taxonID	type	description	language	source
03A887C6FF8DFFB19CF4FAFD2F4E126B.taxon	materials_examined	Type species: Branchiostoma lithobioides Newport, 1845, by subsequent designation of Attems (1930). Diagnosis: Otostigminae with ten pairs of spiracles (on segments 3, 5, 7, 8, 10, 12, 14, 16, 19 and 20). Sixteen to 22 antennal articles, basal 3 to almost 5 glabrous dorsally. Cephalic plate without paramedian sutures on posterior part. Tooth-plates with four to seven teeth; forcipular trochanteroprefemoral process present. Tergite 1 overlapping cephalic plate. Leg 1 with femoral, tibial and tarsal spurs; most of subsequent legs with tibial and tarsal spur (s) only.	en	Joshi, Jahnavi, Karanth, Praveen K., Edgecombe, Gregory D. (2020): The out-of-India hypothesis: evidence from an ancient centipede genus, Rhysida (Chilopoda: Scolopendromorpha) from the Oriental Region, and systematics of Indian species. Zoological Journal of the Linnean Society 189: 828-861
03A887C6FF8DFFBA9CFDF8322B96120A.taxon	diagnosis	Diagnosis: Eighteen antennal articles; basal four to almost five articles glabrous dorsally. Cephalic plate relatively narrow. Coxosternal tooth-plates longer than wide, with four main teeth, with a small cusp on inner part of innermost tooth fused; median suture on forcipular coxosternum longer than tooth plates, extending ~ 20 % length of coxosternum. Sternites with paramedian sutures along 90 – 100 % of their length. Coxopleuron twice the length of sternite of ultimate leg-bearing segment; coxopleural process with two apical spines only. Leg 1 with two tarsal spurs, legs 2 – 20 with one. Material Holotype: BMNH (E) 200084, from Matheran, Maharashtra, India (The Natural History Museum, London, UK). Other specimens: CES 091396, CES 091397, from Matheran, Raigad district, Maharashtra; collected by Jahnavi Joshi in August 2008 – 2009; locality coordinates in the Supporting Information (Table S 1). Description: Length ≤ 70 mm. Eighteen antennal articles, basal 4.5 to five articles glabrous dorsally and five ventrally (Figs 6 A, 7 A). Cephalic plate and tergites smooth. Longitudinal median furrow on anterior 10 % of cephalic plate. Cephalic plate and T 1 brown to dark green, the following tergites brown with dark green-grey transverse band posteriorly; legs pale yellow. Forcipular coxosternal tooth-plates longer than wide, with four main teeth, the outer two teeth distinct, inner two teeth fused with a weak cusp on the innermost tooth; base of tooth-plates defined by oblique sutures diverging at 110 ° (Figs 6 B, 7 B); median suture relatively long, extending 20 – 25 % length of coxosternum. Coxosternum and trochanteroprefemur strongly puncate. Trochanteroprefemoral process bearingoneapicalandonelateraltooth. Tergitessmooth (Figs 6 G, 7 F), with paramedian sutures complete from TT 4 – 5. Tergites fully marginate starting from TT 6 – 8. Paramedian sutures 90 – 100 % length of sternites (Figs 6 F, 7 E); longitudinally ovate median depression in posterior part of sternites. Tergite of ultimate leg-bearing segment ~ 1.2 times wider than long, with parallel lateral margins, rounded posteromedially (Figs 6 H, 7 G). Sternite of ultimate leg-bearing segment slightly shorter than wide; lateral margins nearly straight, moderately convergent posteriorly; weak longitudinal median furrow (Figs 6 C, 7 C). Coxopleuron twice the length of sternite of ultimate leg-bearing segment; coxopleural process with two apical spines. Pores dense, with pore-field terminating distinctly beneath dorsal margin of coxopleuron; non-porose area on coxopleural process a narrow strip almost reaching to opposite posterior margin of sternite of ultimate leg-bearing segment (Figs 6 C, 7 C). Ultimate legs long, prefemur ≤ 4.9 mm, femur ≤ 3.75 mm, tibia ≤ 3 mm, tarsus 1 ≤ 2.5 mm and tarsus 2 ≤ 1 mm; prefemoral spine formula ventrolateral (VL), ventromedial (VM) and dorsomedial (DM) (Figs 6 H, I, 7 G – I). Leg 1 with two tarsal spurs, legs 2 – 20 with one. Leg 1 with a tibial spur and a femoral spur. Distribution: This species was recorded from evergreen forests of Matheran (a plateau at 800 m elevation), Maharashtra, in the northern parts of the Western Ghats. The type specimen (Fig. 6) and two individuals sampled in the present study (Fig. 7) are from the same locality. Forests there are much more fragmented and isolated than those of the central and southern Western Ghats. It is likely that this species might occur on the other adjacent plateaus and forested areas at similar elevations in the northern parts of the Western Ghats. Remarks: This species is part of the R. longipes clade, resolved as sister species to the widespread R. longipes and R. konda from the southern and northern Eastern Ghats. It was retrieved as a distinct species by both GMYC and mPTP (Fig. 2). It is morphologically unique among Indian Rhysida species in the paramedian sutures occupying most of or the entire length of the sternites, a higher number of glabrous antennal articles, confinement of two tarsal spurs to the first one or two leg pairs, relatively long tooth-plates and a longer median suture on the forcipular coxosternum than in other species. It is also unique in its relatively narrow cephalic plate and conspicuous punctae on the forcipular coxosternum.	en	Joshi, Jahnavi, Karanth, Praveen K., Edgecombe, Gregory D. (2020): The out-of-India hypothesis: evidence from an ancient centipede genus, Rhysida (Chilopoda: Scolopendromorpha) from the Oriental Region, and systematics of Indian species. Zoological Journal of the Linnean Society 189: 828-861
03A887C6FF86FFB99E41F83A28421343.taxon	description	(FIGS 8, 9)	en	Joshi, Jahnavi, Karanth, Praveen K., Edgecombe, Gregory D. (2020): The out-of-India hypothesis: evidence from an ancient centipede genus, Rhysida (Chilopoda: Scolopendromorpha) from the Oriental Region, and systematics of Indian species. Zoological Journal of the Linnean Society 189: 828-861
03A887C6FF86FFB99E41F83A28421343.taxon	diagnosis	Diagnosis (based on Indian material): 18 antennal articles. Coxosternal tooth-plates with four main teeth. Sternites with paramedian sutures confined to anterior 5 % of length. Ultimate leg prefemoral spine formula VL 3, VM 3, DM 3. Two tarsal spurs on legs 1 – 4. Tibial spur on legs 1 – 2. Material Holotype: NHMUK 1601663, West Indies. Peninsular Indian material: CES 091395, Matheran, Raigad district, Maharashtra; CES 091404, Mysuru, Mysuru district, Karnataka, both collected by Jahnavi Joshi in 2007 – 2010. Locality coordinates in the Supporting Information (Table S 1). Description: Length ≤ 50 mm. Eighteen antennal articles, basal three articles glabrous dorsally and 2.8 ventrally (Figs 8 A, 9 A). Cephalic plate and tergites smooth. Longitudinal median furrow on anterior 10 % of cephalic plate. Cephalic plate and T 1 brown, the following tergites brown with green pigmentation, legs pale yellow. Forcipular coxosternal tooth-plates wider than long, with four main teeth, separated into two groups, two inner and two outer; base of tooth-plates defined by oblique sutures diverging at 130 ° (Figs 8 B, 9 D). Trochanteroprefemoral process bearing one apical and two or three lateral teeth. Tergites with paramedian sutures complete from TT 4 – 7. Tergites fully marginate starting from TT 7 – 8. Tergites smooth (Figs 8 F, 9 J). Paramedian sutures 5 % length of sternites (Figs 8 G, 9 I); longitudinally ovate median depression in posterior part of sternites (Figs 8 G, 9 I). Tergite of ultimate leg-bearing segment as wide as long, without median furrow on posterior (Figs 8 C, 9 E). Sternite of ultimate leg-bearing segment with lateral margins convex, moderately convergent posteriorly; posterior margin gently concave (Figs 8 E, 9 C, F). Coxopleuron long, 1.6 – 2.0 times the length of sternite of ultimate leg-bearing segment; coxopleural process with two apical spines, one subapical, one lateral spine. Pores dense, pore-field terminating distinctly beneath dorsal margin of coxopleuron; non-porose area on coxopleural process a narrow strip almost reaching to opposite posterior margin of sternite of ultimate leg-bearing segment (Figs 8 E, 9 B, C, F). Ultimate legs long, prefemur ≤ 5 mm, femur ≤ 4 mm, tibia ≤ 3 mm, tarsus 1 ≤ 2.25 mm and tarsus 2 ≤ 1.75 mm; prefemoral spine formula VL 3, VM 3, DM 3 (Figs 8 C, D, 9 G, H). Legs 1 – 4 with two tarsal spurs, legs 5 – 19 with one. Legs 1 – 2 with tibial spur. Leg 1 with femoral spur. Distribution: This is one of the most widespread species occurring throughout the Oriental Region. In peninsular India, it also has a widespread distribution in many habitat types, including evergreen forests, rocky outcrops, dry forests and plantations. The current distribution record of this species according to Siriwut et al. (2018) and the present study is as follows: Southeast Asia and Melanesia: Cambodia; Vietnam; Myanmar – Thai – Malay Peninsula; Philippines; Papua New Guinea. East Asia: China, Taiwan. Indian subcontinent: Nepal; India (Kerala, Karnataka and Maharashtra); Sri Lanka; Pakistan. Indian Ocean: Chagos Island, Madagascar, Seychelles, Mauritius and Yemen. Central and South America: Mexico; Peru; Cuba (Cupul-Magaña, 2015); Haiti (Martínez-Muñoz & Perez-Gelabert, 2018). Remarks: Siriwut et al. (2018) recently assessed R. longipes from across Southeast Asia. They reported high genetic divergence among Indian and Asian R. longipes individuals, but those specimens from India are here recognized as a distinct species (R. pazhuthara). However, five individuals from across peninsular India are part of this widespread R. longipes, with less genetic divergence (Fig. 2). Only two of them are assessed for morphological description, the others being young specimens. The Indian specimens have two tarsal spurs on legs 1 – 4 and a tibial spur on legs 1 – 2, vs. two tarsal spurs on legs 1 – 7 or 8 and a tibial spur on legs 1 – 3 or 4 in material from Southeast Asia. This is one of the most widespread among the Rhysida species assessed in the present study, occupying two biogeographical subregions, peninsular India and Southeast Asia. The genetic divergence in COI (corrected p - distance) within this species was 0.05.	en	Joshi, Jahnavi, Karanth, Praveen K., Edgecombe, Gregory D. (2020): The out-of-India hypothesis: evidence from an ancient centipede genus, Rhysida (Chilopoda: Scolopendromorpha) from the Oriental Region, and systematics of Indian species. Zoological Journal of the Linnean Society 189: 828-861
03A887C6FF85FFA79E4AF9F028911362.taxon	description	(FIG. 10) h t t p: / / z o o b a n k. o r g / u r n: l s i d: z o o b a n k. o r g: a c t: 6 DE 67800 - C 5 C 8 - 4565 - A 2 A 6 - EC 0 B 61 B 711 F 5 Rhysida longipes (CES 07172, CES 07180, CES 07187), Rhysida cf. immarginata (CES 07193 – immature individual), Joshi & Karanth, 2011. Diagnosis: Eighteen antennal articles. Coxosternal tooth-plates with four teeth. Coxopleural process bearing two apical spines, one subapical and one lateral spine. Ultimate leg prefemoral spine formula VL 3, VM 2, DM 3. Two tarsal spurs on legs 1 – 18 (16); tibial spur on legs 1 – 3, variably on legs 4, 5 or 7. Type material Holotype: CES 091094, Pandimatte, Shendurney Wildlife Sanctuary, Kollam district, Kerala, India (8.8732 ° N, 77.1666 ° E, 600 m a. s. l.), collected by Jahnavi Joshi, 24 August 2009. Paratypes: CES 091080 Rockwood Estate, Shendurney Wildlife Sanctuary, Kollam district, Kerala; CES 091095, Pandimatte, Shendurney Wildlife Sanctuary, Kollam district, Kerala; CES 07172, Peppara Wildlife Sanctuary, Thiruvananthapuram district, Kerala. Other material: CES 07180, CES 07187, Ponmudi, Thiruvananthapuram district, Kerala; CES 091076, Rockwood Estate, Shendurney Wildlife Sanctuary, Kollam district, Kerala; CES 091082, Rosemla forest, Shendurney Wildlife Sanctuary, Kollam district, Kerala; CES 091092, Pandimatte, Shendurney Wildlife Sanctuary, Kollam district, Kerala, India; collected by Jahnavi Joshi in July 2008 – 2009. Locality coordinates in the Supporting Information (Table S 1). Etymology: Centipedes are called ‘ pazhuthara ’ in Malayalam, the language spoken in the state of the type locality. It is a noun in apposition. Description: Length ≤ 50 mm. Eighteen antennal articles, basal three articles glabrous dorsally and ventrally (Fig. 10 A). Cephalic plate and T 1 smooth. Longitudinal median furrow on anterior 10 % of cephalic plate. Cephalic plate and T 1 red – brown, the following tergites brown – green, legs reddish yellow. Forcipular coxosternal tooth-plates wider than long, with four main teeth, separated into two groups, the outer two teeth and the inner two teeth; base of tooth-plates defined by oblique sutures diverging at 110 ° (Fig. 10 B); median suture on anterior part of forcipular coxosternum about as long as tooth-plates. Trochanteroprefemoral process bearing two apical teeth and one lateral tooth. Tergites with paramedian sutures complete from T 7. Tergites fully marginate from TT 8 – 10. Tergites mostly smooth, sometimes with lateral depressions (Fig. 10 G). Paramedian sutures 5 – 10 % length of sternites (Fig. 10 F). Tergite of ultimate leg-bearing segment ~ 1.4 times wider than long, with parallel lateral margins (Fig. 10 C). Sternite of ultimate leg-bearing segment slightly shorter than wide, lateral margins mostly straight, moderately convergent posteriorly; posterior margins concave (130 °) (Fig. 10 D). Coxopleuron 1.75 times the length of sternite of ultimate leg-bearing segment; coxopleural process with two apical spines, one subapical spine and one lateral spine. Pores dense, with pore-field reaching close to dorsal margin of coxopleuron; non-porose area on coxopleural process a long, narrow strip almost reaching to opposite posterior margin of sternite of ultimate leg-bearing segment (Fig. 10 D). Ultimate legs long, prefemur ≤ 5 mm, femur ≤ 4 mm, tibia ≤ 3.5 mm, tarsus 1 ≤ 2.75 mm and tarsus 2 ≤ 2 mm; prefemoral spine formula VL 3, VM 2, DM 3 (Figs 10 C, H). Legs 1 – 18 usually with two tarsal spurs, in some specimens (CES 091080) only legs 1 – 16; leg 19 with one tarsal spur, leg 20 lacking tarsal spur (except one individual, CES 07187). Legs 1 – 3 (sometimes 4, 5 or 7) with a tibial spur. Leg 1 with femoral spur. Distribution: This species is found in the southern parts of the Western Ghats in the Agasthyamalai and Shendurney Mountains, south of the Shencottah Gap. This overlaps with another species from the R. immarginata clade, R. aspinsosa. This region represents one of the longest contiguous patches of evergreen forests in the Western Ghats and is known for high species diversity, including other scolopendrid genera, such as Digitipes and Ethmostigmus. Remarks: In species delimitation analyses, GMYC identifies two putative species within this taxon, which are distributed in the Agasthyamalai Hill Range and the Shendurney Wildlife Sanctuary, respectively (Fig. 2), whereas mPTP identifies only one species. The genetic variation within COI (corrected p - distance) is 0.04, and among the two putative species it is 0.02 (Shendurney clade) and 0.04 (Agasthyamalai clade), respectively. There are no distinct morphological characters to distinguish between these two putative species, and given the contiguity of the forests in the Agasthyamalai and Shendurney hill ranges, we treat it is as one. Morphologically, R. puzhuthara can be distinguished from the three other species in the R. longipes species complex by the presence of two tarsal spurs on legs 1 – 18 in most specimens, a distinctive spine formula on the ultimate leg prefemur and a relatively longer coxopleural process than, e. g. R. longipes.	en	Joshi, Jahnavi, Karanth, Praveen K., Edgecombe, Gregory D. (2020): The out-of-India hypothesis: evidence from an ancient centipede genus, Rhysida (Chilopoda: Scolopendromorpha) from the Oriental Region, and systematics of Indian species. Zoological Journal of the Linnean Society 189: 828-861
03A887C6FF9BFFA49E6DF9EC28E01746.taxon	description	(FIG. 11) h t t p: / / z o o b a n k. o r g / u r n: l s i d: z o o b a n k. org: act: 97 E 7286 E- 95 A 8 - 4241 - 8 B 01 - 6 C 8 A 62 B 08 D 5 D Diagnosis: Eighteen antennal articles. Coxosternal tooth-plates with four main teeth, separated into two groups, two outer teeth distinct, inner two / three teeth fused, with a cusp on inner part of innermost tooth. Sternites with paramedian sutures along 5 – 10 % of their length. Coxopleural process bearing two apical, one subapical and one lateral spine. Ultimate leg prefemoral spine formula VL 3, VM 2, DM 3. Two tarsal spurs on legs 1 – 7 (12), remaining legs to leg 20 with one; tibial spur on legs 1 – 4. Type material Holotype: CES 091394, Devmali, Kaorput district, Odisha, India (18.6600 ° N, 82.9900 ° E, 1200 m a. s. l.), collected by Ishan Agarwal and Aniruddha Datta-Roy in September 2010. Paratypes: CES 091029, Sitakund, Mayurbhanj district, Odisha; CES 091030, Bali, Mayurbhanj district, Odisha, India. Locality coordinates in the Supporting Information (Table S 1). Etymology: From ‘ konda ’, meaning hills in Telegu, a language spoken predominantly in Andhra Pradesh. It is a noun in apposition. Description: Length ≤ 60 mm. Eighteen antennal articles, basal three articles glabrous dorsally and ventrally (Fig. 11 A). Cephalic plate and T 1 smooth. Longitudinal median furrow on anterior 10 % of cephalic plate. Cephalic plate and T 1 red – brown, the following tergites red – brown with weak green pigmentation, legs pale yellow – red. Forcipular coxosternal tooth-plates wider than long, with four main teeth, the outer two teeth distinct, the inner two or three fused with a cusp on the innermost tooth; base of tooth-plates defined by oblique sutures diverging at 110 °; median suture about as long as tooth-plates, extending 10 % length of coxosternum (Fig. 11 B, F). Trochanteroprefemoral process bearing two apical teeth and one lateral tooth. Tergites with paramedian sutures complete from TT 4 – 5. Tergites fully marginate from TT 12 – 15. Tergites smooth (Fig. 11 G). Paramedian sutures 5 – 10 % length of sternites (Fig. 11 H). Tergite of ultimate leg-bearing segment ~ 1.8 times wider than long, with parallel lateral margins, rounded posteromedially (Fig. 11 C). Sternite of ultimate leg-bearing segment slightly shorter than wide, lateral margin convex, moderately convergent posteriorly, posterior margins concave (Fig. 11 H). Coxopleuron almost twice the length of sternite of ultimate leg-bearing segment; coxopleural process with two apical, one subapical and one lateral spine. Pores dense, with pore-field nearly reaching dorsal margin of coxopleuron; non-porose area on coxopleural process a narrow strip almost reaching to opposite posterior margin of sternite of ultimate leg-bearing segment (Fig. 11 D, E). Ultimate legs long, prefemur ≤ 5 mm, femur ≤ 4.75 mm, tibia ≤ 4 mm, tarsus 1 ≤ 3 mm and tarsus 2 ≤ 1.5 mm; prefemoral spine formula VL 3, VM 2, DM 3 (Fig. 11 I, J). Legs 1 – 12 (only in holotype) with two tarsal spurs, legs 13 – 20 with one; other three individuals with legs 1 – 7 with two tarsal spurs, legs 8 – 20 with one. Legs 1 – 4 with tibial spur. Leg 1 with femoral spur. Distribution: This species is recorded from hill ranges (1200 – 1600 m a. s. l.) in the moist forests and coffee plantations in central and northern parts of the Eastern Ghats. The mountains of the Eastern Ghats are much more fragmented and isolated than those of the Western Ghats and, apart from the mountain tops, much of the region has predominantly savanna or deciduous forests. More detailed surveys are required through central and northern parts of the Eastern Ghats. This distribution pattern has also been reported in some of the lizards and caecilian species found in similar habitat (Agarwal & Karanth, 2015; Gower et al., 2016). This species has an overlapping range in Chandragiri – Pottangi (Deomali) hill ranges with a species from the R. immarginata clade: R. sp. 2. Remarks: This species is part of the R. longipes species complex, sister species to the widespread R. longipes from peninsular India and Asia. The genetic divergence in COI (corrected p - distance) is high among R. konda, 0.117, and it is retrieved as a distinct species by both GMYC and mPTP (Fig. 2). Additionally, individuals sampled in the present study occupy distinct geographical areas in the Eastern Ghats, suggesting that a detailed survey across these mountain tops might reveal more diversity. Morphologically, it differs from its sister species, R. longipes, by having tibial spurs on the first four pairs of legs, vs. leg 1 only, lateral margination of tergites from TT 10 – 15, vs. TT 7 – 8 for R. longipes, and the ultimate leg prefemur having VM 2 vs. VM 3.	en	Joshi, Jahnavi, Karanth, Praveen K., Edgecombe, Gregory D. (2020): The out-of-India hypothesis: evidence from an ancient centipede genus, Rhysida (Chilopoda: Scolopendromorpha) from the Oriental Region, and systematics of Indian species. Zoological Journal of the Linnean Society 189: 828-861
03A887C6FF98FFA29F9DFDB029E614CD.taxon	description	(FIG. 12)	en	Joshi, Jahnavi, Karanth, Praveen K., Edgecombe, Gregory D. (2020): The out-of-India hypothesis: evidence from an ancient centipede genus, Rhysida (Chilopoda: Scolopendromorpha) from the Oriental Region, and systematics of Indian species. Zoological Journal of the Linnean Society 189: 828-861
03A887C6FF98FFA29F9DFDB029E614CD.taxon	diagnosis	Diagnosis: 20 – 21 (22) antennal articles. Coxosternal tooth-plates with 4 + 4 teeth, the innermost small. Coxopleural process ~ 1.5 times length of sternite of ultimate leg-bearing segment, with two apical spines and one subapical spine. Legs 1 – 18 with two tarsal spurs, legs 19 and 20 with one. Material C E S 0 7 1 0 2, R a m a n a g a r, R a m a n a g a r a d i s t r i c t, Karnataka; CES 07165, Neyyar Wildlife Sanctuary, Thiruvananthapuram district, Kerala; CES 07148, CES 07149, IISc Campus, Bangalore district, Karnataka; CES 08202, Devarayanadurga, Tumkur district, Karnataka; CES 08298, Biligirirangana Hills, Chamarajanagar district, Karnataka; CES 07262, CES 07263, Ranebennur, Haveri district, Karnataka; CES 08995, CES 08999, CES 081000, Thattekad Wildlife Sanctuary, Ernakulam district Kerala; CES 091301, CES 091302, Achankovil Reserve Forest, Pathanamthitta district, Kerala; CES 091387, Javadi hills, Tiruvannamalai district, Tamil Nadu; CES 091390, Chinnar Wildlife Sanctuary, Idukki district, Kerala; CES 091460, Srisailam, Kurnool district, Andhra Pradesh; CES 1517, Palaruvi Reserve Forest, Kollam district, Kerala; CES 1545, Kanakpura, Bangalore district, Karnataka; collected by Jahnavi Joshi in July and August, 2008 – 2009. Locality coordinates in the Supporting Information (Table S 1). Description: Length ≤ 80 mm. Twenty or 21 antennal articles (one specimen with 22), basal three articles glabrous dorsally and ventrally (Fig. 12 A). Cephalic plate and T 1 smooth. Longitudinal median furrow on anterior 10 % of cephalic plate. Cephalic plate and T 1 red – brown, the following tergites brown, legs pale yellow. Forcipular coxosternal tooth-plates wider than longer with four main teeth, the inner two sharing a common base, the outermost shorter in comparison to the second tooth; base of tooth-plates defined by oblique sutures diverging at 130 ° (Fig. 12 B, D). Trochanteroprefemoral process bearing two apical teeth and one or two lateral teeth. Tergites with paramedian sutures complete from TT 4 to TT 8. Tergites not marginated except 21 st, incipient margins as far anteriorly as T 15. Tergites mostly smooth, sometimes with longitudinal depressions (Fig. 12 G). Paramedian sutures 5 – 10 % length of sternites (Fig. 12 H). Tergite of ultimate leg-bearing segment little wider than long, with parallel lateral margins (Fig. 12 I, J). Sternite of ultimate leg-bearing segment about as wide as long, lateral margins convex, moderately convergent posteriorly; posterior margin gently concave; weak longitudinal median furrow (Fig. 12 C, E). Coxopleuron 1.6 times the length of sternite of ultimate leg-bearing segment; coxopleural process with two apical spines and one subapical spine. Pores dense, with pore-field nearly reaching dorsal margin of coxopleuron; non-porose area on coxopleural process a narrow, short strip towards the tip (Fig. 12 C, E). Ultimate legs long, with prefemur ≤ 5 mm, femur ≤ 4.5 mm, tibia ≤ 3.5 mm, tarsus 1 ≤ 3 mm and tarsus 2 ≤ 2 mm (Fig. 12 K – L); prefemoral spine formula VL 2, VM 1, DM 1; two individuals (CES 08999, CES 081000) with fewer prefemoral spines, VL 1, VM 1 (Fig. 12 M, N). Legs 1 – 18 (17) with two tarsal spurs, legs 19 – 20 (18) with one. First two pairs of legs with one tibial and femoral spur. Distribution: This species is one of the most widely distributed in peninsular India, occupying a wide range of habitat types, including wet evergreen, moist deciduous, dry deciduous and scrub forests and rocky outcrops. Remarks: Rhysida trispinosa was described by Jangi & Dass (1984) as a subspecies of R. lithobioides based on a specimen from Pune, Maharashtra (probably from semi-evergreen forests), but including material from other parts of Maharashtra and Tamil Nadu. The type specimens could not be located, but morphological description of the type specimen was compared with all the individuals analysed in the present study. Assignment of our specimens to R. trispinosa emphasizes the eponymous three spines on a moderately long coxopleural process, corresponding numbers of prefemoral spines, and the geographical proximity of some of the new collections and Tamil Nadu records of Jangi & Dass (1984). Specimens studied here agree with the description of R. trispinosa in all respects apart from the type specimen having only the first pair of legs with a tibial spur, whereas almost all of the individuals studied here have the first two pairs of legs with a tibial spur. In our species delimitation analyses, both GMYC and mPTP retrieve multiple species in R. trispinosa. In GMYC, seven distinct units are identified, of which three are represented by single individuals (CES 08217, CES 08999 and CES 101460) and four with multiple individuals (Fig. 3). The mPTP retrieves four distinct units, of which again two are represented by single individuals (Fig. 3). Among the two distinct units, one is represented by 13 individuals from many distinct geographical locations, including CES 08999, which is retrieved as a distinct species by GMYC. The second unit is represented by three individuals from northern Western Ghats and also distinct in GMYC. However, morphologically there are no fixed or distinguishing characters observed; therefore, it has been treated as part of R. trispinosa. There is also high (8.5 %) within-group variation in COI (corrected p - distance).	en	Joshi, Jahnavi, Karanth, Praveen K., Edgecombe, Gregory D. (2020): The out-of-India hypothesis: evidence from an ancient centipede genus, Rhysida (Chilopoda: Scolopendromorpha) from the Oriental Region, and systematics of Indian species. Zoological Journal of the Linnean Society 189: 828-861
03A887C6FF9EFFA39C25FD4D2F7713EF.taxon	description	(FIG. 13)	en	Joshi, Jahnavi, Karanth, Praveen K., Edgecombe, Gregory D. (2020): The out-of-India hypothesis: evidence from an ancient centipede genus, Rhysida (Chilopoda: Scolopendromorpha) from the Oriental Region, and systematics of Indian species. Zoological Journal of the Linnean Society 189: 828-861
03A887C6FF9EFFA39C25FD4D2F7713EF.taxon	diagnosis	Diagnosis: Twenty antennal articles. Coxosternal tooth-plates with 5 + 5 teeth. Coxopleural process relatively short, with two apical spines, lacking subapical, dorsal and lateral spines. Ultimate leg prefemur with VL 1 only or spines lacking completely. Material CES 091308, Achankovil Reserve Forest, Pathanamthitta district, Kerala; CES 07159, Neyyar Wildlife Sanctuary, Thiruvananthapuram district, Kerala; CES 091079, Rockwood Estate, Shendurney Wildlife Sanctuary, Kollam district, Kerala; CES 101451, Periyar Tiger Reserve, Kerala, India; collected by Jahnavi Joshi in July 2008 – 2009. Locality coordinates in the Supporting Information (Table S 1). Description: Length ≤ 45 mm. Twenty antennal articles, basal three articles glabrous dorsally and ventrally (Fig. 13 A). Cephalic plate smooth. Longitudinal median furrow on anterior 10 % of cephalic plate. Cephalic plate and T 1 brown – red, the following tergites brown with green – red pigmentation, legs pale yellow. Forcipular coxosternal tooth-plates with five teeth; teeth are separated into two groups, the outer two teeth and inner three; innermost is the smallest and sometimes fused, base of tooth-plates defined by oblique sutures diverging at 110 ° (Fig. 13 B). Trochanteroprefemoral process bearing two apical and two lateral teeth. Tergites with paramedian sutures complete from TT 5. Tergites not marginated except tergite of ultimate leg-bearing segment. Tergites smooth (Fig. 13 I). Paramedian sutures 5 % length of sternites (Fig. 13 J). Tergite of ultimate leg-bearing segment 1.5 times wider than long, with parallel lateral margins, pointed apex (Fig. 13 C). Sternite of ultimate leg-bearing segment slightly wider than long; lateral margins convex, moderately convergent posteriorly; posterior margins concave with an angle of 160 ° (Fig. 13 D). Coxopleuron 1.4 times the length of sternite of ultimate leg-bearing segment; coxopleural process with two apical spines. Pores dense, with pore-field terminating distinctly beneath dorsal margin of coxopleuron; non-porose area on coxopleural process narrow, short (Fig. 13 D, F). Ultimate legs long, with prefemur ≤ 4.5 mm, femur ≤ 4.5 mm and tibia ≤ 3.5 mm, tarsus 1 ≤ 3 mm and tarsus 2 ≤ 1.5 mm; prefemur with VL 1 (Fig. 13 G, H), except in one individual with VL 1, VM 1. Legs 1 – 8 with two tarsal spurs, legs 19 – 20 with one. Legs 1 – 3 with one tibial spur. Leg 1 with femoral spur. Distribution: This species is found across the Shencottah Gap, in low- and mid-elevation forests in the southern Western Ghats. It has an overlapping distribution south of the Shenkottah Gap with R. pazhuthara The type locality of R. aspinosa is Thattekad Wildlife Sanctuary, Kerala (Balan & Sureshan, 2017), which is further (~ 100 km Periyar Tiger Reserve) north of the localities from which we report it, making it a widely distributed species in the southern Western Ghats. Specimens collected from the type locality of R. aspinosus are placed within R. trispinosa in the molecular phylogeny (Fig. 3). Remarks: Genetic variation within COI (corrected p - distance) in this species was 12 %. In both GMYC and mPTP, it was considered as a distinct species (Fig. 3). Morphologically, R. aspinosa can be distinguished from the three other species in the R. immarginata clade in the southern Western Ghats by the presence of only the two apical spines on the coxopleural process and prefemoral spines at most represented by VL 1 only. It can be distinguished clearly morphologically from another sympatric species from the R. longipes clade, R. pazhuthara, because R. aspinosa has 20 antennal articles vs. R. pazhuthara having 18, and the ultimate leg prefemora of R. aspinosa having only VL 1 spines, whereas R. pazhuthara has VL 3, VM 3 and DM 3.	en	Joshi, Jahnavi, Karanth, Praveen K., Edgecombe, Gregory D. (2020): The out-of-India hypothesis: evidence from an ancient centipede genus, Rhysida (Chilopoda: Scolopendromorpha) from the Oriental Region, and systematics of Indian species. Zoological Journal of the Linnean Society 189: 828-861
03A887C6FF9FFFAE9C93F868281412D2.taxon	description	(FIG. 14) h t t p: / / z o o b a n k. o r g / u r n: l s i d: z o o b a n k. o r g: a c t: 5 F 4 AA 0 BC- 1227 - 4 FBB- 8 ACF-C 7 E 39 D 5 AEBA 4 Diagnosis: Twenty antennal articles. Coxosternal tooth-plates with 4 + 4 to 5 + 5 teeth. Paramedian sutures <5 % length of sternites. Coxopleural process with two apical spines and one dorsal spine. Ultimate leg prefemoral spine formula VL 2, VM 1, DM 1. Type material Holotype: CES 091360, Gunyacha Maal, Phansad Wildlife Sanctuary, Raigad district, Maharashtra, India (18.4152 ° N, 72.9712 ° E, 200 m a. s. l.), collected by Jahnavi Joshi, 16 September 2009. Paratypes: CES 091350, Guptbhima, Bhimashankar Wildlife Sanctuary, Pune district, Maharashtra; CES 091361, Gunyacha Maal, Phansad Wildlife Sanctuary, Raigad district, Maharashtra. Morphology only: CES 091359, Kondhavala, Bhimashankar Wildlife Sanctuary, Pune district, Maharashtra, India. Locality coordinates in the Supporting Information (Table S 1). Etymology: This species is found on lateritic plateaus in the northern Western Ghats, which are called ‘ sada ’ in Marathi, one of the predominant languages spoken in the region. It is a noun in apposition. Description: Length ≤ 40 mm. Twenty antennal articles, basal three articles glabrous dorsally and ventrally (Fig. 14 A). Cephalic plate and T 1 smooth. Longitudinal median furrow on anterior 10 % of cephalic plate. Cephalic plate and T 1 light grey, the following tergites and legs grey with light green; ultimate legs dark grey. Forcipular coxosternal tooth-plates slightly wider than longer, with 4 + 4 main teeth, the outermost smaller than the inner two; innermost tooth variably with a distinct cusp (5 + 4 or 5 + 5); base of tooth-plates defined by oblique sutures diverging at 120 ° (Fig. 14 B). Trochanteroprefemoral process bearing two apical teeth and one lateral tooth. Tergites with paramedian sutures complete from T 5. Tergites not marginated except tergite of ultimate leg-bearing segment. Tergites with longitudinal depressions (Fig. 14 G). Paramedian sutures 5 % length of sternites or indistinct (Fig. 14 F). Tergite of ultimate leg-bearing segment as wide as long, convex, with parallel lateral margins, rounded posteromedially (Fig. 14 H). Sternite of ultimate leg-bearing segment wider than long, with two sutures / depressions on both sides (Fig. 14 D). Coxopleuron moderately long, 1.5 times the length of sternite of ultimate leg-bearing segment; coxopleural process with two apical spines and a dorsal spine; pores dense, with pore-field terminating distinctly beneath dorsal margin of coxopleuron; non-porose area on coxopleural process a narrow, short strip towards the tip (Fig. 14 C, D). Ultimate legs long (Fig. 14 E, H), with prefemur ≤ 4.5 mm, femur ≤ 4 mm, tibia ≤ 3.5 mm, tarsus 1 ≤ 2 mm and tarsus 2 ≤ 1.5 mm; prefemur with large spines, as follows: VL 2, VM 1, DM 1 (Fig. 14 E, H). Legs 1 – 17 (one individual with 14) with two tarsal spurs, legs 18 – 19 with one; leg 20 lacking tarsal spur. Legs 1 – 2 with one tibial spur. Leg 1 with femoral spur. Distribution: This species is endemic to lateritic plateaus of the northern Western Ghats (also called as ferricretes; Thorpe & Watve, 2015), one of the unique and threatened habitats in the Western Ghats. This is home to many endemic species, with relatively narrow distributions (snakes, lizards, plants and caecilians; Thorpe & Watve, 2015). This species is currently known from two distinct localities: Bhimashankar Wildlife Sanctuary at the northern edge of the Western Ghats and Phansad Wildlife Sanctuary, on the west coast, relatively isolated lowland forests from the main mountain range of the Western Ghats. Remarks: Phylogenetically, this species is part of a clade composed of R. lewisi, R. immarginata, R. sp 1 and R. sp. 2, but its position in that clade is not resolved. It is retrieved as a single putative species by GMYC and mPTP (Fig. 3). It is diagnosibly distinct from all of these allied species in having negligible development of paramedian sutures on the sternites, a unique combination of the number of antennal articles (fixed at 20) and the numbers and arrangements of coxopleural and prefemoral spines.	en	Joshi, Jahnavi, Karanth, Praveen K., Edgecombe, Gregory D. (2020): The out-of-India hypothesis: evidence from an ancient centipede genus, Rhysida (Chilopoda: Scolopendromorpha) from the Oriental Region, and systematics of Indian species. Zoological Journal of the Linnean Society 189: 828-861
03A887C6FF92FFAF9E6FFF542B86164B.taxon	description	(FIG. 15) h t t p: / / z o o b a n k. o r g / u r n: l s i d: z o o b a n k. o r g: a c t: B 2 FE 6 A 5 A- 35 B 7 - 48 EE- 9850 - D 2109 E 1 D 2 A 64 Diagnosis: Twenty antennal articles. Prominent median ridges from T 3. Paramedian sutures complete from T 3. Coxopleural process with two apical spines, lacking subapical, dorsal and lateral spines. Ultimate leg prefemur with VL 2 spines only. Type material Holotype: CES 101438, Miao, Chanlang district, Arunachal Pradesh, India (27.48 ° N, 96.20 ° E, 250 m a. s. l.), collected by Jahnavi Joshi, 13 April 2011. Paratypes: CES 101440, Miao, Chanlang district, Arunachal Pradesh; CES 101438, Miao, Chanlang district, Arunachal Pradesh, India. Locality coordinates in the Supporting Information (Table S 1). Etymology: From the Lisu word for centipedes. Lisu is one of the tribes residing in Miao, Arunachal Pradesh. It is a noun in apposition. Description: Length ≤ 30 mm. Twenty antennal articles, basal three articles glabrous dorsally and three ventrally (Fig. 15 A). Cephalic plate smooth. Longitudinal median furrow on anterior 10 % of the cephalic plate. Cephalic plate and T 1 reddish, the following tergites red and pale brown; legs pale yellow. Forcipular coxosternal tooth-plates with 5 + 5 teeth in a group of three inner and two outer, the outermost smaller than inner three large teeth; innermost tooth small; base of tooth-plates defined by oblique sutures diverging at 160 ° (Fig. 15 B, D). Trochanteroprefemoral process bearing two apical teeth and one lateral tooth. Tergites with paramedian sutures complete from T 3. Tergites not marginated but with distinct median ridge starting from T 3. Tergites with a shallow depression on both sides of paramedian suture (Fig. 15 H). Paramedian sutures 10 % length of sternites (Fig. 15 G). Tergite of ultimate leg-bearing segment wider than long, with parallel lateral margins, posterior margin convex but with a nearly transverse median extent (Fig. 15 E). Sternite of ultimate leg-bearing segment slightly longer than wide, lateral margins strongly converging posteriorly, posterior margin concave, with an angle of 110 °; weak longitudinal median furrow (Fig. 15 C). Coxopleural process moderately long; coxopleuron ~ 1.5 times the length of sternite of ultimate leg-bearing segment; coxopleural process with two apical spines, lacking subapical and dorsal or lateral spines. Pores dense, with pore-field terminating distinctly beneath dorsal margin of coxopleuron; non-porose area on coxopleural process narrow, short (Fig. 15 C). Ultimate legs relatively short, with prefemur ≤ 3.5 mm, femur ≤ 3 mm, tibia ≤ 2.25 mm, tarsus 1 ≤ 1.5 mm and tarsus 2 ≤ 0.75 mm. Prefemur with VL 2 (Fig. 15 I, J). Legs 1 – 18 with two tarsal spurs, legs 19 – 20 with one tarsal spur. Leg 1 with a tibial and femoral spur. Distribution: This species is recorded from eastern Arunachal Pradesh (also known as Eastern Himalaya) in northeast India, part of the Himalayas biodiversity hotspot. The forests are contiguous with Myanmar and are biogeographically recognized as a distinct subregion, Indo-Burma (Myers et al., 2000). Remarks: Phylogenetically, this species is distinct and the sister species to other members of the R. immarginata clade (Fig. 3). A total of three individuals from the same locality are sampled and, for two individuals, COI successfully sequenced, depicting no variation between those two individuals. Both GMYC and mMPT retrieve a single distinct species (Fig. 3). The presence of distinct median ridges from T 3 distinguishes this species from the rest of the R. immarginata clade. It is additionally distinct in having paramedian sutures complete from T 3 (vs. TT 4 – 6 in other Indian species of the R. immarginata group). Having prefemoral spines confined to the VL row is shared only with R. aspinosa in this clade, but R. ikhalama has VL 2 vs. VL 1 in R. aspinosa. These two species are grouped in having the coxopleuron bearing only (two) apical spines.	en	Joshi, Jahnavi, Karanth, Praveen K., Edgecombe, Gregory D. (2020): The out-of-India hypothesis: evidence from an ancient centipede genus, Rhysida (Chilopoda: Scolopendromorpha) from the Oriental Region, and systematics of Indian species. Zoological Journal of the Linnean Society 189: 828-861
03A887C6FF93FFAD9C85FD1828081644.taxon	description	(FIG. 16) h t t p: / / z o o b a n k. o r g / u r n: l s i d: z o o b a n k. o r g: a c t: D 2 DD 6016 - 549 F- 45 BF- 9097 - 9 F 6582 A 56 BDC	en	Joshi, Jahnavi, Karanth, Praveen K., Edgecombe, Gregory D. (2020): The out-of-India hypothesis: evidence from an ancient centipede genus, Rhysida (Chilopoda: Scolopendromorpha) from the Oriental Region, and systematics of Indian species. Zoological Journal of the Linnean Society 189: 828-861
03A887C6FF93FFAD9C85FD1828081644.taxon	diagnosis	Diagnosis: Twenty (19) antennal articles, three glabrous. Coxosternal tooth-plates with four main teeth. Coxopleural process with two apical spines, lacking subapical spine. Ultimate leg prefemoral spine formula VL 2, VM 1, DM 1. Legs 1 – 18 (16 / 17) with two tarsal spurs. Type material Holotype: CES 07224, Anshi-Dandeli Tiger Reserve, Karwar district, Karnataka, India (14.9890 ° N, 74.3712 ° E, 600 m a. s. l.), collected by Jahnavi Joshi, 17 August 2008. Paratypes: CES 08908, Kurunjal, Kudremukh National Park, Chikkamagaluru district, Karnataka; CES 091010, Kudremukha Peak, Kudremukh National Park, Chikkamagaluru district, Karnataka, India. Other material: CES 07238, Kumta, Uttara Kannada district, Karnataka; CES 08909, CES 08914, Kurunjal, Kudremukh National Park, Chikkamagaluru district, Karnataka; CES 08920 Tadoli, Kudremukh National Park, Chikkamagaluru district, Karnataka; CES 08950, Shirgaovkar Road, Amboli, Sindhudurg district, Maharashtra; CES 07271, Radhanagari Wildlife Sanctuary, Kolhapur district, Maharashtra; CES 08275, Radhanagari Wildlife Sanctuary, Kolhapur district, Maharashtra, India; collected by Jahnavi Joshi in August – September 2008 – 2009. Locality coordinates in the Supporting Information (Table S 1). Etymology: This species is named for Dr John G. E. Lewis, in appreciation of his exemplary work on Scolopendromorpha. Description: Length ≤ 45 mm. Twenty (19) antennal articles, basal three articles glabrous dorsally and ventrally (Fig. 16 A). Cephalic plate and T 1 smooth. Longitudinal median furrow on anterior 10 % of cephalic plate. Cephalic plate and T 1 brown – dark green, the following tergites brown with red – green pigmentation, legs pale yellow. Forcipular coxosternal tooth-plates wider than long, with four main teeth separated into two groups, two inner and two outer; outermost smaller in comparison to other three teeth; base of tooth-plates defined by oblique sutures diverging at 110 ° (Fig. 16 B, E). Trochanteroprefemoral process bearing two apical and two lateral teeth. Tergites with paramedian sutures complete from TT 4 – 5 in most specimens, rarely from TT 6 or TT 7. Tergites not marginated. Tergites with shallow longitudinal depressions (Fig. 16 G). Paramedian sutures 10 % length of sternites (Fig. 16 H). Tergite of ultimate leg-bearing segment ~ 1.3 times wider than long, with parallel lateral margins, rounded posteromedially (Fig. 16 J). Sternite of ultimate leg-bearing segment slightly shorter than wide, lateral margins gently convex, moderately convergent posteriorly; faint depression near and parallel to lateral margin; strong median depression (Figs 16 D, I). Coxopleural process 1.5 times the length of sternite of ultimate leg-bearing segment, with two apical spines, lacking subapical and dorsal spines. Pores dense, with pore-field terminating distinctly beneath dorsal margin of coxopleuron; non-porose area on coxopleural process a narrow strip almost reaching to opposite posterior margin of sternite of ultimate leg-bearing segment (Fig. 16 D, I). Ultimate legs long, prefemur ≤ 3.5 mm, femur ≤ 3.3 mm, tibia ≤ 3 mm, tarsus 1 ≤ 2.25 mm and tarsus 2 ≤ 1.5 mm; prefemoral spine formula VL 2, VM 1, DM 1 (Fig. 16 I, J). First 18 (less commonly 16 or 17, exceptionally 11) pairs of legs with two tarsal spurs; remaining legs to leg 19 with one; leg 20 lacking tarsal spur. Legs 1 and 2 (exceptionally, leg 1 only) with tibial spur. Leg 1 with femoral spur. Distribution: This species is distributed in the central and northern parts of the Western Ghats. It is currently known from three distinct locations in the former and two in the latter. It is found across a broad elevational range (500 – 1800 m a. s. l.) and in different types of habitats, including low- and high-elevation montane forests, grassland and moist deciduous forests. Remarks: Phylogenetically, R. lewisi is sister to R. immarginata from Southeast Asia, one of the widely distributed species in Asia (Fig. 3). It is retrieved as a putative species by mPTP. In GMYC, there are three distinct units, one single specimen and another two with multiple individuals. Within-group variation in COI (corrected p - distance) is 4.5 %. Morphologically, the three units identified by GMYC cannot be distinguished, and mPTP recovers only one unit. Therefore, it is treated currently as a single species. Morphologically, R. lewisi can be distinguished from its closest relative, R. immarginata, based on the arrangement of spines on the coxopleural process and the number of antennal articles. Rhysida lewisi has only two apical spines, whereas R. immarginata has two apical and one subapical spine. Rhysida lewisi has 20 (19) antennal articles, whereas R. immarginata ranges from 16 to 18 antennal articles. Within the R. immarginata clade, two more distinct species are identified by both species delimitation methods with strong support (Fig. 3): Rhysida. sp. 1 and R. sp. 2. Rhysida sp. 1 is sister to a clade consisting of R. immarginata and R. lewisi, with a broad geographical distribution in the central and southern Western Ghats and the southern Eastern Ghats. However, it is represented by three immature individuals and thus cannot be described morphologically, but provides evidence for another species. More sampling in the Western and Eastern Ghats will be essential to describe this species morphologically. Rhysida sp. 2 is represented by two individuals from the central Eastern Ghats, CES 101400 and CES 101032. Phylogenetically it is distinct, with a PP of one and BS of 100 %, but its position is unresolved, and it is part of a clade consisting of R. immarginata, R. sada, R. lewisi and Rhysida sp. 1.	en	Joshi, Jahnavi, Karanth, Praveen K., Edgecombe, Gregory D. (2020): The out-of-India hypothesis: evidence from an ancient centipede genus, Rhysida (Chilopoda: Scolopendromorpha) from the Oriental Region, and systematics of Indian species. Zoological Journal of the Linnean Society 189: 828-861
