#ZooKeys ZooKeys 1228: 225-273 (2025) DOI: 10.3897/zookeys.1228.143007 Centipedes (Myriapoda, Chilopoda) of Aldabra Atoll (Seychelles) George Popovici'”®, Gregory D. Edgecombe?® 1 Department of Life Sciences, Imperial College London, London SW7 2AZ, UK 2 "Grigore Antipa" National Museum of Natural History, Pavel Dimitrievici Kiseleff St. 1,011341, Bucharest, Romania 3 The Natural History Museum, Cromwell Road, London SW7 5BD, UK Corresponding author: George Popovici (gp522@ic.ac.uk) OPEN Qaccess Academic editor: Pavel Stoev Received: 28 November 2024 Accepted: 13 January 2025 Published: 21 February 2025 ZooBank: https://zoobank. org/3A5C895C-ACF6-4E44-BD53- 6296E1F9EAB1 Citation: Popovici G, Edgecombe GD (2025) Centipedes (Myriapoda, Chilopoda) of Aldabra Atoll (Seychelles). ZooKeys 1228: 220-273. https://doi.org/10.3897/ zookeys.1228.143007 Copyright: © George Popovici & Gregory D. Edgecombe. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International - CC BY 4.0). Abstract Centipedes collected during Royal Society surveys of the arthropod fauna of the Aldabra Atoll in 1968-1975 are identified, described, and illustrated to provide the first checklist to the Aldabran centipede fauna, comprising 12 species. These newly include the lithobio- morph Lamyctes tristani (Pocock, 1893), the scolopendromorphs Scolopendra morsitans Linnaeus 1758, Cryptops cf. japonicus Takakuwa, 1934, Cryptops mauritianus Verhoeff, 1939, and Cryptops nigropictus Takakuwa, 1936, and the geophilomorphs /typhilus cf. tae- niaformis (Lawrence, 1960), Mecistocephalus angusticeps (Ribaut, 1914), Mecistocepha- lus lohmanderi Verhoeff, 1939, Orphnaeus dekanius Verhoeff, 1938, Ribautia cf. paucipes Attems, 1952, and Tuoba sydneyensis (Pocock, 1891). The geophilomorph genera Hovanyx Lawrence, 1960, syn. nov., and Mixophilus Silvestri, 1929, syn. nov., are revised in light of the examined material and hereby designated junior subjective synonyms of Tuoba Cham- berlin, 1920 with the species Geophilus lemuricus Verhoeff, 1939, syn. nov., and Hovanyx waterloti Lawrence, 1960, syn. nov., designated as junior subjective synonyms of T. syd- neyensis. The oryid genus Nycternyssa Crabill, 1959, syn. nov., is revised and designated a junior subjective synonym of Orphnaeus Meinert, 1870. New data on intraspecific mor- phological variation are presented for C. nigropictus, with the validity of Cryptops daszaki Lewis, 2002 being questioned following examination of its type material. The affinities and possible origins of the Aldabran centipede fauna are found to be mainly East African, with several species occurring across other islands in the Western Indian Ocean. Key words: Checklist, taxonomy, Western Indian Ocean Introduction The Western Indian Ocean Islands, delimited at the north by Socotra and at the south by Madagascar, have been identified as a global biodiversity hotspot (Myers et al. 2000; Attorre and Van Damme 2020; Agnarsson et al. 2015). Although extremely variable in size, and geological origin and histo- ry, these islands harbour a terrestrial fauna notable for striking patterns of radiation and endemicity (Legros et al. 2020; Bergsten and Bistro6m 2022; Sherlock et al. 2024). Despite being less intensively studied than their ma- rine faunas, especially in smaller island groups, the terrestrial faunas of the Western Indian Ocean face significant risk of extinction due to the impacts of introduced species (Gerlach 2006; Griffiths and Florens 2006; Cheke 2010; 225 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) Hawlitschek et al. 2011) and habitat degradation (Haverkamp et al. 2017; Ibouroi et al. 2021). Smaller island groups can offer unique insights into un- derstanding the patterns of colonisation and diversification in an area ex- tensively shaped by eustatic changes (Camoin et al. 2004). The paleogeo- graphic history of coral atolls in the Western Indian Ocean in particular has identified the present composition of terrestrial faunas on these islands as recent in origin (Austin et al. 2004; Agnarsson and Kuntner 2012; Steibl et al. 2024) and reflective of multiple colonisations from mainland populations and refugia (Nagy et al. 2003; Kehlmaier et al. 2023). Aldabra Atoll is located in the Western Indian Ocean and has an area of approximately 155 km? and a maximum elevation of 8 m, being the sec- ond largest coral atoll in the world (Plummer 1995). It is composed of four main islands, Grand Terre (114 km?), Malabar (26.5 km?), Picard (9.28 km’), and Polymnie (1.93 km?), in addition to 40 small islets. The nearest large landmasses include the coast of Tanzania (640 km), the northern coast of Madagascar (425 km), Ngazidja (Comoros) (380 km), and Mahé (Sey- chelles) (1130 km). Palaeogeographical research on Aldabra has identified a complete submergence event during the last interglacial period, with any terrestrial fauna that had previously colonized the atoll having almost cer- tainly disappeared approximately 125,000 years before the present (Warren et al. 2005). Re-emergence of a more permanent dry land area, together with signs of a terrestrial fauna including land snails is estimated to have occurred 118,000-80,000 years before the present, with continued fluctua- tions in the sea level exposing a dry land area twice as large as the current one ~ 27,000 years before the present. This was followed by dry land area reduction to its present condition after flooding of the inner lagoon ~ 5000 years before the present (Braithwaite et al. 1973; Hume et al. 2018). The faunal affinities of the Aldabra Atoll have been identified as predominant- ly East African, with general biogeographical patterns indicating dispersal and colonisation from continental East Africa, Madagascar, and the Sey- chelles (Cogan et al. 1971; Hill and Newbery 1982; Lawrence 2015), in con- trast to the Mascarenes and Comoros, in which dispersal from Madagascar seemingly predominates (Agnarsson and Kuntner 2012). Given its young geological age and recent complete submergence, the fauna of Aldabra is comparatively less rich in endemic taxa than the granitic inner islands of the Seychelles (Agnarsson and Kuntner 2012). Previous surveys of the centipede fauna of the Seychelles (Bonato and Mi- nelli 2010; Lewis 2010b; Stoev and Gerlach 2010) have identified three spe- cies from Aldabra, Australobius inflatitarsis Eason, 1978, Mecistocephalus an- gusticeps (Ribaut, 1914), and Tuoba sydneyensis (Pocock, 1891). The threat level faced by endemic Myriapoda within the inner and outer islands of the Seychelles, including the Aldabra group, is significantly higher than the threat level faced by indigenous non-endemic (i.e., widespread) or introduced myr- iapods (Gerlach 2012). Observations on the natural history of large Scolopen- dromorpha in Mauritius and the Rodrigues provide circumstantial evidence on the vulnerability of island centipede populations to introduced predators (Lewis et al. 2010; Tercel et al. 2024). We provide a comprehensive checklist and taxonomic revision of the cen- tipedes from the Aldabra Atoll collected between 1968 and 1975 to provide ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 226 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) historic baseline data to inform future conservation efforts of Aldabran ter- restrial arthropods and shed light on the poorly known centipede fauna of the Western Indian Ocean. Materials and methods Morphological examination Specimens were examined under a Nikon SMZ1270 stereomicroscope and a Leica DMR binocular microscope. Partial dissection was carried out as necessary accord- ing to the protocol outlined by Pereira (2000), with anatomical structures tempo- rarily mounted in glycerol. Where necessary, specimens were cleared by temporary mounting in 80% lactic acid. Drawings were prepared with the aid of acamera lucida. Specimen data All specimens examined are part of the Myriapoda collection of the Natural His- tory Museum, London (NHMUK). Collection data for a total of 181 specimens from the Aldabra Atoll are given in the Results section for each species. Sam- pling was undertaken in 1974 and 1975, apart from four specimens collected in 1968 and three from 1973. In addition to material collected from Aldabra, the following specimens have been examined, either as conspecifics from other geographic regions or for comparison to Aldabra samples: Lamyctes tristani (Pocock, 1893) NHMUK015626352, 29 9, Downtown, Diego Garcia, Chagos Archipelago, 7.263°S, 72.374°E, 26.06.2022, leg. W. Rabitsch, suction sampler; NHMUK015619670, 19, Diego Garcia, Chagos Archipelago, wetland site, 7.310°S, 72.419°E, 22.06.2022, leg. W. Rabitsch, suction sampler. NHMUK01 5087793, syntypes, Tristan da Cunha. Ballophilus maldivensis Pocock, 1906 BMNH #200555, Chilo 1952-.12.11.102, 1 2 (holotype), Midu (2), Addu Atoll, Maldives, 1899-1900, Maldive-Laccadive Expedition 1899-1900. Cryptops daszaki Lewis, 2002 14 (holotype, Spm. 1), fle aux Aigrettes, damp soil against wall of Warden's House, 19 October 1995, leg. S. J. Lewis. Som 4 and 5, in soil under ebony litter, grid square E4, 18.10.1995, leg. J. G. E. Lewis. Cryptops niloticus Lewis, 1967 BMNH(E)#200011 Chilo.1966.9.6.2, 19 (holotype), Blue Nile Bridge, Khar- toum, Sudan, 28.09.1964, leg. J. G. E. Lewis, 13 (allotype), under stone, top of Nile Bank, Blue Nile Bridge, Khartoum, Sudan, 09.11.1962, leg. J. G. E. Lewis, BMNH(E) #200014 Chilo.1966.9.6.11, 1 juvenile (paratype), Blue Nile Bridge, Khartoum, Sudan, 09.11.1962. ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 227 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) Mecistocephalus glabridorsalis Attems, 1900 NHMUK015991408, 2 specimens, Serpent Island, Mauritius, 04.11.1995, leg. P. Daszak & J. Cottingham. Orphnaeus brevilabiatus (Newport, 1845) NHMUK015991423, 12, S. (South) Tenasserim (Great Tenasserim Riv- er, Myanmar), BM1889.7.15.73-4, leg. E. W. Oates; NHMUK015991422, 14, Rangoon (Yangon, Myanmar), BM1889.7.15.73-4, leg. E. W. Oates; NHMUK015991420, 19, Singapore, 18.10.1898, 98.10.18.53-55, leg. H. N. Hidley; NHMUK015991421, 12, Takhamen, Siam (Tak province, Thailand), 1897.9.7.34, leg. S. S. Flower; NHMUK015991412, 14:22, Agraky Hills, Ye- men, leg. G. W. Berry. Orphnaeus dekanius Verhoeff, 1938 NHMUK015991413, 19, Ratnapura, Ceylon (Ratnapura, Sri Lanka), 19.08.1892, 92.8.19.6; NHMUK015991414, 14, Pundulaya, Ceylon (Pundaluoya, Sri Lan- ka), 13.12.1899, 1899.12.13.42; NHMUK015991415, 19, Nundulaya, Ceylon (Nuwara Eliya, Sri Lanka), 13.12.1899, 99.12.13.41; NHMUK015991416, 19, Ceylon (Sri Lanka), 88.55; NHMUK015991417, 23.419, Singapore, 18.10.1898, 98.10.18.53-55, leg. H. N. Hidley; NHMUK015991418, 12119, Singapore, leg. H. N. Hidley; NHMUK015991419, 1, Kenurus, Maldives, 1951.12.11.100, Mal- dive-Laccadive Expedition 1899-1900. Ribautia cf. paucipes Attems, 1952 NHMUK015991409, 1 specimen, fine soil overhanging rock surface, Coin de Mire, Mauritius, 19°56.5'S, 57°37'E, 27.10.1995, leg. J. G. E. Lewis & S. J. Lewis. Tuoba sydneyensis (Pocock, 1891) NHMUK015991411, 3¢4'329, under slabs of tuff rock, Serpent Island, Mauri- tius, 04.11.1995, P Daszak & J. Cottingham. Results Checklist Order Lithobiomorpha Family Henicopidae Genus Lamyctes Meinert, 1868 Lamyctes tristani (Pocock, 1893) Fig. 1 Examined material. 41 specimens: NHMUK015991449, 192, Cing Cases, 16.03.1974; NHMUK015991450, 29, Malabar, 14.02.1975; NHMUK015991451 ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 228 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) _0.25mm____ G D OOsIHAnA Figure 1. Lamyctes tristani (Pocock, 1893) A, B, D NHMUK015991452 A forcipular segment, ventral view B anterior mar- gin of forcipular coxosternite, ventral view C NHMUK015991458, distal end of tibia of leg pair 12, lateral view D distal end of tibia of leg pair 12, lateral view ENHMUK015991456, Female gonopods, ventral view. 19, Picard, 28.02.1975; NHMUK015991452, 39, 1 juvenile, Cing Cases B. F., 20.03.1974; NHMUK015991453, 2°, Picard, 23.02.1974; NHMUK015991454, 22, Cing Cases, 24.03.1974; NHMUK015991458, 32, Malabar, 08.06.1974; NHMUK015991455, 19, 1 juvenile, Aldabra, 30.01.1975; NHMUK015991456, 219, Picard, Summer 1975, leg. V. W. Spaull; NHMUK015991457, 2°, 1 juvenile, Ochna litter, Cing Cases B. F., 23.03.1974. Remarks. The 38 specimens from the Aldabra Atoll (from Cing Cases, Mal- abar, and Picard) for which sex can reliably be determined are all females, such that the population is parthenogenetic. They are likely conspecific with L. tristani as described from the Chagos Archipelago (Popovici et al. 2024), sharing the distinctly reduced spinous process on the tibia of leg pair 12, often taking the form of a rounded bump (Fig. 1C). The stability of this char- acter is however uncertain as specimens from localities in the Aldabra Atoll exhibit conspicuous variation in the shape of the process (Fig. 1D, Table 1). Additional similarities between these two populations include the morpholo- gy of the forcipular coxosternite (Fig. 1A, B), the number of antennal articles, and the number and shape of the spurs on the female gonopods (Fig. 1E). Variability in the characters of antennal article number, number, and distri- bution of coxal pores and shape of the distal spinous process on the tibia of leg pair 12 are listed in Table 1 and compared with putatively conspecific individuals from the Chagos Archipelago, excluding anamorphic stages. ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 229 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) Table 1. Morphological variability of putative L. tristani specimens from Western Indian Ocean localities. Asterisk indi- cating regenerated appendage. Locality Body length | Antennal article Coxal pore formula (legs 12/12 — | Shape of tibial spinous process on (mm) number 13/13 -— 14/14 — 15/15) leg pair 12 Cing Cases (Aldabra) 3.5 20 1 ASA eS Rounded bump Cing Cases (Aldabra) 4 20 F247 Rounded bump Cing Cases (Aldabra) 5 Wg PID D2 Bl3 242 Acuminate, minute Cing Cases (Aldabra) 5 22 3/3'=°3/3 33/3 — 2/2 Acuminate, minute Cing Cases (Aldabra) 5 - 2/2. 7373 13/3)= 2/2 - Cing Cases (Aldabra) 5 22 22 3/33/38 = 272 Rounded bump Cing Cases (Aldabra) 6 23 3/3'°= 3/3 =3/3'= 3/3 Acuminate, minute Malabar (Aldabra) 5.5 24 DEP O42 +3735 Pf 2 - Malabar (Aldabra) 5.5 24 212, =.2/2-Bl3 = 2/2 Rounded bump Malabar (Aldabra) 6 (23)24 242 = 2/2--9/3.-19/2 Rounded bump Malabar (Aldabra) 6 15(20) 3/3°= 8/3:-73/3'=18/3 Asymmetrical, rounded bump on left tibia, acuminate spur on right tibia Malabar (Aldabra) 6 24 3/3: =. 3/3 3/3-—3/3 - Picard (Aldabra) a5 19 17 SSA, Al. = WA Rounded bump Picard (Aldabra) 3.5 20 so Naot al Dread 4 na | Rounded bump Picard (Aldabra) 3.5 20 el: = eae tel Rounded bump Picard (Aldabra) 4 20 1 Sul SL al Rounded bump Picard (Aldabra) 4 20 a ao oes Pra el a Me | Rounded bump Picard (Aldabra) 4 20 1A =a ay = Rounded bump Picard (Aldabra) 4 20 TAS IF Sal aay Rounded bump Picard (Aldabra) 4 21 2[2 > 2/2 = 2/2 = V1 Rounded bump Picard (Aldabra) 4 21 22 =2I/2 ye W| Rounded bump Picard (Aldabra) 4 20 iA Sit a ; Picard (Aldabra) 4.5 24 2/2212 3/38 = 2/2 Acuminate, minute Picard (Aldabra) 4.5 (22)23 Qi. meni 2 3/3 2/2 Rounded bump Picard (Aldabra) 45 21 TDS FP = 979 ; Picard (Aldabra) 5 (24)25 PRAVR A 2f2-— 2/2 Rounded bump Picard (Aldabra) 5 (22)23 Boole =o) a = 2/2 Acuminate, minute Picard (Aldabra) 5 t5* 3/375 S/R of3. tee Acuminate, minute Picard (Aldabra) 5 24 2f2=3/3' 3.8/3.4 2/2 Acuminate, minute Picard (Aldabra) 5 (21)24 2 DISS Slam 212 Acuminate, minute Picard (Aldabra) 5 - 2/23/38 8/3 = 2/2 Acuminate, minute Picard (Aldabra) 5 24 2/2 Bl Or Sam 2LZ Acuminate, minute Picard (Aldabra) 5 20 2/ 22/2 ~3f3°— 2)2 Rounded bump Picard (Aldabra) 5.5 24 3/8 B/s'> 3/3°5 3/8 Acuminate, minute Picard (Aldabra) 6 24 3/3:= 3/3= 3/33/38 Acuminate, reduced in size Picard (Aldabra) 7 (24)25 B/8 5138 28/8 33/3 Acuminate, reduced in size Diego Garcia (Chagos) 6 27 3/3: 3138/S= 3/S- 3/3 Rounded bump Diego Garcia (Chagos) 6.2 (18)20 3/8.=3/3'-.3/3.= 378 Rounded bump Diego Garcia (Chagos) 6.5 (21)24 3/3: 3/3: ~ 3/4 AA Rounded bump ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 Order Scolopendromorpha Family Cryptopidae Genus Cryptops Leach, 1815 Cryptops cf. japonicus Takakuwa, 1934 Fig. 2 Examined material. 35 specimens: NHMUK015991425, 4 specimens, Black Path, Picard, Summer 1975, leg. V. W. Spaull; NHMUK015991429, 1 specimen, 230 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) A —— oS Figure 2. Cryptops cf. japonicus Takakuwa, 1934 A, B, D NHMUK015991433 A anterior margin of forcipular coxosternite, ven- tral view B Calyx of venom gland, lateral view C NHMUK015991430, coxopleuron of ultimate leg-bearing segment, lateral view D femur, tibia, and tarsus 1 of ultimate leg telopodite, lateral view E. NHMUK015991 429, pretarsus of leg pair 8, lateral view. Aldabra, 10.11.1973; NHMUK015991430, 6 specimens, Point Hodoul, Grande Terre, 22.03.1974; NHMUK015991431, 2 specimens, Ile. Malabar, 08.06.1974; NHMUK015991432, 18 specimens, Casuarina and Sideroxylon litter, Anse Ced- res, 12.02.1974; NHMUK015991433, 4 specimens, Esprit, 14.12.1974. Remarks. Cryptops cf. japonicus collected in Aldabra is an unexpected oc- currence for a species otherwise restricted to localities in southern Japan, the Korean peninsula, Manchuria (Takakuwa 1936) and Taiwan (Chao 2008). Despite the description of the morphologically similar C. doriae Pocock, 1891 from nearby localities in the Seychelles (Lewis 2007a), consistent diagnostic morphological characters separate these two populations. Most prominent- ly, specimens from Aldabra only have one pretarsal accessory spur on legs 1-20, > 1/2 the length of the pretarsus (Fig. 2E), in contrast to C. doriae from the Seychelles which has been described with two generally subequal, conspic- uous, accessory spurs that are much shorter relative to the pretarsus, on the same leg pair range (Lewis 2007a). The examined specimens agree with the description provided by Chao (2008) based on specimens collected in Taiwan, in which he notes the low number of coxopleural pores (9) in the immature (“lar- va”) stages, which overlaps with the condition of Aldabra specimens (Fig. 2C), ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 231 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) the presence of 4 setae along the anterior margin of the forcipular coxosternite (Fig. 2A), and the ovoid shape of the calyx of the venom gland (Fig. 2B). All specimens from Aldabra Atoll range from 3-10 mm and exhibit several traits characteristic of juvenile specimens including reduced number of pores on the coxopleuron, indistinct paramedian sutures on tergites, and a reduced number of tibial and tarsal saw teeth (Fig. 2D). Following clearing, no spermatozoa or 00- cytes could be observed in the posterior trunk of specimens. Without additional sampling to confirm the condition of adult specimens it is not possible to com- ment on the presence or absence of sexually mature adults in the present sample. Introduced populations of C. doriae have been described bearing similar neotenic characteristics (reduced body size, number of coxal pores, number of saw teeth), even in sexually mature adults (Lewis 2007b), potentially explaining the morphol- ogy of the Aldabra specimens in light of possible introduction to the atoll. Never- theless, in the absence of additional material and molecular data, our assignment to Cryptops japonicus is only tentative. The status and relationships of different populations identified as C. doriae and related taxa remains to be clarified. Cryptops mauritianus Verhoeff, 1939 Fig. 3 Examined material. NHMUK015991434, 1 specimen, Sideroxylon litter, Anses Coco & Porche, Aldabra, 03.12.1974, V. W. Spaull. leg. Remarks. Cryptops mauritianus has been described by Verhoeff (1939) from Mauritius. Subsequent taxonomic revision of Mauritian Cryptops species (Lewis 2002) completed the summary original description and provided additional illus- trations of material from near the type locality. In the singular Aldabra specimen examined, the presence of minute accessory spurs on the pretarsus of leg pairs 1-20 (Fig. 3G) clearly delimits it from all other Cryptops species known from localities in the Western Indian Ocean (Lewis 2011). Subsequent re-description by Lewis (2002) provided data on an immature (“adolescens”) specimen which matches the condition of the material presently examined (body length 8 mm), allowing us to confidently refer it to C. mauritianus. Cryptops nigropictus Takakuwa, 1936 Fig. 4 Examined material. 12 specimens: NHMUK015991424, 3 specimens, Black Path, Picard, Aldabra, Summer 1975; NHMUK015991 426, 3 specimens, Calophyllum lit- ter, Takamaka, Grande Terre, 14.01.1975; NHMUK015991427, 2 specimens, Sid- eroxylon litter, Au Parc, Aldabra, 14.02.1975. V. W. Spaull leg; NHMUK015991428, 1 specimen, Picard, Aldabra, 28.02.1975; NHMUK015991445, 1 specimen, Sid- eroxylon litter, Ile. Michel, Aldabra, 28.03.1975; NHMUK015991446, 2 speci- mens, Casuarina litter, Picard, Aldabra, 08.05.1974. Remarks. Taxonomic revision of Cryptops species belonging to the “hortensis group” identified C. decoratus Lawrence, 1960, C. melanotypus Chamberlin, 1941, and C. nigropictus Takakuwa, 1936 as a potential spe- cies complex, raising doubts on the taxonomic validity of many morpholog- ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 232 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) 0.1 mm WW LO'O LULU SQ) 0.07 mm Figure 3. Cryptops mauritianus Verhoeff, 1939. NHMUK015991434 A anterior margin of forcipular coxosternite, ventral view B calyx of venom gland. Left forcipule (top), right forcipule (bottom) C coxopleuron of ultimate leg-bearing segment, lateral view D Ultimate leg telopodite, lateral view E tibia and tarsal articles of ultimate leg, lateral view F tibial comb (top), tarsal comb of left ultimate leg (middle), tarsal comb of right ultimate leg (bottom) G pretarsus of leg pair 18, lateral view. ically similar species (Lewis 2011). A reliable diagnostic trait mentioned by Lewis (2011) is the presence of a single long pretarsal accessory spur in C. nigropictus (Fig. 4K), differentiating it from C. decoratus and C. melanoty- pus which bear two small pretarsal accessory spurs on leg pairs 1-20. The Aldabra specimens key out to C. nigropictus in the key provided within the ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 233 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) A 0.05 mm. ) RY !) LAA B 0). mm 0.25 mm 0.25mm Figure 4. Cryptops nigropictus Takakuwa, 19364, F, K, 1, B NHMUK015991 426 H, JNHMUK015991446 G NHMUK01 5991 424 C “Cryptops daszaki” holotype, lle aux Aigrettes, 19.10.1995 D, E paratype, lle aux Aigrettes, 18.10.1995 A, B, C, D, Eanterior margin of forcipular coxosternite, ventral view F calyx of venom gland, lateral view G, H coxopleuron of ultimate leg-bear- ing segment, lateral view I, J tibia and tarsus of ultimate leg telopodite, lateral view K pretarsus of leg pair 8, lateral view. ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 234 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) same article and agree with its revised description. Although originally de- scribed from East Asia, various nominal taxa recorded from East Africa and islands of the Indian Ocean (including Mauritius and Rodrigues) have been placed in synonymy with C. nigropictus (Lewis 2011). Cryptops daszaki Lewis, 2002 was described from several localities in Mau- ritius. Lewis (2002) remarked on the small body size (4.5-7.5 mm) of sexually mature specimens and their apparent juvenile characteristics (low number of coxal pores, reduced setation). The specimens collected from Aldabra with a similar body length to specimens assigned by Lewis to C. daszaki overlap in morphology with respect to the usual lack of subcuticular dark pigment, the number of setae on the anterior margin of the forcipular coxosternite (2 or 3 on each side in C. daszaki, 3 or 4 on each side in C. nigropictus; Fig. 4C versus Fig. 4A, B, D, respectively), the relative length of the single pretarsal accessory spur on legs 1-20 (> 1/2 the length of the pretarsus) and the number of coxal pores (5 or 6inC. daszaki, 4—9 in C. nigropictus) but show greater variation than C. daszaki in the number of saw teeth on the tibia (3-5) and tarsus (2 or 3). The number of tibial and tarsal saw teeth is known to be intraspecifically variable in Cryptops (lorio and Geoffroy 2003; Lewis 2009) and scales allometrically with body size, as exemplified by the largest C. nigropictus specimen in our sample (12 mm, NHMUK015991426; Fig. 41). The only other diagnostic trait given by Lewis (2011) separating C. daszaki and C. nigropictus is the position of the setae near the anterior margin of the forcipular coxosternite, being described as “marginal” in C. daszaki and “sub- marginal” in C. nigropictus. Re-examination of the type specimens of C. daszaki revealed that these setae occupy a submarginal position (Fig. 4C), which may be misinterpreted as marginal depending on the orientation of the specimens. As both of the two putative diagnostic traits separating C. daszaki and C. ni- gropictus are fully encompassed by intraspecific variation within C. nigropictus and the latter species has been recorded from nearby localities (Lewis 2002), we consider C. daszaki a likely junior subjective synonym of C. nigropictus, from which it cannot be reliably distinguished by morphology alone. Similarly, the presence of subcuticular dark pigment cannot be reliably used to sepa- rate putative Cryptops species in Western Indian Ocean and Eastern African localities. This character is variable within our sample, with all specimens with the exception of NHMUK015991428 lacking dark subcuticular pigment. Dark pigmentation is also variable in the type specimens of Cryptops niloticus Lewis, 1967 (present in male allotype but absent in female holotype and in juvenile paratype), a species previously recorded from the Western Indian Ocean (Lewis 2002) but subsequently synonymised with C. nigropictus (Lewis 2011). Family Scolopendridae Genus Scolopendra Linnaeus, 1758 Scolopendra morsitans Linnaeus, 1758 Examined material. 52 specimens: NHMUK015991435, 33 juveniles; Casua- rina litter, Picard, Aldabra, 18.04.1974, leg. V. W. Spaull; NHMUK015991436, 1 specimen, Casuarina, Picard, Aldabra, 04.02.1974; NHMUK015991437, 1 speci- men, Ochna soil, Picard, Aldabra, 15.02.1974; NHMUK015991438; 2 specimens, ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 235 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) mixed scrub, Picard, 24.12.1974, NHMUK015991439, 1 specimen, Thespe- sia litter, Cing Cases, Aldabra, 15.11.1973; NHMUK015991441, 5 specimens, Black Path, Picard, 1975; NHMUK015991442, 1 specimen, Picard, 03.12.1973; NHMUK015991443, 1 specimen, Picard, 18.01.1974; NHMUK015991440, 4 specimens, Picard, 18.11.1974; NHMUK015991444, 1 specimen, Pitfall trap 5, 08.12.1974; NHMUK015991447, 1 specimen, South Island, Aldabra, 13- 20.03.1968, leg. B. Cogan & A. Hutson; NHMUK015991448, 1 specimen, Ile. Mi- chel, 02.1968, leg. B. Cogan & A. Hutson. Remarks. Specimens of S. morsitans Linnaeus, 1758 collected from mainland Africa, originally identified as Scolopendra amazonica Bicherl, 1946 overlap with the specimens from the Aldabra Atoll in several characters (Table 2) (Lewis 1966, 1967, 1968, 1969), but generally have a greater number of glabrous basal anten- nal articles, a character in which specimens from the Aldabra Atoll more close- ly match Indian specimens of S. morsitans previously assigned to S. amazonica (Jangi 1955, 1959). In the absence of newly collected material from which molec- ular data can be collected to evaluate possible interspecific delimitation between different population of S. morsitans, we assign material collected in the Aldabra Atoll to S. morsitans, following the conclusions on interspecific variation within this taxon reached by Wirmli (1975). Subsequent phylogenetic analyses of mo- lecular data for S. morsitans identified multiple lineages within this taxon (Joshi and Karanth 2011; Siriwut et al. 2016), potentially indicating the existence of a cryptic species complex as suggested by recent taxonomic review (Lewis 2010a). Until the global taxonomy of S. morsitans is interrogated using molecular data to establish if this is the case, we classify the examined material as S. morsitans. Additionally, specimens were compared to the original description of the morphologically similar and geographically proximate Scolopendra antanana- rivoensis Kronmiller, 2010. Material from the Aldabra Atoll did not exhibit the characters given as diagnostic for S. antananarivoensis, lacking a longitudinal median depression on sternite 21 and not having a distinctly more elongate coxopleural process (Kronmiller 2010). Table 2. Morphological variability in Scolopendra morsitans populations obtained from literature data (Lewis 1966, 1967, 1969) and examined specimens. Character Subadult and adult body length (mm) Number of antennal articles Number of glabrous antennal articles Complete paramedian sutures begin Lateral margination Coxopleural process spines Lateral coxopleural spine Leg 20 tarsal spur Scolopendra morsitans (India) May begin more anteriorly Scolopendra amazonica Scolopendra amazonica sensu Jangi (= Scolopendra sensu Lewis (= Scolopendra morsitans) (India) morsitans) (Africa) Scolopendra morsitans (Aldabra) 18-113 15-65 13-100 13-78 19 18-21 (17)18-19 Can be > 6 <6 BZ (3.5)4 Mostly T3 Mostly T2 TT2-4 Ts Generally last 5 tergites Last 2-15 tergites Last 4-10 tergites 4 2-6 (3)4-5 Present Present or absent Present or absent Present Present Absent Absent Absent ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 236 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) Order Geophilomorpha Family Geophilidae Genus Ribautia Brolemann, 1909 Ribautia cf. paucipes Attems, 1952 Fig. 5 Examined material. NHMUK015991467, 1 juvenile, Casuarina litter, Picard, Al- dabra, 10.12.1974, leg. V. W. Spaull. Remarks. The sexually immature specimen found in the present sample dis- plays all diagnostic characters that support its assignment to Ribautia, com- prising an elongate cephalic shield, lack of lappets on the first maxillae, the two halves of the second maxillary coxosternite being united by a sclerotised isthmus and the pleural sutures of the forcipular coxosternite being parallel to its lateral edge distally. A potentially novel ontogenetic observation is the incomplete separation of the two halves of the second maxillary coxosternite by an isthmus (Fig. 5B), as is a characteristic of Ribautia. Taking into account the very small size (length 7 mm) and sexual immaturity of the specimen, this may be a character state that becomes more conspicuous in older individuals. Beside the very low number of leg-bearing segments (37), which is shared with R. cf. paucipes reported from the Seychelles (Bonato and Minel- li 2010), the developmental stage of the present specimen does not allow for satisfactory evaluation of potential morphological differences between 0.02 mm LULU 1° wu TO 0.02 mm (a) a BS A\K /y \ Figure 5. Ribautia cf. paucipes Attems, 1952. NHMUK015991467 A head and forcipular segment, ventral view B anterior margin of second maxillary coxosternite, ventral view C calyx of venom gland, lateral view D ultimate leg-bearing and postpedal segments, ventral view. ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 237 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) Ribautia specimens described from Western Indian Ocean localities and type material of Ribautia paucipes described from the environs of Lake Kivu in Cen- tral Africa. Notably, the Aldabra specimen lacks evident denticles on the ante- rior margin of the forcipular coxosternite and the interior margin of the forcip- ular trochanteroprefemur, although these are clearly illustrated in the original description of R. paucipes (Attems 1952: fig. 203). The specimen also lacks conspicuous coxal organs or coxal pores, a character state not noted in R. cf. paucipes recorded from the Seychelles. Ontogenetic variation in the number of coxal pores has been well-documented for other geophilid centipedes (Hor- neland and Meidell 2009; Gregory and Barber 2010; Brena 2014; Stojanovic et al. 2020), increasing with body size at each postembryonic stage, and be- ing absent in comparably sized adolescens stages of some species (Gregory and Barber 2010). Additionally, the second maxillary pretarsus of the present specimen is markedly shorter and less acuminate than illustrated for R. pauci- pes from continental Africa. As the ontogenetic variation of the morphology of the second maxillae in Ribautia is not presently known and in other characters the specimen strongly resembles individuals described from the Seychelles, we maintain its presently assigned identity. Genus Tuoba Chamberlin, 1920 Hovanyx Lawrence, 1960, syn. nov. Mixophilus Silvestri, 1929, syn. nov. Remarks. The monotypic genus Mixophilus was erected by Silvestri (1929) to place a new species of geophilomorph sampled from riparian sites in Ma- dras (Chennai), Southern India. The original description of Mixophilus indicus includes illustrations of the head, forcipular apparatus and ultimate leg-bear- ing segment as well as detailed ecological observations concerning its pre- ferred microhabitats in the type locality. Subsequent physiological investiga- tions revealed a modified tracheal system, comprising possible adaptations to immersion for long periods of time (Rajulu 1970). An affinity to Henia and Chaetechelyne (Dignathodontidae) was suggested, however, within the same section Silvestri (1929) pointed to differences between Mixophilus and both of these genera in “the structure of the labrum” (tripartite in M. indicus but unipar- tite in Dignathodontidae), “the distribution of sternal pores” (transverse band in M. indicus, medial sub-circular/elliptical field in Dignathodontidae), in the last leg-bearing segment (telopodite composed of 7 articles in M. indicus but 6 in Henia), and in the elongation of the forcipular pretergite (longer in M. indicus than in Chaetechelyne). The structure of the labrum described and illustrated for M. indicus, comprising three conspicuous pieces, suggests a close affinity to other members of Geophilidae s. str. An estuarine habitat preference and multiple morphological characters show a near complete overlap between the diagnoses of Tuoba and Mixophilus (Table 3). Complete chitin lines in M. indi- cus may indicate intraspecific or interspecific variability which has been record- ed in Tuoba (Jones 1998) or artifacts of examination under light microscopy. In light of this re-evaluation of its original description, we propose reassignment of M. indicus to Tuoba considering the available data on its morphology, with Tuoba indica comb. nov. as the valid name for specimens on which its original ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 238 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) Table 3. Taxonomically informative invariant and variable morphological characters for the genera Hovanyx, Mixophilus, and Tuoba based on literature data. Characters in boldface represent putative morphological differences. Character Tuoba Chamberlin, 1920 Hovanyx Lawrence, 1960 Mixophilus Silvestri, 1929 Head shape Subquadrate Longer than wide Subquadrate Setation of clypeus Three pairs of setae medially, 10-11 setae Four pairs of setae medially, flanked by a group of 2-4 flanked by a group of 3 or 4 setae on each side setae on each side Medial piece of labrum. Anteriorly recurved (variable) Anteriorly recurved Anteriorly recurved Orientation of tubercles Side pieces of labrum With variable number of : Without tubercles or plumose tubercles or plumose setae setae Maxillae | lappets Absent. External corners with | Absent. External corners with Absent spiniform cuticular projections | spiniform cuticular projections Maxillae II pretarsus Simple, claw-shaped. Variably Simple, claw-shaped Simple, claw-shaped. Reduced reduced in size in size Forcipular coxosternite chitin lines | Complete or nearly complete | Vanishing before reaching the Complete condyles Denticle at the base of the Present Present Present tarsungulum Carpophagus structures Present = : Metasternal pore field shape Transverse band Transverse band. Divides on Transverse band (medially LBS XI-XIl constricted) Pretarsus of walking legs Distinctly elongate Distinctly elongate Distinctly elongate LLBS metasternite shape Wider than long, trapezoidal Wider than long, trapezoidal Wider than long, trapezoidal Coxal organs Multiple opening in single pit Absent Multiple opening in single pit LLBS pretarsus Simple, claw-shaped Simple, claw-shaped Simple, claw-shaped description was based. Consequently, we propose that Mixophilus is the junior subjective synonym of Tuoba syn. nov. Similarities between Tuoba indica comb. nov. and T. sydneyensis exist in elongation of the ultimate legs and reduction of the second maxillary pretarsus, however these characters are shared by sever- al species of Tuoba. The incomplete original description provided by Silvestri does not allow for definitive assignment of Tuoba indica comb. nov. to another species of Tuoba until the type material can be adequately re-described. Similarly, the genus Hovanyx was erected for the species Hovanyx waterloti, described in Lawrence's (1960) catalogue of Malagasy centipedes. An affinity to Dignathodontidae was again proposed based on similarities in the structure of the labrum, originally described for H. waterloti as composed of a single piece and bearing a small number of rudimentary, anteriorly oriented tubercles (“Labre [...] 4 dents pas trés distinctes, quatre courtes dents triangulaires di- rigées vers l’'avant, de chaque cété.”). The diagnosis of Hovanyx singled out the absence of coxal organs (‘...] différe de tous les autres membres de la sous-famille des Dignathodontine par l’absence de pores aux pattes termina- les.”) as the main distinguishing trait separating it from all other dignathodontid genera. However, both the original description and accompanying illustrations suggest a closer affinity to Geophilidae s. str., as multiple other morphological characters (shape of head, forcipular coxosternite, metasternal pore fields) are characteristic of the Geophilidae rather than the Dignathodontidae, and subse- quent taxonomic revision of both families placed Hovanyx under Geophilidae (Bonato 2011). Furthermore, the incomplete original description overlaps al- most entirely with that of T. sydneyensis, a wide-ranging geophilid encountered in littoral sites in the Seychelles (Bonato and Minelli 2010) and the Aldabra ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 239 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) Atoll (present records) close to the type locality of H. waterloti. Shared char- acters include range of leg bearing segment number (41-43), morphology of the labrum, condition of chitin lines on the forcipular coxosternite (incomplete) (Table 3). The only putative differences between H. waterloti and T. sydneyensis are the unipartite labrum and absence of coxal organs in the former (Table 3). These may however be unreliable characters because of inadequate docu- mentation, as published illustrations of Tuoba specimens and the material here illustrated show great variability in the orientation and shape of the labral piec- es, which depending on the degree of sclerotization seen in the specimen and shape and number of denticles on the side pieces, may resemble the labrum of Dignathodontidae under light microscopy. The absence of coxal organs has not been previously reported in any species of Tuoba, however this character shows extensive ontogenetic plasticity within Geophilidae (Horneland and Meidell 2009; Gregory and Barber 2010; Tuf and Danyi 2015; Peretti and Bonato 2016; Stojanovic et al. 2020) and the small size of the holotype and only known specimen as well as variations in clearing and position of the coxopleuron may render the pit inconspicuous. Tuoba sydneyensis (Pocock, 1891a) Figs 6-10 Geophilus (Bothrogeophilus) lemuricus Verhoeff, 1939, syn. nov. Hovanyx waterloti Lawrence, 1960, syn. nov. Examined material. 19 specimens: NHMUK015991475, 1 specimen, South Is- land, Aldabra, 13-—20.03.1968, leg. B. Cogan & A. Hutson; NHMUK015991476, 1 juvenile, Cinq Cases/Point Hodoul Arga, 27.03.1974, leg. V. W. Spaull; NHMUK015991477, 19, inside fallen dead coconut tree, Picard, Aldabra, 23.02.1974, leg. V. W. Spaull; NHMUK015991478, 39, 14, 1 juvenile, Cing Cas- es, Aldabra, 10-17.03.1974, leg. V. W. Spaull; NHMUK015991479, 19, Cyper- us ligularis soil and litter, Dune Patates, Aldabra, 05.06.1974, leg. V. W. Spaull; NHMUK015991480, 14, 1 incomplete, Mystroxylon and Dracaena litter, Gionnet, 03.12.1974, leg. V. W. Spaull; NHMUK015991481, 19, 1, Casuarina litter, Picard, Aldabra, 08.05.1974, leg. V. W. Spaull; NHMUK015991482, 1’, Pemphis litter, Dune D’Messe, Grande Terre, 29.01.1975, leg. V. W. Spaull; NHMUK015991 483, 24, 19, Casuarina, Picard, 27.12.1974, leg. V. W. Spaull.; NHMUK015991 484, 2°, Suriana litter near Point Hodoul, Grande Terre, 22.03.1974, leg. V. W. Spaull; 1 juvenile, Cocos litter, Esprit, Aldabra, 14.12.1974, leg. V. W. Spaull. Remarks. Tuoba sydneyensis has previously been reported from the Sey- chelles (Bonato and Minelli 2010). Material presently described from the Al- dabra Atoll, as well as specimens collected on Serpent Island (Mauritius) and other islands of the Seychelles (Bonato and Minelli 2010) can be assigned as conspecific on the basis of the low number of leg-bearing segments (41-45), elongation of the antennal articles (Fig. 6A) and in the shape of the ultimate leg-bearing segment metasternite (Jones 1998). The only other species of Tuoba hitherto reported from the East African coast are Tuoba poseidonis Verhoeff, 1901 (Zapparoli 1990) and Tuoba sudanensis Lew- is, 1963. The small number of diagnostic characters separating these two species ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 240 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) Figure 6. Tuoba sydneyensis (Pocock, 1891a). NHMUK015991484 A head and antennae, dorsal view B cephalic shield, dorsal view C antennal article XIV, dorsal view. casts doubt on the validity of T- sudanensis or on the Somali record of T. posei- donis. Examination of additional material from the East African coast would be necessary to elucidate the diversity of Tuoba in Northern and Eastern Africa. Both species can be reliably differentiated from T. sydneyensis in the Western Indian Ocean by the larger number of leg-bearing segments (51-53 in T. sudanensis compared to 41-45 in T. sydneyensis), greater elongation of the telopodal lappets of the first maxillae (30% of the length of the telopodite as illustrated for T. sudanensis; compared to minute in T. sydneyensis (Fig. 8B)), shape of the carpophagus structure (with a distinct median “hump” in T. sudanensis; lacking any “hump” medially in T. sydneyensis), point of the midbody transition (sternites 20-22 in T. poseidonis and T. sudanensis; ster- nites 14-15 in T. sydneyensis) and the shape of the ultimate leg-bearing seg- ment metasternite (1.6 x wider than long as illustrated for male T. sudanen- sis; 1.9 x wider than long in male T. sydneyensis, Fig. 10A). The combination of diagnostic characters presented clearly unify the Western Indian Ocean populations of Tuoba under one morphospecies, closely matching the de- scription of T. sydneyensis, which is distinct from Tuoba species recorded in continental Eastern and Northern Africa. Verhoeff (1939) described Geophilus (Bothrogeophilus) lemuricus from La Ponce, Mauritius and remarked that it is closely related to (“[...] nahe verwandt [...!”) Geophilus carpophagus Leach, 1815. Although incomplete, the description provides several characters that allow reliable assignment to T. sydneyensis, with which it agrees in number of leg-bearing segments (47 in G. /emuricus, ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 241 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) Figure 7. Tuoba sydneyensis (Pocock, 1891a) B, C NHMUK015991478 A NHMUK015991484 B clypeus and labrum, ven- tral view C labrum, ventral view. 41-45 in T. sydneyensis from the Western Indian Ocean), the apical claw of the second maxillae being reduced in size and not overtaking surrounding se- tae in length (“[...], Uberragt nicht die Nachbarborsten.”) (Fig. 8B, C), and in the arrangement of the coxal organs on the coxopleuron of the last leg-bearing segment, which are arranged into a rosette opening in a pouch near the edge of the metasternite (“[...] neben dem Endbeinsternit miindet eine Tasche und in diese eine Rosette von Driisen”) (Figs 9A, 10A). We consider G. lemuricus to be a junior subjective synonym of T. sydneyensis, which has been subsequently collected from Mauritius (NHMUK015991411). Family Mecistocephalidae Genus Mecistocephalus Newport, 1843 Mecistocephalus angusticeps (Ribaut, 1914) Fig. 11B Examined material. 11 specimens: NHMUK015991461, 1, Pandanus tectorius soil and litter, Cing Cases, 24.03.1974, leg. V. W. Spaull; NHMUK015991462, 19,2 km N of Cing Cases, 11.03.1974, leg. V. W. Spaull; NHMUK015991463, 6 juveniles, Sideroxylon litter, Cing Cases, 10-17.03.1974, leg. V. W. Spaull; NHMUK015991464, 12, Pandanus tectorius soil and litter, Cinq Cases/Point Hodoul, 27.03.1974, leg. V. W. Spaull; NHMUK015991465, 1, Gionnet, Aldab- ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 242 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) A 0.1 mm WW S00 Wu SZ’0 WU L"O Figure 8. Tuoba sydneyensis (Pocock, 1891a). NHMUK015991484 A Mandible, lateral view B First and second maxillae, ventral view C second maxillary article III and pretarsus, ventral view D forcipular segment, ventral view E right forcipule, ventral view. ra, 03.12.1974, leg. V. W. Spaull; NHMUK015991466, 1<, Casuarina, Aldabra, 04.02.1974, leg. V. W. Spaull. Remarks. Previously recorded from multiple localities near the East African Coast and the Western Indian Ocean (Ribaut 1914; Bonato and Minelli 2010; Popovici et al. 2024), making natural dispersal a likely explanation for the pres- ence of M. angusticeps in Aldabra. Presently examined specimens are morpho- logically indistinguishable from conspecifics recorded from the Seychelles and the Chagos Islands. Although not previously noted, the forcipular cerri were found to be absent in examined specimens (Fig. 11B). In combination with other diagnostic characters (47 leg-bearing segments, conspicuous medial re- duction in clypeal reticulation, large distal trochanteroprefemoral denticle and sternal sulcus not furcate), this allows for easy separation from other syntopic Mecistocephalus spp. within its range. ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 243 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) 0.1 mm Figure 9. Tuoba sydneyensis (Pocock, 1891a) A NHMUK015991484, female ultimate leg-bearing and postpedal seg- ments, ventral view BNHMUK015991478, juvenile ultimate leg-bearing and postpedal segments, ventral view. ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 244 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) Figure 10. Tuoba sydneyensis (Pocock, 1891a), NHMUK01591483 A male ultimate leg-bearing and postpedal segments, ventral view B male gonopods, ventral view C penis, ventral view. Mecistocephalus lohmanderi Verhoeff, 1939 Fig. 11A Examined material. 2 specimens: NHMUK015991459, 19, Black Path, Picard, Aldabra, Summer 1975, leg. V. W. Spaull; NHMUK015991460, 16, Casuarina lit- ter, Picard, 10.12.1974, leg. V. W. Spaull. Remarks. Originally described from Mauritius (Verhoeff, 1939), M. lohman- deri has been found on other Western Indian Ocean Islands (Bonato and Minel- li 2010; Popovici et al. 2024). Similarly to M. angusticeps (Fig. 11B), examined ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 245 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) 0.5mm WLU S°0 Figure 11. A Mecistocephalus lohmanderi Verhoeff, 1939, NHMUK015991460, forcipular segment, dorsal view B Mecis- tocephalus angusticeps (Ribaut, 1914), NHMUK015991466, forcipular segment, dorsal view. specimens lack forcipular cerri (Fig. 11A). Presently examined specimens are most similar to M. lohmanderi specimens collected from the Diego Gar- cia atoll (Popovici et al. 2024). Adults (female 34 mm body length, male 34 mm body length) in the Aldabra sample lack dark subcutaneous pigment patches and maintain the characteristic clypeal setation pattern described in M. lohmanderi from other localities (Bonato and Minelli 2010; Popovici et al. 2024). Similarly, specimens assigned to M. insularis described from So- cotra (Lewis and Wranik 1990) match all diagnostic characters outlined for M. lohmanderi, and can be referred to this taxon, thus extending its range to island localities in the Northwestern Indian Ocean. Records of large adult specimens (70-91 mm) assigned to Mecistocephalus insularis from the Arabian Peninsula (Lewis 1996) are fully consistent with the revised description of Mecistocephalus glabridorsalis Attems, 1900 from the Sey- chelles (Bonato and Minelli 2010) and are almost certainly misidentified M. gla- bridorsalis. In particular, the clypeal morphology illustrated for specimens from Saudi Arabia shares the presence of a small non-areolate insula anterior to the plagulae with specimens from the Seychelles and the arrangement of setae in a transverse line on the areolate part of the clypeus. This morphology has hitherto only been recorded in M. glabridorsalis and M. punctifrons Newport, 1843 (Bon- ato and Minelli 2004), casting further doubt on the true identity and distribution of M. insularis. Insufficient data on morphological variability in M. lohmanderi and the uncertain status of M. insularis records from past literature prevent further inferences on the taxonomic validity and relationships between these species. ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 246 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) Family Oryidae Genus Orphnaeus Meinert, 1870 Nycternyssa Crabill, 1959 syn. nov. Orphnaeus dekanius Verhoeff, 1938 Figs 12-16, 18D-H Examined material. 6 specimens: NHMUK015991469, 1¢, 40 mm, 73 leg-bearing segments, Picard, Aldabra, 08.10.1974; NHMUK015991470, 123, 35 mm, 75 leg-bearing segments, Picard, 09.04.1974; NHMUK015991471, 19, 24 mm, 81 leg-bearing segments, Grande Terre, Aldabra, 03.1974, leg. J. Wilson; NHMUK015991472, 1 juvenile, 13 mm, 81 leg-bearing segments, Grande Terre, Aldabra, 05.1974, leg. J. Wilson; NHMUK015991473, 14,37 mm, 75 leg-bearing segments, Pandanus litter, Aldabra, 22.03.1974; NHMUK015991474, 19, 51 mm, Takamaka (Anse Takamaka), 23-27.02.1968, leg. B. Cogan & A. Hutson. Diagnosis. Medium to large size Orphnaeus species, with 73-81 leg-bearing segments and variable but generally present longitudinal bands of dark pig- ment flanking the central vessel. Mandible with three or four pectinate lamellae. First maxillae with both telopodal and coxosternal lappets present and uniar- ticulate telopodite. Second maxillary pretarsus spatulate, fringed by acuminate hyaline projections. Posterior trunk metasternites with paired pore fields at the posterior end. Pore fields on posterior metasternites, procoxae and metacoxae bordered by dense groups of setae-like projections. Female gonopods uniartic- ulate, medially overlapping, with angled, rounded external margin. Description. Head and antennae. Cephalic plate with broadly rounded an- terior margin and straight posterior margin, overlapping the forcipular tergite. Head approximately as broad as long (NHMUK015991473) to 1.2 x broader than long (NHMUK015991474). Antennae approximately 2.5 x longer than head, weakly tapering distally (Fig. 12A). In older specimens, the tapering of antennae and dorsoventral compression of proximal antennal articles are more clearly visible (Fig. 12C). Antennal article XIV 1.9 x longer than the penultimate, with two clusters of sensilla basiconica arranged in lateral pits. Small, spear- like sensilla present at apical end. Mandibles. Of typical aspect for the genus (Fig. 13C, D). Four conspicuous pectinate lamellae evident, arranged concentrically around distal edge. Prox- imal to the outermost lamella, isolated projections resembling those on the lamellae are present. Two minute sensilla present laterally. Labrum and clypeus. Labrum of typical oryid aspect, with short hairlike hyaline projections on middle part (Fig. 13B). Lateral parts incompletely separated from middle part and clypeus by evident sutures. Clypeus with two pairs of postanten- nal sensilla, a median field of sensilla spanning its mediolateral axis and one pair of prelabral sensilla posterior to these (Fig. 13A). Polygonal reticulation evident. Maxillae. First maxillae with apically rounded, short coxal projections, bearing 8-11 sensilla (Fig. 14A). Telopodite broadly rounded, uniarticulated, of similar size to the coxal projection, bearing 7-12 sensilla. Both telopodal and coxoster- nal pairs of lappets present, with distinct spinous reticulation (Fig. 14C). Cox- osternal pair of lappets completely obscured by second maxillary telopodite in ventral view. Second maxillary coxosternite with shallow, concave, rounded an- terior margin bordered by a row of trichoid sensilla and two groups of trichoid ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 247 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) WW S70 Figure 12. Orphnaeus dekanius Verhoeff, 1938 A, B, D NHMUK015991470 A, C NHMUK015991474. Head and antennae, dorsal view B antennal article XIV, dorsal view D forcipular segment, ventral view. sensilla proximally (Fig. 14A). Metameric pore conspicuous, surrounded by scle- rotised rim. Telopodal articles stout. Pretarsus with spine-like projections around entire exterior margin and two pores opening on its dorsal surface (Fig. 14D, E). Forcipular segment. Forcipular tergite 3.2 x broader than long. Exposed surface of forcipular coxosternite 2.2 x broader than long (Fig. 12D). Chitin lines absent. Anterior margin rounded, deeply concave. Pleural sutures strongly converging posteriorly. Trochanteroprefemur ~ 1.5 x broader than long, with evidently round- ed external face. Tarsungulum stout, large, entirely covered by anterior edge of cephalic plate, with smooth inner concavity. Opening of venom gland channel im- mediately proximal to tip of tarsungulum. All forcipular articles without denticles. Trunk. Last five or six trunk metasternites with two posteriorly located pore fields (Fig. 15D), anteriorly bordered by dense clusters of hairs (Fig. 15A). All other trunk metasternites with four pore fields, two anterior and two posterior, of equal size on metasternites 2—46, the anterior gradually decreasing in size until ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 248 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) 0.1 mm = = ae = 5 Af ra © ‘ / 0.05 mm Figure 13. Orphnaeus dekanius Verhoeff, 1938. NHMUK015991470 A clypeus and labrum, ventral view B labrum, ventral view C, D Mandible, lateral and dorsal views. ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 249 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) LU 10 LULU S00 Figure 14. Orphnaeus dekanius Verhoeff, 1938 NHMUK015991470 A first and second maxillae, ventral view B first max- illary right telopodal lappet, ventral view C first maxillary left telopodal and coxosternal lappets, dorsal view D, E left and right second maxillary pretarsi, ventral view. ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 250 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) WW G0’ eX aS [otk Vig ; i av a Airgas SS AN C™ N AN Qh i ayy Sys PAS AAA ets yt i rl RD MAN W. eee } wh Aaah it x Soe ees ooh ANAS ' Aaa) <8 4 ren WW GC'0 Figure 15. Orphnaeus dekanius Verhoeff, 1938 A, D NHMUK015991474 B, C NHMUK015991470 A metasternites of leg-bearing segments 79 — 80, ventral view B walking leg, pair 74, lateral view C Walking leg pair 74 pretarsus, lateral view D Leg-bearing segments 79 — 81, ventral view. disappearing on metasternites 74-76. One single row of paratergites present beginning from the second leg-bearing segment, becoming very conspicuous on the eighth leg-bearing segment. General setation of sclerites sparse. Ultimate leg-bearing and postpedal segments. Ultimate leg-bearing segment metasternite variably trapeziform, 2.3 x broader than long (Fig. 16A, B). Pos- terior edge with dense field of hairs and occasionally small clusters of pores. Coxopleuron stout, without coxal organs. Telopodite of ultimate leg-pair only ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 251 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) Figure 16. Orphnaeus dekanius Verhoeff, 1938 A, C, DNHMUK015991471 B, ENHMUK015991470 A, B ultimate leg-bear- ing and postpedal segments, ventral view C ultimate leg pair metatarsus, lateral view D female gonopods, ventral view E male gonopods, ventral view. moderately inflated in both males (Fig. 16B) and females (Fig. 16A), with dense fields of hairs present on the ventral side of all articles. Pretarsus absent (Fig. 16C). Metatarsus with small hair-like projections at its apical edge. Female gonopods usually uni-articulated (Figs 15D, 16D), occasionally with an anterior notch or asymmetrical articulation. Lateral edge smooth, rounded, forming strongly acute angle with posterior edge of first genital sternite. Male gonopods biarticulated, bearing 16 setae (Fig. 16E). ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 252 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) Remarks. Orphnaeus dekanius was originally described from Trivandrum (Thiruvanathapuram), India (Verhoeff 1938), and maintained as a valid species under Orphnaeus until its reassignment to Nycternyssa, justified by the descrip- tion of the female gonopods as “uni-articulate”. A detailed re-evaluation of the status of Nycternyssa is provided below. Subsequent to its original description, there is no evidence that other specimens had been assigned to either O. dekanius or N. dekania prior to recent records from the Chagos Archipelago (Popovici et al. 2024). Based on morphology alone, the present specimens are considered conspecific with those collected in the Chagos Archipelago and match the original descrip- tion of O. dekanius. The most salient diagnostic trait allowing for reliable differentiation of O. dekanius from O. brevilabiatus is the presence of dense clusters of minute setae bordering the posterior pore fields of the former (Fig. 15A). Although this character was illustrated by Verhoeff in the original description of O. dekanius (Verhoeff 1938: tafel 8, fig. 61), no mention was made of it in the text of the description. It is unambiguously shared by all specimens studied here from near the type locality of O. dekanius (speci- mens from Sri Lanka listed above under “Specimen data”), and from other localities in the Western Indian Ocean, and is consistent in both sexes in specimens. Setae are sparse in the smallest studied Aldabra specimen (13 mm) but are clustered by a body length of 24 mm. As these clusters of setae are completely absent in specimens identified as O. brevilabiatus from near its type locality in Myanmar (Fig. 17) and other localities in mainland and maritime Southeast Asia, we maintain the validity of O. dekanius and O. bre- vilabiatus even in light of the variability of female gonopodal articulation in the latter (Fig. 18A-C). The incomplete description of Orphnaeus meruinus Attems, 1909 does not allow for reliable separation from O. dekanius, given the inconsistencies in how diagnostic characters for these two taxa are cod- ed in past literature. One salient difference from all O. dekanius specimens in our sample is the greatly inflated ultimate leg telopodite in males assigned to O. meruinus collected in Oman (Lewis and Gallagher 1993), a character that separates these species even when accounting for body size. Family Schendylidae Genus Ityphilus Cook, 1899 Ityphilus cf. taeniaformis (Lawrence, 1960) Figs 19-23 Examined material. NHMUK015991468, 14, Aldabra, 21.03.1974., V. W. Spaull leg. Description. Head and antennae. Antennae conspicuously claviform in shape, medially weakly geniculate, with articles IX-XIV widened (Figs 19A, 20A, C). Articles IX and XIII with clusters of type c sensilla (sensu Pereira 2017) on the distal edge of the dorsal side (Fig. 19C, D). Article XIV with two lateral clusters of sensilla basiconica and a small number of spear-shaped sensilla at its apex (Fig. 19B). Head approximately as wide as forcipular tergite, 1.14 x broader than long. Curved sulcus near posterior margin. Cha- etotaxy of head as in Fig. 20B. ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 253 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) 0.5mm 0.25mm Figure 17. Orphnaeus brevilabiatus (Newport, 1845). NHMUK015991423 A leg bearing segments 77 — 79, ventral view B metasternite of leg-bearing segment 78, ventral view. ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 254 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) Figure 18. Female gonopods of Orphnaeus Meinert, 1870 species A, B, C Orphnaeus brevilabiatus (Newport, 1845). (A = NHMUK015991423 (Myanmar), B = NHMUK015991421 (Thailand), C = NHMUK015991420 (Singapore)) D, E, F, G, H Orphnaeus dekanius Verhoeff, 1938. (D = NHMUK015991415 (Sri Lanka), E = NHMUK015991413 (Sri Lanka), F = NHMUK015991417 (Singapore), G = NHMUK015991418 (Singapore), H = NHMUK015991416 (Sri Lanka)). Scale bars: 0.1 mm. Mandibles. Dentate lamella with seven denticles, only six conspicuous in lat- eral view. Pectinate lamella with approximately 22 hyaline projections (Fig. 21A). Labrum and clypeus. Clypeus with a pair of postantennal setae, a clus- ter of seven medial setae and one prelabral seta. Lateral pieces of labrum narrow, conspicuously sclerotised, lacking any fringes or projections. Medial piece contiguous with clypeus, poorly sclerotised, membranous and lacking conspicuous hairs or projections. Maxillae. First maxillae with evident, triangular coxal projections, each bear- ing one sensillum. Telopodites bearing one sensillum each, conspicuously larg- er than coxal projections and partly covering them (Fig. 21B). Lappets absent. Second maxillary coxosternite with evident but incomplete medial suture, ex- tending for half of its length (Fig. 21C). Each side of the suture bearing one sensillum. Telopodite stout, terminating in large pretarsus. Second maxillary pretarsus spatulate, lateral edges densely fringed (Fig. 21D, E). ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 255 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) UW S0'0 0.1 mm Figure 19. Ityphilus cf. taeniaformis (Lawrence, 1960). NHMUK015991468 A right antenna, dorsal view B antennal article XIV, ventral view C clusters of type c sensilla on antennal article XIII (top) and IX (bottom), dorsal view. Forcipular segment. Exposed face of forcipular coxosternite 2.2 x broader than long (Fig. 22A). Chitin lines present, reaching the condyles. Forcipular trochanteroprefemur 1.25 x longer than broad. Calyx of venom gland elon- gated, ovoid in shape (Fig. 22C). All forcipular articles without denticles (Fig. 22B). Internal margin of tarsungulum smooth. Extended, forcipules do not reach the anterior margin of the head. Trunk. 75 leg-bearing segments. Pore fields located on raised areas in the middle of all metasternites excluding those of leg-bearing segments 1, 74, and 75. Shape of pore field oval, medially constricted and anteriorly bordered by a line of setae (Fig. 22D). Colour of pore field bluish grey, conspicuously pigment- ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 256 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) 0.25mm Figure 20. Ityphilus cf. taeniaformis (Lawrence, 1960) NHMUK015991468 A head and antennae, dorsal view B cephalic shield, dorsal view C right antenna, ventral view. ed relative to surrounding cuticle. Despite the faded colour of the ethanol-pre- served specimen, pigmentation of the pore field is conspicuous and the trunk is generally greenish grey in appearance. Ultimate leg-bearing and postpedal segments. Intercalary pleurites separated from ultimate pretergite by evident sutures. Ultimate metasternite trapezoidal, 1.3 x longer than broad. Coxopleura each with two distinct coxal organs, partially covered by the ultimate metasternite (Fig. 23A). Entire ventral side of ultimate leg-bearing segment covered in setae. Ultimate leg telopodite composed of sev- en articles, all distinctly thickened. Pretarsus absent (Fig. 23B). Metatarsus with a small spine subapically. Intermediate sternite indistinct. First genital sternite with straight posterior margin. Gonopods uni-articulate, flanking penis (Fig. 23C). ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 257 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) 0.2 mm Figure 21. Ityphilus cf. taeniaformis (Lawrence, 1960). NHMUK015991468 A mandible, lateral view B first and second maxillae, ventral view C anterior margin of second maxillary coxosternite, ventral view D second maxillary pretarsus left (top), right (bottom), ventral view. Remarks. The taxonomy of /typhilus remains largely unresolved, especially outside of South America, where different authors have disagreed on its relation to Ballophilus, alternatively considering it a different genus (Attems 1929) ora synonym of Ballophilus (Verhoeff 1939). Recent revision of the genus (Bonato et al. 2007) has maintained the distinction between /typhilus and Ballophilus, but remarked on the close relationship between the two, and on cases in which the presence of complete or nearly complete chitin lines is doubtful, such as in Ityphilus boteltobogensis (Wang, 1955), despite this character being predomi- nantly used to distinguish the two genera. Similarly, several Ballophilus species are described as bearing a cuticular thickening in the usual position of the chi- tin line (Demange 1963; Pereira et al. 1997), further confounding the utility of this character in taxonomy within the Ballophilidae. The presence of a median sulcus in the second maxillary coxosternite has been shown to be unreliable ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 258 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) WI |"O WIL S00 Figure 22. Ityphilus cf. taeniaformis (Lawrence, 1960). NHMUK015991468 A forcipular segment, ventral view B left forcipule, ventral view C calyx of venom gland, ventral view D leg-bearing segment 9, ventral view. in separating Ballophilus and Ityphilus. This character has been described in both Ballophilus (Ribaut 1914; Pereira et al. 1997) and /typhilus (Pereira et al. 2000), in some cases as incomplete (Br6lemann 1909) (Fig. 21B, C), and is not included in the most recent diagnosis of the latter genus (Bonato et al. 2007). Verhoeff (1939) described two Ballophilus species from Mauritius, B. law- rencei Verhoeff, 1939 and B. mauritianus Verhoeff, 1939. Both are known from single specimens but, according to their original description, compare closely with Ballophilus allauadi Ribaut, 1914 described from Eastern Africa. The Al- dabra specimen differs from these species in the presence of pore fields on all but the first and last two leg-bearing segment metasternites (contrasting with the absence of the pore field on the first and the last four metasternites). Addi- tionally, the distal end of the antenna of B. lawrencei is illustrated as markedly less clavate than observed for the Aldabra specimen. Despite this, the /typhilus specimen collected in Aldabra overlaps in the shape of the metasternal pore field, its position on a raised area and in the relative elongation of the ultimate ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 259 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) A R B 0.05 mm Sof uu eae = Ba ae 2 Lyf MY x NH LL AS Oi" ff ms ayy aa Figure 23. Ityphilus cf. taeniaformis (Lawrence, 1960). NHMUK015991468 A ultimate leg-bearing and postpedal seg- ments, ventral view B tip of right ultimate leg telopodite, ventral view C male gonopods, ventral view. leg-bearing segment telopodite and the number of leg-bearing segments with B. lawrencei. As all Mauritian Ballophilus species are known solely from their holotypes, it is not possible to comment on intraspecific variability that may account for the overlap in these traits. Ballophilus maldivensis Pocock, 1906, described from the Maldives, sim- ilarly resembles the Aldabra specimen in the shape of the metasternal pore ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 260 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) fields and their pigmentation. The incomplete original description did not al- low for comparison of any other putative diagnostic characters beside the number of leg-bearing segments (67 in the female holotype), which is lower than that of the Aldabra specimen (75 in a male). Re-examination of the ho- lotype (Fig. 24), the sole known specimen, revealed several features that fur- ther distance it from the Aldabra specimen and bring it closer to the currently accepted diagnosis of Ballophilus. Ballophilus maldivensis lacks chitin lines WW 7'O 0.2 mm Figure 24. Ballophilus maldivensis Pocock, 1906. BMNH #200555 A head and forcipular segment, ventral view B mag- nified view of forcipular coxosternite showing lateral pleurite and condyle, right side, ventral view C metasternite of leg-bearing segment 7, ventral view D ultimate leg-bearing and postpedal segments, ventral view. ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 261 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) or any cuticular thickenings near their position (Fig. 24A, B). Other important differences from /. cf. taeniaformis from Aldabra include the absence of the metasternal pore field from the last four leg-bearing segments and a more strongly enlarged ultimate leg telopodite (Fig. 24D). Lawrence (1960) described three species assigned to Ballophilus from Mad- agascar, of which Ballophilus taeniaformis Lawrence, 1960 overlaps in nearly all diagnostic traits with the Aldabra material examined, differing only in the num- ber of teeth on the dentate lamella of the mandible (7 in the Aldabra specimen compared to eight or nine in B. taeniaformis). Lawrence made no comment on the presence or absence of the chitin line on the forcipular coxosternite for the species he assigned to Ballophilus. As noted above, this character has been used to differentiate between the genera Ballophilus and Ityphilus, although its variability as discussed above and the lack of consensus on the status of Ityph- ilus at the time of Lawrence's original description of B. taeniaformis prompt us to refer the examined specimen from Aldabra to /typhilus. In nearly all characters examined, the singular specimen from Aldabra agrees with the description of Ityphilus melanostigma (Attems 1900) and the subse- quent redescription of this species from specimens collected in the Seychelles (Bonato and Minelli 2010). However, the greatly reduced number of leg-bearing segments (75) relative to the presently known range within /. melanostigma (95- 101) suggest specific distinction, as extensive variation in leg-bearing segment number is not generally known from other species of /typhilus. Other putative dif- ferences to the original description of |. melanostigma include the greater relative enlargement of the ultimate leg pair telopodites in the Aldabra specimen, relative to Ityphilus specimens illustrated from the Seychelles (Bonato and Minelli 2010). Discussion Composition and affinities of the Aldabran centipede fauna With the exception of Australobius inflatitarsis, all centipede species previously recorded from Aldabra are represented in the examined sample. This includes nine species new to the atoll, raising the total number of centipede species known from Aldabra to 12. The high abundance of L. tristani and Cryptops spec- imens in the examined sample is surprising given the lack of previous mentions of their presence on the atoll. Of the recorded species, half are shared with other islands of the Seychelles (Lewis 2010b; Stoev and Gerlach 2010) and approximately a third with both Mauritius and the Chagos Archipelago (Table 4; Verhoeff 1939; Lewis and Daszak 1996; Lewis 2002; Popovici et al. 2024). Some have a wide to near-cos- mopolitan distribution and have been recorded from mainland localities in East Africa (Cryptops nigropictus, Mecistocephalus angusticeps, Scolopendra morsitans; Ribaut 1914; Lewis 1969; Zapparoli 1990) and the Indian subcon- tinent (Orphnaeus dekanius, Scolopendra morsitans; Verhoeff 1938; Joshi and Karanth 2011) or have been recorded from coastal localities across the entire Indian Ocean (Orphnaeus dekanius, Tuoba sydneyenis; Bonato and Minelli 2010; Popovici et al. 2024). The new records of S. morsitans from Aldabra confirm its presence in the outer islands of the Seychelles, which was previously suggest- ed as likely despite the lack of known specimens (Lewis 2007a). ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 262 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) Table 4. Occurrence data of centipede species found on the Aldabra Atoll and in nearby areas. Asterisk indicates that morpho- logically similar congeners have been recorded from the locality but taxonomic revision is required to confirm their identity. Species Australobius inflatitarsis Lamyctes tristani Cryptops cf. japonicus Cryptops mauritianus Cryptops nigropictus Scolopendra morsitans Ityphilus cf. taeniaformis Mecistocephalus angusticeps Mecistocephalus lohmanderi Orphnaeus dekanius Ribautia cf. paucipes Tuoba sydneyensis Locality Chagos Madagascar Maldives Mauritius eSeyen ane roy Bice Archipelago (inner islands) (mainland) Not Not recorded Not recorded Not recorded Recorded Not recorded recorded Recorded Recorded Not recorded Not recorded* Not recorded Not recorded Not Not recorded Not recorded Not recorded Not recorded* Not recorded recorded Not Not recorded Not recorded Recorded Not recorded Not recorded recorded Not Not recorded Not recorded Recorded Recorded Recorded recorded Not Not recorded Not recorded Recorded Not recorded Recorded recorded Not Recorded Not recorded Not recorded Not recorded* Not recorded recorded Recorded Not recorded Not recorded Not recorded Recorded Recorded Recorded Not recorded* Recorded Recorded Recorded Not recorded Recorded Not recorded* Recorded Not recorded Not recorded Not recorded* Not Not recorded Not recorded Not recorded Recorded Not recorded recorded Not Recorded Not recorded Recorded Recorded Not recorded recorded Comparison with literature data and specimens from Mauritius in the NHM collection reveals that there are no centipedes endemic to Aldabra that are obviously diverging in morphology, and most species occur throughout the Western Indian Ocean islands. Overviews of diagnostic morphological characters given above and in previous surveys of Western Indian Ocean centipedes (Bonato and Minelli 2010; Popovici et al. 2024) strongly suggest that these populations may be closely related and part of a distinctive cen- tipede fauna unique to the Western Indian Ocean, characterised by strong dispersive potential rather than localised radiations within island groups. In the absence of molecular data that would allow for the interrogation of population structure as well as additional vetting of the proposed identifi- cations, and a more complete sample of centipedes from the Comoros and Mascarene island groups, these conclusions are only tentative and based on morphological examination of collected specimens. Similarly, the unre- solved taxonomy of the genera Cryptops, Ityphilus, and Lamyctes, particu- larly in the Western Indian Ocean, makes the apparent absence of L. tristani and Ityphilus cf. taeniaformis potentially doubtful from islands in which morphologically similar congeners have been recorded (Bonato and Minelli 2010; Stoev and Gerlach 2010), as future taxonomic revision may reveal that these populations are conspecific. Human-mediated introduction of centipedes to the Aldabra Atoll is diffi- cult to assess in light of the patchy information on its early human habitation (Stoddart 1971). The atoll was apparently uninhabited when a hydrographic survey was conducted in 1878, and settlement at Picard has been continuous but sparsely populated since 1890. Plantings of coconut, maize, sisal, cotton and other crops by the late 19" Century may account for some soil inverte- ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 263 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) brate introductions, including centipedes. Approximately 20% of plant species known from Aldabra are considered to be human-mediated introductions, de- spite a small area devoted to plantations (Renvoize 1971). Human introduc- tions of terrestrial invertebrates are generally poorly characterised across the Western Indian Ocean but have been recorded for the terrestrial gastropod Achatina fulica Férussac, 1821 (Peake 1971) and for terrestrial invertebrates in the Chagos Islands (https://chagosinformationportal.org/uploads/Terres- trial_Inverts_of_BIOT_21_03_18_1.xlsx), where 50% of recorded species are listed as introduced and 2.8% as potentially invasive, as well as suggested for the geophilomorph Tygarrup javanicus Attems, 1929 in the granitic inner islands of the Seychelles (Bonato and Minelli 2010). Being comprised of low to mid-elevation atolls, we consider the Chagos is- lands a useful guidepost for evaluating the possibility of centipede introduc- tions to Aldabra. The cryptic habits of most centipedes pose challenges to accurately assessing present diversity, let alone introduction potential. One isolated record in Feasibility Study for the Resettlement of the British Indian Ocean Territory Draft Report (https://data.parliament.uk/DepositedPapers/ Files/DEP2014-1543/Feasibility_Study_for_the_Resettlement_of_the_BIOT_ Draft_Report.pdf) attests to the import of soil from Sri Lanka on some islands, potentially representing a channel for centipede introduction. The morpho- logical similarity of Chagossian Rhysida longipes to Sri Lankan populations of this species (Popovici et al. 2024) well as its classification as a likely in- vasive (https://chagosinformationportal.org/uploads/Terrestrial_Inverts_of_ BIOT_21_03_18_1.xlsx) both indicate that introduction of this species to its Western Indian Ocean locality may have been human-mediated, in association with ballast or imported soil. The lack of any comprehensive historical record of such soil or vegetation transfers on Aldabra makes it difficult to ascribe the presence of any presently reported centipede species to human introduction. The status of Nycternyssa The genus Nycternyssa was erected by Crabill (1959), comprising the newly described Nycternyssa stheno Crabill, 1959 from Okinawa, Japan and four other taxa previously included in Orphnaeus, namely Nycternyssa conspersa (Verhoeff, 1937a), Nycternyssa dekania dekania (Verhoeff, 1938), Nycternys- sa dekania singaporiensis (Verhoeff, 1937b) and Nycternyssa eidmanni (Ver- hoeff, 1942). The original generic diagnosis states “Orphnaeus differs from Nycternyssa in that the female gonopod is divided into two distinct articles while it is simple and undivided in Nycternyssa”, with both genera being distin- guished from all other related oryids by having only one row of paratergites. No comment was made on characters that differ between male specimens of Orphnaeus and Nycternyssa. Additional characters listed in the diagno- sis of Nycternyssa, based on the description of N. stheno, are of ambiguous significance or shared by Orphnaeus. Crabill (1959) remarked that the first maxillary telopodite is “bipartite”, although the accompanying illustrations provided for N. stheno show it as unambiguously uniarticulated and other- wise identical in aspect to illustrations of this structure given for Orphnaeus (Fig. 14A). The antennae of Nycternyssa are described as barely attenuate and barely flattened proximally, in contrast to the diagnosis of Orphnaeus ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 264 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) in which the proximal end of the antennae is described as flattened (“platt- gedriickt”) (Attems 1929). This difference of appearance is likely a result of the time since collection and fixation medium of examined specimens, as variation within it can be observed within conspecific individuals from the same locality (Fig. 12A, C), and similar deformations of antennal shape in other geophilomorph taxa have likewise been treated as carrying dubious taxonomic value (Danyi and Tuf 2017; Popovici 2024). Subsequent mentions of Orphnaeus restate the biarticulate nature of the female gonopods as characteristic for the genus (Crabill 1968), or do not comment on this character specifically, but stress the uniarticulate female gonopods of Nycternyssa as diagnostic (Bonato 2011). Although this char- acter is regarded as fixed in Orphnaeus in the monograph on Geophilomor- pha compiled by Attems (1929), several earlier accounts of the type species, O. brevilabiatus, do not clearly illustrate biarticulate female gonopods (Sau- ssure and Humbert 1872; Haase 1887), incorrectly identify male specimens as female (Pocock 1889), do not mention the female gonopods in generic keys (Pocock 1893), or illustrate unambiguously uniarticulate female go- nopods (Pocock 1896, Saussure and de Zehntner 1902). Although the type material for this species is presumed to be lost, we examined and illustrate the two specimens mentioned by Pocock (1891b) from areas near the type locality in present-day Myanmar (Burma) (Figs 17, 18A—-C). One of the two specimens with provenance given as S. Tenasserim is an adult female with unambiguously uniarticulate gonopods (Fig. 18A). Notably Pocock (1891b) mentioned, “I have carefully compared the types of brevilabiatus and lineatus [...]”, implying that the condition of the gonopods in the female specimen from S. Tenasserim and that of the type material did not raise any doubt with respect to the identity of these specimens. As no other morphological characters of the near-topotypic specimens illustrated here suggest they are different from O. brevilabiatus as originally described, we consider that the articulation of the female gonopods is likely variable within Orphnaeus. Giv- en that there are no characters to support the reciprocal monophyly of both genera, Nycternyssa is here placed in junior subjective synonymy. Specimens identified as O. brevilabiatus from other localities displayed vari- ably biarticulate female gonopods, a state previously considered typical for this species (Fig. 18A—C). The present observations rise further doubt on the identi- ty of old records of O. brevilabiatus and agree with the reserve of other authors in continuing to treat O. brevilabiatus as a single, pantropical species (Wurmii 1974). As intraspecific variation in the articulation of the female gonopods is not known in Oryidae, it is possible that the specimens assigned to O. brevila- biatus are part of a complex of morphologically similar species. Acknowledgements We thank Jan Beccaloni (The Natural History Museum) for curatorial sup- port and providing access to the collection. We are grateful to Lauren Hughes (The Natural History Museum) for kindly providing lab space, and to Lucio Bonato (Universita degli Studi di Padova) and Pavel Stoev (National Museum of Natural History, Bulgarian Academy of Sciences) for construc- tive reviews of the manuscript. ZooKeys 1228: 225-273 (2025), DOI: 10.3897/zookeys.1228.143007 265 George Popovici & Gregory D. Edgecombe: Centipedes (Myriapoda, Chilopoda) of the Aldabra Atoll (Seychelles) Additional information Conflict of interest The authors have declared that no competing interests exist. Ethical statement No ethical statement was reported. Funding No funding was reported. Author contributions Conceptualization: GP GDE. Investigation: GP. Writing — original draft: GR GDE. Writing - review and editing: GDE, GP. Author ORCIDs George Popovici © https://orcid.org/0000-0003-0662-7472 Gregory D. Edgecombe ® hitps://orcid.org/0000-0002-9591-8011 Data availability All of the data that support the findings of this study are available in the main text. References Agnarsson I, Kuntner M (2012) The Generation of a Biodiversity Hotspot: Biogeography and Phylogeography of the Western Indian Ocean Islands. In: Anamthawat-Jonsson K (Ed.) Current Topics in Phylogenetics and Phylogeography of Terrestrial and Aquat- ic Systems. 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