Corresponding author: Bernhard Klarner (
Academic editor: Pavel Stoev
A new genus
Bonato L, Klarner B, Widyastuti R, Scheu S (2016) The first geophilid centipedes from Malesia: a new genus with two new species from Sumatra (Chilopoda, Geophilidae). ZooKeys 605: 53–71. doi:
The diversity of geophilomorph centipedes (
The situation is especially unsatisfactory for the large island of Sumatra, when compared with the neighbouring Malay peninsula and the other Malesian islands. To the best of our knowledge, all records of geophilomorph centipedes from Sumatra derive from half a dozen papers (
Species of
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Two new species of
Specimens were found in soil samples collected in Sumatra, along a gradient including secondary rainforests, jungle rubber agroforests, rubber, and oil palm plantations. Sampling has been carried out within the interdisciplinary project “Ecological and socioeconomic functions of tropical lowland rainforest transformation systems (Sumatra, Indonesia) – EFForTS”. For details on the study region and experimental design see
The specimens were examined by light microscopy (Leica DMLB) according to standard protocols for geophilomorphs, by placing them in ethylene glycol in temporary mounts (
To evaluate whether similar or possibly related species were already recorded in south-eastern Asia, the entire taxonomic and faunistic literature on centipedes was browsed to retrieve all published records from that area.
Indonesian Institute of Science, Cibinong, Indonesia
Department of Biology, University of Padova, Italy
Relatively small geophilids, less than 1 cm long; cephalic plate distinctly elongate, without frontal line; clypeus with two pairs of setae on the anterior medial part, without a distinct clypeal area; intermediate part of labrum bearing stout tubercles, lateral parts far apart from each other and bearing bristles; first maxillae without lappets; second maxillary coxosternite with anterior margin entire and concave, without anterior projections, neither statuminia nor other distinctly sclerotized parts associated with the metameric pores; second maxillary pretarsus in shape of an elongate claw, more than 3.5 times as long as wide at the basis, sub-conic and slightly bent, with a small sub-basal dorsal bulge; forcipular tergite subtrapezoidal; forcipular coxosternite relatively elongate, the exposed part as wide as or only slightly wider than long, the anterior margin slightly projecting anteriorly, with two very short denticles and a narrow notch inbetween; coxopleural sutures complete, entirely ventral, straight and subparallel for most of their length; chitin-lines absent or hardly distinct; forcipules relatively elongate, the trochanteroprefemur is more than 1.4 times as long as wide, the tarsungulum more than 2.5 times as long as wide; forcipular intermediate articles distinct, without denticles; tarsungulum with at least a distinct basal denticle; trunk metasternites longer than wide, without obvious “carpophagus” pit; whenever present, a single sub-circular, posterior pore-field on all metasternites of the trunk; leg claws with at most a pair of accessory spines, shorter than mid-length of the pretarsus, similar to each other in length; ultimate leg-bearing segment with an entire pleuropretergite, without sulci; ultimate metasternite sub-trapezoid, the setae distributed almost uniformly in the female, unknown in the male; coxopleuron with at least two coxal organs, opening through independent pores on the ventral side; telopodite of the ultimate leg pair approximately 1.8–2.0 times as long as that of the penultimate pair; anal pores distinct.
From “Sunda”, the name in use for the south-eastern part of the continental shelf of Asia, including Sumatra and other islands, and “
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The two new species are confidently recognised as belonging to the family
The two species are here described in a new genus because they do not fit the diagnosis of any other known geophilid genus (Table
Major differences between the species of
Genus/Species | General features | Clypeus | Labrum | Second maxillae | Forcipule | Leg-bearing segments | Ultimate pair of legs | |||||||||
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head and forcipules: distinctly elongate | clypeal area: distinctly present | lateral parts: almost touching each other | coxosternite: statuminia: distinctly present | coxosternite: anterior |
coxosternite: anterior projections: distinctly present | pretarsus: much elongate | pretarsus: distinctly stout | tarsungulum: a second denticle flanking the basal denticle | number of leg pairs | anterior metasternites: pore-fields present | coxopleuron: ventral pores opening into pits | coxopleuron: all ventral pores close to metasternite | telopodite: number of articles | telopodite: distinctly swollen in females | pretarsus: shape | |
+ | - | - | - | - | - | + | - | + | 33–35 | - | - | + | 6 | + | group of spines | |
+ | - | - | - | - | - | + | - | - | 31* | + | - | - | 6 | - | claw | |
+ | + | - | - | + | - | - | - | - | 71* | - | - | - | 5 | - | claw | |
- | ? | - | ? | + | - | ? | ? | - | 41–45 * | + | + | ? | ? | - | claw | |
+ | + | + | - | - | - | + | - | - | 41–91 | + | + | + | 6 | - | claw | |
- | - | - | - | + | - | - | - | - | 47–65 | + | - | + | 6 | - | claw | |
+ | + | - | - | + | - | - | + | - | 47–81 | - | - | - | 6 | ? | claw | |
+ | + | + | + | + | - | - | - | - | 37–75 | - | - | - | 6 | - | claw | |
+ | + | - | + | + | + | - | - | - | 31–125 | + | +/- | +/- | 6 | - | claw/spine | |
+ | + | - | + | + | + | - | + | - | 49–79 | - | - | - | 6 | - | claw | |
+ | + | + | - | + | - | + | - | - | 37–53 | - | - | - | 6 | - | claw | |
- | - | - | - | - | - | - | - | - | 39–73 | + | + | + | 6 | - | claw | |
+ | + | - | ? | ? | ? | ? | ? | - | 33–41 | - | - | + | 6 | - | claw |
The two new species are similar to each other in the minute body size, the head and the forcipules distinctly elongate, the second maxillae provided with very slender claws, as well as in other characters. Nevertheless, uniting the two species in a single genus should be taken as a preliminary, parsimonious arrangement. Actually, we cannot rule out the possibility that most similarities between the two species comprise convergent adaptive characters or shared ancestral conditions. As a matter of fact, body miniaturization evolved independently in different lineages of geophilids (
The forcipules of
The forcipules of the geophilomorphs show great diversity in number, size and pattern of denticles (
The discovery of two geophilid species inhabiting Sumatra is quite unexpected when confronting the known global distribution of the
While many geophilid species in different genera are known living in temperate Asia, Australia and many Pacific islands, only a few claims have been published so far for the entire area comprising Indochina, Malesian islands, and New Guinea, and all these putative records have turned out to refer to misclassified representatives of different families. In particular, the species of
We thank the village heads, local site owners, PT REKI, and Bukit Duabelas National Park for granting access to their properties and the counterparts and local assistants for support. We appreciate the comments from Pavel Stoev, Luis Pereira, and Marzio Zapparoli that helped to improve an earlier version of the manuscript. A collection permit (no. S.07/KKH-2/2013) was recommended by the Indonesian Institute of Sciences (