Tijdschrift voor Entomologie 156 (2013) 35–56 brill.com/tve

Oriental Bolboceras: Introduction to the and of the nigricans group of species (Coleoptera: : ) J. Krikken

The taxonomy of the genus Bolboceras Kirby, 1819 (= Indobolbus Nikolajev, 1979) is discussed. The Oriental members of the genus as currently conceived have not been revised for the past 160 years. Operational groups of species are proposed, including the Bolboceras nigricans group. In this group B. nigricans Westwood, 1848 and five new Oriental group members are here keyed, diagnosed, and illustrated. The new species proposed are: Bolboceras schulzei (Bangladesh, India), B. malabaricum (India), B. extraneum (Bangladesh, India, Pakistan), B. insulare (Sri Lanka), and B. mimicans (Myanmar). In an Appendix a single new species is described in the Bolboceras perpunctatum group, to consolidate the key to the Bolboceras species groups – B. perpunctatum (India). A list of names of all Bolboceras species is given, indicating a formal retransfer of some from Indobolbus to Bolboceras, following a 2006 ruling of the ICZN on the use of the name Bolboceras. Keywords: Coleoptera, Geotrupidae, Bolboceras, genus description, nigricans species group, keys, list of species, recombinations, new species, Oriental Region. J. Krikken, Naturalis Biodiversity Center/National Museum of Natural History, PO Box 9517, NL-2300 RA Leiden, The Netherlands. [email protected]

Introduction head, pronotum, and of male genitalia, indispensable Bolboceras Kirby, 1819 (= Indobolbus Nikolajev, for species identification, are absent or insufficiently 1979) is one of the genera of Bolboceratinae (“bol- detailed. The original material of some of the Bol- bos”, subcosmopolitan subfamily, with ca 600 species boceras species described by Westwood (1848, 1852) in ca 50 genera) still lacking a modern treatment. was not recovered, while both Westwood himself and For the past 160 years hardly anything has been his contemporaries omitted, overlooked, or mixed added to our knowledge of the Oriental fauna of up some available names; their pictures and descrip- this genus as conceived since the ICZN (2006) rul- tions can hardly be qualified as taxonomically use- ing. I looked at Bolboceras (as currently conceived) ful by today’s standards. The identities of some ap- long ago, and, when recently deciding on another at- parently non-congeneric 19th century species-group tempt at clarifying the taxonomy of the genus, I ex- names combined with the name Bolboceras need clar- perienced again that this is, and will remain, a tax- ification. All in all, ca 30 genuine named and un- onomically resistant group. Several species are virtu- named Bolboceras species, mainly Oriental, are now ally unidentifiable without access to both types and known to me, but the actual number may be higher, informative series (i.e. adequately representing in- particularly among the relatives close to B. quadri- traspecific variation and, where applicable, sexual di- dens (Fabricius, 1781) and B. transversale Westwood, morphism). In publications comparative pictures of 1848.

Tijdschrift voor Entomologie 156: 35–56. [ISSN 0040-7496]. brill.com/tve © Nederlandse Entomologische Vereniging. Published by Koninklijke Brill NV, Leiden. Published 15 July 2013. DOI 10.1163/22119434-00002018

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As implied above, in spite of these drawbacks, I am may well have evolved in the subcontinent, in a way not throwing in the towel yet. Some handle on the similar to what has been hypothesized for their rel- taxonomy of Oriental Bolboceras is needed, because atives in Bolbohamatum Krikken, 1980 and other at this moment nothing meaningful can be done for reputedly ancient, Gondwana-based scarab groups the evident lack of verified species names and associ- (the out-of-India hypothesis of some authors, e.g., ated diagnostic information. As a first contribution, Davis in Scholtz et al. 2009). Contrary to Bolbo- therefore, the genus is here defined, provisionally hamatum (13 species in South Asia), which has one subdivided into operational clusters of species, and derived polytypic species ranging from Myanmar to six Oriental species (B. nigricans Westwood, 1848 Java and China, species of Bolboceras have not (yet) plus five new relatives) are keyed and diagnosed in been recorded from regions East and North of Myan- what is called the nigricans group. The species in this mar. There is one record of Bolboceras from the An- group are characterized by K- or X-patterned peri- daman Islands (Chandra 1996). The presence of marginal clypeal ridges culminating in a single an- congeners and other closely related bolbos in sub- teromedian point (Figs 27–32), as opposed to the Saharan Africa (Bolboceratops Krikken, 1978, Bolbaf- roughly trapeziform clypeal surface seen in the bulk froides Nikolajev, 1979, etc., cf. Krikken 1984), and of Bolboceras, and in larger collections usually stand- the absence of bolbos from Madagascar, may indi- ing under B. quadridens as an evidently mixed lot cate that there is more to their historical biogeogra- (see also key to groups of species below, couplets phy than a simple Gondwana-India explanation (cf. 3 and 4). Although the known morphological di- also discussion of Bolboceratops in Carpaneto et al. versity in the nigricans group is adequately repre- 1993). sented in this paper, the taxonomic conclusions can- All these genera seem to evade extensive areas of not yet be final for all taxa, the trivial reason being close multistratal evergreen forest; some of the Bol- that more material is needed. There are in Bolboceras boceras at hand are from upland regions, like the questionable cases of compound polymorphism (ge- higher Himalayan valleys (elevations over 1500 m, cf. ographic + individual [size-related] + sexual, etc.), new species in Appendix, below). The precise role of which cannot be solved without sufficiently infor- Bolboceras in their preferred ecosystems is unknown, mative series, particularly from a larger number of only few significant label data being available. In localities than is currently the case. Local hybridiza- plain savanna in East Africa I collected small series of tion cannot be excluded – I was, for instance, puzzled two Bolboceras species and numerous other bolbos at by the variation in a collection of small forms (cf. B. light, during the rainy season. They all buzz around extraneum sp.n.) from Pusa (Bihar, India) and other the lamp at an audibly higher pitch than do the non- northeastern localities. The application of molecular bolbos among incoming medium-sized scarabs, their techniques could perhaps clarify these puzzles in the arrival thus being announced immediately. Our col- not too distant future. leagues in Asia, particularly India, might be inter- A premise in the intrageneric group distinctions is ested to find out the details of their habits, as noth- that polymorphism in the basic shape of both clypeus ing of substance is known. Bolboceras, like all family and pronotum is indeed limited, but, as experienced members, are certainly ardent soil tunnellers, but in other bolboceratine genera, exceptions cannot be what exactly they feed and breed on (fungi? vegetable ruled out (cf. apparent sexual dimorphism in B. in- debris, humus? dung, in combination with fungi?), sulare sp.n.). Note that members of certain other ge- and how they do this, is still a mystery (similar to nera, i.e. outside the Oriental Region, may have X- what is known of certain North American and Aus- shaped clypeal ridges and/or an anteromedian clypeal tralian bolbos? – cf. papers by Howden et al. between protrusion similar to those seen in the B. nigricans 1955 en 2007, Houston & Bougher 2010, Houston group, but none of these has the full set of other 2012). Remember always to collect and safely store generic features described below. Following this ac- voucher specimens while doing ecological work, else count of the nigricans group, work on other groups naming the species concerned will be virtually im- in Bolboceras and other bolbos is envisaged (see also possible and less progress will be made. Krikken 2011). Apart from the five or six named species recorded Discussion of generic delimitation, from the Palaearctic and Afrotropical Regions, the nomenclature, and position genus Bolboceras, in the sense of the 2006 ruling of Members of the genus Bolboceras Kirby, 1819, the ICZN (Opinion 2138), appears essentially Ori- judged from hundreds of specimens seen, are eas- ental, the majority of the species known to me living ily distinguished from other Oriental bolbos by the in the Indian subcontinent – this then is the geo- combination of the following morphological charac- graphic focus of the present treatment. The genus ters:

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(1) basic outline of clypeus and its perimarginal ica (and these were during most of the past 60 years ridges usually subtrapeziform (modified in combined with Bolboceras), one is in Europe. the nigricans group); During the past decade it has been argued that (2) shape of eye canthi laterally rounded, with bolbos should be placed in a family of their own variably developed anterior protrusion (or (, cf. Scholtz & Grebennikov 2006 angle); and included references). I take, with others, like (3) presence of long transverse interocular ridge; Verdú et al. (2004) and Li et al. (2008), a more (4) basically quadrituberculate (trisulcate) pro- conservative view, considering the jury still out. This notum; also holds for the internal suprageneric taxonomy of (5) pronotal base (medially) marginate; the bolbos: the group (the proposed family) is mor- (6) elytral stria 1 (of 7 between suture and phologically diverse, nomenclaturally confusing, and humeral umbone) not reaching base of possibly better subdividable into tribes and subtribes elytron; than is currently the case. Outsiders are just led astray (7) separation of mesocoxae by relatively simple when looking at the insufficiently explained applica- (non-hooked) metasternal process; tion of (sub)tribal names coined from 1970 on, now (8) structure of antennal club, including lack of compounded by the Bolboceras- ruling (cf. grooves on outer side of last lamella; recent incorrect usage of tribal names in Král et al. (9) structure of aedeagus (including set of stalks 2006; latest situation reviewed by Smith in Bouchard between parameres); et al. 2011; cf. also Krell et al. 2004). Irrespective (10) sexual dimorphism usually limited. of current conclusions, Bolboceras as here defined is In several Bolboceras species the four pronotal the type-genus of its family-group name, and any protrusions may be noticeable only as indistinct intra-subfamily division must take some morpholo- swellings separated by three shallow, punctate depres- gical character states of the genus into account, for sions (i.e. also in “major” individuals of the species instance, the separation of the mesocoxae, the pat- concerned). All Bolboceras agree with the majority tern of elytral striae, and details of the antennal club of Bolboceratinae in being generally semiglobular (additional comments in the generic account below). and unicolour yellow to brown. Among the Bol- boceratinae the peculiar complex structure of the aedeagus appears unique to (in phylogenetic terms, Material and methods likely an autapomorphy of) the genus Bolboceras (cf. In Bolboceras intraspecific and interspecific differ- also Comments, under the generic description). The ences in body size, and development of pronotal pro- metasternal process between the mesocoxae never has trusions and impressions, remain difficult to disen- a fine recurved hook in front, in which Bolboceras dif- tangle, but good series should be informative about fers from Bolbohamatum. polymorphism, B. nigricans being a case in point. To It should be re-emphasized here that the name caution other workers I deliberately left two morphs Bolboceras is now reserved for the species just charac- of nigricans separated in the dichotomous key and terized, being no longer applicable to the Holarctic the synoptic table given below. The criteria to join genus Odonteus Samouelle, 1919 (= Odontaeus,an morphs in a single species include similar aedeagal old, widespread emended spelling) since the recent shape and other salient characters. Only where size ICZN ruling on type species designations already in a series from a particular locality seems constant, mentioned above. In the bolboceratine spectrum of this size and potentially related character differences morphological diversity the two genera, as now con- are considered of diagnostic significance. Sexual di- ceived, could hardly be more different. I consider the morphism indeed appears unusual and slight in Bol- designation of Scarabaeus quadridens Fabricius, 1781 boceras, but, as mentioned above, here also caution as type species of Bolboceras in this ruling unfortu- is warranted (cf. B. insulare sp.n.). Individual colour nate, having preferred any option completely dis- appears only slightly variable; differences may be due pelling the ambiguously applied name Bolboceras – to manner of preservation, or specimen age. A selec- but so be it. tion of 19 diagnostic characters relevant to the nigri- In passing, it may be noted that the known range cans group is listed in Table 1. of the genus Odonteus has only recently been ex- The holotype descriptions of the new species are panded just into the region here under consider- kept short, additional details being better shown by ation, i.e. by the formal description of two Asian the technical pictures. Combining the detailed pic- species, from altitudes around 2000 m in North Pak- tures of the heads (Figs 27–32) and the male geni- istan and North India, respectively (Carpaneto & talia (Figs 33–39) should be particularly helpful in Mignani 2005, Mittal 1998), and more are known. identifying nigricans group species. The terminology Most other Odonteus species occur in North Amer- of externally visible parts of the aedeagi is explained

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Table 1. Synoptic guide to species in the nigricans group.

Characters and character states 1. Perimarginal ridges of clypeus (full-face) 11. Shape of discoparamedian protrusions 1. More K-shaped 1. Mere callosities 2. More X-shaped 2. Distinctly protuberant 3. Trapeziform (in female) 3. Conical (pointed) 2. Anteromedian cross-point of male clypeal ridges 12. Discomedian impression of pronotum posteriorly 1. Hardly protuberant 1. At most moderately defined 2. Protuberant, simply angular 2. Well-defined 3. Protuberant, transverse, rounded 13. Discolateral impressions of pronotum posteriorly 3. Height of interocular elevation on frons 1. At most moderately defined 1. High 2. Well-defined 2. Low 14. Discolateral impressions of pronotum 4. Crest of interocular elevation 1. Very shallow 1. Over half interocular distance long 2. Moderately shallow 2. Up to half interocular distance long 3. Deep 5. Crest of interocular elevation (axial view) 15. Anterolateral point of pronotum 1. In no way tri-undulate/tuberculate 1. (virtually) rectangular 2. Tri-undulate/tuberculate (medially slightly raised) 2. acute 6. Anterolateral edge of eye canthus 16. Density of primary punctation above shallow lateral pronotal fovea 1. Angular, at most slightly elevated 1. Abundant 2. Angulate-tuberculate 2. Sparse 7. Distal (curved) lobe of eye canthus 17. Tip of lateral stalks on median apparatus of aedeagus (lsas, Fig. 3) 1. Shorter than proximal part 1. Straight or slightly curved, apex blunt 2. Equal to or longer than proximal part 2. Robustly hooked, recurved 8. Depression at base of anterior pronotal declivity (amd in Fig. 2) 3. Finely hooked, recurved 1. Virtually absent 18. Parameral tip 2. Slight 1. Acuminate 3. Very distinct 2. Blunt 9. Discomedian impression of pronotum 19. Body length in mm usually 1. Shallow or indistinct 1. Less than 9 mm 2. Deep 2. 9 mm or over 10. Discoparamedian protrusions on pronotum 1. Low 2. High

Characters Species nigricans large small schulzei malabaricum extraneum insulare mimicans

1111122/31 233322-313 31211221 41112122 51112122 62111112 71122222 811-232223 92111112 10 2 2 1 1 1 1 2 11 3 3 2 1 1 1 2 12 2 1 1 1 1 1 1 13 2 1 1 1 1 1 1 14 3 2 2 1 1 1 2 15 1 1 1 2 1 1 1 16 1 1 2 2 1 1 1 17 2 2 1 3 3 3 2 18 1 1 1 1 1 2 1 19 2 2 2 2 1 1-2 2

/ slash separating presence of two discrete states, - hyphen indicating transitional states in Fig. 3. Extruded aedeagi (Figs 35, 39) cannot eas- gical terminology relevant to the characters of head ily be forced into a resting position or vice versa,and and pronotum is explained in Figs 1–2. our pictures all give the situation as is (handling with Punctation densities are roughly classified as pins/pincers may result in damage). The morpholo- sparse (punctures mostly separated by >5 punctural

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Figs 1–4. 1–3, Bolboceras, generalized schemes, with terminology, of 1, head, full-face view (mandibles omitted), 2, pronotum, dorsal view, 3, aedeagus, upper side. – Fig. 4, B. perpunctatum, holotype male, head, full-face view. Abbreviations: amd anteromedian depression of pronotum dpmp discoparamedian protrusion (of pronotum) b backward lateral perimarginal ridge ec eye canthus of clypeus ga genal angle (in front of eye canthus) bm basomedian (part of pronotum) ior interocular ridge bp basal piece (of male genitalia) lat lateral part (of pronotum) c central perimarginal ridge of clypeus lpf lateral pronotal fovea clyp clypeus lsas lateral sclerotized aedeagal stalk dlc discolateral concavity (of pronotum) maa (upper lobe of) median aedeagal apparatus dlp discolateral protrusion (of pronotum) pmas paramedian aedeagal struts dmc discomedian concavity (of pronotum) pol paraocular line/ridge diameters), abundant (2–5 diameters), dense (1–2 off to 1/2 (0.5) mm, body part measurements to diameters), or crowded (<1 diameter); close means 0.1 mm. Measurements of body parts are given that various types of punctures on a particular sur- in the species accounts (holotypes, other selected face are separated by less than twice their diame- specimens), and, consequently, scale bars are omitted ters. Minute (for punctures) means that punctation is from the figures. present, but hardly distinct at less than ×40 magnifi- To facilitate administrative specimen recognition cation. Multi-sized means, that, contrary to a double several locality names follow the label text; a few punctation, punctural diameters on a particular sur- older locality names on handwritten labels remained face vary, but do not show two distinct size classes. obscure (or were absent at all). Additions and/or Marginate means: with distinctly demarcated, more adaptations in brackets. Some labels gave, other data or less raised, narrow marginal surface. aside, only initials of the collector. Measurements must be interpreted with caution in these semiglobular specimens; body parts were Collections measured through the microscope with a calibrated The Bolboceras material used for this nigricans group ocular micrometer scale. Total lengths are rounded review (ca 100 specimens) and for the introduc-

Downloaded from Brill.com10/08/2021 04:54:17PM via free access 40 Tijdschrift voor Entomologie, volume 156, 2013 tory matrix text came from the following collections: usually not sharply separated from (generally sim- ple) tempora, not retuse along pronotal anterome- BMNH The Natural History Museum, dian edge. Anterior edge of eye canthus more or London, UK less straight proximally, thence rounded, usually with CJSB Collection J. Schulze, Berlin, anterolateral angle (strongly protuberant in larger Germany species); distal tip usually rounded (extension of dis- CMNC Canadian Museum of Nature, tal lobe varies according to species); eye canthus Ottawa, Canada and temporal area separated (eyes not completely di- CNMS National Museum, Colombo, Sri vided); dorsally visible area of eye relatively large. Lanka Paraocular ridge delimiting eye canthus from frontal IRSNB Institut Royal des Sciences Naturelles disc, longitudinal, running posteriorly to, or slightly de Belgique, Brussels, Belgium beyond, level of transverse ridge. MHNG Musée d’Histoire Naturelle, Geneva, Pronotum and scutellum. Pronotum with variably Switzerland distinct pairs of discoparamedian and discolateral MNHN Muséum National d’Histoire protrusions (usually four protrusions or callosities Naturelle, Paris, France in a more or less transverse line), separated in most NMPC National Museum, Prague, Czech species by (three) intervening impressions (one disco- Republic median, one discolateral on either side); anterior de- OUMNH Oxford University Museum of clivity never very broadly concave. Pronotum lacking Natural History, Oxford, UK crest just behind apex, apical margin proper raised, PIME PIME Entomological Museum, but not particularly modified; basal area of pronotum Detroit MI, USA also lacking particular protrusion(s); pronotal apex RMNH Naturalis Biodiversity lacking pair of small, deep postocular foveae (though Center/National Museum of Natural heavy concentrations of punctures in shallow de- History, Leiden, The Netherlands pression present in some species). Fovea on lateral SMF Senckenberg Museum, Frankfurt, declivity of pronotum usually slight. Pronotal base Germany marginate (delimiting ridge may be reduced to row SMTD Staatliches Museum für Tierkunde, of punctures at position of scutellum); midline on Dresden, Germany basomedian part of pronotum punctate, not deeply ZMHB Zoological Museum, Humboldt sulcate. Apical edge of pronotum at most fringed University, Berlin, Germany with narrow velum. Posterolateral section of prono- ZSM Zoologische Sammlungen des tal border widely, evenly curved, anterolateral angle Bayrischen Staates, Munich, Germany distinct. Scutellum broad, with sides evenly curved along elytral side (usually slightly sinuate outward at base), apex rounded; relatively short (length/width Genus Bolboceras Kirby, 1819 ratio ca 1, in some species distinctly less); general Description surface of scutellum even, scutellar base and margins Head. Outline of mandibles asymmetrical; outer unmodified. margin of right mandible with rounded lobe (de- Elytron. Elytral base immarginate (not raised-delim- limited by incision). Anterior edge of labrum bis- ited), epipleuron gradually narrowing to and reach- inuate to straight, transverse ridge on surface occa- ing apicosutural angle; disc with 7 punctate striae be- sionally lost in rugulate-punctate sculpture. Outline tween suture and humeral umbone, plus 7 more lat- of clypeal surface (in full-face view) subtrapeziform erally. Striae 3–7 (nearly) extending to elytral base, (longer transverse anterior edge, anterolateral angles stria 1 terminating at side of scutellum, at some dis- distinct), in some species anteromedially more or less tance from base, 2 may be more or less effaced near produced (i.e. perimarginal ridges of clypeal surface base. Striae all (at most) very shallowly impressed, X- or K-patterned in full-face view, forming antero- strial punctures fine, usually distinct; discal inters- median protrusion, nigricans group); anterior edge of triae all equally and at most slightly convex. Humeral clypeus never widely arcuate (anterolateral angles of angle of elytron unmodified. Alae normally devel- clypeal surface not rounded); clypeal disc or clype- oped. ofrontal transition lacking elevation(s); clypeogenal Antenna. Antenna with 11 antennomeres. Antennal angle protruding. Frons with straight transverse in- club not remarkably thickened (not “inflated”); sur- terocular ridge only (length at least half interocular face slightly, evenly convex on both distal and prox- distance, in some species reaching paraocular ridge), imal sides (no grooves). Outline of club generally surrounding surface lacking distinct impression(s), at oval; proximal surface of club antennomere 1 partly most shallowly concave. Vertex lacking elevations,

Downloaded from Brill.com10/08/2021 04:54:17PM via free access Krikken: The Bolboceras nigricans group 41 glabrous, this area gradually passing to pubescent 2011), and the inaequale group reputedly reaches parts (i.e. not delimited by surface feature). Senegal (Paulian 1941). Pectus (ventral parts of thorax). Preprosternal para- median costae distinct, intercostal surface generally Comments slightly convex or flat. Prosternum lacking conspicu- Nikolajev (1979) published the name Indobolbus ous posteromedian protrusion(s), not even a longitu- (type species Scarabaeus quadridens Fabricius, 1781), dinal ridge, not bulbose or otherwise inflated-dilated. assuming, understandable at the time, congeneric Mesocoxae distinctly separated, although interven- type species for both Bolboceras Kirby, 1819 and ing anteromedian process of metasternum rather nar- Odonteus Samouelle, 1819; Krikken (1984) further row, its surface flat or nearly so, with fine peri- implemented Nikolajev’s proposal. This has all been marginal ridge, abruptly retuse in front to mesoster- overruled by the ICZN (2006, see Introduction, num, lacking any particular accessory protrusions. above). The name Bolboceras is to be used as neuter Metasternal disc more or less rhombic in outline. (Krell et al. 2003). From the explicit lectotype des- Legs. Protibia gradually dilated to apex, with oblique ignation by Krell (l.c.) it is clear that B. quadridens pointed external denticles (5–6), strongly dimin- is not a member of the nigricans group as here con- ishing in size proximad; terminal spur of protibia ceived; it has a subtrapeziform clypeus. long, more or less tapering. Protarsomere 1 not elon- Apart from world lists of bolboceratine characters gated. Fossorial elevations on outer side of meso- (Krikken 1977, Howden & Cooper 1977), the above and metatibiae with (bi)dentate, (bi)lobate spinifer- revised generic diagnosis of Bolboceras is based partic- ous crest; both tibiae, apex aside, with single robust ularly on comparisons with other Eurasian supraspe- anteapical elevation, others (proximally) usually no- cific taxa (see also Li et al. 2008). Bolboceras belongs ticeable only as short denticles (their size decreasing in the group of genera with eyes not completely di- proximad). Femora unmodified (profemur not den- vided, anterior clypeal outline not convex-arcuate, tate). and mesocoxae separated (group also including Bol- Genitalia. Aedeagus (Fig. 3) showing pair of moder- bohamatum, Bolbaffroides,andmore)–i.e.thetribe ately sclerotized, usually tapering (outer) parameral or subtribe Bolboceratini/Bolboceratina, depending sheaths (pm in Fig. 3), median apparatus modified, on the overall family-group classification, as dis- i.e. with accessory elements, including a pair of scle- cussed in the Introduction. The groups of species rotized lateral stalks (lsas), movable at base, their tip within Bolboceras (as keyed below) are tentative, il- varying in shape (stalks usually resting in parameral lustrating some supraspecific features of the mor- sheaths, standing out almost rotor-like when aedea- phological diversity. Although this does not immedi- gus extruded). Distal abdominal segments (pygid- ately affect the taxonomy of the nigricans group, the ium) not particularly modified. identity of B. transversale Westwood, 1848, stand- Sexual dimorphism. Slight or absent, possibly with ing for one of the other groups, will need clarifi- some exceptions (see under Comments); outline and cation (type not located, we have seen species like ornamentation of head and pronotum of male and it). Whatever the outcome of an eventual phylo- female usually very similar. genetically supported reclassification, there can be Colours, pilosity, size. Colour more or less uni- no doubt that the species in the various operational colour yellow-brown to brown (apart from in- groups are all closely interrelated, that they must be fuscated ridges, sutures, tips, margins, punctures, kept in one genus, and that this genus may well be etcetera), forebody may be slightly darker than ely- monophyletic. tra. Dorsal side virtually glabrous. Body length usu- While the unique structure of the male genitalia ally 6–16 mm. of Bolboceras supports a monophyly of the genus, an Immature stages. Unknown. in-depth analysis of the homologies of the aedeagal components (and more) might yield new insights, Range with likely consequences for the generic phylogeny Indian subcontinent from Myanmar to Pakistan, of the Bolboceratinae in general and the position of including Himalayas (Sikkim, Ladakh), Near East Bolboceras in particular. In recent papers on other ge- (Beirut: one specimen reported), sub-Saharan Africa nera, comprising potential relatives in the Afrotrop- (northern and eastern parts). Total of 17 species ical fauna (Carpaneto et al. 1992, 1993, Cambe- described (including six new in this paper); total fort 1998, Gussmann & Scholtz 2000), the posi- seen presumably ca 30. The B. nigricans group is tion of these genera vis-à-vis Bolboceras were not known from Myanmar to Pakistan (possibly also in explicitly discussed; Krikken (2011) briefly reviewed East Africa, see under B. nigricans). In Africa the the Afrotropical situation. Socotrabolbus Cambefort, quadridens group occurs in the Northeast (Krikken 1998 may belong in the Bolboceras group of genera,

Downloaded from Brill.com10/08/2021 04:54:17PM via free access 42 Tijdschrift voor Entomologie, volume 156, 2013 as implied by its author. Some characters are indeed B. birmanicum Lansberge, 1885 – Myanmar – re- similar, but differences in some details justify its sep- combined from Indobolbus [keying to quadridens arate position, including a modified abdominal tip – group]. the terminal ventrite is distinctly emarginate. Unfor- B. consanguineum Lansberge, 1885 – Myanmar – re- tunately, the concept of the genus Bolboceras as im- combined from Indobolbus [keying to quadridens plied in Krajcik’s world list (2012) ignores several of group]. the considered efforts of post-Boucomont workers to B. gaujani Fairmaire, 1892 – Djibouti, Somalia, further consolidate the generic classification of the Sudan (Paulian 1941, Krikken 2011). Bolboceratinae. B. inaequale Westwood, 1848 – India; Sudan, Sene- For the sake of completeness couplet 4 of the gal (Paulian 1941, Krikken 2011; Krikken in key to the species groups contains the name of a prep.). species representing the Bolboceras group combining B. kenyense Krikken, 2011 – Kenya. a subtrapeziform clypeus and a heavily punctate B. niloticum Boucomont, 1928 – Sudan, Somalia, pronotum; this is a new species, formally described Ethiopia (Paulian 1941, Krikken 2011). in the Appendix below, as there were no available B. perpunctatum sp.n. (Appendix 1 this paper). – species-group names in this particular group. India. The smaller nigricans group species, with their B. quadridens (Fabricius, 1781, in Scarabaeus)–In- pointed anteromedian clypeal protrusion, seem close dia; Andamans [species identity of this record to to forms in the groups with a subtrapeziform clypeus; be verified]. at this moment there is one likely case of conspeci- B. transversale Westwood, 1848 – India – recom- ficity, i.e. a male with an X-shaped clypeus and a fe- bined (from Indobolbus) [type depositary not es- male with a trapeziform clypeus (cf. Bolboceras insu- tablished]. lare sp.n.). B. trisulcatum Klug, 1845 – India – recombined from Indobolbus [keying to quadridens group]. List of named Bolboceras taxa Note changes in species names below (recombina- Key to groups of Oriental Bolboceras and tions, changes in suffixes). Recombined here means: species in its nigricans group formally transferred back from Indobolbus to Bol- Groups of Bolboceras (males) boceras (due to ICZN Opinion 2138, contrary to 1. Clypeal surface always more or less trapezi- Krikken 1984 and others). Some comments between form, anteromedian perimarginal ridge square brackets. (c in Fig. 1) of clypeal surface distinct, straight, transverse (angle at either end of Genus Bolboceras Kirby, 1919 ridge may be raised), lateral ridges never Type species: Scarabaeus quadridens Fabricius, 1781 convergent to a single median point ...... (ICZN 2006, Opinion 2138; Krell et al. 2003) – ...... groupsofBolboceras not treated here, 2 type material seen in BMNH. – Clypeus with lateral ridges of male clypeus Synonym: Indobolbus Nikolajev, 1979 – same type convergent to single anteromedian, more species. or less elevated point (Figs 27–32); any very short transverse anteromedian eleva- Oriental species in the nigricans group (order of tion convex (i.e. crest not concave and an- treatment in this paper): gular on either side, in axial view) ...... B. nigricans Westwood, 1848 – India, Myanmar, East ...... nigricans group: key to species below Africa; elsewhere? 2. Pronotal disc may have distinct discome- B. schulzei sp.n. – India: NE, Bangladesh. dian impression, but this is usually continu- B. malabaricum sp.n. – India: SW, elsewhere? ing onto anterior declivity, not enclosed by B. extraneum sp.n. – Bangladesh, India: N, Pakistan: saddle connecting discoparamedian protru- S. sionsandlateralsaddleoneitherside...... 3 B. insulare sp.n. – Sri Lanka. – Pronotal disc with discomedian impression, B. mimicans sp.n. –Myanmar. situated mainly behind more or less pointed Other available species-group names referable to discoparamedian tubercles (which are con- Bolboceras, with changes (alphabetical order): nected by transverse saddle; beware of mi- B. berytense Petrovitz, 1963 – Lebanon – recom- nor morphs!), posteriorly usually limited by bined from Indobolbus. rounded to W-shaped rim, in majors later- ally “overflowing” (over obliquely oriented

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saddle) to discolateral impression. (Review 3. Pronotal impressions posteriorly ill defined, Indianspp.inprep.)...... inaequale group discolateral impressions relatively shallow, 3. Pronotal set of four pointed tubercles or discomedian extending onto anterior de- denticles and three intervening impressions clivity. Crest of interocular ridge up to 0.5 distinct, discoparamedians distinctly protu- interocular distance long; more or less tri- berant (frequently conical, not usually con- undulate (axial view). Pronotum distinctly nected by transverse saddle). Body length depressed at base of anterior declivity. Dis- usually over 9 mm. Pronotum lacking close, tal lobe of eye canthus long. Basal genitalic distinct, double or multi-sized punctation capsule of male very large (lateral view) ...... quadridens group ...... B. mimicans – Pronotum (even of quasi major males, se- – Pronotal impressions posteriorly all simply, ries!) with superficial protrusions (at least distinctly defined, discolateral impressions the discolateral ones) separated by three deep. Interocular ridge ca 0.8 interocular very shallow (usually punctate) impressions distance long, crest straight to lateral angle (discomedian impression shallow, poorly (axial view). Pronotum slightly depressed at delimited)...... 4 base of anterior declivity. Distal lobe of eye 4. Pronotum with limited punctation only, canthus very short. Basal genitalic capsule i.e. not an extensive close cover. Small ofmalesmaller...... B. nigricans large down to 5.5 mm, length of majors usually 4. Aedeagal stalks (Fig. 3, lsas) with tips not exceeding 9 mm, pronotal protrusions slightly tapering, not hooked. Pronotum all reduced (anterior declivity poorly pro- distinctly depressed at base of anterior de- nounced)...... transversale group clivity (lateral view). Interocular ridge long – Pronotum with close, distinct, multi-sized ...... B. schulzei or double punctation, almost throughout – Aedeagal stalks with tips hooked-recurved. (less dense on either side of basal midline). Pronotum at most slightly and/or shortly Larger,usuallyover9mmlong...... depressedatbaseofanteriordeclivity...... 5 ...... perpunctatum group 5. Pronotum with very slight protrusions and impressions, discoparamedian protrusions Species in the nigricans group (males) not pointed. Eye canthus with long distal 1. Perimarginal ridges of male clypeus (full- lobe. Interocular ridge usually short ...... face view) K-shaped (anterior surface steep ...... B. malabaricum immediately above labrum). Pronotal dis- – Pronotum with shallow but distinct impres- coparamedian protrusions usually distinct. sions, discoparamedian protrusions conical. Medium-sized and large species, body Eye canthus with short distal lobe. Interoc- length usually over 9 mm. Parameres always ularridgelong...... B. nigricans small acuminate...... 2 6. Interocular elevation ca 0.8 interocular dis- – Perimarginal ridges of male clypeus more tance long, laterally abruptly declivous, X-shaped (median point shifted backward), crest unmodified (axial view). Perimarginal less so in very small specimens with trans- clypeal ridges anteromedially broadly pro- versely arcuate median point. Pronotal dis- tuberant, central tip transverse (if broader, coparamedian protrusions are simple cal- rounded in axial view). Parameral apex losities separated by shallow impressions. acuminate. Bangladesh, N India, S Pakistan Usually small, body length up to ca 10 mm...... B. extraneum Anterior edge of eye canthus simply angu- – Interocular elevation less than 0.5 inte- late. Pronotum not or shortly depressed at rocular distance long, crest more or less base of anterior declivity (lateral view)...... 6 tri-undulate/tuberculate (axial view). Cross- 2. Pronotal discomedian impression deep. Eye point of perimarginal clypeal ridges slightly canthus with tuberculate anterolateral an- protuberant. Parameral apex blunt. Sri gle. Large, body length reaching at least Lanka...... B. insulare 12mm...... 3 – Pronotal discomedian impression shallow or almost effaced. Eye canthus angulate in front but not conspicuously tuberculate. Body length usually less than 12 mm ...... 4

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Species accounts of head (full-face, excluding labrum and mandibles) Bolboceras nigricans Westwood, 1848 2.2, width 3.4. Median length of pronotum (dorsal) Figs 5, 11–14, 27, 33 4.5, maximum width 6.9. Median length of scutel- lum 1.0, maximum width 1.3. Sutural length of Material examined elytra (dorsal) 3.4, maximum width combined 7.0. Type, not sexed (OUMNH), with two original Width genital capsule 1.0. handwritten labels “Bengal”, “nigricans Westw.”, and various subsequently attached type labels, includ- ing one marking it as Coleoptera type 520. Further Variation material as in Annex 1. Note polymorphism indicated above, under Identifi- cation. One specimen 10 mm long. Identification This species and B. mimicans are the largest species in Comments the group. B. nigricans, however, has major and mi- This appears a widespread, variable species, in Asia nor morphs (observed in South Indian series), which, ranging from much of India to Rangoon in Myan- standing isolated, could be mistaken for different mar. The list of material examined above includes species; apart from size and associated development a slightly worn major male from Tanzania (“Dares- of pronotal protrusions and cavities I can see no spe- sal”), which is very similar to the Indian material. cific morphological differences. Confirmation based on additional, preferably fresh The discoparamedian protrusions of B. nigricans material is needed, but an East African occurrence are nearly always conical-pointed, unlike the smaller of Bolboceras nigricans is not unlikely (cf. situation in species diagnosed further below (like B. extraneum Bolboceratops, one species being shared between In- sp.n.). Major individuals have very distinct, pos- dia and East Africa, Carpaneto et al. 1993). Within teriorly well defined discomedian and discolateral India there is indeed variation, but, apart from the pronotal cavities – these may be considerably less Susai Nathan material, many records are based on pronounced in minors; discomedian impression re- singletons with limited pre-1940 data. mains distinct as posteriorly evenly rounded hol- low, in front gradually descending to anterior de- Distribution clivity. Interocular ridge strongly developed, long, Myanmar, India; African records to be verified. with medially simple crest, angulate on either side in major individuals (axial view). Anterolateral pro- trusion of eye canthus very distinct, distal lobe rela- Bolboceras schulzei sp.n. tively short compared to other group members (rela- Figs 6, 15–16, 28, 34 tively longer in minors). Pronotum with heavy con- Material examined centration of punctures behind eyes. Aedeagal lateral Holotype male (BMNH) from Bangladesh, labelled: sclerotized stalks robust, their tip hooked-recurved, “INDIA: \ Dacca. \ 27.viii.1945 \ D. Leston. \ parameres acuminate (Fig. 33). Depressed base of B.M. 1945–86.”, my type label added. Paratypes the anteromedian pronotal declivity short (in lateral as in Annex 1. Females possibly belonging to this view). Clypeus with K-shaped ridges, anteromedian species, excluded from type series. tubercle strongly produced, clypeal sides somewhat roundish (in full-face view). Colour more castaneous Identification than in other species, forebody usually slightly darker The decisive characters of this species seem to be than elytra. Body length usually 11–15 mm. the strongly depressed anteromedian area of the pronotum (i.e. at base of anterior declivity, best Measurements judged in profile) and the non-hooked lateral scle- Measurements in mm, of large male pictured. To- rotized stalks of the aedeagus (Fig. 34). B. schulzei tal body length 14.5. Median length of head (full- is medium-sized, and the pronotal protrusions are face, excluding labrum and mandibles) 2.7, width blunt but distinct. Interocular elevation long com- 4.4. Median length of pronotum (dorsal) 5.3, max- pared to malabaricum. Clypeus with K-shaped peri- imum width 8.8. Median length of scutellum 1.5, marginal ridges, compare Fig. 28 with other full-face maximum width 1.8. Sutural length of elytra (dorsal) pictures of Bolboceras heads. Parameres somewhat 4.5, maximum width combined 9.0. Width genital acuminate. Body length 10–11 mm. Few records, all capsule 1.1. – Measurements in mm, of small male from the Ganges basin in the North of the subconti- pictured. Total body length 11.5. Median length nent.

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Figs 5–10. Habitus of species in the nigricans group, oblique views, of: 5, Bolboceras nigricans,6,schulzei,7, malabaricum,8,extraneum,9,insulare, 10, mimicans. – Holotypes (6–10), Coimbatore (5), all males.

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Figs 11–18. Habitus, dorsal (left row) and lateral views (right row), of 11–12, Bolboceras nigricans large, 13–14, small, 15–16, schulzei, 17–18, malabaricum. – Holotypes (15–18), Coimbatore (11–12) and Anaimalai Hills (13– 14), all males.

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Holotype (male) (dorsal) 3.4, maximum width combined 7.0. Width Body length 10.5 mm. Colour generally uniformly genital capsule 1.0. dark brown, shiny. Labrum with slightly concave anterior margin, Variation transverse ridge distinct. Clypeal surface with supra- Few males available, variation slight. Females have anterolateral angles approximated-contiguous, for- a strongly depressed anteromedian pronotal surface, ming single broad anteromedian tubercle with as in male, excluded from type series. Some of these rounded apex, perimarginal ridges distinct (trans- females may belong to another (sub)species – see also verse near clypeogenal angle), K-patterned (full- under B. malabaricum treated hereafter. face view). Clypeofrons densely, distinctly punc- tate, around interocular elevation sparsely punctate, behind elevation abundantly to sparsely punctate; Distribution secondary punctation indistinct. Anterior edge of Bangladesh, N India. eye canthus straight to slight anterolateral tuber- cle, thence arcuate to tip of long distal lobe; sur- Etymology face rugulate-punctate; paraocular ridge fine, virtu- Dedicated to Joachim Schulze, who, in the Berlin ally straight, posteriorly extending slightly beyond museum, was most supportive during the initial stage side of interocular elevation. Transverse interocular of my studies on bolbos and other scarabs. elevation long, not reaching paraocular ridges, lat- eral slope slight, oblique, crest unmodified, straight Bolboceras malabaricum sp.n. to lateral angle on either side (all axial view). Figs 7, 17–18, 29, 35 Pronotum anteromedially distinctly depressed (lateral view), anterior declivity steep; discoparame- Material examined dian and lateral tubercles on pronotum slightly pro- Holotype male (MNHN) from India, labelled: tuberant; discomedian impression distinct, contin- “MAHÉ \ Chass. Indigènes \ Août”, “COTE uing onto anterior declivity; discolateral impression DE MALABAR \ M. MAINDRON”, ex coll. shallow, oblique, posteriorly not limited by edge; A. Boucomont. Other specimens (Annex 1) excluded basomedian surface with distinct but very shallow, from type series. punctate midline impression; anterolateral angle of pronotum ca 90°. Pronotal surface with double Identification punctation, primary punctation laterally, anteriorly, This is a medium-sized species with strongly re- and along margins abundant, from edge of lateral duced pronotal protrusions, and with its slightly de- (punctate) fovea onto basomedian surface a cluster pressed base of the anteromedian pronotal decliv- of 20–25 distinct punctures; secondary punctures ity short, smaller and less well defined than in B. sparse, minute; base marginate (medially virtually ef- schulzei. Interocular ridge of malabaricum very short faced). Scutellum with scattered, basally dense dou- (Fig. 29), ca 0.5 of interocular distance (crest slightly ble punctation. uneven in axial view). Aedeagal lateral sclerotized Elytra with discal striae shallowly impressed, finely stalks very slender, apex hooked-recurved, parameres punctate; punctures separated by 5–8 punctural di- acuminate. Clypeus with K-shaped ridges. Note that ameters, slightly crenulating interstriae. Elytral inter- anterolateral angle of pronotum of type is distinctly striae very slightly convex, vaguely, sparsely, minutely sharper than in other species. Body length ca 10 mm. punctate. These notes are based on the male type from Protibia with 6 distinct external denticles; apex southwestern India; other specimens slightly differ- unmodified, with robust tapering spur. Meso- and ent (interocular ridge, anterolateral angle of prono- metatibiae with bilobate apical and one complete tum) – see under Variation and comment, below. anteapical bilobate fossorial elevation; their crest fringed with fine spines. Aedeagus, Fig. 34; lateral sclerotized stalks with Holotype (male) slightly curved, tapering, blunt tip (one parameral Body length 10 mm. Colour generally uniformly point in type broken off). dark-brown, shiny. Measurements in mm. Median length of head Labrum with slightly concave anterior margin, (full-face, excluding labrum and mandibles) 1.8, transverse ridge distinct. Clypeal surface with supra- width 3.4. Median length of pronotum (dorsal) 3.9, anterolateral angles approximated-contiguous, for- maximum width 6.2. Median length of scutellum ming single broad anteromedian tubercle with trans- 1.0, maximum width 1.0. Sutural length of elytra versely rounded apex, perimarginal ridges distinct,

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K-patterned (full-face view). Clypeofrons sparsely to excluded from the type-series; in other words, future abundantly, finely but distinctly punctate in front to informative series may indicate that they represent base of interocular elevation, behind elevation also another (sub)species near B. malabaricum as here sparsely to abundantly punctate; punctation multi- conceived – the geographical separation is also a sized. Anterior edge of eye canthus short, straight to point to be kept in mind. See also females under B. distinct anterolateral tubercle, thence arcuate to tip schulzei. Total size range 10–11 mm. of long distal lobe; surface rugulate-punctate; paraoc- ular ridge fine, virtually straight, posteriorly extend- Distribution ing beyond side of interocular elevation. Transverse Southwestern India, possibly occurring also in more interocular elevation short, ca 0.5 of interocular dis- northern regions (different species?). tance, lateral slope slight, oblique, crest medially slightly raised (all axial view). Pronotum anteromedially slightly depressed (lat- Etymology eral view), gradually sloping to steeper anterior de- Species named after the type region of this species, clivity; discoparamedian and lateral tubercles on adjective. pronotum slightly protuberant, vaguely conical; dis- comedian impression and discolateral impressions Bolboceras extraneum sp.n. very shallow; basomedian surface with distinct but Figs 8, 19–20, 30, 36 fine, punctate midline impression; anterolateral an- Material examined gle of pronotum ca 80°. Pronotal surface with double Holotype male (BMNH) from India, labelled: “I. punctation, primary punctation laterally, anteriorly, \ 20.VIII.20 \ At light \ Pusa \ Haq.[?] Coll.”. and along margins abundant, from edge of lateral Paratypes as in Annex 1. Females possibly belonging (punctate) fovea onto basomedian surface a cluster to this species excluded from type series. of 20–25 distinct punctures; secondary punctures sparse, minute; base marginate (medially virtually effaced). Scutellum with scattered, relatively sparse, Identification double punctation. This is a small, light to medium brown species (body Elytra with discal striae shallowly impressed, length usually not over 9 mm) from an area ranging finely punctate; punctures separated by 4–7 punc- from Bangladesh through lowland North India, and tural diameters, slightly crenulating interstriae. Ely- apparently West to South Pakistan. The pronotum tral interstriae scarcely convex, vaguely, very sparsely, has a slight anterior declivity, a short basal anterome- minutely punctate. dian depression (in profile), and its antediscal pro- Protibia with 6 distinct external denticles; apex trusions are reduced to mere callosities (contrary to unmodified, with robust tapering spur. Meso- and minors in, for instance, B. nigricans). Clypeus with metatibiae with bilobate apical and one complete moderately distinct X-shaped perimarginal ridges, anteapical bilobate fossorial elevation; their crest median tubercle usually distinct, its tip shortly, trans- fringed with fine spines. versely rounded (in axial view, may look straight Aedeagus, Fig. 35; lateral sclerotized stalks slender, in dorsal view). Interocular ridge of B. extraneum with rounded-hooked tip (extruded in holotype). with simple crest, not distinctly tri-tuberculate, con- Measurements in mm. Median length of head trary to insulare (in axial view), described further be- (full-face, excluding labrum and mandibles) 1.9, low. Head densely punctate(-rugulate). Pronotal pro- maximum width 3.2. Median length of pronotum trusions reduced to slight convexities. Aedeagal lat- (dorsal) 3.5, maximum width 6.9. Median length of eral sclerotized stalks slender, with equally slender scutellum 1.0, maximum width 1.0. Sutural length tip hooked-recurved (may be concealed in parameral of elytra (dorsal) 3.6, maximum width combined sheath), parameres acuminate. 5.9. Width genital capsule 0.9. Holotype (male) Body length 8 mm. Colour generally uniformly me- Variation and comment dium brown, somewhat rufous, shiny. Apart from the characters expressed in the holotype Labrum with slightly concave anterior margin, (given under Identification, above), two other males transverse ridge distinct. Clypeal surface with supra- differ in details, including the anterolateral corner of anterolateral angles approximated-contiguous, for- the pronotum being (in full-face view) rectangular ming single broad anteromedian tubercle with trans- instead of more acute, and the interocular ridge being versely rounded apex, perimarginal ridges distinct, longer (the tips of their lateral parameral stalks are X-patterned (full-face view). Clypeofrons densely, curved-hooked). These “non-typical” specimens are distinctly punctate(-rugulate) in front to base of in-

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Figs 19–26. Habitus, dorsal (left row) and lateral views (right row), of: 19–20, Bolboceras extraneum, 21–22, insulare, 23–24, mimicans, 25–26, perpunctatus. – Holotypes, all males.

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Figs 27–32. Head upper side, appendages omitted, full-face views (right), of: 27, Bolboceras nigricans, 28, schulzei, 29, malabaricum, 30, extraneum, 31, insulare, 32, mimicans. – Holotypes (28–32), Coimbatore (27); all males. terocular elevation, behind elevation densely punc- Pronotum anteromedially shortly depressed (lat- tate; punctation multi-sized. Anterior edge of eye eral view), anterior declivity slight; discoparamedian canthus short, straight to distinct anterolateral an- and lateral tubercles reduced to mere callosities; dis- gle, thence arcuate to tip of long distal lobe; surface comedian and discolateral impressions shallow; ba- rugulate-punctate; paraocular ridge very fine, virtu- somedian surface with shallow, punctate midline im- ally straight, posteriorly extending slightly beyond pression; anterolateral angle of pronotum ca 90°. side of interocular elevation. Transverse interocular Pronotal surface with double punctation, primary elevation low, long, not reaching paraocular ridges, punctation laterally, anteriorly, and along margins lateral slope slight, oblique, crest unmodified, virtu- abundant, from edge of lateral (punctate) fovea ally straight to lateral angle on either side (all axial onto basomedian surface a cluster of 30–40 dis- view). tinct punctures; secondary punctures sparse, minute;

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Figs 33–39. Aedeagus distal part, upper side, of 33, Bolboceras nigricans, 34, schulzei, 35, malabaricum, 36, extraneum, 37, insulare, 38, mimicans, 39, perpunctatum. – Holotypes (34–39), Coimbatore (33). Median apparatus extruded (35, 39), sclerotized stalks expanded. – Fig. 40, B. perpunctatum holotype, punctation halfway left lateral declivity of pronotum (scale bar = 0.5 mm).

Downloaded from Brill.com10/08/2021 04:54:17PM via free access 52 Tijdschrift voor Entomologie, volume 156, 2013 base marginate (medially virtually effaced). Scutel- shaped clypeal ridges (at least in the male sex, see lum with scattered, relatively sparse, double puncta- under Variation and sexual dimorphism, below) and tion. a relatively short, tri-undulate to tri-tuberculate inte- Elytra with discal striae shallowly impressed, dis- rocular ridge (Fig. 31). Pronotal protrusions strongly tinctly punctate; punctures separated by 3–5 punc- reduced, almost effaced on a gradually sloping an- tural diameters, distinctly crenulating interstriae. terior declivity. Parameral tips blunt (Fig. 37), not Elytral interstriae scarcely convex, vaguely, sparsely, acuminate, as in all other nigricans group members minutely punctate. and in most other Bolboceras. Body length usually 9– Protibia with 6 distinct external denticles; apex 10 mm. unmodified, with robust tapering spur. Meso- and metatibiae with bilobate apical and one complete Holotype (male) anteapical bilobate fossorial elevation; their crest Body length 9 mm. Colour generally uniformly ru- fringed with fine spines. fous brown. Aedeagus, Fig. 36; lateral sclerotized stalks fine, Labrum with slightly concave anterior margin, with hooked-recurved tip. transverse ridge distinct. Clypeal surface with supra- Measurements in mm. Median length of head anterolateral angles approximated-contiguous, for- (full-face, excluding labrum and mandibles) 1.8, ming single anteromedian point, perimarginal ridges maximum width 2.6. Median length of pronotum fine, X-patterned (full-face view). Clypeofrons abun- (dorsal) 2.8, maximum width 4.6. Median length of dantly to sparsely, distinctly punctate in front to base scutellum 0.8, maximum width 0.8. Sutural length of interocular elevation, behind elevation sparsely of elytra (dorsal) 2.7, maximum width combined punctate; secondary punctation present, but indis- 4.8. Width genital capsule 0.8. tinct. Anterior edge of eye canthus short, straight to distinct anterolateral angle, thence arcuate to tip of Variation and comment long distal lobe; surface irregularly abundantly punc- The specimens seen appear all conspecific, looking tate; paraocular ridge very fine, virtually straight, less variable than the previous species, but this may posteriorly extending slightly beyond side of interoc- be partly due to size, and definitely a study of more ular elevation. Transverse interocular elevation low, material is required to confirm this; one yellow- short, lateral slope slight, oblique, crest finely tri- brown male seen from Pakistan. Body length 6.5– tuberculate (all axial view). 8 mm. Some specimens have a more K-shaped set Pronotum anteromedially slightly depressed (lat- of perimarginal ridges, but in these cases the median eral view), anterior declivity slight; discoparamedian point is distinctly transversely arcuate. and discolateral protrusions reduced to mere callosi- ties; discomedian and discolateral impressions ob- solescent; basomedian surface with punctate, hardly Distribution impressed midline; anterolateral angle of pronotum Bangladesh, N India, S Pakistan. ca 90°. Pronotal surface with double punctation, pri- mary punctation laterally, anteriorly, and along mar- Etymology gins abundant, from edge of lateral (punctate) fovea Name, adjective, referring to apparent occurrence onto basomedian surface a cluster of 20–25 distinct outside Indian subcontinent proper – the conspeci- punctures; secondary punctures sparse, minute; base ficity of such specimens has to be verified. marginate (medially virtually effaced). Scutellum with scattered, relatively sparse, double punctation. Bolboceras insulare sp.n. Elytra with discal striae shallowly impressed, Figs 9, 21–22, 31, 37 finely punctate; punctures separated by 4–6 punc- tural diameters, slightly crenulating interstriae. Ely- Material examined tral interstriae scarcely convex, vaguely, very sparsely, Holotype male (BMNH) from Sri Lanka, labelled: minutely punctate. \ \ \ “Halupahani, Haldummulle, Ceylon. 1904– Protibia with 6 distinct external denticles; apex 71”. Paratypes as in Annex 1. unmodified, with robust tapering spur. Meso- and metatibiae with bilobate apical and one complete Identification anteapical bilobate fossorial elevation; their crest Although other groups of Bolboceras are represented fringed with fine spines. on Sri Lanka, this species is so far the only one in Aedeagus, Fig. 37; lateral sclerotized stalks with the nigricans group, and may be an island endemic. slenderhooked-recurved tip; parameres short, blunt. Size of B. insulare varies, but usually rather small to Measurements in mm. Median length of head medium-sized, uniformly brown, with distinctly X- (full-face, excluding labrum and mandibles) 1.7,

Downloaded from Brill.com10/08/2021 04:54:17PM via free access Krikken: The Bolboceras nigricans group 53 maximum width 3.0. Median length of pronotum Holotype (male) (dorsal) 3.5, maximum width 5.7. Median length of Body length ca 14.5 mm. Colour generally uni- scutellum 1.0, maximum width 1.0. Sutural length formly medium brown, shiny. of elytra (dorsal) 2.9, maximum width combined Labrum with slightly concave anterior margin, 5.9. Width genital capsule 0.9. transverse ridge obliterated by rugulate puncta- tion. Clypeal surface with supra-anterolateral angles Variation and sexual dimorphism approximated-contiguous, forming single broad an- From the type-locality and from “Ceylon” (no de- teromedian tubercle with transversely rounded apex, tails) females with a strongly punctate, simply sub- perimarginal ridges distinct, K-patterned (full-face trapeziform clypeus are available which have a local- view). Clypeofrons densely, distinctly punctate to ity label identical to the Bolboceras insulare holotype, near base of interocular elevation, around elevation and an identical “Ceylon” label on a male paratype, with less punctures; punctation multi-sized. Ante- respectively. These females are otherwise quite simi- rior edge of eye canthus short, straight to high an- lar to their male “label counterparts” with X-shaped terolateral tubercle, thence arcuate to tip of long dis- perimarginal ridges, sharing a short tri-dentate in- tal lobe; surface rugulate-punctate; paraocular ridge terocular ridge and almost effaced pronotal protru- fine, virtually straight, posteriorly extending to level sions. It may thus well be that these document an of interocular elevation. Transverse interocular eleva- exception to the prevalent absence of sexual dimor- tion relatively short, lateral slope slight, oblique, crest phism in the genus Bolboceras. However, there cer- tri-undulate, ca 0.5 of interocular distance (all axial tainly are more or less similar “trapeziform males” view). from other Sri Lanka localities, with aedeagi differ- Pronotum anteromedially strongly depressed (lat- ent from insulare. More material is needed. Colour eral view), anterior declivity steep on either side of varies from light to dark brown. Body length of B. discomedian impression; discoparamedian and dis- insulare 8–10 mm. colateral tubercles on pronotum moderately protu- berant, subconical; discomedian impression deep, anteriorly not separated from anterior declivity (as Distribution in B. nigricans) but sloping down to anteromedian Sri Lanka. surface; discolateral impression shallow, posteriorly slightly limited by blunt edge; basomedian surface Etymology with distinct but shallow, densely punctate midline Name of this species based on its insular occurrence, impression; anterolateral angle of pronotum ca 90°. on Sri Lanka; adjective. Pronotal surface with double punctation, primary punctation laterally, anteriorly, and along margins Bolboceras mimicans sp.n. abundant, from edge of lateral (punctate) fovea onto basomedian surface a cluster of 40–50 distinct punc- Figs 10, 23–24, 32, 38 tures; secondary punctures sparse, minute, vague; Material examined base marginate (medially virtually effaced). Scutel- Holotype male (BMNH) from Myanmar, labelled: lum with fine, scattered, relatively sparse punctation. “Burma: \ Rangoon. \ Archbald Coll. \ B.M. 1926– Elytra with discal striae shallowly impressed, finely 391.”. Paratypes as in Annex 1. punctate; punctures separated by 5–7 punctural di- ameters, slightly crenulating interstriae. Elytral in- Identification terstriae scarcely convex, vaguely, sparsely, minutely This is a large species, characterized primarily by its punctate. distinct pronotal protrusions and impressions. Dis- Protibia with 6 distinct external denticles; apex comedian sulcus of B. mimicans broad, long, grad- unmodified, with robust tapering spur. Meso- and ually sloping down over anterior declivity to large metatibiae with bilobate apical and one complete depressed anteromedian surface (Figs 23–24). Inte- anteapical bilobate fossorial elevation; their crest rocular ridge relatively short, with tri-undulate crest, fringed with fine spines. like in B. insulare, which is a much smaller endemic Aedeagus, Fig. 38; lateral sclerotized stalks robust, of Sri Lanka. Lateral sclerotized aedeagal stalks ro- with hooked-recurved tip, parameres narrow in rela- bust, their tip hooked-recurved, basal capsule very tion to basal capsule. robust, parameres in comparison small, finely acumi- Measurements in mm. Median length of head nate (Fig. 38). Clypeus with K-shaped perimarginal (full-face, excluding labrum and mandibles) 2.6, ridges. Body length 11.5–15 mm. maximum width 4.5. Median length of pronotum (dorsal) 5.5, maximum width 8.9. Median length of scutellum 1.3, maximum width 1.6. Sutural length

Downloaded from Brill.com10/08/2021 04:54:17PM via free access 54 Tijdschrift voor Entomologie, volume 156, 2013 of elytra (dorsal) 4.7, maximum width combined concave (axial view); anterior and lateral clypeal declivities 9.4. Width genital capsule 1.3. rugulate-punctate. Most of clypeofrontal surface, including in- terocular ridge, finely, distinctly, densely to crowdedly punc- tate. Anterior edge of eye canthus short, straight to distinct an- Variation terolateral tubercle, thence arcuate to tip of roughly equal-sized Females seen smaller, with less prominent pronotal distal lobe; surface rugulate-punctate; paraocular ridge fine but discomedian sulcus, but the number of specimens is distinct, virtually straight, posteriorly extending beyond level too low to draw conclusions on sexual dimorphism. of interocular elevation. Transverse interocular elevation very distinct, lateral slope gradual, almost reaching raised paraocu- Distribution lar ridge, crest fine, slightly convexly curved (axial view). Myanmar. Pronotum anteromedially not depressed, anterior decliv- ity steep, discomedian impression very shallow, not distinctly separated from adjacent surface; discoparamedian protrusion Comment shortly conical; discolateral impression very shallow, not dis- Bolboceras birmanicum and consanguineum,bothde- tinctly separated from adjacent surface; discolateral protrusion scribed from Minhla in Myanmar (Lansberge 1885), broad, slightly pronounced; basomedian impression shallow belong in the quadridens group. but distinct, and densely, finely punctate; anterolateral angle of pronotum ca 85°; pronotal base distinctly marginate and lined with dense to crowded punctation. Pronotal surface laterally Etymology and anteriorly with generally close multi-sized punctation, pri- Name of this species based on its “mimicking” B. mary punctation dense to crowded, rather coarse; interspersed nigricans. finer punctation sparser, but locally dominant and dense, as on the four pronotal protrusions; surface on either side of punctate basomedian impression shiny, with almost sparse fine puncta- Appendix. The Bolboceras perpunctatum tion only; punctation on shallow postocular impression strong, group crowded. Scutellum with scattered, mostly fine and abundant punctation. Presumably there are more species in this group, judged from Elytra with discal striae shallowly impressed, distinctly the few specimens in the material at hand, and consequently punctate; punctures separated by 2–4 punctural diameters, this notice of a characteristic group representative, apart from slightly crenulating interstriae. Elytral interstriae scarcely con- underpinning the key to the Bolboceras groups of species given vex, vaguely, sparsely, minutely punctate in vaguely microretic- above, also serves to promote more collecting and study. ulate background. Protibia with 6 distinct external denticles; apex unmodified, Bolboceras perpunctatum sp.n. with robust tapering spur. Meso- and metatibiae with bilobate Figs 4, 25–26, 39, 40 apical and one complete anteapical bilobate fossorial elevation, their crest fringed with slender spines. Material examined Aedeagus, Fig. 40 (extruded in holotype); lateral sclerotized \ Holotype male from India (IRSNB), labelled: “Tumlong stalks very robust, with broadly recurved tip; parameres acumi- \ Sikkim Schneider” (printed), ex coll. J.J.E. Gillet. Paratype nate. female with same data. Altitude ca 1600 m (my estimate). Measurements in mm. Median length of head (full-face, excluding labrum and mandibles) 2.4, maximum width 3.9. Identification Median length of pronotum (dorsal) 4.6, maximum width 7.8. This North Indian species differs from all other described Bol- Median length of scutellum 1.2, maximum width 1.6. Sutural boceras by its very strongly punctate forebody. The punctation length of elytra (dorsal) 4.3, maximum width combined 7.9. on the pronotum is very extensive, symmetrically patterned, Width genital capsule 2.2. varied and overall multi-sized (Fig. 40). Pronotal protrusions short, discoparamedians shortly conical, intervening impres- Etymology sions very shallow for such a relatively large species. Lateral Name of this species based on the heavy punctation of the sclerotized aedeagal stalks robust, recurved. The characters of forebody, adjective. potential relatives are difficult to assess due the paucity of the material (few specimens, mainly female); some come from low- land areas (like Calcutta) – additional accounts are anticipated. Acknowledgements Body length of perpunctatus apparently 12–13 mm. This study is based on access to the collections men- tioned above, and I am indebted to the present and Holotype (male) former owners and curating officers for their patient Body length ca 12.5 mm. Colour generally uniformly medium brown, shiny, somewhat opaque due to close punctation on collaboration. Early drafts of this paper were read by forebody and microreticulation on all dorsal parts. my colleagues J. van Tol (Leiden, The Netherlands), Labrum with slightly concave anterior margin, transverse and. F.-T. Krell (Denver CO, USA). I thank all in- ridge distinct. Clypeal surface subtrapeziform (full-face view), volved for their continuous support. with supra-anterolateral angles protuberant, intervening ridge

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Paulian, R., 1941. Coprophaga Africana: Deuxième note. Bolboceras malabaricum Revision des Bolboceras africains. – Revue de Zoologie [BANGLADESH:] Dinajpur: Dhanjuri, vi/1963, Mappelli, 1 et Botanie africaines 35: 1–71. spm, in MHNG. Petrovitz, R., 1963. Neue und interessante Scarabaeidae aus INDIA: Bihar: Pusa, 12/x/1906, Mitra J, at light, 1 spm, dem vorderen Orient II. Teil. – Reichenbachia 1: 235– in BMNH. Malabar: Mahé, viii/1901, 1 holotype,in 267. MNHN Boucomont. Scholtz, C.H., A.L.V. Davis & U. Kryger, 2009. Evolu- Subtotal 3 male specimens, 3 collection records. tionary Biology and Conservation of Dung Beetles. – Sofia/Moscow, Pensoft, 1–567. Bolboceras mimicans Scholtz, C.H. & V.V. Grebennikov, 2005. 12. [MYANMAR:] Burmah: Rangoon, Meggitt FJ, 1 paratype in Scarabaeiformia Crowson, 1960. 13. Scarabaeoidea La- BMNH. Burma: Rangoon, Archbald Coll., 1 holotype,in treille, 1802. – In: Handbuch der Zoologie/Handbook BMNH. Rangoon, Meggitt FJ, 2 paratypes, in USNM. 1 of Zoology (4) 38: 345–429. paratype in SMTD. Birmah: 1 paratype in BMNH. Verdú, J.R., E. Galante, J.-P. Lumaret & F.-J. Cabreo- Subtotal 6 male and female specimens, 5 collection records. Sañudo, 2004. Phylogenetic analysis of Geotrupidae (Coleoptera, Scarabaeoidea) based on larvae. – Syste- Bolboceras nigricans matic Entomology 29: 509–523. INDIA: [Andhra Pradesh:] Hyderabad, vi/1970, 1 spm, Westwood, J.O., 1848 (Oct.). Descriptions of some new or in CMNC. [Madhya Pradesh:] Jabalpur, ix/1957, Susai imperfectly known species of Bolboceras. Proceedings of Nathan P, 1 spm, in CMNC. [Maharashtra:] S Bom- bay [Mumbai], 1 spm, in BMNH. Kadegaon, 1 spm, in the Linnean Society of London 1: 384–387 [same title BMNH. [Karnataka:] Belgaum, 1 spm, in BMNH. [Tamil & text: 1848 (Nov.). – Annals and Magazine of Natural Nadu:] Ammayana [text unclear], Yakonour, Maduré History (2) 2: 353–356]. [Madurai], v/1908, 1 spm, in MNHN Boucomont. Coim- Westwood, J.O., 1852. Descriptions of some new or im- batore, Susai Nathan P, 1400 ft, 2 spms, in CMNC. perfectly known species of Bolboceras,Kirby.–Transac- vi/1950, Susai Nathan P, 1 spm, in CMNC. x/1966, Susai tions of the Linnean Society of London 21: 19–30. Nathan TR, 1400 ft, 1 spm, in CMNC. x/1966, 1400 ft, 1 spm, in CMNC. x/1966, 1400 ft, 4 spms, in CMNC. Received: December 7, 2012 xi/1955, Susai Nathan P, 1 spm, in CMNC. xii/1932, Su- Accepted: February 20, 2013 sai Nathan P, 1 spm, in MNHN. Coimbatore, x/1965, Susai Nathan P, 1400 ft, 1 spm, in ZSM. x/1965, Susai Nathan P,1400 ft, 7 spms, in ZSM. xi/1963, Susai Nathan P, 1400 ft, 1 spm, in RMNH. xi/1964, Susai Nathan P, 1 spm, in USNM. xi/1963, Susai Nathan P,1400 ft, 1 spm, Annex 1. Material examined (label data in RMNH. xi/1964, Susai Nathan P, 1400 ft, 14 spms, in Bolboceras nigricans species group) ZSM. Dindigul, 1 spm, in BMNH. Trichinopoly [Tiruchi- Species in alphabetical order rapalli], Paesler T, 1 spm, in ZMHUB. Madras [Chen- nai], Rankin RW, 1 spm, in BMNH. Anaimalai Hills, Bolboceras extraneum Cinchona, vi/1966, Susai Nathan P, 3500 ft, 1 spm, in [BANGLADESH:] Dinajpur: Dhanjuri, vi/1963, Mapelli, 1 CJSB. Anaimalai Hills, Cinchona, v/1966, Susai Nathan P, paratype in MHNG. 3500 ft, 1 spm, in CJSB. Pondicherry: Karikal, viii/1964, INDIA: 3 paratypes, in BMNH. Fyzabad, Hingston RWG, 1 Susai Nathan P, 5 spms, in ZSM. N Bengal: 1 spm, in paratype in BMNH. Bihar: Pusa, 20/vii/1920, Haq.Coll., BMNH. [INDIA?]: Dakhun [text unclear], 1 spm, in at light, 2, incl. holotype, paratype in BMNH. 01/vii/ BMNH. IND.OR.: 1 spm, in BMNH. Type as in species 1907, HML, at light, 1 paratype in BMNH. 04/viii/1905, account. CHA, at light, 1 paratype in BMNH. 03/viii/1920, Sarkar [MYANMAR:] Rangoon, 1 spm, in SMT. SC, at light, 1 paratype in BMNH. 17/ix/1908, GMC, at [TANZANIA:] Daressal[aam], 1 spm, in MNHN Bouco- light, 1 paratype in BMNH. 05/x/1905, CHA, 1 paratype mont. in BMNH. Bengal: Kalkutta, 1 paratype in SMTD. NO LOCATION: 1 spm, in BMNH (comp. with type by PAKISTAN: Karachi, Bell TR, 1 paratype in BMNH. Arrow). Subtotal 14 male and female specimens, 11 collection Subtotal 58 male and female specimens, 31 collection records. records. Bolboceras schulzei Bolboceras insulare [BANGLADESH]: Dacca, 27/viii/1945, Leston D, 1 holo- [SRI LANKA:] Ceylon, 2 paratypes, in NMPC. Ceylon, Niet- type, in BMNH. Dinajpur: Boldipukur, ix/1958, Pussetto, ner, 1 paratype in ZMHUB. Halupahani: Haldummulle, 1 spm, in MHNG. Dhanjuri, vi/1963, Mappelli, 1 spm, in 1904–171, 3, incl. holotype, paratypes in BMNH. Mal MHNG. vi/1963, Mappelli, 1 spm, in PIME. Oya, xii/1905, 1 paratype in CNMS. Wellawaya, xi/1905, INDIA: West Bengal: Kalkutta [Kolkata], 1 paratype in 2 paratypes, in CNMS. Peradenya, vii/1905, 1 paratype, SMTD. Uttar Pradesh: C, Almora Dn., Kumaon, vii/ x/1905, 1 paratype, both in IRSNB ex Gillet coll. 1920, Champion HG, 1 paratype in BMNH. Subtotal 11 male and female specimens, 7 collection records. Subtotal 6 male and female specimens, 6 collection records.

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