Coleoptera, Scarabaeidae, Scarabaeinae

Zootaxa 3802 (2): 276–284 www.mapress.com /zootaxa / Copyright © 2014 Magnolia Press

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ZOOTAXA

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http://dx.doi.org/10.11646/zootaxa.3802.2.8 http://zoobank.org/urn:lsid:zoobank.org:pub:2797E064-00BF-4113-8F84-0AEDE0C991DC

A new giant species of Deltochilum subgenus Deltohyboma (Coleoptera, Scarabaeidae, Scarabaeinae) from Colombia, with notes on D. spinipes Paulian, 1938 FERNANDO A. B. SILVA1,3 & FERNANDO Z. VAZ-DE-MELLO2 1

Universidade Federal do Pará, Instituto de Ciências Biológicas, Campus Belém. Rua Augusto Corrêa, 01. Guamá. Belém PA 66075110, Brazil. E-mail: [email protected] 2 Muséum National d’Histoire Naturelle, Département Systématique et Évolution, Entomologie, 57 rue Cuvier, F-75231 Paris cedex 05, France. Permanent address: Universidade Federal de Mato Grosso, Instituto de Biociências, Departamento de Biologia e Zoologia. Av. Fernando Correa da Costa, 2367. Boa Esperança. Cuiabá MT 78060-900, Brazil. E-mail: [email protected]. CNPq fellow 3 Corresponding author

Abstract Deltochilum (Deltohyboma) gigante sp. nov., from Parque Nacional Las Cuevas de los Guácharos, Huila, Colombia is described based on differences in external and genital morphology. Its diagnostic characters are provided and illustrated. Lectotype and paralectotypes for D. spinipes, a close relative to the new species, are designated. Key words: Deltochilum gigante, Scarabaeinae, new species, diagnosis, taxonomical remarks

Introduction The genus Deltochilum Eschscholtz, 1822 belongs to the subfamily Scarabaeinae, which includes the insects popularly known as dung beetles (Halffter & Matthews 1966). In the most recent account, the genus included 85 valid species (Génier 2012). The classification of Deltochilum into subgenera is based mainly on elytral carinae (Lane 1946). Kolbe (1893) was the first who classified Deltochilum species by splitting the genus into seven subgenera: Aganhyboma Kolbe, 1893, Calhyboma Kolbe, 1893, Deltochilum s. str., Euhyboma Kolbe, Meghyboma Kolbe, Paedhyboma Kolbe and Telhyboma Kolbe. Further studies continued development of subgeneric classification of Deltochilum that is currently divided into eight subgenera: Aganhyboma; Calhyboma; Deltochilum s. str.; Deltohyboma Lane, 1946; Euhyboma; Hybomidium Shipp, 1897; Parahyboma Paulian, 1938 and Rubrohyboma Paulian, 1938 (Shipp 1897; Paulian 1938, 1939; Pereira & Martínez 1956; Vulcano & Pereira 1964; Génier 2012). The subgenus Deltohyboma, which includes the new species, has approximately 47 valid species, distributed throughout the Neotropics, ranging from Mexico to Argentina (González et al. 2009). The species of this subgenus are characterized by the ninth interstria distinctly carinate; ninth interstrial carina extending at most one-half length of pseudepipleuron; pseudepipleural carina distinctly visible basally; and clypeus bidentate (Vaz-de-Mello et al. 2011). Like other dung beetles, most species of Deltohyboma are usually necrophagous or coprophagous (Vaz-deMello 1999). However, other types of behavior, for example, predation by species of the "valgum" group on diplopods has been recorded (Cano 1998; Génier 2001; Escobar 2003; Halffter 2003; Larsen et al. 2009; Silva et al. 2012). Until now, D. spinipes Paulian, 1938 has been the largest known species of the genus, with a body length ranging from 18 to 24 mm (Paulian 1938). Examination of the collections at the Natural History Museum, London, and the Muséum National d'Histoire Naturelle, Paris, has revealed a new species of Deltochilum that is larger than any hitherto known. In this paper we describe this newly discovered species of the subgenus Deltohyboma and provide notes on its probable closest relative D. spinipes.

276 Accepted by S. Tarasov: 16 Apr. 2014; published: 26 May 2014

Material and methods The material studied is deposited in the following museums: BMNH (Natural History Museum, London, curators Max Barclay and Malcolm Kerley); MNHN (Muséum National d'Histoire Naturelle, curators Olivier Montreuil and Antoine Mantilleri). Photographs of the holotype were taken with a Leica stereomicroscope M205A, using image stacking software (Leica Application Suite, version 3.7.0). The terminology used to describe the endophallic sclerites of the male genitalia is that proposed by Tarasov and Solodovnikov (2011), with some slight changes. For illustration and examination of endophallic sclerites, the endophallus was removed from the aedeagus through the basal foramen of the phallobase. Endophallic sclerites were drawn in dorsal view at rest within the body (seen through the right lateral view of the aedeagus), except for the superior right peripheral sclerite (SRP), which was illustrated at rest in ventral view (seen from the left side of the aedeagus).

Taxonomy Deltochilum (Deltohyboma) gigante sp. nov. (Fig. 1–2) Etymology. The species name refers to the body size, gigante meaning "giant" in both the Portuguese and Spanish languages. The species epithet is a noun in apposition. Diagnosis. The new species is placed in the group of species that comprises D. spinipes, probably D. aequinoctiale (Buquet 1844) (type specimens not examined), and D. speciosissimum Balthasar, 1939. This group is characterized by the following characters: size equal or greater than 13 mm; apical tubercles or carinae on interstriae 3-4-5-6, very weak or lacking on interstria 7; elytra lacking foveate punctures on discal striae. In this group of species, D. aequinoctiale and D. speciosissimum have a microgranulated opaque dorsal integument, while D. spinipes and D. gigante share a smooth and shiny surface of the elytra and pronotum (punctures might be present but surface lacks microgranulation). Deltochilum spinipes differs from D. gigante by its significantly smaller size and the lack of the posterior metasternal spiniform projections as well as the internal tooth on protibia. In addition, D. spinipes has a stronger mesofemur with a larger tubercle on posterior margin, and a conspicuous apical carina on the seventh elytral stria. Furthermore, D. gigante can be differentiated from all the remaining species in this subgenus by its larger body size. Description (Holotype). Colour. Completely metallic blue, with green to violaceous sheen (Figures 1A, B). Length. 24 mm. Head. Dorsal surface with dense foveiform punctures, closer on lateral portions (Figure 1C). Clypeal margin with two small, triangular, sharp, central teeth, separated by approximately five times the width of their basal portion. Margin between clypeal teeth and clypeo-genal suture almost straight. Margin of gena strongly curved outward. Clypeo-genal suture slightly impressed. Clypeo-frontal suture brighter than surrounding surface. Dorsal interocular width eight times the eye width (Figure 1C). Antennae 9-segmented, club 3-segmented. Posterior dorsal margin of head (occipital region of front) completely marginated by fine carina. Thorax. W/L = 1.3; pronotum strongly convex, with a smooth longitudinal depression medially (Figure 1A). Dorsal surface with dense and small foveiform punctures, randomly spaced; punctures larger and closer on anterior angles and lateral portions of pronotum. Anterior margin deeply emarginated. Anterior angles acute, directed forward (Figure 1C). Lateral pronotal impressions with a small rounded tubercle, poorly differentiated from surrounding surface. Margin between anterior and lateral angles oblique, slightly curved outward. Margin between lateral and posterior angles distinctly curved outward. Hypomera not excavated (Figure 1D); lacking carina separating the anterior and posterior area; foveiform punctures denser on anterior and posterior portions, with some long setae on anterior portion; median portion with sparse or absent punctures. Meso- and metepisternum with foveiform punctures. Mesosternum with dense foveiform punctures, almost touching on lateral portion. Mesometasternal suture arched, angulate medially. Metasternum with dense foveiform punctures on lateral and anterior portions; disc smooth, with a strong triangular depression (Figure 1B); anterior portion of depression with a strong spiniform projection; posterior portion with two smaller spiniform projections. Elytra. Elongated, with lateral margins slightly curved outward (Figure 1A). Elytral striae weakly impressed (Figure 1E), with foveiform punctures separated by approximately four or five times their diameter on elytral disc. Interstriae flattened. Interstrial surface with

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setigerous foveiform punctures, randomly spaced. Basal carina of seventh interstria inconspicuous. Basal carina of ninth interstria reaches approximately half the length of elytra. Apical tubercle of second elytral interstria absent. Apical tubercle of third interstria rounded. Apical carina of fifth and sixth interstriae longer than fourth. Apical carina of seventh interstria inconspicuous. Pseudepipleural carina complete, visible for entire length of epipleuron. Abdomen. First sternite expanded medially, occupying 2/3 of the abdominal length (Figure 1F); anterior portion of sternite with two tubercles on its medial portion (Figure 1F). Sternites 2, 3 and 4 not visible medially. Lateral region of sternites 1 – 5 with numerous foveiform punctures closer to anterior margin. Sixth sternite smaller medially than fourth. Pygidium separate from propygidium by carina, with dense setigerous foveiform punctures on its surface, punctures closer along the midline. Pygidium as wide as high, apically rounded, clearly margined. Legs. Protrochanter with a cluster of bristles on the anterior edge. Meso and metatrochanter with a cluster of bristles on the posterior edge (Figure 1B). Femora with numerous foveiform punctures, randomly spaced. Profemur with clusters of bristles on basal third of anterior margin (Figure 1B). Mesofemur widened at the middle portion, with a small denticle on basal third of posterior margin (Figure 1B). Metafemur widened at its apical third. Protibia widened at 2/3 of the internal margin; apical third of the internal margin angularly lobed, with a concave curvature, forming a strong triangular tooth and fourteen spiniform denticles between tibial base and that tooth (Figure 1G). External margin with three teeth on its apical third, apical one larger than basal two (Figure 1G). Meso- and metatibiae arched toward body. Mesotibia with an internal, apical, spur-like denticle, about the length of the first tarsomere; smaller spur shorter than the sum of the length of second and third metatarsomeres; larger spur approximately the sum of three basal tarsomeres in length (Figure 1H). Basal third of metatibia with small denticles on the internal margin (Figure 1A). Internal angle of apex of metatibia prolonged beyond insertion of tarsus in spiniform process, with the same length of tarsus (Figure 2A). Length of metatibial spur less than that of tarsal claws. Protarsus absent. Length of first meso- and metatarsomeres short, about half the second tarsomere, obliquely truncated at apex (Figures 1H, 2A). Tarsal claws without basal teeth. Male genital organ. Symmetrical parameres (Figures 2B, C). Genital sclerite with internal arms elongated and weakly sclerotized (Figure 2D). Endophallus with small basal lobe (Figure 2E). Lamella copulatrix (LC): left and right lobes subrectangular (Figure 2F). Superior right peripheral sclerite (SRP) circular (Figure 2G). Fronto-lateral peripheral sclerite (FLP) and complex of axial and subaxial sclerites (A+AS) as in Figure 2H. Variation: In the collection of the MNHN (Paris), we found a female lacking an abdomen and without locality data. We strongly believe that Paulian (1938) considered this female, 24 mm in length, to be conspecific with D. spinipes specimens due to their body size overlap (see below notes for D. spinipes), because he states in his original description that specimens belonging to this species reach up to 24 mm, and specimens of that size (or near) were found identified as D. spinipes by Paulian in MNHN. Although this female was found in a box of specimens used by Paulian in his revision (Paulian 1938, 1939), since it lacks any label, we cannot be sure that it is a syntype of D. spinipes. Nevertheless, we are confident that this female belongs to the species herein described and have made this unlabeled female the paratype. This female differs from the male holotype in the following aspects: body sheen green, brighter than in holotype. Head with longer clypeal teeth and more protruding clypeo-genal angle. Pronotum smaller in relation to elytra, with four rounded feeble depressions on each side of disc. Metathorax feebly concave on disc, with only indications of anterior and posterior spiniform projections. Profemur less strong, protibiae lacking internal tooth, with reduced basal internal denticles, somewhat dilated in the apical 2/5, and with three external teeth in the apical 2/5, the middle tooth smaller than the others, apical teeth less developed than in male. Meso- and metafemur straighter and weaker than in male. Meso- and metatibia less curved than in male, internal and apical denticles absent. Internal angle of apex of metatibia not prolonged in spiniform process. Middle and hind tibial spurs much longer than in male. Meso- and metatarsi about 1/7 longer than in male. Type material. Holotype: 1 ♂ (white label, handwritten). COLOMBIA: Huila, Nat. Park Las Cuevas, 1,80 dm, 23.V.76, M. Cooper./ 2 (white label, handwritten). Parque Nacional Las Cuevas de los Guácharos, Cloud forest, Forest of Quercus humboldtii/ 3 (white label, printed in black). BMNH {E} 2004275 M. Cooper/ 4 (white label, printed in black). BMNH (E) #943732 / 5 (red label, printed and handwritten in black). HOLOTYPE ♂, Deltochilum gigante sp. nov., Silva and Vaz-de-Mello, des. 2013 (BMNH). Paratype: 1 ♀, no locality label, no abdomen (MNHN). 1 (yellow label, printed in black). PARATYPE./ 2 (white label, printed and handwritten in black) Deltochilum gigante sp. nov., Silva and Vaz-de-Mello, des. 2013 PARAT.

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FIGURE 1. Deltochilum gigante sp. nov. (holotype ♂). (A) dorsal view; (B) ventral view (spiniform projections arrowed); (C) head; (D) hypomera; (E) elytral disc; (F) male abdominal sternites (tubercles arrowed); (G) male protibia; (H) apex of mesotibia (spiniform spurs arrowed).



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FIGURE 2. Deltochilum gigante sp. nov. (holotype ♂). (A) apex of metatibia (arrow shows the internal angle of apex of metatibia prolonged beyond insertion of tarsus); (B) aedeagus (lateral view); (C) aedeagus (dorsal and ventral views); (D) genital sclerite; (E) endophallus; (F) lamella copulatrix (top left lobe; bottom right lobe); (G) superior right peripheral sclerite (SRP); (H) complex of axial and subaxial sclerites (A+AS) (arrow left), fronto-lateral peripheral sclerite (FLP) (arrow right).



Deltochilum (Deltohyboma) spinipes Paulian, 1938 (Fig. 3) Paulian, 1938: 280

Notes. Paulian described this species based on several specimens from his collection as well as the collections of Oberthür, Boucomont, and the British Museum. He designated a pair of specimens from the Oberthür collection as "types" and the others as "cotypes"", but there was no formal Holotype designation and all type material is therefore syntypic. All syntypes in MNHN of D. spinipes have a body length less than 20 mm. However, Paulian states the length for the species as "18-24 mm". Based on this fact, we believe that the 24 mm female found in a separate box (female of D. gigante, see above) was regarded by Paulian as spinipes while describing this species. The type series of spinipes shows variation and we strongly believe it to include more than one species. Therefore, in order to maintain stability of nomenclature, a lectotype is herein designated. Two of the paralectotypes (♀4 and ♀5) appear to be non-conspecific with the remainder of the type series and likely represent two different species. They differ in being blue (not bright green), much darker and dull; one specimen is bigger and duller than the other; both specimens were collected in Ecuador (not Colombia as the remainder of the type series). These two specimens, at a first glance, most likely belong to the species-complex that includes D. aequinoctiale and D. speciosissimum. However, we cannot be sure about the identity of D. aequinoctiale since its type has not yet been located (see Génier 2001). Material examined. Lectotype (here designated): ♂. (old white label, handwritten). Bogota. / 2 (old white label, printed and bordered in black). Ex-Musaeo Mniszech./ 3 (white label, handwritten). D. spinipes n. sp. Type./ 4 (white label, bordered in black) R. PAULIAN Vidit./ 5 (white label, printed in black). MUSÉUM PARIS 1952 coll. R. OBERTHÜR./ 6 (red label, printed in black) SYNTYPE./ 7 (white label, printed in black) MNHN EC2509./ 8 (red label, printed in black) LECTOTYPE./ 9 (white label, handwritten, printed and bordered in black) Deltochilum spinipes Paulian LECTO. F Vaz-de-Mello des. 2013. Figure 3A (copyright MNHN/A. Mantilleri).Paralectotypes: 2 ♂♂, 5 ♀♀ (MNHN): ♂1. 1 (old white label, printed and bordered in black). COLOMBIE COLLECTION LE MOULT. / 2 (white label, handwritten and printed in black). D. spinipes Paul. R. Paulian det./ 3 (white label, printed in black). MUSÉUM PARIS 1952 coll. R. OBERTHÜR./ 4 (red label, printed in black) SYNTYPE./ 5 (white label, printed in black) MNHN EC2508./ 6 (yellow label, printed in black) PARALECTOTYPE./ 7 (white label, handwritten, printed and bordered in black) Deltochilum spinipes Paulian PARALECTO. F Vaz-de-Mello des. 2013. Figures 3B, C, F and G (copyright MNHN/A. Mantilleri). ♂2. 1 (old white label, handwritten). Fusagasugá./ 2 (old white label, handwritten). pretiosum Harold./ 3 (old white label, printed and bordered in black). Ex-Musaeo E. Steinheil./ 4 (white label, bordered in black) R. PAULIAN Vidit./ 5 (yellow label, printed in black) PARALECTOTYPE./ 6 (white label, handwritten, printed and bordered in black) Deltochilum spinipes Paulian PARALECTO. F Vaz-de-Mello des. 2013. ♀1. 1 (old white label, printed and bordered in black). COLOMBIE ANTIOQUIA./ 2 (old white label, printed and bordered in black) COLLON LE MOULT Naturaliste Paris. / 3 (white label, bordered in black) R. PAULIAN Vidit./ 4 (white label, printed in black). MUSÉUM PARIS 1952 coll. R. OBERTHÜR./ 5 (white label, printed in black) Deltochilum spinipes Paulian./ 6 (red label, printed in black) SYNTYPE./ 7 (white label, printed in black) MNHN EC2510./ 8 (yellow label, printed in black) PARALECTOTYPE./ 9 (white label, handwritten, and printed and bordered in black) Deltochilum spinipes Paulian PARALECTO. F Vaz-de-Mello des. 2013. Figures 3D and E (copyright MNHN/A. Mantilleri). ♀2. 1 (old white label, handwritten). Muzo, Columbia./ 2 (old white label, printed and bordered in black). ExMusaeo D. Sharp 1890./ 3 (white label, bordered in black) R. PAULIAN Vidit./ 4 (yellow label, printed in black) PARALECTOTYPE./ 5 (white label, handwritten, printed and bordered in black) Deltochilum spinipes Paulian PARALECTO. F Vaz-de-Mello des. 2013. ♀3. 1 (old white label, printed and bordered in black). COLOMBIE COLLECTION LE MOULT. / 2 (white label, bordered in black) R. PAULIAN Vidit./ 3 (yellow label, printed in black) PARALECTOTYPE./ 4 (white label, handwritten, printed and bordered in black) Deltochilum spinipes Paulian PARALECTO. F Vaz-de-Mello des. 2013.



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FIGURE 3. Deltochilum spinipes. (A) lectotype ♂, dorsal view; (B) paralectotype ♂1, dorsal view; (C) paralectotype ♂1, ventral view; (D) paralectotype ♀1, dorsal view; (E) paralectotype ♀1, ventral view; (F) paralectotype ♂1, aedeagus (lateral view); (G) paralectotype ♂1, aedeagus (dorsal view). Photographic copyright A. Mantilleri (MNHN).



♀4. 1 (white label, printed and bordered in black) Equateur Env. d'Ambato R.P. Irenée Blanc./ 2 (white label, bordered in black) R. PAULIAN Vidit./ 3 (yellow label, printed in black) PARALECTOTYPE./ 4 (white label, handwritten, printed and bordered in black) Deltochilum spinipes Paulian PARALECTO. F Vaz-de-Mello des. 2013./ 5 (white label, handwritten, printed and bordered in black) DIFFERENT of D spinipes lectot. F Vaz-deMello des. 2013. ♀5. 1 (old white label, handwritten) Ecuador Buckley./ 2 (white label, printed and bordered in black) ExMusaeo H.W.Bates 1892./ 3 (white label, bordered in black) R. PAULIAN Vidit./ 4 (yellow label, printed in black) PARALECTOTYPE./ 5 (white label, handwritten, printed and bordered in black) Deltochilum spinipes Paulian PARALECTO. F Vaz-de-Mello des. 2013./ 6 (white label, handwritten, printed and bordered in black) DIFFERENT of D spinipes lectot. F Vaz-de-Mello des. 2013.

Taxonomic remarks An identification key for the species of the subgenus Deltohyboma is not given here because the subgenus is in need of a taxonomical revision and many of its species cannot be reliably identified at this time. Many of the species were superficially described and/or their type specimens have not been examined. The present situation makes the identification of some species impossible at this moment. However, the description, diagnosis, and illustrations of male genitalia and somatic morphology provided here give sufficient information for the reliable identification and differentiation of this new species from all others in the genus.

Acknowledgements Our special thanks are due to Max Barclay for loan of the BMNH specimens, and to Olivier Montreuil and Antoine Mantilleri for providing facilities to FZVM during his stay in Paris, Stéphane Boucher for important historical background, Antoine Mantilleri for photography, Gustavo Ruiz, Adam Brunke, Max Barclay and Sergei Tarasov for valuable suggestions on the text of the manuscript. We thank the Hymenopterist Martin Cooper for presenting very interesting material of Neotropical Coleoptera, including the holotype of our new species, to the BMNH. We also thank CAPES/Pro-equipments, for providing the photographic equipment, and CNPq for providing PQ2 fellowships and research grants for FZVM (304925/2010-1, 302997/2013-0, 405697/2013-9, 484035/2013-4, 202327/2013-2). This research received support from the Synthesys Project http://synthesys3.myspecies.info/ which is financed by the European Community Research Infrastructure Action under the FP7 (GB-TAF 3855).

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