Distinguishing the Sundaland Species in the Onthophagus (Parascatonomus) Aurifex Group (Coleoptera: Scarabaeidae: Scarabaeinae)

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. [email protected]

Introduction usually man-made, monsoonal or soil-determined) In dung and carrion beetle collections from South- habitats. Consequently, full multi-island taxonomic east Asia, submitted for identification over the past overlap among these forest scarabs, is uncommon, three decades by ecological colleagues, we recognized and this seems to be the case here as well: the two several undescribed species of the near-cosmopolitan new North Sumatran forest species appear to have scarab genus Onthophagus Latreille, 1802, one of the very similar but taxonomically different relatives largest genera in the animal kingdom. In this paper on Borneo and elsewhere. In the aurifex group we two new species from North Sumatra (Aceh) are seem to be dealing with complex sets of geographic described, both related to O. aurifex Harold, 1877. and ecological vicariants, and it may well be that Boucomont (1914), in his classic review, made this unresolved character differences (for instance, vari- species the “type” of his group 6, the aurifex group, ation in pilosity and parameral structure) represent including three more species (all mentioned below). more than intraspecific variability. Published critical The Gunung Leuser National Park, where the new records and available collection series for the group, Onthophagus were found in 1972 and 1983, is an however, are limited; hence our inevitable question extensive area of lowland to subalpine ecosystems. marks behind the published regional records below At the time of the collecting activities the Park was (see the species list), indicating points of interest for covered with much pristine forest, home to our nov- future revisers. elties as well as conservation icons like orang-utan, The variation of the group member O. semiaureus elephant, rhino, tapir, and tiger – some now perhaps Lansberge, 1883 is another case in point, and it may locally (nearly) extinct. not be by chance that this multi-island, more variable Onthophagus are no different from many other species occurs mainly in secondary habitats. The Lei- groups of Southeast Asian animals and plants in that den museum has syntypical material of semiaureus, species occurring in primeval forest are, as a rule, less and since this paper is a contribution to disentan- widespread than are their close relatives in open (i.e., gling species identities in the aurifex group, the

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designation of a lectotype for this variable species is plane of the picture (like the head). appropriate. The qualification abundant for microsculptural units (like punctures) usually means: separated by Characteristic for the members of the Onthophagus 2-5 diameters, dense: 1-2 diameters, crowded: less aurifex group of species, as here understood, are than 1 diameter; sparse: means separated by at least the presence of a broad, short tooth on the middle 5 diameters, or limited to certain parts only. The of the clypeal margin, the lack of distinct protru- type of punctation is qualified only if it is decid- sions (like horns and ridges) on the head surface, edly different from a simple punctation. The prefix the punctate (non-granulate) pronotal disc, and the micro- usually refers to sculpture and pilosity distinct strongly angulate pronotal base (for full group diag- at magnifications x40 and higher. The term propec- nosis, see below). Note that in the present definition, tus denotes the underside of the prothorax. for purely operational reasons, we have excluded The morphology of the male genitalia is of foremost O. denticollis Lansberge, 1883, and any of its immedi- importance in species recognition; casual identifiers ate relatives, which lack the tooth on the clypeal mar- should not have the illusion to be able to recognize gin and are smaller. The species of the aurifex group species without critically looking at the aedeagi of have been placed in the subgenus Parascatonomus the males they handle. In particular, in the aurifex Paulian, 1932, for instance by Kon et al. (2000), group, note the more strongly sclerotized distal piece thus deviating from the limited subgeneric concept on each paramere, here termed paramerite; its shape of Balthasar (1963). Meanwhile, many more species and orientation are important characters. Note that were transferred to, or described as new in, various strict viewing angles apply (pointed parts may not other species groups allocated to Parascatonomus. appear as such when viewed from a different angle). Certain authors have gone further by raising Parasca- Ad hoc terminology is explained in the figures. tonomus to full genus (lately, Kabakov 2006, see also Always beware of deceptive intraspecific differences his earlier papers, like Kabakov 1992), but we can- between major males, minor males, and females – use not accept such a far-reaching action without sup- series including fresh major males whenever possible. port from a broad-scale approach to the supraspecific Males have foreshortened abdominal sternites. We classification of the hyperdiverse Onthophagini, with suspect polymorphism in the distribution and nature phylogenetic underpinning. Nevertheless, until this of the pilosity over the body. Also note that older, has happened, the Onthophagus aurifex group can, in worn specimens may have lost (part of) their pilosity. our view, safely be accommodated in an operation- The new material recorded in this paper, currently ally defined subgenus Parascatonomus (our definition kept in the National Museum of Natural History, is implicit in the group diagnosis below). Leiden, The Netherlands (RMNH), was collected by baited pitfall trapping (some aurifex group mem- Technical remarks bers, like other Parascatonomus, have a preference The primary purpose of this paper (part of an ongo- for carrion, but see ecological notes under species ing series on the taxonomy of laparostict scarabs of accounts and in Annex 1). In this paper we distin- the Southeast Asian islands) is the straightforward guish two altitudinal qualifications, lowland and description and diagnosis of the new taxa, and a upland, roughly under and above 1000 m (subalpine comparison with their closest relatives by means of altitudes, above 2000 m, were rarely sampled). an identification key and comparative pictures. Our comments on information from other sources The Sundaland region is here defined as the Malay are placed between square brackets (for instance in Peninsula + Sumatra + Java + Borneo + Bali + nearby the species list in the next section). Characters in the and intervening islands. following group diagnosis are arranged in their esti- Pictures are considered more useful than abstract mated order of taxonomic importance. descriptive text, and consequently, in addition to habitus pictures, at least six elements are depicted for each new species and its closest relative: head (full- The Onthophagus (Parascatonomus) face), pronotum and left elytron (dorsal), protibia aurifex group in Sundaland (upperside), metatibia (underside), and aedeagus Group diagnosis. Clypeus (in both sexes) with sin- (here mainly in lateral view). The photographs are gle anteromedian, reflexed, usually well-delimited not all to the same scale; refer to the species accounts broad tooth (its tip rounded or obtusely angu- for measurements of material pictured. The pilosity, lar), not standing in deep, abruptly delimited as depicted on the legs, is not diagnostic. The term emargination, clypeal edge widely, evenly rounded full-face view is applied where the upper surface of to lateral (subangular) tips of genae. Surface of head the element concerned is maximally parallel to the of male lacking higher protrusions, (at most) with

Downloaded from Brill.com09/25/2021 11:43:50PM via free access Krikken & Huijbregts: Onthophagus aurifex 175 obsolescent clypeofrontal and/or vertexal transverse Key ridge. Pronotal disc and anterior declivity simply 1. Dorsum entirely black, generally matt (pro- punctate (not granulate, rugulate, or asperate). Ely- notal disc and elytra microreticulate), on av- tral interstriae unmodified, simply punctate (pilos- erage larger species. Elytra virtually glabrous. ity in punctation varies, but never with long erect Interstrial punctures fine (distinctness of setae). Metasternum raised in front, prow-shaped, microstubbles dependent on illumination) . . 2 midline carinate. Pronotum without isolated protru- – Forebody shiny, metallic (usually green or sions, anterior declivity may be slightly depressed or cupreous). Elytra dark-brown or black, matt deplanate (in one species, liewi, topped by transverse, (microreticulate), on average smaller species. arcuate crest). Base of pronotum distinctly angulate Interstrial punctures (at least laterally) with medially, largely immarginate. Anterolateral angle of numerous long pale setae (very distinct at prothorax slightly angular, rounded off, not truncate x20) or with microstubbles, or virtually gla- or otherwise modified. Antennae relatively short; brous (at least on disc) ...... 4 scapus not serrate, indistinct from above; club com- 2. Clypeal median denticle on either side de- pacted, club segment 1 somewhat cup-shaped. Para- limited by sinuate emargination (fig. 37). meres relatively complex, apex blunt, with sclerotized Pronotum with (indication of) transverse ar- (darker brown), more or less projecting distal para- cuate ridge topping anterior declivity (large merite. Meso- and metafemora broad, complanate. males). Length 10.5-14.5 mm ...... liewi Meso- and metatibiae with more or less straight (not – Clypeal median denticle lacking sinuate strongly lobate) apex. Metatarsal segment 1 long, emargination on either side (fig. 38). Prono- robust, length approaching segments 2-5 combined. tum with more or less evenly declivous ante- Eyes in full-face view narrowly subelliptic. Protibia rior side ...... 3 with four outer denticles separated by serration; ter- 3. Pronotal and interstrial punctures on elytra minal spur dilated, not simply acuminate. Elytral extra fine. Parameres as in fig. 32., finely striae 6 and 7 of elytra subparallel, straight or evenly multiple-pointed distal paramerite standing curved near humerus (not sinuate). Body waisted. out laterally (fig. 32a). Length 10-14 mm . . Colour of body and legs entirely or partly (brown-) ...... sumawacus black, no light-dark pattern, dorsal side of forebody – Pronotal and interstrial punctures on elytra black(-brown), or conspicuously metallic (green, less fine. Parameres as in fig. 33, with coarse- blue, cupreous). Pygidial base marginate. Sexual ly multiple-pointed distal paramerite (lateral dimorphism slight (abdominal sternites in male view). Length 10-14 mm ...... sarawacus foreshortened). Medium-sized, body length 6-14.5 4. Elytral interstriae laterally at most with mm. (O. denticollis Lansberge, 1883, without clypeal microstubbles or microsetae, disc virtually tooth, omitted from the present group definition, glabrous. Pronotum glabrous. Parameres as the key, and the list.) in fig. 36. Distal paramerite variably, finely multiple-pointed, standing out laterally (fig. 36a). Length 8-12 mm ...... semiaureus Key to species – All elytral interstriae as well as pygidium Important characters evenly covered with distinct, rather long Shape of distal paramerite (denoted dpa in the fig- pale-white to pale-brown subappressed setae ures; orientation; distribution of angular points) and in distinct punctures. Parameres as in figs 34 other parameral elements (like the basolateral plate, & 35. Distal paramerite not multiple-point- denoted blp in the figures) ed, not standing out laterally (indistinct from • Colour and reflection of forebody (head and pro- above)...... 5 notum, simply black versus metallic (degree of 5. Parameres as in fig. 34, with distal paramerite reflection = surface microsculpture, see 3) shifted onto apex (fig. 34a), space between • Microsculpture (pronotum and elytral interstri- paramerite and basolateral plate not excised. ae, size and density of punctation, background Length 10-12 mm ...... semifex microreticulation) – Parameres as in fig. 35, with distal paramer- • Pilosity and micropilosity (particularly on elytra, ite distinct in lateral view, space between par- distribution of setae, setal length) amerite and basolateral plate excised. Length • Shape of clypeal margin (median tooth and any 9-12 mm ...... aurifex emargination on either side)

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1 7

5 11

2 8

3 9 6 12

4 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 24

14 21

15 22 18 25

16 23

19a 19a

dpa

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 declivity unmodified. Prothorax in dorsal outline Exclamation marks denote confirmations based on with sides evenly rounded over at least anterior material actually seen by us; question marks denote 0.7, slightly sinuate posteriorly. Anterior border of verification of material required. pronotum marginate. Anterolateral pronotal angle slightly obtuse, rounded off. Lateral sides of prono- Onthophagus tum simply declivous, marginate. Posterolateral section of pronotum strongly sinuate; pronotal postero- aurifex Harold, 1877 – Borneo (Sarawak!, Kaliman- lateral angles simple, distinct (but very obtuse). Base tan!), Malay Peninsula [?], Sumatra [?, including of pronotum immarginate, very distinctly angulate different taxon?], Java [?], Thailand [?, in Kaba- medially, surface slightly depressed. Pronotum gla- kov & Napolov 1999] brous, generally punctate (fine on disc, punctures liewi Ochi & Kon, 2005 – Borneo (Sabah) laterally and anterolaterally larger; microstubbles in sarawacus Harold, 1877 –Borneo (Sarawak!, Sabah!), them scarcely distinct), abundant to dense (sparser Sumatran West Coast Islands [? Batu Islands] on disc, very fine there). Estimated number of pri- semiaureus Lansberge, 1883 – Java!, Bali!, Borneo mary punctures per 0.25 sq mm on pronotal disc (Sarawak!, Sabah!, Kalimantan!), Sumatra [Cen- 30-35, diameters ca 0.01 mm, mostly separated by tral!, South!] plus West Coast islands [?], Sulawe- 5-10 diameters. Pronotum black, matt, microreticu- si [?], Philippines [?], Thailand [?] late. semifex sp.n. – North Sumatra Elytra black, matt, microreticulate, virtually gla- sumawacus sp.n. – North Sumatra brous, with hardly distinct microstubbles only. Sutural base slightly pressed apart, accommodat- ing pronotal angle. Humeral umbone weakly pro- Species accounts nounced. Punctures of (discal) elytral striae slightly Onthophagus (Parascatonomus) sumawacus impressed (roundish, fine, crenulating interstriae, Krikken & Huijbregts, sp. n. separated by 4-6 diameters); striae distinct, shallowly impressed, narrow, sharply defined. Discal interstriae Figs 1-6, 32 flat, simply, very finely punctate and microreticulate. Holotype. Male, in RMNH, with label data as fol- Interstrial punctures separated by 5-10 diameters, lows: [Indonesia], North Sumatra: Gng Leuser NP: not arranged in rows, more or less scattered, sparse. Lawe Mamas, 10-17/ix/1983, H Räisänen, 40Cb, Antennal lamellae light-brown. Antennal scapus 1530-1550 m, 3°20’N-97°44’S, tropical lower mon- smooth. tane rainforest, pitfall trap, human dung. Paratypes Propectus laterally black, abundantly punctate-setose as listed in Annex 1. to ocellate-punctate-setose (back side). Metaster- num black; anterior lobe granulate, protruding Description (holotype, male) (prow-shaped). Metasternal disc finely, sparsely to Body length ca 12.5 mm. Almost entirely black to abundantly punctate, with microstubbles. Metaster- black-brown, dorsum very matt, more or less seri- num laterally ocellate-punctate-setose, abundant ceous. to crowded. Setae on propectoral sides very long, Head without distinct non-marginal protrusions, abundant to dense. Abdominal venter black, matt, wider than long. Clypeal margin anteriorly distinctly microreticulate, coarsely ocellate-punctate-setose, on reflexed; apex with short tooth, its tip rounded, lat- proximal sternites along base only. Pygidium slightly erally simply sinuate. Clypeofrontal surface gener- convex, marginate along base, black, matt, microre- ally flat. Clypeofrontal transition without distinct ticulate, densely ocellate-punctate-setose, length of elevation (only smooth, slightly raised strip present). setae mostly not exceeding punctural diameter. Clypeofrons black, surface glabrous, generally (more Profemur black-brown, shiny, abundantly punctate, or less transversely) rugulate-punctate. Clypeal bor- partly with long setae, broad, with distinct ridge on der from gena forward rounded; clypeogenal mar- anterior side. Protibia robust, black-brown, shiny, gin (at and near suture) widely rounded. Genae externally with 4 denticles (1 short, 2-4 long, acu- laterally subangulate. Frontovertex entirely devoid minate, 4 slender, all separated by slight serration). of protrusions (basal transverse ridge obsolescent). Proximal serration on outer side of protibia distinct. Eyes (foramen in full-face view) small, more or less Protibia with apex oblique, apico-internal angle narrowly elliptic. Maximum number of facet rows obtuse, with internal side evenly, slightly curved. (transversely) across eye foramen ca 12. Ratio trans- Protibial spur inserted near inner angle, elongate- verse interocular distance/maximum eye width 6.5. acuminate (robust, largely parallel-sided). Meso- and Pronotum generally evenly convex, disc and anterior metafemora broad, complanate, black-brown, shiny,

Downloaded from Brill.com09/25/2021 11:43:50PM via free access Krikken & Huijbregts: Onthophagus aurifex 179 abundantly finely punctate, with microstubbles. mentioned hereafter, the clypeal tooth is poorly Meso- and metatibiae distinctly dilated distad, with delimited on either side, i.e. by a slightly sinuate fossorial protrusions on outer side; fossorial protru- curve (fig. 38). Length 10-14 mm. sions transverse. Mesotibial spurs simply acumi- Measurements in mm of pictured male (Danum). nate (very robust, long). Metatibial apex (virtually) Approximate maximum width of head 3.8; median straight (edge lined with spines). Metatibial spur length of pronotum (dorsal view) 5.0, maximum simply acuminate (very robust, long). Meso- and width 6.8; sutural length of elytra (dorsal view) 5.0, metatibiae black-brown, shiny. Metatarsal segment maximum width of elytra combined 7.1; total length 1 robust (slightly curved, slightly longer than tarsal ca 13. segments 2-5 combined); approximate length pro- portions of terminal spur versus tarsal segments 1-5: Distribution and ecology 29//31/9/6/4/7. Borneo (type locality Sarawak), apparently in low- Measurements in mm. Approximate maximum land forest, mostly in fish baited traps. According to width of head 3.7; median length of pronotum (dor- Davis et al. (2001) an interior forest specialist. Extra- sal view) 4.7, maximum width 6.5; sutural length of Bornean records to be confirmed (for the identity elytra (dorsal view) 5.0, maximum width of elytra of Vietnamese sarawacus-like species, see Kabakov combined 7.0. 1992). Parameres, figs 32, distal paramerite finely multi- angulate. Onthophagus (Paracatonomus) liewi Ochi &

Diagnosis Kon, 2005 Fig. 37 Compared to O. sarawacus, this new species has distinctly finer pronotal and elytral punctures, the intervening microreticulate surface being definitely less Diagnosis reflective. Anterolateral surface of pronotum simply This species is immediately recognizable from the punctate. Dorsum entirely matt, black, i.e. forebody shape of its clypeal apex, pronotal anterior impression not metallic. Elytra virtually glabrous, microstubbles and ridge (in large males), and aedeagus (see various hardly distinct. Aedeagus, figs 32, shape and orienta- figures in Ochi & Kon 2005). The clypeal tooth is tion of distal paramerite characteristic. Clypeal tooth on either side well defined by a sinuate emargination lacking emargination on either side, contrary to (fig. 37). Forebody black, not metallic. Elytra virtu- O. liewi (see below). Length 10-14 mm. ally glabrous, also black. Length 10.5-14.5 mm.

Distribution and ecology Distribution and ecology North Sumatra, both lowland and upland forest Sabah, in upland situations on and around Mt Kina- (350-1550 m altitude), mostly collected in human balu – no further ecological information with the faeces baited traps. original description. Perhaps an upland vicariant of O. sarawacus, and possibly limited in space and/or Derivation of species name time, as we did not encounter O. liewi in our Sabah Composition of Sumatra, home island of the species, samples. and the name of the related species sarawacus. Onthophagus (Parascatonomus) semifex Onthophagus (Parascatonomus) sarawacus Krikken & Huijbregts, sp. n. Harold, 1877 Figs 13-19, 34 Figs 7-12, 33, 38 Holotype. Male, in RMNH, with label data as follows: [Indonesia], North Sumatra: [30 minutes walk Diagnosis and notes up] Mt Bandahara [trail], 3°43’N-97°41’S, 25/vi-5/ Compared to O. sumawacus, this species has coarser vii/1972, J Krikken, no. 23, ca 810 m, multistratal pronotal and elytral punctures, the intervening evergreen forest, from human excrements. Paratypes microreticulate surface being slightly more reflec- as listed in Annex 1. tive. Anterolateral surface of pronotum more or less asperate-punctate. Forebody black, not metal- Description (holotype, male) lic. Elytra virtually glabrous, also black. Aedeagus, Body length ca 11 mm. Head and pronotum cupre- fig. 33, with characteristic distal paramerite. Con- ous, elytra dark-brown, underside and legs largely trary to the equally uniformly black group member brown.

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26 27 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, section of pronotum strongly sinuate; pronotal wider than long. Clypeal margin anteriorly distinctly posterolateral angles simple, distinct (very obtuse). reflexed, apex with short tooth, its tip rounded. Base of pronotum immarginate (except laterally), Clypeofrontal surface generally flat; clypeofrontal angulate medially (very distinctly so), surface slightly transition without distinct elevation (only smooth depressed. Pronotum setose, setae long (only present arcuate strip present). Clypeofrons cupreous, surface on short, narrow, oblique zone on either side of glabrous. Clypeofrons generally transversely rugu- disc), surface with extremely fine, almost indistinct late to crowdedly asperate-punctate. Clypeal border micropunctation. Pronotum generally abundantly from gena forward rounded; clypeogenal margin (at to densely punctate, sparse on disc (punctures fine and near suture) widely rounded). Genae laterally on disc, laterally larger, anterolaterally somewhat subangulate. Frontovertex entirely devoid of protru- asperate, microstubbles scarcely distinct). Estimated sions; vertex a solid plate. Eyes (foramen in full-face number of punctures per 0.25 sq mm on pronotal view) small, more or less narrowly elliptic. Maxi- disc 25-30, their diameters ca 0.01, mostly separated mum number of facet rows (transversely) across eye by 5-10 diameters. Pronotum entirely cupreous. foramen ca 12. Ratio transverse interocular distance/ Elytra dark-brown, matt, abundantly to densely maximum eye width ca 6. setose, setae pale-yellow-brown, length 3-5 punctural Pronotum generally evenly convex, disc and ante- diameters. Humeral umbone weakly pronounced. rior declivity unmodified. Prothorax in dorsal out- Punctures of (discal) elytral striae slightly impressed line with sides evenly rounded over at least anterior (roundish, fine, crenulating interstriae, separated by 0.7, slightly sinuate posteriorly. Anterior border of 3-5 diameters); striae distinct, shallowly impressed, pronotum marginate. Anterolateral pronotal angle narrow, sharply defined. Elytral interstriae all slightly obtuse, rounded off. Lateral sides of pro- distinctly setose, setae distinct, long, subappressed; notum simply declivous, marginate. Posterolateral surface of interstriae feebly convex, simply punctate

Downloaded from Brill.com09/25/2021 11:43:50PM via free access Krikken & Huijbregts: Onthophagus aurifex 181 and microreticulate; interstrial punctures separated or green). Elytra dark-brown (to black), distinctly, by 5-8 diameters, not arranged in rows, more or less abundantly setose. Length 10-12 mm. scattered, sparse, lateral interstriae abundantly punctate-setose. Distribution and ecology Antennal lamellae brown. Antennal scapus smooth. North Sumatra, in lowland forest (350-810 m alti- Propectus laterally brown-cupreous, densely ocellate- tude), almost without exception in human faeces punctate-setose. Metasternum brown; metasternal traps. anterior lobe granulate, protruding, strongly prow- shaped; metasternal disc finely, sparsely to abun- Derivation of species name dantly punctate, glabrous. Metasternum laterally Composition of the names of the related species, ocellate-punctate-setose, dense. Setae on propectoral semiaureus and aurifex. sides very long, abundant to dense. Abdominal venter brown, matt, microreticulate, coarsely ocellate- Onthophagus (Parascatonomus) aurifex punctate-setose, on proximal sternites along base Harold, 1877 only. Pygidium slightly convex, marginate along Figs 20-25, 35 base, cupreous, moderately shiny, densely punctate- setose. Profemur brown, shiny, abundantly punctate, partly Diagnosis and notes with long setae, broad, with distinct ridge on anterior Closely related to semifex. Distal paramerite usually side. Protibia robust, brown, shiny, externally with 4 not multi-angulate, having a different orientation denticles (1 short, 2-4 long, acuminate, 4 slender, compared to semifex (i.e. laterally distinct); other all separated by slight serration). Proximal serration parameral elements also different (compare figs. on outer side of protibia distinct. Protibia with apex 34-35). Forebody shiny, metallic (usually cupreous oblique, apico-internal angle obtuse, with internal or green). Elytra dark brown to black, matt, evenly side evenly, slightly curved. Protibial spur inserted abundantly to densely setose, also on disc. Length near inner angle, elongate-acuminate (robust, largely 9-12 mm. parallel-sided). Meso- and metafemora broad, com- There is variation in the shape of the parameres, as planate, brown, shiny. Mesofemur abundantly punc- illustrated by males from the Upper Kapuas region tate, partly with long setae. Metafemur abundantly in West Kalimantan (fig. 35a) and from the Mulu punctate-setose, setae fine, subappressed. Meso- and region in Sarawak (fig. 35b). metatibiae distinctly dilated distad, with fosso- A male and a female from South Sumatra are ten- rial protrusions on outer side; fossorial protrusions tatively allocated to aurifex: the paramerite of the transverse. Mesotibial spurs simply acuminate (very male is different, being finely angulate, as in semiau- robust, long). Metatibial apex (virtually) straight reus, but the pair has the characteristic elytral pilos- (edge packed with spines). Metatibial spur simply ity of aurifex; we postpone a decision about their acuminate (very robust, long). Meso- and metatib- status (different taxon?), awaiting further material. iae dark-brown, shiny. Metatarsal segment 1 robust O. semiaureus agreeing with material from West Java (very slightly curved, slightly longer than tarsal seg- were found in the same region. The aurifex specimen ments 2-5 combined); approximate length propor- recorded from “Soepajang” in Central Sumatra by tions of terminal spur versus tarsal segments: 1-5: Harold 1887 is a female, has a glabrous elytral disc, 29//30/9/6/3/7. and is here allocated to O. semiaureus (q.v.). No con- Measurements in mm. Approximate maximum firmed records from Sabah. width of head 3.3; median length of pronotum (dor- Measurements in mm of pictured male (Mulu). sal view) 4.0, maximum width 5.8; sutural length of Approximate maximum width of head 3.1; median elytra (dorsal view) 4.5, maximum width of elytra length of pronotum (dorsal view) 3.9, maximum combined 6.1. width 5.4; sutural length of elytra (dorsal view) 4.0, Parameres, figs 34; distal paramerite not multi-angu- maximum width of elytra combined 5.7; total length late or -denticulate, mainly sitting on apical side. ca 10.5.

Diagnosis Distribution and ecology Closely related to aurifex. Distal paramerite not Parts of southern Borneo (type locality Sarawak), multi-angulate, not projecting, its orientation dif- and possibly South Sumatra, in kerangas [heath for- ferent from O. aurifex (i.e. laterally only narrowly est], and in other types of (preferably open?) forest. distinct), other elements also different (compare figs Extra-Bornean records all require verification. 34-35). Forebody shiny, metallic (usually cupreous

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32 33

dpa blp

32a dpa

35a 34

dpa

blp

34a

36 35b

dpa dpa 36a

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 32- 36 = 1 mm. Figs. 37-38. Outline of clypeal margin, male. 37, Onthophagus liewi (after Ochi & Kon 2005); 38, sarawacus. Scale lines = 1 mm.

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Onthophagus (Parascatonomus) semiaureus Distribution and ecology Lansberge, 1883 O. semiaureus seems widespread in or near lowland Figs 26-31, 36 forest or woodland, i.e usually in or near open situations; the majority of the specimens was collected in Diagnosis and notes fish traps. Confirmed from Java, Bali, Sumatra, and The virtual lack of pilosity on the elytral disc of Borneo; possibly elsewhere in Southeast Asia (Mas- semiaureus seems the most useful external character umoto 2002), but verification of all material from of this species, while fresh aurifex and semifex always unconfirmed regions required (see question marks in have abundantly, evenly, distinctly setose elytra. Dis- species list, above). tal paramerite distinctly projecting laterally, finely, variably multi-pointed (figs 36). Forebody shiny, metallic (usually cupreous or green). Elytra dark- Acknowledgements brown to black, matt. Length 9-12 mm. Thanks are due to the following colleagues for The shape of the head is misrepresented in Balthasar submitting specimens: H. Räisänen and I. Hanski (1963), his key implying that the anteromedian pro- (both Finland), A.J. Davis (UK), A.L. van Berge trusion of the clypeus is different from O. aurifex – Henegouwen (The Netherlands). Our colleague this is why semiaureus and aurifex end up in distant J. van Tol (The Netherlands) kindly read a draft of couplets. this paper. Thanks are also due to the organizations The Bornean populations of O. semiaureus may dif- granting research permission, including Sabah Parks, fer slightly from those of Java and nearby regions in the Sabah Forestry Department, and the Indonesian details of the parameral elements (such as the short Institute of Science (LIPI). projections of the distal paramerite). Clearly, as the last word has not been said about O. semiaureus, the lectotype designation and type locality restriction References given below may provide a useful reference point for Balthasar, V., 1963. Monographie der Scarabaeidae und future revisers. Aphodiidae der palaearktischen und orientalischen Measurements in mm of pictured male (Salak). Region (Coleoptera Lamellicornia). 2. – Prague, Approximate maximum width of head 3.1; median Tschechoslowakische Akademie der Wissenschaften: length of pronotum (dorsal view) 3.8, maximum 1-627. width 5.2; sutural length of elytra (dorsal view) 4.0, Boucomont, A., 1914. Les Coprophages de l’Archipel Ma- lais (Coléopt.). – Annales de la Société Entomologique maximum width of elytra combined 5.5; total length de France 73: 238-350. ca 10. Davis, A.J., J.D. Holloway, H. Huijbregts, J. Krikken, A.H. Kirk- Spriggs & S.L. Sutton, 2001. Dung beetles Lectotype designation as indicators of change in the forests of northern Bor- Van Lansberge (1883) mentioned Java and Sumatra neo. – Journal of Applied Ecology 38: 593-616. as type localities; we did not recognize material from Harold, E. von, 1877. Enumération des Lamellicornes Co- Sumatra with labels in van Lansberge’s handwriting prophages rapportés de l’Archipel Malais, de la Nou- in Leiden – this may be elsewhere, or lost, or is it velle Guinée, et de l’Australie boréale par M.M.J. Do- perhaps the female recorded by Harold 1887 from ria, O. Beccari et L.M. d’Albertis. – Annali del Museo “Soepajang” as aurifex (see above; after all, the Cen- Civico di Storia Naturale di Genova 10: 38-109. tral Sumatra Expedition led by Veth took place from Harold, E. von, 1887. Scarabaeidae (Coprini) [= Laparos- 1877-1879). The lectotype in the Leiden museum, ticti]. – In: Snelleman, J.F. & Ritsema, C. (eds), Bij- a female, here designated, is labelled as follows: dragen tot de kennis der fauna van Midden-Sumatra. “Muller,\ Java” [printed], “Semiaureus\ Lansb.” [in 1. – Leiden, Brill: 24-29. Kabakov, O.N., 1992. Taxonomic status of Parascatonomus van Lansberge’s hand], and bears our lectotype label. (Onthophagini, Scarabaeidae), with the description of As the collector, Salomon Müller (1804-1864, name new species from Southeast Asia. – In: Medvedev, L.N. on printed labels always without the umlaut) worked (ed.), Systematization and ecology of insects of Viet- for the Leiden museum on various volcanic moun- nam. – Moscow, Nauka: 196-209 [In Russian]. tains of West Java, the type-locality is here restricted Kabakov, O.N., 2006. Lamellicorn beetles of the subfamily to West Java, using our own (male) material from Scarabaeinae of the fauna of Russia and adjacent coun- Mount Salak, close to Bogor, as a reference (speci- tries. – Moscow, Tovarichestvo nauchnykh izdanii mens labelled accordingly). The aedeagus of a Salak KMK: 1-374 [In Russian]. male is illustrated in figs 36. 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 Ochi, T. & Kon, M., 2005. Notes on the coprophagous Entomologs 37: 58-96. scarab-beetles (Coleoptera: Scarabaeidae) from South- Kon, M., Sakai, S. & Ochi, T., 2000. A new species of the east Asia (VIII). Six new species of Onthophagus (Paras- genus Onthophagus (Coleoptera: Scarabaeidae) from catonomus) and a new subspecies of O. (P.) katoi from Borneo. – Entomological Science 3: 367-371. Borneo. – Entomological Review of Japan 60: 83-98. Lansberge, J.W. van, 1883. Révision des Onthophagus de l’archipel Indo-Neerlandais, avec description des es- pèces nouvelles. – Notes from the Leyden Museum 5: 41-82. Masumoto, K., 2002. New record of Onthophagus semiau- Received: 30 January 2008 reus Lansberge from Thailand. – Elytra 30: 172 Accepted: 16 May 2008

Annex 1. Compilation of material Onthophagus sarawacus examined Material examined: Malaysia: Sabah: Crocker Range: Keningau-Kimanis rd (km19), 19-23/xi/1987, Krikken Label data and status of specimens & Rombaut sa46, 900 m, multistr evergr forest, human The terms pictured and measured in the list refer to speci- excr trap, 2 exx.; Crocker Range: Keningau-Tambunan mens (apart from the holotypes of the new species) pic- rd (km34), 21-24/xi/1987, Krikken & Rombaut sa47, tured in this paper, their measurements being given above, 900 m, evergr scrub (secondary), human excr trap, under Species accounts. Species sequence as in main text. 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 Onthophagus sumawacus envs (E5), 24-28/x/1987, Krikken & Rombaut sa16, Paratypes: Indonesia: Sumatra: Mt Leuser NP: Ketambe, 100-200 m, multistr evergr forest, human excr trap, 1 ex.; 22-29.vi.1983, Räisänen hr11a, 500 m, lowland evergr Danum Valley: Fld Centre envs (W2-3), 20-24/x/1987, forest, human excr trap, 1 ex.; Mt Leuser NP: Ketambe, Krikken & Rombaut sa07b, 100-200 m, multistr ev- 29-4.vi-vii.1983, Räisänen hr11b, 500 m, lowland evergr forest, fish trap, 3 exx.; Danum Valley: Fld Centre ergr forest, human excr trap, 2 exx.; Mt Leuser NP: Keta- envs (W2-3), 24-28/x/1987, Krikken & Rombaut sa14, mbe, 4-10/vii/1983, Räisänen hr10f, 350 m, alluvial for- 100-200 m, multistr evergr forest, human excr trap, 1 ex.; est, human excr trap, 2 exx.; Mt Leuser NP: Ketambe, Danum Valley: Fld Centre envs (W22), 15-18/vi/1991, 4-10/vii/1983, Räisänen hr11c, 500 m, lowland evergr Davis ad393, 100-250 m, multistr evergr forest, human forest, human excr trap, 3 exx.; Mt Leuser NP: Ketambe, excr trap, 1 ex.; Danum Valley: Fld Centre envs (W5), 19-24/ix/1983, Räisänen hr10Ci, 350 m, alluvial for- 20-24/x/1987, Krikken & Rombaut sa06b, 100-200 m, est, human excr trap, 1 ex.; Mt Leuser NP: Lawe Mamas, multistr evergr forest, fish trap, 27 exx.; Danum Valley: Fld 10-17/ix/1983, Räisänen hr40Cb, 1530-1552 m, low- Centre envs (W8), 20-24/x/1987, Krikken & Rombaut er montane forest, human excr trap, 4 exx.; Mt Leuser sa05b, 100-200 m, multistr evergr forest, fish trap, 16 exx. NP: Lawe Mamas, 11-16/ix/1983, Räisänen hr29Cb, (pictured and measured); Kinabalu NP: Poring (Kipungit), 430-450 m, selective logging, human excr trap, 1 ex.; 18-22/i/1986, Krikken pw79a, 700 m, multistr evergr for- Mt Leuser NP: Lawe Mamas, 11-16/ix/1983, Räisänen est, human excr trap, 1 ex.; Kinabalu NP: Poring (Kipun- hr30Bb, 580-600 m, lowland evergr forest, fish trap, 1 ex.; git), 18-22/i/1986, Krikken pw79b, 700 m, multistr ev- Mt Leuser NP: Lawe Mamas, 11-16/ix/1983, Räisänen ergr forest, fish trap, 2 exx.; Kinabalu NP: Poring (Rafflesia hr31Cb, 750-780 m, lowland evergr forest, human excr forest), 13-18/i/1986, Krikken pw77b, 570 m, multistr ev- trap, 3 exx.; Mt Leuser NP: Lawe Mamas, 11-16/ix/1983, ergr forest, fish trap, 1 ex.; Kinabalu NP: Poring (Rafflesia Räisänen hr32Bb, 900-920 m, lowland evergr forest, fish forest), 19-23/i/1986, Krikken pw78b, 570 m, multistr trap, 1 ex.; Mt Leuser NP: Lawe Mamas, 11-16/ix/1983, evergr forest, fish trap, 1 ex.; Kinabalu NP: Poring (Raf- Räisänen hr34Ab, 1190-1210 m, lower montane forest, flesia forest), 8-11/xi/1987, Krikken & Rombaut sa32b, human excr trap, 1 ex.; Mt Leuser NP: Lawe Mamas, 500-600 m, multistr evergr forest, fish trap, 4 exx.; Lahad 18-21/ix/1983, Räisänen hr42A, 1300 m, lower mon- Datu: Lahad Datu, 21-23/iii/1987, van Tol jt870323, tane forest, human excr trap, 1 ex.; Mt Leuser NP: Lawe 170 m, multistr evergr forest, human excr trap, 1 ex.; Mamas, 18-21/ix/1983, Räisänen hr42B, 1300 m, lower Sandakan: Sepilok FR (waterfall trail F482), 1-4/xi/1987, montane forest, fish trap, 1 ex.; Mt Leuser NP: Mt Mamas, Krikken & Rombaut sa25b, 50 m, multistr evergr forest, 20-29/vii/1983, Räisänen hr16a, 550 m, lowland evergr fish trap, 1 ex.; Sandakan: Sepilok FR (waterfall trail F482), forest, human excr trap, 4 exx.; Mt Leuser NP: Mt Mamas, 1-4/xi/1987, Krikken & Rombaut sa26, 50 m, multistr ev- 21-29/vii/1983, Räisänen hr18a, 1150 m, lower montane ergr forest, human excr trap, 1 ex. SARAWAK: Mt Mulu, forest, human excr trap, 1 ex. iii-v/1978, Hanski, 100 m, alluvial forest, fish trap, 4 exx.; Mt Mulu, iii-v/1978, Hanski, 500-800 m, mixed diptero- Subtotal 28 specimens in RMNH, 15 records. carp forest, fish trap, 3 exx. KALIMANTAN: Mt Kenepai, i/1894, Büttikofer, 1 ex.

Subtotal 75 specimens in RMNH, 20 records.

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Onthophagus semifex multistr evergr forest, fish trap, 5 exx.; Sandakan: Se- Paratypes: Indonesia: Sumatra: Mt Leuser NP: Ketambe, pilok FR (waterfall trail F482), 1-4/xi/1987, Krikken & 30-4/v-vi/1983, Räisänen hr2b, 350 m, alluvial forest, ba- Rombaut sa25a, 50 m, multistr evergr forest, human excr nana trap, 1 ex.; Mt Leuser NP: Ketambe, 8-13/vi/1983, trap, 3 exx.; Sandakan: Sepilok FR (waterfall trail F482), Räisänen hr10b, 350 m, alluvial forest, human excr trap, 1-4/xi/1987, Krikken & Rombaut sa25b, 50 m, multistr 1 ex.; Mt Leuser NP: Ketambe, 22-29/vi/1983, Räisänen evergr forest, fish trap, 7 exx.; Sandakan: Sepilok FR (wa- hr11a, 500 m, lowland evergr forest, human excr trap, terfall trail F482), 1-4/xi/1987, Krikken & Rombaut sa26, 3 exx.; Mt Leuser NP: Ketambe, 29-4/vi-vii/1983, Räisänen 50 m, multistr evergr forest, human excr trap, 1 ex.; hr11b, 500 m, lowland evergr forest, human excr trap, Sandakan: Sepilok FR (waterfall trail post 8), 1-4/xi/1987, 4 exx.; Mt Leuser NP: Ketambe, 4-10/vii/1983, Räisänen Krikken & Rombaut sa24b, 50 m, multistr evergr forest, hr11c, 500 m, lowland evergr forest, human excr trap, fish trap, 12 exx.; Sandakan: Sepilok FR Centre (resthouse), 1 ex.; Mt Leuser NP: Lawe Mamas, 11-16/ix/1983, 1-4/xi/1987, Krikken & Rombaut sa27, 50 m, gardens, Räisänen hr29Cb, 430-450 m, selective logging, hu- human excr trap, 1 ex. Sarawak: Mt Mulu, iii-v/1978, man excr trap, 4 exx.; Mt Leuser NP: Lawe Mamas, Hanski, 100 m, alluvial forest, fish trap, 2 exx. Indonesia: 11-16/ix/1983, Räisänen hr30Cb, 580-600 m, lowland ev- Kalimantan: Muarateweh, viii/1934, Sylvanus, 1 ex. Su- ergr forest, human excr trap, 2 exx.; Mt Leuser NP: Lawe matra: Palembang, Knappert, 1 ex.; Supajang, iv/1877, 1 Mamas, 11-16/ix/1983, Räisänen hr30Bb, 580-600 m, ex. (aurifex in Harold 1887). Java: loc. unspecified, Müller, lowland evergr forest, fish trap, 2 exx.; Mt Leuser NP: Mt 1 ex. lectotype; Alas Purwo NP: Trianggulasi, 24-26/ Mamas, 20-29/vii/1983, Räisänen hr16a, 550 m, lowland ii/1996, Huijbregts hh9614b, 50 m, monsoon forest, fish evergr forest, human excr trap, 2 exx. trap, 1 ex.; Ardja Sari, Kerkhoven, 1 ex.; Baluran NP: Bekol, 11-15/ii/1996, Huijbregts hh9607, 50 m, monsoon forest, Subtotal 21 specimens in RMNH, 10 records. 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 Onthophagus aurifex Doesburg, 7 exx.; Mt Papandajan, Louwerens, 1 ex.; Mt Material examined: Malaysia: Sarawak: Mt Mulu, Salak: N slope, 28-31/xii/1985, Krikken pw72a, 900 m, iii-v/1978, Hanski, 150 m, kerangas forest, fish trap, multistr evergr forest, human excr trap, 1 ex.; Mt Salak: 4 exx.; Mt Mulu, iii-v/1978, Hanski, 500-800 m, mixed N slope, 28-31/xii/1985, Krikken pw72b, 900 m, mul- dipterocarp forest, 1 ex.; Mt Mulu, iii-v/1978, Hanski, tistr evergr forest, fish trap, 1 ex. (pictured and measured); 100 m, alluvial forest, fish trap, 1 ex. (pictured and meas- Nongkodjadjar, i/1911, Jacobson, 1 ex. Bali: Bedugul (wa- ured). Indonesia: Kalimantan: Upper Kapuas, Brug, rung crater rim), 24/iii/1986, van Berge Henegouwen & 1 ex. (pictured). Sumatra: Way Kambas NP, 4/v/1986, van Pariwono bh136, 1300 m, cult area (crater rim), human Berge Henegouwen & Pariwono bh163, sealevel, multistr excr trap, 1 ex.; Belimbing: Pura Mekori Forest, 27-31/ evergr forest, fish trap, 2 exx. (mentioned in text above). 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, Subtotal 9 specimens, 5 records, in RMNH. 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 Onthophagus semiaureus with bamboo, fish trap, 1 ex.; Mt Batukau: Pura Luhur Material examined: Malaysia: Sabah: Danum Valley, Forest, 29-31/x/1991, Krikken & de Vries lk42a, 800 m, 13-16/ix/1990, Davis ad174, 100-250 m, forest logged in multistr evergr forest (nr river), human excr trap, 1 ex.; 1989, human excr aerial trap, 1 ex.; Danum Valley: Bole Riv- Mt Batukau: Pura Luhur Forest, 29-31/x/1991, Krik- er, 19-23/x/1987, Krikken & Rombaut sa03b, 100-200 m, ken & de Vries lk42b, 800 m, multistr evergr forest (nr second-growth forest logging rd, fish trap, 8 exx.; Da- river), fish trap, 8 exx.; Mt Batukau: Pura Luhur Forest, num Valley: Bole River, 19-23/x/1987, Krikken & Rom- 29-31/x/1991, Krikken & de Vries lk43a, 800 m, multistr baut sa04b, 100-200 m, second-growth forest, fish trap, evergr forest, human excr trap, 1 ex.; Mt Batukau: Pura 15 exx.; Danum Valley: Fld Centre envs (E5), Luhur Forest, 29-31/x/1991, Krikken & de Vries lk43b, 21-24/x/1987, Krikken & Rombaut sa11b, 100-200 800 m, multistr evergr forest, fish trap, 9 exx. m, multistr evergr forest, fish trap, 5 exx.; Danum Val- Subtotal 144 specimens in RMNH, 35 records. ley: 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), Total 5 taxa, 277 specimens, in 85 records. 20-24/x/1987, Krikken & Rombaut sa06b, 100-200 m,

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