Research Article |
Corresponding author: Menno Schilthuizen ( menno.schilthuizen@naturalis.nl ) Academic editor: Borislav Guéorguiev
© 2018 Menno Schilthuizen, Michel Perreau, Iva Njunjić.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Schilthuizen M, Perreau M, Njunjić I (2018) A review of the Cholevinae from the island of Borneo (Coleoptera, Leiodidae). ZooKeys 777: 57-108. https://doi.org/10.3897/zookeys.777.23212
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The available knowledge of the round fungus beetle subfamily Cholevinae (Leiodidae) from the island of Borneo is reviewed, and the results of newly studied material presented. The currently known 30 species (of which 14 are newly described herein) represent the genera Micronemadus (one species), Catops (one species), Baryodirus (one species), Ptomaphaginus (14 species), and Ptomaphaminus (13 species). The following new species are described: Micronemadus sondaicus Schilthuizen & Perreau, sp. n., Ptomaphaginus grandis Schilthuizen & Perreau, sp. n., P. louis Schilthuizen & Perreau, sp. n., P. muluensis Schilthuizen & Perreau, sp. n., and P. isabellarossellini Schilthuizen, Njunjić & Perreau, sp. n., and Ptomaphaminus kinabatanganensis Njunjić, Schilthuizen & Perreau, sp. n., P. testaceus Schilthuizen & Perreau, sp. n., P. nanus Schilthuizen & Perreau, sp. n., P. marshalli Schilthuizen & Perreau, sp. n., P. hanskii Schilthuizen & Perreau, sp. n., P. sarawacensis Schilthuizen & Perreau, sp. n., P. layangensis Schilthuizen & Perreau, sp. n., P. microphallus Schilthuizen & Perreau, sp. n., and P. alabensis Schilthuizen & Perreau, sp. n. It is expected that the cholevine biodiversity of Borneo is still far from completely known. Nonetheless, provisional identification keys to all species known so far are presented.
caves, Indonesia, Malaysia, scavenging beetles, taxonomy
Borneo is, after New Guinea, the second-largest tropical island of the world. It has never been strongly isolated, having formed part of a larger land mass, known as Sundaland, during marine transgressions in the Pleistocene (
The beetle subfamily Cholevinae (Coleoptera, Staphylinoidea, Leiodidae) consists mostly of small, soil-dwelling scavengers, well represented in the litter fauna of all tropical regions.
In this paper, we provide an overview of the species of Cholevinae currently known from the island of Borneo. By necessity, this is a very preliminary overview, since it is based on comparatively little information.
We provide a brief description for all previously described genera and species, and more extensive descriptions for newly-described species, as well as differential diagnoses for new species that have close congeners in Borneo. Where available, we also refer to DNA sequences in the Barcode of Life Database (BOLD, http://boldystems.org) and to so-called Barcode Index Numbers (BINs;
The collection abbreviations used in the lists of examined material are as follows:
JRUC collection of Jan Růžička, Prague, Czech Republic;
TXEX Taxon Expeditions, Leiden, The Netherlands.
CMPR Collection Michel Perreau, Paris.
Male genitalia were mounted in Euparal after dissection and dehydration in ethanol 95%. Female genitalia were cleared in hot KOH 10% and stained with Azoblack before mounting in DMHF (2,5-DiHydroxyMethylFurane). Photographs of genitalia were taken on a Leitz Diaplan microscope using a Spot Insight IN1820 or a Leica MC170HD camera. High-resolution photonic pictures of external morphological details (Figure
Small, 1.4–2.6 mm. Oval habitus, with the apex of the elytra rounded. Head and pronotum punctate, elytra with transverse strigae. Antenna with antennomeres 4 and 5 very short and wide. Mesosternum with a longitudinal mesoventral process. The four first protarsomeres and the first mesotarsomere dilated in the male. Aedeagus: median lobe triangular, parameres longer than the median lobe, and bent inward.
Holotype: Malaysia, Sabah, Crocker Range Park, Inobong, 5°51.265'N, 116°08.363'E, 500 m elev., 21–23.ix.2012 (leg. M. Schilthuizen, Crocker Range / Kinabalu Expedition,
Length: 2.0–2.6 mm. Colouration: in fully coloured individuals dark brown to black, margins of pronotum and front half of the elytra reddish brown, entirely covered in reddish brown setae; legs, palps, and basal three and final antennomeres light reddish brown; antennomeres 4–10 darker (Figure
Similar to M. pusillimus (Kraatz, 1877), described from Japan, but compared with Japanese specimens of M. pusillimus that we have seen, M. sondaicus is larger, more uniformly coloured (Figure
Micronemadus. a M. sondaicus sp. n. habitus (paratype female, Sarawak, Gunung Mulu) b M. pusillimus (Kraatz) aedeagus, Sarawak, Gunung Mulu c M. sondaicus sp. n. aedeagus Sarawak, Gunung Mulu d M. sondaicus sp. n. endophallus, Sarawak, Gunung Mulu e M. sondaicus sp. n. male urite IX, Sarawak, Gunung Mulu f M. sondaicus sp. n. female sternite VIII, Sarawak, Gunung Mulu.
In the BOLD database, COI sequences are available for the holotype (
Very common and widespread, in primary and secondary forest, 0–1850 m elev. In addition to the type material, we have seen seemingly conspecific material from many other localities in Sundaland, (Sabah, Sarawak, Peninsular Malaysia, Mindanao, Java, Bali) and also from Vietnam.
For many years, we considered this Borneo Micronemadus, which is usually the commonest leiodid in baited traps, as identical to M. pusillimus. However, DNA-barcodes for Japanese individuals (BOLD BIN ABU9390) and Bornean individuals (BOLDBINsABU9391 and ACK0008) display a 17% sequence divergence, strongly suggesting that, despite the only slight morphological differences, these belong to separate, not closely related species. We suspect that M. pusillimus, previously considered a very widespread Asian species (
The name refers to the Sunda region, of which Borneo forms part (sondaicus (L.) = from Sunda). We used the spelling sondaicus, rather than sundaicus, to conform with other specific epithetons, such as Rhinoceros sondaicus Desmarest, 1822.
Fairly robust cholevines of sizes that range from just longer than 2 mm to almost 10 mm. Body fairly convex, pronotum relatively large, but usually narrower than the elytra. Elytra sometimes with traces of parallel longitudinal striae. Antennae with the 8th antennomere usually broader than long. In the male, the four first protarsomeres and the first mesotarsomere are dilated; also, the protibiae often are sexually dimorphic. Aedeagus: median lobe usually symmetric, elongated, lance-shaped, terminally rounded, truncate, acute, or two-pronged. Parameres usually thin, hair-like.
Catops pruinosus Schweiger, 1956: 538, fig. 5; type from Kuatun, Fukien, China.
Catops
solitarius
Szymczakowski, 1961: 129, figs 16–19; type from Sandakan, Sabah, Borneo (in
Sabah. Sandakan (leg. W.B. Pryer,
Length: 3.9 mm. Habitus slender and elongated, somewhat flattened. Body reddish brown; head, centre of the pronotum, and centre of the elytra dark brown. Entire dorsum covered in orange setae. Antenna robust, 7th antennomere as wide as long, 6th, 8th, 9th, and 10th wider than long. Pronotum 1.54 times as wide as long, frontal and caudal margins straight, lateral margins curved, more strongly rostrad than caudad (greatest width of the pronotum slightly behind the centre), caudal angles broadly rounded; surface with dense, rasp-like punctuation and somewhat matte due to microsculpture. Elytra with fine punctuation, shagreened, and with slight indications of longitudinal striae, 1.27 times as wide as the pronotum, 1.3 times as long as jointly wide (length measured from the caudal tip of the scutellum to the apex of the elytra). Male unknown.
Catops pruinosus is known from a large latitudinal expanse along the East Asian coast: Shanghai, Fujian, and North Borneo. The only known record from Borneo (Sabah: Sandakan) is the female holotype of C. solitarius, later synonymized with C. pruinosus (
Catops pruinosus is a member of the C. hilleri group (
Length: 2.2 mm. Winged and with fully developed eyes (Figure
Baryodirus
hammondi
Perreau, 2000 19–20, figs 1–10; type from Mulu, Sarawak (in
See genus description above: the genus is monotypic, and the holotype is the only known specimen of the species.
The biotope and ecology of this species are unknown, but several characters suggest that it could be a commensal of hymenopterans. The compact habitus and the double setation are observed (almost exclusively) in many myrmecophilous leiodid genera or subgenera, such as Ptomaphagus (Echinocoleus) Horn in North America, and Synaulus in North Africa (our observations).
In Borneo, Ptomaphaginus consists of relatively large Ptomaphagini with four dilated male protarsomeres, distinguishable from Ptomaphaminus by the ventral spines of the protibiae being aligned along the latero-external row of equal spines, making a second, more widely spaced row next to the external one, as well as by the metaventral sutures, which are roughly parallel to the axis of the body. Also, in the female, the gonocoxites are elongated, whereas in most Ptomaphaminus, they are reduced.
With Ptomaphaminus Perreau (see below), Ptomaphaginus Portevin makes up the vast majority of the Borneo cholevine diversity. We currently recognise 14 species. However, more diversity is to be expected, both cryptic (e.g., DNA-analysis suggests additional diversity in P. bryanti and related species;
Ptomaphaginus
anas
Schilthuizen & Perreau, 2008: 199, figs 16–17; type from Gombak, Selangor, Peninsular Malaysia (in
(In addition to that given in
(Adapted from
The aedeagus is characteristically shaped: strongly curved, distally tapering into a narrow, flattened, and slightly upturned apex. Ptomaphaginus louis and P. muluensis have a similar aedeagus, which, however, is less strongly curved in the terminal half.
One individual (
Lowland and lower montane forest, up to 1,500 m. Sabah: Batu Punggul, Kinabalu Park HQ, Kibongol; Sarawak: Semongok. Peninsular Malaysia: Gombak, Cameron Highlands.
Ptomaphaginus, habitus, dorsal. a P. anas Schilthuizen & Perreau, Ulu Gombak, male (
Ptomaphaginus
Bryanti
Jeannel, 1936: 56–59, figs 63–66; type from Mt. Matang, Sarawak, Borneo (in
Length 2.8 mm. Winged. Slender build (pronotum 1.56 times wider than long), as wide as the elytra. Aedeagus subterminally with two lateral extensions and a long median processus (the image given by
Very similar to P. bryantioides, from which it differs chiefly (as far as can be discerned from the single P. bryanti individual available) by its more slender habitus.
In the BOLD database, COI barcodes are available for
To date only known from the type location, Mt. Matang in Sarawak. However, given the great molecular distances (
Ptomaphaginus
bryantioides
Schilthuizen & Perreau, 2008: 192–193, figs 20–21; type from Danum Valley, Sabah, Borneo (in
(In addition to that given in
Habitus broad, rectangular, flat, very variable in size, 2.1–3.5 mm. Pronotum on average 1.8 times as wide as long, as wide as the elytra. Elytra as long as wide (length measured from the caudal tip of the scutellum to the apex of the elytra). Winged. Aedeagus with two apical, laterally directed extensions and a long terminal processus. (It appears that in P. bryanti, the lateral extensions may be directed even more ventrad, and the median processus is more widened apically than in P. bryantioides, but the significance of these differences needs to be substantiated.) Male forelegs with long setae on the ventral sides of the femur and tibia. The 3rd, 4th, and 5th visible abdominal ventrite of the male carry a slight central notch and show a depression around these notches. Spermatheca simple, inflated, semicircular (Figure
Distinguishable from other Bornean Ptomaphaginus species with the same aedeagus structure by the more elongated aedeagus with long, narrow processus, and the long setae on the male femur and tibia.
For the following specimens, COI barcodes are available in BOLD:
Widely distributed in Sabah and northern Sarawak, in primary and secondary lowland forest (usually up to 500 m, one exceptional record at 1350 m). Sabah: Kota Kinabalu, Kiansom, Sugud, Kinabalu Park (Poring), Crocker Range Park (Inobong, KK–Tambunan road km 56), Danum Valley, Batu Punggul; Sarawak: Mulu National Park. DNA sequencing (
Ptomaphaginus
burckhardti
Schilthuizen & Perreau, 2008: 202–203, figs 7–8; type from Gunung Kinabalu, Sabah, Borneo (in
(Adapted from
Unique among the Bornean Ptomaphaginus because of its small, slender build, reduced eyes, and long, slender aedeagus.
Only two specimens known (holotype and paratype), from upper montane forest at 2600 m elev. on Gunung Kinabalu.
Ptomaphaginus, habitus, dorsal. a P. louis sp. n., Gunung Mulu, paratype, male (
Ptomaphaginus
caroli
Schilthuizen & Perreau, 2008: 193, figs 14–15, 26; type from Gunung Mas, Sabah, Borneo (in
(Adapted from
Ptomaphaginus caroli has a similar aedeagus as P. bryanti, P. similipes, and P. bryantioides. However, it differs in having a distinctly elongated habitus (elytral index of 1.41) and very short apical ‘wings’ on the aedeagus.
So far, only known from the type specimen, collected in lower montane forest at 1350 m in the Crocker Range of Sabah. The aedeagal shape shows that it belongs within the “bryanti-group”.
The aedeagus of the holotype has been lost shortly after it was first collected and studied (in 2000). Before the loss, sketches were made of the dorsal and lateral view of the aedeagus, which form the basis for the line drawing in Figure
Holotype: Malaysia, Sarawak, Mulu National Park, Slope, TPS 7–9, 29.iv.1978 (P.M. Hammond & J.E. Marshall leg.,
Habitus: very large (3.4–4.0 mm), dark reddish brown, the elytra deeper red than the pronotum, basis and tip of the antenna pale, antennomeres 5–10 much darker; relatively parallel-sided and somewhat convex, head relatively narrow, pronotum 1.67 times as wide as long, slightly wider than the elytra, caudal angles clearly extended. Elytra of moderate length, convex, broadest at the shoulders, in the caudal one-third gently rounded towards the apex, jointly ca. 1.5 times as long as wide (length measured from the caudal tip of the scutellum to the apex of the elytra). Body entirely covered in dense pale yellow setation. Wings present. Antennae slender, 4th antennomere longer than wide, 9th and 10th antennomere square. Male protarsi only slightly dilated, the first four tarsomeres jointly ca. 4 times as long as wide. Aedeagus short and broad, in lateral view only very slightly bent ventrad with a barely perceptible upturned tip at the end, in dorsal view narrowing (in rounded fashion) towards the blunt apex. Stylet short and straight. Spiculum gastrale long, narrow-triangular, with the caudal part rounded. Spermatheca U-shaped, with ca. 6 distinct narrow rings on the proximate leg of the “U”, and ca. 6 additional, indistinct, broader rings on the remainder. Spermiduct long, thin, consisting of 5–6 360° coils.
Externally distinctive by its large size and the colour pattern of the antennae. From equally large P. bryantioides, it may be distinguished by the narrower male protarsi and the longer elytra. Aedeagus cannot be confused with that of any other known Bornean species, but is similar in shape to the one of P. nitens Jeannel, 1936 from Sri Lanka (which, however, is smaller and has much more condensed antennae) and of several species described from China and Taiwan (especially P. pingtungensis Perreau, 1996, P. guangxiensis Wang & Zhou, 2015, P. perreaui Wang & Zhou, 2015, and P. yui Wang & Zhou, 2015), from which P. grandis is distinguished by the unique combination of body size, antennomere proportions, and details of the spiculum gastrale, spermatheca, and aedeagus shape.
Only known from Mulu National Park in Sarawak.
Named grandis for its large size.
Ptomaphaginus, aedeagus, dorsal (left) and lateral (right) view. a, b P. kinabaluensis Schilthuizen & Perreau, Gunung Kinabalu (
Holotype: Malaysia, Sabah, Mt. Kinabalu National Park, Bukit Ular Trail (low), 1800 m elev., multistratum evergreen forest, 2 fish traps, 07–11.xi.1987 (leg. Krikken & Rombaut,
Large (3.3–4.0 mm), winged. Elytra, legs and basis and tip of the antennae dark reddish brown, pronotum, head and antennomeres 5–10 nearly black. Pronotum 1.6 times as wide as long, caudal angles clearly extended, as wide as the elytra at the shoulders. Elytra slender, slightly convex, 1.5 times as long as wide (length measured from the caudal tip of the scutellum to the apex of the elytra). Elytral apex in the female gradually rounded and joining the suture at a right angle; in the male more acute, meeting the suture at a sharp angle, and with a distinct bunch of thick, black, outwardly-curved, spine-like setae. Wings present. Body otherwise covered with dense light grey setation. Male tarsi strongly dilated, tarsomeres 1–4 jointly twice as long as wide. Female tarsi not dilated. Aedeagus very strongly curved ventrad, very convex, with a distinct dorsal keel; apex trilobate; stylet very long and thin, hair-like. Spermatheca semicircular, spermiduct extremely long (in extended condition probably at least 5 mm), consisting of ca. 30 360° loops.
Among Sabah Ptomaphaginus, and more generally, very distinctive by its large size, dark colouration (but see below under Remarks), the spine-like setae on the male elytral apex, and the uniquely shaped aedeagus (the basic design of which, with two lateral flaps at the apex, resembles that of the P. bryanti group, as well as non-Bornean species like P. sinuatus Schilthuizen, 1984). The conspicuous bunch of spines at the elytral apex in the male is shared with three other non-Bornean species, viz. P. riedeli Perreau, 1995, P. pilipennis Perreau, 1991 and P. pilipennoides Perreau, 1991, which, however, differ strongly from P. isabellarossellini in aedeagal shape. In other Bornean species, P. bryantioides and P. similipes, stronger setae at the male elytral apex can also be discerned, but never as conspicuous as in P. isabellarossellini.
Only known from the lower montane forest around Kinabalu Park Headquarters.
It should be noted that specimens of the normally rusty-coloured P. scaphaner Szymczakowski, 1972 from the same collection sample as P. isabellarossellini are also nearly black. This may mean that the dark colouration of P. isabellarossellini is a preservation artefact.
Named in honour of the actress and biologist Isabella Rossellini, whose short movies and stage performances on animal reproduction have popularized theories on the evolution of genitalia. In P. isabellarossellini, the extremely long penis stylet in the male and similarly long spermiduct in the female suggest a long history of sexually antagonistic coevolution, one of the types of selection that appears in Rossellini’s ‘Green Porno’ series on SundanceTV (
Ptomaphaginus, aedeagus, dorsal (left) and lateral (right) view. a, b P. isabellarossellini sp. n. Gunung Kinabalu, holotype (
Ptomaphaginus
kinabaluensis
Schilthuizen & Perreau, 2008: 195, figs 24–25, 29–30; type from Gunung Kinabalu, Sabah, Borneo (in
(In addition to that given in
Length 2.3–3.0 mm. Habitus slender, ovoid. Pronotum 1.68–1.86 times as wide as long, as wide as the elytra. Elytra 1.2–1.3 times as long as their combined width (length measured from the caudal tip of the scutellum to the apex of the elytra). Winged. Aedeagus short and wide, with two elongated apical, laterally directed ‘wings’ and a short terminal processus. Spermatheca thin, broadly bent over a 90° angle, slightly bulbous at the basis and with several indistinct annulations at the terminus. Spermiduct long, narrow, with numerous coils. Antennae short, as long as the width of the head. Long setae on the ventral side of the male profemur and protibia absent. Female elytral apices drawn out, male elytral apices rounded, not truncated. Male with a central extension on the 4th visible abdominal sternite.
Similar in aedeagal shape to P. bryantioides, but very different in habitus, which is more slender in P. kinabaluensis. Furthermore, P. kinabaluensis has only slightly dilated male protarsi, no setae on the male profemur and protibia, and extended elytral apices in the female.
For the following specimens, DNA barcodes are available in BOLD:
In montane forest at 1400–1930 m. Sabah: Kinabalu Park (around Park HQ), Crocker Range Park (Gunung Alab and km 51 KK-Tambunan road). The Crocker Range and Kinabalu populations are genetically very similar (BOLD BIN ACK0160).
Ptomaphaginus, male genital segment (urite IX), dorsal view. a P. anas Schilthuizen & Perreau, Ulu Gombak (
Ptomaphaginus
latimanus
Schilthuizen & Perreau, 2008: 196, figs 22–23; type from Gunung Trus Madi, Sabah, Borneo (in
(Adapted from
Ptomaphaginus latimanus is closely related to P. kinabaluensis, but differs in the habitus, which is much more stocky in P. latimanus. Also, P. kinabaluensis has extended elytral apices in the female, less strongly dilated male protarsi, and a central extension on the male 4th abdominal sternite.
Only known from montane forest at Gunung Trusmadi in Sabah, at 1400 m elev. One bryanti-group female (
Ptomaphaginus, female spermiduct and spermatheca. a P. bryantioides Schilthuizen & Perreau, Gunung Mulu (
Holotype: Malaysia, Sarawak, Mulu National Park, TPS 7–13, 4.v.1978 (leg. P.M. Hammond & J.E. Marshall,
Length 2.0–2.8 mm. Habitus: Light to dark reddish brown; flattened and relatively short and broad, head broad, pronotum 1.6–1.7 times as wide as long, narrower than the elytra, caudal angles almost not extended. Elytra short, gently convex, jointly ca. 1.2 times as long as wide (length measured from the caudal tip of the scutellum to the apex of the elytra). Body entirely covered in dense golden-yellow setation. Wings present. Antennae slender, 4th, 9th, and 10th antennomeres almost as long as wide. Male protarsi slightly dilated, the first four tarsomeres jointly ca. 3.5 times as long as wide. Aedeagus gently bent ventrad, flattened, apically broad and convex but subapically tapering in a rounded fashion into a broad but sharp upturned tip. Stylet long and thin, hair-like; stored in a wide loop in the basal part of the aedeagus. Spiculum gastrale elongate-ovoid, with the caudal part button-shaped, truncated. Spermatheca semicircular, thick, otherwise featureless, “sausage-shaped”. Spermiduct long, thin, consisting of ca. 4–6 360° coils.
Aedeagus in dorsal view very similar to that of P. tarsalis Symczakowski, 1964 from Sumatra, but in lateral view apically clearly more convex and with a shorter stylet. Moreover, the habitus and appendages of P. tarsalis are very stout and thick, whereas those in P. louis are much more slender. Also very similar to P. muluensis, but externally distinguished by the smaller size and more stocky habitus, with shorter elytra, narrower pronotum and the absence of drawn-out caudal pronotal angles. Aedeagus in dorsal view tapering abruptly towards the apex, not as gradually as in P. muluensis; in lateral view, the apex is more convex. Spermatheca distinguished from P. muluensis by the semicircular shape without any distinctive rings.
Only known from Mulu National Park, Sarawak.
Two females from the Mulu locality “Slope” have several rings at the basis of the spermatheca. As they are externally identical to other females of this species, they have been provisionally included in this species, but excluded from the type series. Several specimens infected on the elytra and pygidium with black Laboulbeniales.
We name this species after our friend and colleague Dr. Louis Deharveng (MNHN), in recognition for his logistic and emotional support during the preparation of this paper. The specific epithet is given as a noun in apposition.
Holotype: Malaysia, Sarawak, Mulu National Park, Mixed dipterocarp forest, TPS 7–10, v–viii.1978 (leg. P.M. Hammond & J.E. Marshall,
Length 2.5–3.1 mm. Habitus: reddish brown; relatively parallel-sided and somewhat flattened, head broad, pronotum 1.6 times as wide as long, as wide as the elytra, caudal angles clearly extended. Elytra of moderate length, gently convex, in the caudal one-third gently rounded towards the apex, jointly ca. 1.4 times as long as wide (length measured from the caudal tip of the scutellum to the apex of the elytra). Body entirely covered in dense golden-yellow setation. Wings present. Antennae slender, 4th, 9th, and 10th antennomeres slightly wider than long. Male protarsi slightly dilated, the first four tarsomeres jointly ca. three times as long as wide. Aedeagus gently bent ventrad, in lateral view apically flattened and ending in a bulbous upturned tip, in dorsal view gradually narrowing towards the apex. Stylet long and thin, hair-like. Spiculum gastrale long-ovoid, with the caudal part truncated. Spermatheca J-shaped, with 7–10 narrow rings on the long shaft of the “J”, and 5–7 broader rings on the curved part. Spermiduct long, thin, consisting of at least ten 360° coils.
Very similar to P. louis, but externally distinguished by the larger size and more slender habitus, with longer elytra, broader pronotum and distinctly drawn-out caudal pronotal angles. Aedeagus in dorsal view tapering gradually towards the apex, not as abruptly as in P. louis; in lateral view, the apex is not as convex. Spermatheca distinguished from P. louis by the J-shape with distinctive rings.
Only known from Mulu National Park, Sarawak. Several specimens infected with black Laboulbeniales on the elytra and the pygidium.
Named after Mulu National Park, to date the only locality from which this species is known.
Ptomaphaginus
sabahensis
Schilthuizen & Perreau, 2008: 202, figs 11, 12; type from Gunung Kinabalu, Sabah, Borneo (in
(adapted from
The elongated habitus combined with the rectangular and stocky aedeagus are unique features among the Bornean Ptomaphaginus.
Only known from the male holotype, collected on Gunung Kinabalu at 1580 m elevation.
Ptomaphaginus
scaphaner
Szymczakowski, 1972: 279–300, figs 28–33); type from Cue phuong Ninh binh, Vietnam (in
(in addition to that given in
Length 1.8–2.5 mm. Habitus broad and short, colouration light reddish-brown. Pronotum 1.75 times as wide as long, slightly narrower than the elytra. Elytra 1.2 times as long as jointly wide (length measured from the caudal tip of the scutellum to the caudal tip of the elytra). Wings present. Antennae very broad and short, with antennomeres 9 and 10 twice as broad as long. Aedeagus strongly curved, extremely convex and swollen, ending in a flattened “beak”; stylet long and thin, hair-like, running along the inside of the roof of the convex part of the aedeagus. Spiculum gastrale short, triangular, as long as wide. Spermatheca U-shaped, with 6–7 broad rings along the proximal part, and ca. 5 much narrower rings in the terminal one-quarter. Spermiduct very long and very thin.
Unique among the Bornean Ptomaphaginus by the condensed antennae and the inflated aedeagus with flattened “beak” and long stylet.
This appears to be a very widespread species, now known from Vietnam, Peninsular Malaysia, Borneo, and Java. It is possible, however, that the species consists of a complex of closely related species, given the geographic variation in secondary sexual characters (
Ptomaphaginus
similipes
Schilthuizen & Perreau, 2008: 193–194, figs 18–19, 27–28; type from Crocker Range Park, Sabah, Borneo (in
(In addition to that given in
Length 2.4–3.0 mm. Pronotum 1.69–1.80 times as wide as long, as wide as the elytra. Elytra 1.19–1.23 times as long as wide (length measured from the caudal tip of the scutellum to the apex of the elytra). Wings present. Aedeagus short and broad, with two short apical, laterally directed ‘wings’ and a short terminal processus. Male forelegs usually with long setae on the ventral side of the femur and tibia. Male protarsus completely not dilated, of the same width as in the female. The 3rd, 4th, and 5th visible abdominal ventrite of the male carry a slight central notch and show a depression around these notches. Spermatheca narrow, V-shaped, with ca. 10 indistinct rings. Spermiduct long and thin, consisting of ca. 6 360° loops.
Among other members of the P. bryanti complex, distinguished by the not dilated male protarsi, thin, annulated spermatheca, the relatively small size, and the short and squat aedeagus.
COI barcodes are available for the following specimens:
The male protarsi were slightly dilated in the one male from the Kinabalu Headquarters area.
During the study of this Bornean material, we have refined our concept of the genus Ptomaphaminus, which is why the genus description below is more extensive than for the previous genera.
Species of small size, not exceeding 2 mm. Colour generally brown, partly yellowish or light brown, rarely darker. Dorsal surface covered with short recumbent setae inserted along transverse strigae which also cover the whole dorsum of the body. Head with more or less developed eyes. A significant eye reduction is observed in species living in subterranean environments. Antennae generally slender, the apical club weakly marked. Pronotum transverse, the largest width generally at the base, the sides of the pronotum and the elytra continuously arcuate, of equal width. Elytra generally elongate, the posterior sutural angles rounded in males (Figure
The two species P. latescens and P. testaceus sp. n. have significantly different morphological characters from other species of Ptomaphaminus (not limited to Borneo): a short stylus of the endophallus (limited to half the length of the median lobe) with a symmetric basal loop, female gonocoxites long (more than three times longer than wide), a weakly sclerified spermatheca with a set of transversal rings (similar to structures preventing a collapse under depression, like for the respiratory trachea). Other species have a long stylus developed on most of the length of the aedeagus, female gonocoxites short, sub-square and a more sclerified spermatheca without reinforcing transversal rings. These two species form a distinct species group which possibly represents another genus.
Little information is available on the biology of Ptomaphaminus. Two methods of sampling are successful in obtaining specimens: trapping with pitfall traps baited with meat, cheese, or human excrement (either in epigean or in cave environments) and manual collecting in caves. Species collected in epigean conditions generally have fully developed eyes and flight wings while specimens from caves often have reduced eyes (although presently no anophthalmic Ptomaphaminus are known). The eye reduction observed in species recorded from caves is not correlated with the flight wing reduction, in contrast to palaearctic and nearctic subterranean species of Cholevinae (Leptodirini; Ptomaphagus (Adelops)). Flight wings of P. fagei Perreau, 2009 and P. latescens Szymczakowski, 1964, for example, remain fully functional as observed in Gua Sedepan (Eastern Kalimantan) and caves in the Kinabatangan valley (Sabah) where specimens flew up when lighted by headlamps even in the dark zone deep inside caves. A similar observation was reported by
Species are very similar externally. For each of the species below, we provide only specific diagnoses, without listing any shared generic characters.
Ptomaphaginus
latescens
Szymczakowski, 1964: 140–144, figs 122–132; type from Cave of Durian, Padang Highlands, Sumatra (in
Sabah. Kinabalu Park, Poring Hot Springs, 6°02.894'N, 116°14.957'E, 625 m elev., 15–20.ix.2012 (leg. M. Schilthuizen, Crocker Range / Kinabalu Expedition, in
Length 1.45–1.75 mm. General colour dark brown. Winged. Eyes normally developed. Pronotum 1.56 times as wide as long. Elytra 1.35 times as long as wide. Female sutural angle of elytra expanded backwards into a sharp tooth. Male protarsi 0.5 times as wide as the protibia. Male genital segment with a very elongate and thin spiculum gastrale (Figure
Larger than most other Bornean Ptomaphaminus, with spiniform elytral apices in the female, and a very narrow urite IX in the male; differing from the otherwise similar P. testaceus by the darker head and discus of the elytra, as well as slight differences in the aedeagus.
COI barcodes are available for the following specimen:
Widely distributed in South East Asia. Indonesia: Sumatra (type locality); Kalimantan (Gunung Marang). Malaysia: Continental Malaysia (Batu Caves in Selangor), Sabah (Kinabalu; Lower Kinabatangan), Sarawak (Gunung Jambusan; Gunung Mulu). It has been found in many caves, but also in forest litter and pitfall traps.
Variations can be observed in the curvature of the aedeagus. In the type series from Sumatra, the aedeagus is moderately curved, similarly to the population of Marang Mountains, while the population from Continental Malaysia have a more pronounced curvature. We consider that these differences do not exceed intraspecific variation.
Holotype: Malaysia, Sarawak, 4th Division, Gunung Mulu National Park, mixed dipterocarp forest, v–viii.1978 (leg. P.M. Hammond & J.E. Marshall,
Length: 1.4–1.8 mm. General colour light reddish brown; legs, antenna, and mouthparts yellowish. Winged. Eyes well developed. Pronotum 1.45 times as wide as long. Elytra 1.45 times as long as wide. Female sutural angle of elytra expanded backwards into a sharp tooth. Male protarsi 0.4 times as wide as the apex of protibia. Male genital segment with a very elongate thin spiculum gastrale (Figure
Very similar to P. latescens, but distinct in the external morphology by its significantly smaller size, and its lighter colour. The spermiduct is slightly helical, which is not the case in P. latescens.
Known only from the lowland forest of Gunung Mulu, Sarawak, Malaysia.
Named for its light brown colour (testaceus = brick-coloured).
Ptomaphaminus, aedeagus. a, c, e, g, i, k, n, p, r, t, v, x, z lateral view b, d, f, h, j, l, m, o, q, s, u, w, y dorsal view. a, b P. latescens c, d P. testaceus sp. n., Sarawak, Gunung Mulu e, f P. ater, paratype (CMPR). g, h P. chapmani, Sarawak, Gunung Mulu i, j P. sarawacensis sp. n., Sarawak, Gunung Mulu k, l P. hanskii sp. n., Sarawak, Gunung Mulu m, n P. nanus sp. n., Sarawak, Gunung Mulu o, p P. marshalli n. sp., Sarawak, Gunung Mulu q, r P. fagei holotype (CMPR) s, t P. layangensis sp. n., Sabah, Gunung Kinabalu, Layang-Layang u, v P. kinabatanganensis sp. n., Sabah, Kinabatangan valley, Gua Ikan w, x P. alabensis sp. n., Sabah, Gunung Alab y, z P. microphallus sp. n., Sabah, Kinabalu, Poring Hot Springs.
Ptomaphaminus, a, c, e, g, i, k, n, o, r, t, v, x male genital segments b, d, f, h, j, l, n, p, q, s, u, w, y female genital segments. a, b P. latescens c, d P. testaceus sp. n. e, f P. ater, paratype (CMPR) g, h P. sarawacensis sp. n., Sarawak, Gunung Mulu i, j P. hanskii sp. n., Sarawak, Gunung Mulu k, l P. fagei, paratype (CMPR) m, n P. kinabatanganensis sp. n. Sabah, Lower Kinabatangan, Gua Fico o, p P. layangensis sp. n. Sabah, Gunung Kinabalu, Layang-Layang q P. microphallus sp. n., Sabah, Kinabalu, Poring Hot Springs r, s P. chapmani, Sarawak, Gunung Mulu t, u P. nanus sp. n., Sarawak, Gunung Mulu. v, w P. alabensis sp. n., Sabah, Gunung Alab. x, y P. marshalli sp. n., Sarawak, Gunung Mulu.
Ptomaphaminus
ater
Perreau, 2009: 5, fig. 5; type from Gunung Kinabalu, Sabah, Borneo (in
Ptomaphaginus
ater
:
(In addition to that listed in
Length 1.75–2.00 mm. Large species, dark brown, winged. Eyes reduced, with 25 ommatidia. Pronotum ca. 1.5 times wider than long. Elytra approximately 1.2 times longer than wide. Female apex of the elytra with a sharp sutural angle expanded posteriorly (Figure
For two individuals,
Externally recognizable by its large size, dark colouration, and strongly cuneiform habitus. Apex of the female elytra spiniform. Spermatheca with a sclerotised plate at its apex. Aedeagus blunt-ended.
Known from high altitude on Gunung Kinabalu, above 3000 m. Some specimens were taken under a rocky overhang (Paka cave), others in Panar Laban (type locality) and Gunting Lagadan, without detail on collecting conditions.
The only species in Borneo with an apical sclerotised plate at the apex of the spermatheca (which occurs in several other species outside Borneo).
Holotype: Malaysia, Sabah, Lower Kinabatangan, Batu Batangan, Gua Kolam, 5°27.557'N, 118°06.118'E, 24.ii–1.iii.2016 (leg. I. Njunjić et al., BOR/COL/14218), 1 male. Paratypes: Sabah. Lower Kinabatangan, Batu Batangan, Gua Babi, 5°27.570'N, 118°06.088'E, 24–28.ii.2016 (leg. field course students, BOR/COL/14219–14220), 2 individuals; Lower Kinabatangan, Gua Fico, 5°27.135'N, 118°08.769'E, 18.iii.2015 (leg. I. Njunjić et al., CMPR), 1 female; Lower Kinabatangan, Batu Batangan, Gua Babi, N 5°27.570'N, 118°06.088'E, 24–28.ii.2016 (leg. I. Njunjić et al., CMPR), 1 female; Lower Kinabatangan, Batu Batangan, Gua Ikan, 5°27.558'N, 118°05.891'E, 18.iii.2015 (leg. I. Njunjić et al., CMPR), 2 males.
Length: 1.45–1.60 mm. General colour brown; antenna, mouthparts, and protarsi yellowish, the other tarsi light brown. Winged. Eyes well developed. Pronotum 1.55 times wider than long. Elytra 1.25 times longer than wide (slightly wider in males than in females). Female sutural apex of elytra angular but not protruding backwards. Male protarsi 0.65 times as wide as the apex of protibia. Spiculum gastrale of the male genital segment shortly protruding beyond the apex of epipleurites, significantly dilated (Figure
Female with spiniform elytra; male with long apical hook of the (relatively short but straight) aedeagus, which, however, is not retroverted.
Known exclusively from three caves in the lower Kinabatangan valley: Gua Babi, Gua Fico, and Gua Ikan.
Named after the lower Kinabatangan valley, in which the specimens were collected.
Ptomaphaginus
chapmani
Peck, 1981.
Sarawak. Mulu, Mayday Cave, rotten prawn bait, 24.vii.1980 (
Length 1.40–2.0 mm. Colour light brown. Winged. Eyes with ten ommatidia. Pronotum 1.47 times as wide as long, elytra 1.84 times as long as wide. Female sutural angle of elytra expanded in a sharp apical tooth (Figure
Unique among Bornean Ptomaphaminus in showing a combination of troglomorphic features: reduced eyes and elongated habitus. Aedeagus very similar to that of P. ater, which, however, is darker and has a more strongly cuneiform habitus.
Known only from a single cave (Clearwater cave) in Gunung Mulu, Sarawak, Malaysia.
Ptomaphaminus
fagei
Perreau, 2009.
Kalimantan Timur. Kebupaten Kutai Timur, karst of Mangkalihat, Mt Marang, Gua Sedepan, 8.vi.2002 (leg. M. Perreau, Expédition du Kalimanthrope, TXEX), 1 male, 1 female paratypes.
Length 1.50–1.90 mm. Colour light brown. Winged. Eyes with 15 ommatidia. Body very elongate, parallel-sided. Pronotum approximately 1.6 times wider than long. Elytra approximately 1.4 times longer than wide. Female sutural angle of elytra expanded backwards in a sharp tooth. Male protarsi 0.75 times as wide as the apex of protibia. Spiculum gastrale of the male genital segment expanded, with a triangular apical part (Figure
Among the species with similarly reduced eyes and/or spiniform female elytral apices, P. fagei is unique in having an aedeagus that shows an abrupt narrowing (in lateral view) in the apical third.
Known from two caves of Gunung Marang, Kalimantan, Indonesia: Gua Sedepan and Gua Gala.
Holotype: Malaysia, Sarawak, 4th Division, Gunung Mulu National Park, mixed dipterocarp forest litter, v–viii.1978 (leg. P.M. Hammond & J.E. Marshall, in
Length: 1.28–1.50 mm. General colour brown; tarsi, antenna, mouthparts yellowish. Winged. Eyes well developed. Pronotum1.6 times as wide as long. Elytra 1.2 times as long as wide. Elytral internal angle rounded in male and in female, without noticeable sexual dimorphism. Male protarsi 0.6 times as wide as the protibia. Spiculum gastrale of the male genital segment dilated into a narrow spatula (Figure
Small-sized species with normally developed eyes and non-spiniform female elytra. Spermiduct tightly coiled; aedeagus small; distinguishable from P. marshalli, which has an equally small aedeagus, by the very short hook.
Known from lowland forests of Gunung Mulu, Sarawak, Malaysia.
A very small species, one of the smallest species of the genus.
Named for its very small size (nanus = dwarf).
Holotype: Malaysia, Sarawak, 4th Division, Gunung Mulu National Park, Slope, 29.4.78, Tps 4–6, v–viii.1978 (leg. P.M. Hammond & J.E. Marshall, in
Length: 1.50–1.90 mm. General colour brown; the tarsi, the two first antennomeres, the base of the third antennomere, and of the tibiae yellowish. Winged. Eyes well developed. Pronotum 1.65 times as wide as long. Elytra 1.25 times as long as wide. Female sutural apex of elytra rounded (Figure
Females of this species can be recognized by the apex of the spermatheca, which is conically narrowed, a very distinct morphology compared to all other species, which have a spermatheca with a widely rounded apex.
Known from lowland forests of Gunung Mulu, Sarawak, and of Gunung Kinabalu, Sabah, Malaysia.
Dedicated to J. E. Marshall, one of the collectors of the species during the expedition of the Natural History Museum of London in Sarawak.
Holotype: Malaysia, Sarawak, 4th Division, Gunung Mulu National Park, Slope, 4.5.78, 9.1, 9.3, v–viii.1978 (leg. P.M. Hammond & J.E. Marshall, in
Length: 1.6–2.1 mm. General colour brown; the tarsi and two first antennomeres yellowish. Winged. Eyes well developed. Pronotum 1.65 times as wide as long. Elytra 1.30 times as long as wide. Sutural angle of female elytra angular. Apex of the spiculum gastrale of the male genital segment dilated into a diamond-like form (Figure
Species with normally developed eyes and a long aedeagus. Very similar to P. sarawacensis, from which it cannot be confidently distinguished in the female sex. Males of P. hanskii have a thicker median lobe of the aedeagus than P. sarawacensis.
Known from lowland forests of Gunung Mulu, Sarawak, Malaysia.
The external morphology and the female genital morphology are extremely similar to P. sarawacensis, so that females are nearly impossible to distinguish. However, the male aedeagi of these species are very different.
Named in honour of Ilkka Hanski, the Finnish ecologist who played an important role in the Royal Geographical Society expedition to Mulu of 1978, and who passed away in 2016.
Holotype: Malaysia, Sarawak, 4th Division, Gunung Mulu National Park, Slope, 4.5.78, 9.1, 9.3, v–viii.1978 (leg. P.M. Hammond & J.E. Marshall, in
Length: 1.90–2.35 mm. General colour brown; the tarsi and two first antennomeres yellowish. Winged. Eyes well developed. Pronotum 1.6 times as wide as long. Elytra 1.4 times as long as wide. Sutural angle of female elytra angular (Figure
See under P. hanskii (above).
Known from lowland forests of Gunung Mulu, Sarawak, Malaysia.
Extremely similar to P. hanskii, except in aedeagus shape.
Named for the Malaysian state of Sarawak, where the type locality lies.
Ptomaphaginus sp. n. bryanti complex Merckx et al., 2015: extended data fig. 6.
Holotype: Malaysia, Sabah, Kinabalu Park, Layang-Layang, 2750 m elev., 6°02.748'N, 116°33.632'E, 13–18.ix.2012 (leg. M. Schilthuizen, Crocker Range / Kinabalu Expedition,
Length: 1.6–2.5 mm. General colour dark brown; the tarsi and two first antennomeres lighter. Winged. Eyes well developed. Pronotum 1.6 times as wide as long. Elytra 1.45 times as long as wide. Sutural angle of female elytra rounded, without noticeable sexual dimorphism. Male protarsi 0.75 times as wide as the protibia. Spiculum gastrale of the male genital segment short, moderately dilated at the apex (Figure
Normal-sized species with unreduced eyes and non-spiniform female elytra. Aedeagus with a long, retroverted apical hook. Distinguishable from P. alabensis by the anteriorly widened spiculum gastrale and the broader apex of the median lobe of the aedeagus. Females are distinguishable from P. nanus by their larger size.
COI barcodes are available in BOLD for the holotype,
Known from high altitude (above 2000 m) on Gunung Kinabalu, Sabah, Malaysia.
Named for the type locality on Gunung Kinabalu.
Holotype: Malaysia, Sabah, Kinabalu Park, Poring Hot Springs, 6°02.894'N, 116°41.957'E, 625 m elev., in baited pitfall traps, 15–20.ix.2012, (leg. M. Schilthuizen, Crocker Range / Kinabalu Expedition), 1 male in
Length: 1.2 mm. General colour dark brown; the tarsi and two first antennomeres lighter. Winged. Pronotum 1.55 times as wide as long. Elytra 1.15 times as long as wide. Male protarsi 0.6 times as wide as the protibia. Spiculum gastrale of the male genital segment apically dilated into a short discoid expansion (Figure
The female is unknown, but the male aedeagus shares several features with other species, such as P. marshalli, P. nanus, and P. alabensis (i.e., a relatively short aedeagus with retroverted hook). However, P. microphallus is unique among Borneo Ptomaphaminus by its extremely short (0.22 mm) aedeagus.
Known from the type locality, in Kinabalu Park, Sabah, Malaysia.
Female unknown.
Named for the relatively small male genitalia.
Ptomaphaginus nr. fagei Merckx et al., 2015: extended data figs 2, 6.
Holotype: Malaysia, Sabah, Crocker Range, Gunung Alab, 5°47.766'N, 116°20.504'E, 1930 elev., baited pitfall trap, 17–22.iv.2012 (leg. M. Schilthuizen,
Length: 1.5–1.6 mm. General colour dark brown; the tarsi and two first antennomeres yellowish. Winged. Pronotum 1.6 times as wide as long. Elytra 1.33 times as long as wide. Sutural angle of female elytra rounded, without noticeable sexual dimorphism. Male protarsi 0.8 times as wide as the apex of protibia. Spiculum gastrale of the male genital segment straight, without apical dilation. Median lobe of the aedeagus 4.7 times shorter than the body length. Apex of the median lobe with a ventrally deflexed, arcuate hook. Aedeagus regularly narrowed from base to apex in dorsal view, ventrally straight (Figure
Females are easily recognized by the spermatheca which has a helical base and the spermiduct short, not helical. When helical structures occur in the female genitalia in other species, this affects the spermiduct, and not the spermatheca.
In BOLD, COI barcodes area available for the holotype,
Known from the type locality, on Gunung Alab, Crocker Range, Sabah, Malaysia.
Named for the type locality, Gunung Alab in the Crocker Range.
1 | Mesocoxal cavities attached to one another. Protibiae without a long row of equal spines along the external side. Male first mesotarsomere dilated. Epistomal suture absent | 2 |
– | Mesocoxal cavities separated by a mesoventral process. Protibiae with a long row of equal spines along the external side. Male first mesotarsomere undilated. Epistomal suture present (Ptomaphagini) | 3 |
2 | Elytral surface with punctuation aligned in transverse strigae. Male urite IX entire (Anemadini, Nemadina) | Micronemadus sondaicus sp. n. |
– | Elytral surface with uniformly spaced punctuation, without transverse strigae. Male urite IX reduced (Cholevini, Catopina) | Catops pruinosus Schweiger |
3 | Female protarsi tetramere and widely dilated (Figure |
Baryodirus hammondi Perreau |
– | Female protarsi pentamere and undilated. Mesoventral process narrow, the ventral side sharp-edged (Ptomaphaginina) | 4 |
4 | Ventral spines of protibiae randomly arranged. Metaventral sutures convergent towards the central axis of the body | Ptomaphaminus Perreau |
– | Ventral spines of protibiae aligned along the latero-external row of equal spines, making a second, more widely-spaced row next to the external one. Metaventral sutures roughly parallel to the axis of the body | Ptomaphaginus Portevin |
1 | Body length more than 3.3 mm | 2 |
– | Body length less than 3.1 mm | 4 |
2 | Elytra more than two times as long as the pronotum | 3 |
– | Elytra not more than two times as long as the pronotum | P. bryantioides Schilthuizen & Perreau (exceptionally large individuals) |
3 | Female: apical capsule of spermatheca annulated, U-shaped, spermiduct not extremely long, consisting of 5–6 coils (Figure |
P. grandis sp. n. |
– | Female: apical capsule of spermatheca not annulated, semicircular, spermiduct extremely long, consisting of ca. 30 coils (Figure |
P. isabellarossellini sp. n. |
4 | Aedeagus with two lateral “flaps” at the apex and usually a median processus, the apex thereby appearing bilobate or trilobate (Figs |
5 |
– | Aedeagus tip upturned or flattened, sometimes with a median processus, but not clearly bi- or trilobate (Figs |
10 |
5 | Habitus narrow and elongated, relatively flat. Pronotum 1.5–1.6 times as wide as long, narrowing in a more rectilinear fashion from caudal to rostral (Figure |
6 |
– | Habitus broader. Pronotum 1.7–1.9 times as wide as long, narrowing in a gradual curve from caudal to rostral | 7 |
6 | Aedeagus apically with two long lateral flaps that jointly are more than half the width of the basal part of the aedeagus, and a long median processus. Female unknown | P. bryanti Jeannel |
– | Aedeagus apically with two short, triangular lateral flaps, jointly less than half the width of the basal part of the aedeagus, without a clear median processus (Figure |
P. caroli Schilthuizen & Perreau |
7 | Aedeagus in dorsal view slender, more than two times as long as wide, at the apex with a long (longer than each of the lateral flaps), caudally pointing, median processus (Figure |
P. bryantioides Schilthuizen & Perreau |
– | Aedeagus in dorsal view short and broad, less than two times as long as wide, if at the apex with a caudally pointing median processus, then this is shorter than each of the lateral flaps; spermatheca sometimes with ring-shaped constrictions | 8 |
8 | Male protarsi not dilated, less than one-third the width of the apex of the protibia; as narrow as in the female; spermatheca with multiple, ring-shaped constrictions | P. similipes Schilthuizen & Perreau |
– | Male protarsi moderately to strongly dilated, at least half the width of the apex of the protibia; spermatheca unknown | 9 |
9 | Male protarsi moderately dilated, about half as wide as the apex of the protibia; elytral apices rounded (male) or drawn out (female) | P. kinabaluensis Schilthuizen & Perreau |
– | Male protarsi strongly dilated, as wide as the apex of the protibia; elytral apices truncated in both sexes | P. latimanus Schilthuizen & Perreau |
10 | Antennae short and broad, antennomeres 9 and 10 twice as long as wide; aedeagus inflated and strongly convex; spermatheca with multiple ring-shaped constrictions (Figure |
P. scaphaner Szymczakowski |
– | Antennae slender, antennomeres 9 and 10 square or only slightly wider than long; aedeagus not inflated and highly convex; (spermatheca not known for all species) | 11 |
11 | Eyes reduced (each eye only one-tenth of the width of the head), wingless; female unknown | P. burckhardti Schilthuizen & Perreau |
– | Eyes normally developed (each eye ca. one-sixth of the width of the head), winged | 12 |
12 | Aedeagus in dorsal view quadrangular, two times as long as wide (Figure |
P. sabahensis Schilthuizen & Perreau |
– | Aedeagus in dorsal view narrowed caudally, either gradually or abruptly pointed, more than two times as long as wide | 13 |
13 | Upturned tip of the aedeagus sharp, pointed (Figs |
14 |
– | Upturned tip of the aedeagus broadly flattened and rounded (Figure |
P. anas Schilthuizen & Perreau |
14 | Apex of the aedeagus abruptly narrowed (Figure |
P. louis sp. n. |
– | Apex of the aedeagus gradually narrowed; pronotum as wide as the elytra at the shoulders, caudal angles distinctly drawn out (Figure |
P. muluensis sp. n. |
1 | Basal loop of the endophallus symmetric and stylus short and more strongly sinuate, located in the apical half of the median lobe (Figure |
2 |
– | Basal loop of the endophallus asymmetric and stylus long and weakly sinuate, extended over most of the length of the median lobe. Anterior half of the spiculum gastrale of the male genital segment generally dilated into a paddle-shape (except P. alabensis sp. n.). Apex of aedeagus with a more or less developed hook-shaped expansion visible in lateral view. Female gonocoxites short, approximately as wide as long. Female sutural apex of elytra rounded, angular or spiniform | 3 |
2 | Head and discus of the elytra noticeably darker brown than the rest of the body. Male protarsi more expanded, 0.5 times as wide as the apex of protibia. Spermiduct irregular | P. latescens Szymczakowski |
– | Entire body light reddish brown. Male protarsi less expanded, 0.4 times as wide as the apex of protibia. Spermiduct slightly helical | P. testaceus sp. n. |
3 | Protarsus dilated (males) | 4 |
– | Protarsus undilated (females) | 14 |
4 | Eyes reduced (<50 ommatidia) | 5 |
– | Eyes normally developed (>100 ommatidia) | 7 |
5 | Aedeagus long (more than 0.5 mm). Apical hook of the aedeagus long and retroverted (Figure |
6 |
– | Aedeagus short (~0.35 mm). Median lobe abruptly narrowed in lateral view on the apical third of its length, with a short apical hook. (Figure |
P. fagei Perreau |
6 | Apex of the aedeagus rectangular, the lateral angles sharp (Figure |
P. chapmani Peck |
– | Lateral angles of the apex of the aedeagus rounded (Figure |
P. ater Perreau |
7 | Aedeagus long, more than 0.5 mm | 8 |
– | Aedeagus short, less than 0.4 mm | 9 |
8 | Median lobe of the aedeagus thicker, regularly arcuate along its whole length (Figure |
P. hanskii sp. n. |
– | Median lobe of the aedeagus thinner, slightly angular at the second third of its length (Figure |
P. sarawacensis sp. n. |
9 | Apical hook of the median lobe of aedeagus short (Figure |
P. nanus sp. n. |
– | Apical hook of the median lobe of aedeagus longer | 10 |
10 | Apical hook of the aedeagus retroverted | 11 |
– | Apical hook of the aedeagus not retroverted (Figure |
P. kinabatanganensis sp. n. |
11 | Apical hook of the median lobe of the aedeagus thick (Figure |
P. microphallus sp. n. |
– | Apical hook of the median lobe of the aedeagus thin | 12 |
12 | Apical hook of the median lobe of aedeagus angular (Figure |
P. marshalli sp. n. |
– | Apical hook of the median lobe of aedeagus roundish | 13 |
13 |
Spiculum gastrale of the male genital segment slightly expanded in the anterior part (Figure |
P. layangensis sp. n. |
– |
Spiculum gastrale of the male genital segment not expanded in the anterior part (Figure |
P. alabensis sp. n. |
14 | Sutural apex of elytra sexually dimorphic, rounded in males, angular or spiniform in females (Figure |
15 |
– | Sutural apex of elytra not sexually dimorphic, rounded in females as in males (Figure |
19 |
15 | Eyes reduced (<50 ommatidia). Sutural apex of elytra spiniform (Figure |
16 |
– | Eyes normally developed (>80 ommatidia). Sutural apex of elytra simply angular (Figure |
18 |
16 | Spermiduct not helical (Figure |
P. chapmani Peck |
– | Spermiduct helical | 17 |
17 | Spermatheca with an apical sclerotised plate (Figure |
P. ater Perreau |
– | Spermatheca without apical plate (Figure |
P. fagei Perreau |
18 | Spermiduct not helical (Figure |
P. kinabatanganensis sp. n. |
– | Spermiduct membranous, vaguely helical near the base of the spermatheca (Figure |
P. hanskii sp. n. and P. sarawacensis sp. n. |
19 | Spermiduct not helical | 20 |
– | Spermiduct helical | 21 |
20 | Terminal lobe of spermatheca regularly narrowed, the apex conical (Figure |
P. marshalli sp. n. |
– | Apex of spermatheca rounded, base of the spermatheca helical (but not the spermiduct) (Figure |
P. alabensis sp. n. |
21 | Very small size, <1.3 mm. Spermiduct with many coils tightly compacted (Figure |
P. nanus sp. n. |
– | Normal size, >1.5 mm. Spermiduct with coils regularly spaced, not tightly packed (Figure |
P. layangensis sp. n. |
We are greatly indebted to Louis Deharveng for accommodating us in his lab at the MNHN, Paris. M.S. and I.Nj. acknowledge the support from Sabah Parks (especially Dr. Maklarin Lakim and Mrs. Rimi Repin), the Sabah Wildlife Department, the Sabah Forestry Dept. (especially Dr. Arthur Chung), and the Danau Girang Field Centre (especially Dr. Benoit Goossens and Ms. Danica Stark). We thank Hans Huijbregts (