Distinguishing the Sundaland species in the Onthophagus (Parascatonomus) aurifex group (Coleoptera: Scarabaeidae: Scarabaeinae)

J. Krikken & J. Huijbregts

The taxonomic position and diversity of the aurifex species group in the scarab genus Onthophagus Latreille, 1802, subgenus Parascatonomus Paulian, 1932, is discussed. The O. aurifex group is rediagnosed, and a key to the six known Sundaland representatives of the group is given. Two new species from North Sumatra are described and illustrated: O. semifex and O. sumawacus. A lectotype is designated for O. semiaureus Lansberge, 1883, the type locality here being restricted to West Java. Attention is drawn to the variation in this species and in O. aurifex Harold, 1877. Records are given for members of the aurifex group; several published records need confirmation. The species are associated with dung and carrion. J. Krikken & J. Huijbregts,National Museum of Natural History Naturalis, Postbus 9517, NL-2300 RA Leiden, The Netherlands. krikken@nnm.nl

Introduction
In dung and carrion beetle collections from Southeast Asia, submitted for identification over the past three decades by ecological colleagues, we recognized several undescribed species of the near-cosmopolitan scarab genus Onthophagus Latreille, 1802, one of the largest genera in the animal kingdom. In this paper two new species from North Sumatra (Aceh) are described, both related to O. aurifex Harold, 1877. Boucomont (1914), in his classic review, made this species the type of his group 6, the aurifex group, including three more species (all mentioned below). The Gunung Leuser National Park, where the new Onthophagus were found in 1972 and 1983, is an extensive area of lowland to subalpine ecosystems. At the time of the collecting activities the Park was covered with much pristine forest, home to our novelties as well as conservation icons like orang-utan, elephant, rhino, tapir, and tiger some now perhaps locally (nearly) extinct. Onthophagus are no different from many other groups of Southeast Asian animals and plants in that species occurring in primeval forest are, as a rule, less widespread than are their close relatives in open (i.e.,

usually man-made, monsoonal or soil-determined) habitats. Consequently, full multi-island taxonomic overlap among these forest scarabs, is uncommon, and this seems to be the case here as well: the two new North Sumatran forest species appear to have very similar but taxonomically different relatives on Borneo and elsewhere. In the aurifex group we seem to be dealing with complex sets of geographic and ecological vicariants, and it may well be that unresolved character differences (for instance, variation in pilosity and parameral structure) represent more than intraspecific variability. Published critical records and available collection series for the group, however, are limited; hence our inevitable question marks behind the published regional records below (see the species list), indicating points of interest for future revisers. The variation of the group member O. semiaureus Lansberge, 1883 is another case in point, and it may not be by chance that this multi-island, more variable species occurs mainly in secondary habitats. The Leiden museum has syntypical material of semiaureus, and since this paper is a contribution to disentangling species identities in the aurifex group, the

Tijdschrift voor Entomologie 151: 173185, Figs. 1 38. [ISSN 00407496]. http://www.nev.nl/tve 2008 Nederlandse Entomologische Vereniging. Published 1 December 2008.

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designation of a lectotype for this variable species is appropriate. Characteristic for the members of the Onthophagus aurifex group of species, as here understood, are the presence of a broad, short tooth on the middle of the clypeal margin, the lack of distinct protrusions (like horns and ridges) on the head surface, the punctate (non-granulate) pronotal disc, and the strongly angulate pronotal base (for full group diagnosis, see below). Note that in the present definition, for purely operational reasons, we have excluded O. denticollis Lansberge, 1883, and any of its immediate relatives, which lack the tooth on the clypeal margin and are smaller. The species of the aurifex group have been placed in the subgenus Parascatonomus Paulian, 1932, for instance by Kon et al. (2000), thus deviating from the limited subgeneric concept of Balthasar (1963). Meanwhile, many more species were transferred to, or described as new in, various other species groups allocated to Parascatonomus. Certain authors have gone further by raising Parascatonomus to full genus (lately, Kabakov 2006, see also his earlier papers, like Kabakov 1992), but we cannot accept such a far-reaching action without support from a broad-scale approach to the supraspecific classification of the hyperdiverse Onthophagini, with phylogenetic underpinning. Nevertheless, until this has happened, the Onthophagus aurifex group can, in our view, safely be accommodated in an operationally defined subgenus Parascatonomus (our definition is implicit in the group diagnosis below). Technical remarks The primary purpose of this paper (part of an ongoing series on the taxonomy of laparostict scarabs of the Southeast Asian islands) is the straightforward description and diagnosis of the new taxa, and a comparison with their closest relatives by means of an identification key and comparative pictures. The Sundaland region is here defined as the Malay Peninsula + Sumatra + Java + Borneo + Bali + nearby and intervening islands. Pictures are considered more useful than abstract descriptive text, and consequently, in addition to habitus pictures, at least six elements are depicted for each new species and its closest relative: head (fullface), pronotum and left elytron (dorsal), protibia (upperside), metatibia (underside), and aedeagus (here mainly in lateral view). The photographs are not all to the same scale; refer to the species accounts for measurements of material pictured. The pilosity, as depicted on the legs, is not diagnostic. The term full-face view is applied where the upper surface of the element concerned is maximally parallel to the

plane of the picture (like the head). The qualification abundant for microsculptural units (like punctures) usually means: separated by 2-5 diameters, dense: 1-2 diameters, crowded: less than 1 diameter; sparse: means separated by at least 5 diameters, or limited to certain parts only. The type of punctation is qualified only if it is decidedly different from a simple punctation. The prefix micro- usually refers to sculpture and pilosity distinct at magnifications x40 and higher. The term propectus denotes the underside of the prothorax. The morphology of the male genitalia is of foremost importance in species recognition; casual identifiers should not have the illusion to be able to recognize species without critically looking at the aedeagi of the males they handle. In particular, in the aurifex group, note the more strongly sclerotized distal piece on each paramere, here termed paramerite; its shape and orientation are important characters. Note that strict viewing angles apply (pointed parts may not appear as such when viewed from a different angle). Ad hoc terminology is explained in the figures. Always beware of deceptive intraspecific differences between major males, minor males, and females use series including fresh major males whenever possible. Males have foreshortened abdominal sternites. We suspect polymorphism in the distribution and nature of the pilosity over the body. Also note that older, worn specimens may have lost (part of) their pilosity. The new material recorded in this paper, currently kept in the National Museum of Natural History, Leiden, The Netherlands (RMNH), was collected by baited pitfall trapping (some aurifex group members, like other Parascatonomus, have a preference for carrion, but see ecological notes under species accounts and in Annex 1). In this paper we distinguish two altitudinal qualifications, lowland and upland, roughly under and above 1000 m (subalpine altitudes, above 2000 m, were rarely sampled). Our comments on information from other sources are placed between square brackets (for instance in the species list in the next section). Characters in the following group diagnosis are arranged in their estimated order of taxonomic importance.

The Onthophagus (Parascatonomus) aurifex group in Sundaland

Group diagnosis. Clypeus (in both sexes) with sin-

gle anteromedian, reflexed, usually well-delimited broad tooth (its tip rounded or obtusely angular), not standing in deep, abruptly delimited emargination, clypeal edge widely, evenly rounded to lateral (subangular) tips of genae. Surface of head of male lacking higher protrusions, (at most) with

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obsolescent clypeofrontal and/or vertexal transverse ridge. Pronotal disc and anterior declivity simply punctate (not granulate, rugulate, or asperate). Elytral interstriae unmodified, simply punctate (pilosity in punctation varies, but never with long erect setae). Metasternum raised in front, prow-shaped, midline carinate. Pronotum without isolated protrusions, anterior declivity may be slightly depressed or deplanate (in one species, liewi, topped by transverse, arcuate crest). Base of pronotum distinctly angulate medially, largely immarginate. Anterolateral angle of prothorax slightly angular, rounded off, not truncate or otherwise modified. Antennae relatively short; scapus not serrate, indistinct from above; club compacted, club segment 1 somewhat cup-shaped. Parameres relatively complex, apex blunt, with sclerotized (darker brown), more or less projecting distal paramerite. Meso- and metafemora broad, complanate. Meso- and metatibiae with more or less straight (not strongly lobate) apex. Metatarsal segment 1 long, robust, length approaching segments 2-5 combined. Eyes in full-face view narrowly subelliptic. Protibia with four outer denticles separated by serration; terminal spur dilated, not simply acuminate. Elytral striae 6 and 7 of elytra subparallel, straight or evenly curved near humerus (not sinuate). Body waisted. Colour of body and legs entirely or partly (brown-) black, no light-dark pattern, dorsal side of forebody black(-brown), or conspicuously metallic (green, blue, cupreous). Pygidial base marginate. Sexual dimorphism slight (abdominal sternites in male foreshortened). Medium-sized, body length 6-14.5 mm. (O. denticollis Lansberge, 1883, without clypeal tooth, omitted from the present group definition, the key, and the list.)

Key
1. Dorsum entirely black, generally matt (pronotal disc and elytra microreticulate), on average larger species. Elytra virtually glabrous. Interstrial punctures fine (distinctness of microstubbles dependent on illumination) . . . 2 Forebody shiny, metallic (usually green or cupreous). Elytra dark-brown or black, matt (microreticulate), on average smaller species. Interstrial punctures (at least laterally) with numerous long pale setae (very distinct at x20) or with microstubbles, or virtually glabrous (at least on disc) . . . . . . . . . . . . . . . . . . . . . . 4 2. Clypeal median denticle on either side delimited by sinuate emargination (fig. 37). Pronotum with (indication of ) transverse arcuate ridge topping anterior declivity (large males). Length 10.5-14.5 mm . . . . . . . . . . . liewi Clypeal median denticle lacking sinuate emargination on either side (fig. 38). Pronotum with more or less evenly declivous anterior side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3. Pronotal and interstrial punctures on elytra extra fine. Parameres as in fig. 32., finely multiple-pointed distal paramerite standing out laterally (fig. 32a). Length 10-14 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . sumawacus Pronotal and interstrial punctures on elytra less fine. Parameres as in fig. 33, with coarsely multiple-pointed distal paramerite (lateral view). Length 10-14 mm . . . . . . . . . . . . sarawacus 4. Elytral interstriae laterally at most with microstubbles or microsetae, disc virtually glabrous. Pronotum glabrous. Parameres as in fig. 36. Distal paramerite variably, finely multiple-pointed, standing out laterally (fig. 36a). Length 8-12 mm . . . . . . . . . . . . . semiaureus All elytral interstriae as well as pygidium evenly covered with distinct, rather long pale-white to pale-brown subappressed setae in distinct punctures. Parameres as in figs 34 & 35. Distal paramerite not multiple-pointed, not standing out laterally (indistinct from above). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5. Parameres as in fig. 34, with distal paramerite shifted onto apex (fig. 34a), space between paramerite and basolateral plate not excised. Length 10-12 mm . . . . . . . . . . . . . . . . . . . . semifex Parameres as in fig. 35, with distal paramerite distinct in lateral view, space between paramerite and basolateral plate excised. Length 9-12 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . aurifex

Key to species
Important characters Shape of distal paramerite (denoted dpa in the figures; orientation; distribution of angular points) and other parameral elements (like the basolateral plate, denoted blp in the figures)  Colour and reflection of forebody (head and pronotum, simply black versus metallic (degree of reflection = surface microsculpture, see 3)  Microsculpture (pronotum and elytral interstriae, size and density of punctation, background microreticulation)  Pilosity and micropilosity (particularly on elytra, distribution of setae, setal length)  Shape of clypeal margin (median tooth and any emargination on either side)

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12

10

3a

9a

Figs 1-12. Habitus and body elements of: 1-6, Onthophagus sumawacus (holotype), 1, habitus, 2, head, 3, pronotum, with 3a, left anterolateral corner (full-face), 4, left elytron, 5, protibia, 6, metatibia; 7-12, sarawacus (from Danum), 7, habitus, 8, head, 9, pronotum, with 9a, left anterolateral corner (full-face), 10, left elytron, 11, protibia, 12, metatibia.

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13

20

17 14 21

24

15

22 18 25

16

23

19a dpa

19a

Figs 13-25. Habitus and body elements of: 13-19, Onthophagus semifex (holotype), 13, habitus, 14, head, 15, pronotum, 16, left elytron, 17, protibia, 18, metatibia, 19a, aedeagal tip (full-face), and 19b, aedeagus (lateral view) [compare with fig. 34, dpa = distal paramerite]; 20-25, aurifex (from Mulu), 20, habitus, 21, head, 22, pronotum, 23, left elytron, 24, protibia, 25, metatibia.

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List of species
Exclamation marks denote confirmations based on material actually seen by us; question marks denote verification of material required.

Onthophagus
aurifex Harold, 1877 Borneo (Sarawak!, Kalimantan!), Malay Peninsula [?], Sumatra [?, including different taxon?], Java [?], Thailand [?, in Kabakov & Napolov 1999] liewi Ochi & Kon, 2005 Borneo (Sabah) sarawacus Harold, 1877 Borneo (Sarawak!, Sabah!), Sumatran West Coast Islands [? Batu Islands] semiaureus Lansberge, 1883 Java!, Bali!, Borneo (Sarawak!, Sabah!, Kalimantan!), Sumatra [Central!, South!] plus West Coast islands [?], Sulawesi [?], Philippines [?], Thailand [?] semifex sp.n. North Sumatra sumawacus sp.n. North Sumatra

Species accounts
Onthophagus (Parascatonomus) sumawacus Krikken & Huijbregts, sp. n.
Figs 1-6, 32 Holotype. Male, in RMNH, with label data as fol-

lows: [Indonesia], North Sumatra: Gng Leuser NP: Lawe Mamas, 10-17/ix/1983, H Risnen, 40Cb, 1530-1550 m, 320N-9744S, tropical lower montane rainforest, pitfall trap, human dung. Paratypes as listed in Annex 1. Description (holotype, male) Body length ca 12.5 mm. Almost entirely black to black-brown, dorsum very matt, more or less sericeous. Head without distinct non-marginal protrusions, wider than long. Clypeal margin anteriorly distinctly reflexed; apex with short tooth, its tip rounded, laterally simply sinuate. Clypeofrontal surface generally flat. Clypeofrontal transition without distinct elevation (only smooth, slightly raised strip present). Clypeofrons black, surface glabrous, generally (more or less transversely) rugulate-punctate. Clypeal border from gena forward rounded; clypeogenal margin (at and near suture) widely rounded. Genae laterally subangulate. Frontovertex entirely devoid of protrusions (basal transverse ridge obsolescent). Eyes (foramen in full-face view) small, more or less narrowly elliptic. Maximum number of facet rows (transversely) across eye foramen ca 12. Ratio transverse interocular distance/maximum eye width 6.5. Pronotum generally evenly convex, disc and anterior

declivity unmodified. Prothorax in dorsal outline with sides evenly rounded over at least anterior 0.7, slightly sinuate posteriorly. Anterior border of pronotum marginate. Anterolateral pronotal angle slightly obtuse, rounded off. Lateral sides of pronotum simply declivous, marginate. Posterolateral section of pronotum strongly sinuate; pronotal posterolateral angles simple, distinct (but very obtuse). Base of pronotum immarginate, very distinctly angulate medially, surface slightly depressed. Pronotum glabrous, generally punctate (fine on disc, punctures laterally and anterolaterally larger; microstubbles in them scarcely distinct), abundant to dense (sparser on disc, very fine there). Estimated number of primary punctures per 0.25 sq mm on pronotal disc 30-35, diameters ca 0.01 mm, mostly separated by 5-10 diameters. Pronotum black, matt, microreticulate. Elytra black, matt, microreticulate, virtually glabrous, with hardly distinct microstubbles only. Sutural base slightly pressed apart, accommodating pronotal angle. Humeral umbone weakly pronounced. Punctures of (discal) elytral striae slightly impressed (roundish, fine, crenulating interstriae, separated by 4-6 diameters); striae distinct, shallowly impressed, narrow, sharply defined. Discal interstriae flat, simply, very finely punctate and microreticulate. Interstrial punctures separated by 5-10 diameters, not arranged in rows, more or less scattered, sparse. Antennal lamellae light-brown. Antennal scapus smooth. Propectus laterally black, abundantly punctate-setose to ocellate-punctate-setose (back side). Metasternum black; anterior lobe granulate, protruding (prow-shaped). Metasternal disc finely, sparsely to abundantly punctate, with microstubbles. Metasternum laterally ocellate-punctate-setose, abundant to crowded. Setae on propectoral sides very long, abundant to dense. Abdominal venter black, matt, microreticulate, coarsely ocellate-punctate-setose, on proximal sternites along base only. Pygidium slightly convex, marginate along base, black, matt, microreticulate, densely ocellate-punctate-setose, length of setae mostly not exceeding punctural diameter. Profemur black-brown, shiny, abundantly punctate, partly with long setae, broad, with distinct ridge on anterior side. Protibia robust, black-brown, shiny, externally with 4 denticles (1 short, 2-4 long, acuminate, 4 slender, all separated by slight serration). Proximal serration on outer side of protibia distinct. Protibia with apex oblique, apico-internal angle obtuse, with internal side evenly, slightly curved. Protibial spur inserted near inner angle, elongateacuminate (robust, largely parallel-sided). Meso- and metafemora broad, complanate, black-brown, shiny,

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abundantly finely punctate, with microstubbles. Meso- and metatibiae distinctly dilated distad, with fossorial protrusions on outer side; fossorial protrusions transverse. Mesotibial spurs simply acuminate (very robust, long). Metatibial apex (virtually) straight (edge lined with spines). Metatibial spur simply acuminate (very robust, long). Meso- and metatibiae black-brown, shiny. Metatarsal segment 1 robust (slightly curved, slightly longer than tarsal segments 2-5 combined); approximate length proportions of terminal spur versus tarsal segments 1-5: 29//31/9/6/4/7. Measurements in mm. Approximate maximum width of head 3.7; median length of pronotum (dorsal view) 4.7, maximum width 6.5; sutural length of elytra (dorsal view) 5.0, maximum width of elytra combined 7.0. Parameres, figs 32, distal paramerite finely multiangulate. Diagnosis Compared to O. sarawacus, this new species has distinctly finer pronotal and elytral punctures, the intervening microreticulate surface being definitely less reflective. Anterolateral surface of pronotum simply punctate. Dorsum entirely matt, black, i.e. forebody not metallic. Elytra virtually glabrous, microstubbles hardly distinct. Aedeagus, figs 32, shape and orientation of distal paramerite characteristic. Clypeal tooth lacking emargination on either side, contrary to O. liewi (see below). Length 10-14 mm. Distribution and ecology North Sumatra, both lowland and upland forest (350-1550 m altitude), mostly collected in human faeces baited traps.
Derivation of species name

mentioned hereafter, the clypeal tooth is poorly delimited on either side, i.e. by a slightly sinuate curve (fig. 38). Length 10-14 mm. Measurements in mm of pictured male (Danum). Approximate maximum width of head 3.8; median length of pronotum (dorsal view) 5.0, maximum width 6.8; sutural length of elytra (dorsal view) 5.0, maximum width of elytra combined 7.1; total length ca 13. Distribution and ecology Borneo (type locality Sarawak), apparently in lowland forest, mostly in fish baited traps. According to Davis et al. (2001) an interior forest specialist. ExtraBornean records to be confirmed (for the identity of Vietnamese sarawacus-like species, see Kabakov 1992).

Onthophagus (Paracatonomus) liewi Ochi & Kon, 2005

Fig. 37

Diagnosis

This species is immediately recognizable from the shape of its clypeal apex, pronotal anterior impression and ridge (in large males), and aedeagus (see various figures in Ochi & Kon 2005). The clypeal tooth is on either side well defined by a sinuate emargination (fig. 37). Forebody black, not metallic. Elytra virtually glabrous, also black. Length 10.5-14.5 mm.
Distribution and ecology

Composition of Sumatra, home island of the species, and the name of the related species sarawacus.

Sabah, in upland situations on and around Mt Kinabalu no further ecological information with the original description. Perhaps an upland vicariant of O. sarawacus, and possibly limited in space and/or time, as we did not encounter O. liewi in our Sabah samples.

Onthophagus (Parascatonomus) sarawacus Harold, 1877

Figs 7-12, 33, 38

Onthophagus (Parascatonomus) semifex Krikken & Huijbregts, sp. n.

Figs 13-19, 34

Diagnosis and notes Compared to O. sumawacus, this species has coarser pronotal and elytral punctures, the intervening microreticulate surface being slightly more reflective. Anterolateral surface of pronotum more or less asperate-punctate. Forebody black, not metallic. Elytra virtually glabrous, also black. Aedeagus, fig. 33, with characteristic distal paramerite. Contrary to the equally uniformly black group member

lows: [Indonesia], North Sumatra: [30 minutes walk up] Mt Bandahara [trail], 343N-9741S, 25/vi-5/ vii/1972, J Krikken, no. 23, ca 810 m, multistratal evergreen forest, from human excrements. Paratypes as listed in Annex 1. Description (holotype, male) Body length ca 11 mm. Head and pronotum cupreous, elytra dark-brown, underside and legs largely brown.

Holotype. Male, in RMNH, with label data as fol-

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29

28

30

31

Figs 26-31. Habitus and body elements of: 26-31, Onthophagus semiaureus (from Salak), 26, habitus, 27, head, 28, pronotum, 29, left elytron, 30, protibia, 31, metatibia.

Head without distinct non-marginal protrusions, wider than long. Clypeal margin anteriorly distinctly reflexed, apex with short tooth, its tip rounded. Clypeofrontal surface generally flat; clypeofrontal transition without distinct elevation (only smooth arcuate strip present). Clypeofrons cupreous, surface glabrous. Clypeofrons generally transversely rugulate to crowdedly asperate-punctate. Clypeal border from gena forward rounded; clypeogenal margin (at and near suture) widely rounded). Genae laterally subangulate. Frontovertex entirely devoid of protrusions; vertex a solid plate. Eyes (foramen in full-face view) small, more or less narrowly elliptic. Maximum number of facet rows (transversely) across eye foramen ca 12. Ratio transverse interocular distance/ maximum eye width ca 6. Pronotum generally evenly convex, disc and anterior declivity unmodified. Prothorax in dorsal outline with sides evenly rounded over at least anterior 0.7, slightly sinuate posteriorly. Anterior border of pronotum marginate. Anterolateral pronotal angle slightly obtuse, rounded off. Lateral sides of pronotum simply declivous, marginate. Posterolateral

section of pronotum strongly sinuate; pronotal posterolateral angles simple, distinct (very obtuse). Base of pronotum immarginate (except laterally), angulate medially (very distinctly so), surface slightly depressed. Pronotum setose, setae long (only present on short, narrow, oblique zone on either side of disc), surface with extremely fine, almost indistinct micropunctation. Pronotum generally abundantly to densely punctate, sparse on disc (punctures fine on disc, laterally larger, anterolaterally somewhat asperate, microstubbles scarcely distinct). Estimated number of punctures per 0.25 sq mm on pronotal disc 25-30, their diameters ca 0.01, mostly separated by 5-10 diameters. Pronotum entirely cupreous. Elytra dark-brown, matt, abundantly to densely setose, setae pale-yellow-brown, length 3-5 punctural diameters. Humeral umbone weakly pronounced. Punctures of (discal) elytral striae slightly impressed (roundish, fine, crenulating interstriae, separated by 3-5 diameters); striae distinct, shallowly impressed, narrow, sharply defined. Elytral interstriae all distinctly setose, setae distinct, long, subappressed; surface of interstriae feebly convex, simply punctate

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and microreticulate; interstrial punctures separated by 5-8 diameters, not arranged in rows, more or less scattered, sparse, lateral interstriae abundantly punctate-setose. Antennal lamellae brown. Antennal scapus smooth. Propectus laterally brown-cupreous, densely ocellatepunctate-setose. Metasternum brown; metasternal anterior lobe granulate, protruding, strongly prowshaped; metasternal disc finely, sparsely to abundantly punctate, glabrous. Metasternum laterally ocellate-punctate-setose, dense. Setae on propectoral sides very long, abundant to dense. Abdominal venter brown, matt, microreticulate, coarsely ocellatepunctate-setose, on proximal sternites along base only. Pygidium slightly convex, marginate along base, cupreous, moderately shiny, densely punctatesetose. Profemur brown, shiny, abundantly punctate, partly with long setae, broad, with distinct ridge on anterior side. Protibia robust, brown, shiny, externally with 4 denticles (1 short, 2-4 long, acuminate, 4 slender, all separated by slight serration). Proximal serration on outer side of protibia distinct. Protibia with apex oblique, apico-internal angle obtuse, with internal side evenly, slightly curved. Protibial spur inserted near inner angle, elongate-acuminate (robust, largely parallel-sided). Meso- and metafemora broad, complanate, brown, shiny. Mesofemur abundantly punctate, partly with long setae. Metafemur abundantly punctate-setose, setae fine, subappressed. Meso- and metatibiae distinctly dilated distad, with fossorial protrusions on outer side; fossorial protrusions transverse. Mesotibial spurs simply acuminate (very robust, long). Metatibial apex (virtually) straight (edge packed with spines). Metatibial spur simply acuminate (very robust, long). Meso- and metatibiae dark-brown, shiny. Metatarsal segment 1 robust (very slightly curved, slightly longer than tarsal segments 2-5 combined); approximate length proportions of terminal spur versus tarsal segments: 1-5: 29//30/9/6/3/7. Measurements in mm. Approximate maximum width of head 3.3; median length of pronotum (dorsal view) 4.0, maximum width 5.8; sutural length of elytra (dorsal view) 4.5, maximum width of elytra combined 6.1. Parameres, figs 34; distal paramerite not multi-angulate or -denticulate, mainly sitting on apical side. Diagnosis Closely related to aurifex. Distal paramerite not multi-angulate, not projecting, its orientation different from O. aurifex (i.e. laterally only narrowly distinct), other elements also different (compare figs 34-35). Forebody shiny, metallic (usually cupreous

or green). Elytra dark-brown (to black), distinctly, abundantly setose. Length 10-12 mm. Distribution and ecology North Sumatra, in lowland forest (350-810 m altitude), almost without exception in human faeces traps. Derivation of species name Composition of the names of the related species, semiaureus and aurifex.

Onthophagus (Parascatonomus) aurifex Harold, 1877

Figs 20-25, 35

Diagnosis and notes Closely related to semifex. Distal paramerite usually not multi-angulate, having a different orientation compared to semifex (i.e. laterally distinct); other parameral elements also different (compare figs. 34-35). Forebody shiny, metallic (usually cupreous or green). Elytra dark brown to black, matt, evenly abundantly to densely setose, also on disc. Length 9-12 mm. There is variation in the shape of the parameres, as illustrated by males from the Upper Kapuas region in West Kalimantan (fig. 35a) and from the Mulu region in Sarawak (fig. 35b). A male and a female from South Sumatra are tentatively allocated to aurifex: the paramerite of the male is different, being finely angulate, as in semiaureus, but the pair has the characteristic elytral pilosity of aurifex; we postpone a decision about their status (different taxon?), awaiting further material. O. semiaureus agreeing with material from West Java were found in the same region. The aurifex specimen recorded from Soepajang in Central Sumatra by Harold 1887 is a female, has a glabrous elytral disc, and is here allocated to O. semiaureus (q.v.). No confirmed records from Sabah. Measurements in mm of pictured male (Mulu). Approximate maximum width of head 3.1; median length of pronotum (dorsal view) 3.9, maximum width 5.4; sutural length of elytra (dorsal view) 4.0, maximum width of elytra combined 5.7; total length ca 10.5. Distribution and ecology Parts of southern Borneo (type locality Sarawak), and possibly South Sumatra, in kerangas [heath forest], and in other types of (preferably open?) forest. Extra-Bornean records all require verification.

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dpa

blp

32a

dpa

34

35a

dpa blp 34a 36 35b

dpa 36a dpa

37

38

Figs 32-36. Aedeagus, right side view; some species: projecting right distal paramerite (oblique from above) and tip (full-face); dpa = distal paramerite, blp = basolateral plate. 32, sumawacus (holotype), with 32a, distal paramerite; 33, Onthophagus sarawacus (from Danum); 34, semifex (holotype), with 34a, aedeagal tip; 35, aurifex (35a, from Mulu, 35b, from Upper Kapuas); 36, semiaureus (from Salak), with 36a distal paramerite. Scale line with figs 3236 = 1 mm. Figs. 37-38. Outline of clypeal margin, male. 37, Onthophagus liewi (after Ochi & Kon 2005); 38, sarawacus. Scale lines = 1 mm.

Krikken & Huijbregts: Onthophagus aurifex

183

Onthophagus (Parascatonomus) semiaureus Lansberge, 1883

Figs 26-31, 36

Diagnosis and notes The virtual lack of pilosity on the elytral disc of semiaureus seems the most useful external character of this species, while fresh aurifex and semifex always have abundantly, evenly, distinctly setose elytra. Distal paramerite distinctly projecting laterally, finely, variably multi-pointed (figs 36). Forebody shiny, metallic (usually cupreous or green). Elytra darkbrown to black, matt. Length 9-12 mm. The shape of the head is misrepresented in Balthasar (1963), his key implying that the anteromedian protrusion of the clypeus is different from O. aurifex this is why semiaureus and aurifex end up in distant couplets. The Bornean populations of O. semiaureus may differ slightly from those of Java and nearby regions in details of the parameral elements (such as the short projections of the distal paramerite). Clearly, as the last word has not been said about O. semiaureus, the lectotype designation and type locality restriction given below may provide a useful reference point for future revisers. Measurements in mm of pictured male (Salak). Approximate maximum width of head 3.1; median length of pronotum (dorsal view) 3.8, maximum width 5.2; sutural length of elytra (dorsal view) 4.0, maximum width of elytra combined 5.5; total length ca 10. Lectotype designation Van Lansberge (1883) mentioned Java and Sumatra as type localities; we did not recognize material from Sumatra with labels in van Lansberges handwriting in Leiden this may be elsewhere, or lost, or is it perhaps the female recorded by Harold 1887 from Soepajang as aurifex (see above; after all, the Central Sumatra Expedition led by Veth took place from 1877-1879). The lectotype in the Leiden museum, a female, here designated, is labelled as follows: Muller,\ Java [printed], Semiaureus\ Lansb. [in van Lansberges hand], and bears our lectotype label. As the collector, Salomon Mller (1804-1864, name on printed labels always without the umlaut) worked for the Leiden museum on various volcanic mountains of West Java, the type-locality is here restricted to West Java, using our own (male) material from Mount Salak, close to Bogor, as a reference (specimens labelled accordingly). The aedeagus of a Salak male is illustrated in figs 36.

Distribution and ecology O. semiaureus seems widespread in or near lowland forest or woodland, i.e usually in or near open situations; the majority of the specimens was collected in fish traps. Confirmed from Java, Bali, Sumatra, and Borneo; possibly elsewhere in Southeast Asia (Masumoto 2002), but verification of all material from unconfirmed regions required (see question marks in species list, above).

Acknowledgements
Thanks are due to the following colleagues for submitting specimens: H. Risnen and I. Hanski (both Finland), A.J. Davis (UK), A.L. van Berge Henegouwen (The Netherlands). Our colleague J. van Tol (The Netherlands) kindly read a draft of this paper. Thanks are also due to the organizations granting research permission, including Sabah Parks, the Sabah Forestry Department, and the Indonesian Institute of Science (LIPI).

References
Balthasar, V., 1963. Monographie der Scarabaeidae und Aphodiidae der palaearktischen und orientalischen Region (Coleoptera Lamellicornia). 2. Prague, Tschechoslowakische Akademie der Wissenschaften: 1-627. Boucomont, A., 1914. Les Coprophages de lArchipel Malais (Colopt.). Annales de la Socit Entomologique de France 73: 238-350. Davis, A.J., J.D. Holloway, H. Huijbregts, J. Krikken, A.H. Kirk- Spriggs & S.L. Sutton, 2001. Dung beetles as indicators of change in the forests of northern Borneo. Journal of Applied Ecology 38: 593-616. Harold, E. von, 1877. Enumration des Lamellicornes Coprophages rapports de lArchipel Malais, de la Nouvelle Guine, et de lAustralie borale par M.M.J. Doria, O. Beccari et L.M. dAlbertis. Annali del Museo Civico di Storia Naturale di Genova 10: 38-109. Harold, E. von, 1887. Scarabaeidae (Coprini) [= Laparosticti]. In: Snelleman, J.F. & Ritsema, C. (eds), Bijdragen tot de kennis der fauna van Midden-Sumatra. 1. Leiden, Brill: 24-29. Kabakov, O.N., 1992. Taxonomic status of Parascatonomus (Onthophagini, Scarabaeidae), with the description of new species from Southeast Asia. In: Medvedev, L.N. (ed.), Systematization and ecology of insects of Vietnam. Moscow, Nauka: 196-209 [In Russian]. Kabakov, O.N., 2006. Lamellicorn beetles of the subfamily Scarabaeinae of the fauna of Russia and adjacent countries. Moscow, Tovarichestvo nauchnykh izdanii KMK: 1-374 [In Russian]. Kabakov, O.N. & A. Napolov, 1999. Fauna and ecology of Lamellicornia of subfamily Scarabaeinae (Coleoptera: Scarabaeidae) of Vietnam and some parts of adjacent

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countries: South China, Laos and Thailand. Latvijas Entomologs 37: 58-96. Kon, M., Sakai, S. & Ochi, T., 2000. A new species of the genus Onthophagus (Coleoptera: Scarabaeidae) from Borneo. Entomological Science 3: 367-371. Lansberge, J.W. van, 1883. Rvision des Onthophagus de larchipel Indo-Neerlandais, avec description des espces nouvelles. Notes from the Leyden Museum 5: 41-82. Masumoto, K., 2002. New record of Onthophagus semiaureus Lansberge from Thailand. Elytra 30: 172

Ochi, T. & Kon, M., 2005. Notes on the coprophagous scarab-beetles (Coleoptera: Scarabaeidae) from Southeast Asia (VIII). Six new species of Onthophagus (Parascatonomus) and a new subspecies of O. (P.) katoi from Borneo. Entomological Review of Japan 60: 83-98.

Received: 30 January 2008 Accepted: 16 May 2008

Annex 1. Compilation of material examined

Label data and status of specimens
The terms pictured and measured in the list refer to specimens (apart from the holotypes of the new species) pictured in this paper, their measurements being given above, under Species accounts. Species sequence as in main text.

Onthophagus sarawacus

Paratypes: Indonesia: Sumatra: Mt Leuser NP: Ketambe, 22-29.vi.1983, Risnen hr11a, 500 m, lowland evergr forest, human excr trap, 1 ex.; Mt Leuser NP: Ketambe, 29-4.vi-vii.1983, Risnen hr11b, 500 m, lowland evergr forest, human excr trap, 2 exx.; Mt Leuser NP: Ketambe, 4-10/vii/1983, Risnen hr10f, 350 m, alluvial forest, human excr trap, 2 exx.; Mt Leuser NP: Ketambe, 4-10/vii/1983, Risnen hr11c, 500 m, lowland evergr forest, human excr trap, 3 exx.; Mt Leuser NP: Ketambe, 19-24/ix/1983, Risnen hr10Ci, 350 m, alluvial forest, human excr trap, 1 ex.; Mt Leuser NP: Lawe Mamas, 10-17/ix/1983, Risnen hr40Cb, 1530-1552 m, lower montane forest, human excr trap, 4 exx.; Mt Leuser NP: Lawe Mamas, 11-16/ix/1983, Risnen hr29Cb, 430-450 m, selective logging, human excr trap, 1 ex.; Mt Leuser NP: Lawe Mamas, 11-16/ix/1983, Risnen hr30Bb, 580-600 m, lowland evergr forest, fish trap, 1 ex.; Mt Leuser NP: Lawe Mamas, 11-16/ix/1983, Risnen hr31Cb, 750-780 m, lowland evergr forest, human excr trap, 3 exx.; Mt Leuser NP: Lawe Mamas, 11-16/ix/1983, Risnen hr32Bb, 900-920 m, lowland evergr forest, fish trap, 1 ex.; Mt Leuser NP: Lawe Mamas, 11-16/ix/1983, Risnen hr34Ab, 1190-1210 m, lower montane forest, human excr trap, 1 ex.; Mt Leuser NP: Lawe Mamas, 18-21/ix/1983, Risnen hr42A, 1300 m, lower montane forest, human excr trap, 1 ex.; Mt Leuser NP: Lawe Mamas, 18-21/ix/1983, Risnen hr42B, 1300 m, lower montane forest, fish trap, 1 ex.; Mt Leuser NP: Mt Mamas, 20-29/vii/1983, Risnen hr16a, 550 m, lowland evergr forest, human excr trap, 4 exx.; Mt Leuser NP: Mt Mamas, 21-29/vii/1983, Risnen hr18a, 1150 m, lower montane forest, human excr trap, 1 ex. Subtotal 28 specimens in RMNH, 15 records.

Onthophagus sumawacus

Material examined: Malaysia: Sabah: Crocker Range: Keningau-Kimanis rd (km19), 19-23/xi/1987, Krikken & Rombaut sa46, 900 m, multistr evergr forest, human excr trap, 2 exx.; Crocker Range: Keningau-Tambunan rd (km34), 21-24/xi/1987, Krikken & Rombaut sa47, 900 m, evergr scrub (secondary), human excr trap, 1 ex.; Danum Valley: Fld Centre envs (E5), 21-24/x/1987, Krikken & Rombaut sa11b, 100-200 m, multistr evergr forest, fish trap, 3 exx.; Danum Valley: Fld Centre envs (E5), 24-28/x/1987, Krikken & Rombaut sa16, 100-200 m, multistr evergr forest, human excr trap, 1 ex.; Danum Valley: Fld Centre envs (W2-3), 20-24/x/1987, Krikken & Rombaut sa07b, 100-200 m, multistr evergr forest, fish trap, 3 exx.; Danum Valley: Fld Centre envs (W2-3), 24-28/x/1987, Krikken & Rombaut sa14, 100-200 m, multistr evergr forest, human excr trap, 1 ex.; Danum Valley: Fld Centre envs (W22), 15-18/vi/1991, Davis ad393, 100-250 m, multistr evergr forest, human excr trap, 1 ex.; Danum Valley: Fld Centre envs (W5), 20-24/x/1987, Krikken & Rombaut sa06b, 100-200 m, multistr evergr forest, fish trap, 27 exx.; Danum Valley: Fld Centre envs (W8), 20-24/x/1987, Krikken & Rombaut sa05b, 100-200 m, multistr evergr forest, fish trap, 16 exx. (pictured and measured); Kinabalu NP: Poring (Kipungit), 18-22/i/1986, Krikken pw79a, 700 m, multistr evergr forest, human excr trap, 1 ex.; Kinabalu NP: Poring (Kipungit), 18-22/i/1986, Krikken pw79b, 700 m, multistr evergr forest, fish trap, 2 exx.; Kinabalu NP: Poring (Rafflesia forest), 13-18/i/1986, Krikken pw77b, 570 m, multistr evergr forest, fish trap, 1 ex.; Kinabalu NP: Poring (Rafflesia forest), 19-23/i/1986, Krikken pw78b, 570 m, multistr evergr forest, fish trap, 1 ex.; Kinabalu NP: Poring (Rafflesia forest), 8-11/xi/1987, Krikken & Rombaut sa32b, 500-600 m, multistr evergr forest, fish trap, 4 exx.; Lahad Datu: Lahad Datu, 21-23/iii/1987, van Tol jt870323, 170 m, multistr evergr forest, human excr trap, 1 ex.; Sandakan: Sepilok FR (waterfall trail F482), 1-4/xi/1987, Krikken & Rombaut sa25b, 50 m, multistr evergr forest, fish trap, 1 ex.; Sandakan: Sepilok FR (waterfall trail F482), 1-4/xi/1987, Krikken & Rombaut sa26, 50 m, multistr evergr forest, human excr trap, 1 ex. SARAWAK: Mt Mulu, iii-v/1978, Hanski, 100 m, alluvial forest, fish trap, 4 exx.; Mt Mulu, iii-v/1978, Hanski, 500-800 m, mixed dipterocarp forest, fish trap, 3 exx. KALIMANTAN: Mt Kenepai, i/1894, Bttikofer, 1 ex. Subtotal 75 specimens in RMNH, 20 records.

Krikken & Huijbregts: Onthophagus aurifex

185

Onthophagus semifex

Paratypes: Indonesia: Sumatra: Mt Leuser NP: Ketambe, 30-4/v-vi/1983, Risnen hr2b, 350 m, alluvial forest, banana trap, 1 ex.; Mt Leuser NP: Ketambe, 8-13/vi/1983, Risnen hr10b, 350 m, alluvial forest, human excr trap, 1 ex.; Mt Leuser NP: Ketambe, 22-29/vi/1983, Risnen hr11a, 500 m, lowland evergr forest, human excr trap, 3 exx.; Mt Leuser NP: Ketambe, 29-4/vi-vii/1983, Risnen hr11b, 500 m, lowland evergr forest, human excr trap, 4 exx.; Mt Leuser NP: Ketambe, 4-10/vii/1983, Risnen hr11c, 500 m, lowland evergr forest, human excr trap, 1 ex.; Mt Leuser NP: Lawe Mamas, 11-16/ix/1983, Risnen hr29Cb, 430-450 m, selective logging, human excr trap, 4 exx.; Mt Leuser NP: Lawe Mamas, 11-16/ix/1983, Risnen hr30Cb, 580-600 m, lowland evergr forest, human excr trap, 2 exx.; Mt Leuser NP: Lawe Mamas, 11-16/ix/1983, Risnen hr30Bb, 580-600 m, lowland evergr forest, fish trap, 2 exx.; Mt Leuser NP: Mt Mamas, 20-29/vii/1983, Risnen hr16a, 550 m, lowland evergr forest, human excr trap, 2 exx. Subtotal 21 specimens in RMNH, 10 records.

Material examined: Malaysia: Sarawak: Mt Mulu, iii-v/1978, Hanski, 150 m, kerangas forest, fish trap, 4 exx.; Mt Mulu, iii-v/1978, Hanski, 500-800 m, mixed dipterocarp forest, 1 ex.; Mt Mulu, iii-v/1978, Hanski, 100 m, alluvial forest, fish trap, 1 ex. (pictured and measured). Indonesia: Kalimantan: Upper Kapuas, Brug, 1 ex. (pictured). Sumatra: Way Kambas NP, 4/v/1986, van Berge Henegouwen & Pariwono bh163, sealevel, multistr evergr forest, fish trap, 2 exx. (mentioned in text above). Subtotal 9 specimens, 5 records, in RMNH.

Onthophagus aurifex

Material examined: Malaysia: Sabah: Danum Valley, 13-16/ix/1990, Davis ad174, 100-250 m, forest logged in 1989, human excr aerial trap, 1 ex.; Danum Valley: Bole River, 19-23/x/1987, Krikken & Rombaut sa03b, 100-200 m, second-growth forest logging rd, fish trap, 8 exx.; Danum Valley: Bole River, 19-23/x/1987, Krikken & Rombaut sa04b, 100-200 m, second-growth forest, fish trap, 15 exx.; Danum Valley: Fld Centre envs (E5), 21-24/x/1987, Krikken & Rombaut sa11b, 100-200 m, multistr evergr forest, fish trap, 5 exx.; Danum Valley: Fld Centre envs (W2-3), 20-24/x/1987, Krikken & Rombaut sa07b, 100-200 m, multistr evergr forest, fish trap, 7 exx.; Danum Valley: Fld Centre envs (W5), 20-24/x/1987, Krikken & Rombaut sa06b, 100-200 m,

Onthophagus semiaureus

multistr evergr forest, fish trap, 5 exx.; Sandakan: Sepilok FR (waterfall trail F482), 1-4/xi/1987, Krikken & Rombaut sa25a, 50 m, multistr evergr forest, human excr trap, 3 exx.; Sandakan: Sepilok FR (waterfall trail F482), 1-4/xi/1987, Krikken & Rombaut sa25b, 50 m, multistr evergr forest, fish trap, 7 exx.; Sandakan: Sepilok FR (waterfall trail F482), 1-4/xi/1987, Krikken & Rombaut sa26, 50 m, multistr evergr forest, human excr trap, 1 ex.; Sandakan: Sepilok FR (waterfall trail post 8), 1-4/xi/1987, Krikken & Rombaut sa24b, 50 m, multistr evergr forest, fish trap, 12 exx.; Sandakan: Sepilok FR Centre (resthouse), 1-4/xi/1987, Krikken & Rombaut sa27, 50 m, gardens, human excr trap, 1 ex. Sarawak: Mt Mulu, iii-v/1978, Hanski, 100 m, alluvial forest, fish trap, 2 exx. Indonesia: Kalimantan: Muarateweh, viii/1934, Sylvanus, 1 ex. Sumatra: Palembang, Knappert, 1 ex.; Supajang, iv/1877, 1 ex. (aurifex in Harold 1887). Java: loc. unspecified, Mller, 1 ex. lectotype; Alas Purwo NP: Trianggulasi, 24-26/ ii/1996, Huijbregts hh9614b, 50 m, monsoon forest, fish trap, 1 ex.; Ardja Sari, Kerkhoven, 1 ex.; Baluran NP: Bekol, 11-15/ii/1996, Huijbregts hh9607, 50 m, monsoon forest, human excr trap, 4 exx.; Banyuwangi, 1911, MacGillavry, 1 ex.; Kederi: Songbolong, Louwerens, 1 ex.; Mt Kawi, vii/1934, van Doesburg, 7 exx.; Mt Kawi, viii/1934, van Doesburg, 7 exx.; Mt Papandajan, Louwerens, 1 ex.; Mt Salak: N slope, 28-31/xii/1985, Krikken pw72a, 900 m, multistr evergr forest, human excr trap, 1 ex.; Mt Salak: N slope, 28-31/xii/1985, Krikken pw72b, 900 m, multistr evergr forest, fish trap, 1 ex. (pictured and measured); Nongkodjadjar, i/1911, Jacobson, 1 ex. Bali: Bedugul (warung crater rim), 24/iii/1986, van Berge Henegouwen & Pariwono bh136, 1300 m, cult area (crater rim), human excr trap, 1 ex.; Belimbing: Pura Mekori Forest, 27-31/ x/1991, Krikken & de Vries lk41a, 630 m, multistr evergr forest (remnant), human excr trap, 4 exx.; Belimbing: Pura Mekori Forest, 27-31/x/1991, Krikken & de Vries lk41b, 630 m, multistr evergr forest (remnant), fish trap, 22 exx.; Candikuning: Kebun Raya Bali, 28-1/x-xi/1991, Krikken, Huijbregts, de Vries lk45b, 1350 m, second-growth forest with bamboo, fish trap, 1 ex.; Mt Batukau: Pura Luhur Forest, 29-31/x/1991, Krikken & de Vries lk42a, 800 m, multistr evergr forest (nr river), human excr trap, 1 ex.; Mt Batukau: Pura Luhur Forest, 29-31/x/1991, Krikken & de Vries lk42b, 800 m, multistr evergr forest (nr river), fish trap, 8 exx.; Mt Batukau: Pura Luhur Forest, 29-31/x/1991, Krikken & de Vries lk43a, 800 m, multistr evergr forest, human excr trap, 1 ex.; Mt Batukau: Pura Luhur Forest, 29-31/x/1991, Krikken & de Vries lk43b, 800 m, multistr evergr forest, fish trap, 9 exx. Subtotal 144 specimens in RMNH, 35 records. Total 5 taxa, 277 specimens, in 85 records.