, 98°55'47" E, 448 m; 2 &male;&male;, MHNG 5 G, Lamphun Province, Mae Tha District, Doi Khuntan National Park, 1100 m, evergreen hill forest, 18°29'35.83"N, 99°17'6.43"E, 8.X.1992, leg. A. Schwendinger; 1 &male;, MHNG 5 I, Province Chiang Mai, Chiang Dao District, Doi Chiang Dao, 400 m, 7.VII.1987, leg. P. Schwendinger; 1 &male;, ZFMK MYR3499, Chiang Mai Province, District Chiang Dao, Padeng Lodge, 03.VII.2014, leg. S. Huber; 1 &male;, ZFMK MYR3500, Chiang Mai Province and District, Doi Suthep, behind tourist market, 18°48'03'' N, 98°55'05'' E, 2040 m, leg. 29.VI.2014, S. Huber; 1 &male;, ZFMK MYR4907, Chiang Mai Province, Chiang Dao, 19°23'54''N, 98°55'47'' E, 448 m, 14–16.VIII.2015. Remarks. This species, as already mentioned previously (Srisonchai et al. 2021), is more widespread than previously thought (Fig. 2A). We observed a lot of intraspecific variation in the genetic barcode (Sup. file 1) as well as in the telopods of Z. phrain, so there might be cryptic species under this name.

Zephronia erawani Bhansali &Wesener, new species Figures 2A, 12, 13, 14. Diagnosis. The position of the organ of Tömösváry at the brim and not in the antennal groove identifies this species as a member of the Zephronia senso-stricto group with which it also aligns weakly genetically. The sensory cones on the palpi of the gnathochilarium are arranged in a single field, not in separate clusters and the body is covered in a dense coat of golden setae like in several other Thai Zephronia. The greyish-brown color resembles those of Z. chrysomallos sp. nov. and Z. panhai. The male second leg mesal coxal margin is extended into a membranous process (Fig. 13B arrow), a character shared only with Z. golovatchi and Z. enghoffi. Z. erawani sp. nov. differs from both Z. golovatchi and Z. enghoffi in the shorter marginal bristles of the endotergum which end well before the margin, while the bristles are projecting above the two in both other species. Z. erawani sp. nov. differs genetically in the COI barcoding fragment from Z. panhai by a p-distance of 11.6%, from Z. chrysomallos sp. nov. by a p-distance of 11.9% and from Z. lannaensis by 12.6–13.3% (Sup. file 1). Derivatio nominis. Named after the type locality, the Erawan waterfalls. Erawan is a mythological elephant, similar to the Hindi Airavata. Noun in apposition. Material examined (total: 1 &male;) Holotype: THAILAND: 1 &male;, NHMD K56-9, Kanchanaburi Province, Si Sawat District, 50 km W of Kanchanaburi, Erawan Waterfall, [probably: 14°21'31.32"N, 99° 8'26.23"E, 377 m], 6.VI.1990, leg. A. R. Rasmussen & B. Helwigh. Description (based on holotype) Size. Length: 31.4 mm. Width of thoracic shield 14.8 mm, of widest segment (6) 16.5 mm. Height of thoracic shield 9.3 mm, of highest segment (6) 9.8 mm. Colour. Faded after 30 years in 75% ethanol. Collum medium brown, head and legs with traces of green. Antenna green. Tergites and anal shield pale greyish-brown, darker brown towards posterior margin. Thoracic shield-like tergite with darker groove and margins. Anal shield colour like tergite. Head. Number of ommatidia 95. Tömösváry organ placed on the rim of antennal groove. Antennae short, not reaching to center of head (Fig. 12A–D). Antennomere lengths: 1>2=3=4=5

Zephronia lannaensis Likhitrakarn & Golovatch, 2021 in Likhitrakarn et al. 2021 New locality data: 3 &male;&male;, ZFMK MYR4911, Chiang Mai Province, Mae Rim District, Mae Sa Valley, 18°53'54"N, 98°51'16"E, 650 m, 31.VIII–03.IX.2015; 2 &male;&male;, NHMD K57B, Chiang Mai Province and District, Doi Suthep, Mae Sa Waterfall, 29.V.1991, leg. C.C. Kinze; 1 &male;, MHNG 3 B; Chiang Mai Province & District, Doi Suthep, 800 m, evergreen gallery forest, 12.V.1986, leg. P. Schwendinger; 1 &female;, ZFMK MYR3498, Chiang Mai Province and District, foot of western side of Doi Suthep, 18°48'30'' N, 98°56'49'' E, 360 m. 05.VII.2014, leg S. Huber; 1 &male;, ZFMK MYR3501, Chiang Mai Province, Mae Rim District, Traidhos School Campus, 18°57'57'' N, 98°55'05'' E, 330 m, 30.VI.2014, leg. S. Huber; 3 &male;&male;, ZFMK MYR4912, Chiang Mai Province, Mae Rim District, Mae Sa Valley, 18°53'54'' N, 98°51'16'' E, 650 m, 31.VIII–03.IX.2015. Remarks. The localities mentioned above lie close to the type locality (Fig. 2A)., Published as part of Bhansali, Sneha & Wesener, Thomas, 2022, New Thai giant pill-millipede species, with new genetic barcoding data (Diplopoda Sphaerotheriida, Zephroniidae), pp. 357-380 in Zootaxa 5105 (3) on page 376, DOI: 10.11646/zootaxa.5105.3.2, http://zenodo.org/record/6332731, {"references":["Likhitrakarn, N., Golovatch, S. I., Srisonchai, R. & Sutcharit, C. (2021) Two New Species of the Giant Pill-Millipede Genus Zephronia Gray, 1832 from Thailand (Diplopoda: Sphaerotheriida: Zephroniidae). Tropical Natural History, 21, 12 - 26."]}

Sphaerobelum meridionalis Bhansali & Wesener, new species Figures 2A, 3, 4, 5, 6A. Diagnosis. Differs from all other species of the genus Sphaerobelum aside from S. aesculus in the shape of the posterior telopod, where there is a swelling at the tip of the immovable finger, but the swelling does not extend above the margin (Fig. 5B). Both species also share numerous other characteristics, such as general size and color, the glabrous, leather-like surface of the tergites and the shape of the anal shield, the low number of ventral spines on the legs, as well as the absence of apical spines on leg 3. S. meridionalis sp. nov. differs from S. aesculus in several characters, such as the presence of a distinct mesal process on the prefemur of midbody legs (Fig. 4B, arrow) (absent in S. aesculus), a mesal coxal process on the first coxae (Fig. 4A) (absent in S. aesculus), the telopoditomeres 3 and 4 of the anterior telopods being clearly separated (Fig. 4F) (being almost fused in S. aesculus) as well as some minor characters such as the lower number of ventral spines on the first tarsi (1 versus 4 in S. aesculus) and the slightly higher number of apical cones on the antenna (68–73 versus 52–56 in S. aesculus) (Fig. 3B). S. meridionalis sp. nov. differs from S. aesculus by an uncorrected p-distance on the COI gene of 17% (Sup. file 1). Derivatio nominis. meridionalis, Latin for southern, adjective used as noun. Noun in apposition. Material examined (total: 1 &male;) Holotype THAILAND: 1 &male; (fragmented), MHNG 4 B-2, Yala Province Yala, Bannang Sata District, Bang Lang National Park, near Than To Waterfall, [probably 6°11'47.50"N, 101° 9'50.90"E], 150 m, 1.II.1991, leg. P. Schwendinger. Description (based on holotype) Size. Length 26.8 mm. Width of thoracic shield 13.3 mm, of widest segment 14.4 mm. Height of thoracic shield 7.9 mm, Height of highest segment (8) 8.8mm. Color. Faded after 30 years in 75% ethanol. Head medium brown. Antenna pale light brown with traces of green and a brown border. Last antennomere distinctly light green. Legs pale brown, tarsi greenish. Tergites and anal shield medium brown with green color, ventral side light green. Posterior margin of tergites medium brown. Thoracic shield-like tergites but groove greenish. Head. Number of ommatidia low, 61–65. Tömösváry organ located in antennal groove. Antennae short, reaching to center of head, antennomere lengths: 6>1>3=4>2=5. Last antennomere apically slightly swollen (Fig. 3A), number of apical cones 73/68 (Fig. 3B). Mandible. Not dissected. Gnathochilarium. Lingual lamella with numerous long setae. Palpi with sensory cones arranged in single field. Collum. Setae distributed towards borders, more concentrated at lateral ends. Glabrous on posterior and median region. Thoracic shield. Grooves wide and deep, with 5 or 6 sclerotized ledges along inner ridge. Surface like tergites. Tergites. Paratergite tips on posterior half projecting backwards. Tergites glabrous with dull leather-like surface. Endotergum (Fig. 6A). With a regular flat margin. Outer zone with three rows of irregular marginal setae, not extending beyond posterior margin, but reaching 4/5 of outer area. Middle section with a single row of distant, oval cuticular impressions. Distance between impressions twice the diameter. Inner area with numerous rows of setae, widely distributed, shorter than marginal setae. First stigmatic plate (Fig. 4A). Large, larger than coxa, with a well-rounded apex. Pleurites. Pleurite 1 weakly projecting posteriorly with a sharp apex. Pleurite 2 short with well-rounded apex, not projecting. Legs. Ventral spines on leg 1 1/1, on 2 2/2, on 3 4/4. Apical spine absent on leg 3. Three apical and 6 or 7 ventral spines on midbody legs (Fig. 4B). Inner margin of femur with up to 20 small teeth, but not excavated. Femur 1.7, tarsus 3.8 times longer than wide. Prefemur with conspicuous mesal process (Fig. 4B arrow). First coxae with conspicuous mesal process (Fig. 4A). Anal shield. Well-rounded and glabrous. Ventral side with single medium length locking carina, placed close to margin. Male sexual characters. Gonopore (Fig. 4C) large, consisting of large membranous opening located directly at mesal margin of second coxa, covered posteriorly by a single semicircular sclerotized plate. Anterior telopods (Figs 4D–G). Syncoxite with few setae. Podomere 1 rectangular 1.2 times longer than wide, more densely covered with setae in apical half than at basal margin, with longer setae medially, basal margin glabrous. Podomere 2 with wide process visible in anterior view, protruding to half-length of podomere 3. Process curved, well-rounded, apically slightly tapering (Fig. 4D). Podomere 3 cylindrical, almost twice as long as wide, latero-apical process juxtaposed to process of podomere 2. One spine visible in lateral view. Podomere 4 clearly separated from podomere 3, cylindrical, of half-length of podomere 3, with one spine below apex visible in posterior and lateral view. Posterior telopods (Fig. 5). Syncoxite inner horns (not drawn): well-rounded, apically tapering. Podomere 1 rectangular, as long as wide, with moderately long setae. Podomere 2 with a single, large triangular membranous lobe at base of immovable finger. Immovable finger wide, straight, slightly wider anteriorly and apex tapering towards movable finger in a small process. Process with sclerotized round spots in anterior view. Single large oval membranous spot visible in posterior view. Surface of finger glabrous except for margins. Podomere 3 circa 3 times longer than wide. Membranous ledge with a single spine, Posterior side with a row of 7 small crenulated teeth. Glabrous except for several long setae clustered at lower inner margin. Podomere 4 short, apically tapering, slightly curved towards immovable finger, ca. 2.5 times longer than wide. Membranous ledge and 2 spines present mesal margin. Glabrous. Distribution Only known from the area near the Than To Waterfall in the very south of Thailand (Fig. 2A)., Published as part of Bhansali, Sneha & Wesener, Thomas, 2022, New Thai giant pill-millipede species, with new genetic barcoding data (Diplopoda Sphaerotheriida, Zephroniidae), pp. 357-380 in Zootaxa 5105 (3) on pages 364-366, DOI: 10.11646/zootaxa.5105.3.2, http://zenodo.org/record/6332731

Zephronia phrain Likhitrakarn & Golovatch in Likhitrakarn et al. 2021 New locality data: 1 &male;, SMF, Chiang Mai Province, Chiang Dao District 20 km NNE of Chiang Dao, Tham Houay Luk, 19°32'18''N, 99°2'44''E, 623 m, outside cave, by hand, leg. 26.VI.2014, P. Jäger & E. Grall; 1 &male;, SMF, Chiang Mai Province, Chiang Dao District, Doi Chiang Dao, Ma Lee's Resort, 19°24'12''N, 98°55'34''E, 470 m, garden, by hand, by day and night, leg. 24–30.VI.2014, P. Jäger & E. Grall; 1 &male;, 1 &female;, SMF, Chiang Mai Province, Chai Prakan District, near Si Dong Yen, ThamNgam [=pretty cave], 19°39'19.46''N, 99°6'11.10''E, 660 m, inside cave and in front of cave, by day, by hand, leg. 29.VI.2014, P. Jäger & E. Grall; 1 &male;, ZFMK MYR4907, Chiang Mai, Chiang Dao, 19°23'54" N, 98°55'47" E, 448 m; 2 &male;&male;, MHNG 5 G, Lamphun Province, Mae Tha District, Doi Khuntan National Park, 1100 m, evergreen hill forest, 18°29'35.83"N, 99°17'6.43"E, 8.X.1992, leg. A. Schwendinger; 1 &male;, MHNG 5 I, Province Chiang Mai, Chiang Dao District, Doi Chiang Dao, 400 m, 7.VII.1987, leg. P. Schwendinger; 1 &male;, ZFMK MYR3499, Chiang Mai Province, District Chiang Dao, Padeng Lodge, 03.VII.2014, leg. S. Huber; 1 &male;, ZFMK MYR3500, Chiang Mai Province and District, Doi Suthep, behind tourist market, 18°48'03'' N, 98°55'05'' E, 2040 m, leg. 29.VI.2014, S. Huber; 1 &male;, ZFMK MYR4907, Chiang Mai Province, Chiang Dao, 19°23'54''N, 98°55'47'' E, 448 m, 14–16.VIII.2015. Remarks. This species, as already mentioned previously (Srisonchai et al. 2021), is more widespread than previously thought (Fig. 2A). We observed a lot of intraspecific variation in the genetic barcode (Sup. file 1) as well as in the telopods of Z. phrain, so there might be cryptic species under this name., Published as part of Bhansali, Sneha & Wesener, Thomas, 2022, New Thai giant pill-millipede species, with new genetic barcoding data (Diplopoda Sphaerotheriida, Zephroniidae), pp. 357-380 in Zootaxa 5105 (3) on pages 375-376, DOI: 10.11646/zootaxa.5105.3.2, http://zenodo.org/record/6332731, {"references":["Likhitrakarn, N., Golovatch, S. I., Srisonchai, R. & Sutcharit, C. (2021) Two New Species of the Giant Pill-Millipede Genus Zephronia Gray, 1832 from Thailand (Diplopoda: Sphaerotheriida: Zephroniidae). Tropical Natural History, 21, 12 - 26.","Srisonchai, R., Sutcharit, C. & Likhitrakarn, N. (2021) The giant pill-millipede genus Zephronia Gray, 1832 from Thailand, with a redescription of Z. siamensis Hirst, 1907 and descriptions of three new species (Diplopoda, Sphaerotheriida, Zephroniidae). ZooKeys, 1067, 19 - 56. https: // doi. org / 10.3897 / zookeys. 1067.72369"]}

Zephronia panhai Srisonchai, Sutcharit & Likhitrakarn, 2021 New locality data: 1 &female;, ZFMK MYR8116, Ratchaburi Province, Ratchaburi and Photharam District, 18–20 km WNW of Ratchaburi, N: 13°35'17"–13°43'07", E: 99°40'07"–99°46'21", 30–80 m; 1 &female;, ZFMK MYR8118, same data as previous; 1 &male;, 1 &female;, 3 imm., ZFMK MYR8128, same data as previous; 5 &male;&male;, 4 &female;&female;, ZFMK MYR8134, same data as previous; 4 &male;&male;, 1 &female;, MHNG 3 A, Ratchaburi Province, Chombueng District, Tham Khao Bin Forest Park, 70 m, secondary forest on limestone, 13°35'12''N, 99°40'01''E, 27/ 28.VI.2011, leg. P. Schwendinger, sample TH-11/01. Remarks. The localities mentioned above are close to the type locality (Fig. 2A)., Published as part of Bhansali, Sneha & Wesener, Thomas, 2022, New Thai giant pill-millipede species, with new genetic barcoding data (Diplopoda Sphaerotheriida, Zephroniidae), pp. 357-380 in Zootaxa 5105 (3) on page 376, DOI: 10.11646/zootaxa.5105.3.2, http://zenodo.org/record/6332731, {"references":["Srisonchai, R., Sutcharit, C. & Likhitrakarn, N. (2021) The giant pill-millipede genus Zephronia Gray, 1832 from Thailand, with a redescription of Z. siamensis Hirst, 1907 and descriptions of three new species (Diplopoda, Sphaerotheriida, Zephroniidae). ZooKeys, 1067, 19 - 56. https: // doi. org / 10.3897 / zookeys. 1067.72369"]}

Zephronia chrysomallos Bhansali & Wesener, new species Figures 2A–B, 6B, 7, 8, 9, 10, 11. Diagnosis. The position of the organ of Tömösváry at the brim and not in the antennal groove identifies this species as a member of the Zephronia sensu stricto group, with which it also aligns weakly genetically (Fig. 1). The sensory cones on the palpi of the gnathochilarium are arranged in a single field, not in separate clusters (Fig. 8D) and the body is covered in a dense coat of golden setae like in several other Thai Zephronia. The greyish brown tergites color resembles those of Z. erawani sp. nov. and Z. panhai (Fig. 2B). The male second leg mesal coxal margin is not extended into a membranous process, a character shared only with Z. lannaensis and Z. panhai. Zephronia chrysomallos sp. nov. differs from Z. lannaensis in the color (grey versus brown), telopod characters as well as the endotergum. Z. chrysomallos sp. nov. differs from Z. panhai in being much larger in size (40 mm versus 20 mm), the higher number of ventral spines on the walking legs (10–12 versus 7–9), the higher number of apical cones on the antenna (78–88 versus around 50), the higher number of rows of marginal bristles on the endotergum (3 versus 2) and the shape of the first pleurite, which is weakly extended and well-rounded in Z. chrysomallos sp. nov. but with a long sharp process in Z. panhai. Z. chrysomallos sp. nov. differs genetically in the COI barcoding fragment from Z. panhai by a p-distance of 10.8%, from Z. erawani sp. nov. by a p-distance of 11.9% and from Z. lannaensis by 14.8–15.2% (Sup. file 1). Derivatio nominis. chrysomallos, Greek, meaning Golden Wool, and also the flying sheep ram Chrysomallos from Greek mythology, which was sacrificed to Zeus in order to obtain the famous Golden Fleece. Noun in apposition. Material examined (total: 7 &male;&male;) Holotype THAILAND: 1 &male;, ZFMK MYR8826, Kanchanaburi Province, Sai Yok District, Sai Yok Noi Waterfall, 14°14'12'' N, 99°03'46'' E, 150 m. Paratypes THAILAND: 1 &male;, ZFMK MYR11366, CT scan voucher, same data as for holotype; 1 &male;, ZFMK MYR11365, same data as for holotype. Other material THAILAND: 4 &male;&male;, CUMZ, Kanchanaburi Province, Sai Yok District, Sai Yok Noi Waterfall (Khao Phang WF), 14°14'21'' N, 99°03'28'' E, ca. 160 m a.s.l., 11.VII.2009, leg. ASRU members. Description (based on holotype) Size. Length: ca. 40 mm. Width of thoracic shield 16.3 mm, of widest segment (8th) 16.9 mm. Height of thoracic shield 9.4 mm, of highest segment (8th) 11.5 mm. Color (from living specimens). Head, collum and anterior margin of thoracic shield dark brown to black. Antenna dark brown, legs pale brown. Tergites and anal shield greyish-brown (fading to light brown in ethanol), darker towards posterior margin. Thoracic shield-like tergite with darker anterior and posterior margins. (Fig. 2B). Head (Fig. 7D). Number of ommatidia more than 90. Tömösváry organ placed on rim of antennal groove. Antennae (Fig. 7A–D) short, axe-shaped (Fig. 8A–C), not reaching to center of head (Fig. 7C), antennomere lengths: 6>1>5=4=3=2. Antennomere 1 basally with scales (Fig. 8A). Antennomere 6 apically wide, and flattened, axe-shaped, with 78/88 apical cones (Fig. 8C). Epipharynx. With a large inner tooth. Gnathochilarium (Figs 7D, 8D–F, 9A). Lamellae linguales with numerous long setae, more towards the margins. Stipes and mentum with numerous long setae in a regular pattern (Fig. 9A). Gnathochilarium palpi with sensory cones arranged in single field (Fig. 8E). 'Pillows' of central pads with two types of sensory cones (Fig. 8F). Mandible (Fig. 9B). External tooth undivided, inner tooth with 3 lobes. Pectinate lamellae with 6 or 7 rows. Condylus at anterior margin with two distinct ridges. Collum (Figs 7A, C). With few setae distributed towards the margins, more concentrated towards lateral ends. Thoracic shield. Thoracic shield grooves wide and shallow (Figs 7A–C), without sclerotized ledges. First stigmatic plate (Fig. 10A). Smaller than coxa, with rounded sub-triangular apex. Tergites. Tergites densely covered in short, dense setae visible at higher magnification, with dull orange skinlike surface (Fig. 2B). Paratergite tips only weakly projecting backwards (Fig. 7B). Pleurites. Pleurite 1 well-rounded, weakly projecting. Pleurite 2 well-rounded, not projecting (Fig. 7C). Endotergum (Fig. 6B). With a regular flat margin. Outer area with two dense rows of marginal setae, reaching 4/5 of outer area. Middle section with a single row of rounded cuticular impression, distance between impression 2–3 times their diameter. Inner area without setae or spines. Legs. Ventral spines on leg 1 2/2, at 2 4 /4, at 3 5/5. Leg 3 with single apical spine. Mid-body legs with 3 or 4 apical and 10–12 ventral spines (Fig. 10B). Inner margin of femur with 12–15 small triangular teeth, concentrated at apical half, but not excavated. Femur 1.5, tarsus 3.3 times longer than wide. Anal shield. Well-rounded and like tergites covered with short dense setae visible at higher magnification (Fig. 2B). Ventral side with dark-colored medium-sized locking carina placed slightly closer to pleurite than margin (Fig. 7C). Male sexual characters. Male gonopore consisting of large membranous opening located directly at mesal margin of coxa 2 (Fig. 10C). Coxae 2 without membranous processes. Anterior telopods (Figs 10D–F). Syncoxite setaceous. Podomere 1 rectangular, as long as wide. Few setae, mainly at margins and in apical half. Podomere 2 in anterior view as wide as but slightly narrower as podomere 1. Process massive, visible in anterior view (Fig. 10E), relatively short and wide. Process protruding above basal half of podomere 3, triangular (Figs 10E, F). Podomere 3 cylindrical, 1.4 longer than wide, with one spine in the mesal membranous region, apically with 4 crenulated teeth (Fig. 10D). Podomere 4 clearly separated from podomere 3, conical bearing a black spot at apex. Meso-posterior margin with two spines. Posterior telopods (Fig. 11). Syncoxite inner horns (not drawn): apex well rounded, apically diverging. Podomere 1 rectangular, as wide as long, with short setae. Podomere 2 with long and narrow immovable finger, 5 times longer than wide. Immovable finger slightly wider posteriorly and apically very slightly curved towards movable finger. Immovable finger glabrous, remaining part of podomere 2 setose. Membranous lobe bearing 2 elongated processes, fused at base. Podomere 3 in posterior view glabrous, in anterior view basal half setose. With a row of 17 or 18 large crenulated square teeth on posterior side (Fig. 11B). Membranous ledge on podomere 3 with single spine. Podomere 4 on both sides glabrous, slightly curved towards immovable finger, membranous lobe with 2 spines. Immovable finger shorter than movable finger. Distribution Z. chrysomallos sp. nov. is currently only known from the type locality near the Sai Yok Noi Waterfall (Fig. 2A) and might be microendemic, as are other, closely related species in nearby forests., Published as part of Bhansali, Sneha & Wesener, Thomas, 2022, New Thai giant pill-millipede species, with new genetic barcoding data (Diplopoda Sphaerotheriida, Zephroniidae), pp. 357-380 in Zootaxa 5105 (3) on pages 367-370, DOI: 10.11646/zootaxa.5105.3.2, http://zenodo.org/record/6332731

Genus Sphaerobelum Verhoeff, 1924 Remarks. For a recent revision and a key see Semenyuk et al. (2018), Wesener (2019), Semenyuk et al. (2020), Zhao et al. (2020), Rosenmejer et al. (2021). Type species. Sphaerobelum clavigerum Verhoeff, 1924 (subsequently designated by Jeekel 1971: 28), from Vietnam. Species included. 20 (including the one described below). Distribution. Vietnam, Thailand, Laos, China., Published as part of Bhansali, Sneha & Wesener, Thomas, 2022, New Thai giant pill-millipede species, with new genetic barcoding data (Diplopoda Sphaerotheriida, Zephroniidae), pp. 357-380 in Zootaxa 5105 (3) on page 361, DOI: 10.11646/zootaxa.5105.3.2, http://zenodo.org/record/6332731, {"references":["Verhoeff, KW. (1924) Results of Dr. E. Mjoberg's Swedish scientific expeditions to Australia 1910 - 1913. Myriapoda, Diplopoda. Arkiv for zoologi, 16 (5), 1 - 142.","Semenyuk, I., Golovatch, S. I. & Wesener, T. (2018) Four new species of giant pill-millipedes from Vietnam (Sphaerotheriida, Zephroniidae). Zootaxa, 4459 (3), 535 - 550. https: // doi. org / 10.11646 / zootaxa. 4459.3.7","Wesener, T. (2019) First records of giant pill-millipedes from Laos (Diplopoda, Sphaerotheriida, Zephroniidae). Zootaxa, 4563 (2), 201 - 248. https: // doi. org / 10.11646 / zootaxa. 4563.2.1","Semenyuk, I., Golovatch, S. I. & Wesener, T. (2020) Some new or poorly-known Zephroniidae (Diplopoda, Sphaerotheriida) from Vietnam. In: Korsos, Z. & Danyi, L. (Eds.), Proceedings of the 18 th International Congress of Myriapodology, Budapest, Hungary. ZooKeys, 930, pp. 37 - 60. https: // doi. org / 10.3897 / zookeys. 930.47742","Zhao, Y., Yu, J. & Liu, W. (2020) A molecular-based phylogeny of the millipede genus Sphaerobelum Verhoeff, 1924, with the first record of the genus from mainland China (Diplopoda: Sphaerotheriida: Zephroniidae). Annales de la Societe entomologique de France, New Series, 56, 341 - 348. https: // doi. org / 10.1080 / 00379271.2020.1811153","Rosenmejer, T., Enghoff, H., Moritz, L. & Wesener, T. (2021) Integrative description of new giant pill-millipedes from southern Thailand (Diplopoda, Sphaerotheriida, Zephroniidae). European Journal of Taxonomy, 762, 208 - 232. https: // doi. org / 10.5852 / ejt. 2021.762.1457"]}

Family Zephroniidae Gray, 1842 Remarks. See Wesener (2016) for a catalogue of the family., Published as part of Bhansali, Sneha & Wesener, Thomas, 2022, New Thai giant pill-millipede species, with new genetic barcoding data (Diplopoda Sphaerotheriida, Zephroniidae), pp. 357-380 in Zootaxa 5105 (3) on page 361, DOI: 10.11646/zootaxa.5105.3.2, http://zenodo.org/record/6332731

Genus Zephronia Gray, 1832 Type species. Zephronia ovalis Gray, 1832 (by monotypy). Species included. 48, including the two described below (Wesener 2016a, 2019; Semenyuk et al. 2018, 2020; Likhitrakarn et al. 2021; Rosenmejer et al. 2021; Srisonchai et al. 2021). Distribution. NE India, Nepal, Myanmar, with a few species also in other SE Asian countries., Published as part of Bhansali, Sneha & Wesener, Thomas, 2022, New Thai giant pill-millipede species, with new genetic barcoding data (Diplopoda Sphaerotheriida, Zephroniidae), pp. 357-380 in Zootaxa 5105 (3) on page 367, DOI: 10.11646/zootaxa.5105.3.2, http://zenodo.org/record/6332731, {"references":["Gray, G. (1832) The Myriapods (Myriapoda - Mitosata). In: Griffith, E. & Pidgeon, E. (Eds.), The Class Insecta arranged by the Baron Cuvier, London, 2, pp. 1 - 796.","Semenyuk, I., Golovatch, S. I. & Wesener, T. (2018) Four new species of giant pill-millipedes from Vietnam (Sphaerotheriida, Zephroniidae). Zootaxa, 4459 (3), 535 - 550. https: // doi. org / 10.11646 / zootaxa. 4459.3.7","Semenyuk, I., Golovatch, S. I. & Wesener, T. (2020) Some new or poorly-known Zephroniidae (Diplopoda, Sphaerotheriida) from Vietnam. In: Korsos, Z. & Danyi, L. (Eds.), Proceedings of the 18 th International Congress of Myriapodology, Budapest, Hungary. ZooKeys, 930, pp. 37 - 60. https: // doi. org / 10.3897 / zookeys. 930.47742","Likhitrakarn, N., Golovatch, S. I., Srisonchai, R. & Sutcharit, C. (2021) Two New Species of the Giant Pill-Millipede Genus Zephronia Gray, 1832 from Thailand (Diplopoda: Sphaerotheriida: Zephroniidae). Tropical Natural History, 21, 12 - 26.","Rosenmejer, T., Enghoff, H., Moritz, L. & Wesener, T. (2021) Integrative description of new giant pill-millipedes from southern Thailand (Diplopoda, Sphaerotheriida, Zephroniidae). European Journal of Taxonomy, 762, 208 - 232. https: // doi. org / 10.5852 / ejt. 2021.762.1457","Srisonchai, R., Sutcharit, C. & Likhitrakarn, N. (2021) The giant pill-millipede genus Zephronia Gray, 1832 from Thailand, with a redescription of Z. siamensis Hirst, 1907 and descriptions of three new species (Diplopoda, Sphaerotheriida, Zephroniidae). ZooKeys, 1067, 19 - 56. https: // doi. org / 10.3897 / zookeys. 1067.72369"]}

Family Zephroniidae Gray, 1843 Remarks See Wesener (2016a) for a catalogue of the family., Published as part of Rosenmejer, Trine, Enghoff, Henrik, Moritz, Leif & Wesener, Thomas, 2021, Integrative description of new giant pill-millipedes from southern Thailand (Diplopoda, Sphaerotheriida, Zephroniidae), pp. 108-132 in European Journal of Taxonomy 762 (1) on page 114, DOI: 10.5852/ejt.2021.762.1457, http://zenodo.org/record/5211725, {"references":["Wesener T. 2016 a. The giant pill-millipedes, order Sphaerotheriida - An annotated species catalogue with morphological atlas and list of apomorphies (Arthropoda: Diplopoda). Bonn Zoological Bulletin - Supplementum 63: 1 - 104."]}

Zephronia viridisoma Rosenmejer & Wesener sp. nov. urn:lsid:zoobank.org:act: 272E06C8-2DE5-4722-A7D5-D85002D68424 Figs 2B, 6B – 12 Diagnosis Posterior telopod typical for the genus, not differing from those of other Zephronia species. Small (25– 28 mm long) green species (Fig. 2B), surface appearing glabrous, dull, with a single medium sized locking carina at the anal shield and a strongly projecting pleurite 1 (Fig. 9A). One of the few Zephronia species with just a single apical spine on the legs (Fig. 9B), differing in this character from all other described Thai Zephronia species which have 2–5 apical tarsal spines. Male antennomere 6 swollen (Fig. 7A) but not axe-shaped, with Holotype THAILAND • &male;; Nakhon Si Thammarat Province, Sichon District, Khao Lark Waterfall; 9°03′6″ N, 99°47′24″ E; 25 Aug. 2007; Chulalongkorn University expedition of millipede workshop leg.; dense jungle on limestone; NHMD 621695. Paratypes THAILAND • 2 &male;&male;, 4 &female;&female;; same collection data as for holotype; NHMD 621696 • 1 &male;; same collection data as for holotype; ZFMK MYR8786 • 1 &female;; same collection data as for holotype; CT scan voucher; ZFMK MYR8787. Description (based on holotype) SIZE. Length 25.6 mm. Width of thoracic shield 12.1 mm, of widest segment (8) 13 mm. Height of thoracic shield 7.1 mm, of highest segment (8) 7.7 mm. COLOUR. Head medium brown, faded from green. Antennae medium brown. Legs medium brown, tarsal claws apically dark brown. Tergites light brown, faded green medially, with darker brown posterior margin. Paratergite tips medium brown with darker edges. Anal shield dark green, edges faded to brown (Fig. 2B). HEAD (Fig. 7A–C). Number of ommatidia 50 (Fig. 7B). Organ of Tömösváry placed midway between ocelli and antennal groove (Fig. 7A), with its typical coral-like inner structure (Fig. 7C). ANTENNAE (Fig.7A,D). Reaching leg pair3,6visible antennomeres.Antennomere lengths:6>1>2=3>4=5. Antennomere 6 apically swollen, number of apical cones 32/44. EPIPHARYNX (Fig. 8A). With an extraordinarily large inner tooth. MANDIBLE (Fig. 8E–F). Inner tooth 3-cusped. Pectinate lamellae with 6 or 7 rows of teeth. Condylus at anterior margin with two ridges. GNATHOCHILARIUM (Fig. 8B–D). Lamellae linguales with numerous long setae, medially glabrous. Stipes and mentum with numerous long setae in a regular patter. Sensory cones on gnathochilarium palps in clusters (Fig. 8D). COLLUM. With short setation, evenly distributed across surface. THORACIC SHIELD. Thoracic shield grooves wide and deep, with 9 sclerotized ledges along inner ridge of grooves. TERGITES (Fig. 9D). Tergites glabrous, with dull orange skin like surface. At high magnification tiny setae and knobs become visible. Paratergite tips on posterior half projecting backwards (Fig. 12A–B). ENDOTERGUM (Fig. 6B). With a regular flat margin. Outer zone with three dense rows of irregular marginal setae, some of them extending beyond posterior margin. A single row of rounded cuticular impressions present next to marginal ridge. Intersegmental membrane smooth, without cones and with very few setae. STIGMATIC PLATE (Fig. 10A). First plate with a rounded sub-triangular apex. More posterior plates similar to those of other representatives of the family, half-covered by the pleurites (Fig. 12C). PLEURITES (Figs 9A, 12D). Pleurite 1 projecting strongly, with sharp apex. Pleurite 2 projecting slightly less than 1 and with more rounded apex (Fig. 12D). Projection absent from pleurite 3 (Fig. 12D). LEGS. Ventral spines on leg 1 1/1, on leg 2 3/2, on leg 3 4/4. Apical spine on leg 3 absent. A single apical spine and 5 or 6 ventral spines on midbody legs (Fig. 9B). Femur regularly shaped, 2.2 times longer than wide, sclerotized ledge of medium length, inner margin apically with 5 or 6 small rounded triangular spines on ventral side (Fig. 9B–C). Tarsus 4.2 times longer than wide. ANAL SHIELD. Well-rounded and glabrous. Ventral side with dark coloured. A medium-sized locking carina placed slightly closer to pleurite than margin (Fig. 12D). MALE GONOPORE (Fig. 10B). Inconspicuous, consisting of large membranous opening located directly at mesal margin. ANTERIOR TELOPODS (Fig. 10C–E). With long setae medially on the first 2 podomeres, covering posterior part on podomere 2. Podomere 1 rectangular, as long as wide. Podomere 2 in anterior view as wide as but slightly narrower than podomere 1. Immovable finger barely visible in anterior view (Fig. 10D), short, not protruding up to podomere 4 (Fig. 10C). Podomere 3 long and wide, with rounded projection on posterior side, bearing 3 crenulated teeth. Podomere 4 very narrow and short, with 3 large spines near apex, and a smaller spine near edge to podomere 3. POSTERIOR TELOPODS (Fig. 10F–G). Podomere 2 setose, immovable finger straight, apically tapering, with rows of small circular sclerotized spots. Membranous lobe between podomere 3 and 4 bearing 2 elongated processes, fused at base. Podomere 3 with 8 or 9 small crenulated teeth on posterior side. Membranous lobe on podomere 3 with single spine. Podomere 4 slightly curved towards immovable finger, membranous lobe with 2 spines. Setation on podomere 2 primarily on anterior side. Description of female paratype Length 25.6 mm. Width of thoracic shield 12.3 mm, of widest tergite (8) 13.7 mm. Height of thoracic shield: 8.4 mm, of highest tergite (8) 9.3 mm. Antennae reaching leg pair 3. Apical cones left/right: 25/25. Subanal plate well rounded. Vulvae (Fig. 11): with a large pointed operculum (Op). External lateral plate (EP) and inner mesal plate (IP) fused at bottom. Another female paratype was used for microcomputed tomography. The resulting volume rendering is shown in Fig. 12. While this approach revaled no significant additional details it serves to illustrate morphology of the specimen in an excellent way, much better than any photograph., Published as part of Rosenmejer, Trine, Enghoff, Henrik, Moritz, Leif & Wesener, Thomas, 2021, Integrative description of new giant pill-millipedes from southern Thailand (Diplopoda, Sphaerotheriida, Zephroniidae), pp. 108-132 in European Journal of Taxonomy 762 (1) on pages 121-127, DOI: 10.5852/ejt.2021.762.1457, http://zenodo.org/record/5211725

Zephronia viridisoma Rosenmejer & Wesener sp. nov. urn:lsid:zoobank.org:act: 272E06C8-2DE5-4722-A7D5-D85002D68424 Figs 2B, 6B – 12 Diagnosis Posterior telopod typical for the genus, not differing from those of other Zephronia species. Small (25– 28 mm long) green species (Fig. 2B), surface appearing glabrous, dull, with a single medium sized locking carina at the anal shield and a strongly projecting pleurite 1 (Fig. 9A). One of the few Zephronia species with just a single apical spine on the legs (Fig. 9B), differing in this character from all other described Thai Zephronia species which have 2–5 apical tarsal spines. Male antennomere 6 swollen (Fig. 7A) but not axe-shaped, with Derivatio nominis Named after the overall green colour of living individuals of the species, noun in apposition. Material examined Holotype THAILAND • &male;; Nakhon Si Thammarat Province, Sichon District, Khao Lark Waterfall; 9°03′6″ N, 99°47′24″ E; 25 Aug. 2007; Chulalongkorn University expedition of millipede workshop leg.; dense jungle on limestone; NHMD 621695. Paratypes THAILAND • 2 &male;&male;, 4 &female;&female;; same collection data as for holotype; NHMD 621696 • 1 &male;; same collection data as for holotype; ZFMK MYR8786 • 1 &female;; same collection data as for holotype; CT scan voucher; ZFMK MYR8787. Description (based on holotype) SIZE. Length 25.6 mm. Width of thoracic shield 12.1 mm, of widest segment (8) 13 mm. Height of thoracic shield 7.1 mm, of highest segment (8) 7.7 mm. COLOUR. Head medium brown, faded from green. Antennae medium brown. Legs medium brown, tarsal claws apically dark brown. Tergites light brown, faded green medially, with darker brown posterior margin. Paratergite tips medium brown with darker edges. Anal shield dark green, edges faded to brown (Fig. 2B). HEAD (Fig. 7A–C). Number of ommatidia 50 (Fig. 7B). Organ of Tömösváry placed midway between ocelli and antennal groove (Fig. 7A), with its typical coral-like inner structure (Fig. 7C). ANTENNAE (Fig.7A,D). Reaching leg pair3,6visible antennomeres.Antennomere lengths:6>1>2=3>4=5. Antennomere 6 apically swollen, number of apical cones 32/44. EPIPHARYNX (Fig. 8A). With an extraordinarily large inner tooth. MANDIBLE (Fig. 8E–F). Inner tooth 3-cusped. Pectinate lamellae with 6 or 7 rows of teeth. Condylus at anterior margin with two ridges. GNATHOCHILARIUM (Fig. 8B–D). Lamellae linguales with numerous long setae, medially glabrous. Stipes and mentum with numerous long setae in a regular patter. Sensory cones on gnathochilarium palps in clusters (Fig. 8D). COLLUM. With short setation, evenly distributed across surface. THORACIC SHIELD. Thoracic shield grooves wide and deep, with 9 sclerotized ledges along inner ridge of grooves. TERGITES (Fig. 9D). Tergites glabrous, with dull orange skin like surface. At high magnification tiny setae and knobs become visible. Paratergite tips on posterior half projecting backwards (Fig. 12A–B). ENDOTERGUM (Fig. 6B). With a regular flat margin. Outer zone with three dense rows of irregular marginal setae, some of them extending beyond posterior margin. A single row of rounded cuticular impressions present next to marginal ridge. Intersegmental membrane smooth, without cones and with very few setae. STIGMATIC PLATE (Fig. 10A). First plate with a rounded sub-triangular apex. More posterior plates similar to those of other representatives of the family, half-covered by the pleurites (Fig. 12C). PLEURITES (Figs 9A, 12D). Pleurite 1 projecting strongly, with sharp apex. Pleurite 2 projecting slightly less than 1 and with more rounded apex (Fig. 12D). Projection absent from pleurite 3 (Fig. 12D). LEGS. Ventral spines on leg 1 1/1, on leg 2 3/2, on leg 3 4/4. Apical spine on leg 3 absent. A single apical spine and 5 or 6 ventral spines on midbody legs (Fig. 9B). Femur regularly shaped, 2.2 times longer than wide, sclerotized ledge of medium length, inner margin apically with 5 or 6 small rounded triangular spines on ventral side (Fig. 9B–C). Tarsus 4.2 times longer than wide. ANAL SHIELD. Well-rounded and glabrous. Ventral side with dark coloured. A medium-sized locking carina placed slightly closer to pleurite than margin (Fig. 12D). MALE GONOPORE (Fig. 10B). Inconspicuous, consisting of large membranous opening located directly at mesal margin. ANTERIOR TELOPODS (Fig. 10C–E). With long setae medially on the first 2 podomeres, covering posterior part on podomere 2. Podomere 1 rectangular, as long as wide. Podomere 2 in anterior view as wide as but slightly narrower than podomere 1. Immovable finger barely visible in anterior view (Fig. 10D), short, not protruding up to podomere 4 (Fig. 10C). Podomere 3 long and wide, with rounded projection on posterior side, bearing 3 crenulated teeth. Podomere 4 very narrow and short, with 3 large spines near apex, and a smaller spine near edge to podomere 3. POSTERIOR TELOPODS (Fig. 10F–G). Podomere 2 setose, immovable finger straight, apically tapering, with rows of small circular sclerotized spots. Membranous lobe between podomere 3 and 4 bearing 2 elongated processes, fused at base. Podomere 3 with 8 or 9 small crenulated teeth on posterior side. Membranous lobe on podomere 3 with single spine. Podomere 4 slightly curved towards immovable finger, membranous lobe with 2 spines. Setation on podomere 2 primarily on anterior side. Description of female paratype Length 25.6 mm. Width of thoracic shield 12.3 mm, of widest tergite (8) 13.7 mm. Height of thoracic shield: 8.4 mm, of highest tergite (8) 9.3 mm. Antennae reaching leg pair 3. Apical cones left/right: 25/25. Subanal plate well rounded. Vulvae (Fig. 11): with a large pointed operculum (Op). External lateral plate (EP) and inner mesal plate (IP) fused at bottom. Another female paratype was used for microcomputed tomography. The resulting volume rendering is shown in Fig. 12. While this approach revaled no significant additional details it serves to illustrate morphology of the specimen in an excellent way, much better than any photograph.

Sphaerobelum aesculus Rosenmejer & Wesener sp. nov. urn:lsid:zoobank.org:act: 84D3714E-5CF8-4063-8A4D-40BF8CBEBFB0 Figs 2A, 3–5, 6A Diagnosis Differs from all other species of the genus Sphaerobelum in the shape of the posterior telopod, where there is a swelling at the tip of the immovable finger, but the swelling does not extend above the margin (Fig. 5B arrow). Such a swelling is currently unknown from any other giant pill-millipede species. Derivatio nominis Named after the horse chestnut tree Aesculus hippocastanum L., for the resemblance of the rolled-up female to a horse chestnut. Noun in apposition. Material examined Holotype THAILAND • &male;; Phuket Island, Forest in Kathu district; 14 Feb.1989. M. Andersen and A.R. Rasmussen leg.; in and under old logs; NHMD 621693. Other material THAILAND • 1 &female;; Nakhon Si Thammarat Prov., Khao Luang NP; 8°43′25.2″ N, 99°40′7.7″ E, 355 m a.s.l.; 10–12 Oct. 2003; ATOL Expedition 2003 leg.; NHMD 621694. Description (based on holotype) SIZE. Length 23.8 mm. Width of thoracic shield 10.5 mm, of widest segment (9) 11.1 mm. Height of thoracic shield 6.2 mm, Height of highest segment (7) 7 mm. COLOUR. Head, antenna, legs and tergites golden light brown. Posterior margin of tergites and paratergite tips medium brown. Anal shield medially brown, with lighter colour at edges (Fig. 2A). HEAD. Number of ommatidia 55. Antennae (Fig. 3A–C): Antennomere lengths: 6>1>2=3=4=5. Antennae short, barely reaching first leg pair. Sixth antennomere apically slightly swollen (Fig. 3A), number of apical cones 52/56 (Fig. 3B). Mandible: not dissected. Gnathochilarium: lingual lamella with numerous long setae, medially glabrous. COLLUM. With few setae spread thinly along borders. THORACIC SHIELD. Thoracic shield grooves wide and deep, with 3 sclerotized ledges along inner ridge. TERGITES. Paratergite tips on posterior half projecting backwards. Tergites glabrous with dull leather-like surface (Fig. 3D–E). At high magnification tiny setae become visible. ENDOTERGUM (Fig. 6A). With a regular flat margin. Outer zone with two rows of irregular MARGINAL SETAE. Not extending beyond posterior margin, but reaching &frac23; of the outer area. Anterior part of marginal ridge flat. Intersegmental membrane smooth, without cones and with a row of setae with large glabrous gaps. FIRST STIGMATIC PLATE (Fig. 4A). With a well-rounded apex. PLEURITES. Pleurite 1 projecting posteriorly with sharp apex. Pleurite 2 with rounded apex projecting slightly. LEGS. Ventral spines on leg 1 4/4, on leg 2 3/5, on leg 3 3/4. Apical spine absent on leg 3. Three apical and 4 or 5 ventral spines on midbody legs (Fig. 4C). Inner margin of femur with 8–12 small teeth, but not excavated. Femur 2, tarsus 4 times as long as wide. ANAL SHIELD. Well-rounded and glabrous. Ventral side with a single small locking carina, placed &frac13; from pleurite. MALE GONOPORE. Located at mesal margin of coxa (Fig. 4B), large, covered by a membranous plate. ANTERIOR TELOPODS (Fig. 4D–F). Podomere 1 rectangular, as long as wide, covered with setae anteriorly, with extra long setae medially, and posteriorly glabrous. Podomere 2 with long immovable finger visible laterally in anterior view. Finger curving inwards against podomere 3. Podomere 3 cylindrical twice as long as wide. Suture between podomere 3 and 4 barely visible in posterior and lateral views. Podomere 4 very short, just ¼ of length of podomere 3, with one short and dark spine on apex. POSTERIOR TELOPODS (Fig. 5A–B). Podomere 1 as long as wide, covered in setae anteriorly, and nearly glabrous posteriorly. Podomere 2 with wide immovable finger. Immovable finger straight, only slightly tapering towards apex, bearing an oval membranous spot on posterior side. The membrane between podomere 2 and 3 with single membranous lobe, angled. Podomere 3 with sparse setation. Podomere 3 ca 3 times longer than wide, towards immovable finger with membranous ledge, a single spine, posterior side with row of 9 or 10 crenulated teeth. Podomere 4 short, straight, ca 2.5 times as long as wide, with a short membranous ledge and 2 spines towards immovable finger. Female from Khao Luang NP Length 45 mm. Width of thoracic shield 23 mm, of widest segment (9) 24.6 mm. Height of thoracic shield 12.6 mm, of highest segment (8) 15.2 mm.Antennae reaching leg pair 2. Apical cones on antennae 43/34. Subanal plate well rounded. Vulvae (Fig. 5C): small, with a narrow operculum (Op), only reaching bottom part of prefemur. External lateral plate (EP) and inner mesal plate (IP) almost completely fused. Distribution If the female from Khao Luang (Fig. 2A) is indeed conspecific with the male holotype, the species appears to have a wide area of distribution stretching from Phuket Island at least 160 km to the east (Fig. 2). Unpublished data from a larger inventory project (see, e.g., Wesener et al. 2021) of giant pill-millipedes in the surroundings of Krabi, half way between Phuket Island and Khao Luang, did not recover this species among the numerous specimens, hinting at a patchy distribution and specific microhabitat requirements of S. aesculus sp. nov., Published as part of Rosenmejer, Trine, Enghoff, Henrik, Moritz, Leif & Wesener, Thomas, 2021, Integrative description of new giant pill-millipedes from southern Thailand (Diplopoda, Sphaerotheriida, Zephroniidae), pp. 108-132 in European Journal of Taxonomy 762 (1) on pages 114-120, DOI: 10.5852/ejt.2021.762.1457, http://zenodo.org/record/5211725

Genus Sphaerobelum Verhoeff, 1924 Type species Sphaerobelum clavigerum Verhoeff, 1924, from Vietnam. Other taxa included 18 species including the one described below (Semenyuk et al. 2018, 2020; Wesener 2019; Zhao et al. 2020). Distribution Vietnam, Thailand, Laos, China., Published as part of Rosenmejer, Trine, Enghoff, Henrik, Moritz, Leif & Wesener, Thomas, 2021, Integrative description of new giant pill-millipedes from southern Thailand (Diplopoda, Sphaerotheriida, Zephroniidae), pp. 108-132 in European Journal of Taxonomy 762 (1) on page 114, DOI: 10.5852/ejt.2021.762.1457, http://zenodo.org/record/5211725, {"references":["Semenyuk I., Golovatch S. I. & Wesener T. 2018. Four new species of giant pill-millipedes from Vietnam (Sphaerotheriida: Zephroniidae). Zootaxa 4459 (3): 535 - 550. https: // doi. org / 10.11646 / zootaxa. 4459.3.7","Semenyuk I., Golovatch S. I. & Wesener T. 2020. Some new or poorly-known Zephroniidae (Diplopoda, Sphaerotheriida) from Vietnam. ZooKeys 930: 37 - 60. https: // doi. org / 10.3897 / zookeys. 930.47742","Wesener T. 2019. First records of giant pill-millipedes from Laos (Diplopoda, Sphaerotheriida, Zephroniidae). Zootaxa 4563 (2): 201 - 248. https: // doi. org / 10.11646 / zootaxa. 4563.2.1","Zhao Y., Yu J., & Liu W. 2020. A molecular-based phylogeny of the millipede genus Sphaerobelum Verhoeff, 1924, with the first record of the genus from mainland China (Diplopoda: Sphaerotheriida: Zephroniidae). Annales de la Societe entomologique de France (NS) 56 (4): 341 - 348."]}

Genus Zephronia Gray, 1832 Type species Zephronia ovalis Gray, 1832. Other taxa included 44 species, including the one described below (Wesener 2016a, 2019; Semenyuk et al. 2018, 2020). Distribution NE India, Nepal, Myanmar, with a few species also in SE Asia. Remark Likhitrakarn et al. (2021) provided an updated diagnosis of this genus including the presence of several apical tarsal spines. The new species described here necessitates a slight modification, as it has only one apical tarsal spine., Published as part of Rosenmejer, Trine, Enghoff, Henrik, Moritz, Leif & Wesener, Thomas, 2021, Integrative description of new giant pill-millipedes from southern Thailand (Diplopoda, Sphaerotheriida, Zephroniidae), pp. 108-132 in European Journal of Taxonomy 762 (1) on page 120, DOI: 10.5852/ejt.2021.762.1457, http://zenodo.org/record/5211725, {"references":["Wesener T. 2016 a. The giant pill-millipedes, order Sphaerotheriida - An annotated species catalogue with morphological atlas and list of apomorphies (Arthropoda: Diplopoda). Bonn Zoological Bulletin - Supplementum 63: 1 - 104.","Wesener T. 2019. First records of giant pill-millipedes from Laos (Diplopoda, Sphaerotheriida, Zephroniidae). Zootaxa 4563 (2): 201 - 248. https: // doi. org / 10.11646 / zootaxa. 4563.2.1","Semenyuk I., Golovatch S. I. & Wesener T. 2018. Four new species of giant pill-millipedes from Vietnam (Sphaerotheriida: Zephroniidae). Zootaxa 4459 (3): 535 - 550. https: // doi. org / 10.11646 / zootaxa. 4459.3.7","Semenyuk I., Golovatch S. I. & Wesener T. 2020. Some new or poorly-known Zephroniidae (Diplopoda, Sphaerotheriida) from Vietnam. ZooKeys 930: 37 - 60. https: // doi. org / 10.3897 / zookeys. 930.47742","Likhitrakarn N., Golovatch S. I., Srisonchai R. & Sutcharit C. 2021. Two new species of the giant pillmillipede genus Zephronia Gray, 1832 from Thailand (Diplopoda: Sphaerotheriida: Zephroniidae). Tropical Natural History 21: 12 - 26."]}

Sphaerobelum aesculus Rosenmejer & Wesener sp. nov. urn:lsid:zoobank.org:act: 84D3714E-5CF8-4063-8A4D-40BF8CBEBFB0 Figs 2A, 3–5, 6A Diagnosis Differs from all other species of the genus Sphaerobelum in the shape of the posterior telopod, where there is a swelling at the tip of the immovable finger, but the swelling does not extend above the margin (Fig. 5B arrow). Such a swelling is currently unknown from any other giant pill-millipede species. Derivatio nominis Named after the horse chestnut tree Aesculus hippocastanum L., for the resemblance of the rolled-up female to a horse chestnut. Noun in apposition. Material examined Holotype THAILAND • &male;; Phuket Island, Forest in Kathu district; 14 Feb.1989. M. Andersen and A.R. Rasmussen leg.; in and under old logs; NHMD 621693. Other material THAILAND • 1 &female;; Nakhon Si Thammarat Prov., Khao Luang NP; 8°43′25.2″ N, 99°40′7.7″ E, 355 m a.s.l.; 10–12 Oct. 2003; ATOL Expedition 2003 leg.; NHMD 621694. Description (based on holotype) SIZE. Length 23.8 mm. Width of thoracic shield 10.5 mm, of widest segment (9) 11.1 mm. Height of thoracic shield 6.2 mm, Height of highest segment (7) 7 mm. COLOUR. Head, antenna, legs and tergites golden light brown. Posterior margin of tergites and paratergite tips medium brown. Anal shield medially brown, with lighter colour at edges (Fig. 2A). HEAD. Number of ommatidia 55. Antennae (Fig. 3A–C): Antennomere lengths: 6>1>2=3=4=5. Antennae short, barely reaching first leg pair. Sixth antennomere apically slightly swollen (Fig. 3A), number of apical cones 52/56 (Fig. 3B). Mandible: not dissected. Gnathochilarium: lingual lamella with numerous long setae, medially glabrous. COLLUM. With few setae spread thinly along borders. THORACIC SHIELD. Thoracic shield grooves wide and deep, with 3 sclerotized ledges along inner ridge. TERGITES. Paratergite tips on posterior half projecting backwards. Tergites glabrous with dull leather-like surface (Fig. 3D–E). At high magnification tiny setae become visible. ENDOTERGUM (Fig. 6A). With a regular flat margin. Outer zone with two rows of irregular MARGINAL SETAE. Not extending beyond posterior margin, but reaching &frac23; of the outer area. Anterior part of marginal ridge flat. Intersegmental membrane smooth, without cones and with a row of setae with large glabrous gaps. FIRST STIGMATIC PLATE (Fig. 4A). With a well-rounded apex. PLEURITES. Pleurite 1 projecting posteriorly with sharp apex. Pleurite 2 with rounded apex projecting slightly. LEGS. Ventral spines on leg 1 4/4, on leg 2 3/5, on leg 3 3/4. Apical spine absent on leg 3. Three apical and 4 or 5 ventral spines on midbody legs (Fig. 4C). Inner margin of femur with 8–12 small teeth, but not excavated. Femur 2, tarsus 4 times as long as wide. ANAL SHIELD. Well-rounded and glabrous. Ventral side with a single small locking carina, placed &frac13; from pleurite. MALE GONOPORE. Located at mesal margin of coxa (Fig. 4B), large, covered by a membranous plate. ANTERIOR TELOPODS (Fig. 4D–F). Podomere 1 rectangular, as long as wide, covered with setae anteriorly, with extra long setae medially, and posteriorly glabrous. Podomere 2 with long immovable finger visible laterally in anterior view. Finger curving inwards against podomere 3. Podomere 3 cylindrical twice as long as wide. Suture between podomere 3 and 4 barely visible in posterior and lateral views. Podomere 4 very short, just ¼ of length of podomere 3, with one short and dark spine on apex. POSTERIOR TELOPODS (Fig. 5A–B). Podomere 1 as long as wide, covered in setae anteriorly, and nearly glabrous posteriorly. Podomere 2 with wide immovable finger. Immovable finger straight, only slightly tapering towards apex, bearing an oval membranous spot on posterior side. The membrane between podomere 2 and 3 with single membranous lobe, angled. Podomere 3 with sparse setation. Podomere 3 ca 3 times longer than wide, towards immovable finger with membranous ledge, a single spine, posterior side with row of 9 or 10 crenulated teeth. Podomere 4 short, straight, ca 2.5 times as long as wide, with a short membranous ledge and 2 spines towards immovable finger. Female from Khao Luang NP Length 45 mm. Width of thoracic shield 23 mm, of widest segment (9) 24.6 mm. Height of thoracic shield 12.6 mm, of highest segment (8) 15.2 mm.Antennae reaching leg pair 2. Apical cones on antennae 43/34. Subanal plate well rounded. Vulvae (Fig. 5C): small, with a narrow operculum (Op), only reaching bottom part of prefemur. External lateral plate (EP) and inner mesal plate (IP) almost completely fused. Distribution If the female from Khao Luang (Fig. 2A) is indeed conspecific with the male holotype, the species appears to have a wide area of distribution stretching from Phuket Island at least 160 km to the east (Fig. 2). Unpublished data from a larger inventory project (see, e.g., Wesener et al. 2021) of giant pill-millipedes in the surroundings of Krabi, half way between Phuket Island and Khao Luang, did not recover this species among the numerous specimens, hinting at a patchy distribution and specific microhabitat requirements of S. aesculus sp. nov.

Zoosphaerium libidinosum species-group Remarks Species can be identified as members of the Z. libidinosum species-group based on morphological characters of the posterior telopods, in which the third telopoditomere carries more than 15 large crenulated teeth as well as two large membranous lobes (Wesener & Wägele 2008). The species-group now contains 16 described species distributed in all types of forests all over Madagascar, including the arid spiny bush (Wesener 2009). Key to species of the Zoosphaerium libidinosum species-group 1. Anal shield without locking carinae (in small, immature individuals sometimes traces of carinae are visible).........................................................................................................................................2 – Anal shield with two obvious locking carinae..............................................................................7 2. Operculum of vulva at least weakly notched apically. Fifth antennomere with field of sensilla basiconica apically............................................................................................................................3 – Operculum of vulva well-rounded apically. No sensilla basiconica present on fifth antennomere...............................................................................................................................................4 3. Tergites dark green, in alcohol almost blackish, posterior margin light brown to yellow. Tarsus of walking legs broad, only 3.0 times as long as wide. Third podomere of anterior telopods with four large, sclerotized teeth. Endotergal marginal bristles strongly extending beyond tergite......................................................................................................................... Z. haackeri Wesener, 2009 – Tergites almost white, with light, shiny-green field dorso-laterally on each side. Tarsus of walking legs of usual size, 3.8 times as long as wide. Third podomere of anterior telopods with single large, sclerotized tooth. Endotergal marginal bristles slightly extending beyond tergite................................................................................................................................................ Z. album Wesener, 2009 4. Anterior telopod second podomere process slightly extending beyond third podomere. Third podomere laterally with plateau and mesally with a sharp, prominent process. Tergites light green with distinct, light brown posterior margin. Male harp with two weakly-developed stridulation ribs............................................................................................................ Z. discolor Wesener, 2009 – Anterior telopod second podomere process shorter than third podomere. Third podomere of anterior telopods only laterally with plateau....................................................................................5 5. Tergites dark, almost blackish green, posteriorly with distinct and wide light brown margin. Anal shield covered with numerous short hairs. Endotergum with a single row of marginal bristles, slightly extending beyond tergite.............................................................. Z. mitoho Wesener, 2009 – Tergites with dark green, light green pattern, posteriorly with distinct, thin light brown margin. Anal shield glabrous. Endotergum with two or more rows of marginal bristles, clearly extending beyond apical margin.......................................................................................................................6 6. Endotergum with ≥ 3 rows of marginal bristles, strongly extending beyond tergite. Third podomere of anterior telopods laterally with a triangular projection. Tarsi of walking legs>3.7 times as long as wide. Male harp with two strongly developed stridulation ribs............................................................................................................. Z. libidinosum (de Saussure & Zehntner, 1897) – Endotergum with two rows of marginal bristles, strongly extending beyond tergite. Third podomere of anterior telopods laterally without triangular projection. Tarsi of walking legs 3.5 times as long as wide. Male harp with two weakly developed stridulation ribs........................... Z. silens sp. nov. 7. Second and third podomeres of posterior telopods covered with numerous minute hairs. Antennae with four apical cones; tergites glossy, glabrous...............................................................................8 – Second and third podomeres of posterior telopods glabrous or covered with long setae...........10 8. Tergites black. Anterior telopod with one stridulation rib.................................... Z. nigrum sp. nov. – Tergites jade-like green. Anterior telopod at least apically with 2 stridulation ribs........................9 9. Tergites glossy, completely jade-like green. Male harp with two basally fused stridulation ribs. Third podomere of anterior telopods mesally with sharp, prominent process. Endotergum with one row of rounded cuticular impressions. Fixed finger of posterior telopod basally without membranous lobe..................................................... Z. anomalum (de Saussure & Zehntner, 1902) – Tergites glossy, jade-like green, with shiny dark green pattern. Male harp with two fully separate stridulation ribs. Third podomere of anterior telopods without process. Endotergum without cuticular impressions. Fixed finger of posterior telopod with basal membranous lobe..................................................................................................................................... Z. pulchellum Wesener, 2009 10. Posterior end of anal shield in both sexes excavated..................................................................11 – Posterior end of anal shield well-rounded....................................................................................14 11. Tergites glossy, green. Male harp with two weakly-developed stridulation ribs. Third podomere of anterior telopods with a single large sclerotized tooth............ Z. endemicum Wesener, 2009 – Tergites matte, brownish. Male harp with two well-developed stridulation ribs........................12 12. Locking carinae on anal shield of similar length. Third podomere of posterior telopods with ca 28 large sclerotized teeth. Antenna with more than 40 apical cones............................................................................................................................................................ Z. xerophilum Wesener, 2009 – Posterior locking carina at least two times as long as anterior carina. Third podomere of posterior telopods with less than 25 large sclerotized teeth. Antenna with less than 30 apical cones......................................................................................................................................................13 13. Posterior locking carina of anal shield more than three times as long as anterior carina....................................................................................................... Z. arborealis Wesener & Sierwald, 2005 – Posterior locking carina ca twice as long as anterior one..... Z. lambertoni (Brolemann, 1922) 14. Anal shield glabrous, golden. Tergites with unique pitch-black/reddish-brown pattern. Third podomere of anterior telopods without large sclerotized teeth. First antennomere without sensilla basiconica. Sclerotized teeth present on antennomeres 1–4. Endotergal marginal bristles not extending beyond tergite margin................................................................................. Z. aureum Wesener, 2009 – Anal shield covered with few, minute hairs of dirty brownish color. Tergites with light brown, blackish coloration pattern. Third podomere of anterior telopods laterally with at least a single large sclerotized tooth. First antennomere with sensilla basiconica. Sclerotized teeth present on antennomeres 1–5. Endotergal marginal bristles slightly extending beyond tergite margin.........15 15. Legs short, femur 1.45 times as long as wide, tarsus 3.2 times as long as wide. Third pair of legs without apical spine. Third podomere of posterior telopods with ca 18 large sclerotized teeth. Endotergum with two rows of marginal bristles. Posterior locking carina 4 times as long as anterior carina. Antenna with 25 or fewer apical cones......... Z. priapus (de Saussure & Zehntner, 1897) – Legs of usual size, femur 1.9 times as long as wide, tarsus 4.2 times as long as wide. Third pair of legs with apical spine. Third podomere of posterior telopods with ca 22 large sclerotized teeth. Endotergum with three rows of marginal bristles. Locking carinae of equal size. Antenna with 35–86 apical cones........................................... Z. pseudopriapus Wesener, 2009, Published as part of Wesener, Thomas & Sagorny, Christina, 2021, Seven new giant pill-millipede species and numerous new records of the genus Zoosphaerium from Madagascar (Diplopoda, Sphaerotheriida, Arthrosphaeridae), pp. 1-48 in European Journal of Taxonomy 758 (1) on pages 7-9, DOI: 10.5852/ejt.2021.758.1423, http://zenodo.org/record/5075824, {"references":["Wesener T. & Wagele J. W. 2008. The giant pill-millipedes of Madagascar: revision of the genus Zoosphaerium (Myriapoda, Diplopoda, Sphaerotheriida). Zoosystema 30 (1): 5 - 82.","Saussure H. de & Zehntner L. 1897. Atlas de l'histoire naturelle des Myriapodes. Histoire physique, naturelle et politique de Madagascar 27 (53): 1 - 12.","Saussure H. de & Zehntner L. 1902. Myriapodes de Madagascar. In: Grandidier A. (ed.) Histoire physique, naturelle et politique de Madagascar 27 (53): i - viii, 1 - 356.","Brolemann H. W. 1922. Liste des myriapodes de l'Academie Malagache de Tananarive. Bulletin de la Societe zoologique de France 47: 223 - 278. https: // doi. org / 10.5962 / bhl. part. 3470"]}

Zoosphaerium voahangy sp. nov. urn:lsid:zoobank.org:act: 11B9C806-A111-48FC-AC36-035385125232 Figs 2E, 3, 5E, 9 Diagnosis The position of the locking carinae close to the last pleurite identifies Z. voahangy sp. nov. as a member of the Z. platylabum species-group. The uniquely slender and elongated posterior telopods in combination with the presence of a single stridulation rib on the male harp and more than 35 apical cones on the antennae, are characters only shared with Z. broelemanni Wesener, 2009. Zoosphaerium voahangy sp. nov. differs from Z. broelemanni in the differently shaped anterior telopods, spination and teeth number of the movable finger of the posterior telopod (see key), as well as in characters of the endotergum, which has two dense rows of long marginal bristles in the former, but a single row of short bristles in the latter. Etymology ‘Voahangy’, noun in apposition, to honor the great collector and Malagasy scientist Voahangy Soarimalala. Material examined Holotype MADAGASCAR – Mahajanga • &male;; Analalava, 5.5 km NNE of Ambinda Village; 14°18.6′ S, 47°54.9′ E; alt. 120 m; dry deciduous forest; 8 Dec. 2004; V. Soarimalala leg.; coll. no. VS-508; FMNH- INS 3119921. Paratype MADAGASCAR – Mahajanga • &male;; same collection data as for holotype; FMNH-INS 3119919. Description BODY LENGTH. Male (holotype): length 26.3 mm, width of thoracic shield 16.1 mm, of tergite 7 16.7 mm (widest), height of thoracic shield 8.9 mm, of tergite 7 9.3 mm (highest). COLORATION. Head, collum, thoracic shield and tergites light brown with dark posterior margins. Anal shield entirely light brown (Fig. 2E). Antennae dark green, legs basally light brown, apically green. HEAD. Eyes with more than 65 ommatidia. Antennae short, protruding back to leg 5. Length of antennomeres 1=2 =3=4 =5Male sexual characters GONOPORE. Covered with a single undivided, apically rounded, membranous, basally sclerotized plate covering ¼ of surface of coxa. Gonopore located basally on joint (Fig. 9B). ANTERIOR TELOPODS. Harp with single stridulation rib located mesally, long and well developed (Fig. 9D). First podomere 1.1 times as wide as long. Second podomere process lobe-like and slightly curved with rounded edges, protruding up to half of third podomere height, apically with sclerotized spots (Fig. 9F). Third podomere long, almost as long as first; cavity mesally with numerous sclerotized spots and three thin, sclerotized spines located apically (Fig. 9E–F); one sclerotized spot located close to tip, laterally with a row of ca 11 crenulated teeth (Fig. 9E). Podomeres 1–3 covered on both sides with numerous long, isolated hairs. POSTERIOR TELOPODS. Movable finger slender and long, with a well-rounded tip, not tapering towards tip; hollowed-out inner margin with 3 or 4 sclerotized spines located apically, posterior aspect with ca 25 small sclerotized teeth (Fig. 9G). Movable finger 4 times as long as wide, slightly longer than fixed finger. Fixed finger straight, only apically curved towards movable finger, not tapering, more slender than third podomere. Fixed finger at margin with numerous sclerotized spots (Fig. 9H). First podomere in posterior view almost glabrous, podomeres 2 and 3 covered on both sides with numerous long, isolated hairs. Female Unknown. Distribution and ecology The only known habitat of Z. voahangy sp. nov., the forest of Analalava (Fig. 3), is a small (2 km south to north, 750 m east to west) fragment of disturbed and partly degraded lowland rainforest, isolated by 30 km from Betampona, the only other remaining lowland rainforest fragment in a 100 km radius., Published as part of Wesener, Thomas & Sagorny, Christina, 2021, Seven new giant pill-millipede species and numerous new records of the genus Zoosphaerium from Madagascar (Diplopoda, Sphaerotheriida, Arthrosphaeridae), pp. 1-48 in European Journal of Taxonomy 758 (1) on pages 28-30, DOI: 10.5852/ejt.2021.758.1423, http://zenodo.org/record/5075824

Unidentifiable Zoosphaerium spp. (mainly immatures) Material examined MADAGASCAR – Antsiranana • 1 spec.; Marojejy Res.; 4 km NNW of Manantenina; 14°26′ S, 49°45′ E; alt. 700 m; 10–16 Nov. 1993; J. Coddington, N. Scharff, S. Larcher, C. Griswold and R. Andriamasimanana leg.; NHMD • 1 ♀; Montagne dʼAmbre; 12°32′ S, 49°10′ E; alt. 1100 m; 23–28 Nov. 1993; J. Coddington, N. Scharff, S. Larcher, C. Griswold and R. Andriamasimanana leg.; NHMD • 1 spec.; same collection data as for preceding; NHMD • 2 specs; N Ankarana Reserve, base of tsingy summit; 12.8461° S, 49.15° E; alt. 284 m; dry deciduous forest on limestone; 8 Apr. 2007; Ken Emberton leg.; coll. no. MBI-2038; FMNH-INS 000 103 908 • 1 spec.; N Ankarana Reserve, local tsingy, summit; 12.8465° S, 49.1503° E; alt. 294 m; dry deciduous forest on limestone; 8 Apr. 2007; Ken Emberton leg.; coll. no. MBI-2037; FMNH-INS 000 103 909 • 1 spec.; NE Ankarana Reserve, roadside; 12.8329° S, 49.2286° E; alt. 331 m; dry deciduous forest on lava; 14 Apr. 2007; Ken Emberton leg.; coll. no. MBI- 2066; FMNH-INS 000 103 904 • 2 specs; E Ankarana Reserve, talus and rocks below 20 m tsingy cliff; 12.8609° S, 49.2267° E; alt. 294 m; dry deciduous forest on limestone; 14 Apr. 2007; Ken Emberton leg.; coll. no. MBI-2068; FMNH-INS 000 103 911 • 1 spec.; NE Ankarana Reserve, local limestone summit; 12.8738° S, 49.2249° E; alt. 315 m; dry deciduous forest on limestone; 14 Apr. 2007; Ken Emberton leg.; coll. no. MBI-2064; FMNH-INS 000 103 920 • 4 specs; NE Ankarana Reserve, talus below tsingy visible from road; 12.8617° S, 49.2266° E; alt. 266 m; dry deciduous forest on limestone; 14 Apr. 2007; Ken Emberton leg.; coll. no. MBI-2062; FMNH-INS 000 103 922 • 2 spec.; NE Analamerana, near summit; 12.6833° S, 49.5333° E; alt. 36 m; litter-loam pockets in limestone brocks, dry deciduous forest on limestone; 23 Apr. 2007; Ken Emberton leg.; coll. no. MBI-2072; FMNH-INS 000 103 929 • 3 specs; NE Analamerana, 160 m downslope E from summit; 12.6833° S, 49.5333° E; limestone outcrop, dry deciduous forest on limestone; 25 Apr. 2007; Ken Emberton leg.; coll. no. MBI-2086; FMNH-INS 000 103 921 • 1 spec.; NE Analamerana, on W bank of Antafiamantsina River, at waterfall, 3 rd rapid up; 12.6833° S, 49.5333° E; dry deciduous forest on limestone; 26 Apr. 2007; Ken Emberton leg.; coll. no. MBI-2099; FMNH-INS 000 103 903 • 7 specs; NE Analamerana, small tsingy 180 m above river, ca 30 minute walk; 12.6833° S, 49.5333° E; dry deciduous forest on limestone; 26 Apr. 2007; Ken Emberton leg.; coll. no. MBI-2094; FMNH-INS 000 103 917 • 2 specs; N Cap d'Ambre, Pointe du Phoque; 11.9698° S, 49.2845° E; alt. 11 m; spring-bottle tree stand, dry deciduous forest on limestone; 4 May 2007; Ken Emberton leg.; coll. no. MBI-2108; FMNH-INS 000 103 910 • 2 specs; NW of Antalaha, S of Mahadera, Mt. Ambohodrapaka; 14.6805° S, 50.0325° E; alt. 270 m; partially cut rainforest on granite; 29 Jul. 2007; Ken Emberton leg.; coll. no. MBI-2403; FMNH-INS 000 103 906 • 2 specs; same collection data as for preceding; coll. no. MBI-2404; FMNH-INS 000 103 907 • 1 spec.; NW of Antalaha, S of Mahadera, Mt Ambohodrapaka; 14.6836° S, 50.031° E; alt. 369 m; partially cut rainforest on granite; 29 Jul. 2007; Ken Emberton leg.; coll. no. MBI-2417; FMNH-INS 000 103 924 • 1 spec.; NW of Antalaha; 14. 4123° S, 50.1755° E; alt. 1790 m; 30 Jul. 2007; Ken Emberton leg.; coll. no. MBI-2439; FMNH-INS 000 103 905 • 2 specs; NW of Antalaha, S of Mahadera, Mt Ambohodrapaka; 14.6873° S, 50.0301° E; alt. 499 m; rainforest, on quartzite; 30 Jul. 2007; Ken Emberton leg.; coll. no. MBI-2430; FMNH-INS 000 103 915 • 1 spec.; NW of Antalaha, S of Mahadera, Mt. Ambohodrapaka; 14.6872° S, 50.0293° E; alt. 545 m; 30 Jul. 2007; Ken Emberton leg.; coll. no. MBI-2434; FMNH- INS 000 103 918 • 1 spec.; NW of Antalaha, SW of Andranofotsy, Mt Ambodivohitra; 14.8157° S, 49.9634° E; alt. 487 m; rainforest on quartzite; 10 Aug. 2007; Ken Emberton leg.; coll. no. MBI- 2459; FMNH-INS 000 103 913 • 2 specs; NW of Antalaha, SW of Andranofotsy, Mt Ambodivohitra; 14.8157° S, 49.9634° E; alt. 487 m; rainforest on quartzite; 10 Aug. 2007; Ken Emberton leg.; coll. no. MBI-2459; FMNH-INS 000 103 919 • 2 specs; NW of Antalaha, SW of Andranofotsy, Mt Ambodivohitra; 14.8166° S, 49.9622° E; alt. 575 m; rainforest, pandanus, palms, sphagnum on quartzite; 11 Aug. 2007; Ken Emberton leg.; coll. no. MBI-2472; FMNH-INS 000 103 912 • 1 spec.; NW of Antalaha, S of Antsahanoro, riverside hill rainforest; 14.8434° S, 50.1372° E; alt. 82 m; on quartzite; 15 Aug. 2007; Ken Emberton leg.; coll. no. MBI-2498; FMNH-INS 000 103 916 • 1 spec.; NW of Antalaha, S of Antsahanoro; 14.8445° S, 50.1359° E; alt. 131 m; riverside hill rainforest on quartzite; 15 Aug. 2007; Ken Emberton leg.; coll. no. MBI-2512; FMNH-INS 000 103 934 • 2 specs; Mt Bobanora, about 40 km W of Vohemar; 13.2141° S, 49.7574° E; alt. 320 m; viny rainforest on basalt talus; 26 Aug. 2007; Ken Emberton leg.; coll. no. MBI-2550; FMNH-INS 000 103 914 • 1 spec.; Manongorivo Reserve, slope high above Berato; 14.0366° S, 48.2882° E; alt. 897 m; rainforest on metamorphic boulders; 17 Oct. 2007; Ken Emberton leg.; coll. no. MBI-2611; FMNH-INS 000 103 933 • 2 specs; Manongorivo Reserve, Monongorivo River junction with small tributary; 13.9846° S, 48.2891° E; alt. 321 m; rainforest on granite; 20 Oct. 2007; Ken Emberton leg.; coll. no. MBI-2619; FMNH-INS 000 103 941 • 5 specs; Manongorivo Reserve, riverside flat area below cliffs, Monongorivo River; 13.9818° S, 48.2858° E; alt. 281 m; rainforest on granite; 20 Oct. 2007; Ken Emberton leg.; coll. no. MBI-2620; FMNH-INS 000 103 943. – Mahajanga • 1 spec.; Namoroka Reserve, southern valley on NE side of tsingy; 16.469° S, 45.3647° E; alt. 142 m; dry deciduous forest on limestone; 25 Sep. 2007; Ken Emberton leg.; coll. no. MBI-2567; FMNH-INS 000 103 936 • 2 specs; Namoroka Reserve, southern isolated tsingy canyon; 16.4674° S, 45.3806° E; alt. 432 feet; dry deciduous forest on limestone; 27 Sep. 2007; Ken Emberton leg.; coll. no. MBI-2574; FMNH-INS 000 103 928 • 2 specs; Namaroka Reserve, southern isolated tsingy; 16.4871° S, 45.3371° E; alt. 435 m; dry deciduous forest on limestone; 27 Sep. 2007; Ken Emberton leg.; coll. no. MBI-2572; FMNH-INS 000 103 927 • 9 spec.; S Namaroka National Park, low tsingy baobobs; 16.4565° S, 45.4011° E; alt. 115 m; dry deciduous forest on limestone; 29 Sep. 2007; Ken Emberton leg.; coll. no. MBI-2578; FMNH-INS 000 103 923 • 1 spec.; Anjajavy, isolated steep limestone hill near Antsangabe; 15.0643° S, 47.2373° E; alt. 30 m; dry deciduous forest on limestone; 8 Oct. 2007; Ken Emberton leg.; coll. no. MBI-2592; FMNH-INS 000 104 000 • 1 spec.; Anjajavi, side of long tsingy hill near dry mudflat; 15.026° S, 47.2712° E; alt. 22 m; dry deciduous forest on limestone; 11 Oct. 2007; Ken Emberton leg.; coll. no. MBI-2595; FMNH-INS 000 103 940. – Toamasina • 1 spec.; Andriantanely Massif; 18.6907° S, 48.8063° E; alt. 589 m; rainforest; 19 May 2008; Ken Emberton leg.; coll. no. MBI-2665; FMNH-INS 000 103 932 • 1 spec.; Andriantantely Massif, streamside cliff base; 18.6997° S, 48.806° E; alt. 301 m; in rainforest, on granite; 22 May 2008; Ken Emberton leg.; coll. no. MBI-2693; FMNH-INS 000 103 930 • 1 spec.; Makira Reserve, W of Andaparaty; 15.2302° S, 49.5975° E; alt. 430 m; rainforest; 23 Nov. 2008; Ken Emberton leg.; coll. no. MBI-3152; FMNH-INS 000 103 939 • 2 specs; N of Manona littoral forest; 16.6028° S, 49.8037° E; alt. 23 m; 2 Dec. 2008; Ken Emberton leg.; coll. no. MBI-3158; FMNH-INS 000 103 937. – Toliara • 2 specs; W of Ehasity; 24.154° S, 47.1821° E; alt. 347 m; rainforest; 18 Aug. 2008; Ken Emberton leg.; coll. no. MBI-2966; FMNH-INS 000 103 938 • 1 spec.; Mount Ivohibe; 24.5607° S, 47.2065° E; alt. 457 m; rainforest; 31 Aug. 2008; Ken Emberton leg.; coll. no. MBI-2900; FMNH-INS 000 103 931 • 1 spec.; W of Ehasity rainforest; 24.1514° S, 47.178° E; alt. 479 m; 18 Sep. 2008; Ken Emberton leg.; coll. no. MBI-2964; FMNH-INS 000 103 942., Published as part of Wesener, Thomas & Sagorny, Christina, 2021, Seven new giant pill-millipede species and numerous new records of the genus Zoosphaerium from Madagascar (Diplopoda, Sphaerotheriida, Arthrosphaeridae), pp. 1-48 in European Journal of Taxonomy 758 (1) on page 42, DOI: 10.5852/ejt.2021.758.1423, http://zenodo.org/record/5075824

Zoosphaerium masoala sp. nov. urn:lsid:zoobank.org:act: F62E0E9E-172D-4076-BD01-509AD42BAE89 Figs 2F, 3, 10A, 11 Diagnosis The shape and position of the locking carinae, as well as the slender posterior telopods place Z. masoala sp. nov. in the Z. platylabum species-group. Zoosphaerium masoala sp. nov. differs from all other species of the group in the combination of the presence of two stridulation ribs on the male harp, slender posterior telopods in which both fingers reach almost the same length and>10 apical cones on the antenna. Etymology ‘Masoala’, noun in apposition, after the forest and peninsula of Masoala, the largest remaining area of lowland rainforest on Madagascar, for which the species described here is the first recorded giant pillmillipede species (Fig. 3). Material examined Holotype MADAGASCAR – Toamasina • &male;; Région Sava, Parc National de Masoala, Forêt de Tampolo, 1.2 km S of Ambodiforaha village; 15°43′45.6″ S, 49°57′49.4″ E; alt. 15 m; pitfall buckets, littoral forest; 11 Dec. 2015; V. Soarimalala leg.; coll. no. VS-2582; FMNH-INS 3195651. Description Male BODY LENGTH. Length 23.9 mm, width of thoracic shield 14.2 mm (widest), height of thoracic shield 7.2 mm (highest). COLORATION. Head, collum, thoracic shield and tergites dark brown to black. Grooves of thoracic shield greenish, anal shield lighter brown (Fig. 2F). Antennae and legs green. HEAD. Eyes with more than 50 ommatidia. Antennae short, protruding back to leg 4. Length of antennomeres 1>2>3=4 =5Male sexual characters GONOPORE. Covered with a single undivided, rounded, apically membranous, basally sclerotized plate, covering ¼ of surface of coxa. Gonopore located basally on joint (Fig. 11B). ANTERIOR TELOPODS. Harp with two stridulation ribs, mesal rib smaller than lateral one, both ribs well developed, located in mesal corner (Fig. 11D). First podomere 1.1 times as wide as long. Second podomere process lobe-like and slightly curved, with rounded edges, protruding up to half of third podomere height, apically with sclerotized spots. Third podomere long, as long as first; cavity mesally with numerous sclerotized spots and three thin, sclerotized spines; one sclerotized spot located close to tip; laterally with up to 12 crenulated teeth. Podomeres 1–3 covered on both sides with numerous long, isolated hairs. POSTERIOR TELOPODS. Movable finger with a well-rounded tip, tapering towards tip; hollowed-out inner margin without lobes but with 5 sclerotized spines, 2 located apically, 2 at midpoint and 1 basally (Fig. 11H). Posterior aspect of movable finger with ca 14 small sclerotized teeth (Fig. 11G). Movable finger slightly longer than fixed finger, the latter slender (4 times as long as wide), apically curved towards former. Fixed finger basally with a single spine, at margin with numerous sclerotized spots. First podomeres almost glabrous, movable and immovable finger covered on both sides with numerous long, isolated hairs, only tips of chela glabrous. Female Unknown. Distribution and ecology This species is currently known only from the vast lowland rainforests of the Masoala Peninsula on the east coast (Fig. 3). Remarks Despite several attempts, no DNA could be extracted from the male holotype. Therefore, its degree of similarity to the two much larger females known and sequenced from the Masoala area (KY399024 and KY399025) could not be assessed (see Fig. 1)., Published as part of Wesener, Thomas & Sagorny, Christina, 2021, Seven new giant pill-millipede species and numerous new records of the genus Zoosphaerium from Madagascar (Diplopoda, Sphaerotheriida, Arthrosphaeridae), pp. 1-48 in European Journal of Taxonomy 758 (1) on pages 30-33, DOI: 10.5852/ejt.2021.758.1423, http://zenodo.org/record/5075824

Zoosphaerium silens sp. nov. urn:lsid:zoobank.org:act: 2FA555CB-B805-401A-8730-6DFB6525C585 Figs 2B, 3, 5B, 6 Diagnosis The posterior telopods (Fig. 6H���I) identify Z. silens sp. nov. as a member of the Z. libidinosum speciesgroup. Zoosphaerium silens sp. nov. shares the absence of locking carinae with a number of spiny forest species. It differs from all other species except Z. libidinosum in the color of the tergites, the glabrous anal shield and the presence of at least two rows of very long endotergal bristles. Zoosphaerium silens sp. nov. differs from Z. libidinosum in the wider tarsus (3.5 versus 3.8 times as long as wide), the presence of only two rows of endotergal bristles (three rows in Z. libidinosum) and the absence of a lateral triangular projection on the third podomere of the anterior telopods (present in Z. libidinosum). Z. silens sp. nov. differs by more than 14% uncorrected p-distance in the COI gene from Z. album and Z. libidinosum. Etymology ���Silens��� (= ���quiet���), noun in apposition, after the reduced stridulation ribs in the male. Material examined Holotype MADAGASCAR ��� Toliara ��� &male;; Parc National de Kirindy Mite, 25.6 km SW of Belo sur Mer; 20��56.6��� S, 43��52.3��� E; alt. 10 m; dry deciduous forest; 9 Mar. 2007; V. Soarimalala leg.; coll. no. VS- 1348; FMNH-INS 3119887. Paratype MADAGASCAR ��� Toliara ��� &female;; same collection data as for holotype; FMNH-INS 3119887. Other material MADAGASCAR ��� Toliara ��� 4 &male;&male;; R��serve Sp��ciale d��Andranomena; 20��10���31��� S, 44��28���25��� E; alt. 18 m; dry deciduous forest; 12 Mar. 2010; V. Soarimalala leg.; coll. no. VS-1815; FMNH-INS 3119875 ��� 1 &male;; same collection data as for preceding; ZFMK MYR11137. Description BODY LENGTH. Largest female (paratype): length ca 27.2 mm, width of thoracic shield 14.3 mm, of tergite 8 14.8 mm (widest), height of thoracic shield 8.2 mm, of tergite 8 10.1 mm (highest). COLORATION. Some discoloration may have occurred because of preservation in ethanol. Tergites dorsally orange to brown, especially at posterior margin, laterally green except for paratergite depressions which are yellow. Anal shield light brown to yellowish dorsally as well as posteriorly, laterally green. Thoracic shield green, but grooves and margins yellow-brownish (Fig. 2B). Head and antennae green, legs basally light brown, apically green. HEAD. Wide and short. Eyes with>75 ommatidia. Aberrant ocellus located in antennal groove. Antennae short, with cylindrical joints, protruding posteriorly to leg pair 3. Length of antennomeres 1= 2=3= 4=5Male sexual characters GONOPORE. Covered with a single undivided, rounded membranous plate, very wide, located on basal half of coxa (Fig. 6A). ANTERIOR TELOPODS. Harp with two weak stridulation ribs (Fig. 6E). First podomere with numerous setae mainly at lateral margin, as wide as long. Process of second podomere as long as basal fifth of third podomere, not visible in anterior view, with apical mound of a few sclerotized spots juxtaposed to third podomere. Third podomere with an apical sclerotized spot and a lateral crenulated tooth, mesally with an elevated area carrying sclerotized spots (Fig. 6D, F); lower area carrying 4 spines (Fig. 6D, F). Second and third podomeres sparsely setose. POSTERIOR TELOPODS.Movable finger weakly curved; hollowed-out inner margin with two non-sclerotized lobes and ca five smaller sclerotized spines, posterior aspect with ca 15 large crenulated teeth. Fixed finger wider than movable finger, with slightly curved tip; basally with single membranous lobe, on inner margin covered with a few sclerotized spots. Second and third podomeres lacking long hair, almost completely glabrous. First podomere glabrous except for a few setae at mesal margin. Female sexual characters Washboard with two short interrupted stridulation ribs on each half (Fig. 6G). Vulva covering more than &frac23; of coxa surface. Operculum large, ending before coxal margin. Apical margin of operculum wellrounded, reaching apicalmost point medially. Mesal plate reaching up to half of operculum��s height (Fig. 6J). Distribution and ecology Zoosphaerium silens sp. nov. is currently known from the western dry forest of Kirindy-Mite as well as from the dry forest of Andranomena further north (Fig. 3). These populations differ by an uncorrected p-distance of 2.8% (Table 2)., Published as part of Wesener, Thomas & Sagorny, Christina, 2021, Seven new giant pill-millipede species and numerous new records of the genus Zoosphaerium from Madagascar (Diplopoda, Sphaerotheriida, Arthrosphaeridae), pp. 1-48 in European Journal of Taxonomy 758 (1) on pages 15-17, DOI: 10.5852/ejt.2021.758.1423, http://zenodo.org/record/5075824

Zoosphaerium discolor Wesener, 2009 Material examined MADAGASCAR – Mahajanga • 1 &female;; Forêt de Beanka, 4.9 km S of Ambinda; 18°03.7′ S, 44°31.5′ E; alt. 320 m; slightly disturbed dry deciduous tsingy forest; 27 Oct. 2009; V. Soarimalala leg.; coll. no. VS-1783; FMNH-INS 3119880 • 1 &female;; Majunga, Reserve forestière Beanka, 50.7 km E of Maintirano; 17°52′49″ S, 44°28′08″ E; alt. 140 m; tropical dry forest on tsingy, general collecting; 28–31 Oct. 2009; B.L. Fisher et al. leg.; coll. no. BLF23002; CASENT9068288 • 1 &male;; Forêt de Beanka; 18°01.25′ S, 44°30.08′ E; alt. 220 m; slightly disturbed dry deciduous forest; 14 Jan. 2010; Z.H. Harimpiria leg.; FMNH-INS 3119897B. Remarks This species, previously known from the western dry forests of Bemaraha and Kirindy (Wesener 2009), has now also been recorded from Beanka (Fig. 13C)., Published as part of Wesener, Thomas & Sagorny, Christina, 2021, Seven new giant pill-millipede species and numerous new records of the genus Zoosphaerium from Madagascar (Diplopoda, Sphaerotheriida, Arthrosphaeridae), pp. 1-48 in European Journal of Taxonomy 758 (1) on pages 40-41, DOI: 10.5852/ejt.2021.758.1423, http://zenodo.org/record/5075824

Zoosphaerium villosum Wesener & Sierwald, 2005 Material examined MADAGASCAR – Toamasina • 1 &male;; Parc National de Zahamena, Tetezambatana forest, near jct of Nosivola and Manakambahiny Rivers; 17°44′35″ S, 48°43′46″ E; alt. 860 m; general collecting in camp, rainforest; 18–19 Feb. 2009; B.L. Fisher et al. leg.; coll. no. BLF 21973; CASENT 9068292. Remarks Another locality for this species (Fig. 13C), which appears to be common in northeastern Malagasy lowland rainforests, is added (cf Wesener et al. 2010b).

Zoosphaerium ambatovaky sp. nov. urn:lsid:zoobank.org:act: 42C1A931-D533-4FF6-B76A-CAB8F7E1461D Figs 2C, 3, 5C, 7 Diagnosis The presence of only 7 or 8 ventral spines on the walking legs in Z. ambatovaky sp. nov., especially in combination with a glossy green color, two stridulation ribs on the male harp and more than 10 apical cones on the antenna, is only shared with Z. muscorum Wesener & Bespalova, 2010. The new species differs from Z. muscorum in the presence of an apical spine on the third leg pair (absent in Z. muscorum), the movable finger of the posterior telopod being 2.5 times as long as wide (2 times as long as wide in Z. muscorum) and the second locking carina of the anal shield being 2–3 times as long as the first (4–5 times as long as the first in Z. muscorum). Etymology ‘Ambatovaky’, noun in apposition, after the forest of Ambatovaky, the only known locality for this species (Fig. 3). Material examined Holotype MADAGASCAR – Toamasina • &male;; Rés. Speciale Ambatovaky, Sandrangato River; 16°46′22″ S, 49°15′56″ E; alt. 450 m; rainforest; 20 Feb. 2010; B.L. Fisher et al. leg.; coll. no. BLF24311; CASENT 9068294 /1. Description Male BODY LENGTH. Holotype male: length ca 44 mm, width of thoracic shield 23.1 mm, of tergite 5 24.4 mm (widest), height of thoracic shield 14.1 mm (highest). COLORATION. Some discoloration may have occurred because of preservation in ethanol. Head, collum, thoracic shield and tergites shiny green with a conspicuous narrow light posterior margin (Fig. 2C). Anal shield of a different color, jade-like green. Antennae green, legs basally dark green, apically brownish. HEAD. Eyes with> 95 ommatidia. Antennae long, with cylindrical joints, protruding posteriorly to leg pair 4. Length of antennomeres 1>2> 3=4= 5 GNATHOCHILARIUM AND MANDIBLE. Not dissected. STIGMATIC PLATES. First stigmatic plate triangular, apex well rounded, nearly as wide as at base, slightly turned towards coxa. Hair most abundant on apical margin, very few hairs on basal and lateral margins (Fig. 7C). PLEURITES. First pleurite well-rounded, not projecting. COLLUM. Glabrous. THORACIC SHIELD. Smooth and glabrous, without protuberances. TERGITES. Surface glabrous and shiny, lacking small pits. Tips of paratergites of midbody tergites projecting posteriorly. ENDOTERGUM. Inner section with numerous short triangular spines and longer setae. Middle area above spines with a single row of sparse circular cuticular impressions slightly varying in size. Apically with 1–2 rows of short marginal bristles, protruding to &frac23; of distance of tergite margin (Fig. 5C). Bristles with numerous small spines. ANAL SHIELD. Male anal shield bell-shaped, without hairs, glabrous. Underside with two black locking carinae, posterior one 2–3 times as long as anterior one, both located close to anal shield margin. LEGS. Leg 1 with 4 ventral spines, leg 2 with 6 or 7, leg 3 with 6. First two leg pairs without an apical spine, spine present on leg 3. Leg pairs 4–21 with 7 or 8 ventral spines and an apical spine. On leg 9, femur 2 and tarsus 4.5 times as long as wide. All podomeres with setae (Fig.). Male sexual characters GONOPORE. Covered with a single undivided, rounded membranous plate, covering ¼ of surface of coxa. Gonopore located on basal half of joint (Fig. 7B). ANTERIOR TELOPODS. Harp with two stridulation ribs, one long and well pronounced rib located laterally of a less pronounced and shorter mesal rib (Fig. 7D). First podomere with only a few setae, restricted to apical parts of lateral margins, 1.5 times as wide as long. Process of second podomere nearly as long as basal &frac23; of third podomere, not visible in anterior view, and with an apical mound of a few sclerotized spots juxtaposed to third podomere (Fig. 7E–F). Third podomere with an apical brown sclerotized spot and a field of elevated sclerotized spots at inner margin below brown spot; two small spines located below sclerotized spot (Fig. 7F); depression lateral to field of spots with one small sclerotized spine, lateral margin with up to 6 crenulated teeth (Fig. 7E–F). Second podomere densely setose, third podomere with setae mainly apically and laterally. POSTERIOR TELOPODS. Movable finger (podomere 3) 2.5 times as long as wide, weakly curved towards immovable finger (process of podomere 2). Shallowly hollowed-out inner margin with five sclerotized spines, three positioned apically and two in middle of cavity next to small membranous lobe. Additional sclerotized spine at base of movable finger. Posterior aspect with ca 12 small crenulated teeth (Fig. 7H). Immovable finger thinner and shorter than movable finger, curved towards the latter. Sclerotized spine located on anterior side near base of immovable finger. Inner part of immovable finger apically covered with a few sclerotized spots (Fig. 7I). Movable finger glabrous on both sides except for a few hairs basolaterally. Second podomere in posterior view setose, in anterior view with a few setae on finger. Female Unknown. Distribution and ecology Zoosphaerium ambatovaky sp. nov. is currently known only from the type locality, a lowland rainforest close to the Ille Sainte Marie (Fig. 3). This species seems to be closely related to Z. muscorum from the montane forest of Anjanaharibe-Sud, the only species with which it shares the presence of only 7 or 8 ventral spines on the walking legs, as well as a glossy green color, two stridulation ribs on the male harp and more than 10 apical cones on the antenna. A female Zoosphaerium from the same forest (CASENT 9068291B) belongs to a different species. The forest of Ambatovaky is isolated and shrinking rapidly (Rakotondrasoa et al. 2017).

Zoosphaerium ambrense Wesener, 2009 Material examined MADAGASCAR – Antsiranana • 1 &female;; Montagne dʼAmbre, 2 km away from park entrance; 12°32′ S, 49°10′ E; alt. 1000 m; 23–28 Nov. 1993; J. Coddington, N. Scharff, S. Larcher, C. Griswold and R. Andriamasimanana leg.; NHMD • 1 &male;, 2 &female;&female;; same collection data as for preceding; alt. 1100 m; NHMD • 1 &male;; same collection data as for preceding; NHMD • 1 &male;; same collection data as for preceding; NHMD • 1 &male;; same collection data as for preceding; NHMD • 1 &male;; same collection data as for preceding; NHMD. Remarks We provide collection information for several more specimens of Z. ambrense that were sampled at the type locality., Published as part of Wesener, Thomas & Sagorny, Christina, 2021, Seven new giant pill-millipede species and numerous new records of the genus Zoosphaerium from Madagascar (Diplopoda, Sphaerotheriida, Arthrosphaeridae), pp. 1-48 in European Journal of Taxonomy 758 (1) on page 41, DOI: 10.5852/ejt.2021.758.1423, http://zenodo.org/record/5075824

Zoosphaerium corystoides Wesener, 2009 Material examined MADAGASCAR – Antsiranana • 2 &male;&male;, 2 specs; Montagne dʼAmbre; 12°32′ S, 49°10′ E; alt. 1100 m; 23–28 Nov. 1993; J. Coddington, N. Scharff, S. Larcher, C. Griswold and R. Andriamasimanana leg.; NHMD • 1 &male;; same collection data as for preceding; NHMD. Remarks Additional specimens were collected at the type locality, Montagne dʼAmbre. Zoosphaerium corystoides is currently classified as Least Concern (Rudolf & Wesener 2017d) on the IUCN Red List., Published as part of Wesener, Thomas & Sagorny, Christina, 2021, Seven new giant pill-millipede species and numerous new records of the genus Zoosphaerium from Madagascar (Diplopoda, Sphaerotheriida, Arthrosphaeridae), pp. 1-48 in European Journal of Taxonomy 758 (1) on pages 41-42, DOI: 10.5852/ejt.2021.758.1423, http://zenodo.org/record/5075824, {"references":["Rudolf E. & Wesener T. 2017 d. Zoosphaerium corystoides. The IUCN Red List of Threatened Species 2017: e. T 65524417 A 65527885. Available from https: // doi. org / 10.2305 / IUCN. UK. 2017 - 1. RLTS. T 65524417 A 65527885. en [accessed 7 Apr. 2020]."]}

Zoosphaerium beanka sp. nov. urn:lsid:zoobank.org:act: 7F9589AA-AD5D-473F-A7C1-FB94F30FF8AD Figs 2D, 3, 5D, 8 Diagnosis The shape and position of the locking carinae place Z. beanka sp. nov. in the Z. platylabum speciesgroup, but the telopods of Z. beanka sp. nov. have almost no similarities to those of any other described giant pill-millipede species except Z. tsingy. Both species share characters of the posterior telopods, namely the presence of spines and at least one large membranous lobe on the movable finger, and an uncurved, wide immovable finger (2.7 times as long as wide) that is parallel to the movable finger. Zoosphaerium beanka sp. nov. differs from the latter in the light brown color (olive in Z. tsingy), the presence of only four apical cones on the antenna (> 15 in Z. tsingy) and the differently shaped anterior telopods, in which telopoditomere 3 lacks crenulated teeth (present in Z. tsingy). Etymology ‘Beanka’, noun in apposition, after the type and only known locality for this species, the dry forest of Beanka (Fig. 3). Material examined Holotype MADAGASCAR – Mahajanga • &male;; Forêt de Beanka; 18°01′25″ S, 44°30′08″ E; alt. 220 m; slightly disturbed dry deciduous forest; 14 Jan. 2010; Z.H. Harimpitia leg.; coll. no. Z.H.H-031; FMNH-INS 3119897. Description BODY LENGTH. Male (holotype): length 36.7 mm, width of thoracic shield up 22.1 mm (widest), height of thoracic shield 11.8 mm (highest). COLORATION. Head, collum, thoracic shield, tergites and anal shield light brown (Fig. 2D). Antennae green, legs light brown. HEAD. Eyes with more than 75 ommatidia. Antennae short, protruding back to leg 3. Length of antennomeres 1>2=3 =4=5 Male sexual characters GONOPORE. Covered with a single undivided, rounded membranous plate, covering ¼ of surface of coxa. Gonopore located centrally on joint (Fig. 8B). ANTERIOR TELOPODS. Harp with two stridulation ribs, mesal rib smaller than lateral one, both ribs well developed (Fig. 8F). First podomere 1.2 times as wide as long. Second podomere process lobe-like and slightly curved, with rounded edges, protruding up to half of third podomere height, apically with sclerotized spots. Third podomere long, as long as first; cavity mesally with numerous sclerotized spots and three thin, sclerotized spines; one sclerotized spot located close to tip. Podomeres 1–3 covered on both sides with numerous long, isolated hairs. POSTERIOR TELOPODS. Movable finger with a well-rounded tip, not tapering towards tip; hollowed-out inner margin with one non-sclerotized lobe and 3 or 4 sclerotized spines, posterior aspect with ca 14 small sclerotized teeth. Movable finger slightly longer than fixed finger, the latter straight, as wide as third podomere. Fixed finger at margin with numerous sclerotized spots. All podomeres covered on both sides with numerous, long isolated hairs, only tips of chela glabrous (Fig. 8G–H). Female Unknown. Distribution and ecology Zoosphaerium beanka sp. nov. is currently known only from the dry forest of Beanka (Fig. 3). With most known dry forest giant pill-millipede species being widespread (Wesener & Wägele 2008; Wesener 2009), this species can be expected in surrounding dry forest fragments., Published as part of Wesener, Thomas & Sagorny, Christina, 2021, Seven new giant pill-millipede species and numerous new records of the genus Zoosphaerium from Madagascar (Diplopoda, Sphaerotheriida, Arthrosphaeridae), pp. 1-48 in European Journal of Taxonomy 758 (1) on pages 25-27, DOI: 10.5852/ejt.2021.758.1423, http://zenodo.org/record/5075824, {"references":["Wesener T. & Wagele J. W. 2008. The giant pill-millipedes of Madagascar: revision of the genus Zoosphaerium (Myriapoda, Diplopoda, Sphaerotheriida). Zoosystema 30 (1): 5 - 82."]}

Zoosphaerium aureum Wesener, 2009 Material examined MADAGASCAR – Antsiranana • 1 &male;; Montagne dʼAmbre; 12°32′ S, 49°10′ E; alt. 1100 m; 23–28 Nov. 1993; NHMD. Remarks We provide collection information for one additional specimen from the type locality. Zoosphaerium aureum is currently classified as Least Concern (Rudolf & Wesener 2017b) on the IUCN Red List., Published as part of Wesener, Thomas & Sagorny, Christina, 2021, Seven new giant pill-millipede species and numerous new records of the genus Zoosphaerium from Madagascar (Diplopoda, Sphaerotheriida, Arthrosphaeridae), pp. 1-48 in European Journal of Taxonomy 758 (1) on page 41, DOI: 10.5852/ejt.2021.758.1423, http://zenodo.org/record/5075824

Zoosphaerium nigrum sp. nov. urn:lsid:zoobank.org:act: 4F170F43-C7A9-44A4-A004-47FD58FDBA2F Figs 2A, 3–4, 5A Diagnosis Posterior telopods (Fig. 4H–I) identify this species as a member of the Z. libidinosum species-group. Within this group, Zoosphaerium nigrum sp. nov. shares a number of characters with only Z. anomalum (de Saussure & Zehntner, 1902) and Z. pulchellum Wesener, 2009. These include the presence of two locking carinae, four apical cones on each antenna, and shiny, glossy tergites. Moreover, the second and third podomeres of the posterior telopods are covered with numerous minute hairs. Zoosphaerium nigrum sp. nov. differs from these two species in the black coloration (jade-like green in the others), the presence of only a single weak stridulation rib on the anterior telopod and differently shaped podomeres. Etymology ‘Nigrum’, adjective, after the dark colour of the species. Material examined Holotype MADAGASCAR – Fianarantosa • &male;; Parc National Befotaka-Midongy, Papango, 28.5 km S of Midongy-Sud, Mount Papango; 23°50′27″ S, 46°57′27″ E; alt. 1250 m; montane rainforest; 17 Nov. 2006; B.L. Fisher et al. leg.; coll. no. BLF14945; CASENT 9068289. Description Male BODY LENGTH. Holotype male: length ca 28 mm, width of thoracic shield 13.1 mm, of tergite 7 13.5 mm (widest), height of thoracic shield 7.1 mm, of tergite 7 7.4 mm (highest). COLORATION. Some discoloration may have occurred because of preservation in ethanol. Tergites, head, anal shield and appendages dark green to black (Fig. 2A). Posterior margin of tergites with a thin brown margin. HEAD. Wide and short. Eyes with>120 ommatidia. Aberrant ocellus located in antennal groove. Antennae short, with cylindrical joints, reaching posteriorly to leg pair 3. Length of antennomeres 1= 2=3= 4=5Male sexual characters GONOPORE. Covered with a single undivided, rounded membranous plate (Fig. 4B). ANTERIOR TELOPODS. Harp with a single weak stridulation rib (Fig. 4E). First podomere with only a few setae, restricted to apical parts of lateral margins, 1.5 times wider than long. Process of second podomere as long as &frac23; of third podomere, not visible in anterior view, with apical mound of a few sclerotized spots juxtaposed to third podomere. Third podomere without sclerotized teeth, but with a conspicuous process carrying sclerotized spots (Fig. 4F–G); base of process with four spines (Fig. 4G). Second and third podomeres sparsely setose. POSTERIOR TELOPODS.Movable finger weakly curved; hollowed-out inner margin with two non-sclerotized lobes and ca three smaller sclerotized spines, posterior aspect with ca 29 crenulated teeth. Fixed finger thinner than movable finger, with slightly curved tip; basally with single membranous lobe, on inner margin covered with numerous sclerotized spots. Second and third podomeres without any long hair, but on both sides densely covered with very small pits, one very small seta standing in each pit. First podomere without those pits but with some isolated, standard longer hairs. Female Unknown. Distribution and ecology Zoosphaerium nigrum sp. nov. is currently known only from the montane rainforests of Midongy- Befotaka (Fig. 3) where it coexists with another species of the genus, Z. spinopiligerum sp. nov. (see below), as well as with a species of the genus Sphaeromimus (Moritz & Wesener 2017)., Published as part of Wesener, Thomas & Sagorny, Christina, 2021, Seven new giant pill-millipede species and numerous new records of the genus Zoosphaerium from Madagascar (Diplopoda, Sphaerotheriida, Arthrosphaeridae), pp. 1-48 in European Journal of Taxonomy 758 (1) on pages 9-14, DOI: 10.5852/ejt.2021.758.1423, http://zenodo.org/record/5075824, {"references":["Saussure H. de & Zehntner L. 1902. Myriapodes de Madagascar. In: Grandidier A. (ed.) Histoire physique, naturelle et politique de Madagascar 27 (53): i - viii, 1 - 356.","Moat J. & Smith P. 2007. Atlas of the Vegetation of Madagascar Vegetation / Atlas de la Vegetation de Madagascar. Royal Botanic Gardens, Kew (bilingual edition).","Moritz L. & Wesener T. 2017. Integrative description of two new species of Malagasy chirping giant pillmillipedes, genus Sphaeromimus (Diplopoda, Sphaerotheriida, Arthrosphaeridae). European Journal of Taxonomy 381: 1 - 25. https: // doi. org / 10.5852 / ejt. 2017.381"]}

Zoosphaerium ambatovaky sp. nov. urn:lsid:zoobank.org:act: 42C1A931-D533-4FF6-B76A-CAB8F7E1461D Figs 2C, 3, 5C, 7 Diagnosis The presence of only 7 or 8 ventral spines on the walking legs in Z. ambatovaky sp. nov., especially in combination with a glossy green color, two stridulation ribs on the male harp and more than 10 apical cones on the antenna, is only shared with Z. muscorum Wesener & Bespalova, 2010. The new species differs from Z. muscorum in the presence of an apical spine on the third leg pair (absent in Z. muscorum), the movable finger of the posterior telopod being 2.5 times as long as wide (2 times as long as wide in Z. muscorum) and the second locking carina of the anal shield being 2–3 times as long as the first (4–5 times as long as the first in Z. muscorum). Etymology ‘Ambatovaky’, noun in apposition, after the forest of Ambatovaky, the only known locality for this species (Fig. 3). Material examined Holotype MADAGASCAR – Toamasina • &male;; Rés. Speciale Ambatovaky, Sandrangato River; 16°46′22″ S, 49°15′56″ E; alt. 450 m; rainforest; 20 Feb. 2010; B.L. Fisher et al. leg.; coll. no. BLF24311; CASENT 9068294 /1. Description Male BODY LENGTH. Holotype male: length ca 44 mm, width of thoracic shield 23.1 mm, of tergite 5 24.4 mm (widest), height of thoracic shield 14.1 mm (highest). COLORATION. Some discoloration may have occurred because of preservation in ethanol. Head, collum, thoracic shield and tergites shiny green with a conspicuous narrow light posterior margin (Fig. 2C). Anal shield of a different color, jade-like green. Antennae green, legs basally dark green, apically brownish. HEAD. Eyes with> 95 ommatidia. Antennae long, with cylindrical joints, protruding posteriorly to leg pair 4. Length of antennomeres 1>2> 3=4= 5Male sexual characters GONOPORE. Covered with a single undivided, rounded membranous plate, covering ¼ of surface of coxa. Gonopore located on basal half of joint (Fig. 7B). ANTERIOR TELOPODS. Harp with two stridulation ribs, one long and well pronounced rib located laterally of a less pronounced and shorter mesal rib (Fig. 7D). First podomere with only a few setae, restricted to apical parts of lateral margins, 1.5 times as wide as long. Process of second podomere nearly as long as basal &frac23; of third podomere, not visible in anterior view, and with an apical mound of a few sclerotized spots juxtaposed to third podomere (Fig. 7E–F). Third podomere with an apical brown sclerotized spot and a field of elevated sclerotized spots at inner margin below brown spot; two small spines located below sclerotized spot (Fig. 7F); depression lateral to field of spots with one small sclerotized spine, lateral margin with up to 6 crenulated teeth (Fig. 7E–F). Second podomere densely setose, third podomere with setae mainly apically and laterally. POSTERIOR TELOPODS. Movable finger (podomere 3) 2.5 times as long as wide, weakly curved towards immovable finger (process of podomere 2). Shallowly hollowed-out inner margin with five sclerotized spines, three positioned apically and two in middle of cavity next to small membranous lobe. Additional sclerotized spine at base of movable finger. Posterior aspect with ca 12 small crenulated teeth (Fig. 7H). Immovable finger thinner and shorter than movable finger, curved towards the latter. Sclerotized spine located on anterior side near base of immovable finger. Inner part of immovable finger apically covered with a few sclerotized spots (Fig. 7I). Movable finger glabrous on both sides except for a few hairs basolaterally. Second podomere in posterior view setose, in anterior view with a few setae on finger. Female Unknown. Distribution and ecology Zoosphaerium ambatovaky sp. nov. is currently known only from the type locality, a lowland rainforest close to the Ille Sainte Marie (Fig. 3). This species seems to be closely related to Z. muscorum from the montane forest of Anjanaharibe-Sud, the only species with which it shares the presence of only 7 or 8 ventral spines on the walking legs, as well as a glossy green color, two stridulation ribs on the male harp and more than 10 apical cones on the antenna. A female Zoosphaerium from the same forest (CASENT 9068291B) belongs to a different species. The forest of Ambatovaky is isolated and shrinking rapidly (Rakotondrasoa et al. 2017)., Published as part of Wesener, Thomas & Sagorny, Christina, 2021, Seven new giant pill-millipede species and numerous new records of the genus Zoosphaerium from Madagascar (Diplopoda, Sphaerotheriida, Arthrosphaeridae), pp. 1-48 in European Journal of Taxonomy 758 (1) on pages 21-23, DOI: 10.5852/ejt.2021.758.1423, http://zenodo.org/record/5075824, {"references":["Wesener T., Bespalova I. & Sierwald P. 2010 b. Madagascar's living giants: discovery of five new species of endemic giant pill-millipedes from Madagascar (Diplopoda: Sphaerotheriida: Arthrosphaeridae: Zoosphaerium). African Invertebrates 51 (1): 133 - 161.","Rakotondrasoa L. O., Malaisse F. & Bogaert J. 2017. Modelisation de la dynamique du paysage forestier de la Reserve Speciale d'Ambatovaky (Nord-Est de Madagascar). Tropicultura 35 (4): 312 - 324."]}

Genus Zoosphaerium Pocock, 1895 Zoosphaerium Pocock, 1895: 410. Globotherium Brolemann, 1922: 230 (formal synonymization in Wesener & Wägele 2008: 9). Heligmasoma Chamberlin, 1921: 58 (formal synonymization in Wesener & Wägele 2008: 9). See Wesener (2016) for a summary; five additional species have been described since then (Sagorny & Wesener 2017; Wesener & Anilkumar 2020). Type species Sphaerotherium neptunus Butler, 1872. Other species included 66 and 11 nomina dubia (including the species described below). Distribution Endemic to Madagascar. Remarks The diverse genus Zoosphaerium has been divided into six species-groups (Wesener & Wägele 2008; Wesener 2009). Six additional species are currently not assignable to any group (Sagorny & Wesener 2017). Four species are only known from females and an additional 11 species are nomina dubia. A key to the species-groups has been published elsewhere (Wesener 2009)., Published as part of Wesener, Thomas & Sagorny, Christina, 2021, Seven new giant pill-millipede species and numerous new records of the genus Zoosphaerium from Madagascar (Diplopoda, Sphaerotheriida, Arthrosphaeridae), pp. 1-48 in European Journal of Taxonomy 758 (1) on pages 6-7, DOI: 10.5852/ejt.2021.758.1423, http://zenodo.org/record/5075824, {"references":["Pocock R. I. 1895. Description of new genera of Zephronidae, with brief preliminary diagnoses of some new species. Annals and Magazine of Natural History, Series 6 16 (95): 409 - 415. https: // doi. org / 10.1080 / 00222939508680293","Brolemann H. W. 1922. Liste des myriapodes de l'Academie Malagache de Tananarive. Bulletin de la Societe zoologique de France 47: 223 - 278. https: // doi. org / 10.5962 / bhl. part. 3470","Wesener T. & Wagele J. W. 2008. The giant pill-millipedes of Madagascar: revision of the genus Zoosphaerium (Myriapoda, Diplopoda, Sphaerotheriida). Zoosystema 30 (1): 5 - 82.","Chamberlin R. V. 1921. New Chilopoda and Diplopoda from the East Indian region. Annals and Magazine of Natural History, Series 9 7 (37): 50 - 87. https: // doi. org / 10.1080 / 00222932108632489","Wesener T. 2016. The giant pill-millipedes, order Sphaerotheriida - an annotated species catalogue with morphological atlas and list of apomorphies (Arthropoda: Diplopoda). Bonn Zoological Bulletin - Supplementum 63: 1 - 104.","Sagorny C. & Wesener T. 2017. Two new giant pill-millipede species of the genus Zoosphaerium endemic to the Bemanevika area in northern Madagascar (Diplopoda, Sphaerotheriida, Arthrosphaeridae). Zootaxa 4263 (2): 273 - 294. https: // doi. org / 10.11646 / zootaxa. 4263.2.4","Wesener T. & Anilkumar P. A. 2020. The millipedes collected by the Museum \" La Specola \" on Madagascar 1989 / 1991, with the description of three new species of giant pill-millipedes (Diplopoda, Sphaerotheriida, Arthrosphaeridae). In: Korsos Z. & Danyi L. (eds) Proceedings of the 18 th International Congress of Myriapodology, Budapest, Hungary. ZooKeys 930: 3 - 35. https: // doi. org / 10.3897 / zookeys. 930.47620"]}

Zoosphaerium neptunus (Butler, 1872) Sphaerotherium digitale de Saussure & Zehntner, 1897: pl. 5 fig. 16. Sphaerotherium pygidiale de Saussure & Zehntner, 1902: 42. See Wesener & Wägele (2008) for a revision and Wesener (2016) for a complete bibliography. Material examined MADAGASCAR – Antsiranana • 3 &male;&male;; SAVA, Parc National de Marojejy, 6.5 km NW of Manantenina village; 14°27′21.2″ S, 49°46′29.8″ E; alt. 780 m; disturbed lowland humid forest, pitfalls; 31 May 2016; V. Soarimalala leg.; FMNH-INS 2858681 •> 30 juvs; Marojejy; 14°26′ S, 49°45′ E; alt. 700 m; 10–17 Nov. 1993; NHMD. – Toamasina • 1 &female;; Reserve Betampona, Camp Rendrirendry, 34.1 km 332° from Toamasina; 17°55.26″ S, 049°11.59″ E; alt. 390 m; general collecting in camp, rainforest; 28 Nov. 2005; Brian L. Fisher et al. leg.; coll. no. BLF 13126; CASENT 9032818 • 1 &female;; Reserve Nationale Integrale Betampona, Betampona, 35.1 km NW of Toamasina; 17°55.05″ S, 49°12.03″ E; alt. 500 m; rainforest, general collecting; 15–17 Dec. 2007; Brian L. Fisher et al. leg.; coll. no. BLF 19514; CASENT 9068298 • 1 &female;; Region Atsinanana, Reserve Naturelle Integrale, along Piste Principale (PP); 17°54.8′ S, 49°11.6′ E; alt. 400–500 m; in relatively intact lowland humid forest, pitfall 2, bucket 6; 10 Nov. 2015; Steven M. Goodman leg.; FMNH-INS SMG-19237 • 1 &female;; same collection data as for preceding; by hand; coll. no. SMG-19235; FMNH-INS 2858602 • 1 &female;; same collection data as for preceding; pitfall 3, bucket 5; 11 Nov. 2015; coll. no. SMG-19246; FMNH-INS 2858597 • 1 &female;; same collection data as for preceding; coll. no. SMG-19247; FMNH-INS 2858600 • 1 &female;; same collection data as for preceding; pitfall 1; 12 Nov. 2015; coll. no. SMG-19255; FMNH-INS 2858599 • 1 &female;; same collection data as for preceding; pitfall 3; coll. no. SMG-19256; FMNH-INS 2858604 • 1 &male;; same collection data as for preceding; pitfall 2; coll. no. SMG-19254; FMNH-INS 2858593 • 1 &male;; same collection data as for preceding; pitfall 1; 13 Nov. 2015; coll. no. SMG-19263; FMNH-INS 2858594 • 1 &male;; same collection data as for preceding; coll. no. SMG-19264; FMNH-INS 2858591 • 1 &female;; same collection data as for preceding; pitfall 3; 15 Nov. 2015; coll. no. SMG-19271; FMNH-INS 2858603 • 1 &female;; Region Atsinanana, Reserve Naturelle Integrale, along Piste Principale (PP) between 1650 and 2550 m; 17°54.62′ S, 49°11.70′ E; alt. 460–520 m; in undisturbed lowland humid forest, pitfall 4, bucket 8; 17 Nov. 2015; Steven M. Goodman leg.; coll. no. SMG-19289; FMNH-INS 2858598 • 1 &male;; same collection data as for preceding; bucket 11; coll. no. SMG-19291; FMNH-INS 2858590 • 1 &female;; same collection data as for preceding; pitfall 5, bucket 3; coll. no. SMG-19290; FMNH-INS 2858596 • 1 &male;; same collection data as for preceding; pitfall 4, bucket 1; 21 Nov. 2015; coll. no. SMG-19313; FMNH- INS 2858592. Remarks We provide further locality information including the lowland forest of Betampona and lowland forests in Marojejy for this relatively widespread species of Zoosphaerium (Fig. 13A). The gigantic females of Z. neptunus are currently the largest Sphaerotheriida known, reaching the size of a tennis ball or small orange when rolled up. The juveniles of this species show swarming behavior around Andasibe (Wesener & Schütte 2010), a so far unique behavior for giant pill-millipedes., Published as part of Wesener, Thomas & Sagorny, Christina, 2021, Seven new giant pill-millipede species and numerous new records of the genus Zoosphaerium from Madagascar (Diplopoda, Sphaerotheriida, Arthrosphaeridae), pp. 1-48 in European Journal of Taxonomy 758 (1) on pages 37-38, DOI: 10.5852/ejt.2021.758.1423, http://zenodo.org/record/5075824, {"references":["Saussure H. de & Zehntner L. 1897. Atlas de l'histoire naturelle des Myriapodes. Histoire physique, naturelle et politique de Madagascar 27 (53): 1 - 12.","Saussure H. de & Zehntner L. 1902. Myriapodes de Madagascar. In: Grandidier A. (ed.) Histoire physique, naturelle et politique de Madagascar 27 (53): i - viii, 1 - 356.","Wesener T. & Wagele J. W. 2008. The giant pill-millipedes of Madagascar: revision of the genus Zoosphaerium (Myriapoda, Diplopoda, Sphaerotheriida). Zoosystema 30 (1): 5 - 82.","Wesener T. 2016. The giant pill-millipedes, order Sphaerotheriida - an annotated species catalogue with morphological atlas and list of apomorphies (Arthropoda: Diplopoda). Bonn Zoological Bulletin - Supplementum 63: 1 - 104.","Wesener T. & Schutte K. 2010. Swarming behaviour and mass occurrences in the world's largest giant pill-millipede species, Zoosphaerium neptunus, on Madagascar and its implication for conservation efforts (Diplopoda: Sphaerotheriida). Madagascar Conservation and Development 5 (2): 89 - 94."]}

Zoosphaerium platylabum species-group Remarks Species can be identified as members of the Zoosphaerium platylabum species-group based on the number of locking carinae (always 2) of the anal shield that are located close to the last pleurite, as well as on morphological characters of the female vulva, whose apical processes are usually sharpedged (with the exception of those in Z. tsingy Wesener, 2009). The posterior telopods (Wesener & Wägele 2008) show two vastly different modifications: the movable finger is either very thick, or quite elongated and slender. The species-group now contains 6 +3 described species distributed in humid as well as western dry forests all over Madagascar. Species of this group are conspicuously absent from the southern spiny forest ecosystem (Wesener 2009). Key to species of the Zoosphaerium platylabum species-group 1. Movable finger of posterior telopods very thick, Z. platylabum (de Saussure & Zehntner, 1902) – Movable finger 1.6 times as long as wide, at inner margin without lobe. First stigmatic plate large, lobe triangular and tapering towards well-rounded tip. Antenna with 23–35 apical cones.................................................................................................... Z. pseudoplatylabum Wesener, 2009 3. Anterior edges of thoracic shield slightly projecting towards head. Movable finger shorter than fixed finger........................................................................................... Z. solitarium Wesener, 2009 – Anterior edges of thoracic shield well-rounded. Movable finger at least slightly longer than fixed finger..................................................................................................................................................4 4. Male harp with two stridulation ribs. Movable finger 3 times as long as wide.....................5 – Male harp with a single stridulation rib. Movable finger>4 times as long as wide................7 5. Movable finger on posterior telopods with five spines and no membranous lobes. Immovable finger of posterior telopod slender, 4 times as long as wide, apical part tapering and curved towards movable finger. Only 7 or 8 ventral spines on legs. Found in eastern rainforests, Masoala Peninsula.............................................................................................................................. Z. masoala sp. nov. – Movable finger on posterior telopods with spines and at least one large, triangular membranous lobe. Immovable finger of posterior telopod wide, 2.7 times as long as wide, not curved, parallel to movable finger, apically only weakly tapering. 10 or more ventral spines on legs. Found in western dry forests..........................................................................................................................................6 6. Light brown. Antennae with four apical cones. Third podomere of anterior telopods lacking crenulated teeth..................................................................................................... Z. beanka sp. nov. – Olive green. Antennae with>15 apical cones. Third podomere of anterior telopods laterally with 2 or 3 crenulated teeth...................................................................... Z. tsingy Wesener, 2009 7. Tergites shiny black, appendages red. Anal shield strongly developed, shaped like a helmet. Fixed finger of posterior telopod slightly curved towards movable finger. Operculum of vulva deeply notched, divided into two sharp ‘hornsʼ. Female washboard on each side with a single stridulation rib.................................................................................... Z. corystoides Wesener, 2009 – Tergites light brown or olive-brownish, appendages brown or green. Anal shield well-rounded. Fixed finger of posterior telopod parallel to movable finger.........................................................8 8. Up to 30 mm long. Movable finger of posterior telopod with 25 teeth, three spines located at apex. Endotergum with two dense rows of long marginal bristles, protruding above tergite margin. (Female unknown)............................................................................... Z. voahangy sp. nov. – Up to 55 mm long. Movable finger of posterior telopod with ca 15 teeth, three spines located at regular intervals along entire margin. Endotergum with a single row of short, dense marginal bristles, protruding up to &frac23; of distance towards tergite margin. Operculum of vulva only very shallowly notched, forming two sharp horns. Female washboard with four stridulation ribs......................................................................................................................... Z. broelemanni Wesener, 2009, Published as part of Wesener, Thomas & Sagorny, Christina, 2021, Seven new giant pill-millipede species and numerous new records of the genus Zoosphaerium from Madagascar (Diplopoda, Sphaerotheriida, Arthrosphaeridae), pp. 1-48 in European Journal of Taxonomy 758 (1) on pages 24-25, DOI: 10.5852/ejt.2021.758.1423, http://zenodo.org/record/5075824, {"references":["Wesener T. & Wagele J. W. 2008. The giant pill-millipedes of Madagascar: revision of the genus Zoosphaerium (Myriapoda, Diplopoda, Sphaerotheriida). Zoosystema 30 (1): 5 - 82.","Saussure H. de & Zehntner L. 1902. Myriapodes de Madagascar. In: Grandidier A. (ed.) Histoire physique, naturelle et politique de Madagascar 27 (53): i - viii, 1 - 356."]}

Zoosphaerium spinopiligerum sp. nov. urn:lsid:zoobank.org:act: B5E6A6B0-F750-4227-8F0B-4F6417FDF88D Figs 2G, 3, 10B, 12 Diagnosis The greatly elongated posterior telopods identify Zoosphaerium spinopiligerum sp. nov. as a member of the Z. piligerum species-group. Zoosphaerium spinopiligerum sp. nov. shares the shape of the vulva and the anal shield, as well as a similar posterior telopod with Z. pseudopiligerum. Zoosphaerium spinopiligerum sp. nov. differs from the latter in the non-glossy tergites, differences in the shape of the anterior telopods, as well as the higher number of ventral spines on the legs (see key above). Etymology ���Spinopiligerum���, noun in apposition, as a reference to the higher number of ventral spines on the legs compared to related species. Material examined Holotype MADAGASCAR ��� Fianarantsoa ��� &male;; Parc National de Midongy-Befotaka, 6 km N of Befotaka village; 23��46.7��� S, 47��06.4��� E; alt. 815 m; dense humid transitional lowland and montane forest; 8 Feb. 2008; V. Soarimalala leg.; coll. no. VS-1442; FMNH-INS 3119885. Other material MADAGASCAR ��� Fianarantsoa ��� 1 &female;; Parc National de Midongy-Befotaka, 11.5 km SW of Befotaka village; 23��53.3��� S, 46��53.3��� E; alt. 1053 m; dense humid transitional lowland and montane forest; 27 Feb. 2008; V. Soarimalala leg.; coll. no. VS-1536; FMNH-INS 3119888. Description BODY LENGTH. Male: length 37.1 mm, width of thoracic shield 22.3 mm (widest), height of thoracic shield 11.1 mm (highest). Female (paratype): length 28.8 mm, width of thoracic shield 16.8 mm, height of thoracic shield up to 8.7 mm. COLORATION. Tergites brown with dark brown posterior margin (Fig. 2G). Collum dark brown to green. Head dark green with yellow-brown markings. Anal shield brown. Antennae dark green, legs basally light brown, apically dark green. HEAD. Eyes with more than 80 ommatidia. Antennae long and slender, protruding back to leg 3. Length of antennomeres 1 =2= 3=4 =5Male sexual characters GONOPORE. Covered with a single undivided, rounded, apically membranous, basally sclerotized plate, covering �� of surface of coxa. Gonopore located basally on joint (Fig. 12B). ANTERIOR TELOPODS. Harp with two stridulation ribs, both ribs of same size and well developed, located in mesal corner (Fig. 12D). First podomere 1.3 times as wide as long. Second podomere process lobe- like and slightly curved, with rounded edges, protruding up to &frac23; of third podomere height, apically with sclerotized spots, visible in anterior view (Fig. 12D���F). Third podomere narrow, half as long as first; cavity mesally with numerous sclerotized spots and three thin, sclerotized spines (Fig. 12E���F); one sclerotized spot located close to tip; laterally with up to 4 crenulated teeth. Podomeres 1���3 on both sides with numerous long, isolated hairs. POSTERIOR TELOPODS (FIGS). Movable finger strongly curved, apically with invagination. Hollowed-out inner margin with one large triangular membranous lobe and 3���5 smaller sclerotized spines (Fig. 12H). Posterior aspect of movable finger with ca 11 small, sclerotized teeth (Fig. 12I). Immovable finger slender, with weakly curved tip, slightly shorter than movable finger. Anterior side of immovable finger basally with a membranous lobe and a sclerotized spine, at tip with numerous small sclerotized spots (Fig. 12I). All three podomeres of posterior telopods on both sides glabrous. Inner horns of syncoxite apically curved posteriorly. Tips sharp, entire horns covered with numerous short hairs. Female sexual characters Subanal plate wide, slightly triangular. Washboard with well-developed stridulation ribs, with 3 ribs on each half (Fig. 12G). Vulva large, covering more than &frac23; of coxa (Fig. 12J). Operculum small and low, ending far before coxal margin. Apical margin of operculum medially notched, lateral tip twice as high as mesal one, well rounded. Mesal plate long, extending beyond operculum. Distribution and ecology Zoosphaerium spinopiligerum sp. nov. occurs in sympatry with Z. nigrum sp. nov. and Sphaeromimus midongy Moritz & Wesener, 2017 in the remote mountainous forests of Midongy in the southeast of Madagascar (Fig. 3). Remarks The female specimen examined here is only tentatively placed in Z. spinopiligerum sp. nov. as its COI barcode shows quite a p-distance to the male holotype (Table 2). Such a distance is far higher than usually present in millipedes. New locality data for Zoosphaerium species in Madagascar, Published as part of Wesener, Thomas & Sagorny, Christina, 2021, Seven new giant pill-millipede species and numerous new records of the genus Zoosphaerium from Madagascar (Diplopoda, Sphaerotheriida, Arthrosphaeridae), pp. 1-48 in European Journal of Taxonomy 758 (1) on pages 34-37, DOI: 10.5852/ejt.2021.758.1423, http://zenodo.org/record/5075824, {"references":["Moritz L. & Wesener T. 2017. Integrative description of two new species of Malagasy chirping giant pillmillipedes, genus Sphaeromimus (Diplopoda, Sphaerotheriida, Arthrosphaeridae). European Journal of Taxonomy 381: 1 - 25. https: // doi. org / 10.5852 / ejt. 2017.381"]}

Zoosphaerium smaragdinum Wesener, 2009 Material examined MADAGASCAR – Antsiranana • 3 juvs; Marojejy; 14°26′ S, 49°45′ E; alt. 700 m; 10–17 Nov. 1993; NHMD. Remarks Three more specimens have been collected from the type locality. Zoosphaerium smaragdinum is currently classified as Near Threatened (Rudolf & Wesener 2017c) on the IUCN Red List., Published as part of Wesener, Thomas & Sagorny, Christina, 2021, Seven new giant pill-millipede species and numerous new records of the genus Zoosphaerium from Madagascar (Diplopoda, Sphaerotheriida, Arthrosphaeridae), pp. 1-48 in European Journal of Taxonomy 758 (1) on page 41, DOI: 10.5852/ejt.2021.758.1423, http://zenodo.org/record/5075824, {"references":["Rudolf E. & Wesener T. 2017 c. Zoosphaerium smaragdinum. The IUCN Red List of Threatened Species 2017: e. T 65525770 A 65528000. Available from https: // doi. org / 10.2305 / IUCN. UK. 2017 - 1. RLTS. T 65525770 A 65528000. en [accessed 7 Apr. 2020]."]}

Microsphaerotherium anjozorobe Wesener, 2009 Material examined MADAGASCAR – Antananarivo • 1 &female;; Analamanga, 6.7 km SE of Anjozorobe village, Andasin’i Jaonera; 18°25.3′ S, 47°56.6′ E; alt. 1250 m; in disturbed montane forest, pitfalls 1–3; 19–23 Feb. 2016; Steven M. Goodman leg.; FMNH-INS 2858679. Remarks This is another record of this rare miniaturized species (Wesener 2009), previously known only from the type series collected near Andranomay. This species is currently classified as Critically Endangered (Rudolf & Wesener 2017e) on the IUCN Red List., Published as part of Wesener, Thomas & Sagorny, Christina, 2021, Seven new giant pill-millipede species and numerous new records of the genus Zoosphaerium from Madagascar (Diplopoda, Sphaerotheriida, Arthrosphaeridae), pp. 1-48 in European Journal of Taxonomy 758 (1) on pages 43-44, DOI: 10.5852/ejt.2021.758.1423, http://zenodo.org/record/5075824, {"references":["Rudolf E. & Wesener T. 2017 e. Microsphaerotherium anjozorobe. The IUCN Red List of Threatened Species 2017: e. T 65524228 A 65527805. Available from https: // doi. org / 10.2305 / IUCN. UK. 2017 - 1. RLTS. T 65524228 A 65527805. en [accessed 7 Apr. 2020]."]}

stellatum Carl, 1912c: 122-124, text fig. 10 [Castanotherium]. Loka und Umgebung bis 1300 m üb. M., Süd-Celebes (coll. Sarasin). More than one &female;. The MHNG collection contains two specimens in alcohol (MHNG-ARTO-14304). The identification labels have “ types Loka Celebes mérid. coll. Sarasin” and “ types Celebes mérid. coll. Sarasin” written on them respectively, indicating that the specimens are syntypes. There are seven syntypes (referred to as “ Typen ” in the NMB catalogue) in the NMB (inventory number NMB- DIPL-00120a). Castanotherium stellatum Carl, 1912, Zephroniidae, Published as part of Hollier, John, Stöckli, Eduard, Wesener, Thomas, Mesibov, Robert, Decker, Peter & Sierwald, Petra, 2020, An annotated list of the millipede (Diplopoda) species described by Johann Carl, pp. 183-240 in Revue suisse de Zoologie 127 (1) on page 231, DOI: 10.35929/RSZ.0015, http://zenodo.org/record/5743454, {"references":["Carl J. 1912 c. Die Diplopoden-Fauna von Celebes. Revue suisse de Zoologie 20: 73 - 206."]}

Scolodesmus iuliformis iuliformis (Carl, 1905), Paradoxosomatidae iuliforme volzi Carl, 1913a: 202-204, fig. 1 [Strongylosoma]. Yonni und Falaba (nordöstliches Sierra-Leone). Unspecified number of &male; and &female;. The MHNG collection contains seven specimens in alcohol (MHNG-ARTO-14540). There is a separate vial containing gonopods (MHNG-ARTO-14541). The original identification labels in the jar have “ Sierra Leone W. Volz” written on them, indicating that the specimens are syntypes. There is also an undated typewritten label indicating that Hoffman identified the specimens as Scolodesmus iuliformis (Carl). There may be other syntypes in the NMBE. Scolodesmus volzi volzi (Carl, 1913), Paradoxosomatidae japonicus Carl, 1902: 614-616, pl. 11, figs 38-39 [Polydesmus]. Japan (Genfer Museum). Two &male;. The MHNG collection contains parts of at least six broken specimens (MHNG-ARTO-14542). There is a discrepancy between the original description, which mentions &female; characters, and the list of specimens (two &male;) which follow it. The type series is restricted to the two &male; which are presumably amongst these specimens. The original identification labels have “ Japon &male; &female; ” written on them and there is a separate pencil label reading “ Japon ”. There is also an undated typewritten label indicating that Hoffman identified the specimens as Epanerchodus japonicus (Carl). Polydesmus japonicus Carl, 1902 is a junior homonym of P. japonicum Peters, 1864. The replacement name Polydesmus carli is proposed here for P. japonicus Carl, 1902., Published as part of Hollier, John, Stöckli, Eduard, Wesener, Thomas, Mesibov, Robert, Decker, Peter & Sierwald, Petra, 2020, An annotated list of the millipede (Diplopoda) species described by Johann Carl, pp. 183-240 in Revue suisse de Zoologie 127 (1) on page 209, DOI: 10.35929/RSZ.0015, http://zenodo.org/record/5743454, {"references":["Carl J. 1905. Diplopodes de la Guinee espagnole. Memorias de la Sociedad Espanola de Historia Natural 1 (15): 261 - 284.","Carl J. 1913 a. Westafrikanische Diplopoda. Revue suisse de zoologie 21: 201 - 224.","Carl J. 1902. Exotische Polydesmiden. Revue suisse de Zoologie 10: 563 - 679.","Peters W. C. H. 1864. Ubersicht der im Koniglich zoologischen Museum befindlichen Myriopoden aus der Familie der Polydesmi, sowie Beschreibungen einer neuen Gattung, Trachyjulus, der Juli und neuer Arten der Gattung Siphonophora. Monatsberichte der Koniglich Preussischen Akademie der Wissenschaften zu Berlin 1864: 529 - 551."]}

velutinus xanthopleurus Carl, 1909a: 249-250 [Sphaeropoeus]. Sumatra (Coll. W. Morton und Museum Bern). Unspecified series. The MHNG collection contains one specimen in alcohol (MHNG-ARTO.18449). The specimen is accompanied by a vial containing gonopods. The identification labels in the jar do not give any locality data, but the specimen is probably a syntype. There are two specimens collected by Morton in Sumatra in the MCZL which are syntypes, and presumably at least one other syntype in the NMBE. A junior synonym of Sphaeropoeus velutinus Carl, 1906, Zephroniidae, Published as part of Hollier, John, Stöckli, Eduard, Wesener, Thomas, Mesibov, Robert, Decker, Peter & Sierwald, Petra, 2020, An annotated list of the millipede (Diplopoda) species described by Johann Carl, pp. 183-240 in Revue suisse de Zoologie 127 (1) on page 235, DOI: 10.35929/RSZ.0015, http://zenodo.org/record/5743454, {"references":["Carl J. 1909 a. Neue Diplopoden. Revue suisse de Zoologie 17: 249 - 71.","Carl J. 1906. Diplopoden aus dem Malayischen Archipel. Zoologische Jahrbucher Abteilung fur Systematik, Geographie und Biologie der Tiere 24: 227 - 240."]}

pilosum Carl, 1912c:114-116,textfig.5[Castanotherium]. Bontorio und Umgebung; Bowanglangi, 1200-1500 m. Süd-Celebes (coll. Sarasin). Three &female;. No specimens found in the MHNG. The three syntypes (three &female; referred to as “ Typen ” in the NMB catalogue) are in the NMB (inventory numbers NMB-DIPL-00116a and NMB-DIPL-00116b). Castanotherium pilosum Carl, 1912, Zephroniidae, Published as part of Hollier, John, Stöckli, Eduard, Wesener, Thomas, Mesibov, Robert, Decker, Peter & Sierwald, Petra, 2020, An annotated list of the millipede (Diplopoda) species described by Johann Carl, pp. 183-240 in Revue suisse de Zoologie 127 (1) on page 223, DOI: 10.35929/RSZ.0015, http://zenodo.org/record/5743454, {"references":["Carl J. 1912 c. Die Diplopoden-Fauna von Celebes. Revue suisse de Zoologie 20: 73 - 206."]}

ornatum Carl, 1912c: 116-118, pl. 6, figs 32-33, 41, text fig. 6 [Castanotherium]. Bontorio und Umgebung, Süd-Celebes (coll. Sarasin). Unspecified number of &male; and &female;. The MHNG collection contains two specimens in alcohol (MHNG-ARTO-14303). One specimen has a pin running most of the length of the body. The identification labels have “ Celebes coll. Sarasin” and “ types Celebes coll. Sarasin” written on them respectively, indicating that the specimens are syntypes. There are three syntypes (referred to as “ Typen ” in the NMB catalogue) in the NMB (inventory number NMB-DIPL-00117a). Castanotherium ornatum Carl, 1912, Zephroniidae, Published as part of Hollier, John, Stöckli, Eduard, Wesener, Thomas, Mesibov, Robert, Decker, Peter & Sierwald, Petra, 2020, An annotated list of the millipede (Diplopoda) species described by Johann Carl, pp. 183-240 in Revue suisse de Zoologie 127 (1) on page 221, DOI: 10.35929/RSZ.0015, http://zenodo.org/record/5743454, {"references":["Carl J. 1912 c. Die Diplopoden-Fauna von Celebes. Revue suisse de Zoologie 20: 73 - 206."]}

simplex Carl, 1909a: 250-252, fig. 21 [Sphaeropoeus (Castanotherium)]. Java (Coll. L. Zehntner). Unspecified number of &male; and &female;. The MHNG collection contains five specimens in alcohol (MHNG-ARTO-18446). The specimens are accompanied by a vial containing gonopods. The two largest specimens have a pin running the length of the body. The identification labels in the jar have “ Java Zehntner” and “ Java, Dr L Zehntner” written on them respectively, indicating that the specimens are syntypes. Castanotherium simplex (Carl, 1909), Zephroniidae, Published as part of Hollier, John, Stöckli, Eduard, Wesener, Thomas, Mesibov, Robert, Decker, Peter & Sierwald, Petra, 2020, An annotated list of the millipede (Diplopoda) species described by Johann Carl, pp. 183-240 in Revue suisse de Zoologie 127 (1) on page 228, DOI: 10.35929/RSZ.0015, http://zenodo.org/record/5743454, {"references":["Carl J. 1909 a. Neue Diplopoden. Revue suisse de Zoologie 17: 249 - 71."]}

suspectum Carl, 1912c: 109 [Castanotherium]. Mapane am Golf von Tomini, Central-Celebes (coll. Sarasin). One damaged &female;. No specimens found in the MHNG. The &female; holotype is in the NMB (inventory number NMB-DIPL-00113a). Castanotherium suspectum Carl, 1912, Zephroniidae, Published as part of Hollier, John, Stöckli, Eduard, Wesener, Thomas, Mesibov, Robert, Decker, Peter & Sierwald, Petra, 2020, An annotated list of the millipede (Diplopoda) species described by Johann Carl, pp. 183-240 in Revue suisse de Zoologie 127 (1) on page 232, DOI: 10.35929/RSZ.0015, http://zenodo.org/record/5743454, {"references":["Carl J. 1912 c. Die Diplopoden-Fauna von Celebes. Revue suisse de Zoologie 20: 73 - 206."]}

boetonense Carl, 1912c: 119-120, text fig. 8 [Castanotherium]. Insel Boëton, südöstl. von Celebes (coll. Elbert). One &female;. No specimens found in the MHNG. The holotype was expected to be in the SMF but could not be found. Castanotherium boetonense Carl, 1912, Zephroniidae, Published as part of Hollier, John, Stöckli, Eduard, Wesener, Thomas, Mesibov, Robert, Decker, Peter & Sierwald, Petra, 2020, An annotated list of the millipede (Diplopoda) species described by Johann Carl, pp. 183-240 in Revue suisse de Zoologie 127 (1) on page 190, DOI: 10.35929/RSZ.0015, http://zenodo.org/record/5743454, {"references":["Carl J. 1912 c. Die Diplopoden-Fauna von Celebes. Revue suisse de Zoologie 20: 73 - 206."]}