The millipede family Paradoxosomatidae in the Philippines, with a description of Eustrongylosoma penevi sp.n., and notes on Anoplodesmus anthracinus Pocock, 1895, recorded in Malaysia and Sri Lanka for the first time (Diplopoda, Polydesmida)

Abstract The Philippine fauna of the family Paradoxosomatidae is reviewed and shown to comprise only 12 certain species (+ one dubious), definitely only a fraction of the real diversity to be expected from such a large tropical archipelago. Two new combinations are proposed: Euphyodesmus philippina (Nguyen Duc & Sierwald, 2010), comb. n. ex Desmoxytes Chamberlin, 1923, and Luzonomorpha polilloensis (San Juan & Lit, 2010), comb. n. ex Prionopeltis Pocock, 1895. The first representative of the large, basically Papuan genus Eustrongylosoma Silvestri, 1896 is described from Luzon, Philippines: Eustrongylosoma penevi sp. n. It differs from the other congeners in certain details of gonopod structure, as well as by the particularly long legs. Based on a restudy of the types of Strongylosoma luzoniense Peters, 1864, from Luzon, the species is shown to be a new senior subjective synonym of Helicorthomorpha orthogona (Silvestri, 1898), syn. n. This formally results also in Helicorthomorpha luzoniensis (Peters, 1864), comb. n. Anoplodesmus anthracinus Pocock, 1895 is illustrated and briefly redescribed, based on material from State Pulau Penang, Malaysia, which represents the first formal record of the species in that country. This species is also new to the fauna of Sri Lanka. A review of the Anoplodesmus species reported from Sri Lanka, nearly all of them dubious, is presented.


Introduction
The family Paradoxosomatidae is one of the largest and the most diverse in Diplopoda, and it has long been known to dominate the fauna of the Indo-Australian region (Jeekel 1968). However, only very few paradoxosomatid species have hitherto been recorded in the Philippines. Among them is an unidentified species of the large, mostly Papuan genus Eustrongylosoma Silvestri, 1896, reported by Hoffman (1978) from Mindanao. The present paper reviews and updates the paradoxosomatid fauna of the Philippines, including a description of the first Philippine Eustrongylosoma. The checklist presented below is highly condensed and skeletal on purpose, because a complete catalogue of the Diplopoda of the Philippines is in preparation (Korsós, pers. comm.). Here we also provide descriptive notes on the Southeast Asian species Anoplodesmus anthracinus Pocock, 1895, hitherto known only from Myanmar, but currently found in Malaysia and Sri Lanka as well.
Most of the material is housed in the collection of Diplopoda of the National Museum of Natural History, Sofia (NMNHS), with only a single paratype of Eustrongylosoma penevi sp. n. donated to the Zoological Museum, Moscow State University, Russia (ZMUM), as indicated hereafter.

Diagnosis
Most similar to E. exiguum Hoffman, 1978, from Papua New Guinea, and E. kuekenthali (Attems, 1897), from Borneo and Sulawesi, sharing the presence of a prominent distal spine on the gonopod femorite. Different from all congeners by the transverse orientation of the spine and noticeably long legs in the male (Hoffman 1978, Golovatch 1997.

Etymology
Honours our good friend and colleague Lyubomir Penev, biologist and founder of the Biodiversity Data Journal and Pensoft Publishers.

Notes
The species is hitherto known only from its type locality, Mt Polis Checkpoint on the road Banaue -Sagada (Fig. 3), where it was found close to a human settlement, under wooden plates and logs (Fig. 4).

Notes
This species was originally described from Yangon (= Rangoon), Myanmar (Pocock 1895). Attems (1937) synonymized it with Jonespeltis splendidus Verhoeff, 1936, from southern India, but Jeekel (1965) revalidated the latter species and returned A. anthracinus to its original scope. Furthermore, Jeekel provided very useful illustrations and a detailed redescription of the species, based on a part of the type series. Hoffman (1973) gave more illustrations of the gonopods, based on a paratype of A. kathanus (Chamberlin, 1921), from Katha, north of Yangon, Myanmar, and synonymized it with A. anthracinus.
Our record of A. anthracinus in the State of Pulau Penang, Malaysia considerably extends the range of this species to the south. The studied sample agrees well with the description provided by Jeekel (1965) and Hoffman (1973) in most characters (Figs 6f,  7), including humps in ♂ femora 5 and 6, as well as a process surmounting a hump in ♂ femur 7. Only slight variations have been noticed in the shapes of paraterga and sternal lobe between ♂ coxae 4. The same can be said about the samples from Sri Lanka which are also identified as A. anthracinus.
These are the first formal records of the species in Malaysia and Sri Lanka (Figs 8, 10). However, actually they might well represent introductions. In fact, in Malaysia the species was observed and collected in a highly agricultural and urbanized area, in a b Figure 7.
a: male leg 5, lateral view b: male leg 7, lateral view close proximity to experimental rice fields (Fig. 9), while in Sri Lanka, the collecting locality is a human settlement.  It is noteworthy that Sri Lanka hosts several formal species of Anoplodesmus, nearly all very similar to one another: • Anoplodesmus saussurii (Humbert, 1865), originally described from Sri Lanka, later recorded also in Fiji and Mauritius (Jeekel 1965, Jeekel 1972, Jeekel 1980a). The only meaningful difference from A. anthracinus is said to lie in the absence of a ventral hump in ♂ femur 5. However, given considerable variation in the presence or absence of this character, when such a hump in A. anthracinus can either be present in or absent from ♂ femur 4 (Attems 1937, Jeekel 1965, its status versus the older name A. saussurii is to be questioned. • Anoplodesmus luctuosus (Peters, 1864), from Rambodde; A. inornatus (Humbert, 1865), A. layardi (Humbert, 1865), A. thwaitesii (Humbert, 1865) and A. humberti (Carl, 1902, all from Paradeniya; and A. sabulosus Attems, 1898, from Kandy. All of them have been described from Sri Lanka, still known only from that island. Some of these taxa are however dubious, being based on female or even juvenile material, but most could be included into a key (Jeekel 1965). Regrettably, the first couplet in the key is purely geographic, separating the species from Myanmar and Sumatra from those described from Sri Lanka and India (Jeekel 1965). As one can see from the presently known distributions of A. saussurii and A. anthracinus, this distinction does not hold, also strongly suggesting several introductions through human agency. The only feasible solution lies in collecting new and/or spotting topotypic museum samples of the still enigmatic A. inornatus and A. layardi from Paradeniya, and of A. sabulosus from Kandy, to properly compare them to their type material. In addition, bar-coding could help tracing genetic relationships. Last, but not least, a few congeneric species, most of which also very similar to A. anthracinus, are known to occur in southern India as well. • Since Anoplodesmus is a senior synonym of Paranedyopus ( Golovatch 2000, Golovatch 2013, the sole erstwhile component species of the latter genus from Sri Lanka, A. simplex (Humbert, 1865), from Pundaloya ( Jeekel 1980c), must be considered as well. However, like any former Paranedyopus species, A. simplex shows reduced paraterga and more elaborate gonopods (Golovatch 2013). In other words, A. simplex is quite distinct from the above congeners from Sri Lanka which all have strongly developed paraterga and highly simple gonopods. In contrast, it seems to be more similar to A. rufocinctus (Carl, 1932) and A. subcylindricus (Carl, 1932), both latter taxa from southern India (Jeekel 1980c Attems, 1931 Euphyodesmus philippina (Nguyen Duc & Sierwald, 2010), comb. n.

Notes:
The identity of this species, described from Palawan Island in the genus Desmoxytes Chamberlin, 1923 (Nguyen Duc and Sierwald 2010), has recently been discussed and shown to actually represent the basically Australian subfamily Australiosomatinae (Golovatch et al. 2012). The species has thereby remained referred to as "Desmoxytes" philippina, the genus name being put in quotation marks to emphasize the wrong original assignment. Here we take the opportunity to allocate it properly at least at the subfamily level, choosing the Bornean Euphyodesmus as perhaps the best candidate genus (Golovatch 1996).

Genus Luzonomorpha Hoffman, 1973
Nomenclature: Type-species: Prionopeltis montana Chamberlin, 1921 Notes: This strictly Philippine genus has recently been reviewed, and most of its species have been keyed (Jeekel 2000).

Luzonomorpha acutangula (Newport, 1844)
Notes: Described as Polydesmus acutangulus from an unspecified locality in the Philippines, it has sometimes been quoted in the original spelling (e.g. Jeekel 2000), material contained only females, the identification seems to be highly dubious, better to be ignored altogether.