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Taxonomic paper

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)

expand article infoSergei Golovatch, Pavel Stoev,§
† Institute for Problems of Ecology and Evolution, Russian Academy of Sciences, Moscow, Russia
‡ National Museum of Natural History, Bulgarian Academy of Sciences, Sofia, Bulgaria
§ Pensoft Publishers, Sofia, Bulgaria
Open Access

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: E. 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.

Keywords

Millipedes, checklist, Luzon, State Pulau Penang, new species, new record

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.

Taxon treatments

Eustrongylosoma penevi Golovatch & Stoev, 2013, sp. n.

Materials   Download as CSV 
Holotype:
  1. island: Luzon Island; country: Philippines; stateProvince: Mountain Province; verbatimLocality: Mt Polis Checkpoint on the road Banaue – Sagada; verbatimElevation: 1800-1900 m; locationRemarks: under stones and logs; verbatimLatitude: 16°57'58"N; verbatimLongitude: 121°1'37"E; eventDate: 6 July 2012; individualCount: 1; sex: male; recordedBy: P. Stoev & L. Penev; institutionCode: NMNHS
Paratypes:
  1. island: Luzon Island; country: Philippines; stateProvince: Mountain Province; verbatimLocality: Mt Polis Checkpoint on the road Banaue – Sagada; verbatimElevation: 1800-1900 m; locationRemarks: under stones and logs; verbatimLatitude: 16°57'58"N; verbatimLongitude: 121°1'37"E; eventDate: 6 July 2012; individualCount: 4; sex: male; recordedBy: P. Stoev & L. Penev; institutionCode: NMNHS
  2. island: Luzon Island; country: Philippines; stateProvince: Mountain Province; verbatimLocality: Mt Polis Checkpoint on the road Banaue – Sagada; verbatimElevation: 1800-1900 m; locationRemarks: under stones and logs; verbatimLatitude: 16°57'58"N; verbatimLongitude: 121°1'37"E; eventDate: 6 July 2012; individualCount: 1; sex: female; recordedBy: P. Stoev & L. Penev; institutionCode: NMNHS
  3. island: Luzon Island; country: Philippines; stateProvince: Mountain Province; verbatimLocality: Mt Polis Checkpoint on the road Banaue – Sagada; verbatimElevation: 1800-1900 m; locationRemarks: under stones and logs; verbatimLatitude: 16°57'58"N; verbatimLongitude: 121°1'37"E; eventDate: 6 July 2012; individualCount: 1; lifeStage: juvenile; recordedBy: P. Stoev & L. Penev; institutionCode: NMNHS
  4. island: Luzon Island; country: Philippines; stateProvince: Mountain Province; verbatimLocality: Mt Polis Checkpoint on the road Banaue – Sagada; verbatimElevation: 1800-1900 m; locationRemarks: under stones and logs; verbatimLatitude: 16°57'58"N; verbatimLongitude: 121°1'37"E; eventDate: 6 July 2012; individualCount: 1; sex: male; recordedBy: P. Stoev & L. Penev; institutionCode: ZMUM

Description

Length 18-22 (♂) or 23 mm (♀), width of midbody pro- and metazona 1.1-1.3 and 1.5-1.7 mm (♂), or 1.9 and 2.1 mm (♀), respectively. Holotype ca 22 mm long, width of pro- and metazona 1.3 and 1.6 mm, respectively. Coloration black to light grey-brown (Fig. 1a). Pattern mostly cingulate due to a large light grey band on prozona dorsally in front of stricture extending down until level of paraterga (Fig. 1a, b). Legs light grey-brown. Antennae increasingly infuscate distad, from light brown to blackish (Fig. 1a).

Figure 1.

Eustrongylosoma penevi sp. n.

a♂ paratype, habitus, lateral view
bmidbody segments, dorsal view
c♂ paratype 3 segment 10, lateral view. Scale bar: 1.0 mm.
dleft half of segment 10, dorsal view. Scale bar: 1.0 mm.

Body submoniliform. Antennomeres 2 to 6 subequal in length, antennae rather short, reaching behind segment 3 (♂) or 2 (♀) when stretched dorsally. Tegument generally smooth and shining. In width, segments 2 and 3 < collum = 4 < 5-17, thereafter body gently tapering towards telson. Paraterga (Fig. 1b, c, d) modestly developed, keel-shaped, set low (at about 1/3rd of metazonal height), thinner in poreless, thicker in pore-bearing, segments, slightly reaching behind tergal margin only in segments 2 and 17-19, mostly slightly pointed, delimited by a complete and deep sulcus only dorsally, ventral sulcus being incomplete, developed only in posterior quarter to 1/5th extent. Head densely setose on clypeus and frons, bare on vertex. Collum semilunar, bearing two transverse rows of 2+2 setae, one row along front margin, second row in the middle; lateral edges broadly rounded. Metaterga faintly rugulose, a little more clearly so in postsulcus halves, surface below paraterga microgranular in segments 2-7. Tergal setae rather long, about 1/5th of metatergal length, arranged in two rows of 2+2 in each, one in front of, second behind sulcus. Sulcus starting from segment 5, deep, almost reaching the bases of paraterga. Stricture between pro- and metazona finely and densely ribbed. Ozopores lying close to caudal end of paraterga in a shallow ovoid groove, lateral, only partly visible from above. Pleurosternal carinae poorly developed ridges visible only in segments 2-4. Seta at about midway of each paratergum mostly broken off. Axial line wanting. Epiproct (Fig. 2a) subtruncate, pre-apical lateral papillae small. Hypoproct (Fig. 2b) semi-circular. Sternal lobe between coxae 4 subtrapeziform, densely setose (Fig. 2c). Legs very long and slender, about 2.0 (♂) or 1.3 (♀) times as long as midbody height; ♂ tarsal brushes traceable until about legs of segment 15, thereafter thinning out.

Figure 2.

Eustrongylosoma penevi sp. n. Scale bar: 1.0 mm.

aepiproct
bhypoproct
csternal lobe between coxae 4, caudal view
dright gonopod, mesal view
eright gonopod, lateral view
fright gonopod, dorsal view

Gonopods rather simple (Fig. 2d, e, f): coxite long, subcylindrical, bare; prefemoral portion small, about 1/3rd as long as femorite, the latter slender, ventral lobe somewhat better developed than dorsal one, apicolateral lobe (l) rounded, well developed, with a long transverse spine (s) at base. Solenophore subcircular, with a subterminal lobule.

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).

Figure 3.

Map of Luzon showing the type locality of Eustrongylosoma penevi sp. n. Original map from: Alexander Altenhof (Kater Begemot) via Wikimedia Commons.

Figure 4.

A view of Mt Polis Checkpoint, the type locality of Eustrongylosoma penevi sp. n. Red arrow indicates the exact place where species was found.

Anoplodesmus anthracinus Pocock, 1895

Materials   Download as CSV 
  1. higherGeography: Malay Peninsula; country: Malaysia; stateProvince: State Pulau Penang; verbatimLocality: Station MARDI Seberang Perai; locationRemarks: agriculture area, in close proximity to experimental rice fields, under wooden board; verbatimLatitude: 5°32'24''N; verbatimLongitude: 100°28'11''E; eventDate: 15 June 2011; individualCount: 1; sex: male; behavior: copulation observed; recordedBy: P. Stoev & L. Penev; institutionCode: NMNHS
  2. higherGeography: Malay Peninsula; country: Malaysia; stateProvince: State Pulau Penang; verbatimLocality: Station MARDI Seberang Perai; locationRemarks: agriculture area, in close proximity to experimental rice fields, under wooden board; verbatimLatitude: 5°32'24''N; verbatimLongitude: 100°28'11''E; eventDate: 15 June 2011; individualCount: 2; sex: females; behavior: copulation observed; recordedBy: P. Stoev & L. Penev; institutionCode: NMNHS
  3. country: Sri Lanka; stateProvince: Sabaragamuwa Prov.; verbatimLocality: Millenium Foundation Orphanage; verbatimElevation: 90 m; verbatimLatitude: 7°16'40"N; verbatimLongitude: 80°23'12"E; eventDate: 19-22.XII.2012; individualCount: 1; sex: male; recordedBy: I. Melnik; institutionCode: ZMUM
  4. country: Sri Lanka; stateProvince: Sabaragamuwa Prov.; verbatimLocality: Millenium Foundation Orphanage; verbatimElevation: 90 m; verbatimLatitude: 7°16'40"N; verbatimLongitude: 80°23'12"E; eventDate: 19-22.XII.2012; individualCount: 2; sex: females; recordedBy: I. Melnik; institutionCode: ZMUM

Description

Length ca 33 mm, width of pro- and metazona 2.8 and 3.8 mm, respectively (♂), or 25, 3.0 and 4.0 mm, respectively (♀). Colour pattern highly vivid (Fig. 5), shiny blackish to dark brown, with contrasting yellowish paraterga and the immediately adjacent regions. Paraterga very well-developed, set rather high (about 1/4th of midbody height) (Fig. 6a), callus wide (Fig. 6b), thicker in pore-bearing paraterga. Pleurosternal carinae longitudinally arched ribs, increasingly poorly developed towards telson to totally disappear in segment 15. Legs only slightly enlarged in male, rather long and slender, about 1.3 (♂) or 0.9 (♀) times as long as midbody height. ♂ legs 5 (Fig. 7a) and 6 with large femoral humps, ♂ femur 7 with a humped process even greater than in leg 6 (Fig. 7b). Epiproct subtruncate (Fig. 6c). Hypoproct roundly subtrapeziform (Fig. 6d). Sternal lamina between ♂ coxae 4 semi-circular (Fig. 6e).

Figure 5.

Anoplodesmus anthracinus Pocock, ♂.

ahabitus, lateral view
bhabitus, in situ, dorsal view
Figure 6.

Anoplodesmus anthracinus Pocock, ♂.

asegment 10, lateral view. Scale bar: 2.0 mm
bleft half of segment 10, dorsal view. Scale bar: 2.0 mm
cepiproct. Scale bar: 2.0 mm
dhypoproct. Scale bar: 2.0 mm
esternal lobe between coxae 4, caudal view. Scale bar: 2.0 mm
fright gonopod, mesal view. Scale bar: 1.0 mm
Figure 7.

Anoplodesmus anthracinus Pocock, ♂.

amale leg 5, lateral view
bmale leg 7, lateral view

Gonopods very simple (Fig. 6f): coxite with a few strong setae distodorsally, prefemoral part prominent, only slightly shorter than acropodite; femorite with a strong ventral tooth (a), solenophore bipartite, with two apical lobes (b, c), lobe c supporting a short solenomere (sl).

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 close proximity to experimental rice fields (Fig. 9), while in Sri Lanka, the collecting locality is a human settlement.

Figure 8.

Map of the southern part of Malay Peninsula showing the new locality of Anoplodesmus anthracinus. Original map from Wikimedia Commons.

Figure 9.

A view of Station MARDI Seberang Perai where A. anthracinus was found.

Figure 10.

Map of Sri Lanka showing the new locality of Anoplodesmus anthracinus. Original map from: Uwe Dedering, via Wikimedia Commons.

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).
  • Anoplodesmus anthracinus, new to the fauna of Sri Lanka.

Checklist of the Philippine Paradoxosomatidae

Subfamily Australiosomatinae

Tribe Antichiropodini

Genus Euphyodesmus Attems, 1931

Nomenclature: 
Type-species: Euphyodesmus gracilis 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 Desmoxytesphilippina, 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).

Subfamily Paradoxosomatinae

Tribe Orthomorphini

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), but we prefer to follow Hoffman (1973), who clearly spelled the name in the feminine gender.

Luzonomorpha infulata (Wang, 1951)

Notes: 
Described from Mindanao Island.

Luzonomorpha montana (Chamberlin, 1921)

Notes: 
Described from several places in Luzon.

Luzonomorpha pallidula Jeekel, 2000

Notes: 
Known from Mindoro Island.

Luzonomorpha picea (Brandt, 1839)

Notes: 
Described from Manila.

Luzonomorpha polilloensis (San Juan & Lit, 2010), comb. n.

Notes: 

Known from Polillo Island (San Juan and Lit 2010). This species was originally assigned to Prionopeltis Pocock, 1895, a genus long known to be invalid (Jeekel 1968). In fact it definitely belongs to Luzonomorpha.

Luzonomorpha quatuorputeus (Wang, 1951)

Notes: 
Known both from Mindanao and Luzon islands.

Genus Orthomorpha Bollman, 1893

Nomenclature: 
Type-species: Polydesmus beaumontii Leguillou, 1841

Orthomorpha coarctata (De Saussure, 1860)

Notes: 

This pantropical, definitely anthropochore species has often been referred to as a distinct genus, Asiomorpha Verhoeff, 1939, but we prefer to regard O. coarctata as a species of Orthomorpha (see Likhitrakarn et al. 2011). In the Philippines, it has been recorded in Mactan and Cebu islands (Wang 1955).

Tribe Sulciferini

Genus Chondromorpha Silvestri, 1897

Nomenclature: 
Type-species: Chondromorpha severini Silvestri, 1897

Chondromorpha xanthotricha (Attems, 1898)

Notes: 

In the Philippines, this nearly pantropical, definitely anthropochore species has only been recorded in Luzon (Wang 1953).

Tribe Eustrongylosomatini

Genus Eustrongylosoma Silvestri, 1896

Eustrongylosoma penevi Golovatch & Stoev, 2013, sp. n.

Notes: 
Known from Luzon Island.

Tribe Tectoporini

Genus Helicorthomorpha Attems, 1914

Nomenclature: 
Type-species: Strongylosoma holstii Pocock, 1895

Helicorthomorpha luzoniensis (Peters, 1864), comb. n.

Notes: 

Originally described from Luzon, without a more precise locality (Peters 1864), this species, previously considered dubious (Jeekel 1968), appears to actually represent a new senior subjective synonym of H. orthogona (Silvestri, 1898). The syntypes (1 ♂, 1 ♀), labelled "Bosoboso, Luzon, leg. Martens" (Moritz and Fischer 1978), have been revised and returned to the Museum für Naturkunde in Berlin as а lectotype (♂) and а paralectotype (♀). Lectotype designation is necessary to ensure the species to be based on male material. In addition, unlike the paralectotype, which is an incomplete female, the lectotype is complete. The name luzoniensis is preferred because of its priority, being in use in the last 50 years and thus not representing a nomen oblitum (Moritz and Fischer 1978). This widespread species, previously referred to as H. orthogona, is known to occur from China to New Guinea (Jeekel 2009). In the Philippines it has been recorded from Luzon and Mindanao islands (Wang 1951, Jeekel 1980b).

Species incertae sedis

Orthomorpha bisulcata Pocock, 1895

Notes: 

This species, originally described from Myanmar (Pocock 1895), has been recorded under this name from the Philippines, based on material shipped from the Philippines and intercepted by quarantine in the Hawaiis (Wang 1951). Because that material contained only females, the identification seems to be highly dubious, better to be ignored altogether.

Acknowledgements

The collecting trips of the second author to the Philippines and Malaysia were funded by the German Federal Ministry of Education and Research (BMBF) within the international research programme “Sustainable Land Management” LEGATO Project (Land-use intensity and ecological engineering – assessment tools for risks and opportunities in irrigated rice based production systems, www.legato-project.net). Specimens were collected with permission from the Department of Environment and natural Resources of the Republic of Philippines (Wildlife Gratuitous Permit No. DENR-CAR-001-13). Special thanks go to Zoltán Korsós (Hungarian Natural History Museum, Budapest) for sharing with us his unpublished catalogue draft, and to Jason Dunlop and Anja Friederichs (Museum für Naturkunde in Berlin) for sending us on loan the type material of Strongylosoma luzoniense under their care. Bob Mesibov (Queen Victoria Museum and Art Gallery, Tasmania, Australia), Nesrine Akkari (Danish Natural History Museum, Copenhagen) and Teodor Georgiev (Pensoft) provided helpful comments on the manuscript. We are most grateful to the Bulgarian-Russian Interacademician Exchange Programme which allowed SG to visit NMNHS in 2012 and 2013.

References