Gromphas lacordairei : Hamel-Leigue et al., 2006: 6 Hamel-Leigue et al., 2009: 61 Gromphas amazonica : Cupello and Vaz-de-Mello 2013: 463 A new species and the phylogeny of the South American genus Gromphas Brullé, 1837 (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini) Cupello, Mario Vaz-de-Mello, Fernando Z. Journal of Natural History 2015 2015-09-30 50 943 969   Gromphas amazonica : Cupello and Vaz-de-Mello 2013: 463   6KSH6 Cupello Cupello [341,656,1164,1189] Insecta Scarabaeidae Gromphas GBIF Animalia Coleoptera 3 946 Arthropoda species jardim   ( Figures 1 –7)      Gromphas lacordairei: Hamel-Leigue et al., 2006: 6, fig. 49,50;  Hamel-Leigue et al., 2009: 61(part), 49 (part), fig. 14 (part), figs 27,28.     Gromphas amazonica: Cupello and Vaz-de-Mello 2013: 463(all the fifth paragraph of ‘Intraspecific variation and taxonomic discussion ’section).    Type specimens   Holotype:  BOLÍVIA: BENI: Moxos, Río Ichiguita,  155 m, 15°08 ’S, 65°18 ʹW,  20.V.2005, C. Hameland T. Vidaurrecols. –male ( OUMNH) [ “ BOLIVIA: Beni, Rio Ichiguita,   155 m., 15º08 ʹS 65º18 ʹO,  20.v.2005, Sabana. Trap.cebo heces humano. prep./col.: C. Hamel, T. Vidaurre ”, “  Gromphas lacordaireiBrullé, 1834det. A. C. Hamel.OUMNH-2006-097 ”, “ Trap7 ”, “HOLOTIPO ”, “ HOLOTYPE.  Gromphas jardim sp. nov. Cupello& Vaz-de-Mellodes. 2014 ♂ ”] ( Figure 1A –D, F). Aedeagusextracted and genital capsule glued in a triangular label and internal sac placed in a microvial with glycerine, all pinned with the holotype.   Figure 1.  Gromphas jardim sp. nov.(A –D) Holotype. (A) Dorsal view. (B) Lateral view. (C) Aedeagus. From left to right, dorsal, lateral and ventral views. (D) Labels. (E) Female paratype from Bolivia. (F) Dorsal view of head (holotype). (G) Frontal view of cephalic projection (male paratype).   Figure 2.Protibia of  Gromphas jardim sp. nov.(A, B) Male protibia: (A) Ventral view, (B) Dorsal view. Red arrow indicates apical tubercle modified in a tapered spur. (C) Female protibia. Note that the apical tuft of setae is much denser and longer in male than in female.   Figure 3.Lateral view of pronotum. (A)  Gromphas jardim sp. nov.(B)  Gromphas amazonica. Note that the granulation extends much more posteriorly in  G. jardim, reaching the pronotal posterior margin, than in  G. amazonica, wherein the granulation is absent or rudimentary in posterolateral region after lateral fossa.   Figure 4.Male paratype of  Gromphas jardim sp. nov.and its labels (braces ‘{ ’joining two sides of a same label). We believe that the French zoologist and explorer Alcide d ’Orbigny was the collector of this specimen in 1832 (see the text).   Paratypes:  BRAZIL: MATO GROSSO: Cáceres,  10 October 2008, E . Silvacol . – 1 female( CEMT; specimen identified in Cupello and Vaz-de-Mello (2013, pp. 463 –464) as  G. amazonica).   BOLÍVIA: BENI: Moxos, Río Ichiguita,  155 m, 15°08 ’S, 65°18 ʹW,  19 May 2005, C.  Hameland T.  Vidaurrecols. –  1 female( MNRJ) and 1 female( OUMNH).  COCHABAMBA: Territoryof the Yuracarépeople ( “ Juacares Indians ”), north side of the Cordillera de Cochabamba( “ Cortillera de Cochabamba ”), without date and collector (probably collected by Alcided ’ Orbignyin 1832; see comments below) – 1 male( BMNH).    Etymology The specific name, a noun in apposition, is a patronym honouring Arlindo da Silva Jardim ( 1923 –2014), Brazilian aviator and grandfather of the first author. Having grown up in the small, rural village of Dom Viçoso, Minas Gerais, Arlindo Jardim achieved his childhood dream and flew professionally worldwide for over four decades. He will remain as a source of inspiration for MC.    Description  Colour. Anterior region of clypeus black; remainder of head and pronotum with dark olive green and copper metallic reflections. Elytra, metasternum, ventral surface of legs and pygidium dark olive green with metallic sheen and silky appearance. Ventrites entirely black or black with week metallic green reflections.  Head.Margin of clypeus with four lobes ( Figure 1F) and distinctly upturned. Genae and frons completely granulate, including region adjacent to eyes ( Figure 1F). Cephalic projection a raised carina with converging sides and emarginate apex in major specimens ( Figure 1G); apex narrower than distance between apices of apical lobes of clypeus ( Figure 1F).  Thorax.Pronotum convex; lateral region with dense granulation reaching the posterior margin ( Figure 3A), density of granulation decreasing posteromedially; posteromedian region smooth or with strongly effaced granulation ( Figures 1A, E, 4); posterior fossae apparent only as two very shallow and sometimes only weakly indicated impressions removed from the pronotal posterior margin ( Figures 1E, 4). Posterior margin of pronotum rounded. Mesosternum with dense pilosity. Metasternum with fine and sparse punctation at centre. Anteromedian angle of metasternum convex and with globose apex; area in front of anteromedian angle with evident setae.  Legs.Protibia slightly narrower in males than in females ( Figure 2); in ventral view, longitudinal carina simple in both sexes ( Figure 2A). Space between protibial lateral teeth deeper in males than in females ( Figure 2). Protibial spur with apex strongly expanded and curved downward ( Figure 2). Inner apical angle of protibia with a tuft of setae longer and denser in males than in females; in males, tubercle of inner apical angle developed as a short and tapered spur independent of apical tuft of setae ( Figure 2). Apical protarsomere with a long, distal spiniform prolongation. Mesotarsi and metatarsi with apical tarsomere slightly curved at apex. Metatibia very broad and robust. Metatibial spur with apex distinctly curved.  Elytra. Striae very fine and, especially striae 1 –4, carinulate from base to half or apical two-thirds of elytra. Sutural margin glossy and only sparsely punctate; basal half of sutural margin with sheen extending laterally onto first or second interstria.  Abdomen. Pygidium lacking basal margin and with irregular sculpture. Groove of propygidium extending to base of pygidium. Abdominal sternites microsculptured and sparsely punctuated.  Aedeagus.Apex of phallobase, in ventral view, with membranous area expanded triangularly in the middle ( Figure 1C). Medial sclerite only slightly curved, almost flat.   Measurements (four specimens: two males and two females)  TL: AV: 15.2; MX: 16.3; MN: 13.9. PL: AV: 12.3; MX: 13; MN: 11.7. PW: AV: 8.5; MX: 9; MN: 8.   Intraspecific variation and taxonomic discussion At a first glance,  G. jardimresembles superficially  G. amazonicaand, to a lesser degree,  G. inermis, and, in fact, has been confused with these species both in collections and recent publications. Deposited at the BMNH, the oldest specimen known to us bears four labels with different identifications ( Figure 4): an older, which by the calligraphy we assign to Charles O. Waterhouse, former curator of entomology at the BMNH, has written ‘Gormphas amazonicusBates ’, while the other two more modern labels identify that specimen, respectively, as  G. amazonicaand ‘  G. lacordaireiBrullé, 1834 ʹ, an unavailable name referring to  G. lacordairii Burmeister, 1874, junior synonym of  G. inermis(see more in Cupello and Vaz-de- Mello 2013). The fourth label has handwritten the word ‘Coproides’, but the remaining information is effaced and completely unreadable. d ’ Olsoufieff (1924)examined a specimen of  G. amazonicain the Muséum national d ’Histoire naturelle, Paris, labelled ‘coproidesDej. Cayenne (coll. Mniszech) ’and probably the unavailable name ‘coproides’was used before the description of  G. amazonicaby Bates (1870)as a name in litteristo refer to this species. The three specimens found in OUMNH, in turn, including the holotype, are part of a large series of dung beetles recently collected in Boliviaand the basis for the works of Hamel-Leigue et al. (2006, 2009); they were identified and illustrated in these publications as ‘  G. lacordaireiBrullé ’. The geographical distribution and probably the other information present for ‘  G. lacordaireiBrullé ’in Hamel-Leigue et al. (2006, 2009) have mixed data belonging in fact to  G. jardimand  G. inermis. Similarly, as said in the Introduction of the present work, in Cupello and Vaz-de-Mello (2013), we provisionally identified the specimen (now paratype) from Cáceres as a  G. amazonica. Now, in possession of a greater number of specimens, the differences between  G. jardim, G. inermisand  G. amazonicabecame much clearer.   Gromphas jardimshares only with  G. amazonica, G. inermisand  G. dichroathe characters: genae and frons granulated adjacent to eyes ( Figure 1F), absence of pronotal prominence, protibiae narrower in males than in females ( Figure 2), and protibial spur expanded at apex ( Figure 2); only with  G. amazonicaand  G. inermis, G. jardimshares the character margin of clypeus with four lobes ( Figure 1F). Probably this last characteristic, which is an apomorphy shared by them (see the phylogenetic analysis below), was the main cause for the past misidentifications. Yet  G. jardimis easily differentiated from  G. inermisby having metatibial spur distinctly curved apically (straight in  G. inermis), posterior margin of pronotum rounded (projected at middle in  G. inermis), elytral striae carinulate (simple in  G. inermis), and metasternum and sutural margin of elytra with fine and sparse punctation (dense punctation in  G. inermis); furthermore, pronotal hump and sutural margin of elytra raised are present in major specimens of  G. inermisbut absent in  G. jardim( Figure 1B). On the other hand, the medial sclerite of the internal sac of  G. jardimis very similar to that of  G. inermisand no significant difference between them was found ({fig. 59}). From  G. amazonica, G. jardimis differentiated most easily by the shape of the apical tubercle of male protibia, which, although much more developed in  G. jardimthan the tiny and almost imperceptible tubercle of the other four species of  Gromphas, is still much smaller than that of  G. amazonica; in  G. jardim, the tubercle has the shape of a tapered spur and is separated from the apical tuft of setae, which rests adjacent to the spur ( Figure 2A,B); in  G. amazonica, the spur is long, laterally flattened and curved and has the tuft of setae on its dorsal surface as a row of setae. The shape of the cephalic projection of  G. jardimis similar to that of  G. inermis, i.e. it is narrower than the distance between the apices of the apical lobes of clypeus ( Figure 1F), while that of  G. amazonicahas the equivalent width of that distance. Other differences between  G. jardimand  G. amazonicaare: the colour, which is dark olive green and has metallic reflections in  G. jardim, but black, dark blue, dark green or reddish-brown and never has metallic reflections in  G. amazonica; and the pronotal granulation, which penetrates more the posterior portion of the pronotum and, in lateral view, reaches the posterior margin in  G. jardim( Figure 3A), whereas in  G. amazonicathe granulation is restricted to the anterior portion of the pronotum and never reaches the posterior margin ( Figure 3B). The form of the granules of the head and pronotum is also distinct between the two species, being wider and flattened in  G. amazonicaand more rounded and smaller in  G. jardim(this second form is very similar to that of  G. inermis). Finally, the longitudinal carina of the ventral surface of protibia is simple in both sexes of  G. jardim, resembling  G. aeruginosaand  G. lemoinei, but is distinct to that of  G. amazonica, G. inermisand  G. dichroa, which, in males, has a row of tubercles on its basal half and, in females, is simple. The constant presence of the posterior pronotal fossae in  G. jardimalso distinguishes this species from  G. inermisand  G. amazonica, in which these fossae are usually absent. The spiniform projection at the apex of apical protarsomere was not observed in one of the three females of  G. jardimexamined by us, and we believe that this is due to the wear, as happens in some  G. amazonica, the only other species of  Gromphasthat has this kind of apical protarsomere ( Cupello and Vaz-de-Mello 2013). On the other hand, the nature of the posterior pronotal fossae varies: in the two males observed, the fossae are clearly marked and easily visible to the naked eye, whereas those of the three females are much less marked and almost imperceptible. Whether this is a case of individual or sexual variation is difficult to say until the examination of a larger number of specimens of both sexes.   Geographic distribution  Brazilian subregion: South Braziliandominion: Rondônia province.   BRAZIL: MATO GROSSO: Cáceres.   BOLÍVIA: BENI: Moxos. COCHABAMBA: ‘Territory of the Yuracaré people, north side of the Cordillera de Cochabamba ’( Figure 5).    Comments While the holotypeand the three female paratypesof  G. jardimwere collected in the 21st century and have label information detailed enough to permit an easy understanding of their origin, the male paratypeis much older and has a puzzling history. Only one of the five labels attached to this specimen before our work has information about its provenance ( Figure 4). This label is circular and has ‘ Bolivia ’written on one side and ‘46/76 ʹ, on the other side. According to Max Barclay (pers. comm.), ‘ 1846 –76refers to a collection, all with the same data, acquired in 1846 and including 325 Coleopteraand 250 Lepidoptera ’, and these collecting data are ‘Territory of Juacares Indians (north side of the Cortillera ( sic) de Cochabamba) ’. We believe that ‘Juacares Indians ’refers to the Yuracaré, an indigenous people resident on the north side of the Cordillera de Cochabamba, in the department of Cochabamba, Bolivia. To our knowledge, the only European naturalist who crossed this remote region before 1846 was the French zoologist and explorer Alcide d ’Orbigny ( 1802 –1857), who visited a Yuracaré village on May 28 1832and stayed there for 4 days ( Papavero 1971). Indeed, d ’Orbigny described and illustrated in detail this people in his great work Voyage dans l’ Amérique Méridionale(d ’ Orbigny 1835 –1847). So we believe he was the probable collector of the male paratypeof  G. jardim. This finding is also interesting because, if correct, it indicates that not all insects collected by Alcide d ’Orbigny are deposited in the Muséum National d ’Histoire Naturelle, Paris, France, as suggested by Horn and Kahle (1936)and Evenhuis (1997).   Figure 5.Updated distribution of the six species of  Gromphas.   Bionomics   Thelabel data indicate that the holotypeand the two paratypesfrom Beni, Bolivia, were collected in trapsbaited with human faeces in open habitats.  Thesespecimens also had some unidentified phoretic mites attached to their legs, especially to the metatarsi.  Therecorded months for  G . jardimare May and October. 2831865328 2005-05-20 OUMNH C. Hamel & T. Vidaurre & Beni & Sabana. Trap. & Hamel, T & Trap & Cupello & Vaz-de-Mello & Aedeagus Bolivia A. C. Hamel. 155 -15.133333 Rio Ichiguita 1286 65.3 Moxos 3 946 156 1 Beni holotype 2831865440 [160,909,977,1001] 2008-10-10 Caceres Brazil MATO GROSSO 6 949 1 Mato Grosso paratype 2831865404 [908,1019,977,1001] 2008-10-10 Silva Brazil true MATO GROSSO 6 949 1 Mato Grosso paratype 2831865402 CEMT Brazil 6 949 1 1 Mato Grosso paratype 2831865412 2005-05-19 Bolivia 155 -15.133333 Rio Ichiguita 1286 -65.3 Moxos 6 949 1 Beni paratype 2831865357 [271,432,1081,1105] Bolivia Hamel 6 949 1 Beni paratype 2831865382 [449,606,1081,1105] Bolivia Vidaurre 6 949 1 Beni paratype 2831865318 [651,1152,1081,1105] MNRJ, OUMNH Bolivia 6 949 2 2 Beni paratype 2831865476 BMNH Territory & Yuracare & Juacares Indians & de Cochabamba & Alcide & Orbigny Bolivia Cortillera de Cochabamba Cordillera de Cochabamba 6 949 1 1 Cochabamba paratype 2831865369 [160,957,975,999] South Brazilian 9 952 1 Rondonia 2831865378 Brazil MATO GROSSO 9 952 1 Mato Grosso 2831865307 [382,649,1009,1033] Bolivia BENI 9 952 1 Beni 2831865333 Bolivia The 10 953 2 Beni holotype 2831865444 Bolivia These 10 953 1 Beni holotype 2831865464 Bolivia The 10 953 1 Beni holotype