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Taxonomic diversity of the genus Ochicanthon in Sundaland (Coleoptera: Scarabaeidae: Scarabaeinae)

J. Krikken & J. Huijbregts

The Sundaland representatives of the Oriental genus Ochicanthon Vaz-de-Mello, 2003 (= Phacosoma Boucomont, 1914) are reviewed, on the basis of a study of 1100 specimens and the literature. The genus is rediagnosed, its species are listed, and the Sundaland species are keyed and diagnosed; 13 new species are described from Sabah & Sarawak (11 new species), the Malay Peninsula (1), and Java (1), bringing the total to 36. Taxonomic, biogeographic, and ecological aspects are discussed, and research questions summarized. Ochicanthon species are associated with dung and carrion. They are hypothesized to be essentially Indo-Gondwanan, having dispersed from India to the East, but not into Wallacea and beyond. At least one species is flightless. The following species from Malaysia and Indonesia are new: Ochicanthon edmondsi (Borneo), rombauti (Borneo), crockermontis (Borneo), crypticus (Borneo), peninsularis (Malay Peninsula), neglectus (Borneo), dulitmontis (Borneo), tambunan (Borneo), mulu (Borneo), kimanis (Borneo), danum (Borneo), javanus (Java), hanskii (Borneo). New records of described species are given. The type locality of O. dytiscoides (Boucomont, 1914) is restricted to Sabah by designating a lectotype for this species. Suspected synonyms are briefly discussed, and attention is drawn to the poorly resolved complex of species around O. dytiscoides. A note on Scarabaeinae erroneously recorded from Sulawesi is added, the record of the unidentifiable Ochicanthon punctatus (Boucomont, 1914) being one of them. The recommendation is given not to describe new species-group taxa based on females only, unless they show evidently unique characters. J. Krikken* & J. Huijbregts, National Museum of Natural History Naturalis, Postbus 9517, NL-2300 RA Leiden, The Netherlands. [email protected]

Introduction – all attributes that suffice to distinguish them with- Until a few years ago the small scarabs treated in this out problems from other Oriental Scarabaeinae. paper figured in the literature and in collections un- They appear all to be associated with carrion and/or der the genus-group name Phacosoma Boucomont, excrements. Locally, at least in Southeast Asia, they 1914, a widely used junior homonym, for which the may be quite common in forest environments, being replacement name Ochicanthon Vaz-de-Mello, 2003 easily attracted to baited ground traps – even in for- was proposed. As currently understood Ochicanthon est remnants, as on Singapore Island. comprises a variety of more or less slender-legged This paper increases the number of known Ochican- Canthonini, essentially Oriental in distribution, thon species with 13 to a total of 36. Long series were most of them characterized by the combination of obtained during both our own field campaigns and long first tarsal segments on the middle and hind those of ecological colleagues, mainly on Borneo. On legs, a bidentate clypeus, an evenly, slightly convex analyzing ecological samples some decades ago (see dorsal side, a broadly produced, roundish protibial for instance Hanski & Krikken 1991), we already apex, and a double-crested elytral pseudepipleuron recognized several new Ochicanthon (= Phacosoma)

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species, but their taxonomic description had to be argumentation in General discussion). delayed due to our other obligations. Meanwhile, Most specimens were collected by pitfall trapping manuscript names and codes circulate, even a on ground surface level, using either human faeces genus-group name which should have replaced or fish as bait. All in all, about 1100 Ochicanthon Phacosoma, and some of the new Sundaland species specimens belonging to over 20 species were seen by were described by others (Ochi et al. 1996 through us, and all relevant published descriptions and illus- 2007). Nevertheless, as indicated, there are still trations of Ochicanthon were checked, and the infor- many undescribed species left, and more are likely mation processed to the best of our ability. Efforts to turn up in the future, as will be clear from this to access types of some species recently described paper. When we started looking at our Ochicanthon remained unsuccessful. Potentially related genera material the variation in several of the characters were screened to assess their taxonomic position seemed confusing and rather due to intraspecific in relation to Ochicanthon (see General discussion, polymorphism, but on going through the long series below). mentioned, we gradually learned that the operational units recognized from the outset nearly all represent- Characters and descriptive terminology ed species in their own right, as confirmed by the The diagnoses and descriptions follow a standard virtually constant, characteristic shape of the male pattern, taking into account the diagnostic value of genitalia of these units, and sometimes by their ecol- the Ochicanthon characters resulting from a character ogy. Although a few species seem to be more wide- analysis. The format of the character list (Appendix 1) spread, many species may have a limited distribution and other descriptive information follows from the and ecology, one being flightless – a phenomenon use of a DELTA oriented system (Dallwitz 2005) not unusual among canthonines (see General discus- tuned to our scarab diversity database. The list makes sion below). Records of Ochicanthon species located our use of characters to distinguish the known Sun- far from their type localities, particularly with species daland species transparent, and includes characters inhabiting primeval forest environments or sites with possibly significant in a future phylogeny reconstruc- a known history of ecological isolation, must always tion; the synoptic table facilitates multi-character be critically verified, starting with an analysis of the identification. structure of the male genitalia. Ecological remarks following the diagnoses are based This taxonomic review, part of a series on the tax- on both published reports and our own records. onomy of laparostict scarabs of the Indo-Australian Comments at the end of each of the species accounts islands, more than doubles the number of usually start with some comparative morphological Ochicanthon species known from Sundaland. We features, followed by any further comments. present a generic diagnosis, a list of the known spe- Some additional terminological explanation is in or- cies, a character analysis, a key to 23 Sundaland spe- der (for colour pattern coding and aedeagus elements cies, accounts of these species, including descriptions see next section). of 13 new species. A compilation of the material Interstriae are the intervals between the elytral striae; seen, including numerous new records, is appended. note that Ochicanthon are characterized by the pres- The systematic position of the genus, its species-level ence of a broad pseudepipleuron. Our term meta- diversity, some biogeographic aspects, and the ensu- tarsus denotes the entire tarsus of the hind leg, as ing research questions, are discussed at the end of distinct from metatarsal segment 1. this paper. The qualification abundant for microsculptural units (like punctures) usually means: separated by 2-5 diameters, dense: 1-2 diameters, crowded: less Material, methods, conventions than 1 diameter; sparse: at least 5 diameters. Estimates may be imprecise due to peripunctural depressions Scope and information sources (particularly on rugulate surfaces). Different types The substance of this paper concerns straightforward of punctures (like annulate or ocellate) are specified morphological taxonomy, i.e. the diagnosis and de- only if they are distinctly and unambiguously dif- scription of the new Sundaland species alluded to in ferent from simple punctures (a central micro-pit the introduction, and their insertion in an identifi- [potentially] supporting a seta, for instance, does not cation key. The Ochicanthon material on which this qualify them to be called ocellate). Attributes with study is based was nearly all collected in the context the prefix micro- are defined as those distinct at mag- of tropical forest biodiversity projects on Borneo. nifications of about 40, or higher. Some parts may We are aware that a phylogenetic analysis should fol- be characteristically microstriolate or microreticulate low, but this is at present considered premature (see (appearing sericeous, more or less matt).

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Marginate is a general term, meaning that the mar- the parameres may have sclerotized pieces; in some gin of the body element concerned is set off from the species a piece in the right paramere is developed into general surface (by being raised, narrowly reflexed, or a more or less curved surface (bottom sclerite), with whatever). The pronotum usually shows a fold paral- a pointed (usually basomedian) expansion; this ex- lel and close to the posterior section of the lateral pansion may take the form of a lobe or hook (in the border (opposite base of pseudepipleural crest), here dytiscoides group). The detailed configuration of the termed paramarginal fold. parameres differs among species, and an assessment Total body length measurements given are approxi- of this configuration is indispensable in morpho- mate (different body part postures may create impre- logical species identification. Proposing new species cision). All body part measurements taken in strictly based only on females lacking distinctly unique fea- dorsal view. Ratio interocular distance / (single) tures is absolutely to be avoided. maximum eye width measured along transverse axes. Length proportions of the metatibial terminal spur Illustrations versus the tarsal segments 1-5 given by the numbers As the differences between the species cannot all be s//a/b/c/d/e, as estimated with an ocular scale in the represented by abstract text and coding, compare microscope. the homologous elements in the pictures. We have It should be noted that some published descriptions chosen for comparative photographs of the various of Ochicanthon, also recent ones, are nearly useless, body parts of Ochicanthon (usually head, prono- lacking the detail required, lacking pictures compen- tum, elytron, protibia, metatibia and -femur), sup- sating this, and/or being based on a single unchar- plemented with diagrammatic line drawings of the acteristic female. Note that Paulian (1980, 1983, male genitalia (membranous structures in line draw- 1987) did not sex his material. ings indicated with streaks). In the captions note the different meanings of full-face versus other views; Colour patterns and aedeagus full-face here means: body part surface positioned In Sundaland some groups of Ochicanthon spe- maximally parallel to the plane of the picture; dor- cies can be distinguished, one being characterized sal and ventral views are those from above and be- by a patterned pronotum and/or ditto elytra (the low, parallel to the plane formed by the median and O. dytiscoides group). The basic pronotal and elytral transverse body axes. Body pilosity (and spurs) in colour patterns in this group are coded according to photographs of body parts (not in habitus pictures) the approximate position and fusion of the mark- may be (partially) removed to clearly show contours. ings, as follows (Fig. 3): In some pictures the body element is slightly tilted. • Pronotal lighter marks (yellow-orange-reddish) Refer to the measurements in the descriptions for on dark background: PL (= posterolateral), size indications. ML (= mediolateral), AL (= anterolateral), C (= central), BM (= basomedian), PBM (= para- Distribution and ecology basomedian), PAM (= para-apicomedian); these As for geographic terminology, Sundaland is here lighter marks may be expanded-confluent or re- defined as the Malay Peninsula + Borneo + Sumatra duced, according to (sub)species. + Java + Bali, plus intervening and adjacent smaller • Three sets of lighter elytral marks (with their islands, i.e. minus Sulawesi. The locations of place interstrial extension) are given as: B (= basal), names are usually easily traceable with current topo- M (= postmedial), A (apical); these marks may graphic tools (including travel guidebooks and the also be expanded-confluent or reduced over the Microsoft Encarta Atlas), but some are specified various interstriae 2-7 (as indicated by interstrial (Appendix 2). The generic distribution as given on numbers from the suture; comma means sepa- the map (Fig. 2) is large-scale and approximate, rated, hyphen means confluent; mark numbers with indications of gaps requiring further explora- between square brackets refer to vague patches). tion. The limit between lowland versus upland is The males in one group of species (the gangkui group) (somewhat arbitrarily) set at an altitude of 1000 m have their parameres more or less excised from the (no records from subalpine environments available). apex, usually in an asymmetric way, forming branch- Much of the material examined (Appendix 2) is es and lobes (when deeply excised a superior lobe or labelled with sample codes, here, in addition to the # branch may develop, distinct in lateral view). Many symbol, frequently preceded by (lower case) project Ochicanthon species show a superior sclerotized, usu- or collector initials. The indication “multistr evergr ally paired strut, distinct in upper side view, sitting forest” on many of the labels means multistratal ever- between or over the parameres, varying in shape green forest (a term used in the classification of Fos- (length) and degree of sclerotization. The bottom of berg 1967 to replace the ambiguous term rainforest).

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Note that some species may be restricted to special on apex. Postocular part of propectus evenly deeply forest types (like riverine, heath [kerangas] and lime- concave, separated from other propectoral parts by stone forest). Aliases were used to handle operational well-pronounced ridge. Tarsi present, slender. Meso- taxonomic units from ecological samples, and refer- and metatarsal segments 1 much longer than next ences are given to occurrences of these aliases in pub- segment. Head broader than long, clypeal apex with lished reports. paramedian denticles separated by widely rounded emargination (denticles may also laterally be lim- Types and deposition of material ited by emargination). Genae laterally subangular All specimens seen (including the types) are record- to rounded, border very weakly rounded or straight ed in Appendix 2, most of these being at present (and virtually uninterrupted) to clypeal denticles. kept in the National Museum of Natural History Tarsal claws simple. Mesometasternal suture straight, Naturalis, Leiden (RMNH), The Natural History transverse. Anal and anteanal abdominal sternite Museum, London (BMNH), and the National Mu- normally separated (not fused). seum of Wales, Cardiff (NMWC). The Muséum National d’Histoire Naturelle, Paris (MNHN) will Description receive paratypes of a number of new species. Re- Head. Head about as wide as long, apex usually distribution of material to other institutional col- 2-dentate, with or without variably deep adja- lections (including local ones) will take place in due cent emargination; in one known species quasi-4- course. Ecological samples are kept elsewhere, and dentate. Clypeus emarginate between the more or specimens in these samples may bear manuscript less angular, more or less reflexed paramedian den- names or codes (see previous section for aliases used ticles. Clypeofrontal surface generally very slightly by ecologists, or consult us). convex. Clypeofrontal transition without any eleva- The label data of some types are quoted verbatim in tion; sutures laterally distinct. Clypeal border from this paper, as follows: text lines are separated by \, eye-canthus forward rounded or virtually straight, at label upper- and underside text by \\, texts on dif- most with very slight protrusion or incision at su- ferent labels are enclosed in quotation marks; our ture; margin not reflexed. Genae projecting beyond comments or additions in square brackets [p] means eyes (in full-face view), their tip rounded to subangu- printed text, [h] handwritten text. lar. Frontovertex entirely devoid of protrusions. Eye ventrally globose, usually large, smaller (reduced) in certain species. Eyes (foramen in full-face view) usu- Ochicanthon Vaz-de-Mello ally small, more or less narrowly elliptic or reniform, Ochicanthon Vaz-de-Mello, 2003: nom. n. for Phacosoma to large, broadly elliptic. Boucomont, 1914, preoccupied by Phacosoma Jukes- Pronotum. Prothorax slightly less wide or about as Browne, 1912 (Mollusca). wide as elytra (which usually cover abdomen com- Phacosoma Boucomont, 1914, Type-species: Phacosoma dytiscoides Boucomont, 1914 (designated by Arrow pletely). Pronotum generally evenly convex, base, 1931). disc and anterior declivity unmodified (midline may Recent synopses: Balthasar 1963 [key to species]; be very slightly impressed near base). Prothorax in Ochi, et al. 1997, 2006 [key to species Borneo]; dorsal outline nearly parallel-sided, usually slightly Hanboonsong, et al. 2001 [key to species Thailand] convergent rostrad. Anterolateral pronotal angle rounded or angular (in some species straight or con- Generic diagnosis cave between two distinct angles). Lateral sides of Posterolateral section of pronotal surface with para- pronotum simply declivous. Posterolateral section marginal fold closely parallel to lateral border, run- of pronotum with (variably distinct) paramarginal ning forward from base at level of pseudepipleural fold running from level of pseudepipleural ridge for- ridge. Elytral surface moderately convex, with stria ward, sometimes extended into (angled) groove (fold 7 laterally limited by pronounced (usually anteriorly and groove may be vague, depending on coarseness geminate) pseudepipleural ridge, pseudepipleuron of microsculpture and angle of view). Pronotal pos- with two distinct striae. Procoxal cavity small, not terolateral angles more or less distinct, obtuse. Base distinctly transverse. Base of elytra from humerus of pronotum immarginate, usually evenly widely to humerus fitting well to pronotal base (at most rounded (at most a very slight indication of a me- a minimal pronotal-elytral constriction in dorsal dian angle). Pronotal sides finely marginate or im- view). Protibia with three large external denticles marginate, dorso-ventral transition sharp, simple. and proximal serration, inner side unmodified; apex Scutellum indistinct. extended, more or less lobiform (apex more or less Elytra. Disc generally evenly convex, punctate- rounded, short, not digitate), spur sitting externally striate, elytra combined usually slightly wider than

Downloaded from Brill.com10/11/2021 05:43:19AM via free access Krikken & Huijbregts: Ochicanthon in Sundaland 425 prothorax. Elytron between suture and humeral sec- slender, straight or curved (in some species very long, tion of pseudepipleural ridge with 7 punctate striae. slender, curved); without fossorial protrusions on Total number of elytral striae (including juxtepi- outer side. Both meso- and metatibia with acuminate pleural one) 9 – an additional stria on the pseude- terminal spurs. Metatibial apex somewhat lobate- pipleuron may be obliterated by the (anteriorly dentate or distal section of tibia more characteristi- usually duplicated) pseudepipleural ridge. Humeral cally modified. Tarsal claws fine to very fine, more umbone slight or absent. Anteapical elytral umbone or less sickle-shaped, not with extra denticle on un- vague, if distinct at all. Apicosutural angle of elytron derside. Metatarsal segment 1 distinctly longer than (sub)rectangular. Posterior declivity of elytron un- segment 2 (usually about twice as long), slender, modified. Elytral epipleuron narrow. Pseudepipleu- straight. ron reaching elytral apex (with 2 striae, including General. Body outline usually elliptic, medium-sized juxtepipleural one). Striae on elytral disc all reach- to small, length 2.5-10 mm. Dorsum evenly convex, ing base, distinct, shallowly impressed to superficial. distinctly patterned (usually reddish/orange to yel- Interstriae flat to feebly convex. low marks on brown or black) or uniformly dark- Ventral side. Antenna with nine segments. Anten- coloured (brown or black), underside brown or black, nal club relatively large, simply lamellate. Anten- head appendages and legs usually lighter (brownish nal scapus smooth, slender, long. Labial palpus to yellowish); some species with faint metallic lustre. three-segmented, unmodified. Maxillary palpus Usually much of surface strongly punctate, with mi- four-segmented, unmodified. Propectus with dis- crosetae and/or fine bristles, or just microstubbles. tinctly delimited, deeply concave postocular con- Aedeagi very diverse, from simply subsymmetrical, cavity on either side. Procoxal cavities not strongly pincer-shaped, to complex, asymmetrical, excised transverse, leaving free space on the side of the (usually with variably developed superior struts); propectus. Metasternal anterior lobe virtually flat. bottom may have more or less complex sclerite(s). Mesometasternal suture straight. Mesosternum (lon- Sexual dimorphism. Where dimorphism occurs this is gitudinally) short. Metasternal anterior lobe broad, mainly evident in shape of legs; then female protib- widely separating parallel mesocoxae, not protrud- ial and metatibial dentation relatively unmodified; ing. Metasternal disc flat to slightly convex, lacking female pro- and metafemur without extra angles, concavities. Abdominal sternites normally connate, dentation, lobes; metatibia ditto. Pygidial shape may usually 6 segments visible. Abdominal venter simply differ, arrangement of distal sternites may be slightly transversely convex to almost flat. Hind wings fully more compact in males. developed (with at least one exception). Pygidium Note. This description of the genus has been com- transverse, slightly convex to very convex (location posed with special reference to the Sundaland species of greatest convexity varies), lacking grooves or other at hand. well defined impressions; apical border widely arcuate, apex and base marginate. Distribution and ecology Legs. Profemur unmodified or produced on anterior Range mapped (Fig. 2): Oriental Region (trans- side (into angle or tooth). Procoxa small, not dis- gression into China; not known for certain from tinctly transversely set in prothorax. Protibia with Wallacea). Gap between southern and northern 3 larger external denticles (with or without some in- India, genus still unknown from most of Sumatra, tervening serration), shape of apico-external denticle Philippines, etc. varies, but usually acuminate. Proximal serration on Attracted to dung and carrion, inhabiting various outer side of protibia distinct. Protibial apex more forest types, and possibly more open habitats, up or less expanded beyond apico-external denticle, in to 2150 m (India); also collected from soil litter or some species lobiform, underside with slightly pro- forest debris. Most species are flyers, some getting tuberant end of longitudinal ridge. Internal side of into flight interception traps; at least one species is protibia evenly, slightly concave-curvilinear (full- flightless. face view). Protibial spur inserted away from inner angle, elongate-acuminate. Protarsus fine, length List of Ochicanthon species-group taxa not or slightly exceeding distal width of protibia, fine, segment 1-4 short, 5 longer. Mesocoxae widely Sundaland species arranged and numbered as in the separated, subparallel. Metacoxae (sub)contiguous. Species accounts, other species alphabetically ar- Femora without well-defined supplementary lon- ranged. All species-group names followed by author gitudinal ridge on their underside; metafemur may names between parentheses were formerly combined be produced on posterior side (angulate or lobate). with Phacosoma. Meso- and metatibiae distinctly dilated distad,

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Ochicanthon Vaz-de-Mello, 2003 • O. loebli (Paulian, 1980) – India (S) Phacosoma Boucomont, 1914 • O. nitidus (Paulian, 1980) – India (S) • O. obscurus (Boucomont, 1920) – Myanmar O. dytiscoides group (S), India (NE), Thailand, Laos, Vietnam (C) • O. ochii (Hanboonsong & Masumoto, 2001) O. dytiscoides subgroup – Thailand (N) 1. O. edmondsi sp.n. – Borneo (Sarawak) • O. philippinensis (Ochi, 1990) – Philippines 2. O. dytiscoides (Boucomont, 1914) – Borneo (Mindanao) (Sarawak, Sabah) • O. thai (Paulian, 1987) (as ssp. of tristoides) 3. O. masumotoi (Ochi & Araya, 1996) – Borneo – Thailand (N), Vietnam (N) (Sabah, Sarawak) Phacosoma fallcilaetum Masumoto, 1988 4. O. maryatiae Ochi et al., 2006 – Borneo • O. thailandicus (Masumoto,1989) – Thailand (Sabah) (N), China (Yunnan, Löbl & Smetana 2006) 5. O. rombauti sp.n. – Borneo (Sabah) • O. tristis (Arrow, 1931) – India (SW) • O. tristoides�� (Paulian, 1983) – India (NE) O. cambeforti subgroup 6. sp.n. – Borneo (Sabah) O. crockermontis Key to Sundaland Ochicanthon species 7. O. crypticus sp.n. – Borneo (Sarawak) 8. O. peninsularis sp.n. – Malay Peninsula + The key should primarily aid in the identification to Singapore species, and does not reflect the taxonomy implied in 9. O. kikutai Ochi et al., 2006 – Borneo (Sabah) this paper. See also the species groupings as indicated 10. O. neglectus sp.n. – Borneo (Sarawak) under the relevant headings in the section Species 11. O. dulitmontis sp.n. – Borneo (Sarawak) accounts and the relevant text under the respective 12. O. cambeforti (Ochi, Kon & Kikuta, 1997) species headings. Appendix 1 gives a synoptic table – Borneo (E Kalimantan) of the characters. Ochicanthon punctatus omitted O. simboroni Ochi et al., 2006 syn. n. from the key. 13. O. karasuyamai Ochi et al., 2007 – Sumatra (N) 1. Elytra uniform, entirely brown or black, un- O. woroae group patterned ...... 2 14. O. woroae Ochi et al., 2006 – Borneo (E Kali- – Elytra predominantly black-brown, with mantan) distinct pattern of (orange-)yellow marks . . . 15. O. tambunan sp.n. – Borneo (Sabah) ...... 10 16. O. mulu sp.n. – Borneo (Sabah) – Elytra as well as pronotum predominantly shining (orange-)yellow, symmetric black O. gangkui group marks limited ...... 6. O. crockermontis 17. O. gangkui (Ochi, Kon & Kikuta, 1997) 2. Alae present; pronotal-elytral transition (in – Borneo (Sabah) profile) gradual, habitus deplanate; clypeal 18. O. kimanis sp.n. – Borneo (Sabah) denticles separated by emargination down 19. O. danum sp.n. – Borneo (Sabah) to their base ...... 3 20. O. parantisae (Ochi, Kon & Kikuta, 1997) – Alae absent; pronotal-elytral transition (in – Borneo (Sabah) profile) with dip, habitus distinctly convex; 21. O. javanus sp.n. – Java clypeal denticles fused at base; internal side of male metabia angularly dilated halfway O. hanskii group its length ...... 22. O. hanskii 22. O. hanskii sp.n. – Borneo (Sabah) 3. Borneo ...... 4 – Java ...... 21. O. javanus Species not placed in group 4. Clypeal denticles adjoined laterally by sim- 23. O. hikidai (Ochi, Kon & Kikuta, 1997) ple concave curve ...... 5 – Borneo (W Kalimantan) – Clypeal denticles adjoined laterally by 24. O. punctatus (Boucomont, 1914) – Sumatra? shallow (sinuate) emargination; dorsum with coarse setae; parameres subsymmetric Species outside Sundaland (Fig. 130) ...... 16. O. mulu • O. cingalensis (Arrow, 1931) – Sri Lanka – Clypeal paramedian denticles adjoined lat- • O. deplanatus (Paulian, 1983) – India (NE) erally by deep emargination, producing ex- • O. laetus (Arrow, 1931) – India (S) tra denticle on either side . . . 23. O. hikidai

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5. Parameres subsymmetric (in upper side and – Protibial denticles (male) all simply, sharp- lateral view) ...... 6 ly acuminate; parameres different from – Parameres distinctly excised and/or asym- Fig. 123 ...... 14 metric, complex (lateral view) ...... 7 14. Parameres subsymmetric, elongate, their 6. Discal elytral interstriae with numerous apex pointed (upper side view, Fig. 124), scattered microsetae; pronotum with nu- not excised above (lateral view); discal ely- merous scattered microsetae; dorsal puncta- tral interstrial punctation abundant; clypeal tion more crowded, surface matt ...... denticles adjoined laterally by simple con- ...... 15. O. tambunan cave curve ...... 17 – Discal elytral interstriae mainly with scat- – Parameres asymmetric, short, upper side tered microstubbles; pronotum glabrous, or of right paramere excised (lateral view, nearly so; dorsum densely, distinctly punc- Fig. 127), complex; discal elytral interstrial tate, shining ...... 14. O. woroae punctation dense to crowded; elytral inter- 7. Left paramere very deeply excised, inferior striae shining between punctures, without branch elongate-expanded ...... 8 microreticulation; clypeal denticles laterally – Left paramere more shallowly excised, infe- virtually straight; Borneo . 12. O. cambeforti rior branch short ...... 9 15. Pronotal (orange-)yellow marks limited to 8. Right paramere sinuate-lobate (lateral view, small lateral patch on either side; anterola- Fig.132) ...... 17. O. gangkui teral border of pronotum concave, limited – Right paramere very widely arcuate (lateral by secondary angle; large, length at least 8 view, Fig. 133) ...... 18. O. kimanis mm, dorsum generally shining ...... 9. Pronotum glabrous, or nearly so; right para- ...... 1. O. edmondsi mere simply widely sinuate (Fig. 134) ...... – Pronotal (orange-)yellow marks extensive, ...... 19. O. danum (orange-)yellow predominant ...... – Pronotum with numerous scattered micro- ...... 2. O. dytiscoides setae; both parameres with deep rounded – Pronotal (orange-)yellow marks abundant, excision ...... 20. O. parantisae but (orange-)yellow not predominant . . . 16 10. Pronotum uniformly (brown-)black (at 16. Pronotal-elytral transition (in profile) grad- most with small, very vague marks) . . . 11 ual; small; discal elytral interstriae all flat or – Pronotum with distinct symmetric pattern very slightly convex; pygidium convex along of (orange-)yellow and (brown-)black . . . . . 13 apex, basal surface flat or concave (in pro- 11. Body size small (length usually much less file); dorsum generally shining ...... 18 than 8 mm) ...... 12 – Pronotal-elytral transition (in profile) – Body size large (length over 8 mm); metafe- abrupt; body size large (length usually at or mur (posterior side) and -tibia (internal over 8 mm); discal elytral interstriae 3 and side) without mini-crenulation; parameres 5 slightly more elevated; pygidium more or relatively short; dorsum not microreticulate . less evenly convex (in profile); dorsum gen- ...... 11. O. dulitmontis erally matt ...... 5. O. rombauti 12. Pygidium convex along apex, basal surface 17. Elytra interstriae matt, due to distinct mi- flat or concave (in profile); apical elytral croreticulation; Malay Peninsula ...... (orange-)yellow absent; discal interstrial ...... 8. O. peninsularis punctation dense to crowded; dorsum be- – Elytral interstriae shining, lacking microre- tween punctures shining, without microreticulation; Borneo ...... 9. O. kikutai ticulation; Borneo ...... 10. O. neglectus 18. Smaller species, with small bottom sclerite – Pygidium more or less conical; discal l inter- on aedeagus (lacking large backward hook) . . . strial punctation abundant; dorsum entirely ...... 3. O. masumotoi matt, due to microreticulation; Sumatra ...... – Larger species, with large, angular bottom ...... 13. O. karasuyamai sclerite, point protruding to basal piece of 13. Protibial denticles (male) all acuminate; aedeagus ...... 4. O. maryatiae parameres short (Fig.123); interstriae shining, basal marks on elytra starting on interstria 2 ...... 7. O. crypticus – Protibial denticles (male) 1-2 acuminate, but apico-external denticle broad, basal half or more parallel-sided ...... 15

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Fig. 1. Ochicanthon edmondsi, female of a new species from forest in Sarawak.

Species accounts

(a) The Ochicanthon dytiscoides group (a1) The Ochicanthon dytiscoides subgroup At least elytra (usually also pronotum) with symmetric colour pattern (orange-yellow versus brown- 1. Ochicanthon edmondsi sp. n. black). Parameres usually subsymmetric (in upper Figs 1, 4-9, 118 side view). Metatibia of (major) males slightly angularly dilated (internally) near apex. Profemur of Type-material. Male holotype (RMNH): Malaysia, (major) males angularly expanded in front. with our type label, and the following label data: Two subgroups distinguishable in Sundaland: males Sarawak: 4th Division: Gunung Mulu NP, iii/1978, with broadened, deflexed apico-external denticle on I. Hanski, baited pitfall trap, LMR [lower montane protibia (dytiscoides subgroup), or with simply acu- rainforest], 800-1700 m, #668. minate denticle (cambeforti subgroup). 15 paratypes listed in Appendix 2.

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AL PAM ML PL C PBM BM B

Fig. 2. Ochicanthon, approximate range of the genus Fig. 3. Ochicanthon, coding of basic pronotal and (Asia). elytral colour pattern in the dytiscoides group.

Description (holotype, male) with metallic lustre, posterolaterally with yellow- Body length ca 8.8 mm. Colour generally black, with orange spot on either side. yellow marks on pronotum and elytra. Elytra black-brown, shining, with yellow-orange pat- Clypeal margin anteriorly bidentate (tips angular, tern, on disc with abundant appressed, long micro- median emargination widely rounded). Clypeo- setae. Pseudepipleuron reaching elytral apex (broad, frontal surface generally feebly convex, black-brown, pseudepipleural crest with 2 fine ridges in front). shining, with metallic lustre; glabrous, with crowd- Elytral striae distinct, shallowly impressed; punc- ed, distinct, simple punctation. Clypeal border from tures of discal striae slightly impressed (crenulating gena forward widely rounded. Gena laterally round- interstriae, separated by 1-3 times their diameter); ed. Eyes (foramen in full-face view) medium-sized, interstriae flat to feebly convex, simply punctate, moderately elliptic. Maximum number of facet punctures scattered, crowded. rows (transversely) across eye foramen ca 9. Ratio Antennal lamellae brown. transverse interocular distance/maximum eye width Propectus laterally black-brown, virtually smooth, ca 6.0. microstriolate. Mesosternum virtually smooth. Pronotum generally evenly convex, disc and anterior Metasternal anterior lobe and disc densely declivity unmodified, slightly deplanate. Protho- punctate-microsetose, setae very short. Metasternum rax in dorsal outline nearly parallel-sided, slightly black-brown, laterally with annulate punctation (on convergent rostrad. Anterior border of pronotum microreticulate surface), dense. Abdominal venter unmodified. Anterolateral pronotal angle (nearly) black-brown, proximal sternites with narrow punc- rectangular. Anterolateral section of pronotum very tate basal zone, distal sternites densely punctate- distinctly angulate at ca 0.2 of length behind anterior microsetose, setae long, non-punctate microstriolate angle. Lateral sides of pronotum simply declivous. surface matt. Hind wings fully developed. Pygidium Posterolateral section of pronotum modified as in evenly, slightly convex, marginate along base, black- all congeners, with distinct fold parallel and close brown, densely punctate-microsetose, punctures to border. Pronotal posterolateral angles simple, fine. distinct. Base of pronotum immarginate, evenly, Profemur brownish, projecting in front (distinct very widely rounded, surface unmodified. Pronotal obtuse angle near apex). Protibia externally with border finely marginate in front. Pronotum micro- 3 denticles (apico-external denticle very broad, large- setose, setae mostly long, appressed; generally with ly parallel-sided, with rounded [worn?], deflexed simple crowded punctation. Estimated number of tip). Proximal serration on outer side of protibia dis- punctures per 0.25 sq mm on pronotal disc ca 50- tinct, fine. Protibia with rounded, protruding apex; 60, punctures ca 0.06 mm. Pronotum black-brown, internal side evenly, slightly curved. Protibial spur

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inserted away from inner angle, simply acuminate, Discal elytral interstrial punctation dense to crowd- fine. Protibia dark brown. Protarsi slender. Meso- ed; interstriae shining, without microreticulation, femur slender, unmodified. Metafemur posteriorly all flat or very slightly convex, with numerous scat- with slight projecting angle at 0.3 from base and tered microsetae. Pygidium slightly evenly convex, larger, but very obtuse angle at 0.2-0.3 from apex. brown to black, (virtually) uniform. Protibial den- Meso- and metafemur dark brown. Meso- and ticles (male) 1-2 acuminate, apico-external denticle metatibiae long, curved, without fossorial protru- very robust, extremely broad, its base largely parallel- sions on outer side. Mesotibia simply curved. Meta- sided. Profemur (male) anteriorly strongly lobate- tibia strongly curved, gradually dilated to apex; apex dentate near apex. Metafemur (male) posteriorly modified, obtusely angular internally. Meso- and doubly lobate(-dentate), short blunt denticle near metatibia dark brown. Meso- and metatarsi slender. base. Metatibia long, slender, curved, dilated near Metatarsal segment 1 distinctly longer than seg- apex. Metatibia internally (male) continuous in- ment 2 (approximate proportions spur//segments ternally (not angulate halfway), angulate-dilated at 1-5 7//29/13/9/7/11). short distance from apex. Metafemur posteriorly Aedeagus large, more or less symmetric, as in and metatibia internally not multidenticulate. Alae Fig. 118; bottom sclerite of right paramere distinct, present. Parameres subsymmetric (in upper side curled, with strong basomedian point. view), long, dilated distally, somewhat spatuliform Measurements in mm. Maximum width of head 2.5. (full-face upper side); bottom sclerite of right para- Pronotal dorsal median length 2.6, maximum width mere well-developed, tip projecting basomedially 4.5. Dorsal sutural length of elytra 4.6, maximum (distinct in profile). Pronotal-elytral transition (in width combined 5.1. profile) gradual, habitus deplanate. Body size large Colour patterns. Pronotal lighter marks present: (length usually at or over 8 mm). PL pair only. Elytral lighter marks present: B (5-7), Both sexes known. Length 8-10 mm. M (5-7), A (2-3, 5-7). Distribution and ecology Variation and sexual dimorphism Borneo: Sarawak. Protibial dentation of female simply acuminate. An- Hanski (1983, as Phacosoma sp. n. B) collected this gles on anterior side of profemur and posterior side species in carrion baited pitfall traps at Mulu, at 800- of metafemur in female absent. Distal section of fe- 1700 m altitude, in what was termed lower montane male metatibia unmodified. rain forest (no dung traps were operated on the sites concerned). Derivation of species name Dedicated to the distinguished American scarabaeol- Comments ogist W.D. Edmonds, who contributed much to the O. edmondsi may well be the largest member of the taxonomy and ecology of New World Scarabaeinae, genus; it is easily recognizable from the characteris- particularly the Phanaeini. tically shaped, anterolaterally concave prothoracic border (creating a secondary anterolateral angle), Diagnosis and a well-defined yellow versus black-brown dorsal Clypeal denticles separated by emargination down colour pattern, although only a small yellow patch to their base, adjoined laterally by simple concave on either side of pronotum is present; apico-exter- curve. Clypeogenal border from lateral tip to ante- nal denticle on male protibia remarkably broad, de- rior denticles widely convex-curvilinear. Clypeofron- flexed; parameres large, distally dilated (in full-face tal surface densely to crowdedly punctate. Eye size view). moderate, foramen narrowly elliptic. Pronotal border anterolaterally (at ca 0.2 behind anterior angle) 2. Ochicanthon dytiscoides (Boucomont, 1914) conspicuously angularly bent inward. Paramarginal Figs 16-21, 119 fold on posterolateral surface of pronotum distinct. Pronotum densely to crowdedly punctate, with numerous scattered microsetae. Pronotal (orange-) Diagnosis yellow marks limited to small but distinct lateral Clypeal denticles separated by emargination down patch on either side. Elytra with distinct symmet- to their base, adjoined laterally by simple concave ric pattern of (brown-)black and (orange-)yellow curve. Clypeogenal border from lateral tip to an- marks. Basal elytral (orange-)yellow B (5-7) present. terior denticles widely convex-curvilinear. Clypeo- Postmedial elytral (orange-)yellow M (5-7) present. frontal surface densely to crowdedly punctate. Apical elytral (orange-)yellow A (2-3, 5-7) present. Eye size moderate, foramen narrowly elliptic.

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4 10 8

5 11

6 12

7 13 9

Figs 4-15. Ochicanthon, habitus (male holotypes) and body parts. – 4-9, O. edmondsi, habitus, oblique (4); head, full-face (5); pronotum, dorsal (6); left elytron, dorsal (7); protibia, upper side (8); metatibia and -femur, underside (9); 10-15, O. rombauti, habitus, oblique (10); head, full-face (11); pronotum, dorsal (12); left elytron, dorsal (13); protibia, upper side (14); metatibia and -femur, underside (15).

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Pronotal border anterolaterally (at ca 0.2 behind an- Comments and lectotype designation terior angle) more or less abruptly rounded. Paramar- O. dytiscoides is a somewhat variably patterned spe- ginal fold on posterolateral surface of pronotum short, cies, possibly limited to Borneo; it is recognizable moderately distinct. Pronotum densely to crowdedly from its largely yellow pronotum, densely to crowd- punctate, with numerous scattered microsetae, with edly punctate, non-sericeous elytral interstriae, and symmetric pattern of (orange-)yellow and (brown-) dilated apico-external protibial denticle in the male black. Pronotal (orange-)yellow extensive on disc. sex. The closely related O. masumotoi (as here con- Elytra with symmetric pattern of (brown-)black and ceived) has a largely dark pronotum, but appears (orange-)yellow marks. Basal elytral (orange-)yellow B otherwise very similar to dytiscoides. The forms (5-7) present. Postmedial elytral (orange-)yellow M around dytiscoides need revision, taking both mor- (5-7) present. Apical elytral (orange-)yellow A phological and molecular characters from different (3, [5-7]) present. Discal elytral interstrial puncta- populations into account. Pending the outcome of tion dense to crowded, interstriae between punctures such work, some action to stabilize the nomenclature shining, without microreticulation, interstriae all flat seems appropriate. or very slightly convex, with numerous scattered mi- Boucomont (1914), when describing his Phacosoma crosetae. Pygidium convex along apex, basal surface dytiscoides, apparently had two species before him, flat (in profile), largely (orange-)yellow, but may be one from Kinabalu (agreeing with the present con- more or less infuscated. Protibial denticles (male) cept of dytiscoides), and one from Singapore (agreeing 1-2 acuminate, apico-external denticle broad, its with the present concept of peninsularis, see below). base more or less parallel-sided. Profemur (male) an- We have examined type material, and hereby desig- teriorly lobate-dentate near apex. Metafemur (male) nate a lectotype, effectively restricting the type local- posteriorly lobate-dentate at 0.3-0.5 from base. ity to North Borneo. The male lectotype is labelled as Metatibia long, slender, curved, dilated near apex. follows: “N. BORNÉO\KINA BALU\COLL. V.DE Metatibia internally (male) continuous internally POLL” [p], “MUSEUM PARIS [p]\Boucomont (not angulate halfway), angulate-dilated at short [h]”, “Typus” [p in box, red label], “Boucomont distance from apex (male). Metafemur posteriorly det. 191[p]4\ Phacosoma n.g.\dytiscoides n.sp.[h]”, and metatibia internally not multidenticulate. Alae “jk7912”[h] (in MNHN). Handwriting on labels, present. Parameres subsymmetric, long, slender, ta- apart from last label, by Boucomont. Male genitalia pering (lateral and upper side view), bottom of right of lectotype extracted, on a card, on the pin with paramere with basomedially pointed, curled sclerite. the type. The indication Kinabalu on the type label Pronotal-elytral transition (in profile) gradual, habi- needs not necessarily refer to the mountain. tus deplanate. Body size small. The information in Ochi et al. (1997) on the iden- Both sexes known. Length 5-7 mm. tity of O. dytiscoides seemed inconsistent, which was the primary reason for us to re-examine Bouco- Distribution and ecology mont’s Kinabalu specimen just cited. The protibial Borneo. New data listed in Appendix 2. apico-external denticle of the lectotype is broadened, We have seen much O. dytiscoides-like material from not simply acuminate, as suggested in their paper Sabah, Brunei and Sarawak. Most specimens were (by their Fig. 20). This inconsistency was recog- collected in pitfall traps baited with either carrion nized in Ochi et al. (2006), referring their mate- or human faeces. A few specimens were collected on rial to O. kikutai (q.v.), without settling the matter fungi and fruit. Extensive trapping with flight inter- completely: this is done by the present lectotype ception traps in Danum Valley (Sabah) yielded only designation. one specimen. All specimens were collected in lowland forest at 20-300 m altitude. At Mulu (Sarawak) 3. Ochicanthon masumotoi (Ochi & Araya, O. dytiscoides was only found in kerangas (heath for- 1996) est), while Davis characterizes this species in Danum Figs 22-27, 120 Valley as a riverine/edge specialist – all the more reason to take a closer look at the taxonomic status of the various populations (see comments hereafter). Diagnosis Generally, this species seems to have a preference Clypeal denticles separated by emargination down for open forest types. (Mentioned in Hanski 1983; to their base, adjoined laterally by simple concave Davis 1999, Davis et al. 2000, Davis 2000, Davis curve. Clypeogenal border from lateral tip to an- et al. 2001.) terior denticles widely convex-curvilinear. Clypeo- frontal surface densely to crowdedly punctate. Eye size moderate, foramen narrowly elliptic. Pronotal

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16 22 20

17 23

18 24

19 25

Figs 16-27. Ochicanthon, habitus (males) and body parts. – 16-21, O. dytiscoides, habitus, oblique (16); head, full-face (17); pronotum, dorsal (18); left elytron, dorsal (19); protibia, upper side (20); metatibia and -femur, underside (21); 22-27, O. masumotoi, habitus, oblique (22); head, full-face (23); pronotum, dorsal (24); left elytron, dorsal (25); protibia, upper side (26); metatibia and -femur, underside (27).

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border anterolaterally (at ca 0.2 behind anterior an- Comments gle) more or less abruptly rounded. Paramarginal O. masumotoi was originally based on a single female fold on posterolateral surface of pronotum short, from Sabah (without further locality specification), moderately distinct. Pronotum densely to crowd- and may (as conceived here) be recognizable from its edly punctate, with numerous scattered microse- patterned, largely brownish-black pronotum, crowd- tae, with symmetric pattern of (orange-)yellow and edly punctate, non-sericeous elytral interstriae, and (brown-)black. Pronotal (orange-)yellow marks may dilated apico-external protibial denticle in the male be abundant, but not predominant (small on disc). sex (provided that the present male-female associa- Elytra with distinct, usually symmetric pattern of tion is correct). In their key Ochi et al. (2006) men- (brown-)black and (orange-)yellow marks. Basal tion body size and eye distance in relation to eye elytral (orange-)yellow B (5-7) present. Postmedial width as important characters. elytral (orange-)yellow M (5-7) present. Apical ely- In this review we have simply assigned every tral (orange-)yellow A (3-7) present. Discal elytral O. dytiscoides-like specimen with a dark, patterned interstrial punctation crowded, interstriae shining pronotum to O. masumotoi, being aware that the between punctures, without microreticulation, in- two species are very close, and suspecting that some terstriae all flat or very slightly convex, interstriae or all of the material may eventually turn out to be with numerous scattered microsetae. Pygidium con- conspecific – with the material now available definite vex along apex, basal surface flat (in profile), largely conclusions cannot be drawn. The aedeagi of the two (orange-)yellow, but may be more or less infuscated. species are very similar. Protibial denticles (male) 1-2 acuminate, apico- See also comments under O. dytiscoides above. external denticle broad, its base parallel-sided. Pro- femur (male) anteriorly lobate-dentate near apex. Metafemur (male) posteriorly lobate-dentate at 4. Ochicanthon maryatiae Ochi, Ueda & Kon, 0.3-0.5 from base. Metatibia long, slender, curved, 2006 dilated near apex. Metatibia internally (male) continuous internally (not angulate halfway), angulate Diagnosis at short distance from apex. Metafemur posteri- Clypeal denticles separated by emargination down orly and metatibia internally not multidenticulate. to their base, adjoined laterally by simple concave Alae present. Parameres subsymmetric, long, slen- curve. Clypeogenal border from lateral tip to an- der, tapering to fine point (in lateral and upper side terior denticles widely convex-curvilinear. Clypeo- view), bottom of right paramere with basomedially frontal surface densely to crowdedly punctate to pointed, curled sclerite. Pronotal-elytral transition rugulate-punctate. Eye moderate, foramen moderate- (in profile) gradual, habitus deplanate. Body size ly or narrowly elliptic. Pronotal border anterolaterally small. (at ca 0.2 behind anterior angle) more or less abruptly Both sexes known. Length 4-6.5 mm. rounded. Paramarginal fold on posterolateral surface of pronotum distinct, (angularly) extended in front. Distribution and ecology Pronotum densely to crowdedly punctate, with nu- Borneo. New data listed in Appendix 2. merous scattered microsetae, with symmetric pattern We have seen O. masumotoi from both Sabah and of (orange-)yellow and (brown-)black (disc in some Sarawak. Most specimens were collected in pitfall specimens lighter). Pronotal (orange-)yellow marks traps baited with carrion, smaller numbers with hu- abundant, (orange-)yellow usually not predominant. man excrements, fungi, or with flight interception Elytra predominantly black-brown, with (usually traps. In the Mulu region (Sarawak) it was collected symmetric) pattern of (orange-)yellow marks. Basal at 715-1070 m altitude, in both high mixed dip- elytral (orange-)yellow B(5-7) present. Mediolateral terocarp forest and lower montane rain forest – in elytral (orange-)yellow M(5-7) present. Distal elytral Hanski’s (1983) terminology. Although on the sites (orange-)yellow A(5-7) present. Discal elytral inter- concerned both carrion and dung traps were oper- strial punctation dense to crowded, interstriae shiny, ated, O. masumotoi was only collected in fish and without microreticulation, interstriae all flat or very meat traps. Davis characterizes the occurrence of slightly convex, interstriae with numerous scattered this species in the Danum Valley lowland forest as microsetae. Pygidium convex along apex, basal sur- a riverine/edge specialist. (Mentioned in Hanski face flat or concave (in profile). Protibial denticles 1983, as Phacosoma sp.n. F, in Davis et al. 2000, as (male) 1-2 acuminate, apico-external denticle broad, Phacosoma pictum MS, in Davis 1999, 2000; Davis its base parallel-sided. Metafemur (male) posteriorly et al. 2001, as Phacosoma sp. 3.) lobate-dentate at 0.3-0.5 from base. Metatibia long, slender, curved, dilated near apex. Internal side of

Downloaded from Brill.com10/11/2021 05:43:19AM via free access Krikken & Huijbregts: Ochicanthon in Sundaland 435 metatibia (male) continuous (not angulate halfway), Prothorax in dorsal outline nearly parallel-sided, very angulate-dilated at short distance from apex (male). slightly sinuate, waisted at base. Anterior border of Metafemur posteriorly and metatibia internally not pronotum unmodified. Anterolateral pronotal angle multidenticulate. Meso- and metatarsi long, relative- obtuse, rounded off; anterolateral section of prono- ly slender. Alae present. Parameres roughly subsym- tal border rounded behind anterior angle, adjacent metric with pointed apex (in upperside view), long, surface slightly depressed. Lateral sides of pronotum slender, tapering (lateral view); bottom sclerite large, simply declivous. Posterolateral section of pronotum angular tip pointing well backward (distinct in pro- modified as in all congeners, with distinct fold paral- file). Pronotal-elytral transition (in profile) gradual. lel and close to border, distinctly extended (sulcate) Habitus deplanate. Body rather large. in front. Pronotal posterolateral angles rounded. Both sexes known. Length 7.2-8.1 mm. Base of pronotum immarginate, evenly, very widely rounded, with surface unmodified. Pronotal border Distribution finely marginate in front. Pronotum microsetose, Borneo: Sabah. setae short, erect; surface generally microreticulate and with crowded, more or less rugulate, ocellate Comments punctation. Estimated number of punctures per This recently described species should, in its group, 0.25 sq mm on pronotal disc 55-65, punctures immediately be distinguishable by the combination ca 0.04 mm (away from midline, depressed midline of its colour pattern, relatively large size, and charac- less punctate). Pronotum brown-black, faintly me- teristic aedeagal structure, including a large, angu- tallic, with vague yellow-orange mark on either side. lar bottom sclerite (Ochi et al. 2006, Figs 19a-d). Elytra black-brown, with vague yellow-orange pat- O. maryatiae was collected only at Poring, on the tern, matt, general surface slightly uneven, wrin- slopes of Mt Kinabalu (Sabah), at 1200 m altitude. kly. Humeral and anteapical umbones indistinct. Our collecting efforts around Poring Hot Springs Pseudepipleuron reaching elytral apex (broad, matt, (baited pitfall trapping, sifting cattle dung, etc.) did pseudepipleural crest with 2 fine ridges in front). not yield any Ochicanthon, but this may be due to a Elytral surface with erect numerous microsetae; difference in location. striae superficial, punctures of discal striae slightly impressed (slightly crenulating interstriae, separated by 3-4 times their diameter); interstriae flat to fee- 5. Ochicanthon rombauti sp.n. bly convex (basal surface of 3 and 5 raised), simply Figs 10-15, 121 punctate and microreticulate, punctation scattered, Type-material. Male holotype (RMNH): Malaysia, crowded to dense. with our type label, and the following label data: Antennal lamellae light-brown. Sabah: Crocker Range: Keningau-Kimanis road Propectus laterally black-brown, largely smooth, (km25), 18-23/xi/1987, J. Krikken & E. Rombaut, shining, vaguely punctate. finely marginate in front. 1300 m, multistr evergr forest, 8 human excr traps, Mesosternum virtually smooth. Metasternal anterior #sa45. lobe and disc densely punctate-microsetose, setae 72 paratypes listed in Appendix 2. very short; metasternum black-brown, laterally with dense, annulate punctation (on vaguely microstri- Description (holotype, male) olate surface). Abdominal venter black-brown, matt, Body length ca 8.2 mm. Colour generally brown- abundantly punctate-microsetose distally, setae black, with yellow-orange pattern on pronotum and short, proximally with microstriolation, matt; basal elytra. margin of proximal sternites punctate. Hind wings Clypeal margin anteriorly bidentate (tips angu- present. Pygidium very convex (near apex), under- lar, short, median emargination widely rounded). side with fine median ridge, marginate along base, Clypeofrontal surface generally feebly convex, black-brown, very densely punctate-microsetose, brown-black, matt, glabrous, with crowded, distinct, punctures very distinct. ocellate punctation. Clypeal border from gena for- Profemur brown, projecting in front (slight, obtuse ward widely rounded. Gena laterally rounded. Eyes denticle at ca 0.3 from apex). Protibia externally with (foramen in full-face view) medium-sized, moderate- 3 denticles (apico-external denticle broad, apex ob- ly elliptic. Maximum number of facet rows (trans- tusely angular, largely parallel-sided). Proximal serra- versely) across eye foramen ca 10. Ratio transverse tion on outer side of protibia, distinct, fine. Protibia interocular distance/maximum eye width ca 8.0. with rounded, protruding apex, internal side evenly, Pronotum generally evenly, moderately convex- slightly curved. Protibial spur inserted away from in- declivous in front, midline vaguely depressed. ner angle, simply acuminate, fine. Protibia brown.

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Protarsi slender. Mesofemur slender, unmodified. interstrial surface sericeous, due to microreticula- Metafemur posteriorly with slight protrusion at tion, with abundant, short, more or less erect mi- 0.3 from base, otherwise unmodified. Meso- and crosetae; interstriae 3 and 5 slightly elevated at base. metafemur brown. Meso- and metatibiae long, Pygidium (male) convex near apex, its underside me- curved, without fossorial protrusions on outer side. dially finely carinate, basal surface concave; brown Metatibia strongly curved, dilated near apex (with to black, base may be lighter. Protibial denticles slight, obtuse apico-internal angle). Meso- and (male) 1-2 acuminate, apico-external denticle broad, metatibia brown. Meso- and metatarsi slender. Meta- its base parallel-sided. Profemur (male) anteriorly tarsal segment 1 distinctly longer than segment 2 lobate-dentate near apex. Metafemur (male) poste- (approximate proportions riorly lobate-dentate at 0.3-0.5 from base. Metatibia spur//segments 1-5 10//24/13/9/7/12). long, slender, curved, dilated near apex. Metatibia Aedeagus large, more or less symmetric, as in internally (male) continuous internally (not angu- Fig. 121; bottom sclerite of right paramere dis- late halfway), slightly angulate near apex. Metafemur tinct, tip sharply pointed mesad, halfway parameral posteriorly and metatibia internally not multiden- length. ticulate. Alae present. Parameres subsymmetric (in Measurements in mm. Maximum width of head 2.4. upper side view), long, slender, pincer-shaped; bot- Pronotal dorsal median length 2.7, maximum width tom sclerite of right paramere distinct, with sharp 4.3. Dorsal sutural length of elytra 4.0, maximum point about halfway parameral length. Pronotal- width combined 4.9. elytral transition (in profile) abrupt, habitus strongly Colour patterns. Pronotal lighter marks very vague: convex. Body size large (length usually at or over PL only. Elytral lighter marks vague: B ([3-4]-5-7), 8 mm). M (5-7), A (2-3, 5-7). Both sexes known. Length 8-9 mm.

Variation and sexual dimorphism Distribution and ecology Protibial dentation of female simply acuminate. Borneo: Sabah. Angles on anterior side of profemur and posterior All specimens were collected in upland forest in the side of metafemur in female absent. Distal section of Crocker Range, in human faeces baited pitfall traps female metatibia unmodified. at 1300 m altitude (no carrion traps were set in this area). Derivation of species name Dedicated to our able colleague E. Rombaut, who Comments accompanied Krikken on the 1987 trip during which O. rombauti is a large, matt, vaguely patterned Sabah this species was collected. species, with wrinkly elytra, related to the preceding set of species on account of its broad apico-external Diagnosis protibial denticle (in the male sex). The swollen gen- Clypeal denticles separated by emargination down eral appearance, with the pronotal-elytral dip (dis- to their base, adjoined laterally by simple concave tinct in profile), suggests flightlessness (as is the case curve. Clypeogenal border from lateral tip to ante- in O. hanskii, q.v.), but the hind wings of rombauti rior denticles widely convex-curvilinear. Clypeofron- seem normally developed and functional. tal surface crowdedly punctate. Eye size small (note underside of eye). Pronotal border anterolaterally (at ca 0.2 behind anterior angle) simply rounded. Para- (a2) The Ochicanthon cambeforti subgroup marginal fold on posterolateral surface of pronotum distinct, angularly extended in front (sulcate). Pro- notum with very shallow midline depression, crowd- 6. Ochicanthon crockermontis sp.n. edly punctate, with numerous more or less erect Figs 28-33, 122 scattered microsetae, largely blackish, vague posterolateral patch only. Pronotal (orange-)yellow marks Type-material. Male holotype (RMNH): Malaysia, extensive, (orange-)yellow not predominant, vague. with our type label, and the following label Elytra with symmetric pattern of (brown-)black and data: Sabah: Crocker Range: Kota Kinabalu- (orange-)yellow marks. Basal elytral (orange-) Tambunan road (km64), 21-24/xi/1987, J. Krikken yellow B ([3-4]-5-7) present. Postmedial elytral & E. Rombaut, 1150 m, multistr evergreen forest, (orange-)yellow M (5-7) present. Apical elytral 3 human excr traps, #sa49a. (orange-)yellow A (2-3, 5-7) present. Discal elytral 5 paratypes listed in Appendix 2. surface slightly uneven; interstrial punctation dense,

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Description (holotype, male) Hind wings fully developed. Pygidium very convex Body length ca 5.8 mm. Colour generally orange, near apex, remainder flat, marginate along base, with black marks on pronotum and elytra. largely yellowish, densely punctate-microsetose, Clypeal margin anteriorly bidentate (tips angular, punctures fine. median emargination widely rounded). Clypeo- Profemur black-brown, projecting in front (distinct frontal surface generally feebly convex, brown, very angle near apex). Protibia externally with 3 denticles shining, somewhat metallic, glabrous, with crowded, (apico-external denticle simply tapering). Proximal distinct, fine, simple, shallow punctation. Clypeal serration on outer side of protibia distinct, fine. Pro- border from gena forward widely rounded. Gena lat- tibia with rounded, protruding apex, internal side erally rounded. Eyes (foramen in full-face view) me- evenly, slightly curved. Protibial spur inserted away dium-sized, moderately elliptic. Maximum number from inner angle, simply acuminate, fine. Protibia of facet rows (transversely) across eye foramen ca 7. brown. Protarsi slender. Mesofemur slender, un- Ratio transverse interocular distance/maximum eye modified. Metafemur posteriorly with very slight width ca 9.6. angle at 0.3 from base, otherwise unmodified. Meso- Pronotum generally evenly convex, disc and anterior and metafemur black-brown. Meso- and metatibiae declivity unmodified, slightly deplanate. Protho- long, curved, without fossorial protrusions on outer rax in dorsal outline nearly parallel-sided, slightly side. Metatibia strongly curved, abruptly dilated near convergent rostrad. Anterior border of pronotum apex (with slight internal angle); apex lobate-dentate. unmodified. Anterolateral pronotal angle (nearly) Meso- and metatibia brown. Meso- and metatarsi rectangular. Anterolateral section of pronotum virtu- slender. Metatarsal segment 1 distinctly longer than ally straight, more or less angular at ca 0.2 of length segment 2 (approximate proportions spur//segments behind anterior angle. Lateral sides of pronotum 1-5 10//20/9/6/5/6). simply declivous. Posterolateral section of pronotum Aedeagus more or less symmetric, as in Fig. 122; modified as in all congeners, with distinct fold paral- bottom sclerite of right paramere double curled, pro- lel and close to border, sulcate. Pronotal posterolater- jecting basally on either side. al angles simple, distinct, obtuse. Base of pronotum Measurements in mm. Maximum width of head 1.8. immarginate, narrow margin is metallic green; base Pronotal dorsal median length 1.7, maximum width overall very widely, evenly rounded, with surface un- 3.0. Dorsal sutural length of elytra 3.1, maximum modified. Pronotal border finely marginate in front. width combined 3.7. Pronotum with microsetae, generally with simple, Colour patterns unusual. Pronotum orange-yellow, dense to crowded punctation. Estimated number of with few dark symmetric marks. Elytra orange- punctures per 0.25 sq mm on pronotal disc 60-65, yellow, with few dark symmetric marks. punctures ca 0.04 mm. Pronotum largely orange, very shining, with few symmetric, somewhat blurred Variation and sexual dimorphism black marks. Angles on anterior side of profemur and posterior Elytra largely orange, very shining, with few sym- side of metafemur in female absent. Distal section of metric, somewhat blurred black marks, on disc with female metatibia unmodified abundant microsetae. Pseudepipleuron reaching elytral apex (broad, pseudepipleural crest with 2 fine Derivation of species name ridges in front). Elytral striae distinct, shallowly im- Named after the type locality, situated in the Crocker pressed; punctures of discal striae slightly impressed Range. (crenulating interstriae, separated by 2-3 times their diameter). Elytra with abundant microstubbles; Diagnosis interstriae flat to feebly convex, simply punctate Clypeal denticles separated by emargination down to (making slightly annulate impression), punctation their base, adjoined laterally by simple concave curve. scattered, dense. Clypeogenal border from lateral tip to anterior den- Antennal lamellae brown. ticles widely convex-curvilinear. Clypeofrontal sur- Propectus laterally black-brown, microstriolate. face crowdedly punctate. Eye size moderate, foramen Mesosternum virtually smooth. Metasternal ante- narrowly elliptic. Pronotal border anterolaterally rior lobe and disc densely punctate-microsetose, straight (virtually angular at ca 0.2 behind anterior setae very short; metasternum black-brown, later- angle). Paramarginal fold on posterolateral surface ally with dense, annulate punctation. Abdominal of pronotum short, distinct. Pronotum crowd- venter black-brown, matt, abundantly punctate- edly punctate, with microsetae. Pronotal (orange-) microsetose distally, setae short; proximally with yellow very extensive, covering most of surface. Elytra microstriolation and vague basal punctulation, matt. also with extensive (orange-)yellow, dark marks

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somewhat blurred. Discal elytral interstrial puncta- moderately elliptic. Maximum number of facet tion dense to crowded, interstriae shining, without rows (transversely) across eye foramen ca 10. Ratio microreticulation. Interstriae all flat or very slightly transverse interocular distance/maximum eye width convex, mainly with scattered microsetae. Pygidium ca 5.0. convex along apex, basal surface flat (in profile), large- Pronotum generally evenly convex, disc and anterior ly (orange-)yellow. Protibial external denticles (male) declivity unmodified, slightly deplanate. Protho- all acuminate. Profemur (male) anteriorly lobate- rax in dorsal outline nearly parallel-sided, slightly dentate near apex. Metafemur (male) posteriorly lo- convergent rostrad. Anterior border of pronotum bate-dentate at 0.3-0.5 from base. Metatibia long, unmodified. Anterolateral pronotal angle (nearly) slender, curved, dilated near apex. Metatibia internally rectangular. Anterolateral section of pronotum sub- (male) continuous internally (not angulate halfway), angular at ca 0.2 of length behind anterior angle. slightly angulate-dilated at short distance from apex Lateral sides of pronotum simply declivous. Postero- (male). Metafemur posteriorly and metatibia inter- lateral section of pronotum modified as in all conge- nally not multidenticulate. Alae present. Parameres ners, with distinct fold parallel and close to border, subsymmetric (in upper side view), short, broad, anteriorly somewhat sulcate. Pronotal posterolateral tapering (lateral view); superior strut of aedeagus angles simple, distinct, obtuse. Base of pronotum distinctly elongate; bottom sclerite distinctly curled. immarginate, evenly, very widely rounded, with Pronotal-elytral transition (in profile) gradual, habitus surface unmodified. Pronotal border finely margin- deplanate. Body size small. ate in front. Pronotum with microsetae; generally Both sexes known. Length 5.5-6 mm. with simple, dense punctation. Estimated number of punctures per 0.25 sq mm on pronotal disc 60-65, Distribution and ecology punctures ca 0.04 m\\\\m. Pronotum black-brown, Borneo: Sabah. shining, with metallic lustre and with yellow-orange All specimens were collected in upland forest in the pattern. Crocker Range, in human faeces baited pitfall traps, Elytra black-brown, with yellow-orange pattern, at 1150 m altitude (no carrion traps were set in the shining, with abundant microsetae. Pseudepipleuron area). reaching elytral apex (broad, crest with 2 fine ridges in front). Elytral striae distinct, shallowly impressed; Comments punctures of discal striae slightly impressed (crenu- O. crockermontis is a uniquely patterned species, its lating interstriae, separated by 3-4 times their dorsum being conspicuously shining orange-yellow, diameter); interstriae flat to feebly convex, sim- with a limited, somewhat vague, symmetric dark ply punctate, punctures scattered, dense (almost pattern; anterolateral section of pronotal border rugulate). straight. Antennal lamellae brown. Propectus laterally brown, microstriolate, matt. Mesosternum virtually smooth. Metasternal anterior 7. Ochicanthon crypticus sp.n. lobe and disc densely punctate-microsetose, setae Figs 34-39, 123 very short; metasternum black-brown, laterally with Type-material. Male holotype (RMNH): Malaysia, dense annulate punctation (on microreticulate sur- with our type label, and the following label data: face). Abdominal venter black-brown, matt, abun- Sarawak: 4th Division: Gunung Mulu NP, iii-v/1978, dantly punctate-microsetose distally, setae short, I. Hanski, 100-500 m, baited pitfall trap, MD [mixed proximally with microstriolation and vague punctu- dipterocarp] forest, #413. lation. Hind wings fully developed. Pygidium very 48 paratypes listed in Appendix 2. convex near apex, remainder flat, marginate along base, largely yellow, densely punctate-microsetose, Description (holotype, male) punctures fine. Body length ca 5.7 mm. Colour black-brown, with Profemur brown-yellow, projecting in front (distinct yellow-orange-mark on pronotum and elytra. denticle near apex). Protibia with 3 external denti- Clypeal margin anteriorly bidentate (tips angular, cles, all acuminate (apico-external denticle also sim- median emargination widely rounded). Clypeofron- ply tapering). Proximal serration on external side of tal surface generally feebly convex, brown, shining, protibia distinct, fine. Protibia with rounded, pro- with faint metallic lustre; glabrous, with crowded, truding apex, internal side evenly, slightly curved. distinct, simple punctation. Clypeal border from Protibial spur inserted away from inner angle, simply gena forward widely rounded. Gena laterally round- acuminate, very fine. Protibia brown. Protarsi slen- ed. Eyes (foramen in full-face view) medium-sized, der. Mesofemur slender, unmodified. Metafemur

Downloaded from Brill.com10/11/2021 05:43:19AM via free access Krikken & Huijbregts: Ochicanthon in Sundaland 439 posteriorly with extremely slight angle at 0.3 from denticles (male) all simply acuminate. Profemur base, otherwise unmodified. Meso- and metafe- (male) anteriorly lobate-dentate near apex. Metafe- mur brown. Meso- and metatibiae brown, long, mur (male) posteriorly at most lobate-dentate at 0.3- curved, without fossorial protrusions on outer side. 0.5 from base. Metatibia long, slender, curved, dilat- Mesotibia simply curved. Metatibia strongly curved, ed near apex. Metatibia internally (male) continuous abruptly dilated near apex (with internal angle); internally (not angulate halfway), angulate-dilated at apex lobate-dentate. Meso- and metatarsi slender. short distance from apex (male). Metafemur poste- Metatarsal segment 1 distinctly longer than seg- riorly and metatibia internally not multidenticulate. ment 2 (approximate proportions spur vs. segments Alae present. Parameres subsymmetric (in upper side 1-5 8//17/7/6/4/7). view), short, broad, tapering (lateral view); bottom Aedeagus more or less symmetric, as in Fig. 123; sclerite distinct. Habitus deplanate, body size small. bottom sclerite of right paramere distinct, flat, with Both sexes known. Length 5.5-6 mm. slight basomedian point. Measurements in mm. Maximum width of head 1.9. Distribution and ecology Pronotal dorsal median length 1.8, maximum width Borneo: Sarawak. 3.1. Dorsal sutural length of elytra 3.0, maximum All specimens were collected by Hanski (1983, width combined 3.7. as Phacosoma sp.n. A) in pitfall traps baited with Colour patterns. Pronotal lighter marks present: either carrion or human excrement, at Mulu, at PL-ML, AL, [BM], [PBM], [PAM]. Elytral lighter 100-670 m altitude, mainly in what was termed marks present: B(2-7), M(5-7)-A(2-7). mixed dipterocarp forest; a small number was found in alluvial forest. Variation and sexual dimorphism Angles on anterior side of profemur and posterior Comments side of metafemur in female absent. Distal section O. crypticus is a small species, mainly recognizable of female metatibia unmodified. Protibial denticles from its elytral colour pattern (shining, yellow marks vary in length. Pygidium may be yellow or brown, on elytral base, from interstria 2 on), short, virtu- or intermediate. ally unmodified parameres, and simply acuminate, rather short protibial denticles. Derivation of species name This species was initially hidden in a multi-species 8. Ochicanthon peninsularis sp.n. set under the name O. dytiscoides. Figs 40-45, 124 Diagnosis Type-material. Male holotype (BMNH): Malaysia, Clypeal denticles separated by emargination down with our type label, and the following label data: to their base, adjoined laterally by simple concave Malaysia: Pahang: Fraser’s Hill, 29/x-3/xi/1977, curve. Clypeogenal border from lateral tip to an- B. Bendell. terior denticles widely convex-curvilinear. Clypeo- 4 paratypes listed in Appendix 2. frontal surface densely to crowdedly punctate. Eye size moderate, foramen rather narrowly elliptic. Description (holotype, male) Pronotal border anterolaterally (at ca 0.2 behind Body length ca 6.7 mm. Colour generally black- anterior angle) abruptly rounded. Paramarginal brown, partly matt, pronotum and elytra with fold on posterolateral surface of pronotum distinct. yellow-orange marks. Pronotum densely to crowdedly punctate, with mi- Clypeal margin anteriorly bidentate (tips angular, crosetae, with symmetric pattern of (orange-)yellow median emargination widely rounded). Clypeo- and (brown-)black. Pronotal (orange-)yellow marks frontal surface generally feebly convex, brown, shiny, extensive, but not predominant. Elytra with sym- glabrous, with crowded, distinct, simple punctation. metric pattern of (brown-)black and (orange-)yellow Clypeal border from gena forward widely rounded. marks. Basal elytral (orange-)yellow B (3-7 ) present. Eyes (foramen in full-face view) medium-sized, Postmedial elytral (orange-)yellow M (5-7) present. rather broadly elliptic. Maximum number of facet Apical elytral (orange-)yellow A (2-7) present. Dis- rows (transversely) across eye foramen ca 9. Ratio cal elytral interstrial punctation dense to crowded, transverse interocular distance/maximum eye width interstriae shining, without microreticulation, all flat ca 7.6. or very slightly convex, mainly with scattered mi- Pronotum generally evenly convex, disc and anterior crosetae. Pygidium convex along apex, basal surface declivity unmodified, slightly deplanate. Protho- flat (in profile), largely yellow to brown. Protibial rax in dorsal outline nearly parallel-sided, slightly

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28 34 32

29 35 38

30 36

31 37 39

Figs 28-39. Ochicanthon, habitus (male holotypes) and body parts. – 28-33, O. crockermontis, habitus, oblique (28), head, full-face (29); pronotum, dorsal (30); left elytron, dorsal (31); protibia, upper side (32); metatibia and -femur, underside (33); 34-39, O. crypticus, habitus, oblique (34); head, full-face (35); pronotum, dorsal (36); left elytron, dorsal (37); protibia, upper side (38); metatibia and -femur, underside (39).

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40 46 44

41 47 50

42 48

43 49 45

Figs 40-51. Ochicanthon, habitus and body parts. – 40-45, O. peninsularis (male holotype), habitus, oblique (40); head, full-face (41); pronotum, dorsal (42); left elytron, dorsal (43); protibia, upper side (44); metatibia and -femur, underside) (45); 46-51, O. karasuyamai (female), habitus, oblique (46); head, full-face (47); pronotum, dorsal (48); left elytron, dorsal (49); protibia, upper side (50); metatibia and -femur, underside (51).

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convergent rostrad. Anterior border of pronotum than segment 2 (approximate proportions spur// unmodified. Anterolateral pronotal angle (nearly) segments 1-5 10//25/11/9/6/10). rectangular. Anterolateral section of pronotum Aedeagus more or less symmetric, as in Fig. 124; bot- abruptly rounded at ca 0.2 of length behind anterior tom sclerite of right paramere distinct, curled, with angle. Lateral sides of pronotum simply declivous. slight basomedian point. Posterolateral section of pronotum modified as in Measurements in mm. Maximum width of head 2.2. all congeners, with distinct fold parallel and close to Pronotal dorsal median length 2.1, maximum width border, anteriorly somewhat sulcate. Pronotal poste- 3.6. Dorsal sutural length of elytra 3.5, maximum rolateral angles simple, distinct, obtuse. Base of pro- width combined 4.4. notum immarginate, evenly, very widely rounded, Colour patterns. Pronotal lighter marks present: with surface unmodified. Pronotal border finely PL-ML-AL-BM-PBM-PAM. Elytral lighter marks marginate in front. Pronotum microsetose, setae present: B(2-7), M(5-7)-A(2-7). moderately long; largely yellow, shiny, with metallic lustre and with symmetric black marks; generally Variation and sexual dimorphism with simple, crowded punctation. Estimated number Angles on anterior side of profemur and posterior of punctures per 0.25 sq.mm on pronotal disc 55- side of metafemur in female absent. Distal section 65, punctures 0.05 mm. of female metatibia unmodified. Colour pattern of Elytra black-brown, with yellow-orange pattern, pronotum and interstrial punctation of elytra very matt (sericeous), on disc with abundant, moder- variable. ately long microsetae. Pseudepipleuron reaching elytral apex (broad, pseudepipleural crest with 2 fine Derivation of species name ridges in front). Elytral striae distinct, shallowly im- Occurring on the peninsular extension of continen- pressed; punctures of discal striae slightly impressed tal Southeast Asia. (crenulating interstriae, separated by 1-3 diameters); interstriae flat to feebly convex, simply punctate, Diagnosis punctures scattered, abundant. Clypeal denticles separated by emargination down to Antennal lamellae brown. their base, adjoined laterally by simple concave curve. Propectus laterally black-brown, microstriolate. Clypeogenal border from lateral tip to anterior den- Mesosternum virtually smooth. Metasternal anterior ticles widely convex-curvilinear. Clypeofrontal sur- lobe and disc densely punctate-microsetose, setae face crowdedly punctate. Eye size moderate, foramen very short; metasternum black-brown, laterally with narrowly elliptic. Pronotal border anterolaterally (at dense, annulate punctation (on microreticulate sur- ca 0.2 behind anterior angle) more or less abruptly face). Abdominal venter black-brown, matt, abun- rounded. Paramarginal fold on posterolateral surface dantly punctate-microsetose distally, setae short, of pronotum distinct. Pronotum crowdedly punc- proximally with microstriolation and base of ster- tate, with numerous scattered microsetae, with sym- nites with vague punctulation, matt. Hind wings metric pattern of (orange-)yellow and (brown-)black. fully developed. Pygidium very convex (near apex, Pronotal (orange-)yellow marks variably extensive. base depressed), marginate along base, yellow, mar- Elytra with symmetric pattern of (brown-)black gins infuscated, densely punctate, punctures fine. and (orange-)yellow marks. Basal elytral (orange-) Profemur brown, projecting in front (distinct den- yellow B (2-7) present. Postmedial elytral (orange-) ticle near apex). Protibia externally with 3 denticles yellow M (5-7) present. Apical elytral (orange-) (apico-external denticle simply tapering). Proximal yellow A (2-7) present. Discal elytral interstrial serration on outer side of protibia distinct, fine. Pro- punctation abundant to dense, interstriae sericeous, tibia with rounded, protruding apex, internal side due to microreticulation between punctures, inter- evenly, slightly curved. Protibial spur inserted away striae all flat or very slightly convex, interstriae with from inner angle, simply acuminate, fine. Protibia numerous scattered microsetae. Pygidium convex brown. Protarsi slender. Mesofemur slender, un- along apex, basal surface flat or concave (in profile), modified. Metafemur posteriorly with angle at 0.3 largely (orange-)yellow. Protibial denticles (male) all from base, distal-posterior edge expanded, curved. simply acuminate, slender. Profemur (male) anteri- Meso- and metafemur brown. Meso- and metatibiae orly lobate-dentate near apex (distinct). Metafemur brown, long, curved, without fossorial protrusions (male) posteriorly lobate-dentate at 0.3-0.5 from on outer side. Mesotibia simply curved. Metatibia base to doubly lobate(-dentate). Metatibia long, slen- strongly curved, abruptly dilated close to apex (with der, curved, dilated near apex. Metatibia internally internal angle); apex lobate-dentate. Meso- and met- (male) continuous internally (not angulate halfway), atarsi slender. Metatarsal segment 1 distinctly longer angulate-dilated at short distance from apex (male).

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Metafemur posteriorly and metatibia internally not or concave (in profile). Protibial denticles (male) all multidenticulate. Alae present. Parameres subsym- simply acuminate, slender. Profemur (male) ante- metric (in upper side view), rather long, slender, riorly lobate-dentate near apex. Metafemur (male) tapering (lateral view); bottom sclerite distinct. posteriorly doubly lobate(-dentate). Metatibia long, Pronotal-elytral transition (in profile) gradual, habi- slender, curved, dilated near apex. Internal side of tus deplanate. Body size small. metatibia (male) continuous (not angulate halfway), Both sexes known. Length 6-7 mm. angulate-dilated at short distance from apex (male). Metafemur posteriorly and metatibia internally not Distribution multidenticulate. Meso- and metatarsi long, rela- Malay Peninsula, incl. Singapore. tively slender. Alae present. Parameres subsymmetric (in upperside view), long, slender, tapering (lateral Comments view); bottom sclerite distinct. Pronotal-elytral tran- O. peninsularis is a patterned species, with pro- sition (in profile) gradual. Habitus deplanate. Body tibial denticles all simply acuminate in both sexes; size small. background microsculpture of elytra distinctly Both sexes known. Length 7.2-7.5 mm. microreticulate (giving the interstriae a sericeous appearance), and elytral basal and apical marks ex- Distribution tensive. Also occurring on Singapore Island (Bukit Borneo: Sabah. Timah), whence we have a male specimen very similar to the one described from the same area by Comments Boucomont (1914) under P. dytiscoides. Consider- O. kikutai is a patterned species with all acuminate ing its variability and wide range possibly a set of protibial denticles, close to O. peninsularis, but lack- (sub)species – more material required. O. kikutai ing the microreticulation on the elytral interstriae may be a close relative, but is distinguished easily by characteristic of that species. The aedeagi of the its non-microreticulate elytra and different aedeagus. two species are also different (see Ochi et al. 2006, Figs 20a-d). Ochi et al. (1997) misinterpreted as , rectifying this in Ochi et al. 9. Ochicanthon kikutai Ochi, Ueda & Kon, O. kikutai dytiscoides 2006 (2006). It was found only at Sayap (Sabah), at 1000 m altitude.

Diagnosis 10. Ochicanthon neglectus sp.n. Clypeal denticles separated by emargination down Figs 52-57, 125 to their base, adjoined laterally by simple concave curve. Clypeogenal border from lateral tip to anteri- Type-material. Male holotype (RMNH): Malaysia, or denticles widely convex-curvilinear. Clypeofrontal with our type label, and the following label data: surface densely to crowdedly punctate to rugulate- Sarawak: 4th Division: Gunung Mulu NP, iii-v/1978, punctate. Eye moderate, foramen moderately or nar- I. Hanski, 400-600 m, pitfall trap, fish bait, lime- rowly elliptic. Pronotal border anterolaterally (at ca stone forest, #204. 0.2 behind anterior angle) more or less angular. Para- 11 paratypes listed in Appendix 2. marginal fold on posterolateral surface of pronotum short, moderately distinct to distinct, (angularly) Description (holotype, male) extended in front. Pronotum densely to crowdedly Body length ca 6.8 mm. Colour generally black- punctate, with microstubbles, with symmetric pat- brown, elytra with yellow-orange marks. tern of (orange-)yellow and (brown-)black. Prono- Clypeal margin anteriorly bidentate (tips angular, tal (orange-)yellow marks extensive, (orange-)yellow median emargination widely rounded). Clypeo- predominant. Elytra predominantly black-brown, frontal surface generally feebly convex, black-brown, with (usually symmetric) pattern of (orange-)yellow shining, glabrous, with dense to crowded, distinct, marks. Basal elytral (orange-)yellow B(5-7) present. simple punctation. Clypeal border from gena for- Mediolateral elytral (orange-)yellow M(5-7) present. ward widely rounded. Gena laterally rounded. Eyes Distal elytral (orange-)yellow A(2-7) present. Dis- (foramen in full-face view) medium-sized, moderate- cal elytral interstrial punctation dense to crowded, ly elliptic. Maximum number of facet rows (trans- interstriae shining between punctures, without versely) across eye foramen ca 9. Ratio transverse microreticulation, interstriae all flat or very slightly interocular distance/maximum eye width ca 5.6. convex, interstriae with numerous scattered micro- Pronotum generally evenly convex, disc and an- setae. Pygidium convex along apex, basal surface flat terior declivity unmodified, slightly deplanate.

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52 58 56

53 59 62

54 60

55 61 57

Figs 52-63. Ochicanthon, habitus (holotypes) and body parts. – 52-57, O. neglectus, habitus, oblique (52); head, full-face (53); pronotum, dorsal (54); left elytron, dorsal (55); protibia, upper side (56); metatibia and -femur, underside (57); 58-63, O. dulitmontis, habitus, oblique (58); head, full-face (59); pronotum, dorsal (60); left elytron, dorsal (61); protibia, upper side (62); metatibia and -femur, underside (63).

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64 69 68

66 67 65

Figs 64-69. Ochicanthon, habitus and body parts (male holotype). – O. cambeforti, habitus, oblique (64); head, full-face (65); pronotum, dorsal (66); left elytron, dorsal (67); protibia, upper side (68); metatibia and -femur, underside (69).

Prothorax in dorsal outline nearly parallel-sided, apex (broad, pseudepipleural crest with 2 fine ridges slightly convergent rostrad. Anterior border of pro- in front). Elytral striae distinct, shallowly impressed; notum unmodified. Anterolateral pronotal angle punctures of discal elytral striae slightly impressed (nearly) rectangular. Anterolateral section of pro- (crenulating interstriae, separated by 2-4 times their notum abruptly rounded at ca 0.2 of length behind diameter); surface with appressed, long microsetae; anterior angle. Lateral sides of pronotum simply de- interstriae flat to feebly convex, simply punctate, clivous. Posterolateral section of pronotum modified punctures scattered, crowded. as in all congeners, with distinct fold parallel and Antennal lamellae brown. close to border, anteriorly somewhat sulcate. Pro- Propectus laterally black-brown, microstriolate. notal posterolateral angles simple, distinct, obtuse. Mesosternum virtually smooth. Metasternal anterior Base of pronotum immarginate, evenly, very widely lobe and disc densely punctate-microsetose, setae rounded, with surface unmodified. Pronotal border very short; metasternum black-brown, laterally with finely marginate in front. Pronotum entirely black- dense, annulate punctation (on microreticulate sur- brown, microsetose, setae long, appressed; surface face). Abdominal venter black-brown, matt, with generally with simple punctation, crowded. Estimat- vaguely punctate basal zone on proximal sternites, ed number of punctures per 0.25 sq.mm on pronotal abundantly punctate-microsetose distally, setae disc ca 55-65, punctures ca 0.04 mm. long, with microstriolation, matt. Hind wings fully Elytra black-brown, with yellow-orange pattern, developed. Pygidium very convex near apex, mar- moderately shining, on disc with abundant ap- ginate along base, black-brown, densely punctate- pressed microsetae. Pseudepipleuron reaching elytral microsetose, punctures fine.

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Profemur brown, projecting in front (denticle virtually absent (2-7 lighter brown, but indistinct). near apex). Protibia externally with 3 denticles Discal elytral interstrial punctation dense to crowd- (apico-external denticle simply tapering). Proximal ed, interstriae shining, without microreticulation, serration on outer side of protibia distinct, fine. Pro- interstriae all flat or very slightly convex, with nu- tibia with rounded, protruding apex, internal side merous scattered microsetae. Pygidium convex near evenly, slightly curved. Protibial spur inserted away apex, basal surface flat (in profile), brown to black, from inner angle, simply acuminate, fine. Protibia (virtually) uniform. Protibial denticles (male) all dark brown. Protarsi slender. Mesofemur slender, simply acuminate, slender. Profemur (male) ante- unmodified. Metafemur posteriorly with angle at riorly lobate-dentate near apex (slight). Metafemur 0.3 from base, followed by row of fine, irregular (male) posteriorly lobate-dentate at 0.3-0.5 from denticulations. Meso- and metafemur dark brown. base. Metatibia long, slender, curved, dilated near Meso- and metatibiae long, curved, without fossorial apex. Metatibia internally (male) continuous in- protrusions on outer side. Mesotibia simply curved. ternally (not angulate halfway), angulate-dilated at Metatibia strongly curved, gradually dilated to apex, short distance from apex (male). Metafemur posteri- basal half with inner surface irregularly denticulate; orly and metatibia internally with longitudinal row apex modified (angular). Meso- and metatibia dark of fine denticles. Alae present. Parameres subsym- brown. Meso- and metatarsi slender. Metatarsal seg- metric (in upper side view), long, slender, tapering ment 1 distinctly longer than segment 2 (approximate (lateral view); bottom sclerite distinct. Pronotal- proportions spur//segments 1-5 10//24/11/7/6/10). elytral transition (in profile) gradual, habitus Aedeagus more or less symmetric, as in Fig. 125; bot- deplanate. Body size small. tom sclerite of right paramere distinct, with slight Both sexes known. Length 7-8 mm. basomedian point. Measurements in mm. Maximum width of head 2.1. Distribution and ecology Pronotal dorsal median length 2.2, maximum width Borneo: Sarawak. 3.5. Dorsal sutural length of elytra 3.4, maximum All specimens were collected by Hanski (1983, as width combined 6.2. Phacosoma sp.n. G) in fish baited pitfall traps at Colour patterns. Pronotal lighter marks absent. Ely- Mt Api in Mulu, in limestone forest. In this area tral lighter marks present: B(5-7), M(5-7)-A([2-7]). only fish baited traps were set, at altitudes of 400 and 650 m, and O. neglectus was collected at both alti- Variation and sexual dimorphism tudes. Angles on anterior side of profemur and posterior side of metafemur in female absent. Distal section Comments of female metatibia unmodified. Female also lacking O. neglectus is a small species from limestone for- metafemoral and -tibial denticulation. est in Sarawak, characterized by an all black pronotum, and finely multidenticulate (mini-crenulate) Derivation of species name posterior-interior sides of the metafemora and This species was initially missed, being hidden in a metatibiae in the male sex (best seen from above). multi-species set under the name dytiscoides. O. dulitmontis has also a uniformly dark pronotum, but is presumably larger (one male known), and Diagnosis lacks the multidenticulate ornamentation on the Clypeal denticles separated by emargination down to hind legs. their base, adjoined laterally by simple concave curve. Clypeogenal border from lateral tip to anterior den- 11. Ochicanthon dulitmontis sp.n. ticles widely convex-curvilinear. Clypeofrontal sur- Figs 58-63, 126 face crowdedly punctate. Eye size moderate, foramen narrowly elliptic. Pronotal border anterolaterally (at Type-material. Male holotype only (BMNH): ca 0.2 behind anterior angle) more or less abruptly Malaysia, with our type label, and the following label rounded (subangular). Paramarginal fold on poste- data: Sarawak: Mt Dulit, 18/x/1932, B.M. Hobby & rolateral surface of pronotum short, distinct. Prono- A.W. Moore (Oxford Univ. Exped.), 4000 ft, moss tum densely to crowdedly punctate, with numerous forest. scattered microsetae, uniform (brown-)black. Elytra with symmetric pattern of (brown-)black and (or- Description (holotype, male) ange-)yellow marks. Basal elytral (orange-)yellow B Body length ca 8.2 mm. Colour generally black- (5-7) present. Postmedial elytral (orange-)yellow M brown, elytra with yellow-orange marks. (5-7) present. Apical elytral (orange-)yellow (A) Clypeal margin anteriorly bidentate (tips angular,

Downloaded from Brill.com10/11/2021 05:43:19AM via free access Krikken & Huijbregts: Ochicanthon in Sundaland 447 median emargination widely rounded). Clypeo- from inner angle, simply acuminate, fine. Protibia frontal surface generally feebly convex, black-brown, brown. Protarsi slender. Mesofemur slender, un- matt, glabrous, with crowded, distinct, simple punc- modified. Metafemur posteriorly with denticle at tation. Clypeal border from gena forward widely 0.3 from base, followed by distinctly convex, sim- rounded. Gena laterally rounded. Eyes (foramen in ple curve. Meso- and metafemur brown. Meso- and full-face view) medium-sized, moderately elliptic. metatibiae long, curved, without fossorial protrusions Maximum number of facet rows (transversely) across on outer side. Mesotibia simply curved. Metatibia eye foramen ca 10. Ratio transverse interocular strongly curved, gradually dilated to apex, inner sur- distance/maximum eye width ca 8.9. face lacking denticulation; apex modified (angular). Pronotum generally evenly convex, disc and anterior Meso- and metatibia brown. Meso- and metatarsi declivity unmodified, slightly deplanate. Protho- slender. Metatarsal segment 1 distinctly longer than rax in dorsal outline nearly parallel-sided, slightly segment 2 (approximate proportions spur//segments convergent rostrad. Anterior border of pronotum 1-5 12//30/14/10/9/12). (Legs partly missing.) unmodified. Anterolateral pronotal angle (nearly) Aedeagus more or less symmetric (poorly preserved), rectangular. Anterolateral section of pronotum sub- as in Fig. 126. angular at ca 0.2 of length behind anterior angle. Measurements in mm. Maximum width of head 2.6. Lateral sides of pronotum simply declivous. Pos- Pronotal dorsal median length 2.5, maximum width terolateral section of pronotum modified as in all 4.3. Dorsal sutural length of elytra 4.0, maximum congeners, with distinct fold parallel and close to width combined 5.2. border. Pronotal posterolateral angles simple, distinct, Colour patterns. Pronotal lighter marks absent. Ely- obtuse. Base of pronotum immarginate, evenly, very tral lighter marks present: B (5-7), M (5-7), A (3). widely rounded, with surface unmodified. Pronotal border finely marginate in front. Pronotum entirely Derivation of species name black-brown, microsetose, setae long, appressed; Named after the type locality, situated in the moun- surface generally with simple punctation, crowded. tainous hinterland of Sarawak. Estimated number of punctures per 0.25 sq.mm on pronotal disc 60-65, punctures ca 0.04 mm. Diagnosis Elytra dark brown, with yellow-orange pattern, Clypeal denticles separated by emargination down moderately shining, on disc with abundant ap- to their base, adjoined laterally by simple concave pressed microsetae. Pseudepipleuron reaching elytral curve. Clypeogenal border from lateral tip to ante- apex (broad, pseudepipleural crest with 2 fine ridges rior denticles widely convex-curvilinear. Clypeofron- in front). Punctures of discal elytral striae slightly tal surface crowdedly punctate. Eye size moderate, impressed (crenulating interstriae, separated by foramen narrowly elliptic. Pronotal border anterola- 2-3 times their diameter). Elytral striae distinct, terally (at ca 0.2 behind angle) more or less abruptly shallowly impressed. Elytra with microsetae only rounded (subangular). Paramarginal fold on poste- (appressed, long). Interstriae flat to feebly convex, rolateral surface of pronotum short, distinct. Prono- simply punctate, punctures scattered, crowded. tum crowdedly punctate, with numerous scattered Antennal lamellae brown. microsetae, uniform (brown-)black. Elytra with Propectus laterally black-brown, microstriolate. symmetric pattern of (brown-)black and (orange-) Mesosternum virtually smooth. Metasternal anterior yellow marks. Basal elytral (orange-)yellow B (5-7) lobe and disc densely punctate-microsetose, setae present. Postmedial elytral (orange-)yellow M (5-7) very short; metasternum black-brown, laterally with present. Apical elytral (orange-)yellow limited, A (3) dense, annulate punctation (on microreticulate sur- present. Discal elytral interstrial punctation crowd- face). Abdominal venter black-brown, matt, abun- ed, interstriae shining, without microreticulation, dantly punctate-microsetose distally, setae long, with interstriae all flat or very slightly convex, interstriae microstriolation, matt. Hind wings fully developed. with numerous scattered microsetae. Pygidium con- Pygidium very convex (near apex), marginate along vex along apex, basal surface flat (in profile) to more base, brown, densely punctate-microsetose, punc- or less pointed, brown to black, (virtually) uniform. tures fine. Protibial denticles (male) all simply acuminate, Profemur brown, projecting in front (large denti- slender. Profemur (male) anteriorly lobate-dentate cle near apex). Protibia externally with 3 denticles near apex. Metafemur (male) posteriorly lobate- (apico-external denticle simply tapering). Proximal dentate at 0.3-0.5 from base. Metatibia long, slen- serration on outer side of protibia distinct, fine. Pro- der, curved, dilated near apex. Metatibia internally tibia with rounded, protruding apex, internal side (male) continuous internally (not angulate halfway), evenly, slightly curved. Protibial spur inserted away angulate-dilated at short distance from apex (male).

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Metafemur posteriorly and metatibia internally not lobate-dentate near apex. Metafemur (male) poste- multidenticulate. Alae present. Parameres subsym- riorly doubly lobate(-dentate), distal-posterior lobe metric (in upper side view), long, slender, tapering rounded. Metatibia long, slender, curved, dilated (lateral view). Pronotal-elytral transition (in profile) near apex. Metatibia internally (male) continuous gradual, habitus deplanate. Body size large (length at internally (not angulate halfway), angulate-dilated at or over 8 mm). short distance from apex (male). Metafemur poste- One male available, female unknown. Length be- riorly and metatibia internally not multidenticulate. tween 8-8.5 mm. Alae present. Parameres distinctly excised above, asymmetric. Left paramere broad, tapering. Right Distribution and ecology paramere widely sinuate (with narrow apex, in lateral Borneo: Sarawak. view); bottom sclerite small, flat, with distinct ba- Collected in moist upland forest. somedian projection. Pronotal-elytral transition (in profile) gradual, habitus deplanate. Body size small. Comments Possibly both sexes known (diagnosis above prima- O. dulitmontis presumably is a large species (one rily based on male holotype of cambeforti figured). male available), possibly related to the much smaller Length 5-6.5 mm. neglectus, and different in the details of the legs (hind femoral and tibial multidenticulation absent) and Distribution the aedeagus (parameres short, plump); note that our Borneo: East Kalimantan. Type data of cambeforti in information on aedeagal characters need verification. Appendix 2. Boucomont saw this specimen, labelling it dytiscoides (“comp. au type”). Comments In spite of its asymmetric parameres, everything else in (like the patterned dorsum) points 12. Ochicanthon cambeforti (Ochi, Kon & O. cambeforti Kikuta, 1997) to a position in the dytiscoides group; the combination of the shape of the head and the aedeagus, plus Figs 64-69, 127 the extensive basal and apical yellow elytral marks, = O. simboroni Ochi, Ueda & Kon, 2006 syn. n. should render this Southeast Bornean species easily recognizable. We saw the single known male speci- Diagnosis men, ca 6.5 mm long, which is kept in Paris (ex Clypeal denticles separated by emargination down collection H.W. Bates > R. Oberthur). Apart from to their base, laterally virtually straight. Clypeogenal the region of origin (Indonesian East Kalimantan) border from lateral tip to anterior denticles virtually no further details available. straight (gena shortly rounded). Clypeofrontal sur- From the same region, i.e. from a partly forested low- face densely to crowdedly punctate. Eye size moder- land area near Balikpapan, comes the single female ate, foramen rather narrowly elliptic. Pronotal bor- type on which O. simboroni is based, which, in the der anterolaterally (at ca 0.2 behind anterior angle) original description (Ochi et al. 2006), is compared shortly rounded. Paramarginal fold on posterola- with O. cambeforti, and presumably rightly so; we teral surface of pronotum short, distinct. Pronotum tentatively synonymize both species, as they appear densely to crowdedly punctate, with numerous scat- at least extremely close. The length of the female is tered microsetae, with symmetric pattern of (orange-) given as 5.1 mm. yellow and (brown-)black. Pronotal (orange-)yellow marks extensive, (orange-)yellow not predominant. 13. Ochicanthon karasuyamai Ochi, Kon & Elytra with symmetric pattern of (brown-)black Kawahara, 2007 and (orange-)yellow marks. Basal elytral (orange-) Figs 46-51 yellow B (2-7) present. Postmedial elytral (orange-) yellow M (5-7) present. Apical elytral (orange-) yellow A (2-7) present. Discal elytral interstrial Diagnosis punctation dense to crowded, interstriae shining, Clypeal denticles separated by emargination down without microreticulation, interstriae all flat or very to their base, adjoined laterally by simple concave slightly convex, interstriae with numerous scattered curve. Clypeogenal border from lateral tip to anteri- microsetae. Pygidium convex along apex, basal sur- or denticles widely convex-curvilinear. Clypeofrontal face flat or concave (in profile), brown to black, (vir- surface densely ocellate-punctate. Eye size mod- tually) uniform. Protibial denticles (male) all sim- erate, foramen narrowly elliptic. Pronotal border ply acuminate, slender. Profemur (male) anteriorly anterolaterally (at ca 0.2 behind anterior angle) more

Downloaded from Brill.com10/11/2021 05:43:19AM via free access Krikken & Huijbregts: Ochicanthon in Sundaland 449 or less abruptly rounded; anterolateral corners of pro- The original material of O. karasuyamai came from notum depressed. Paramarginal fold on posterolateral the Sibolangit area, which is ca 100 km (as the crow surface of pronotum distinct (extending into groove flies) from where the Leuser females were found. in front). Pronotum densely ocellate-punctate, gla- The description of karasuyamai more or less fits our brous, or nearly so, uniform (brown-)black. Pronotal (female) specimens; differences may be due to vari- base with shallow midline depression. Elytra with ation and/or to choices in terminology. To facilitate symmetric pattern of (brown-)black and (orange-) verification of the identity of the Leuser material a yellow marks. Basal elytral (orange-)yellow B (5-7) detailed description of one of our females is given present. Postmedial elytral (orange-)yellow M (5-7) below. The male characters in the diagnosis above are present. Apical elytral (orange-)yellow A (2-3, 5-7) based on the original description of karasuyamai. present. Discal elytral interstrial punctation ocellate, fine, abundant, intervening surface matt, due Material examined. Female figured (RMNH) has the to microreticulation, all flat or very slightly convex, following label data: with numerous scattered microsetae. Pygidium more N Sumatra: Gunung Leuser NP: Lawe Mamas, or less pointed, brown to black, (virtually) uniform, 1983, H. Räisänen, 800-820 m, #32Cb. More speci- or with vague lighter patches. Pronotal-elytral transi- men data on karasuyamai in Appendix 2. tion (in profile) gradual, habitus deplanate. Major males have the usual apico-internal angular expan- Description (female figured) sion of the metatibia. Body size varies considerably. Body length ca 6.7 mm. Colour generally matt Protibial denticles (male) all acuminate. Profemur black-brown, with yellow-orange elytral marks. (male) anteriorly lobate-dentate near apex. Metafe- Clypeal margin anteriorly bidentate (tips angular, mur (male) posteriorly lobate-dentate at 0.3-0.5 from median emargination widely rounded). Clypeo- base. Metatibia long, slender, curved, dilated near frontal surface generally feebly convex, black-brown, apex. Internal side of metatibia (male) continuous matt, glabrous, with dense, distinct but slightly (not angulate halfway), angulate-dilated at short dis- superficial, ocellate punctation; punctation less tance from apex (male). Metafemur posteriorly and dense between eyes. Clypeal border from gena for- metatibia internally not multidenticulate. Parameres ward widely rounded. Gena laterally rounded. Eyes distinctly excised, asymmetric; superior strut, and (foramen in full-face view) medium-sized, moderate- hook on bottom sclerite distinct. Pronotal-elytral ly elliptic. Maximum number of facet rows (trans- transition (in profile) gradual. Habitus deplanate. versely) across eye foramen 8-9. Ratio transverse Body size small. interocular distance/maximum eye width ca 8.0. Both sexes known. Length 5.8-8.2 mm. Pronotum generally evenly convex, disc and anterior declivity unmodified, slightly deplanate, basal half Distribution and ecology with vague midline depression. Prothorax in dorsal Sumatra. New data listed in Appendix 2. outline nearly parallel-sided, slightly convergent ros- Our four females were collected in pitfall traps bait- trad. Anterior border of pronotum unmodified. An- ed with human faeces, in one area (Lawe Mamas) in terolateral pronotal angle obtuse, abruptly rounded Mt Leuser NP, in multistratal evergreen forest. In at ca 0.2 of length behind anterior angle. Lateral this area Räisänen sampled an extensive transect sides of pronotum simply declivous. Posterolateral with pitfall traps baited either with carrion or human section of pronotum modified as in all congeners, excrement, at altitudes between 150 and 2500 m with distinct fold parallel and close to border, anteri- – our material was collected at 580-600 and 900- orly extended, sulcate. Pronotal posterolateral angles 920 m altitude, while no other Ochicanthon species shortly rounded. Base of pronotum immarginate, seemed to be present. (A manuscript name used by evenly, very widely rounded. Pronotal border finely us for karasuyamai is Phacosoma leusermontis.). The marginate in front. Pronotum with microstubbles; type locality Sibolangit is at 1200 m. entirely black-brown, matt, microreticulate, generally with fine, dense, ocellate punctation. Estimated Comments number of punctures per 0.25 sq.mm on pronotal O. karasuyamai is a North Sumatran species with a disc 45-50, punctures 0.03 mm. plump, entirely matt appearance; its pronotum is Elytra black-brown, with yellow-orange pat- completely black (or nearly so); dorsal punctation, tern, matt, microreticulate, on disc with abundant for instance on elytral interstriae, much finer than microstubbles. Pseudepipleuron reaching elytral in other species. The somewhat similar peninsularis apex (broad, pseudepipleural crest with 2 fine has a distinctly patterned pronotum, a different ridges in front). Elytral striae distinct, shallowly aedeagus, and a coarser dorsal punctation. impressed; punctures of discal elytral striae slightly

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Diagnosis impressed (crenulating interstriae, separated by 2- Clypeal denticles separated by emargination down 4 times their diameter); elytral surface with abun- to their base, adjoined laterally by simple concave dant appressed microsetae; interstriae flat to feebly curve. Clypeogenal border from lateral tip to ante- convex, with fine, abundant, scattered ocellate rior denticles widely convex-curvilinear. Clypeofron- punctation. tal surface densely punctate (punctation fine). Eye Antennal lamellae brown. size moderate, foramen moderately elliptic. Prono- Propectus laterally black-brown, matt, indistinctly tal border anterolaterally (at ca 0.2 behind anterior annulate-punctate, microreticulate. Mesosternum angle) simply rounded. Paramarginal fold on poste- virtually smooth. Metasternal anterior lobe and rolateral surface of pronotum vaguely distinct. Pro- disc densely ocellate-punctate, with very short mi- notum abundantly to densely punctate, glabrous, or crosetae; metasternum black-brown, laterally with nearly so, uniform black or brown, shining. Elytra annulate punctation (superficial, on microreticulate- uniform, entirely brown or blackish. Discal elytral microstriolate surface), abundant. Abdominal venter interstrial punctation fine, somewhat irregular, abun- black-brown, matt, densely annulate distally, with dant to dense, interstriae shining, without microre- very short microsetae, with microreticulation, matt. ticulation, all flat or very slightly convex, interstriae Hind wings fully developed. Pygidium very convex mainly with scattered microstubbles. Pygidium more (subconical), marginate along base, black-brown, or less evenly convex (in profile), brown to black, abundantly annulate-microsetose, setae very short, (virtually) uniform. Protibial denticles all simply punctures superficial. acuminate, slender. Profemur anteriorly unmodified. Profemur brown, unmodified in front (female). Metafemur posteriorly unmodified. Metatibia very Protibia externally with 3 relatively short denticles slightly curved, internally continuous (not angulate (apico-external denticle simply tapering). Proximal halfway), gradually dilated to apex. Alae present. serration on outer side of protibia distinct, fine. Pro- Parameres subsymmetric (in upper side view), short, tibia with protruding, slightly pointed apex, internal broad, tapering (lateral view), on top separated by side evenly, slightly curved. Protibial spur inserted elongate strut. Pronotal-elytral transition (in profile) away from inner angle, simply acuminate (fine). gradual, habitus deplanate. Body size small. Protibia brown. Protarsi slender. Mesofemur slender, Both sexes known. Length 3.5-4.5 mm. unmodified (female). Meso- and metafemur brown. Meso- and metatibiae long, curved, without fossorial Distribution and ecology protrusions on outer side. Metatibia strongly curved, Borneo. New data in Appendix 2. gradually dilated to apex; apex lobate-dentate. Meso- We have seen much material from lowland forest in and metatibia brown. Meso- and metatarsi slender. Sabah and Sarawak. Most specimens were collected Metatarsal segment 1 distinctly longer than seg- in pitfall traps baited with human faeces, or in flight ment 2 (approximate proportions spur//segments interception traps; smaller numbers in fish baited pit- 1-5 8//25/10/7/6/10). fall traps. At Mulu (Sarawak) O. woroae was collected Measurements in mm. Maximum width of head 2.2. at 110-150 m altitude in what was termed mixed Pronotal dorsal median length 2.1, maximum width dipterocarp forest. Davis characterizes this species in 3.5. Dorsal sutural length of elytra 3.2, maximum the Danum Valley lowland forest as an interior forest width combined 4.3. specialist. (Mentioned in Hanski 1983, as Phacosoma Colour patterns. Pronotal lighter marks absent. sp. n. E, in Davis et al. 2000, as Phacosoma paneloides Elytral lighter marks present: B (5-7), M (5-7), MS, in Davis 1999, Davis 2000, Davis et al. 2001, as A (2-7). Phacosoma sp. 2.)

Comments is a small, shining black-brown species, (b) The Ochicanthon woroae group O. woroae with subsymmetric, characteristically shaped, rela- Dorsum virtually uniformly brown-black. Parameres tively short parameres (Fig. 128); the dorsal puncta- simple, roughly symmetric, never deeply excised- tion of woroae is simple, less dense, but more distinct lobate (lateral view). than in tambunan and other uniformly black-brown Bornean species. The type locality is in Indonesian East Kalimantan, 14. Ochicanthon woroae Ochi, Ueda & Kon, 2006 not far from Balikpapan, so it seems that this species occurs all around lowland Borneo. Although the Figs 70-75, 128 populations from the various parts of Borneo appear

Downloaded from Brill.com10/11/2021 05:43:19AM via free access Krikken & Huijbregts: Ochicanthon in Sundaland 451 conspecific, we add here a detailed description of the disc abundantly to densely, finely punctate, shin- male figured in this paper, for reference purposes. ing. Metasternum dark-brown. Metasternum laterally with annulate punctation, crowded. Abdominal Material examined. Male figured (RMNH) has the venter dark-brown, matt, sparsely (proximally) to following label data: densely (distally) finely punctate(-microsetose), im- Sabah: Danum Valley, 1992, 5.01’N-117.47’E, punctate surface microstriolate. Hind wings reduced. A.J. Davis, #0660. More specimen data on woroae Pygidium very convex, marginate along base, brown, in Appendix 2. finely microsetose, crowdedly punctate. Profemur brown, unmodified. Protibia smooth, Description (male figured) externally with 3 denticles. Proximal serration on Body length 3.6 mm. Colour generally shining dark- outer side of protibia distinct, fine. Protibia apically brown. rounded, protruding, internal side evenly, slightly Clypeal margin anteriorly bidentate (tips blunted, curved. Protibial spur inserted away from inner an- median emargination widely rounded). Clypeo- gle, simply acuminate. Protibia brown. Protarsi very frontal surface generally feebly convex, dark-brown, slender. Meso- and metafemur slender, unmodified. shining, glabrous, with dense, distinct, simple punc- Meso- and metafemur brown. Meso- and metatibiae tation. Clypeal border from gena forward widely distinctly dilated distad, without fossorial protru- rounded. Gena laterally rounded. Eye size ventral- sions on outer side. Mesotibia simply, slightly curved. ly large, globose. Eyes (foramen in full-face view) Metatibia simply, slightly curved, apex lobate-dentate medium-sized, moderately elliptic. Maximum to (virtually) straight. Meso- and metatibia brown. number of facet rows (transversely) across eye fo- Meso- and metatarsi slender. Metatarsal segment 1 ramen 8. Ratio transverse interocular distance/ distinctly longer than segment 2 (approximate pro- maximum eye width ca 8.6. portions spur//segments 1-5 9//15/7/6/4/7). Pronotum generally evenly convex, disc and anterior Aedeagus more or less symmetrical, as in Fig. 128. declivity unmodified, slightly deplanate. Protho- Measurements in mm. Maximum width of head 1.3. rax in dorsal outline nearly parallel-sided, slightly Pronotal dorsal median length 1.2, maximum width convergent rostrad. Anterior border of pronotum 2.1. Dorsal sutural length of elytra 2.0, maximum unmodified. Anterolateral pronotal angle rounded. width combined 2.5. Lateral sides of pronotum simply declivous. Poste- rolateral section of pronotum modified as in all con- 15. Ochicanthon tambunan sp.n. geners, with vague fold parallel and close to border. Figs 76-81, 129 Pronotal posterolateral angles simple, distinct, obtuse. Base of pronotum immarginate, evenly, very Type-material. Male holotype (NMWC): Malay- widely rounded, with surface unmodified. Pronotal sia, with our type label, and the following label sides finely marginate, simple. Pronotal border finely data: Sabah: Crocker Range: 5.50’N-116.00’E, marginate in front. Pronotum glabrous, generally 7-11/viii/1991, A.H. Kirk-Spriggs, 1280 m, 2 pitfall with simple, dense punctation. Estimated number traps baited with dung, #ks084. of punctures per 0.25 sq.mm on pronotal disc 164 paratypes listed in Appendix 2. 60-65, punctures ca 0.01 mm. Pronotum dark- brown, generally shining. Description (holotype, male) Elytra entirely dark-brown, generally shining, vir- Body length ca 3.7 mm. Colour generally brown, tually glabrous at x50. Pseudepipleuron reach- matt. ing elytral apex (broad, pseudepipleural crest with Clypeal margin anteriorly bidentate (tips acute, me- fine ridge in front, stria 7 bordered by fine ridge). dian emargination widely rounded). Clypeofrontal Elytral striae distinct, shallowly impressed; punc- surface generally feebly convex, brown, moderately tures of discal striae slightly impressed (crenulating shining, glabrous, with dense to crowded, distinct, interstriae, separated by 3-5 diameters); interstriae fine, simple punctation. Clypeal border from gena flat, simply punctate (pseudepipleuron coarsely forward widely rounded. Gena laterally rounded. rugulate-punctate, shining); punctures fine, scat- Eyes small, foramen (in full-face view) medium- tered, abundant. sized, moderately elliptic. Maximum number of Antennal lamellae brown. facet rows (transversely) across eye foramen 7-8. Propectus laterally brown, glabrous, moderately Ratio transverse interocular distance/maximum eye shining, virtually impunctate. Mesosternum shin- width 7.5. ing, virtually impunctate. Metasternal anterior lobe Pronotum generally evenly convex, disc and anterior abundantly, finely punctate, shining. Metasternal declivity unmodified, slightly deplanate. Prothorax in

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70 76 74

71 77 80

72 78

73 79 75

Figs 70-81. Ochicanthon, habitus and body parts. – 70-75, woroae (male), habitus, oblique (70); head, full-face (71); pronotum, dorsal (72); left elytron, dorsal (73); protibia, upper side (74); metatibia and -femur, underside (75); 76-81, O. tambunan (male holotype), habitus, oblique (76); head, full-face (77); pronotum, dorsal (78); left elytron, dorsal (79); protibia, upper side (80); metatibia and -femur, underside (81).

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82 88 86

83 89 92

84 90

85 91 87

Figs 82-93. Ochicanthon, habitus and body parts. – 82-87, O. mulu, habitus (male holotype), oblique (82); head, full-face (83); pronotum, dorsal (84); left elytron, dorsal (85); protibia, upper side (86); metatibia and -femur, underside (87); 88-93, O. gangkui (male), habitus, oblique (88); head, full-face (89); pronotum, dorsal (90); left elytron, dorsal (91); protibia, upper side (92); metatibia and -femur, underside (93).

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dorsal outline nearly parallel-sided, slightly conver- longer than segment 2 (approximate proportions gent rostrad. Anterior border of pronotum unmodi- spur//segments 1-5 7//13/6/5/4/6). fied. Anterolateral pronotal angle rounded, abruptly Aedeagus more or less symmetric, as in Fig. 129. rounded at ca 0.2 behind anterior angle. Lateral sides Measurements in mm. Maximum width of head 1.3. of pronotum simply declivous. Posterolateral section Pronotal dorsal median length 1.3, maximum width of pronotum modified as in all congeners, with in- 2.0. Dorsal sutural length of elytra 1.7, maximum distinct fold parallel and close to border. Pronotal width combined 2.4. posterolateral angles simple, distinct, obtuse. Base of pronotum immarginate, evenly very widely round- Variation and sexual dimorphism ed, with surface unmodified. Pronotal sides finely Slight, quantitative. marginate, simple. Pronotal border scarcely marginate in front. Pronotum microsetose, setae long, ap- Derivation of species name pressed. Pronotum generally with simple, crowded Named after the type locality, i.e. one of the localities punctation. Estimated number of punctures per in the Crocker Range region. 0.25 sq.mm on pronotal disc 80, punctures 0.01 mm. Pronotum brown, generally matt, interpunc- Diagnosis tural spaces very narrow. Clypeal denticles separated by emargination down Elytra entirely brown, generally matt, with long, to their base, adjoined laterally by simple concave fine appressed microsetae in rugulate punctation. curve. Clypeogenal border from lateral tip to anteri- Pseudepipleuron reaching elytral apex (broad, or denticles widely convex-curvilinear. Clypeofrontal pseudepipleural crest with fine ridge in front, stria surface crowdedly punctate. Eye size small, foramen 7 bordered by fine ridge). Punctures of discal elytral small, elliptic. Pronotal border anterolaterally (at ca striae slightly impressed (crenulating interstriae, but 0.2 behind anterior angle) abruptly rounded. Para- partly lost in more or less irregular surface, separated marginal fold on posterolateral surface of pronotum by 1-2 diameters). Elytral striae distinct, shallowly vaguely distinct. Pronotum crowdedly punctate, al- impressed (somewhat indistinct in irregular sur- most rugulate, with numerous scattered microsetae, face). Elytra with abundant microsetae. Interstriae uniform (brown-)black, matt due to punctation. flat, crowdedly, simply punctate, somewhat rugulate Elytra uniform, entirely brown or black. Discal ely- (pseudepipleuron coarsely rugulate-punctate, slight- tral interstrial punctation crowded, superficially rug- ly shining). ulate, shining. Interstrial surfaces flat or very slightly Antennal lamellae brown. convex, with abundant, scattered, short microse- Propectus laterally brown, glabrous, shining, sparsely tae. Pygidium more or less evenly convex (in pro- punctate. Mesosternum shining, posteriorly finely file), brown to black, (virtually) uniform. Protibial punctate. Metasternal anterior lobe and disc irregu- denticles) all simply acuminate, slender. Profemur larly, abundantly, finely punctate-microsetose, later- anteriorly unmodified. Metafemur posteriorly un- ally with annulate punctation, crowded. Abdominal modified. Metatibia straight to very slightly curved, venter brown, rather matt, finely, densely punc- distal half dilated. Metatibia internally continuous tate-microsetose, proximal sternites with microstri- (not angulate halfway), gradually dilated to apex. olate posterior margin. Hind wings fully developed. Metafemur posteriorly and metatibia internally not Pygidium slightly convex, marginate along base, multidenticulate. Alae present. Parameres subsym- brown, finely microsetose, crowdedly punctate. metric (in upper side view), short, broad, tapering Profemur brown, unmodified. Protibia smooth, (lateral view). Pronotal-elytral transition (in profile) externally with 3 denticles. Proximal serration on gradual, habitus deplanate. Body size small. outer side of protibia distinct, fine. Protibia apically Both sexes known. Length 3.5-4 mm. rounded, protruding, internal side evenly, slightly curved. Protibial spur inserted away from inner an- Distribution and ecology gle, simply acuminate, long, acute). Protibia light Borneo: Sabah. brown. Protarsi very slender. Meso- and metafemur All specimens were collected in upland forest in the slender, unmodified. Meso- and metafemur brown. Crocker Range, in baited pitfall traps, set between Meso- and metatibiae distinctly dilated distad, with- ca 1250 and 1465 m altitude. The species seems to out fossorial protrusions on outer side. Mesotibia prefer human excrement over fish. simply, slightly curved. Metatibia simply, slightly curved; apex lobate-dentate to (virtually) straight. Comments Meso- and metatibia brown. Mesotarsi slender. O. tambunan appears a small, all black-brown Metatarsi slender. Metatarsal segment 1 distinctly upland species with somewhat rugulate elytra,

Downloaded from Brill.com10/11/2021 05:43:19AM via free access Krikken & Huijbregts: Ochicanthon in Sundaland 455 superficially similar to O. gangkui and its relatives, rugulate-punctate). Elytral striae superficial; punc- but with a very different, subsymmetric aedeagus tures of discal striae slightly impressed (strongly (Fig. 129), much shorter than in woroae. crenulating interstriae, separated by 1-2 times their diameter); interstriae flat, annulate-punctate to rugulate; interstrial punctures arranged in rows (less so 16. Ochicanthon mulu sp.n. on basodiscal surface), more or less crowded. Figs 82-87, 130 Antennal lamellae brown. Type-material. Male holotype (RMNH): Malaysia, Propectus laterally brown, crowdedly annulate-punc- with our type label, and the following label data: tate-microsetose. Mesosternum shining, sparsely an- Sarawak: 4th Division: Gunung Mulu NP, iii-v/1978, nulate-punctate. Metasternal anterior lobe and disc I. Hanski, ca 150 m, baited pitfall trap, kerangas, densely, finely annulate-punctate, with very fine #224. setae, black-brown. laterally with dense annulate 11 paratypes listed in Appendix 2. punctation. Abdominal venter brown, matt, entirely finely annulate-punctate (distal sternites) or along Description (holotype, male) base (proximal sternites), punctures microsetose, Body length ca 4.0 mm. Colour generally brownish, impunctate parts microstriolate. Hind wings fully matt. developed. Pygidium strongly convex, marginate Clypeal margin anteriorly bidentate (tips acute, lat- along base, brown, with fine setae, densely annulate- erally delimited by distinct sinuate curve, median punctate. emargination widely rounded). Clypeofrontal sur- Profemur brown, unmodified. Protibia smooth, with face generally feebly convex, brown, generally matt, 3 acuminate but relatively short external denticles. glabrous, with crowded, distinct, annulate puncta- Proximal serration on outer side of protibia distinct, tion. Clypeal border from gena forward virtually fine. Protibia apically rounded, slightly protrud- straight. Clypeogenal margin (at suture) slightly dis- ing, internal side evenly, slightly curved. Protibial continuous. Genae laterally rounded to subangulate, spur inserted away from inner angle, simply acu- with shallowly concave surface. Eye size ventrally minate (long, acute). Protibia light brown. Protarsi large, globose. Eyes (foramen in full-face view) very very slender, short. Mesofemur slender, unmodi- large, elliptic-circular, short, widely open behind. fied; metafemur slender, posterior side unmodified. Maximum number of facet rows (transversely) across Meso- and metafemur brown. Meso- and metatibiae eye foramen ca 12. Ratio transverse interocular dis- distinctly dilated distad. Mesotibia simply, slightly tance/maximum eye width ca 3.4. curved. Metatibia nearly straight, slightly, gradually Pronotum generally evenly convex, disc and anterior dilated distad (lacking internal hook); apex lobate- declivity unmodified. Prothorax in dorsal outline dentate to (virtually) straight. Meso- and metatibia nearly parallel-sided, slightly convergent rostrad. brown. Meso- and metatarsi relatively compact and Anterior border of pronotum unmodified. Anterola- complanate. Metatarsal segment 1 distinctly longer teral pronotal angle rounded. Anterolateral section of than segment 2 (approximate proportions spur// pronotum not depressed. Lateral sides of pronotum segments 1-5 7//11/5/4/4/5). simply declivous. Posterolateral section of pronotum Aedeagus more or less symmetric, as in Fig. 130. modified as in all congeners, with fold parallel and Measurements in mm. Maximum width of head 1.3. close to border. Pronotal posterolateral angles sim- Pronotal dorsal median length 1.3, maximum width ple, distinct, obtuse. Base of pronotum immarginate, 2.0. Dorsal sutural length of elytra 1.8, maximum evenly, very widely rounded, with surface unmodi- width combined 2.4. fied. Pronotal sides finely marginate, ridge obsolescent (or absent) posteriorly. Pronotal border scarcely Variation and sexual dimorphism marginate in front. Pronotum brown, generally matt, Slight, quantitative. Colour may be rufous brown, microsetose, setae long (short recurved pale bristles, also dependent on lighting length about twice punctural diameter). Pronotum generally with crowded annulate punctation. Esti- Derivation of species name mated number of punctures per 0.25 sq.mm on pro- Named after the type locality, a National Park in notal disc ca 80, punctures ca 0.01 mm. Sarawak. Elytra entirely brown, moderately shining; with erect, apically more or less recurved pale bristles (dis- Diagnosis tally in 2 rows). Humeral umbone distinct. Pseude- Clypeal denticles separated by emargination down pipleuron reaching elytral apex (broad, very coarsely to their base, adjoined laterally by shallow (sinuate)

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emargination. Clypeogenal border from lateral Diagnosis tip anteriorly widely convex-curvilinear. Clypeo- Clypeal denticles separated by emargination down frontal blackish-brown surface densely to crowdedly to their base, adjoined laterally by simple concave annulate-punctate. Eye size large, foramen broadly curve. Clypeogenal border from lateral tip to an- elliptic. Pronotal border anterolaterally (at ca 0.2 be- terior denticles widely convex-curvilinear. Clypeo- hind anterior angle) simply rounded. Paramarginal frontal surface crowdedly punctate. Eye size large, fold on posterolateral surface of pronotum moder- foramen moderately elliptic. Pronotal border ante- ately distinct, extending into vague groove. Prono- rolaterally (at ca 0.2 behind anterior angle) abruptly tum crowdedly annulate-punctate, with numer- rounded. Paramarginal fold on posterolateral surface ous scattered microsetae, uniform blackish-brown. of pronotum short, moderately distinct. Pronotum Elytra uniform, entirely blackish-brown. Discal crowdedly punctate, glabrous, or nearly so, uniform elytral interstrial punctation dense to crowded, an- (brown-)black. Elytra uniform, entirely brown or nulate; interstriae moderately shining, all flat or very black. Discal elytral interstrial punctation crowded slightly convex, interstriae abundantly, distinctly se- to almost rugulate, interstriae slightly matt, due to tose (setae more or less in rows). Pygidium more or crowded microsculpture; interstriae all flat or very less evenly convex (in profile), brown to black, (vir- slightly convex, with numerous scattered microsetae. tually) uniform (strongly convex). Protibial denticles Pygidium slightly evenly convex (in profile), brown all simply acuminate, slender. Profemur anteriorly to black, uniform. Protibial denticles all simply unmodified. Metafemur posteriorly unmodified. acuminate, slender. Profemur anteriorly unmodi- Metatibia straight to very slightly curved, internally fied. Metafemur posteriorly unmodified. Metatibia continuous (not angulate halfway), gradually di- straight to very slightly curved, dilated distad, in- lated to apex. Meso- and metatarsi short, relatively ternal side continuous (not angulate halfway). Alae compact. Alae present. Parameres subsymmetric (in present. Parameres distinctly excised, asymmetric, upper side view), elongate, broad, tapering (lateral complex. Left paramere very deeply excised, infe- view), separated on top by elongate strut. Pronotal- rior branch elongate-expanded. Right paramere elytral transition (in profile) gradual, habitus de- sinuate, lobate at base, inferior branch also elongate- planate. Body size small. expanded (lateral view); strut topping aedeagus dis- Both sexes known. Length 3.5-4.5 mm. tinct. Pronotal-elytral transition (in profile) gradual, habitus deplanate. Punctures of dorsum matt. Body Distribution and ecology size small. Borneo: Sarawak. Both sexes known. Length 4-4.5 mm. All specimens were collected by Hanski (1983, as Phacosoma sp.n. D) using pitfall traps baited with ei- Distribution and ecology ther fish or human excrement, at Mulu, in kerangas Borneo: Sabah. New data listed in Appendix 2. [heath forest] at an altitude of 150 m. Known from moist forest on Mt Kinabalu and Mt Trus Madi, at altitudes between 1200 and Comments 1550 m. Specimens were collected in pitfall traps O.mulu is a small, all rufous to blackish-brown spe- baited with either fish or human faeces. cies characterized by large eye foramina, clypeal denticles delimited by a lateral emargination, coarsely Comments annulate-setose dorsum, and short, tapering, sub- O. gangkui is an all black-brown, more or less matt symmetric parameres (Fig. 130). species, best recognizable from the structural details of its asymmetric parameres (see diagnosis above and Fig. 132). The series we collected comes from the same area as the types, not far from the Mt Kinabalu (c) The Ochicanthon gangkui group NP headquarters. Dorsum virtually uniformly brown-black. Parameres (lateral view) more or less deeply, usually asymmetri- 18. Ochicanthon kimanis sp.n. cally excised, complex. Figs 94-99, 133 Type-material. Male holotype (RMNH): Ma- 17. Ochicanthon gangkui (Ochi, Kon & Kikuta, 1997) laysia, with our type label, and the following label data: Sabah: Crocker Range: Keningau- Figs 88-93, 132 Kimanis road (km25), 18-23/xi/1987, J. Krikken & E. Rombaut, 1300 m, multistr evergr forest,

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8 human excr traps, #sa45. base, brown, glabrous, crowdedly punctate. 10 paratypes listed in Appendix 2. Profemur brown, unmodified. Protibia externally with 3 denticles. Proximal serration on outer side Description (holotype, male) of protibia very fine. Protibia apically rounded, Body length ca 4.3 mm. Colour generally dark- protruding, internal side evenly, slightly curved. brown. Protibial spur inserted away from inner angle, sim- Clypeal margin anteriorly bidentate (tips acute, me- ply acuminate (long, acute). Protibia light brown. dian emargination widely rounded). Clypeofrontal Protarsi very slender. Meso- and metafemur slender, surface generally feebly convex, dark-brown, moder- unmodified, brown. Meso- and metatibiae distinctly ately shining, glabrous, with crowded, distinct, sim- dilated distad, without fossorial protrusions on outer ple punctation. Clypeal border from gena forward side. Meso- and metatibia simply, slightly curved. widely rounded. Gena laterally rounded. Eye size Metatibial apex lobate-dentate. Meso- and metatibia ventrally large, globose. Eyes (foramen in full-face brown. Meso- and metatarsi slender. Metatarsal seg- view) large, broadly elliptic. Maximum number of ment 1 distinctly longer than segment 2 (approximate facet rows (transversely) across eye foramen ca 11. proportions spur//segments 1-5 7//14/6/5/4/7). Ratio transverse interocular distance/maximum eye Aedeagus asymmetrical, as in Fig. 133. width 4.1. Measurements in mm. Maximum width of head 1.4. Pronotum generally evenly convex, disc and anterior Pronotal dorsal median length 1.2, maximum declivity unmodified, slightly deplanate. Protho- width 2.1. Dorsal sutural length of elytra 2.3, maxi- rax in dorsal outline nearly parallel-sided, slightly mum width combined 2.5. convergent rostrad. Anterior border of pronotum unmodified. Anterolateral pronotal angle rounded. Variation and sexual dimorphism Lateral sides of pronotum simply declivous. Postero- Slight, quantitative. lateral section of pronotum modified as in all congeners, with distinct fold parallel and close to border. Derivation of species name Pronotal posterolateral angles simple, distinct. Base Named after the type locality, i.e. one of the localities of pronotum immarginate, evenly rounded (very in the Crocker Range region. widely so), with surface unmodified. Pronotal sides finely marginate, simple. Pronotal border uninter- Diagnosis rupted (very finely marginate in front). Pronotum Clypeal denticles separated by emargination down virtually glabrous. Pronotum generally with simple, to their base, adjoined laterally by simple concave crowded punctation. Estimated number of punctures curve. Clypeogenal border from lateral tip to an- per 0.25 sq.mm on pronotal disc ca 100, punctures terior denticles widely convex-curvilinear. Clypeo- 0.01 mm. Pronotum dark-brown, generally matt frontal surface densely to crowdedly punctate. Eye due to crowded punctation, narrow interpunctural size large, foramen broadly elliptic. Pronotal bor- surfaces shining. der anterolaterally (at ca 0.2 behind anterior angle) Elytra entirely dark-brown, generally matt, with short simply rounded. Paramarginal fold on posterolateral microsetae, in slightly rugulate punctation. Pseude- surface of pronotum short, moderately distinct. Pro- pipleuron reaching elytral apex (broad, pseudepip- notum crowdedly punctate, glabrous, or nearly so, leural crest with 2 fine ridges in front). Punctures uniform (brown-)black. Elytra uniform, entirely of discal elytral striae slightly impressed (crenulating brown or black. Discal elytral interstrial puncta- interstriae, but partly lost in more or less rugulate tion crowded to almost rugulate, interstriae slightly surface, separated by 1-2 diameters). Elytral striae matt, due to rugulate microsculpture, interstriae all mostly distinct, shallowly impressed. Interstriae flat flat or very slightly convex, with numerous scattered or very slightly convex. Pseudepipleuron rugulate- microsetae. Pygidium more or less evenly convex (in punctate. profile), brown to black, uniform. Protibia smooth, Antennal lamellae brown. denticles all simply acuminate, slender. Profemur Propectus laterally brown, glabrous, shining, sparsely anteriorly unmodified. Metafemur posteriorly un- punctate. Mesosternum shining, posteriorly finely modified. Metatibia straight to very slightly curved, punctate. Metasternal anterior lobe and disc irregu- distal half dilated. Metatibia internally continuous larly, abundantly, finely punctate, glabrous, brown, internally (not angulate halfway), gradually dilated laterally with dense annulate punctation. Abdominal to apex. Alae present. Parameres distinctly excised, venter brown, matt, virtually glabrous, irregularly asymmetric, complex. Left paramere very deeply punctate, microstriolate. Hind wings fully devel- excised, lobate inferior branch elongate-expanded. oped. Pygidium slightly convex, marginate along Right paramere also deeply, but more widely

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excised, lacking lobe formed by sinus; inferior branch marginate in front. Pronotum microsetose, setae long elongate-expanded. Strut topping aedeagus distinct. (with micro-stubbles, scarcely distinct at 50). Pro- Pronotal-elytral transition (in profile) gradual, habi- notum generally with simple, crowded punctation. tus deplanate. Body size small. Estimated number of punctures per 0.25 sq.mm on Both sexes known. Length 4-4.5 mm. pronotal disc 80-90, punctures 0.03 mm. Pronotum dark-brown, generally shining. Distribution and ecology Elytra entirely dark-brown, generally shining, with Borneo: Sabah. microstubbles in crowded, somewhat rugulate punc- All specimens were collected in upland forest in the tation, setae laterally longer. Pseudepipleuron reach- Crocker Range, in human faeces baited pitfall traps, ing elytral apex (broad, pseudepipleural crest with at 1300 m altitude (no carrion traps were set in the 2 fine ridges in front). Punctures of discal elytral area). striae slightly impressed (crenulating interstriae, but indistinct in more or less rugulate surface, separat- Comments ed by 1-3 diameters); striae distinct, shallowly im- O. kimanis is an all black-brown, somewhat matt pressed; interstriae with crowded, simple punctation species, found in an area more than a hundred km and abundant microsetae; surface flat; pseudepipleu- SW of Mount Kinabalu; it is best to be recognized ron coarsely rugulate-punctate, somewhat shining. from the structural details of its asymmetric para- Antennal lamellae brown. meres (see diagnosis above and Fig. 133). Propectus laterally brown, glabrous, shining, sparsely punctate. Mesosternum shining, scarcely punctate. Metasternal anterior lobe and disc abundantly, finely 19. Ochicanthon danum sp.n. punctate, shining, dark-brown, laterally with annu- Figs 100-105, 134 late, crowded punctation. Abdominal venter dark- Type-material. Male holotype (RMNH): Malaysia, brown, shining, finely, densely punctate, proximal with our type label, and the following label data: sternites with microstriolate posterior margin. Hind Sabah: Danum Valley: 5.01’N-117.47’E, 1991, wings fully developed. Pygidium moderately, evenly A.J. Davis, #0787. convex, marginate along base, brown, finely micro- 131 paratypes listed in Appendix 2. setose, crowdedly punctate. Profemur brown, unmodified. Protibia smooth, Description (holotype, male) externally with 3 denticles. Proximal serration on Body length ca 3.5 mm. Colour generally dark- outer side of protibia distinct, fine. Protibia api- brown. cally rounded, protruding, with internal side evenly, Clypeal margin anteriorly bidentate (tips acute, slightly curved. Protibial spur inserted away from in- median emargination widely rounded). Clypeo- ner angle, simply acuminate (long, acute). Protibia frontal surface generally feebly convex, dark-brown, brown. Protarsi very slender. Meso- and metafemur shining, glabrous, with crowded, distinct, fine, sim- slender, unmodified, brown. Meso- and metatibiae ple punctation. Clypeal border from gena forward distinctly dilated distad, without fossorial protrusions widely rounded. Gena laterally rounded. Eye size on outer side. Meso- and metatibia simply, slightly ventrally large, globose. Eyes (foramen in full-face curved, apex lobate-dentate to (virtually) straight. view) large, broadly elliptic. Maximum number of Meso- and metatibia brown. Meso- and metatarsi facet rows (transversely) across eye foramen 12-13. slender. Metatarsal segment 1 distinctly longer than Ratio transverse interocular distance/maximum eye segment 2 (approximate proportions spur//segments width ca 3.1. 1-5 7//13/6/5/4/7). Pronotum generally evenly convex, disc and anterior Aedeagus asymmetrical, as in Fig. 134. declivity unmodified, slightly deplanate. Prothorax in Measurements in mm. Maximum width of head 1.3. dorsal outline nearly parallel-sided, slightly conver- Pronotal dorsal median length 1.1, maximum width gent rostrad. Anterior border of pronotum unmodi- 1.8. Dorsal sutural length of elytra 1.8, maximum fied. Anterolateral pronotal angle rounded. Lateral width combined 2.3. sides of pronotum simply declivous. Posterolateral section of pronotum modified as in all congeners, Variation and sexual dimorphism with vague fold parallel and close to border. Pro- Slight, quantitative. A male from Mount Danum, notal posterolateral angles simple, distinct, obtuse. higher than the other sites (nearly 1100 m), has Base of pronotum immarginate, evenly, very widely slightly different genitalia, and is excluded from the rounded, with surface unmodified. Pronotal sides type-series. finely marginate, simple. Pronotal border scarcely

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20. Ochicanthon parantisae (Ochi, Kon & Derivation of species name Kikuta, 1997) Named after the type locality, Danum Valley, a well- known forest ecology study site. Diagnosis Diagnosis Clypeal denticles separated by emargination down Clypeal denticles separated by emargination down to their base, adjoined laterally by simple concave to their base, adjoined laterally by simple concave curve. Clypeogenal border from lateral tip to ante- curve. Clypeogenal border from lateral tip to anteri- rior denticles widely convex-curvilinear. Clypeofron- or denticles widely convex-curvilinear. Clypeofrontal tal surface densely to crowdedly punctate. Eye size surface crowdedly punctate. Eye size large, foramen large, foramen broadly elliptic. Pronotal border an- broadly elliptic. Pronotal border anterolaterally (at terolaterally (at ca 0.2 behind anterior angle) more or ca 0.2 behind anterior angle) simply rounded. Para- less abruptly rounded. Paramarginal fold on poste- marginal fold on posterolateral surface of pronotum rolateral surface of pronotum short, moderately dis- short, vaguely distinct. Pronotum crowdedly punc- tinct. Pronotum densely to crowdedly punctate, with tate, glabrous, or nearly so, uniform (brown-)black. numerous scattered microsetae, uniform (brown-) Elytra uniform, entirely brown or black. Discal ely- black. Elytra uniform, entirely brown or black. Dis- tral interstrial punctation dense to crowded, may be cal elytral interstrial punctation (asperate-)rugulate, slightly rugulate, interstriae shining, interstriae all interstriae shining, without microreticulation, inter- flat or very slightly convex, with numerous scattered striae all flat or very slightly convex, interstriae with microsetae. Pygidium more or less evenly convex (in numerous scattered microsetae. Pygidium more or profile), brown to black, uniform. Protibial denticles less evenly convex (in profile), brown to black, uni- all simply acuminate, slender. Profemur anteriorly form. Protibial denticles all simply acuminate, slen- unmodified. Metafemur posteriorly unmodified. der. Profemur anteriorly unmodified. Metafemur Metatibia straight to very slightly curved, internally posteriorly unmodified. Metatibia straight to very continuous (not angulate halfway), gradually dilated slightly curved, distal half dilated. Metatibia inter- to apex. Parameres distinctly excised, asymmetric, nally continuous internally (not angulate halfway), complex. Left paramere more moderately excised, gradually dilated to apex. Meso- and metatarsi short, inferior branch short, right paramere widely sinu- relatively compact. Alae present. Parameres distinctly ate, strut topping aedeagus distinct. Pronotal-elytral excised, complex, excision of either rounded, with transition (in profile) gradual, habitus deplanate. short, non-lobiform superior branch. Pronotal- Body size small. elytral transition (in profile) gradual, habitus de- Both sexes known. Length 4.2 mm. planate. Body size small. Both sexes known. Length 4.5-5 mm. Distribution and ecology Borneo: Sabah. Distribution and ecology Virtually all specimens were collected in lowland for- Borneo: Sabah. est at Danum Valley (eastern Sabah) at an altitude Recorded from altitudes of 1200-1400 m. between 100 and 250 m. This species was almost exclusively collected in human faeces baited pitfall Comments traps. Extensive deployment of flight interception This Ochicanthon is reputedly related to the ques- traps yielded only one specimen, carrion baited traps tionable punctatus (see below); at any rate its ae- did not yield any specimens. Davis characterizes this deagus is different from other known males in the species in Danum Valley as a riverine/edge specialist. gangkui group, particularly by the evenly rounded Mentioned in Davis et al. 2000, as Phacosoma danum excision, without invading lobes, between the short MS, in Davis 1999; Davis 2000, Davis et al. 2001, inferior and superior parameral branches (on either as Phacosoma sp. 1. side) – see Figs 2-4 in Ochi, Kon & Kikuta (1997).

Comments 21. Ochicanthon javanus sp.n. O. danum is an all black-brown species, best to be Figs 106-111, 135 recognized from the structural details of its asymmetric parameres (see diagnosis above and Fig. 134). Type-material. Male holotype (RMNH): Indonesia, with our type label, and the following label data: W Java: Mt Puncak: Telaga Warna, 27-30/xii/1985, J. Krikken, 1600 m, multistr evergr forest (degraded), 2 fish traps, # pw71b.

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94 100 98

95 101

104

96 102

97 103 99

105

Figs 94-105. Ochicanthon, habitus (male holotypes) and body parts. – 94-99, O. kimanis, habitus (94), oblique; head, full-face (95); pronotum, dorsal (96); left elytron, dorsal (97); protibia, upper side (98); metatibia and -femur, underside (99); 100-105, O. danum, habitus (100), oblique; head, full-face (101); pronotum, dorsal (102); left elytron, dorsal (103); protibia, upper side (104); metatibia and -femur, underside (105).

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106 112 110

107 113

116

108 114

109 115 111

117

Figs 106-117. Ochicanthon, habitus (male holotypes) and body parts. – 106-111, O. javanus, habitus, oblique (106); head, full-face (107); pronotum, dorsal (108); left elytron, dorsal (109); protibia, upper side (110); metatibia and -femur, underside (111); 112-117, O. hanskii, habitus, oblique (112); head, full-face (113); pronotum, dorsal (114); left elytron, dorsal (115); protibia, upper side (116); metatibia and -femur, underside (117).

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Description (holotype, male) outer side of protibia distinct, fine. Protibia apically Body length ca 3.5 mm. rounded, distinctly protruding, internal side evenly, Clypeal margin anteriorly bidentate (tips acute, me- slightly curved. Protibial spur inserted away from indian emargination widely rounded). Clypeofrontal ner angle, simply acuminate. Protibia brown. Protarsi surface generally feebly convex, black, moderately very slender. Meso- and metafemur slender, unmodi- shining, with microstubbles, in crowded, distinct, fied, brown. Meso- and metatibiae distinctly dilated simple punctation. Clypeal border from gena for- distad, without fossorial protrusions on outer side. ward widely rounded. Gena laterally rounded. Eye Meso- and metatibia simply, slightly curved, brown; size ventrally small; eyes (foramen in full-face view) metatibial apex lobate-dentate. Meso- and metatarsi moderately elliptic. Maximum number of facet slender. Metatarsal segment 1 distinctly longer than rows (transversely) across eye foramen ca 7. Ratio segment 2 (approximate proportions spur//segments transverse interocular distance/maximum eye width 1-5 3//12/6/5/5/6). ca 7.5. Aedeagus asymmetrical, as in Fig. 135. Pronotum generally evenly convex, disc and anterior Measurements in mm. Maximum width of head 1.3. declivity unmodified, slightly deplanate. Protho- Pronotal dorsal median length 1.1, maximum width rax in dorsal outline nearly parallel-sided, slightly 2.0. Dorsal sutural length of elytra 1.7, maximum convergent rostrad. Anterior border of pronotum width combined 2.3. unmodified. Anterolateral pronotal angle obtuse to rounded. Lateral sides of pronotum simply declivous. Variation and sexual dimorphism Posterolateral section of pronotum modified as in all Variation quantitative, mainly evident in punctural congeners, with short fold parallel and close to bor- sizes and densities, and in body length and colour, der. Pronotal posterolateral angles simple, distinct, possibly also in the shape of the eye foramen. There obtuse. Base of pronotum immarginate, evenly, very may be subspecific differences between populations widely rounded, with surface unmodified. Prono- on Java (for instance, West versus East), but for a tal sides finely marginate, simple. Pronotal border proper assessment more material is required – speci- very finely marginate in front. Pronotum virtually mens from East Java excluded from type-series. glabrous, generally crowdedly punctate, somewhat rugulate entirely black, generally shining. Estimated Derivation of species name number of punctures per 0.25 sq.mm on pronotal Named after its home island, Java. disc 60-65, punctures 0.04 mm. Elytra entirely black, generally slightly shining, with Diagnosis microstubbles and microsetae in abundant fine punc- Clypeal denticles separated by emargination down tation in rugulate surface. Pseudepipleuron reaching to their base, adjoined laterally by simple concave elytral apex (broad, pseudepipleural crest with 2 curve. Clypeogenal border from lateral tip to ante- fine ridges in front). Elytral striae largely distinct, rior denticles widely convex-curvilinear. Clypeofron- superficially impressed; punctures of discal striae tal surface densely to crowdedly punctate. Eye size also superficially impressed (crenulating interstriae, large, foramen moderately elliptic. Pronotal border basomedian surface, separated by 1-2 diameters); anterolaterally (at ca 0.2 behind anterior angle) sim- interstriae flat, rugulate-punctate (pseudepipleuron ply rounded. Paramarginal fold on posterolateral slightly rugulate, shining). surface of pronotum short, vaguely distinct. Pro- Antennal lamellae brown. notum densely to crowdedly punctate, glabrous, or Propectus laterally (brown-)black, virtually glabrous, nearly so, uniform (brown-)black, shining. Elytra irregularly, abundantly punctate. Mesosternum uniform, entirely brown or black. Discal elytral in- largely smooth. Metasternal anterior lobe and disc terstrial punctation simple to rugulate, interstriae irregularly, abundantly, coarsely punctate, virtually shining, without microreticulation, all flat or very glabrous. Metasternum (brown-)black, laterally slightly convex, mainly with scattered microstubbles. with fine, dense annulate punctation. Abdomi- Pygidium more or less evenly convex (in profile) nal venter (brown-)black, matt, virtually glabrous, (near apex), brown to black, uniform. Protibial den- vaguely microstriolate, distal sternites finely, densely ticles all simply acuminate, slender. Profemur anteri- punctate-microsetose. Hind wings fully developed. orly unmodified. Metafemur posteriorly unmodified. Pygidium very convex (subconical), marginate Metatibia straight to very slightly curved, internally along base, black, rugulate-punctate, with short continuous (not angulate halfway), gradually dilated microsetae. to apex. Alae present. Parameres distinctly excised, Profemur brown-black, unmodified. Protibia ex- asymmetric, complex. Left paramere very deeply ternally with 3 denticles. Proximal serration on excised, inferior branch elongate-expanded; right

Downloaded from Brill.com10/11/2021 05:43:19AM via free access Krikken & Huijbregts: Ochicanthon in Sundaland 463 paramere deeply excised, lobate; strut topping aedea- Ratio transverse interocular distance/maximum eye gus distinct. Pronotal-elytral transition (in profile) width ca 10.5. gradual, habitus deplanate. Body size small. Pronotum generally evenly convex, disc and ante- Both sexes known. Length 3.5-4.5 mm. rior declivity unmodified. Prothorax in dorsal outline nearly parallel-sided, in lateral outline showing Distribution and ecology pronotal-elytral dip. Anterior border of pronotum Java. unmodified. Anterolateral pronotal angle widely Most specimens were collected in moist upland for- rounded. Anterolateral section of pronotum distinct- est (and forest remnants) on the slopes of Java’s vol- ly depressed, particularly behind eyes. Lateral sides canoes, at altitudes of 1200-1600 m, by baited pitfall of pronotum simply declivous (apart from anterior trapping with fish as well as human faeces. depression). Posterolateral section of pronotum with fold parallel to border indistinct. Pronotal postero- Comments lateral angles simple, distinct, obtuse. Base of pro- O. javanus is a small, variable, all black-brown spe- notum immarginate, evenly, very widely rounded, cies, possibly occurring all over Java, from West to with surface unmodified. Pronotal sides immargin- East; differing from other gangkui group members ate. Pronotal border scarcely marginate in front. (from Borneo) in the structural details of the asym- Pronotal surface setose, setae long and more or less metric parameres (see diagnosis above and Fig. 135), squamiform; short, recurved pale bristles changing eye foramen size, pilosity, microsculpture, etc. to ditto scales from sides and apex to base); pilosity standing in annulate, crowded punctation (midline at base partly free); colour brown, anterolaterally rufous, generally matt, due to sculpture and pilosity. (d) The Ochicanthon hanskii group Estimated number of punctures per 0.25 sq.mm on Dorsum virtually uniformly brown-black. Parameres pronotal disc 65-70, punctures 0.04 mm. simple, roughly symmetric. Elytra with distinct rows Elytra entirely brown, moderately shining, with of bristles. Inner side of male metatibia angularly di- erect, apically more or less squamiform and recurved lated about halfway. The single known species also pale bristles. Humeral and anteapical umbones ab- shows attributes apparently related to the loss of its sent; elytral base shallowly depressed. Pseudepip- hind wings. leuron reaching elytral apex (broad, very coarsely rugulate-punctate, pseudepipeural crest narrow, lacking paired ridges in front). Punctures of discal elytral 22. Ochicanthon hanskii sp.n. striae slightly impressed (slightly crenulating inter- Figs 112-117, 131 striae, separated by 1-2 times their diameter), striae Type-material. Male holotype (RMNH): Malaysia, superficial; interstriae flat, mostly annulate-punctate, with our type label, and the following label data: seta-bearing punctures arranged in two distinct rows, Sarawak: 4th Division: Gunung Mulu NP, iii-v/1978, abundant to dense, shining between punctures. I. Hanski, LMR [lower montane rainforest], baited Antennal lamellae yellow-brown. pitfall trap, 800-1700 m, #678. Propectus laterally brown, abundantly annulate- 386 paratypes listed in Appendix 2. punctate-microsetose, microstriolate. Mesoster- num shining, finely annulate-punctate. Metaster- Description (holotype, male) nal anterior lobe and disc abundantly, finely Body length ca 4.6 mm. Colour generally brown. annulate-punctate, with very fine setae; metaster- Clypeal margin anteriorly bidentate (tips acute, num brown, laterally with annulate punctation, sitting on a short upright lobe, intervening emargin- dense. Abdominal venter brown, matt, entirely finely ation consequently less deeply rounded than in other annulate-punctate (distal sternites) or along base species). Clypeofrontal surface generally feebly con- (proximal sternites), lateral punctures microsetose, vex, brown, generally matt, glabrous, with crowded, impunctate parts microstriolate. Hind wings absent. distinct, ocellate punctation. Clypeal border from Pygidium slightly convex, marginate along base, gena forward very widely rounded. Clypeogenal brown, with fine setae, densely annulate-punctate. margin (at suture) slightly discontinuous. Genae Profemur brown, unmodified. Protibia with short laterally rounded to subangulate; with shallowly con- denticles. Protibia smooth, externally with 3 short cave surface. Eye size ventrally small. Eyes (foramen denticles. Proximal serration on outer side of pro- in full-face view) medium-sized, moderately ellip- tibia distinct, fine. Protibia apically rounded, only tic, short, widely open behind. Maximum number slightly protruding, internal side evenly, slightly of facet rows (transversely) across eye foramen 6-7. curved. Protibial spur inserted away from inner

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118 119 120 121

122 125 126

127a

123 124 127c 127b

Figs 118-127. Ochicanthon, aedeagus, upper side full-face. – 118, O. edmondsi; 119, O. dytiscoides; 120, O. masumotoi; 121, O. rombauti; 122, O. crockermontis; 123, O. crypticus; 124, O. peninsularis; 125, O. neglectus; 126, O. dulitmontis; 127, O. cambeforti, aedeagus, lateral views (a, right, b, left side), upper side full-face (c). Scale line = 0.5 mm, all same scale.

angle, simply acuminate (long, acute). Protibia light from apex. Metatibial apex lobate-dentate to (virtu- brown. Protarsi very slender, short. Mesofemur ally) straight. Meso- and metatibia brown. Mesotarsi slender, unmodified. Metafemur slender, posterior- slender. Metatarsi slender. Metatarsal segment 1 dis- inferior ridge angularly expanded near base. Meso- tinctly longer than segment 2 (approximate propor- and metafemur brown. Meso- and metatibiae dis- tions spur//segments 1-5 6//13/6/5/4/6). tinctly dilated distad. Mesotibia simply, slightly Aedeagus more or less symmetric, as in Fig. 131. curved. Metatibia nearly straight, proximally narrow, Measurements in mm. Maximum width of head 1.6. slender, abruptly dilated with interior angle at ca 0.3 Pronotal dorsal median length 1.4, maximum width

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128a 129a 130a 131a

128b 129b 130b 131b

132a 133a 134a

132b 133b 134b

135a 135b

Figs 128-135. Ochicanthon, aedeagus, lateral views (a, right, b, left side). – 128, O. woroae;�� 129, O. tambunan; 130, O. mulu;�� 131, O. hanskii;�� 132, O. gangkui;�� 133, O. kimanis;�� 134, O. danum;�� 135, O. javanus. Scale line = 0.5 mm, all same scale.

2.5. Dorsal sutural length of elytra 2.3, maximum Derivation of species name width combined 2.9. Named after one of the collectors, the distinguished ecologist I. Hanski, who, with Y. Cambefort, co- Variation and sexual dimorphism edited a most useful volume on dung beetle ecology Angles on anterior side of profemur and posterior (1991). side of metafemur in female absent. Internal side of female metatibia continuous, lacking the distinct an- Diagnosis gle of the male. Clypeal denticles slightly fused at base, emargination

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(e) Position uncertain and/or unusual (males less deep than in other species, adjoined laterally by unknown) simple concave curve. Clypeogenal border from lateral tip to anterior denticles widely convex-curvilinear. Clypeofrontal surface blackish-brown, crowdedly 23. Ochicanthon hikidai (Ochi, Kon & ocellate-punctate. Eye foramen short, small; under- Kikuta, 1997) side of eye also small. Pronotal border anterolaterally (at ca 0.2 behind anterior angle) simply, widely rounded. Paramarginal fold on posterolateral surface Diagnosis of pronotum virtually effaced. Pronotum crowdedly Clypeal denticles separated by emargination down to annulate-punctate, with numerous scattered setae, their base, adjoined laterally by deep emargination, uniform blackish-brown. Elytra uniform, entirely producing extra denticle on either side. Clypeogenal blackish-brown. Discal elytral interstrial puncta- border from lateral tip to anterior denticles virtually tion dense, interstriae shining, without microreticu- straight. Clypeofrontal surface densely to crowdedly lation, all flat or very slightly convex, mostly with punctate. Eye size large, foramen broadly elliptic. 2 rows of (squamiform) bristles. Pygidium more or less Pronotal border anterolaterally (at ca 0.2 behind evenly convex (in profile), brown to black, uniform. anterior angle) simply rounded. Paramarginal fold Protibia relatively broad, apex slightly expanded, on posterolateral surface of pronotum short, moder- denticles all shortly acuminate. Profemur anteriorly ately distinct. Pronotum densely to crowdedly punc- unmodified. Metafemur (male) posteriorly obtusely tate, with numerous scattered microsetae, uniform angulate at 0.3-0.5 from base. Metatibia straight to (brown-)black. Elytra uniform, entirely brown or very slightly curved, in male distal half abruptly di- black. Discal elytral interstrial punctation dense to lated, internal side angulate halfway its length. Meso- crowded, interstriae shining, without microreticula- and metatarsi short, relatively compact. Alae absent. tion, interstriae all flat or very slightly convex, in- Parameres subsymmetric (in upper side view), short, terstriae with numerous scattered microsetae. Pygid- broad, tapering (lateral view). Pronotal-elytral transi- ium more or less evenly convex (in profile), brown tion (in profile) with dip, habitus relatively convex. to black, (virtually) uniform. Metatibia straight to Body size small. very slightly curved, distal half dilated. Metatibia Both sexes known. Length 4.5-6 mm. internally (male) gradually dilated to apex. Metafe- mur posteriorly and metatibia internally not multi- Distribution and ecology denticulate. Meso- and metatarsi short, relatively Borneo: Sarawak. compact. Alae present. Pronotal-elytral transition (in Most specimens of this remarkable flightless species profile) gradual, habitus deplanate. Body size small. were collected by Hanski (1983, as Phacosoma sp.n. C) Male unknown. Length 4.0-4.7 mm. in carrion baited pitfall traps at Mulu (Sarawak) at 1320-1800 m altitude, in what was termed upper Distribution montane rain forest. Smaller numbers were found at Borneo: West Kalimantan. human dung, in flood refuse (150-200 m), and in litter. Comments This blackish Southwest Bornean species has four Comments protrusions on its clypeal margin, setting it, with- O. hanskii is a uniformly brown species from upland in the genus, completely apart. Note that only one Sarawak, easily recognizable from its general body female is known, from Mt Bawang in Indonesian shape, pilosity, clypeal dentation, angulate metat- Kalimantan (no altitude given). ibiae (in the male sex), etc. The hind wings seem to be always completely absent. The conspicuous dor- 24. Ochicanthon punctatus (Boucomont, 1914) sal scale-like bristles are easily brushed off. If there is any closer relative in the preceding groups at all, that might be O. mulu. Note that the paramarginal Diagnosis pronotal fold seems effaced and that the doubleness Clypeal denticles separated by emargination down to of the pseudepipleural crest is absent; nevertheless, their base, adjoined laterally by deep emargination, there can be no doubt that this species is congeneric producing extra denticle on either side. Clypeogenal with the preceding species. border from lateral tip to anterior denticles virtually straight. Eyes (foramen in full-face view) moderately large. Dorsum (especially elytra) more or less shining between punctation. Pronotum entirely

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(brown-)black. Pronotal sides anteriorly simply gaps, or whatever. Although the biology of Ochican- rounded. Elytral interstriae with dense, scattered thon species is still scarcely known, the middle and microsetae (may be very inconspicuous). Elytra en- hind legs of some (not all) species are long and curved, tirely (brown-)black. Elytral interstriae punctate. suggesting an adaptation to translocating food and/ Pronotal-elytral transition (in profile) gradual, ha- or breeding material – a behavioural trait so familiar bitus deplanate. Alae fully developed. Protibial den- in telecoprid Scarabaeinae; genuine ball rolling has ticles all simply acuminate, slender. Pygidium con- not been reported. Supposedly related small cantho- vex. Pygidium (brown-)black. Profemur anteriorly nines are known to indeed transport material, and unmodified (not dentate). Metafemur posteriorly to produce ovoid, egg-containing brood balls – see, unmodified (not dentate). Metatibia at most slightly for instance, Paulian (1976a, b). For a general review curved, with distal-internal section unmodified. of the nesting behaviour of Scarabaeinae see Halffter Body size small. & Edmonds (1982). Immatures of Ochicanthon are Only one female known. Length 3.5 mm still unknown.

Distribution Generic monophyly, supraspecific relationships, Erroneously from North Sulawesi; see Supplemen- biogeography tary note, after the General discussion; type data also Ochicanthon are structurally quite homogeneous, in Appendix 2. and (at least the Sundaland taxa) constitute a monophyletic group, with potential relatives outside Asia, Comments i.e. among other Old World members of the pre- O. punctatus is an all black-brown species, apparently dominantly austral tribe Canthonini (we disregard mislabelled, though presumably from Sundaland, the New World fauna here, as we see no direct re- and likely to belong to the gangkui or woroae group; lationships, cf. latest review of Halffter & Martinez with the single female at hand the species is as yet (1966-1977), and subsequent additions). unidentifiable. Further morphological and/or mo- The paramarginal pronotal fold and the detailed lecular work and ensuing nomenclatural decisions configuration of the pseudepipleuron may be among required. Omitted from the key to the species. the apomorphies uniting all Ochicanthon (in the odd, Phacosoma punctatum (sic) is mentioned in flightless O. hanskii the pronotal fold appears obso- Monaghan et al. (2007), in their molecular trees lete). The elytral striation pattern (7+2 striae), with being placed near certain Malagasy canthonines, as the broad elytral pseudepipleuron, the shape of the suggested in the discussion hereafter. tarsal claws, and the long first metatarsal segment, should all be helpful in the search for the closest relatives of Ochicanthon, with one important notion: General discussion there are no known close relatives in the Oriental and adjacent Palaearctic Regions (simply because Scope none of the other canthonines there has pseudepip- The objectives of this discussion are 1) to review the leura or anything else qualifying as synapomorphic). taxonomic and biogeographic patterns emerging Formerly, some non-Oriental Canthonini were from this study of Ochicanthon, 2) put forward some placed in or near Phacosoma, but meanwhile they are testable hypotheses on the evolution of the group, all, and rightly so, placed elsewhere, in other genera. and 3) formulate a number of ensuing research ques- Nevertheless, the groups to which they are currently tions. In view of the extent of these questions, a full assigned may include genuine relatives of Ochican- cladistic analysis down to species level is considered thon. Old World faunal synopses to consider here premature. include: Lebis (1953), Matthews (1974), Paulian (1975, 1976a, b, 1985, 1991), Paulian & Pluot- Biological background Sigwalt (1985), and Scholtz & Howden (1987). The small scarabs of the genus Ochicanthon are wide- From our scan of both these publications and recent spread in southern Asia (Fig. 2), and in parts of Sun- descriptions of generic novelties, plus an inspection daland they are quite common in both lowland and of relevant specimens, we conclude that suitable can- upland forest ecosystems, being attracted to ground didates for a closer phylogenetic relationship with traps baited with faeces or carrion; some species are Ochicanthon may be found among Paulian’s (1976b) active flyers, others are flightless. Their known gener- Afrotropical-Madagascan “Canthonina Longitar- ic distribution is more or less mosaic – due to simple ses”, rather than among Australasian-Pacific genera. ignorance, recent ecological changes (including hab- Actually, Paulian indeed placed Ochicanthon in the itat loss), historically and geologically determined same league as his “Longitarses”, based on their

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similarities in the characters mentioned above (note, usage of the terms varies among authors; (ii) the ab- however, that some of his generic characterizations solute homology of the development of pseudepip- are very sketchy and illustrations limited). A mod- leura in various genera, along with other characters ern reclassification of the rich Malagasy canthonine of the elytra, has not been settled; (iii) in some gen- fauna, started by Montreuil around 2002, might well era the elytral and claw characters do not correlate as shed light on the position of Ochicanthon; for a taste indicated in the previous paragraph (see Oficanthon of what is to come, see also Monaghan et al. (2007). Paulian, 1985 and Penalus Paulian, 1985, both from New Guinea). On the intrageneric levels, within the Oriental Re- gion, the Indian Subregion, the obvious source region Sundaland species: tentative grouping and for the Southeast Asian members of Ochicanthon, has distribution a morphologically diverse but poorly known fauna The ranges of most species appear to be limited, and (8 species, few records). There are considerable gaps one species, O. hanskii from Sarawak, is flightless; al- in the known western distribution of the genus, but, though Ochicanthon may be ancient as a group (how based on our experience in Sundaland, with ad- ancient is, of course, uncertain), this flightlessness equate sampling many novelties might be discovered may well have evolved more recently, i.e. measured – scarabaeologists in the region could do a useful job on a geological time scale. In practical terms limited here. The described Indian-Sri Lankan species are in ranges mean that the collecting density has to be need of revision in relation to the Southeast Asian both high and with an adequate ecological resolution fauna and, with the supplementary exploration sug- to fully appreciate a local Ochicanthon fauna, and, gested, we predict the discovery of elements basic to as already indicated above for the continental range the Sundaland groups dealt with in this paper. Apart of the genus, even on a large scale, gaps are consid- from the Indian connection, the Sundaland fauna erable. A few species seem to be more widespread, has close relatives nearby, for instance in Thailand particularly certain lowland species, like the Bornean (see discussion of species groups below). O. dytiscoides and woroae, and perhaps obscurus in Everything would, of course, be turned upside down continental Southeast Asia (this is apparently a spe- if Southeast Asia would prove to be the cradle of cies of open habitats, Kabakov & Napolov 1999). Ochicanthon, for instance, if oddities, like O. hanskii, O. woroae seems a widespread, active flyer, being would (say, on molecular characters) prove to be frequently encountered in flight interception traps. older (i.e. closer to the root of the Ochicanthon phyl- Scholtz (2000) discussed various aspects of flightless ogeny) than the other groups discussed here. scarabs, which include many canthonines (see also All in all, irrespective of our chequered knowledge, Bell et al. 2004, for a relevant molecular view on present-day Ochicanthon lineages seem to have a flightless canthonines in Australia). long standing in the Oriental Region, and they may We estimate that there are at least ten more unde- well represent a heritage from Indo-Gondwanan scribed Ochicanthon species in Sundaland – let alone ages, now ranging from the Indian Subregion into the remainder of the Oriental Region. Some regions Southeast Asia (for the general context of Scarabaei- show a very limited, others, like Borneo, a remark- nae history and classification see also Scholtz & Gre- ably rich Ochicanthon fauna (see Table 1) – the ques- bennikov (2005) and included references). tion, as indicated before, is of course: why – is this a Until recently some authors (including Matthews sampling artefact, or is it genuine? 1974) have maintained, formally or informally, So far, only one species, not directly related to our two groups within the Canthonini, the “primitive” Sundaland species, was recorded from the Phil- Mentophilina (with pseudepipleura and indentate ippines (Mindanao: Mount Apo). The fauna of tarsal claws, not genuine ball rollers) and the “de- Java seems poor (we recognize only one species), rived” Canthonina (no pseudepipleura, with dentate and no Ochicanthon were found on Bali and the tarsal claws, genuine ball rollers) – and clearly, in this Lesser Sundas (although the Javan species reaches the simplified scheme, Ochicanthon would belong to the eastern tip of Java). Our intensive explorations on first group. We are sceptical, however, as to the phy- Sulawesi and beyond neither yielded Ochicanthon; logenetic validity of this dual division, and look for- the single known specimen recorded from Sulawesi ward to future critical analyses of the world fauna of seems to be part of a mislabelled lot (see note be- what appears to be a polyphyletic group Canthonini low). Sulawesi and other eastern islands, have an (see also Philips et al. 2004, Monaghan et al. 2007). assortment of canthonine species belonging to Our scepsis is strengthened by three morphologi- the very widespread, heterogeneous Afro-Indo- cal points: (i) the distinction between claws that are Pacific Panelus-Lepanus complex and to typically dentate, subdentate or indentate is not clear-cut, and Australasian-Pacific canthonine elements (Matthews

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Table 1. Comparison of total scarab specimen numbers collected in four different Sundaland areas in relation to Ochicanthon numbers. Flight interception traps were only used in Danum valley and are therefore omitted from this table. As for the Leuser material, only a reference collection was available.

Area Altitude Total number of Total number of Ochicanton scarab dung beetles Ochicanton specimens species

Sarawak: Mt Mulu NP 50- 2370 5897 554 8 Sabah: Crocker range 900- 1500 1556 255 4 Sabah: Danum Valley 100- 300 16700 183 4 Sumatra: Mt Leuser NP 150- 2500 18300 x 1

1974, Paulian 1985, 1991, Huijbregts & Krikken another belonging to either group b or c, and with an 2007, etc.). uncertain geographic origin (O. punctatus). In summary, adding our knowledge of (the eastern We are content that our operational groups (a, b, c) limits of) the range of Ochicanthon to the possible constitute natural (monophyletic or paraphyletic) Afrotropical-Madagascan-Indian connection dis- clusters. As indicated above, this is particularly based cussed above, there is every reason to hypothesize on genital structure, colour patterns, nature of dor- that the genus invaded pre-Quaternary Southeast sal pilosity, and shape of leg elements. The gangkui Asia, including “proto-Borneo”, from the West, subgroup members (c), with strongly modified para- sequently profusely speciating there, but not crossing meres, may well be descendants of (a member of) into Wallacea. the woroae group (b), the members of which have relatively simple parameres – morphoclinal transi- Four operational, possibly phylogenetically accept- tions are quite conceivable. able groups of species may be distinguished within As for the possible ancestry of the flightless O. hanskii Sundaland Ochicanthon, and, in the preceding sec- (d), it could well have a more direct relationship tion Species accounts, the species have been arranged with the flight-capable, peculiarly setose O. mulu or accordingly: another species in the woroae group (b), but this con- (a) The symmetrically patterned (orange-yellow sideration would be based rather on overall similar- versus brown-black) members of the dytiscoides ity than on synapomorphism or any other phyloge- group, with a usually subsymmetric aedeagus netic consideration; their parameres are unmodified. (showing more or less modified parameral bot- O. hanskii has not just lost its hind wings: the pre- toms) and angularly dilated metatibial apex in sumably flightlessness-related pronotal-elytral shape the male sex; two subgroups are here distin- is clearly different from its flight-capable relatives in guished (on protibial dentation and aedeagus group (b). It is peculiar that in the dytiscoides group details); this group comprises here 13 known a large species with a similar shape (O. rombauti) species. seems fully winged and capable of flight. (b) The small, uniformly brown to black members of Intra-group species-level relationships must, for the the woroae group with a subsymmetric aedeagus, time being, remain in the domain of speculation, with unmodified legs and all winged; includes additional material, also from outside Malaysian three known species. Borneo, as well as the application of other techniques (c) The small, uniformly brown to black members (see below), being required. of the gangkui group with a distinctly asymmet- The relationships of the four Sundaland groups ric aedeagus, lacking coarser bristles and scales, (a-d) with Ochicanthon elsewhere are not im- with unmodified legs; includes five species. mediately clear, were it alone for the fact that the (d) The uniformly brown to black members of the taxonomically important structure of the male geni- hanskii group with a subsymmetric aedeagus, talia of several non-Sundaland species has not been serially arranged, coarse interstrial bristles, and studied. Species with colour patterns and leg shapes peculiarly shaped metatibiae; includes the single basically similar to those in the dytiscoides group (a) known flightless species O. hanskii. occur in continental Southeast Asia (in Thailand, see Two species are placed outside these four groups Hanboonsong & Masumoto 2001), and these are (under e): both are known from females only, one undoubtedly closely related to Sundaland members having a characteristic clypeal shape (O. hidikai), of the group.

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Some species seem to constitute sets of closely re- being essentially Indo-Gondwana-based, with lated geographical or ecological vicariants; ecological subsequent differentiation in Oriental subre- gradients are lowland vs. upland, and heath forest gions? vs. limestone forest. Samples from such different situations were initially only suspected to represent Some of these questions could certainly be answered different species, but they were confirmed in their by working from research stations with easy ac- status by morphological characters, above all, by the cess to Ochicanthon populations – Borneo being a detailed shape of the male genitalia. In Sundaland, prime candidate. To local scientists, therefore, we only the forms united under or close to O. dytiscoides, recommend to use their resources and opportuni- and O. woroae, appear to be more widespread low- ties to systematically collect material, and do asso- land species, within Borneo. Yet, here also caution ciated field and lab work, as these are likely to be is required, as there is some variation which might rewarding. Applying complementary molecular prove to be geographic (i.e., subspecific or popula- techniques might be most helpful in reconstructing tional), and, moreover, the status of the forms in the the evolution of Ochicanthon, and estimating a time dytiscoides complex remains a moot point. frame (see the example of Bell et al. 2004, and, in a very different ecosystem setting, Sole et al. 2005). Research questions With the present -taxonomic paper we just hope In conclusion, the scarab genus Ochicanthon proves to have pushed Ochicanthon slightly further into a to be a most interesting taxonomic and evolutionary ß-taxonomic future. research challenge, with questions on various levels of integration, such as: • can we pinpoint the closest relative (sister group) Supplementary note on Boucomont’s of Ochicanthon, perhaps in non-Oriental faunas, 1914 Sulawesi records among the Afrotropical-Madagascan “Longitar- ses” of Paulian? On examining material in the Paris museum it be- • what are the relationships of the Sundaland fau- came clear that seven species in the Scarabaeinae na with the species elsewhere in the vast generic recorded by Boucomont (1914) from Sulawesi were range, from Sri Lanka to China? based on material having all the same label “Ton- • what is the precise biology of Ochicanthon, what dano \ coll. C”; Tondano is a place in North Sulawe- could be its bearing on speciation, what do their si visited by several insect collectors. Our colleague immature stages look like, are their characters Y. Cambefort could not shed any light on the mys- taxonomically complementary to those of the terious collector or collection C of 1899. The species adults? are: • which factors selected for an ecological differen- Ochicanthon punctatus (Boucomont, 1914) (as tiation, apparent in the specialization of certain Phacosoma) species (ecosystem type, soil type, altitudinal oc- Cassolus peninsularis Arrow, 1907 currence, etc.)? Onthophagus aphodioides Lansberge, 1883 • which ecological processes favoured the loss Onthophagus denticollis Lansberge, 1883 of flight in certain species (regional, ecological Onthophagus rorarius Harold, 1877 isolation?), is this indeed an ecological phenom- Onthophagus rudis Sharp, 1875 enon, or do historical or geological factors (also) Onthophagus vulpes Harold, 1877 play a role? North Sulawesi was particularly well surveyed during • would further systematic baited pitfall trap- Project Wallace (1985). However, none of the above ping campaigns (particularly outside Malaysian species was rediscovered in the region. Furthermore, Borneo!) indeed reveal much more taxonomic none of the species we found to be common in North diversity, as predicted, and where? Sulawesi during 1985 were found in the collection of • can the taxonomy of the dytiscoides complex be the Paris museum with a label “Tondano \ coll. C”. It resolved, are there a few variable species, or are is therefore assumed that all the specimens with these more taxa involved? specific data were mislabelled. Most of the species are • could character mapping of the known spe- found on Borneo, Sumatra and the Malay Peninsula. cies and any additional taxa produce a well- As O. denticollis is known only from Sumatra, this supported phylogeny, a cladogram, a phylogeog- island seems the most likely origin of the material, raphy? or, at any rate, one of the Sunda Islands. The reputed • would the answers to the previous and related occurrence of Ochicanthon on Sulawesi is based on questions support the hypothesis of Ochicanthon the type of Phacosoma punctatum from Tondano.

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For the time being it is assumed that Ochicanthon Davis, A.J., 1999. Species packing in tropical forests: is not represented on Sulawesi (we collected all over diel flight activity of rainforest dung-feeding beetles Sulawesi). (Coleoptera: Aphodiidae, Scarabaeidae, Hybosori- dae) in Borneo. – Raffles Bulletin of Zoology 47 (2): 473-486. Acknowledgements Davis, A.J., 2000. Does reduced-impact logging help Sincere thanks are due to our ecological colleagues preserve biodiversity in tropical rainforests? A case for contributing a wealth of material from their re- study from Borneo using dung beetles (Coleoptera: search in the Southeast Asian islands, in particular Scarabaeoidea) as indicators. – Environmental Ento- Ilkka Hanski and Hannu Räisänen (both Finland), mology 29 (3): 467-475. Andrew Davis (UK), and Ashley Kirk-Spriggs (South Davis, A.J., J.D. Holloway, H. Huijbregts, J. Krikken, Africa, formerly UK). The names of other collectors A.H. Kirk-Spriggs & S.L. Sutton, 2001. Dung bee- appear in Appendix 2. tles as indicators of change in the forests of northern Ed Rombaut (Netherlands) rendered invaluable Borneo. – Journal of Applied Ecology 38 (3): 593-616. technical assistance to the first author during a trip Davis, A.J., H. Huijbregts & J. Krikken, 2000. The role to Sabah in 1987. Frenk Driessen (Netherlands) pro- of local and regional processes in shaping dung beetle duced the habitus drawing of Ochicanthon edmondsi. communities in tropical forest plantations in Borneo. Rienk de Jong and Jan van Tol (Netherlands), and – Global Ecology and Biogeography 9 (4): 281-292. Olivier Montreuil (France) commented on a draft of Fosberg, F.R., 1967. Classification of vegetation for general this paper. Peter Hammond and his colleagues sup- purposes. In: Peterken, G.F., Guide to the checksheet ported our work in The Natural History Museum, for IBP areas. – Blackwell, Oxford, IBP Handbook 4: London, and facilitated the loan of specimens. Yves 73-120. Cambefort supported our work in the Muséum Halffter, G. & W.D. Edmonds, 1982. The nesting behav- National d’Histoire Naturelle, Paris, and Olivier ior of dung beetles (Scarabaeinae). An ecological and Montreuil arranged the loan of Paris material. Shuhei evolutive approach. – Publicaciones Instituto de Eco- Nomura of the National Science Museum, Tokyo, logia Mexico 10: 1-176. supplied literature. Halffter, G. & A. Martinez, 1966. Revision monograph- Our thanks are also due to the authorities who grant- ica de los Canthonina Americanos (Coleoptera: Scarabaeidae). (1a. parte). – Revista de la Sociedad ed research permissions, and to local scientists who Mexicana de Historia Natural 27: 89-177. provided support; sponsoring organizations included Halffter, G. & A. Martinez, 1967. Revision monographica Sabah Parks and the Forestry Department (both in de los Canthonina Americanos (Coleoptera: Scara- Sabah, Malaysia), and the Indonesian Institute of baeidae). (2a. parte). – Revista de la Sociedad Mexi- Science (LIPI, Indonesia). cana de Historia Natural 28: 79-116. Halffter, G. & A. Martinez, 1968. Revision monographica de los Canthonina Americanos (Coleoptera: Scara- References baeidae). (3a. parte). – Revista de la Sociedad Mexi- Arrow, G.J., 1931. Fauna of British India. Coleoptera cana de Historia Natural 29: 209-290. Lamellicornia. Part III. – Taylor & Francis, London, Halffter, G. & A. Martinez, 1977. Revision monographica i-xii, 1-428. de los Canthonina Americanos (Coleoptera: Scara- Balthasar, V., 1963. Monographie der Scarabaeidae und baeidae). IV parte. Clave para generos y subgeneros. Aphodiidae der palaearktischen und orientalischen – Folia Entomologica Mexicana 38: 29-107. Region (Coleoptera Lamellicornia). 1. – Tsjechoslow- Hanboonsong, Y. & K. Masumoto, 2001. Dung bee- akische Akademie der Wissenschaften, Prague, 1-391. tles (Coleoptera, Scarabaeidae) of Thailand. Part 4. Bell, K.L., D.K. Yeates, C. Moritz & G.B. Monteith, 2004. Genera Phacosoma, Cassolus and Parachorius (Can- Molecular phylogeny and biogeography of the dung thonini and Dichotomini). – Elytra 29(1): 129-140. beetle genus Temnoplectron Westwood (Scarabaeidae: Hanski, I., 1983. Distributional ecology and abundance Scarabaeinae) from Australia’s wet tropics. – Molecular of dung and carrion-feeding beetles (Scarabaeidae) in Phylogenetics and Evolution 31(2): 741-753. tropical rain forests in Sarawak, Borneo. – Acta Zoo- Boucomont, A., 1914. Les Coprophages de l’Archipel logica Fennica 167: 1-45. Malais (Coléopt.). – Annales de la Societé Ento- Hanski, I. & J. Krikken, 1991. Dung beetles in tropical mologique de France 73: 238-350. forests in South-East Asia. In Hanski, I. & Y. Cambe- Boucomont, A., 1920. Coléoptères Coprophages nouveaux fort (eds), Dung beetle ecology. – Princeton University d’Asie et de Malaisie (Scarabaeidae). – Annales de la Press, Princeton New Jersey: 179-197. Societé Entomologique de France 88: 307-320. Huijbregts, J. & J. Krikken, 2007. First record of the ge- Dallwitz, M.J., 2005. Overview of the DELTA System. nus Amphistomus from the Moluccas (Coleoptera: – http://delta-intkey.com/. [Last accessed 10 Sep Scarabaeidae: Scarabaeinae). – Tijdschrift voor Ento- 2007]. mologie 150: 31-37. Downloaded from Brill.com10/11/2021 05:43:19AM via free access 472 Tijdschrift voor Entomologie, volume 150, 2007

Jukes-Browne, A.J., 1912. The genus Dosinia and its sub- Paulian, R., 1976b. Révision des Canthonina Longitarses divisions. – Proceedings of the Malacological Society (Col. Scarabaeidae) de Madagascar. – Annales de la of London 10: 95-106. Societé Entomologique de France (NS)12: 453-479. Kabakov, O.N. & A. Napolov, 1999. Fauna and ecology of Paulian, R., 1980. Coléoptères Scarabaeidae Canthoni- Lamellicornia of subfamily Scarabaeinae (Coleoptera, nae d’Inde du Sud. – Revue Suisse de Zoologie 87: Scarabaeidae) of Vietnam and some parts of adjacent 57-65. countries: South China, Laos and Thailand. – Latvijas Paulian, R., 1983. Sur quelques coléoptères Scarabaeoidea Entomologs 37: 58-96. de la region Orientale. – Revue Suisse de Zoologie 90: Lebis, E., 1953. Révision des Canthoninae de Madagascar 615-622. (Col. Scarab.). – Mémoires de l’Institut Scientifique Paulian, R., 1985. Les coléoptères Scarabaeidae cantho- de Madagascar 3: 107-252. nines de Nouvelle-Guinée. – Annales de la Societé Löbl, I. & A. Smetana (Eds), 2006. Catalogue of Palae- Entomologique de France 21: 219-238. arctic Coleoptera. 3. – Apollo, Stenstrup Denmark, Paulian, R., 1987. Notes sur les coléoptères Scarabaeoidea 1-690. du Museum de Genève. 3. – Revue Suisse de Zoologie Masumoto, K., 1988. Coprophagid-beetles from north- 94: 717-724. west Thailand (II). – Entomological Review of Japan Paulian, R., 1991. Les coléoptères Scarabaeoidea de Nou- 43: 135-143. velle-Caledonie. – Faune Tropicale 29: 1-164. Masumoto, K., 1989. Coprophagid-beetles from north- Paulian, R. & D. Pluot-Sigwalt, 1985. Les canthonines west Thailand (III). – Entomological Review of Japan de Nouvelle-Caledonie (Coleoptera, Scarabaeidae). 44: 31-43. Etude systematique et biogeographique. – Bulletin du Matthews, E.G., 1974. A revision of the scarabaeine dung Muséum National d’Histoire Naturelle Section A Zo- beetles of Australia. II. Tribe Scarabaeini. – Australian ologie Biologie et Ecologie Animales 6: 1091-1133. Journal of Zoology, Supplementary Series 24: 1-211. Philips, T.K., E. Pretorius & C.H. Scholtz, 2004. A phy- Monaghan, M.T., D.J.G. Inward, T. Hunt & A.P. Vogler, logenetic analysis of dung beetles (Scarabaeinae: Scara- 2007. A molecular phylogenetic analysis of the Scara- baeidae): unrolling an evolutionary history. – Inverte- baeinae (dung beetles). – Molecular Phylogenetics and brate Systematics 18: 53-88. Evolution, in press. Scholtz, C.H., 2000. Evolution of flightlessness in Scara- Ochi, T., 1990. Two new coprophagous beetles (Coleop- baeoidea (Insecta, Coleoptera). – Mitteilungen aus tera, Scarabaeidae) from the Philippines. – Elytra 18: dem Museum für Naturkunde in Berlin, Deutsche 215-219. Entomologische Zeitschrift 47: 5-28. Ochi, T. & K. Araya, 1996. Studies on the copropha- Scholtz, C.H. & V.V. Grebennikov, 2005. 12. Scarabaei- gous scarab beetles from East Asia. 3. (Coleoptera, formia Crowson, 1960. 13. Scarabaeoidea Latreille, Scarabaeidae). – Giornale Italiano di Entomologia 8: 1802. – Handbuch der Zoologie (4)38: 345-429. 1-15. Ochi, T., M. Kon & M. Kawahara, 2007. A new species Scholtz, C.H. & H.F. Howden, 1987. A revision of the of the genus Ochicanthon (Coleoptera, Scarabaeidae) African Canthonina (Coleoptera: Scarabaeidae: Scara- from Sumatra, Indonesia. – Kogane 8: 85-88. baeinae). – Journal of the Entomological Society of Ochi, T., M. Kon & T. Kikuta, 1997. Studies on the fam- Southern Africa 50: 75-119. ily Scarabaeidae (Coleoptera) from Borneo 3. The ge- Sole, C.L., C.H. Scholtz & A.D.S. Bastos, 2005. nus Phacosoma Boucomont, with descriptions of four Phylogeography of the Namib Desert dung beetles new species. – Giornale Italiano di Entomologia 8: Scarabaeus (Pachysoma) MacLeay (Coleoptera: Scara- 251-260. baeidae). – Journal of Biogeography 32: 75-84. Ochi, T., A. Ueda & M.Kon, 2006. Ochicanthon (Coleop- Vaz-de-Mello, F.Z., 2003. Ochicanthon, a new name for tera, Scarabaeidae) from Borneo with descriptions of Phacosoma Boucomont (Coleoptera, Scarabaeidae), four new species. – Elytra 34: 309-325. preoccupied with Phacosoma Jukes-Browne (Mollus- Paulian, R., 1975. Sur quelques Canthonina (Coléop- ca). – Coleopterists Bulletin 57: 25-26. tères Scarabéides) montagnards de Madagascar. – An- nales de la Societé Entomologique de France (NS)11: 221-252. Paulian, R., 1976a. Coprophaga Africana. III. Observa- tions sur les Canthonina malgaches (Coleoptera Scara- Received: 29 January 2007 baeidae). – Revue de Zoologie Africaine 90: 121-161. Accepted: 10 July 2007

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Appendix 1 4. glabrous, or nearly so 10. pronotal colour 1. uniform (brown-)black (no trace of Synoptic table of Ochicanthon characters (orange-)yellow marks) 2. with symmetric pattern of (orange-)yellow Characters and their states and (brown-)black 1. clypeal denticles (shape) 11. pronotal (orange-)yellow marks 1. separated by emargination down to their base 1. limited to small lateral patch on either side 2. fused at base (emargination less deep) 2. abundant, (orange-)yellow not predominant 2. clypeal denticles (shape, delimitation, number) 3. extensive, (orange-)yellow predominant 1. adjoined laterally by simple concave curve 12. elytral colour (pattern blurred in some species) 2. adjoined laterally by shallow (sinuate) emar- 1. uniform, entirely brown or black gination 2. predominantly black-brown, with (usually 3. adjoined laterally by deep emargination, pro- symmetric) pattern of (orange-)yellow marks ducing extra denticle on either side 3. predominantly (orange-)yellow, symmetric- 4. laterally virtually straight black marks limited 3. clypeogenal border from lateral tip to anterior 13. basal elytral (orange-)yellow (B) denticles (outline) 1. absent 1. widely convex-curvilinear 2. present 2. virtually straight 14. postmedial elytral (orange-)yellow (M) 4. clypeofrontal surface (punctation) 1. absent 1. abundantly punctate 2. present 2. densely to crowdedly punctate 15. distal elytral (orange-)yellow (A) 3. rugulate-punctate 1. absent 5. eye (size foramen and eye underside) 2. present 1. large, foramen broadly elliptic (inner border 16. discal interstrial punctation straight or convex) 1. abundant 2. moderate, foramen moderately or narrowly 2. dense to crowded elliptic (inner border more or less concabe) 3. (asperate-)rugulate 3. small (underside of eye also small) 17. discal interstriae (reflection) 6. pronotal border anterolaterally (at ca 0.2 behind 1. shining, without microreticulation anterior angle, full-face view) 2. sericeous, due to microreticulation 1. more or less abruptly rounded (may be sub- 3. matt, due to (asperate-)rugulate microsculp- angular) ture 2. simply rounded 18. discal interstriae (elevation) 3. angularly bent inward (straight or concave in 1. all flat or very slightly convex dorsal view) 2. 3 and 5 slightly elevated 7. paramarginal fold on posterolateral surface of 19. discal interstriae (pilosity) pronotum 1. with numerous scattered microsetae (may be 1. short, moderately distinct very inconspicuous) 2. distinct, (angularly) extended in front 2. abundantly, relatively coarsely setose or squa- 3. extended in front with two branches mose (more or less in rows) 4. obsolescent (effaced, may be lost in micro- 3. mainly with scattered microstubbles sculpture) 4. glabrous, or nearly so 8. pronotum (discal punctation) 5. mostly with two distinct rows of bristles 1. abundantly punctate 20. pygidium (shape, specify male or female) 2. densely to crowdedly punctate 1. more or less evenly convex (in profile) 3. rugulate-punctate 2. convex along apex, basal surface flat or con- 9. pronotum (discal pilosity) cave (in profile) 1. with numerous scattered microsetae (may be 3. more or less pointed (may be subconical) very inconspicuous) 21. pygidium (predominant colour) 2. abundantly, relatively coarsely setose or squa- 1. brown to black, (virtually) uniform mose 2. largely (orange-)yellow 3. with microstubbles 3. with distinct light patches

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Synoptic table of Ochicanthon characters. x not scored (unknown or not applicable); - from-to symbol; & and symbol (more states); / or symbol (more states).

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40

1 O. edmondsi 1 1 1 2 2 3 2 2 1 2 1 2 2 2 2 2 1 1 1 1 1 3 2 3 2 1 2 1 2 1 1 3 x x 1 1 2 1 1 8-10 2 O. dytiscoides 1 1 1 2 2 1 1 2 1 2 3 2 2 2 2 2 1 1 1 2 2 3 2 2 2 1 2 1 2 1 1 2 x x 1 1 1 1 1 5-7 3 O. masumotoi 1 1 1 2 2 1 1 2 1 2 2 2 2 2 2 2 1 1 1 2 2 3 2 2 2 1 2 1 2 1 1 2 x x 1 1 1 1 1 4-6.5 4 O. maryatiae 1 1 1 2-3 2 1 2 2 1 2 2 2 2 2 2 2 1 1 1 2 x 3 2 2 2 1 2 1 2 1 1 2 x x 1 1 1 1 1 7.2-8.1 5 O. rombauti 1 1 1 2 2 2 2 2 2 2 1 2 2 2 2 2 1-2 2 1 2 1 3 2 2 2 1 1-2 1 2 1 1 2-3 x x 2 2 2 1 1 8-9 6 O. crockermontis 1 1 1 2 2 1 1 2 1 2 3 3 x x x 2 1 1 1 2 2 1 2 2 2 1 1-2 1 2 1 1 1 x x 1 1 1 1 1 5.5-6 7 O. crypticus 1 1 1 2 2 1 1 2 1 2 2 2 2 2 2 2 1 1 1 2 1&3 1 2 1 2 1 2 1 2 1 1 1 x x 1 1 1 1 1 5.5-6 8 O. peninsularis 1 1 1 2 2 2 1 2 1 2 2/3 2 2 2 2 2 2 1 1 2 2 1 2 2 2 1 2 1 2 1 1 2 x x 1 1 1 4 2 6-7 9 O. kikutai 1 1 1 2-3 2 2 1-2 2 3 2 3 2 2 2 2 2 1 1 1 2 x 1 2 3 2 1 2 1 2 1 1 2 x x 1 1 1 1 1 7.2-7.5 10 O. neglectus 1 1 1 2 2 1 1 2 1 1 x 2 2 2 1 2 1 1 1 2 1 1 2 2 2 1 2 2 2 1 1 2 x x 1 1 1 1 1 7-8 11 O. dulitmontis 1 1 1 1-2 2 1 1 2 1 1 x 2 2 2 1 2 1 1 1 2-3 1 1 2 2 2 1 2 1 2 1 1 2 x x 1 1 2 1 3 8-8.5 12 O. cambeforti 1 4 2 2 2 1-2 1 2 1 2 2 2 2 2 2 2 1 1 1 2 1 1 2 3 2 1 2 1 2 1 2 x x x 1 1 1 1 1 6.5-6 13 O. karasuyamai 1 1 1 1-2 2 2 1 2 3 1 x 2 2 2 2 1 2 1 1 3 1/3 2 2 2 2 1 2 1 1 1 2 x x x 1 1 1 2 1 5.8-8.2 14 O. woroae 1 1 1 2 2 2 1 1 4 1 x 1 x x x 2 1 1 3/4 1 1 1 1 1 1 1 1 1 2 1 1 1 x x 1 1 1 1 1 3.5-4 15 O. tambunan 1 1 1 2 2 1 1 2 1 1 x 1 x x x 2 3 1 1 1 1 1 1 1 1 1 1 1 2 1 1 2 x x 1 1 1 1 1 3.5-4.5 16 O. mulu 1 2 1 2 1 2 1 2 1 1 x 1 x x x 2 1 1 2 1 1 1-2 1 1 1 1 1 1 1 1 1 2 x x 1 1 1 1 1 3.5-4.5 17 O. gangkui 1 1 1 2 1 1 1 2 1 1 x 1 x x x 2 3 1 1 1 1 1 1 1 1 1 1 1 2 1 2 x 1 3 1 1 1 1 1 4-4.5 18 O. kimanis 1 1 1 2 2 2 1 2 4 1 x 1 x x x 2 3 1 1 1 1 1 1 1 1 1 1 1 2 1 2 x 1 2 1 1 1 1 1 4-4.5 19 O. danum 1 1 1 2 1 2 1 2 4 1 x 1 x x x 2 1 1 1 1 1 1 1 1 1 1 1 1 2 1 2 x 2 2 1 1 1 1 1 4-4.5 20 O. parantisae 1 1 1 2 1 1 1 2 1 1 x 1 x x x 3 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 x 2 x 1 1 1 1 1 4.5-5 21 O. javanus 1 1 1 2 1 2 1 2 4 1 x 1 x x x 2 1 1 1 1 1 1 1 1 1 1 1 1 2 1 2 x 1 3 1 1 1 3 1 3.5-4.5 22 O. hanskii 2 1 1 2 3 2 4 2 2 1 x 1 x x x 2 1 1 5 1 1 2 1 2 1 2 1 1 1 3 1 1 x x 2 2 1 1 1 4.5-6 23 O. hikidai 1 3 2 2 1 2 1 2 1 1 x 1 x x x 2 1 1 1 1 1 x x x 1 x 1 1 1 1 x x x x 1 1 1 1 2 4.0-4.7 24 O. punctatus 1 1 2 2 2 2 1 2 1 1 x 1 x x x 2 1 1 1 1 1 x x x 1 x x 1 x 1 x x x x 1 1 1 x 2 3.5

22. protibial denticles (male) (shape, avoid abraded 1. not angulate near apex specimens) 2. angulate-dilated at short distance from apex 1. all simply acuminate, slender (male) 2. all shortly acuminate 28. metafemur posteriorly and metatibia internally 3. 1-2 acuminate, apico-external denticle broad, (denticulation) its base parallel-sided 1. not multidenticulate 23. profemur (male) anteriorly (angle, denticle) 2. with longitudinal row of fine denticles (mini- 1. unmodified crenulate) 2. lobate-dentate near apex 29. meso- and metatarsi (relative length) 24. metafemur (male) posteriorly (angles, denticles) 1. short, relatively compact and complanate 1. unmodified 2. long, relatively slender 2. lobate-dentate at 0.3-0.5 from base 30. alae (reduction) 3. doubly lobate(-dentate) (also protruding at ca 1. present 0.8 from base) 2. distinctly reduced 25. metatibia (curve and dilatation) 3. absent 1. straight to very slightly curved, dilated 31. parameres (general shape) distad 1. roughly symmetric (in upper side view) 2. long, slender, curved, dilated near apex 2. distinctly excised, asymmetric, complex 26. internal side of metatibia (male) (shape of metat- 32. parameres (general shape) ibia 25-1)) 1. short, broad, tapering (lateral view) 1. continuous (not angulate halfway) 2. long, slender, tapering (lateral view) 2. abruptly angular halfway its length 3. long, dilated distally (full-face upper side) 27. internal side of metatibia (male) (ventral view, 33. left paramere (shape details) obtuse internal angle) 1. very deeply excised, inferior branch elongate-

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Synoptic table of Ochicanthon characters. x not scored (unknown or not applicable); - from-to symbol; & and symbol (more states); / or symbol (more states).

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40

expanded (into lobe) 39. sexes 2. more shallowly excised, inferior branch short 1. both known 34. right paramere (shape details) 2. male unknown 1. sinuate, lobate at base (lateral view) 3. female unknown 2. widely sinuate 40. length in mm (usually in 0.5 mm) 3. deeply excised, lobate 35. pronotal-elytral transition (in profile) 1. gradual 2. abrupt (with a dip) Appendix 2 36. habitus 1. deplanate Compilation of material examined 2. relatively convex 37. total length (i.e. average of series in half mm) 1. body size small (length usually under Label data 8 mm) Species sequence as in the section Species accounts in 2. body size large (length usually at or over the main text (using same species numbers). Missing 8 mm) species numbers: no specimens seen. 38. known range All specimens of new species recorded below, and 1. Borneo seen and labelled by us, are types, unless explicitly 2. Sumatra excluded. For holotypes see also under heading of 3. Java new species concerned. The totals for species include 4. Malay Peninsula (+Singapore) males and females, unless mentioned otherwise. 5. Philippines RMNH National Museum of Natural History 6. continental Asia Naturalis, Leiden, The Netherlands

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BMNH The Natural History Museum, London, trap, 1 ex.; Danum Valley: Fld Centre envs, vii.1986, United Kingdom Malumphy, 200-300 m, multistr evergr forest, on fungi, NMWC National Museum of Wales, Cardiff, Unit- 1 ex.; Danum Valley: Nature Trail, 14-17.ix.1990, Davis #ad140-164, 100-250 m, multistr evergr forest, hu- ed Kingdom man excr trap, 1 ex.; Danum Valley: Segama River, MNHN Muséum National d’Histoire Naturelle, 17-20.v.1990, Davis #ad001-054, 100-250 m, multistr ev- Paris, France ergr forest, human excr trap, 4 exx.; Danum Valley: Segama River, 24-27.viii.1990, Davis #ad115-124, 100-250 m, multistr evergr forest, human excr trap, 1 ex.; Ula Segama 1. Ochicanthon edmondsi Reserve, 24-26.iii.1992, Davis #ad793, 100-250 m, forest logged in 1989, flight interception trap, 2 exx. Malay- sia, Sarawak: Mt Mulu, iii-v.1978, Hanski, 800-1700 m, Types. Malaysia, Sarawak: Mt Mulu, iii-v.1978, Hanski, lower montane rain forest, meat trap, 60 exx.; Mt Mulu, 800-1700 m, lower montane rain forest, carrion trap, 16 iii-v.1978, Hanski, 500-800 m, mixed dipterocarp forest, exx. incl. Holotype (RMNH). meat trap, 4 exx. Total 16 exx., 1 record (RMNH). Total 75 exx., 9 records (NMWC, RMNH).

2. Ochicanthon dytiscoides 5. Ochicanthon rombauti Malaysia, Sabah: Danum Valley: Fld Centre envs, vii.1986, Malumphy, 200-300 m, multistr evergr forest, on fungi, Types. Malaysia, Sabah: Crocker Range: Keningau- 2 exx.; Danum Valley: Fld Centre envs (W10-W15), Kimanis rd (km25), 18-23.xi.1987, Krikken & Rombaut 20-23.vii.1986, Malumphy #cm120, 100-200 m, multistr #sa44, 1300 m, multistr evergr forest edge, human excr evergr forest, chicken trap, 1 ex.; Danum Valley: Fld Centre trap, 4 exx. (RMNH); Crocker Range: Keningau-Kimanis envs (W3N3), 7-10.vii.1990, Davis #ad074-078, 100-250 rd (km25), 18-23.xi.1987, Krikken & Rombaut #sa45, m, multistr evergr forest, human excr trap, 1 ex.; Danum 1300 m, multistr evergr forest, human excr trap, 69 exx. Valley: Fld Centre envs (W5-W10/S), 12-15.vii.1986, incl. Holotype (RMNH). Malumphy #cm075, 100-200 m, multistr evergr forest, fish trap, 1 ex.; Danum Valley: Fld Centre envs (W5-W10/S), Total 73 exx., 2 records (RMNH). 12-15.vii.1986, Malumphy #cm089, 100-200 m, multistr evergr forest, fish trap, 1 ex.; Danum Valley: Fld Centre envs (W5-W10/S), 12-15.vii.1986, Malumphy #cm090, 100-200 m, multistr evergr forest, fish trap, 1 ex.; Danum 6. Ochicanthon crockermontis Valley: Segama River, 17-20.v.1990, Davis #ad001-054, 100-250 m, multistr evergr forest, human excr trap, 3 exx.; Types. Malaysia, Sabah: Crocker Range: Kota Kinabalu- Danum Valley: Segama River, 5-8.vii.1990, Davis #ad063- Tambunan rd (km64), 21-24.xi.1987, Krikken & 071, 100-250 m, multistr evergr forest, human excr trap, Rombaut #sa49a, 1150 m, multistr evergr forest, human 1 ex.; Kinabalu, [Waterstradt ?], 1 ex. Lectotype (MNHN); excr trap, 6 exx. incl. Holotype (RMNH). Ula Segama Reserve, 13-19.vii.1990, Davis #ad079-085, 100-250 m, forest logged in 1989, human excr trap, 1 ex.; Total 6 exx., 1 record (RMNH). Ula Segama Reserve, 13-16.ix.1990, Davis #ad167-173, 100-250 m, forest logged in 1989, human excr trap, 1 ex.; Ula Segama Reserve, 24-26.iii.1992, Davis #ad793, 100- 7. Ochicanthon crypticus 250 m, forest logged in 1989, flight interception trap, 1 ex. Brunei: Telisai, 22km SW of, 17-20.xi.1980, Edmonds, Types. Malaysia, Sarawak: Mt Mulu, iii-v.1978, Hanski, 50 m, forest, human faeces, 1 ex.; Telisai, 5km E of, 12- 100-500 m, mixed dipterocarp forest, fish trap, 28 exx. 30.xi.1980, Edmonds, 20 m, forest, human faeces, 2 exx.; (RMNH); Mt Mulu, iii-v.1978, Hanski, 100 m, alluvial Telisai, 5km E of, 12-30.xi.1980, Edmonds, 20 m, forest, forest, fish trap, 5 exx. (RMNH); Mt Mulu, iii-v.1978, fruit trap, 1 ex. Malaysia, Sarawak: Mt Mulu, iii-v.1978, Hanski, 100-500 m, mixed dipterocarp forest, baited trap, Hanski, 150 m, kerangas forest, fish trap, 21 exx. 16 exx. incl. Holotype (RMNH).

Total 40 exx., 16 records (MNHN, NMWC, RMNH). Total 49 exx., 3 records (RMNH).

3. Ochicanthon masumotoi 8. Ochicanthon peninsularis

Malaysia, Sabah: Danum Valley: Bole River, 23-25.ix.1987, Types. Malaysia, Peninsular: Pahang: Fraser’s Hill, Kirk-Spriggs #ks15-1, 100 m, multistr evergr forest, fish 29-3.x-xi.1977, Bendell, 3 exx. incl. Holotype (BMNH), trap, 1 ex.; Danum Valley: Bole River, 28-30.ix.1987, MNHN; Selangor: Bangi Universiti Kebangsaan, Kirk-Spriggs #ks25-1, 100 m, multistr evergr forest, fish 6.viii.1996, Chan Yoong Yoong, ground trap, 1 ex.

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(RMNH) Singapore: Bukit Timah NP, 7.vi.1976, Hoog- Davis #ad619-627, 100-250 m, multistr evergr forest, moed, high forest, human faeces, 1 ex. (RMNH). human excr trap, 3 exx. (RMNH); Danum Valley: Fld Total 5 exx., 3 records (BMNH, MNHN, RMNH). Centre envs (W3N3), 7-10.vii.1990, Davis #ad074-078, 100-250 m, multistr evergr forest, human excr trap, 1 ex. (RMNH); Danum Valley: Fld Centre envs (W3N3), 26-29.v.1991, Davis #ad344-348, 100-250 m, multistr 10. Ochicanthon neglectus evergr forest, human excr trap, 1 ex. (RMNH); Danum Valley: Fld Centre envs (W3N3), 21-23.i.1992, Davis Types. Malaysia, Sarawak: Mt Mulu NP: Mt Api, #ad640, 100-250 m, multistr evergr forest, flight intercep- iii-v.1978, Hanski, 400-600 m, limestone forest, fish trap, tion trap, 1 ex. (RMNH); Danum Valley: Fld Centre envs 12 exx. incl. Holotype (RMNH). (W3N3), 24-27.i.1992, Davis #ad660, 100-250 m, multistr evergr forest, flight interception trap, 4 exx. incl. refer- Total 12 exx., 1 record (RMNH). ence specimen (RMNH); Danum Valley: Fld Centre envs (W3N3), 27-30.i.1992, Davis #ad661, 100-250 m, multistr evergr forest, flight interception trap, 3 exx. 11. Ochicanthon dulitmontis (RMNH); Danum Valley: Fld Centre envs (W3N3), 30- 2.i-ii.1992, Davis #ad683, 100-250 m, multistr evergr for- Type. Malaysia, Sarawak: Mt Dulit, 18.x.1932, Hobby & est, flight interception trap, 3 exx. (RMNH); Danum Val- Moore, 1200 m, moss forest, 1 ex. Holotype (BMNH). ley: Fld Centre envs (W3N3), 2-5.ii.1992, Davis #ad684, 100-250 m, multistr evergr forest, flight interception trap, Total 1 ex. (male only), 1 record (BMNH). 2 exx. (RMNH); Danum Valley: Fld Centre envs (W3N3), 8-11.ii.1992, Davis #ad685-689, 100-250 m, multistr evergr forest, human excr trap, 1 ex. (RMNH); Danum 12. Ochicanthon cambeforti Valley: Fld Centre envs (W3N3), 28-31.iii.1992, Davis #ad803-812, 100-250 m, multistr evergr forest, human excr trap, 2 exx. (RMNH); Danum Valley: Fld Type. Indonesia, Kalimantan: SE Borneo: loc. unspeci- Centre envs (W8), 7-10.iv.1991, Davis #ad215-223, fied, 1 ex. Holotype (MNHN). 100-250 m, multistr evergr forest, fish trap, 1 ex. (RMNH); Total 1 ex. (male only), 1 record (MNHN). Danum Valley: Fld Centre envs (W8), 18-21.vii.1991, Davis #ad449-458, 100-250 m, multistr evergr forest, human excr trap, 3 exx. (RMNH); Danum Valley: Fld 13. Ochicanthon karasuyamai Centre envs (W8), 27-30.vii.1991, Davis #ad460-469, 100-250 m, multistr evergr forest, human excr trap, 1 ex. (RMNH); Danum Valley: Fld Centre envs (W8), Indonesia, Sumatra: Mt Leuser NP: Lawe Mamas, 24-27.i.1992, Davis #ad650-659, 100-250 m, multistr 11-16.ix.1983, Räisänen #hr32Cb, 900-920 m, lowland ev- evergr forest, human excr trap, 1 ex. (RMNH); Danum ergr forest, human excr trap, 3 exx. (RMNH); Mt Leuser Valley: Fld Centre envs (W8), 29-3.ii-iii.1992, Davis NP: Lawe Mamas, 16-22.ix.1983, Räisänen #hr30Cc, #ad738, 100-250 m, multistr evergr forest, flight intercep- 580-600 m, lowland evergr forest, human excr trap, 1 ex. tion trap, 1 ex. (RMNH); Danum Valley: Fld Centre envs (RMNH). (W8), 3-6.iii.1992, Davis #ad758, 100-250 m, multistr evergr forest, flight interception trap, 3 exx. (RMNH); Total 4 exx. (females only), 2 records (RMNH). Danum Valley: Fld Centre envs (W8), 6-9.iii.1992, Davis #ad759, 100-250 m, multistr evergr forest, flight interception trap, 1 ex. (RMNH); Keningau area: Nabawan 14. Ochicanthon woroae (site D), 15-17.xi.1987, Krikken & Rombaut #sa39, 450 m, podzol forest, human excr trap, 1 ex. (RMNH); Malaysia, Sabah: Danum Valley: Fld Centre envs Ula Segama Reserve, 14-17.vii.1990, Davis #ad086-092, (W0-W5), 3-6.vii.1986, Malumphy #cm009, 100-200 m, 100-250 m, forest logged in 1988, human excr trap, 1 ex. multistr evergr forest, fish trap, 1 ex. (NMWC); Danum (RMNH); Ula Segama Reserve, 28-31.viii.1990, Davis Valley: Fld Centre envs (W10), 18-22.ix.1987, Kirk-Spriggs #ad126-132, 100-250 m, forest logged in 1988, human excr #ks13-2, 260 m, multistr evergr forest, human excr trap, trap, 2 exx. (RMNH); Ula Segama Reserve, 16-19.iv.1991, 1 ex. (NMWC); Danum Valley: Fld Centre envs (W10), Davis #ad240-249, 100-250 m, forest logged in 1989, 9-12.v.1991, Davis #ad299-308, 100-250 m, multistr human excr trap, 1 ex. (RMNH); Ula Segama Reserve, evergr forest, human excr trap, 1 ex. (RMNH); Danum 3-6.viii.1991, Davis #ad481-490, 100-250 m, forest Valley: Fld Centre envs (W10), 1-4.viii.1991, Davis logged in 1989, human excr trap, 5 exx. (RMNH); Ula #ad471-480, 100-250 m, multistr evergr forest, human Segama Reserve, 9-12.ii.1992, Davis #ad693-702, 100- excr trap, 1 ex. (RMNH); Danum Valley: Fld Centre envs 250 m, forest logged in 1989, human excr trap, 2 exx. (W2-3), 20-24.x.1987, Krikken & Rombaut #sa07a, (RMNH); Ula Segama Reserve, 24-26.iii.1992, Davis 100-200 m, multistr evergr forest, human excr trap, 1 ex. #ad793, 100-250 m, forest logged in 1989, flight inter- (RMNH); Danum Valley: Fld Centre envs (W22), ception trap, 4 exx. (RMNH); Ula Segama Reserve, 29- 15-18.vi.1991, Davis #ad387-395, 100-250 m, mul- 1.iii-iv.1992, Davis #ad814, 100-250 m, forest logged tistr evergr forest, human excr trap, 2 exx. (RMNH); in 1989, flight interception trap, 2 exx. (RMNH); Ula Danum Valley: Fld Centre envs (W22), 9-12.ix.1991, Segama Reserve: Bole Kecil River, 9-12.iv.1992, Davis

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#ad817, 100-250 m, forest logged in 1981, flight intercep- 19. Ochicanthon danum tion trap, 1 ex. (RMNH); Ula Segama Reserve: Bole Kecil River, 12-15.iv.1992, Davis #ad818, 100-250 m, forest Types. Malaysia, Sabah: Danum Valley: Danum River, logged in 1981, flight interception trap, 1 ex. (RMNH). 27-30.viii.1991, Davis #ad533-542, 100-250 m, mul- Malaysia, Sarawak: Mt Mulu, iii.1978, Collins, 200 m, tistr evergr forest, human excr trap, 1 ex. (RMNH); Da- mixed dipterocarp forest, baited trap, 1 ex.; Mt Mulu, num Valley: Fld Centre envs (W0), 14-17.v.1991, Davis iii-v.1978, Hanski, 100-500 m, mixed dipterocarp forest, #ad312-321, 100-250 m, multistr evergr forest, human human excr trap, 3 exx.; Mt Mulu, iii-v.1978, Hanski, excr trap, 2 exx. (RMNH); Danum Valley: Fld Cen- 100-500 m, mixed dipterocarp forest, fish trap, 1 ex. tre envs (W3N3), 16-19.v.1992, Davis #ad871-880, Total 64 exx., 36 records (BMNH, NMWC, RMNH). 100-250 m, multistr evergr forest, human excr trap, 1 ex. (RMNH); Danum Valley: Mt Danum, 7-9.x.1987, Kirk- Spriggs #ks48-2, 293 m, multistr evergr forest, human 15. Ochicanthon tambunan excr trap, 1 ex. (NMWC); Danum Valley: Nature Trail, 28-1.vi-vii.1991, Davis #ad397-426, 100-250 m, multistr evergr forest, human excr trap, 1 ex. (RMNH); Danum Val- Types. Malaysia, Sabah: Crocker Range NP, 7-11.viii.1991, ley: Nature Trail, 6-9.ix.1991, Davis #ad586-614, 100-250 Kirk-Spriggs #ks53-91, 1463 m, multistr evergr forest, m, multistr evergr forest, human excr trap, 1 ex. (RMNH); montane, fish trap, 12 exx. (NMWC); Crocker Range NP, Danum Valley: Nature Trail, 18-21.iv.1992, Davis #ad821, 7-11.viii.1991, Kirk-Spriggs #ks54-92, 1249 m, multistr 100-250 m, multistr evergr forest, human excr trap, 2 exx. evergr forest, montane, human excr trap, 148 exx. incl. (RMNH); Danum Valley: Segama River, 17-20.v.1990, Holotype (NMWC); Crocker Range: Kota Kinabalu- Davis #ad001-054, 100-250 m, multistr evergr forest, hu- Tambunan rd (km56), 21-24.xi.1987, Krikken & man excr trap, 1 ex. (RMNH); Danum Valley: Segama Rombaut #sa48a, 1350 m, multistr evergr forest, human River, 5-8.vii.1990, Davis #ad063-071, 100-250 m, multi- excr trap, 4 exx. (RMNH); Crocker Range: Kota Kinabalu- str evergr forest, human excr trap, 1 ex. (RMNH); Danum Tambunan rd (km56), 21-24.xi.1987, Krikken & Valley: Segama River, 24-27.viii.1990, Davis #ad115-124, Rombaut #sa48b, 1350 m, multistr evergr forest, fish trap, 100-250 m, multistr evergr forest, human excr trap, 5 exx. 1 ex. (RMNH). (RMNH); Danum Valley: Segama River, 15-18.iv.1991, Total 165 exx., 4 records (NMWC, RMNH). Davis #ad229-239, 100-250 m, multistr evergr forest, human excr trap, 8 exx. (RMNH); Danum Valley: Segama River, 28-31.vi.1991, Davis #ad427-437, 100-250 m, multistr evergr forest, human excr trap, 14 exx. (RMNH); Da- 16. Ochicanthon mulu num Valley: Segama River, 3-6.ix.1991, Davis #ad565-575, 100-250 m, multistr evergr forest, human excr trap, 2 exx. Types. Malaysia, Sarawak: Mt Mulu, iii-v.1978, Hanski, (RMNH); Danum Valley: Segama River, 31-3.i-ii.1992, 150 m, kerangas forest, fish trap, 8 exx. (RMNH); Mt Davis #ad672-682, 100-250 m, multistr evergr forest, hu- Mulu, iii-v.1978, Hanski, 150 m, kerangas forest, human man excr trap, 5 exx. (RMNH); Danum Valley: Segama excr trap, 4 exx. incl. Holotype (RMNH). River, 15-18.ii.1992, Davis #ad716, 100-250 m, multistr evergr forest, flight interception trap, 1 ex. (RMNH); Total 12 exx., 2 records (RMNH). Danum Valley: Segama River, 8-11.v.1992, Davis #ad838-848, 100-250 m, multistr evergr forest, human excr trap, 11 exx. (RMNH); Ula Segama Reserve, 17. Ochicanthon gangkui 16-19.iv.1991, Davis #ad240-249, 100-250 m, forest logged in 1989, human excr trap, 3 exx. (RMNH); Ula Segama Reserve, 30-2.v-vi.1991, Davis #ad351-360, Malaysia, Sabah: Kinabalu NP: Silau Silau (canteen 100-250 m, forest logged in 1978, human excr trap, slope), 16-23.i.1986, Krikken #pw86a, 1540 m, multistr 1 ex. (RMNH); Ula Segama Reserve, 3-6.viii.1991, Davis evergr forest, fish trap, 9 exx.; Mt Kinabalu, 24-25.ix.1977, #ad481-490, 100-250 m, forest logged in 1989, hu- Bacchus, 1550 m, dung trap, 1 ex.; Trus Madi, 24.viii.1977, man excr trap, 20 exx. (RMNH); Ula Segama Reserve, Bacchus, 1200 m, dung trap, 1 ex. 9-12.ii.1992, Davis #ad693-702, 100-250 m, forest logged Total 11 exx., 3 records (BMNH, RMNH). in 1989, human excr trap, 11 exx. (RMNH); Ula Segama Reserve, 14-17.ii.1992, Davis #ad705-714, 100-250 m, forest logged in 1989, human excr trap, 1 ex. (RMNH); Ula Segama Reserve: Bole Kecil River, 5-8.iii.1992, Davis 18. Ochicanthon kimanis #ad748-757, 100-250 m, forest logged in 1981, human excr trap, 2 exx. (RMNH); Ula Segama Reserve: Bole Kecil Types. Malaysia, Sabah: Crocker Range: Keningau- River, 22-25.iii.1992, Davis #ad783-792, 100-250 m, for- Kimanis rd (km25), 18-23.xi.1987, Krikken & Rombaut est logged in 1981, human excr trap, 29 exx. incl. Holo- #sa44, 1300 m, multistr evergr forest edge, human excr type (RMNH); Ula Segama Reserve: Bole Kecil River, trap, 1 ex. (RMNH); Crocker Range: Keningau-Kimanis 14-17.v.1992, Davis #ad861-870, 100-250 m, forest rd (km25), 18-23.xi.1987, Krikken & Rombaut #sa45, logged in 1981, human excr trap, 3 exx. (RMNH); Ula 1300 m, multistr evergr forest, human excr trap, 10 exx. Segama Reserve: Bole River, 5-8.viii.1991, Davis #ad493- incl. Holotype (RMNH). 502, 100-250 m, forest logged in 1981, human excr trap, Total 11 exx., 2 records (RMNH). 5 exx. (RMNH). Total 132 exx., 25 recordsDownloaded (NMWC, from RMNH). 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Excluded from type series. Malaysia, Sabah: Danum Index to species names Valley: Mt Danum, 7-9.x.1987, Kirk-Spriggs #ks52-2, 1093 m, multistr evergr forest, human excr trap, 1 ex. aphodioides, Onthophagus ...... 470 Total 1 ex. (male only), 1 record (NMWC). cambeforti, Ochicanthon ...... 426, 427, 448, 477 cingalensis, Ochicanthon ...... 426 crockermontis, Ochicanthon . . . . 426, 436, 476 21. Ochicanthon javanus crypticus, Ochicanthon ...... 426, 427, 438, 476 danum, Ochicanthon ...... 426, 427, 458, 478 Types. Indonesia, Java: Mt Puncak: Telaga Warna, denticollis, Onthophagus ...... 470 27-30.xii.1985, Krikken #pw71a, 1600 m, multistr evergr deplanatus, Ochicanthon ...... 426 forest, human excr trap, 16 exx. (RMNH); Mt Puncak: dulitmontis, Ochicanthon . . . 426, 427, 446, 477 Telaga Warna, 27-30.xii.1985, Krikken #pw71b, 1600 dytiscoides, Ochicanthon . . . 426, 427, 430, 476 m, multistr evergr forest, fish trap, 16 exx. incl. Holotype (RMNH); Mt Puncak: Telaga Warna, 15-17.iii.1986, van edmondsi, Ochicanthon ...... 426, 427, 448, 476 Berge Henegouwen, 1500 m, submontane forest, hu- fallcilaetum, Phacosoma ...... 426 man excr trap, 3 exx. (RMNH); Mt Salak: N slope, 28- gangkui, Ochicanthon . . . . 426, 427, 456, 478 31.xii.1985, Krikken #pw72a, 900 m, multistr evergr hanskii, Ochicanthon ...... 426, 463, 479 forest, human excr trap, 3 exx. (RMNH); Mt Tangkuban hikidai, Ochicanthon ...... 426, 466 Prahu, iv.1937, Drescher, 1200-1500 m, 1 ex. (RMNH). javanus, Ochicanthon ...... 426, 459, 479 Total 39 exx., 5 records (RMNH). karasuyamai, Ochicanthon . . 426, 427, 448, 477 kikutai, Ochicanthon ...... 426, 427, 443 Excluded from type series. Indonesia, Java: Banyu- kimanis, Ochicanthon . . . . 426, 427, 456, 478 wangi, 1941, MacGillavry, 1 ex.; Ijen Plateau: SE slope, laetus, Ochicanthon ...... 426 23-1.ii-iii.1996, Huijbregts, 1250 m, multistr evergr for- loebli, Ochicanthon ...... 426 est, fish trap, 1 ex.; Ijen Plateau: SE slope, 23-1.ii-iii.1996, maryatiae, Ochicanthon ...... 426, 427, 434 Huijbregts, 1250 m, multistr evergr forest, human excr trap, 1 ex. masumotoi, Ochicanthon ...... 426, 427, 432, 476 mulu, Ochicanthon ...... 426, 455, 478 Total 3 exx., 3 records (RMNH). neglectus, Ochicanthon ...... 426, 427, 443, 477 nitidus, Ochicanthon ...... 426 obscurus, Ochicanthon ...... 426 22. Ochicanthon hanskii ochii, Ochicanthon ...... 426 parantisae, Ochicanthon ...... 426, 427, 459 Types. Malaysia, Sarawak: Mt Mulu, iii-v.1978, Hanski, peninsularis, Cassolus ...... 470 1700-2180 m, upper montane rain forest, fish trap, peninsularis, Ochicanthon . . . . . 426, 427, 439, 476 246 exx. (RMNH); Mt Mulu, iii-v.1978, Hanski, philippinensis, Ochicanthon ...... 426 800-1700 m, lower montane rain forest, carrion trap, punctatus, Ochicanthon ...... 426, 466, 470, 479 129 exx. incl. Holotype (RMNH); Mt Mulu, iii-v.1978, Hanski, 500-800 m, mixed dipterocarp forest, carrion rombauti, Ochicanthon . . . . 426, 427, 435, 476 trap, 1 ex. (RMNH); Mt Mulu: Site H, v-viii.1978, Ham- rorarius, Onthophagus ...... 470 mond & Marshall, 1650 m, lower montane forest, litter, rudis, Onthophagus ...... 470 1 ex. (BMNH); Mt Mulu: Site I, v-viii.1978, Hammond simboroni, Ochicanthon ...... 426, 427, 448 & Marshall, 1860 m, upper montane forest, litter, 2 exx. tambunan, Ochicanthon . . . 426, 427, 451, 478 (BMNH); Mt Mulu: nr Camp 1, v-viii.1978, Hammond thai, Ochicanthon ...... 426 & Marshall, 150-200 m, flood refuse, 1 ex. (BMNH); Mt Mulu: nr Camp 4, v-viii.1978, Hammond & thailandicus, Ochicanthon ...... 426 Marshall, 1800 m, upper montane forest, human faeces, tristis, Ochicanthon ...... 426 7 exx. (BMNH). tristoides, Ochicanthon ...... 426 vulpes, Onthophagus ...... 470 Total 387 exx., 7 records (BMNH, RMNH). woroae, Ochicanthon . . . . . 426, 427, 450, 477

24. Ochicanthon punctatus

Type. Indonesia, Sulawesi: Tondano, 1899, Collector C, 1 ex. Holotype (MNHN). Total 1 ex. (female only), 1 record (MNHN) [assumed to be mislabelled]. Total 20 taxa, 1108 exx.

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