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A new species of giant pill-millipede is described and illustrated from Guizhou Provinces, southern China: Sphaerobelum huzhengkuni n. sp. This is the first record of the genus Sphaerobelum in Chin...

Three new species of the giant pill-millipede family Zephroniidae are described from southern Vietnam: Sphaerobelum pumatensesp. nov., Sphaeropoeus honbaensissp. nov. and Sphaeropoeus bidoupensissp. nov. Two species, Sphaerobelum bicorne Attems, 1938 and Sphaeropoeus maculatus (Verhoeff, 1924), are redescribed, the former from new material, the latter from type material with lectotype designation. A new transfer is proposed: Zephronia manca Attems, 1936, to the genus Sphaeropoeus Brandt, 1833, giving the new combination, Sphaeropoeus manca (Attems, 1936) comb. nov.

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 ♂) Holotype THAILAND: 1 ♂ (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

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"]}

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 • ♂; 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 ♀; 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 ⅔ 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 ⅓ 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."]}

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 • ♂; 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 ♀; 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 ⅔ 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 ⅓ 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.

Two new species of giant pill-millipedes, Zephronia viridisoma Rosenmejer & Wesener sp. nov. and Sphaerobelum aesculus Rosenmejer & Wesener sp. nov., are described based on museum samples from southern Thailand. Zephronia viridisoma sp. nov. comes from Khao Lak, while the type locality of S. aesculus sp. nov. is on Phuket Island. Both species are described integratively, combining light microscopy, scanning electron microscopy, multi-layer photography, micro-CT scans and genetic barcoding. Genetic barcoding was successfully conducted for holotypes of both new species, which could be added to a dataset of all published sequences of the family Zephroniidae, including all described species from Thailand, Laos and Cambodia up to 2020. Genetic barcoding of the COI gene revealed another female of S. aesculus sp. nov., 160 km east of the type locality. Both new species are genetically distant from all other Zephroniidae from Thailand and surrounding countries, showing uncorrected p-distances of 16.8–23.1%. A virtual cybertype of a paratype of Z. viridisoma sp. nov. was created and made publically accessible.

Two giant pill millipedes, Sphaerobelum clavigerum Verhoeff, 1924 and Sphaerobelum hirsutum Verhoeff, 1924, were described from unindicated localities in northern Vietnam. The paper presents the recorded localities, descriptions and colour illustrations of those giant pill millipedes. S. clavigerum Verhoeff, 1924 is recorded from Cuc Phuong National Park (Ninh Binh Province), and distinguished from its congeners by body being shiny, glabrous, anterior telopods 3-segmented, process of 2nd podonomere of posterior telopods slender, not apically strongly swollen. In contrast, S. hirsutum Verhoeff, 1924 is found in Tam Dao National Park and Me Linh Biodiversity Station (both in Vinh Phuc Province), and recognized by body being densely hairy, process of 2nd podonomere of posterior telopods apically strongly swollen, spherical, wider than basis of 3rd podomere. In addition, a key to genera of giant pill millipedes is also provided for the Vietnam fauna.

Sphaerobelum lachneeis new species Figures 6E, 17, 18 Material examined: Type specimens. 1 M holotype (MHNG LT-10/12) from Laos, Oudomxai Province, ca 3 km E of Tad Lak 11, SE of Oudomxai city (= Muang Xai), (N20°36'20", 102°01'16" E), 1020 m, primary forest, leg. 18.IX.2010, P. Schwendinger. 1 F paratype (MHNG LT-10/12), same data as holotype; 1 M paratype (ZFMK MYR8099), same data as holotype. Diagnosis: S. lachneeis n. sp. belongs to a group of Sphaerobelum species in which the mesal margin of the femur is unmodified (Fig. 17B). S. lachneeis n. sp. shares only with S. separatum, S. truncatum and S. peterjaegeri n. sp. the absence of a "hook" at telopoditomere 4 (of the posterior telopods Figs 18C, D). S. lachneeis n. sp. differs from all those species in the presence of an only weakly separated telopoditomere 3 and 4 of the anterior telopods (Figs 18A, B), very long setae at the tergites (Fig. 6E), and a very long mesal extension at the apex of the immovable finger of the posterior telopods (Fig. 18C). Description. Based on holotype and both paratypes. Measurements: Body length: holotype male: length ca 23.5 mm. Width, of thoracic shield = 11.8 mm, of tergite 8 = 12.4 mm (= broadest). Height, of thoracic shield = 6.8 mm (= highest). Female: length ca 26.9 mm. Width, of thoracic shield = 13.3 mm, of tergite 7 = 13.9 mm (=broadest). Height, of thoracic shield = 7.5 mm, of tergite 8 = 8.5mm (= highest). Coloration: in preserved specimens dark olive-green, paratergite depressions and groove of thoracic shield lighter green, as ventral side (Fig. 6E). Antennae green, legs brown- olive greenish. Head: densely setose with very long setae (setae as long as width of tergite). Eyes with ca. 50 ocelli. Aberrant ocellus located inside antennal groove. Antennae short, with rounded joints, extending posteriorly to leg-pair 3. Lengths of antennomeres: 1>2=3=4=5

Sphaerobelum bolavensis new species Figures 6A, 7, 8, 9A, 10. Material examined: Type specimens. 1 M holotype (ZMUC 00040256) from Laos, Champasak Province, Bolaven Plateau, Ban Houayteuay, 1200 m (N 15°4.655', E 106°16.848') disturbed montane forest, leg. 10.XII.2007, by hand, S. Tarasov. 1 F paratype (ZFMK MYR8100), same data as holotype. Non-type material: 1 F (MHNG LT-10/24) from Laos, Champasak Province, Bolaven Plateau, 3 km S of Ban Nong Luang, Tad Kameud, 1270 m (N 15°04'23", 106°12'36" E), undisturbed evergreen hill forest, 1.X.2010, leg. P. Schwendinger. Diagnosis: S. bolavensis n. sp. belongs to a group of Sphaerobelum species in which the mesal margin of the femur is extended and apically tipped with several teeth (Fig. 8A). Sphaerobelum bolavensis n. sp. and S. bicorne from Vietnam are the only species with an extension only in the apical part of the femur. Sphaerobelum bolavensis n. sp. possesses a female vulval operculum only projecting in a single mesal process (Fig. 8B); in contrast, S. bicorne has two processes. Sphaerobelum bolavensis n. sp. has a telopoditomere 4 of the posterior telopod straight, apically without a recessed hook (Figs 10E, F); in contrast S. bicorne possesses a hook. Sphaerobelum bolavensis n. sp. has a short locking carina on the anal shield; whereas the carina is long in S. bicorne. Description. Based on holotype male and paratype female. Measurements: Holotype male. Body length ca 32.9 mm. Width, of thoracic shield = 14.2 mm, of tergite 8 = 14.9 mm (= broadest). Height, of thoracic shield = 7.1 mm, of tergite 7 = 8.2 mm (= highest). Female: body length = ca 28.8 mm. Width, of thoracic shield = 12.6 mm, of tergite 8 = 13.6 mm (=broadest). Height, of thoracic shield = 7.6 mm, of tergite 7, 8.3 mm (= highest). Coloration: in preserved specimens dark green, tergites with black posterior margin. Head and collum chestnut brown. Antennae orange, legs brown (Fig. 6A). Head: Eyes with>70 ocelli. Aberrant ocellus located inside antennal groove. Antennae short, with rounded joints, extending posteriorly to leg-pair 3. First antennomere with cuticular scales (Fig. 7A). Lengths of antennomeres: 1>2=3=4=5

Sphaerobelum splendidum new species Figures 6H, 9D, 22. Material examined: 1 M holotype (SMF) from Laos, Bolikhamsay Province, Lak Sao (1) (N18°11'09.0", 104°53'55.0"E), 500 m; cave entrance, rocks, vegetation, leg. 8.XI.2009, at day, by hand, P. Jäger & S. Bayer. Diagnosis: S. splendidum n. sp. belongs to a group of Sphaerobelum species in which the mesal margin of the femur is unmodified (Fig. 22B). S. splendidum n. sp. shares only with S. hirsutum, S. clavigerum, S. nigrum n. sp. and S. schwendingeri n. sp. the presence of a "hook" at telopoditomere 4 of the posterior telopods (Figs 22G, H). S. splendidum n. sp. differs from S. hirsutum in the presence of glabrous tergites, from S. clavigerum in the presence of a short locking carina on the anal shield. Differs from S. nigrum n. sp. in the presence of a cylindrical antennomere 6, the presence of only 5–7 ventral spines on the tarsi (8–10 ventral spines in the aother species) (Fig. 22B), and the presence of a strongly curved telopoditomere 4 of the posterior telopod, all characters only shared with S. laoticum n. sp. and S. schwendingeri n. sp. S. splendidum n. sp. differs from both in the following characters: Anterior telopod, telopoditomeres 3 and 4 clearly separated (Figs 22 D–F) (separation only partly visible in S. laoticum n. sp. and S. schwendingeri n. sp.), antennae and legs brown (both red in S. laoticum n. sp. and S. schwendingeri n. sp.). Description: Measurements: Body length 21.4 mm. Width, of thoracic shield = 11.3 mm (= broadest). Height, of thoracic shield = 5.9 mm (= highest). Coloration: Head and collum chestnut brown. Tergites black olive green, posterior tergite margin light brown (Fig. 6H). Antennae and legs dirty brown. Head: with 40–50 ocelli. Antennae extending posteriorly to leg 5. Size of antennomeres 1=2=3=4=5Gnathochilarium: usual for the order, sensory cones located in clusters. Mandibles: not investigated. First stigmatic plate: small, well-rounded, inconspicuous (Fig. 22A). Laterotergites: 1 and 2 with a short, but sharp, spine-like projection. Collum: only short setae at margin. Thoracic shield: Shallow grooves with few setae, surface glabrous. Margin towards grooves with 2 anterior and 3 or 4 posterior keels. Tergites: shiny, polished, at high magnification chagrinated. Paratergite tips weakly projecting posteriorly (Fig. 6H). Tergite surface glabrous except for paratergite grooves. Endotergum (Fig. 9D): inner section without setae or spines. Single, sparse row of slightly elliptical large cuticular impressions, distance between impressions slightly larger than their diameter. Externally 2–3 rows of medium length marginal bristles, protruding at most up to tergite margin (Fig. 9D). Legs: leg-pair 1 with 2 ventral spines, leg pair 2 with 3, leg pair 3 with 4 or 5, leg pairs 4–21 with only 5–7. Apical spine present at tarsi 3–21. In leg 9, femur 1.6 times, tarsus 3.3 times longer than wide (Fig. 22B). Coxae 1 and 2 without a coxal process (Figs 22A, C). Anal shield: wellrounded, glabrous, underside with a single very small, almost dot-like, locking carina. Male sexual characters: Male gonopore large, covered by small, inconspicuous plate (Fig. 22C). Anterior telopods (Figs 22 D–F): telopoditomere 1 rectangular, as long as wide. Telopoditomere 2 large, as long as telopoditomeres 3 and 4 combined. Process of telopoditomere 2 located posteriorly and laterally, visible in anterior view. Process of telopoditomere 2 slender, projecting almost as high as apical part of telopoditomere 4, apical part well-rounded. Telopoditomere 3 as long as wide, almost twice as long as telopoditomere 4, inconspicuous. telopoditomere 3 and 4 fused in posterior view, divided by a suture in anterior and lateral views. Telopoditomere 4 well-rounded, tapering apically. Posterior aspect with one black sclerotized spot and 2 spines. All podomeres apically covered with long setae. Posterior telopods (Figs 22G, H): telopoditomere 1 elongated, twice as long as wide. Immovable finger (process of telopoditomere 2) as long as movable finger, consisting of telopoditomeres 3 and 4. Immovable finger with a characteristic, distally swollen apex, massively swollen and curved towards movable finger. Apically broader than base. Telopoditomere 3 elongated, towards immovable finger at midpoint with a triangular extension carrying two large spines. Telopoditomere 4 tapering, apically strongly curved towards immovable finger, with thinly projecting apex. At inner margin with a large spine at midpoint and 1 or 2 smaller spines apically. Entire telopoditomere 1 and 2 at both sides covered with long setae, except for the immovable finger. Telopoditomere 3 only baso-laterally with a few setae, remaining part as well as telopoditomere 4 glabrous. Derivatio nominis: splendidum, noun in apposition, based on the shiny appearance of this species., Published as part of Wesener, Thomas, 2019, First records of giant pill-millipedes from Laos (Diplopoda, Sphaerotheriida, Zephroniidae), pp. 201-248 in Zootaxa 4563 (2) on pages 232-233, DOI: 10.11646/zootaxa.4563.2.1, http://zenodo.org/record/2601072

Sphaerobelum laoticum new species Figures 6I, 23, 24. Material examined: Type material. 1 M holotype (SMF) from Laos, Vientiane Province, Vang Vieng, W. of Nam Song (N18°55'46.86", 102°20'56.82"E), 324 m, Tham Nam Or Khem, in cave, leg. by hand, 28.VII.2014, P. Jäger & S. Münnich. Other material examined: 1 imm. M (SMF) from Laos, Vientinae Province, Van Vieng, Ban Sok Say (N19°1'20.30", 102°26'54.11"E), 295 m, in vegetation, leg. by hand, P. Jäger & S. Packheiser, 23.VII.2018. 1 imm. F (SMF) from Laos, Vientinae Province, Van Vieng, N of Ban Phoxay, Tham Hoi and Tham Loup, in cave (N19°02.350', 102°25.423'E), 256 m, leg. by hand, P. Jäger & S. Packheiser, 23.VII.2018. Diagnosis: S. laoticum n. sp. belongs to a group of Sphaerobelum species in which the mesal margin of the femur is unmodified (Fig. 23B). S. laoticum n. sp. shares only with S. hirsutum, S. clavigerum, S. nigrum n. sp., S. splendidum n. sp. and S. schwendingeri n. sp. the presence of a "hook" at telopoditomere 4 of the posterior telopods (Figs 24E, F). S. laoticum n. sp. differs from S. hirsutum in the presence of glabrous tergites, from S. clavigerum in the presence of a short locking carina on the anal shield. It differs from S. nigrum n. sp. in the presence of a cylindrical antennomere 6, the presence of only 5–7 ventral spines on the tarsi (8–10 in the other species) (Fig. 23B), and the presence of a strongly curved telopoditomere 4 of the posterior telopod, all characters only shared with S. splendidum n. sp. and S. schwendingeri n. sp. For a differentiation of S. laoticum n. sp. with S. splendidum n. sp. see above. S. laoticum n. sp. differs from S. schwendingeri n. sp., which occurs almost in sympatry, in a genetic distance of 13.6% in the COI gene, the process of telopoditomere 2 of the anterior telopods projecting as high as basal 1/3 (3/ 4 in S. schwendingeri n. sp.) of telopoditomere 3+4 (Fig. 22F); anal shield with a single short locking carinae (absent in S. schwendingeri n. sp.); palpi of gnathochilarium, sensory cones located in single field (in several clusters in S. schwendingeri n. sp.). Description: Based on male holotype Measurements: body length 24.4 mm. Width, of thoracic shield = 12.7 mm (= widest). Height, of thoracic shield = 7.4 mm (= highest). Coloration: Head and collum black. Tergites black (fading to brown in longer preserved specimens), paratergite depressions olive green (Fig. 6I). Antennae and legs red. Head: with ca. 65 ocelli. Antennae extending posteriorly to leg 5. Size of antennomeres 1=2=3=4=5

Sphaerobelum schwendingeri new species Figure 25 Material examined: Type material. 1 M holotype (MHNG LT 10/03) from Laos;, Vientiane Province, trail to Tham Pou Kham, W. of Vang Vieng (N18°55'32", 102°23'58"E), 240 m, secondary forest on limestone hill, leg. 7.IX.2010, P. Schwendinger. 1 M paratype (MHNG LT 10 /03), same data as previous. 1 F paratype (SMF), from Laos, Vientiane Province, Vang Vieng, Tham Pou Kham, 260 m (N18°55.549', 102°23.734'E), outside cave, close to cave stream, at rocks, leg. by hand, 13.III.2007, P. Jäger & F. Steinmetz. Other material examined: 1 M, 1 imm. F (SMF) from Laos, Vientiane Province, Van Vieng, across Nam Song, Tham Chang, in or in front of cave (N18°54.550', 102°26.527'E), 270 m, leg. by hand, P. Jäger & S. Packheiser, 22.Vii.2018. 1 M (ZFMK MYR8103), same data as previous. Diagnosis: S. schwendingeri n. sp. belongs to a group of Sphaerobelum species in which the mesal margin of the femur is unmodified (Fig. 25B). S. schwendingeri n. sp. shares only with S. hirsutum, S. clavigerum, S. nigrum n. sp., S. splendidum n. sp. and S. laoticum n. sp. the presence of a "hook" at telopoditomere 4 of the posterior telopods(Figs 25F, G). S. schwendingeri n. sp. differs from S. hirsutum in the presence of glabrous tergites, from S. clavigerum in the presence of a short locking carina on the anal shield. It differs from S. nigrum n. sp. in the presence of a cylindrical antennomere 6, the presence of 8–10 ventral spines on the tarsi (Fig. 25B), and the presence of a strongly curved telopoditomere 4 of the posterior telopod, all characters only shared with S. splendidum n. sp. and S. laoticum n. sp. For a differentiation of S. schwendingeri n. sp. with S. splendidum n. sp. see above. For a differentiation to S. laoticum n. sp. see above. Description: based on male holotype and paratypes of both sexes. Measurements: holotype male: body length 27.3 mm. Width, of thoracic shield = 13.3 mm (= widest). Height, of thoracic shield = 7.4 mm high (= highest). Female: Body length: broken and rolled up, ca. 37 mm. Width, of thoracic shield = 18 mm (unclear of broadest, broken). Coloration: in preserved specimens tergites black, posterior third olive. Head and collum black, anterior half of head green. Antennae and legs orange-red. Head: Eyes with ca. 70 ocelli. Last antennomere thickened. Apical disc with 73/71 apical cones (male), or 36/ 37 (female). All antennomeres covered by long setae. Organ of Tömösváry located inside antennal groove. Gnathochilarium: structure typical of the order. Palpi sensory cones located in several clusters. Mandibles: not dissected. Stigmatic plates: first stigmatic plate widely rounded, apex well-rounded, slightly curved towards coxa 1 (Fig. 25A). Laterotergites: laterotergite 1 and 2 slightly elongated into a short, broad process. Collum: with glabrous surface, margins with few isolated setae. Thoracic shield: surface glabrous, but covered with small pits like tergites, setae only in grooves. Shallow grooves beset with numerous long setae, slope towards groove without keels. Tergites: glabrous, shiny. Anterior third with a few setae only visible when the specimen is rolled up, producing a golden sheen. Tips of paratergites of midbody tergites projecting posteriorly. Endotergum: inner section without any setae or spines. Middle area with a single row of sparse elliptical, cuticular impressions, distance between impressions larger than their diameter. Apically 3-4 dense rows of short marginal bristles, the longest just protruding up to half of length to tergal margin. Bristles not smooth, but with numerous small spicules. Anal shield: sexually dimorphic, in female slightly flattened, in male weakly bell-shaped. Glabrous. Underside without locking carinae. Legs: leg-pair 1 with 2 ventral spines, leg-pair 2 with 6, leg-pair 3 with 6. First two legpairs without an apical spine. Leg-pairs 4–21 with 6–8 ventral spines and one dorso-apical spine. In leg 9 femur 1.7 times, tarsus 3.8 times longer than wide (Fig. 25B). All podomeres densely setose. Coxa with a well rounded process. Coxa process absent at first leg (Fig. 25A) and sharp-edged at second (Fig. 25C). Female sexual characters: Vulva large, covering mesal 2/3 of coxa, extending mesally to anterior third of prefemur (Fig. 25C). Operculum rounded, central part recessed, mesal margin broadly projecting into a sharpedged triangle slightly as higher than remaining operculum. Subanal plate: not examined. Male sexual characters: gonopore large, covered by small, inconspicuous plate. Anterior telopods (Figs 25D, E): Consisting of three telopoditomere above syncoxite. Last telopoditomeres completely fused. Telopoditomere 1 rectangular, slightly longer than wide. Telopoditomere 2 large, as long as telopoditomere 3. Process of telopoditomere 2 located posteriorly, apex visible in anterior view. Process of telopoditomere 2 wide, projecting to length of telopoditomere 3, strongly curved, apically with a well-rounded tip. Telopoditomere 3 massive, cylindrical, straight, apically slightly tapering. Posterior side with three large spines. In anterior view all telopoditomeres covered by long setae, in posterior view telopoditomeres 2 and 3 only with marginal setae. Posterior telopods (Figs 25F, G): telopoditomere 1 rectangular, slightly longer than wide. Immovable finger (process of telopoditomere 2) shorter than movable finger, consisting of telopoditomeres 3 and 4. Immovable finger with a spine in anterior view, a characteristic, distally swollen apex, slightly swollen. Apically broader than base, at tip in anterior view with a few large, crenulated teeth. Telopoditomere 3 elongated, at midpoint with a triangular extension carrying a spine towards immovable finger. Telopoditomere 4 tapering, anterior half strongly curved 90° towards immovable finger, which it overlaps. At inner margin with two larger spines. Entire telopoditomere 1 at both sides covered with long setae. Telopoditomere 2 anterior side completely covered by setae except for tip of immovable finger, posterior side glabrous except for lateral margin. Telopoditomere 3 only basally in anterior view with setae, remaining part, as well as telopoditomere 4 glabrous. Derivatio nominis: schwendingeri, noun in apposition, after the Arachnologist Peter Schwendinger, great collector of arthropods in Southeast Asia., Published as part of Wesener, Thomas, 2019, First records of giant pill-millipedes from Laos (Diplopoda, Sphaerotheriida, Zephroniidae), pp. 201-248 in Zootaxa 4563 (2) on pages 238-239, DOI: 10.11646/zootaxa.4563.2.1, http://zenodo.org/record/2601072

Sphaerobelum nigrum new species Figures 6G, 20, 21 Material examined: Type specimens: 1 M holotype (SMF) from Laos, Champasak Province, Muang Bachieng, Ban Lak 35, Tad Etu (N15°11'37.7", 106°06'06.3"E), 810 m, coffee plantation with remnant of secondary forest, leg. in vegetation, at night, by hand; 16.XI.2012; P. Jäger. Paratype: 1 F (SMF) from Laos, Champasak Province, Muang Bachieng, Ban Lak 35, Tad Etu (N15°11'37.7", 106°06'06.3"E), 810 m, coffee plantation with remnant of secondary forest, close to river, at rock, leg. at day, by hand, 25.XI.2009, P. Jäger & S. Bayer. Diagnosis: S. nigrum n. sp. belongs to a group of Sphaerobelum species in which the mesal margin of the femur is modified (Fig. 20B). S. nigrum n. sp. shares only with S. hirsutum, S. clavigerum, S. splendidum n. sp. and S. schwendingeri n. sp. the presence of a "hook" at telopoditomere 4 of the posterior telopods(Figs 21E, F). S. nigrum n. sp. differs from S. hirsutum in the presence of glabrous tergites, from S. clavigerum in the presence of a short locking carina on the anal shield. Differs from S. splendidum n. sp. and S. schwendingeri sp. in the presence of an axe-shaped antennomere 6, and the presence of only a short hook at the telopoditomere 4 (Figs 21E, F) of the posterior telopods (strongly curved telopoditomere 4 in the other two species). Description. Based on holotype male and paratype female. Measurements: Body length: holotype male: length ca 27.8 mm. Width, of thoracic shield = 16.1 mm, of tergite 7 = 16.8 mm (= broadest). Height, of thoracic shield = 9.8 mm, of tergite 7 = 10.1 mm (= highest). Female: length ca 34.7 mm. Width, of thoracic shield = 15.7 mm, of tergite 7 = 17.4 mm (= broadest). Height of thoracic shield = 9.3 mm, of tergite 7 = 9.8 mm (= highest). Coloration: in preserved specimens yellow with dark brown spots or black spots/bands. Head and collum black (Fig. 6G). Antennae and legs dark brown to black. Head: Eyes with ca. 55 ocelli. Antennae short, with rounded joints, extending posteriorly to leg-pair 4. Lengths of antennomeres: 1=2=3=4=5

Sphaerobelum phouloei new species Figures 6B, 9B, 11, 12. Sphaerobelum sp. II 1 M Material examined: Type specimen. 1 M holotype (ZMUC 00040257) from Laos, Houaphan Province, Phou Loei, way from camp 1 to camp 2, ~ 1200 m, leg. 9.VIII.2008; S. Tarasov. Diagnosis: S. phouloei n. sp. belongs to a group of Sphaerobelum species in which the mesal margin of the femur is extended in several teeth (Fig. 11B). S. phouloei n. sp. shares only with S. denticulatum n. sp. a curved telopoditomere 4 of the posterior telopods which overlaps the immovable finger (Figs 12E, F). S. phouloei n. sp. differs in several characters from S. denticulatum n. sp.: male antenna with only 30–35 apical cones (> 50 in S. denticulatum n. sp.), prefemur mesal margin well-rounded (indentated in S. denticulatum n. sp.), tergites covered with short setae (glabrous in S. denticulatum n. sp.), legs orange (brown in S. denticulatum n. sp.). Description. Measurements: Body length: holotype male: length ca 25.8 mm. Width, of thoracic shield 13.3 mm, of tergite 8 = 14.1 mm (= broadest). Height, of thoracic shield = 8.9 mm, of tergite 8 = 9.3 mm (= highest). Coloration: in preserved specimens anterior 2/3 of tergites dark olive green, posterior 1/3 black. Head, ventral side, anal shield orange (Fig. 6B). Legs and antennae orange. Head: Eyes with 65–70 ocelli. Aberrant ocellus located inside antennal groove. Antennae short, with rounded joints, extending posteriorly to leg-pair 4. First antennomere with cuticular scales. Lengths of antennomeres: 1=2=3Gnathochilarium: structure typical of the order. Palpi sensory cones located in several clusters. Mandibles: not dissected. Stigmatic plates: first stigmatic plate widely rounded, apex well-rounded, curved towards coxa 1 (Fig. 11A). Laterotergites: laterotergite 1 and 2 with a broad, stout projection. Collum: with glabrous surface, margins with few isolated setae. Thoracic shield: surface glabrous unlike tergites, setae only in grooves. Shallow grooves beset with numerous long setae, slope towards groove with 3 weak anterior and 2 or 3 posterior keels. Tergites: anterior 2/3 of surface densely setose with short setae, posterior 1/3 glabrous. Tips of paratergites of midbody tergites projecting posteriorly (Fig. 6B). Endotergum (Fig. 9B): inner section lacking any spines or setae. Middle area with a single row of large, sparse, elliptical, cuticular impressions. Distance between impressions as wide as their diameter. Apically, 1–2 sparse rows of long marginal bristles, strongly protruding beyond tergal margin. Bristles not smooth, but with numerous small spinicles. Anal shield: large, surface completely covered by tiny setae. Underside with a single, long, black locking carina, located close to last laterotergite. Legs: leg-pair 1 with 2 ventral spines, leg-pair 2 with 5, leg-pair 3 with 6 or 7. First two leg-pairs without an apical spine. Leg pairs 4–21 with 9 or 10 ventral spines and one dorso-apical spine. In leg 9, femur 1.3 times, tarsus 3.7 times longer than wide (Fig. 11B). All podomeres densely setose. Coxa with a large and marginally toothed process. Coxa process absent at first leg (Fig. 11B) and sharply projecting at second (Fig. 12A). Prefemur at apical margin with a projection laterally and mesally. Lateral projection triangular and sharply edged, juxtaposed to coxal process. Femur extended mesally into a dentate margin featuring 12–14 teeth. Female unknown. Male sexual characters: gonopore covered with a single, undivided, circular, sclerotized plate (Fig. 12A). Anterior telopods (Figs 12 B–D): consisting of 4 telopoditomeres above syncoxite, but telopoditomere 3 and 4 only divided by thin suture. Telopoditomere 1 rectangular, as long as wide. Telopoditomere 2 large, almost as long as telopoditomere 3 and 4 combined. Process of telopoditomere 2 located posteriorly, not visible in anterior view. Process of telopoditomere 2 wide, projecting to half of telopoditomere 4, apically with a well-rounded tip. Telopoditomere 3 cylindrical, straight, as long as telopoditomere 4. Telopoditomere 4 cylindrical, well-rounded, posterior side with a black sclerotized spot and a small, triangular spine. Telopoditomeres 1–4 in anterior view covered with long setae. In posterior view all telopoditomeres with fewer setae. Posterior telopods (Figs 12E, F): telopoditomere 1 elongated, twice as long as wide. Immovable finger (process of telopoditomere 2) shorter than movable finger, consisting of telopoditomeres 3 and 4. Immovable finger with a characteristic, distally swollen apex, well rounded, apex therefore wider than base. Immovable finger in anterior view with a large spine, at tip opposite to tip of telopoditomere 4 with sclerotized spots. Telopoditomere 3 rectangular, towards immoveable finger with a triangular expansion carrying a spine. Telopoditomere 4 reaching 2/3 of length of telopoditomere 3, apically strongly tapering into acute tip, very slightly curved towards immovable finger. Inner margin with single spine towards immovable finger. Telopoditomere 1 at both sides covered by setae. Telopoditomeres 2 and 3 in anterior view covered by long setae, in posterior view mostly glabrous. Telopoditomere 4 only with marginal setae. Derivatio nominis: phouloei, noun in apposition, after the Phou Loei Mountain, the type locality (Fig. 5)., Published as part of Wesener, Thomas, 2019, First records of giant pill-millipedes from Laos (Diplopoda, Sphaerotheriida, Zephroniidae), pp. 201-248 in Zootaxa 4563 (2) on pages 215-216, DOI: 10.11646/zootaxa.4563.2.1, http://zenodo.org/record/2601072

Sphaerobelum denticulatum new species Figures 6C, 13 Material examined: 1 M holotype (MHNG LT-10/12) from Laos, Oudomxai Province, ca 3 km E of Tad Lak 11, SE of Oudomxai city (= Muang Xai) (N20°36'20"; 102°01'16" E), 1020 m, undisturbed evergreen hill forest, leg. 18.IX.2010, P. Schwendinger. Diagnosis: S. denticulatum n. sp. belongs to a group of Sphaerobelum species in which the mesal margin of the femur is extended with several teeth (Fig. 13A). S. denticulatum n. sp. shares only with S. phouloei n. sp. a curved telopoditomere 4 of the posterior telopods which overlaps the immovable finger (Figs 13D, E). See above for a separation of these two species. Description. Measurements: Body length: length ca 22.7 mm. Width, of thoracic shield = 11.5 mm (= broadest). Height, of thoracic shield = 6.3 mm, tergite 7 = 6.9 mm (= highest). Coloration: in preserved specimens tergites black, thoracic shield grooves and paratergite depressions dark green. Head and collum black (Fig. 6C). Antennae dirty green, antennomere 6 yellow. Legs light brown, tarsus darker. Head: Eyes with ca. 55 ocelli. Antennae short, extending back to leg 3, with rounded joints. Lengths of antennomeres: 1>2=3=4=5

Sphaerobelum spinatum new species Figures 6D, 9C, 14, 15, 16 Material examined: Type specimens. 1 M holotype (ZMUC 00040258) from Laos, Vientiane Province, Phou Khao Khouay (N 18°20.369' N, 102°48.523'E), 700–800 m, strongly disturbed primary evergreen forest, leg. 26- 31.V.2008, A. Solodovnikov & J. Pedersen; 1 F paratype; ZMUC00040259; same data as paratype; 1 M, 1 F paratype (ZFMK MYR8101 & MYR8102), same data as holotype. Diagnosis: S. spinatum n. sp. belongs to a group of Sphaerobelum species in which the mesal margin of the femur is extended with several teeth (Fig. 15B). S. spinatum n. sp. shares only with S. cattiense and S. konkakinhense a straight telopoditomere 4 (Figs 16E, F). S. spinatum n. sp. differs in several characters from both other species: mesal margin of prefemur with indentations like the femur (Fig. 15B), male antenna with 70 in S. denticulatum n. sp.), tergites shiny, but setose (tergites covered with setose pits in the other species), first laterotergite strongly projecting, extended into a sharp tip (first laterotergite only weakly projecting in the other two species). Description. Based on holotype and both paratypes. Measurements: Body length: holotype male: length ca 26.9 mm. Width, of thoracic shield = 13.6 mm (= broadest). Height, of thoracic shield = 7.5 mm (= highest). Female: length ca 32.5 mm. Width, of thoracic shield = 14.5 mm, of tergite 7 = 15.7 mm (=broadest). Height, of thoracic shield = 7.7 mm, of tergite 9 = 9.1mm (= highest). Coloration: in preserved specimens black, paratergite depressions and groove of thoracic shield dark green (Fig. 6D). Antennae brown-orange, legs light brown, but prefemur and femur green. Head: Eyes with>60 ocelli. Aberrant ocellus located inside antennal groove. Antennae short, with rounded joints, extending posteriorly to leg-pair 5. Lengths of antennomeres: 1>2=3=4=5

Sphaerobelum peterjaegeri new species Figures 6F, 19 Material examined: 1 M, holotype (SMF SD553) from Laos, Luang Prabang Province, SE Luang Prabang, Nam Khan, Ban Pak Bak, Houay Kho, 328 m, (N19°44'09.1", 102°16'37.5"E), bank of stream, under stone, moist sand and gravel, leg. 28.II.2008, P. Jäger. Diagnosis: S. peterjaegeri n. sp. belongs to a group of Sphaerobelum species in which the mesal margin of the femur is unmodified (Fig. 19B). S. peterjaegeri n. sp. shares only with S. separatum, S. truncatum and S. lachneeis n. sp. the absence of a "hook" at telopoditomere 4 of the posterior telopods (Figs 19G, H). S. peterjaegeri n. sp. differs from S. separatum in the absence of a lateral extension on podomere 3 of the anterior telopod, as well as the absence of an incurved process on telopoditomere 4 of the posterior telopods. S. peterjaegeri n. sp. is very similar to S. truncatum, but differs in several characters: 80 in S. truncatum), antennae and legs orange-red (yellow in S. truncatum), femur 1.7 times longer than wide (2.0 times longer than wide in S. truncatum). Description. Measurements: Body length: holotype male: length ca 23.8 mm. Width of thoracic shield = 11.6 mm (= broadest). Height of thoracic shield = 6.9 mm (= highest). Coloration: in preserved specimens brown to orange (Fig. 6F). Collum dark olive brown, same as head. Antennae and legs orange. Head: Eyes with ca. 60 ocelli. Aberrant ocellus located inside antennal groove. Antennae short, with rounded joints, extending posteriorly to leg-pair 3. First antennomere with cuticular scales. Lengths of antennomeres: 1=2=3=4=5

Key to species of Sphaerobelum (based mainly on male characters): 1. Femur of legs at mesal margin extended, with teeth (Fig. 13A). Apical margin of prefemur modified, extended into a process..................................................................................................... 2 - Femur of legs not extended, well-rounded (Fig. 17B). Prefemur unmodified........................................ 9 2. Teeth on femur only at apex: 4–6 (Fig. 8A). Anterior telopod consisting of 3 podomeres............................. 3 - Teeth on whole mesal margin of femur:>10. Anterior telopod always consisting of 4 telopoditomeres................... 4 3. Female vulval operculum strongly projecting in two processes. Posterior telopod with telopoditomere 4 tip curved towards immovable finger like a small hook..................................................... S. bicorne Attems, 1938 - Female vulva projecting in a mesal process (Fig. 8B). Posterior telopod: telopoditomere 4 straight, not hooked (Fig. 10E)......................................................................................... S. bolavensis n. sp. 4. Posterior telopod: telopoditomere 4 apex weakly curved towards and overlapping immovable finger. Anterior side of immovable finger with sclerotized spots opposite to tip of telopoditomere 4 (Fig. 12E). Immovable finger in anterior view with a large sclerotized spine...................................................................................... 5 - Posterior telopod: telopoditomere 4 straight, not curved. No sclerotized spots or a spine at immovable finger (Fig. 16E)......................................................................................................... 6 5. Male antennae with 30–35 apical cones. Tergites covered with short setae. Legs orange. Prefemur mesal margin rounded (Fig. 11B)................................................................................... S. phouloei n. sp. - Male antennae with 65–70 apical cones. Tergites glabrous. Legs light brown. Prefemur mesal margin with indentations like the femur (Fig. 13A)..................................................................... S. denticulatum n. sp. 6. Male antennae red or brown, with>70 apical cones. Tergites matte. First laterotergite weakly projecting posterior-laterally....................................................................................................... 7 - Male antennae yellow, with 50–55 apical cones. Tergites shiny, with setae. First laterotergite strongly projecting posterior-laterally................................................................................. S. spinatum n. sp. 7. Tergites glabrous, matte. Endotergum bristles only protruding slightly midway towards margin. Anterior telopod: telopoditomeres 3 and 4 not well-separated in posterior view (Figs 21 B–D)..................................... S. nigrum n. sp. - Tergites hairy, with pits. Endotergum bristles long, protruding above tergite margin. Anterior telopod: telopoditomeres 3 and 4 well separated........................................................................................ 8 8. Colour dark olive, posterior tergal margins with a broad, light-brown band, antennae and legs conspicuously orange. Prefemur without lateral projection. Apical margins of vulval operculum equally converging towards a mesal tip. Tip of laterotergite 1 slightly extended, well-rounded. Male antennomere 6 thickened...................... S. cattiense Semenyuk et al., 2018 - Colour dark chestnut brown, posterior tergal margins black, legs dark brown, tips light brown. Lateral margin of prefemur with a sharp projection juxtaposed to coxal process. Apical margin of vulval operculum recessed centrally. Tip of laterotergite 1 slightly extended, sharp. Male antennomere 6 axe-shaped....................... S. konkakinhense Semenyuk et al., 2018 9. Posterior telopod without 'hook'......................................................................... 10 - Posterior telopod: telopoditomere 4 apically always with a short recurved 'hook' (Fig. 21E)........................... 13 10. Tergites covered with very long setae. Setae longer than width of tergites (Fig. 6E). Immovable finger of posterior telopod with very long mesal extension (Fig. 18C). Antennae green. Anterior telopod suture between telopoditomeres 3 and 4 weak, not visible in anterior view (Figs 18A, B).......................................................... S. lachneeis n. sp. - Tergites covered with short setae. Immovable finger of posterior telopod swollen, but not with such a long lateral extension. Anterior telopod: telopoditomeres 3 and 4 well separated...................................................... 11 11. Podomere 3 of anterior telopod distally with a lateral extension. Posterior telopod: telopoditomere 4 distally with an incurved process overlapping with process of podomere 2. Marginal setae of terga not reaching posterior margin. Tergite posterior margin well-rounded, not tesselated..................................................... S. separatum Attems, 1953 - Podomere 3 of anterior telopod cylindrical, without lateral extension. Posterior telopod: telopoditomere 4 without incurved process, but with 3 separate spines. Marginal setae of terga reaching posterior margin. Tergite posterior margin tesselated...................................................................................................... 12. 12. Antennae with>80 apical cones. Legs: femur 2x longer than wide. Tarsi with 7 or 8 ventral spines. Antennae and legs yellow. Vulval operculum well-rounded, not extending............................. S. truncatum Wongthamwanich et al., 2012 - Antennae with S. peterjaegeri n. sp. 13. Tergites glabrous, at least posterior 2/3 of each tergite....................................................... 14. - Tergites covered with short setae. Dirty olive-brown. Head punctured and densely hirsute........ S. hirsutum Verhoeff, 1924 14. Anal shield with a short, or completely reduced locking carina. Posterior telopod: immovable finger, swollen at inner and mesal margin. Telopoditomere 4 at mid-point strongly curved and overlapping immovable finger (Fig. 22G)............ 15 - Anal shield with long locking carina,> 3X length of tergite carina. Posterior telopod: immovable finger, swollen only at inner margin; mesal margin straight..................................................... S. clavigerum Verhoeff, 1924 15. Anterior telopod: telopoditomeres 3 and 4 clearly separated (Figs 21 B–D). Antennae and legs brown......................................................................................................... S. splendidum n. sp. - Anterior telopod: telopoditomeres 3 and 4 separation only partly visible in anterior view (Fig. 22D). Antennae and legs red..................................................................................................... 16 16. Process of telopoditomere 2 projecting as high as basal 1/3 of telopoditomere 3+4 (Fig. 22F). Anal shield with a single short locking carinae. Palpi of gnathochilarium, sensory cones located in single field........................ S. laoticum n. sp. - Process of telopoditomere 2 projecting as high as basal 3/4 of telopoditomere 3+4 (Fig. 25E). Anal shield without locking carinae. Palpi of gnathochilarium, sensory cones located in several clusters........................ S. schwendingeri n. sp., Published as part of Wesener, Thomas, 2019, First records of giant pill-millipedes from Laos (Diplopoda, Sphaerotheriida, Zephroniidae), pp. 201-248 in Zootaxa 4563 (2) on page 211, DOI: 10.11646/zootaxa.4563.2.1, http://zenodo.org/record/2601072, {"references":["Attems, C. (1938) Die von Dr. C. Dawydoff in franzosisch Indochina gesammelten Myriopoden. Memoires du Museum National d'Histoire Naturelle, N. S. 6 (2), 187 - 353.","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","Attems, C. (1953) Myriopoden von Indochina, Expedition von Dr. C. Dawydoff (1938 - 1939). Memoires du Museum national d'histoire naturelle, N. S., serie A, 5 (3), 133 - 230.","Wongthamwanich, N., Panha, S., Sierwald, P., Wesener, T. & Thirakhupt, K. (2012) A new species of the giant pill-millipede genus Sphaerobelum Verhoeff, 1924 from northern Thailand, with an extensive description and molecular characters (Diplopoda: Sphaerotheriida: Zephroniidae). Zootaxa, 3220, 29 - 43","Verhoeff, K. W. (1924) Results of Dr. E. Mjoberg's Swedish scientific expedition to Australia 1910 - 1913. 34. Myriapoda: Diplopoda. Arkif for Zoologi, 16 (5), 1 - 142, pls. 1 - 5."]}

Before this study, giant pill-millipedes (order Sphaerotheriida) were unknown from Laos despite their presence in all surrounding countries. As by-catch from collections by arachnologists, 31 specimens of Sphaerotheriida from Laos became available for study. The sample included 14 species. Three species were only represented by females, which are described but cannot be named. Of the remaining 11 species, a single species belongs to the genus Zephronia Gray, 1832: Z. laotica n. sp.; and the other ten belong to the genus Sphaerobelum Verhoeff, 1924: S. bolavensis n. sp., S. phouloei n. sp., S. denticulatum n. sp., S. spinatum n. sp., S. lachneeis n. sp., S. peterjaegeri n. sp., S. nigrum n. sp., S. splendidum n. sp., S. laoticum n. sp., and S. schwendingeri n. sp. This more than doubles the known diversity of Sphaerobelum. Here, I integratively describe these species, combining morphology and DNA barcodes with a molecular analysis including all Zephroniidae species deposited on GenBank—including the only giant pill-millipede species known from Cambodia, Zephronia dawydoffi Attems, 1953. An updated determination key to the species of the genus is presented. Zephronia laotica n. sp. belongs to the monophyletic Zephronia sensu stricto group, which is confirmed by molecular barcoding. In contrast, most species of Sphaerobelum are in a weakly supported clade. Genetically, Sphaerobelum species differ greatly from one another, with most p-distances >15%. The lowest observed p-distance (9.8%) is between S. truncatum Wongthamwanich et al. 2012 from Thailand and S. peterjaegeri n. sp.

Sphaerobelum cattiense Semenyuk, Golovatch and Wesener, sp. nov. Figs 1A, 2, 3, 4A Material examined: Type specimens. Holotype male (ZMUM), Vietnam, Dong Nai Province, Cat Tien National Park, monsoon lowland tropical forest (N 11°26' E 107°21', 100–200 m. a.s.l.), litter, floor and under logs, 22– 24.VI.2011, leg. I.I. Semenyuk. Paratypes: 1 males, 2 females (ZMUM), same data, together with holotype; 1 male, 1 female, 1 ex. of undetermined sex (ZFMK MYR 6516, 6519, 6565, same data as holotype, but collected in VI.2017). Other material. 2 males, 2 females, 1 immature (topotypes), ZFMK MYR 6566, Vietnam, Dong Nai Province, Cat Tien National Park, Lagerstroemia calyculata and Afzelia xylocarpa forest, VIII.2011, leg. E. Galoyan. Diagnosis. Medium-sized, 25–30 mm long and 15–17 mm wide, dark olive-brown Sphaerobelum with a microsetose surface and orange antennae and appendages. Modified femur with a dentate mesal margin (Fig. 2D), a character shared only with S. bicorne and S. konkakinhense sp. nov. Differing from both other congeners in the vulval operculum which lacks a recessed apical margin and projects in a single, well-rounded, mesal tip (Fig. 2I). Immovable finger of posterior telopod swollen apically, but extending only mesally (Figs 2E, F). Also differing from S. bicorne and S. konkakinhense sp. nov. in the absence of a lateral projection on the prefemur of the walking legs. Description. Measurements: Holotype male. Body length ca 30 mm, width of thoracic shield = 14.6 mm, of tergite 9 = 15.2 mm (= broadest), height of thoracic shield = 8.5 mm, of tergite 9 = 9.3 mm (= highest). Male paratypes ca 25 or 29 mm long, maximal width ca 17 or 15 mm, respectively. Females: length ca 27.5–30 mm, width of thoracic shield = 14.3–16 mm, of tergite 9 = 15.3–17 mm (=broadest), height of thoracic shield = 8.6–9.0 mm, of tergite 9 = 9.3–10 mm (= highest). Coloration: in live specimens, ventral side including head and all tergites blackish brown; collum, tergites and posterior part of anal shield brown. Thoracic shield brown at margins. Legs and antennae dark orange. In preserved specimens, dark olive to olive brown with contrasting, narrow, darkish, orange bands at caudal margins of all tergites (Fig. 1A). Head dark brown to dark olive-brown. Antennae and legs contrasting bright orange. Head: wide and short, partly micropunctate. Clypeolabral region very densely setose. Occipital region finely and confusedly striolate and with a row of longer hairs near caudal edge. Eyes with>65 ocelli. Aberrant ocellus located inside antennal groove. Antennae short, with rounded joints, extending posteriorly to leg-pair 2. First antennomere with cuticular scales (Fig. 3A). Lengths of antennomeres: 1>2=3=4=5

Four new species of giant pill-millipedes (Sphaerotheriida) are described from southern Vietnam: Sphaerobelum cattiense sp. nov., Sphaerobelum konkakinhense sp. nov., Zephronia konkakinhensis sp. nov. and Zephronia montis sp. nov. An updated generic diagnosis of Sphaerobelum Verhoeff, 1924 and a key to its species are provided.

Sphaerobelum cattiense Semenyuk, Golovatch and Wesener, sp. nov. Figs 1A, 2, 3, 4A Material examined: Type specimens. Holotype male (ZMUM), Vietnam, Dong Nai Province, Cat Tien National Park, monsoon lowland tropical forest (N 11°26' E 107°21', 100–200 m. a.s.l.), litter, floor and under logs, 22– 24.VI.2011, leg. I.I. Semenyuk. Paratypes: 1 males, 2 females (ZMUM), same data, together with holotype; 1 male, 1 female, 1 ex. of undetermined sex (ZFMK MYR 6516, 6519, 6565, same data as holotype, but collected in VI.2017). Other material. 2 males, 2 females, 1 immature (topotypes), ZFMK MYR 6566, Vietnam, Dong Nai Province, Cat Tien National Park, Lagerstroemia calyculata and Afzelia xylocarpa forest, VIII.2011, leg. E. Galoyan. Diagnosis. Medium-sized, 25–30 mm long and 15–17 mm wide, dark olive-brown Sphaerobelum with a microsetose surface and orange antennae and appendages. Modified femur with a dentate mesal margin (Fig. 2D), a character shared only with S. bicorne and S. konkakinhense sp. nov. Differing from both other congeners in the vulval operculum which lacks a recessed apical margin and projects in a single, well-rounded, mesal tip (Fig. 2I). Immovable finger of posterior telopod swollen apically, but extending only mesally (Figs 2E, F). Also differing from S. bicorne and S. konkakinhense sp. nov. in the absence of a lateral projection on the prefemur of the walking legs. Description. Measurements: Holotype male. Body length ca 30 mm, width of thoracic shield = 14.6 mm, of tergite 9 = 15.2 mm (= broadest), height of thoracic shield = 8.5 mm, of tergite 9 = 9.3 mm (= highest). Male paratypes ca 25 or 29 mm long, maximal width ca 17 or 15 mm, respectively. Females: length ca 27.5–30 mm, width of thoracic shield = 14.3–16 mm, of tergite 9 = 15.3–17 mm (=broadest), height of thoracic shield = 8.6–9.0 mm, of tergite 9 = 9.3–10 mm (= highest). Coloration: in live specimens, ventral side including head and all tergites blackish brown; collum, tergites and posterior part of anal shield brown. Thoracic shield brown at margins. Legs and antennae dark orange. In preserved specimens, dark olive to olive brown with contrasting, narrow, darkish, orange bands at caudal margins of all tergites (Fig. 1A). Head dark brown to dark olive-brown. Antennae and legs contrasting bright orange. Head: wide and short, partly micropunctate. Clypeolabral region very densely setose. Occipital region finely and confusedly striolate and with a row of longer hairs near caudal edge. Eyes with>65 ocelli. Aberrant ocellus located inside antennal groove. Antennae short, with rounded joints, extending posteriorly to leg-pair 2. First antennomere with cuticular scales (Fig. 3A). Lengths of antennomeres: 1>2=3=4=5

Sphaerobelum konkakinhense Semenyuk, Golovatch and Wesener, sp. nov. Figs 1B, 4B, 5, 6 Material examined: Type specimens. Holotype male (ZMUM), Vietnam, Gia Lai Province, Kon Ka Kinh National Park, near summit (N 14°12' E 108°18', 1400–1500 m. a.s.l.), mixed tropical cloud forest, slopes, litter and under logs, V.2017, leg. I.I. Semenyuk. Paratypes: 1 male, 1 female, ZFMK MYR 6611, 6612, same data as holotype; 1 female (ZMUM), same data as holotype. Diagnosis. Medium-sized, up to 30 mm long, chestnut Sphaerobelum with a setose surface and basally dark, apically light brown appendages. Modified femur with a dentate mesal margin, modified prefemur with a lateral process (Fig. 5D), female operculum at apical margin depressed centrally (Fig. 5H), all these characters shared only with S. bicorne. Sphaerobelum konkakinhense sp. nov. differs from S. bicorne in the presence of the anterior telopod with four telopoditomeres (vs. 3 in S. bicorne) and a much less strongly invaginated female operculum. Process of anterior telopod visible also laterally in anterior view, with a rectangular apex (Fig. 5B). Immovable finger of posterior telopod swollen apically, extending mesally and laterally (Figs 5F, G). Description. Measurements: Holotype male. Body length ca 30 mm, width of thoracic shield = 13.4 mm, of tergite 8 = 14.0 mm (= broadest), height of thoracic shield = 8 mm (= highest). Female (largest): length ca 28.6 mm, width of thoracic shield = 13.4 mm, of tergite 9 = 14.1 mm (= broadest), height of thoracic shield = 7.8 mm, of tergite 9 = 8.4 mm (= highest). Coloration: in live specimens, head and collum dark brown. Tergites paler dark brown with grey hue (Fig. 1B). Ventral surface of body clothed with short, yellowish, fur-like setae. Legs and antennae much paler, greyish, with yellowish setae. In preserved specimens, head dark brown. Antennae and legs dark brown, legs distal to postfemora, as well as antennomere 6 contrasting light brown. Head: wide and short. Eyes with>60 ocelli. Aberrant ocellus located inside antennal groove. Antennae short, with rounded joints, extending posteriorly to leg-pair 2. Antennomeres 1 and 2 with few sclerotized scales (Fig. 6A). Sizes of antennomeres 1>2=3=4=5

Key to species of Sphaerobelum (based mainly on male characters): 1. Anterior telopod with 3 telopoditomeres................................................................... 2 - Anterior telopod with 4 telopoditomeres.................................................................... 5 2. Tergites shiny, glabrous. Process of podomere 2 of posterior telopod swollen only at inner margin. Endotergum with 3–4 dense rows of short marginal bristles.................................................... S. clavigerum Verhoeff, 1924 - Tergites hairy, process of podomere 2 of posterior telopod strongly swollen apically, spherical, broader than base of podomere.................................................................................................... 3 3. Inner margin of femur extended, with teeth. Lateral margin of prefemur with a sharp projection juxtaposed to coxal process. Body coloration black. Head smooth. Process of podomere 2 of posterior telopods truncated. Operculum of vulva two-horned, both horns blunt.................................................................... S. bicorne Attems, 1938 - Inner margin of femur smooth, prefemur without a process. Body coloration dirty olive-brown; head punctured and densely hirsute......................................................................... S. hirsutum Verhoeff, 1924 4. Inner margin of femur extended, with teeth (Figs 2D, 5D). Locking carina on anal shield short......................... 6 - Inner margin of femur smooth. Locking carina on anal shield very long........................................... 7 5. Body coloration dark olive, posterior margin uniformly light brown, antennae and legs conspicuously orange (Fig. 1A). Prefemur without lateral projection (Fig. 2D). Apical margins of vulval operculum equally converging towards a mesal tip (Fig. 2I). Tip of laterotergite 1 slightly extended, well-rounded. Male antennomere 6 thickened (Fig. 3A)......... S. cattiense sp. nov. - Body coloration dark chestnut brown, posterior tergal margins black, legs dark brown, tips light brown (Fig. 1B). Lateral margin of prefemur with a sharp projection juxtaposed to coxal process (Fig. 5D). Apical margin of vulval operculum recessed centrally (Fig. 5H). Tip of laterotergite 1 slightly extended, sharp. Male antennomere 6 axe-shaped (Fig. 6A).............................................................................................. S. konkakinhense sp. nov. 6. Podomere 3 of anterior telopod distally with a lateral extension; podomere 4 of posterior telopod distally with an incurved process overlapping with process of podomere 2. Marginal setae of terga not reaching posterior margin.................................................................................................... S. separatum Attems, 1953 - Podomere 3 of anterior telopod cylindrical, without lateral extension; podomere 4 of posterior telopod without incurved pro- cess, but with 3 separate spines. Marginal setae of terga reaching the posterior margin................................................................................................... S. truncatum Wongthamwanich et al., 2012, Published as part of Semenyuk, Irina, Golovatch, Sergei I. & Wesener, Thomas, 2018, Four new species of giant pill-millipedes from Vietnam (Sphaerotheriida, Zephroniidae), pp. 535-550 in Zootaxa 4459 (3) on page 543, DOI: 10.11646/zootaxa.4459.3.7, http://zenodo.org/record/1458879, {"references":["Verhoeff, K. W. (1924) Results of Dr. E. Mjoberg's Swedish scientific expedition to Australia 1910 - 1913. 34. Myriapoda: Diplopoda. Arkif for zoologi, 16 (5), 1 - 142.","Attems, C. (1938) Die von Dr. C. Dawydoff in franzosisch Indochina gesammelten Myriopoden. Memoires du Museum national d'Histoire naturelle, New Series, 6 (2), 187 - 353.","Attems, C. (1953) Myriopoden von Indochina, Expedition von Dr. C. Dawydoff (1938 - 1939) Memoires du Museum national d'Histoire naturelle, New Series, A, 5, 133 - 230.","Wongthamwanich, N., Panha, S., Sierwald, P., Wesener, T. & Thirakupt, K. (2012) A new species of the giant pill-millipede genus Sphaerobelum Verhoeff, 1924 from northern Thailand, with an extensive description and molecular characters (Diplopoda: Sphaerotheriida: Zephroniidae). Zootaxa, 3220, 29 - 43."]}

Sphaerobelum truncatum Wongthamwanich, n. sp. Figs. 5 ���11 Material examined. Holotype male, CUMZ 2010.11, Thailand, Nan Province, Song Khwae District, Na Rai Luang Sub-district, Pang Hi Village; secondary forest; Latitude: 19.3962, Longitude: 100.6951; coll. N. Wongthamwanich; hand collecting; 22 August 2010. Paratypes: 4 males, CUMZ 2010.12 ���15, 4 females, CUMZ 2010.16 ��� 19, 1 male, FMNH-INS 0 0 0 0 0 72 674, 1 female, FMNH-INS 0 0 0 0 0 72 673, 1 male, ZFMK Myr 0 0 1, 1 female, ZFMK Myr 0 0 2, same data as holotype. Etymology. The specific epithet is a Latin adjective, meaning terminating abruptly, and refers to the truncated state of the lateral end of the third podomere on the anterior telopods (Fig. 11 C). Diagnosis. Sphaerobelum truncatum differs from all other Sphaerobelum species in the unique combination of the following characters of the anterior telopods (Figs. 11 A���C): four podomeres above syncoxite, second podomere posteriorly with a process (Figs. 11 B, C) and one spine on the anterior side of the inner margin (Fig. 11 C), third and fourth podomeres located posteriorly juxtaposed to process of second podomere, third podomere distally truncate, fourth podomere with three sclerotized spines all located on small thin separated sclerotized plates: one spine at apex, the other two on posterior side (Figs. 11 B, C). Anterior telopods without sclerotized teeth. S. truncatum differs from the closely related species S. separatum Attems, 1953 in the presence of a well-rounded fourth podomere on the posterior telopod (Figs. 11 D, E), while the tip of the posterior telopod in the latter species is prolonged into a thin process and curved towards the process of the second podomere. S. truncatum is the only giant pill-millipede species known to have a regularly ���square-wavy��� margin on the endotergum (Fig. 9 B). COI Sequence (paratype, FMNH-INS 0 0 0 0 0 72 674): Accession # JN 885184. See supplementary material. Description. Measurements: Males: length 18.6 ���24.0 mm, width at the seventh tergite 10.2���12.6 mm, width of thoracic shield 9.9���12.2 mm, height of thoracic shield 5.7���6.8 mm. Sizes of volvated male (for epimorphic stadia, Figs. 4 A, B): length (tergite 5 ���tergite 11 or 12) 12.1���13.9 mm, width (tergite 7) 10.2���12.6 mm, and height (thoracic shield���tergite 8) 10.7���12.6 mm. Females: length 21.2���30.4 mm, width at the seventh tergite 11.5���15.2 mm, width of thoracic shield 11.0��� 14.3 mm, height of thoracic shield 6.2���8.1 mm. Sizes of volvated female: length 13.0��� 17.6 mm, width 11.5���15.2 mm, and height 11.5���15.4 mm. Habitus (Fig. 5 C): Overall shape elliptical, tapering slightly from sixth tergite towards head. Coloration (Figs. 5 A���C): head dark brown, collum, thoracic shield, tergite, and anal shield black. Antennae and legs light yellow. Head: trapezoid, anterior part of the head with many long setae, posterior part densely dimpled; anterior margin of labrum with a single tooth. Eye field round, width approximately one mm, consisting of more than 60 densely-packed ocelli. No sclerotized crest/ridge between antenna socket and eye field. T��m��sv��ry organ (Fig. 5 D) between eye field and antenna socket, next to, but separated from eye field. Antennae (Figs. 6 A���C): with six visible antennomeres; basal antennomere with a few thinly scattered sclerotized nodules (Fig. 6 C); distal antennomere enlarged, round and club-shaped, with 81���84 sensory cones in males, 36���42 in females. Antennomere lengths: 6> 1> 2 = 3> 4 = 5. Mandibles (Figs. 7 A, B): with a single external tooth, a three-lobed internal tooth, six rows of pectinate lamellae, number of teeth on pectinate lamellae declining from apical to proximal lamella, molar plate with one visible ridge. Gnathochilarium (Fig. 7 C): with numerous long setae on the lingual lamella. Inner palpi with numerous elongate and sharp sensory cones (Figs. 7 D, E), reduced lateral palpi with a field of 6 sensory cones (Figs. 7 D, F). Collum: anterior and posterior margin with a few setae, center part sparsely setae, densely covered by dimplelike impressions. Thoracic shield: ridge on lateral lobe absent, accompanied by a few long setae; center part of anterior margin covered sparsely with long setae. Tergites 3���12 (Fig. 8): Anterior margins dominated by small conical spines (Fig. 8 C). Two submarginal rows of round sclerotized nodules: nodules of anterior-most row large and isolated, those of second row small and close together (Fig. 8 B). Anterior fifth of tergite with smooth zone, followed by a zone of 3���5 irregular rows of long setae, and then by an area of short setae (Fig. 8 B). Central four fifths of tergite dominated by short setae, nodules and slightly curved long setae inserted in large dimples (Figs. 8 A, D). Posterior margins with a few thick setae (Fig. 8 A). Endotergum (Fig. 9): with a regularly ���square-wavy��� margin (Fig. 9 B), outer zone with three rows of irregular marginal setae not reaching posterior margin. A single row of small elliptical cuticular impression present next to marginal ridge. Intersegmental membrane smooth, without cones or setae (Fig. 9 A). Anal shield: densely covered with long setae; round in both sexes. Inner side with three or four striae. Ventral side with one long black locking carina (Fig. 5 C, Lc). Legs: first pair of legs without coxal lobe (Fig. 5 F), first tarsi with two ventral spines and without apical spine; second pair of legs with small coxal lobes in male (Fig. 5 H), second tarsi with four ventral spines and without apical spine; third leg pair with acute coxal lobes, third tarsi with five or six ventral spines and one apical spine; ninth pair of legs with seven ventral spines and one apical spine on tarsus, femur with one long ridge (Fig. 11 G). Femur of ninth pair of leg 2.0 times longer than wide, tarsus 4.8 times longer than wide (Fig. 11 G). Stigmatic plate of first leg (Fig. 5 F): rounded, short and weakly curved, but forming a steep angle towards the coxa. Female vulva (Figs. 5 G, 11 H): consisting of two basal plates which are fused proximally and divided distally. Operculum pointed, basal part of operculum surrounding apical part of basal plates, distally protruding above coxa to basal half of prefemur. Vulva covers half (1 / 2) of coxa width, vulva is one-third (1 / 3) longer than coxa. Subanal plate (Fig. 10 A): brown, semicircular with a central shallow notch, laterally with four rib-like structures. Male gonopore (Fig. 10 B): covered by undivided sclerotized plate. Anterior telopods (Figs. 11 A���C): syncoxite with small spines, telopodite consisting of four podomeres, all podomeres covered by long setae. First podomere width equals height. Second podomere with posterior lobe-like, curved process with rounded edges; process protruding up to end of third podomere; anterior side of process with a single spine at inner margin. Third podomere distally truncate. Fourth podomere as long as third podomere, with three sclerotized spines located on small thin sclerotized fields, one spine at apex and the other two located on posterior side. Sclerotized teeth absent. Posterior telopods (Figs. 11 D, E): inner horns with sharp-edged tips, slightly curved backwards. Telopodite consisting of four podomeres. First podomere parallelogram-shaped. Second and third podomere dominated by short setae, margins with long setae. Process of second podomere weakly curved, distally glabrous, posteriorly apically enlarged and swollen (Fig. 11 D), anteriorly apically concave, spatulate, with a single sclerotized spine (Fig. 11 E). Moveable finger consisting of third podomere and small fourth podomere, slightly curved. Third podomere at inner margin with one small light colored spine, without any crenulated teeth. Fourth podomere at inner margin with two small sclerotized spines located in single brown sclerotized field. Intraspecific variation. Only seven males and six females of S. truncatum were available for analysis. However, some characters presented a clear difference between the sexes, such as the body length of females that tended to be longer than that of the males, while males have more sensory cones on the distal antennae than females. Life history. Adult specimens were collected in late August 2010 during the rainy season. Numerous eggs were found in an adult female, with most of them aggregated into many small clusters located between the intestine and the body wall, from the second leg pairs to posterior end of the body. The eggs as preserved in 70 % ethanol were orange in color. Mature eggs were round in shape with a diameter of about 2���2.5 mm. Distribution and ecology. The new species is only known from the type locality at 440���450 m altitude. There is a small stream running along the secondary dry evergreen forest and the forest floor is covered by laterite soil with saplings and seedlings, including a leaf litter of up to 3 cm depth. At the time of collection, the weather was a light precipitation, 92 % relative humidity, 25.5���26 ��C and 27 ��C soil and air temperature respectively. Most specimens were found on a 60 degree slope and only a few specimens were found on a flat plain area. Animals were found rolled up above the forest ground close to tree bases, in shallow soil pockets and under leaf litter. FIGURE 11. Sphaerobelum truncatum n. sp., A-G holotype, H female paratype, drawing, A: right anterior telopod, anterior view; B: last three podomeres of right anterior telopod, posterior view; C: last three podomeres of right anterior telopod, lateral view; D: last three podomeres of right posterior telopod, posterior view; E: last three podomeres of right posterior telopod, anterior view; F: stigmatic plate of 9 th left leg; G: 9 th left leg, posterior view; H: second right coxa with vulva, posterior view. Arrows point to the second podomere process. Abbreviations: Cx = coxa; ST = stigmatic plate; O = operculum; EP = external plate of vulva; IP = mesal plate of vulva. Roman numbers refer to number of podomere. Scale bars = 1 mm. Behavior. Sphaerobelum truncatum seems to be more active than Zephronia sp. They unroll their body in less than 5 minutes after capturing in the hand, which has never been observed for Zephronia sp. Sphaerobelum truncatum is well camouflaged, having similar color patterns to various objects in the environment, such as dried leaves and fruit seeds. The animals can also hide during the day by digging into the soil., Published as part of Wongthamwanich, Nattarin, Panha, Somsak, Sierwald, Petra, Wesener, Thomas & Thirakhupt, Kumthorn, 2012, A new species of the giant pill-millipede genus Sphaerobelum Verhoeff, 1924 from northern Thailand, with an extensive description and molecular characters (Diplopoda: Sphaerotheriida: Zephroniidae), pp. 29-43 in Zootaxa 3220 on pages 34-42, DOI: 10.5281/zenodo.212755, {"references":["Attems, C. (1953) Myriopoden von Indochina, Expedition von Dr. C. Dawydoff (1938 - 1939). Memoires du Museum national d'histoire naturelle, N. S., serie A, 5 (3), 133 - 230."]}

Key to species of Sphaerobelum 1. Anterior telopod with 3 podomeres........................................................................ 2 - Anterior telopod with 4 podomeres....................................................................... 4 2. Tergites shiny, glabrous.......................................................... S. clavigerum Verhoeff, 1924 - Tergites hairy........................................................................................ 3 3. Dirty olive-brown; head punctured and densely hairy; process of 2 nd podomere of posterior telopod apically strongly swollen, spherical, wider than basis of 3 rd podomere............................................ S. hirsutum Verhoeff, 1924 - Black; head smooth; process of 2 nd podomere of posterior telopods truncated; operculum of vulva two-horned, both horns blunt............................................................................ S. bicorne Attems, 1938 4. Podomere 3 of anterior telopod distally with lateral extension; podomere 4 of posterior telopod distally with incurved process, overlapping with process of podomere 2; marginal setae of tergite not reaching posterior margin.. S. separatum Attems, 1953 - Podomere 3 of anterior telopod cylindrical, without lateral extension; podomere 4 of posterior telopod without incurved pro- cess, but with 3 separated spines; marginal setae of tergite reaching posterior margin................. S. truncatum n. sp., Published as part of Wongthamwanich, Nattarin, Panha, Somsak, Sierwald, Petra, Wesener, Thomas & Thirakhupt, Kumthorn, 2012, A new species of the giant pill-millipede genus Sphaerobelum Verhoeff, 1924 from northern Thailand, with an extensive description and molecular characters (Diplopoda: Sphaerotheriida: Zephroniidae), pp. 29-43 in Zootaxa 3220 on page 34, DOI: 10.5281/zenodo.212755, {"references":["Verhoeff, K. W. (1924) Results of Dr. E. Mjoberg's Swedish scientific expedition to Australia 1910 - 1913. 34. Myriapoda: Diplopoda. Arkif for Zoologi, 16 (5), 1 - 142, pl. 1 - 5.","Attems, C. (1938) Die von Dr. C. Dawydoff in franzosisch Indochina gesammelten Myriopoden. Memoires du Museum National d'Histoire Naturelle, N. S. 6 (2), 187 - 353.","Attems, C. (1953) Myriopoden von Indochina, Expedition von Dr. C. Dawydoff (1938 - 1939). Memoires du Museum national d'histoire naturelle, N. S., serie A, 5 (3), 133 - 230."]}

Genus Sphaerobelum Verhoeff, 1924 A complete synonymy is provided by Jeekel (2001: 17) Type species. Sphaerobelum clavigerum Verhoeff, 1924 Other species included. S. hirsutum Verhoeff, 1924; S. bicorne Attems, 1938; S. separatum Attems, 1953, S. truncatum n. sp. Diagnosis. Males of Sphaerobelum species can be distinguished from all other giant pill-millipede genera in the world by the distally swollen process of the second joint of the posterior telopod. Comments. A comprehensive description of the genus Sphaerobelum cannot be given at the moment. However, all species currently assigned to the genus display the following unique combination of characters: posterior telopods consisting of four podomeres and with a large process on the 2 nd podomere which features a greatly enlarged, swollen-looking tip (Fig. 11 D). The latter feature has been described for all four species included in the genus and figured for three of them (Verhoeff, 1924, fig. 36 for S. clavigerum; Attems, 1938, figs. 18, 19 for S. bicorne; Attems, 1953, fig. 27 for S. separatum). Special attention should be given to the minute T��m��sv��ry organ, which is separated by a sclerotized crest from the eye field at least in some Zephronia species, a structure not present in Sphaerobelum., Published as part of Wongthamwanich, Nattarin, Panha, Somsak, Sierwald, Petra, Wesener, Thomas & Thirakhupt, Kumthorn, 2012, A new species of the giant pill-millipede genus Sphaerobelum Verhoeff, 1924 from northern Thailand, with an extensive description and molecular characters (Diplopoda: Sphaerotheriida: Zephroniidae), pp. 29-43 in Zootaxa 3220 on page 34, DOI: 10.5281/zenodo.212755, {"references":["Verhoeff, K. W. (1924) Results of Dr. E. Mjoberg's Swedish scientific expedition to Australia 1910 - 1913. 34. Myriapoda: Diplopoda. Arkif for Zoologi, 16 (5), 1 - 142, pl. 1 - 5.","Jeekel, C. A. W. (2001) A bibliographic catalogue of the Asiatic Sphaerotheriida (Diplopoda). - Myriapod Memoranda, 3, 5 - 38.","Attems, C. (1938) Die von Dr. C. Dawydoff in franzosisch Indochina gesammelten Myriopoden. Memoires du Museum National d'Histoire Naturelle, N. S. 6 (2), 187 - 353.","Attems, C. (1953) Myriopoden von Indochina, Expedition von Dr. C. Dawydoff (1938 - 1939). Memoires du Museum national d'histoire naturelle, N. S., serie A, 5 (3), 133 - 230."]}

As a first step towards an inventory of the giant pill-millipedes in Thailand, a new species of the genus Sphaerobelum Verhoeff, 1924, S. truncatum n. sp. is described from Nan Province, northern Thailand. A determination key is presentedfor all five known Sphaerobelum species. Clear morphological differences between S. truncatum n. sp. and the other four Sphaerobelum species were found on the anterior telopods. For the first time in Sphaerobelum, the partial mitochondrialCOI gene was sequenced for S. truncatum n. sp. and compared with distance, maximum parsimony and maximum likeli-hood methods to those of species from other giant pill-millipede genera. Sphaerobelum truncatum n. sp. was found to dif-fer from all other analyzed giant pill-millipedes, including species of Zephronia Gray, 1832, by 22–30%, includingnumerous amino acid changes, supporting the separate status of Sphaerobelum among other giant pill-millipede genera.Maximum likelihood and parsimony analyses support the placement of Sphaerobelum in the Zephroniidae. Figures of allrelevant structures of Sphaerobelum truncatum n. sp. are provided to allow the use of these characters in future descriptions of species of the family Zephroniidae.