Dendropaemon (Titthopaemon) denticollis Felsche, 1909

François Génier & Patrick Arnaud, 2016, Dendropaemon Perty, 1830: taxonomy, systematics and phylogeny of the morphologically most derived phanaeine genus (Coleoptera: Scarabaeidae, Scarabaeinae, Phanaeini), Zootaxa 4099 (1) : 81-85

41. Dendropaemon (Titthopaemon) denticollis Felsche, 1909

( Figs. 41 , 108–109, 133, 155, 161)

Dendropemon denticollis Felsche 1909 , Deut. Ent. Zeit. 1909: 758 (original description)

Dendropemon denticollis: Gillet 1911 , Col. Cat. 38: 88 (catalogue)

Dendropaemon denticollis: Olsoufieff 1924 , Insecta 13: 129 (monograph)

Dendropaemon denticollis: Pessôa & Lane 1936 , Rev. Biol. Hygiene 7: 90 (identification key, comment) Dendropaemon denticollis denticollis: Blut 1939 , Arch. Naturg. (N.F.) 8: 276 (monograph) Dendropaemon denticollis lividus Blut 1939 , Arch. Naturg. (N.F.) 8: 276 (original description) new synonymy Dendropaemon denticollis: Pessôa & Lane 1941 , Arq. Zool. S. Paulo 2: 491 (identification key, distribution) Dendropemon denticolle: Blackwelder 1944 , U.S. Nat . Mus. Bull. 185: 210 (checklist)

Dendropemon denticolle lividum: Blackwelder 1944 , U.S. Nat . Mus. Bull. 185: 210 (checklist) Dendropaemon (D.) denticollis: Edmonds 1972 , Univ. Kansas Sc. Bull. 49: 850 (comment taxonomy) Dendropaemon denticolle : Vaz-de-Mello 2000, Hac. Proy. CYTED: 192 (faunistic)

Dendropaemon (D.) denticollis: Arnaud 2002 , Col. Monde 28: 15 (mention)

Dendropaemon denticollis: Philips et al. 2004 , Insect Syst. Evol. 35: 51 (phylogeny)

Dendropaemon denticolle: Barbosa 2008 , Comm. Scar.: 19 (biology)

Dendropaemon (D.) denticollis : Vaz-de-Mello & Génier 2009, Col. Bull. 63: 364 (biology) Dendropaemon denticollis lividus: Hamel-Leigue et al. 2009 , Kempffiana 5: 49 (faunistic)

Type locality. Jatahy, Provinz Goyas.

Diagnosis. The only species in the genus with a denticle on the anterior pronotal margin adjacent to the eyes.

Description. Female holotype. Body. Body small, length 9.0 mm, maximum width 4.5 mm; body subrectangular in dorsal view; dorsum narrowly flat. Color. Dorsal surface brown, glossy, with green metallic sheen; head brown along anterior edge of clypeus, metallic green on remaining surface; pronotum with green metallic sheen except on small areas along anteromedian carina and adjacent to lateral fossae; elytra with uniform green metallic sheen; ventrum light to dark brown; pygidium with greenish metallic sheen; legs light to dark brown. Head. Clypeus broadly arcuate, anterior portion upturned; clypeal teeth acutely triangular; clypeal median emargination broadly u-shaped, clypeal edge emarginate on external side of each clypeal tooth, clypeal teeth ventral surface lacking carina, clypeal margin ill-defined, lacking sharp carina posteriorly, clypeal surface with short and blunt transverse rugulae; clypeogenal suture well-defined, bluntly carinate internally; genal surface with small and blunt irregular tubercles, lacking distinct transverse carina, simply convex; clypeofrontal carina rather low, approximately 4 times wider than high, straight in dorsal view, simply carinate, clypeofrontal carina apical edge straight in frontal view; eyes large in dorsal view, interocular ratio 2.7. Pronotum. Pronotum transverse in dorsal view, pronotal width/length ratio 1.4; disc of pronotum minutely punctate throughout, with a sharply defines longitudinal sulcus on posterior half; pronotal anterior margin wider and flat lateral to eyes; anterior portion with a broad sharp and nearly straight carina medially; anterior angles surface with blunt rugulae, nearly smooth along anterior margin, similar to lateral margin along posterior edge of anterior margin; lateral fossae oval, simple; lateral portions slightly explanate; pronotal basal fossae very small and rounded; posterior margin ill-defined, lacking setae. Elytra. Elytra approximately as long as wide in dorsal view, elytral combined width/length ratio 0.9; elytral base lacking distinct margin, simply convex; elytral striae 1–4 moderately wide, evenly impressed throughout, elytral striae 5 similar to 4 on disc, strial punctures minute, adjacent strial edge encroaching on interval, stria 1 weakly impressed apically, going straight to elytral apical margin; interstriae slightly convex, minutely punctate throughout, surface glossy. Thoracic sterna. Proepisternal carina absent; metasternal median lobe bluntly angularly produced anteromedially, ventral ridge ill-defined, v-shaped. Legs. Profemur posterior surface convex, glabrous and punctate internally, posterointernal margin rather thick, evenly developed, internal edge rather wide, with a contiguous row of setae along anterointernal edge, remaining surface smooth. Protibia with four teeth on lateral edge; internal basal angle bluntly lobate; anterior surface with long aligned row of setae internally, surface glossy or feebly microsculptured between punctures; posterior surface with some ill-defined irregular punctures externally to median carina, surface finely and irregularly microsculptured between punctures, with a single interrupted setal row along lateral teeth. Mesofemur unmodified on anterointernal edge apically. Mesotibia rather short, gradually widening toward apex in anterior view; anteroapical edge slightly sinuate in anterior view, anteroapical row of setae complete; apicoanterior edge circularly indented internally; external edge more or less rounded, with several large elongate setiferous punctures. Mesotarsus similar in shape to metatarsus, 3-segmented, first segment moderately elongate, approximately two times as long as wide at apex. Metafemur internal edge nearly straight and lateral edge arcuate, lacking distinct depressed area anterointernally before apex, apicoposterior edge unmodified, anterior surface with a well-defined sulcus on apical half. Metatibia moderately slender, slightly widening toward apex in anterior view, anterior surface with distinct row of setae, surface glossy, metatibial posterior surface flat between longitudinal row of setae and lateral edge, with ill-defined microsculpture. Metatarsus 3-segmented ( Fig. 133 ), first segment short, approximately as long as wide at apex, with anterointernal carina well defined and almost reaching apical edge. Abdominal sternites. Sternites 3–6 longitudinally flat; sternites 4–6 with 1–3 unaligned rows of setae laterally, glabrous medially; sternite 7 approximately longitudinally flat medially, subequal in length to segment 6 along midline; pygidium finely punctate on disc.

Measurements (65 males, 106 females). Length: male 7.0–10.5 (8.7±0.7), female 7.5–10.5 (9.0±0.7) mm.

Primary type data.

Dendropaemon denticollis Felsche (Fig. 155). Holotype female (MTD): [Jatahy/ Prov.Goyas]; [Coll. C. Felsche/ Kauf 20, 1918] green card; [Typus] red card; [Staatl. Museum für/ Tierkunde Dresden]; denticollis / m./ Paraguay] handwritten; [WORLD/ SCARAB./ DATABASE/ WSD 00016835]; [HOLOTYPE / Dendrop[a]emon / denticollis / Felsche, 1909] red card; [ Dendropaemon ♀/ denticollis / Felsche, 1909 / vid. Génier & Arnaud, 2009].

Dendropaemon denticollis lividus Blut. Holotype male (ZMHB): [ BOLIVIA / S Cruz de la Sierra] partly handwritten; [♂]; [Typus] red card; [ Dendropaemon / dentic. lividus/ Boliv. ♂ Type.] handwritten, black border; [HOLOTYPUS / Dendropaemon / denticollis lividus Blut, 1939 / labelled by MNHUB 2009] red card; [WORLD/ SCARAB./ DATABASE/ WSD00016585]; [ Dendropaemon ♂/ denticollis / Felsche. 1909 / deét. Génier & Arnaud,2009]

Material examined. BOLIVIA: SANTA CRUZ, Provincia Andrés Ibáñez, Santa Cruz de la Sierra, (17°48'S, 63°10'W), [no date], coll. [anonymous]— 1 male (holotype) ( ZMHB); Río Ichilo, elev. 350 m, X.1926, coll. F. Steinbach— 1 female ( CMNC); SANTA CRUZ, Yoay, (20°17'S, 63°6'W), xii.1932, coll. [anonymous]— 1 female ( CAS); Provincia Andrés Ibáñez, Santa Cruz de la Sierra, (17°48'S, 63°10'W), xi.1910, coll. J. Steinbach— 1 male ( WDEC); Provincia Chiquitos, Santiago, (18°20'17''S, 59°35'37''W), xi.1959, coll. [anonymous]— 1 female ( CMNC); Provincia Sara, [unspecified locality], [no date], coll. Steinbach— 1 male ( CMNC); BRAZIL: same locality, [no date], coll. [anonymous]— 1 female, 2 males ( IRSNB, MNHN); DISTRITO FEDERAL, Brasilia, elev. 1100 m (15°47'S, 47°55'W), iv.2000, coll. N. Dégallier— 1 female ( CMNC); same locality, xii.2000, coll. N. Dégallier— 1 female ( CMNC); same locality, xi.1999, coll. N. Dégallier— 1 male ( CMNC); same locality, iv.1999, coll. N. Dégallier— 1 male ( CMNC); same locality, ix.1999, coll. N. Dégallier— 1 male ( CMNC); same locality, iii.2000, coll. N. Dégallier— 2 males ( CMNC); GOIÁS, Aragarças, (15°53'50''S, 52°13'48''W), 30.iii.1953, coll. M. Alvarenga— 1 male ( GHCM); Aruanã, Rio Araguaya, (14°55'27''S, 51°4'37''W), v.1967, coll. Dirings— 1 female ( MZSP); Bom Jardim de Goiás, (16°12'10''S, 52°10'32''W), ii.1998, coll. Vaz-de-Mello— 1 female ( CEMT); Campinas, (14°18'52''S, 49°9'30''W), ii.1938, coll. Dr. Nick— 1 female ( CMNC); same locality, i.1938, coll. Dr. Nick— 1 female ( CMNC); Goiânia, (16°41'S, 49°15'W), xii.1992, coll. Crossare— 1 female ( CPFA); same locality, 20.ii.1986, coll. P.B. Silva— 1 female ( CEMT); Goiatuba, (18°0'40''S, 49°22'10''W), i.1947, coll. Guérin— 1 female ( CMNC); same locality, ii.1947, coll. J. Guérin— 1 female ( CMNC); Jataí, (17°53'S, 51°43'W), [no date], coll. [anonymous]— 6 females, 2 males ( BMNH, IRSNB, MTD, ZMHB); Leopoldo de Bulhões, (16°36'52''S, 48°44'26''W), xi.1937, coll. Dr. Nick— 1 female ( BDGC); Mineiros, (17°34'43''S, 52°32'32''W), [no date], coll. [anonymous]— 1 female ( MNHN); Niquelândia, (14°27'S, 48°27'W), x.1994, coll. [anonymous]— 1 female, 1 male ( WDEC); Rio Verde, (17°47'50''S, 50°54'0''W), xii.1945, coll. Zellibor— 1 female ( CMNC); Trindade, (16°38'52''S, 49°29'53''W), [no date], coll. Ch. Pujol— 1 female, 1 male ( MNHN); [unspecified locality], [no date], coll. Baer— 1 female ( MNHN); MARANHÃO, Posto Avançado do Mel, Parque Estadual Mirador, Mirador, (6°43'50''S, 44°58'59''W), 30–31.v.2011, coll. F. Limeira-de-Oliveira, A.A. Santos & T.T.A. Silva— 1 male ( CEMT); MATO GROSSO, Bela Vista, (16°2'S, 56°53'W), 23.iv.1957, coll. J. Toledo— 1 female ( MNRJ); Cuiabá, (15°35'45''S, 56°5'49''W), [no date], coll. [anonymous]— 2 females ( ZMHB); Diamantino, Alto Rio Arinos, x.1999, coll. E. Furtado— 2 females, 2 males ( CEMT); E. Furtado casa, Vale de Solidão, Diamantino, (14°22'14''S, 56°7'59''W), 21.x.2008, coll. E. Furtado— 1 female ( CEMT); same locality, 24.x.2008, coll. E. Furtado— 5 females, 4 males ( CEMT); Fazenda Mutuca, Cuabá, (15°18'52.2''S, 55°58'13.08''W), 20.xii.2008, coll. [anonymous]— 1 female ( CEMT); Fazenda Santhidi, Distrito da Guia, Municipio de Cuiabá, 4.x.2008, coll. L.R. Silva— 1 female ( CEMT); Fazenda Santidi, Cuiabá, elev. 250 m (15°23'6''S, 56°6'42''W), 28.ix.2008, coll. L.R. Silva— 1 female, 1 male ( CEMT); Ferreiros, i.1947, coll. [anonymous]— 1 female ( GHCM); Poconé, (16°15'25''S, 56°37'29''W), 21.x.1953, coll. [anonymous]— 1 female ( CPFA); Rosário Oeste, (14°50'S, 56°25'W), i.1972, coll. Dirings— 1 female ( CMNC); Sul de Mato Grosso, ii.1957, coll. Salvarenga— 1 female ( CMNC); Vale da Solidão (area 2), Municipio Diamantino, (14°21'50''S, 56°7'23''W), ii.2001, coll. E. Furtado— 9 females, 2 males ( CEMT); same locality, iii.2001, coll. E. Furtado— 5 females, 5 males ( CEMT); Vale da Solidão, Municipio Diamantino, (14°21'52''S, 56°7'23''W), 27.ii.2009, coll. D.C.T. Oliveira— 1 female ( CEMT); MATO GROSSO DO SUL, Campo Grande, (20°26'30''S, 54°39'0''W), x.1947, coll. A. Maller— 1 male ( AMNH); same locality, iii.1986 – iii.1987, coll. Bianchin et al.— 1 male ( CEMT); Corghinho Quinta do Sol, (19°49'57''S, 54°49'45''W), ii.2011, coll. L. O. Bavutti— 3 males ( CEMT); Corumbá, (19°0'35''S, 57°39'17''W), i.1946, coll. J. Guérin— 1 male ( CMNC); Fazenda Califórnia, Bodoquena [1], (20°41'46''S, 56°52'55''W), iii.2011, coll. L. O. Bavutti— 1 male ( CEMT); Fazenda Califórnia, Bodoquena [2], (20°41'4''S, 56°51'36''W), iii.2011, coll. L. O. Bavutti— 1 male ( CEMT); Fazenda Califórnia, Bodoquena [3], (20°41'5''S, 56°51'33''W), iii.2011, coll. L. O. Bavutti— 1 female ( CEMT); Horto Nova Palmito, International Paper, Três Lagoas, 5.iii.1993, coll. C.A.H. Flechtmann— 1 female (MEFEIS); same locality, 5.iii.1993, coll. C.A.H. Flechtmann— 1 male (MEFEIS); Horto Rio Verde, Três Lagoas Agroflorestal, Três Lagoas, (20°55'21''S, 52°8'21''W), 26.x.1993, coll. C. Flechtmann— 1 female ( CEMT); same locality, 9.xi.1993, coll. C. Flechtmann— 1 female ( CEMT); Salobra, (12°14'S, 57°8'W), x.1941, coll. J. Guérin— 1 male ( GHCM); Três Lagoas, (20°45'35''S, 51°41'42''W), xii.1982, coll. K. Hudepohl— 2 females ( CPFA); UNESP Farm [=Fazenda Experimental da Universidade Estadual Paulista, câmpus de Ilha Solteira], Selvíria, (20°22'55.41''S, 51°24'39.3''W), 23.x.1989, coll. C.A.H. Flechtmann— 1 male (MEFEIS); same locality, 12.ix.1991, coll. C.A.H. Flechtmann— 1 female (MEFEIS); same locality, 10.x.1991, coll. C.A.H. Flechtmann— 8 females, 2 males (MEFEIS); same locality, 17.x.1991, coll. C.A.H. Flechtmann— 1 female (MEFEIS); same locality, 22.x.1991, coll. C.A.H. Flechtmann— 1 male (MEFEIS); same locality, 21.xi.1991, coll. C.A.H. Flechtmann— 1 female (MEFEIS); same locality, 9.v.1992, coll. C.A.H. Flechtmann— 1 male (MEFEIS); same locality, 10.x.1992, coll. C.A.H. Flechtmann— 1 male (MEFEIS); same locality, 23.x.1992, coll. C.A.H. Flechtmann— 2 females, 1 male (MEFEIS); same locality, 3.i.1993, coll. C.A.H. Flechtmann— 1 male (MEFEIS); same locality, 30.v.1993, coll. C.A.H. Flechtmann— 1 male (MEFEIS); same locality, 21.iii.1996, coll. C.A.H. Flechtmann— 1 female (MEFEIS); same locality, 18.iv.1996, coll. C.A.H. Flechtmann— 1 female (MEFEIS); same locality, 17.ix.1997, coll. C.A.H. Flechtmann— 1 female ( CEMT); same locality, 7.x.1999, coll. C.A.H. Flechtmann— 1 female (MEFEIS); same locality, 6.iv.2000, coll. C.A.H. Flechtmann— 1 male (MEFEIS); same locality, 7.ix.2000, coll. C.A.H. Flechtmann— 1 female ( CEMT); same locality, 21.ix.2000, coll. C.A.H. Flechtmann— 1 female ( CEMT); same locality, 23.xi.2000, coll. C.A.H. Flechtmann— 1 female ( CEMT); same locality, 29.ix.2005, coll. V.G. Tabet— 1 male (MEFEIS); same locality, 9.i.2007, coll. F.W. Mesquita— 1 male (MEFEIS); same locality, 16.ix.2009, coll. C.A.H. Flechtmann— 1 male (MEFEIS); same locality, 4.ii.2010, coll. C.A.H. Flechtmann— 1 female (MEFEIS); Usina Santa Helena, Nova Andradina, (21°59'32''S, 53°26'28''W), 18.x.2012, coll. G. Coutinho— 2 males ( CEMT); MINAS GERAIS, Araxá, (19°35'36''S, 46°56'27''W), xi, coll. [anonymous]— 1 female ( MNRJ); same locality, [no date], coll. [anonymous]— 1 female ( MTD); Barro Preto, Belo Horizonte, (19°55'18''S, 43°57'7''W), [no date], coll. Ch. Pujol— 1 female ( MNHN); Conceição dos Ouros, (22°25'S, 45°48'W), ii.2002, coll. [anonymous]— 1 female ( COBF); Fazenda do Riacho Fundo, Campos de Diamantina, xii.1902, coll. E. Gounelle— 1 female ( MNHN); Fazenda Pontinha, Cordisburgo, elev. 700 m (19°8'53''S, 44°12'1''W), i.1994, coll. F.Z. Vaz de Mello— 1 female ( MZLU); same locality, 6.i.2001, coll. F. Vaz de Mello (2001-01)— 1 male ( CMNC); Irara, [no date], coll. Dr. Bach— 2 males ( ZMHB); Lavras, (21°14'45''S, 44°59'59''W), 11.xi.2004, coll. R.C. Avelar— 1 female ( CEMT); Uberaba, (19°45'S, 47°56'W), [no date], coll. [anonymous]— 2 females, 2 males ( CPFA, IRSNB); SÃO PAULO, Araras, (22°21'28''S, 47°23'6''W), 15.iv.1981, coll. J. Borges— 1 male ( CMNC); Batatais, (20°53'30''S, 47°35'0''W), i.1948, coll. [anonymous]— 1 female, 1 male ( BMNH); Fazenda Contribução, Matão, (21°35'S, 48°22'W), 22.xi.1973, coll. [anonymous]— 3 females ( WDEC); Fazenda da Pedra, Rio Tamandúa, Ribeirão Preto, 5–8.xii.1953, coll. Travassos & Zago— 1 male ( CMNC); Pirassununga, (21°59'45''S, 47°25'37''W), x.1970, coll. A. Bello— 1 female ( CEMT); same locality, 14.x.1948, coll. Schubart— 1 female ( WDEC); same locality, 23.x.1970, coll. Bello— 1 male ( CEMT); Ribeirão Preto, (21°10'S, 47°49'W), ix.1954, coll. Duret— 1 female ( CMNC); same locality, xi.1954, coll. Barretto— 1 female ( MZSP); [NO DATA]: -., coll. [anonymous]— 1 male ( MNHN); PARAGUAY: AMAMBAY, Parque Nacional Cerro Corá, (22°37'41''S, 56°12'28''W), 12–16.x.1981, coll. J. Kochalka— 1 male ( FSCA); same locality, 1–2.vi.2006, coll. B. Garcete— 1 female ( WDEC); ASUNCIÓN, Trinidad, x.1948, coll. Martínez— 1 female ( CMNC); CONCEPCIÓN, Campo Zanja Morotí, (22°31'48''S, 57°13'48''W), xii.2005, coll. C. Aguilar— 1 female ( WDEC); same locality, 20.x.2006, coll. C. Aguilar— 1 male ( WDEC).

Natural history. A series of recently collected specimens using light trap set in cerrado, Eucalyptus grandis stand, mata ciliar, Brachiaria decumbens pasture and Brachiaria decumbens pasture—Atlantic forest edge. Two specimens found in Guzera bovine dropping and a single specimen collected in a pitfall trap baited with bovine dung.

Remarks. Females are extremely similar to males externally, only differing by the anterior pronotal carina being slightly more sinuous; the carina is less sinuous and subtuberculate medially in males. Parameres (Figs. 108– 109) simply rounded apically in dorsal view, rather dorsoventrally compressed and with few fine raspy granules apically. Variation, as usual, occurs in the extent of the metallic markings on the head and pronotum as well as the color of the markings. Although most individuals are green, some are blue to dark violet-blue and some others are light green with coppery tinge in some areas, especially on pronotum.

At higher magnification, the denticle of the pronotal margin show a porous surface medially and we suggest here that it is most likely the opening of an exocrine gland. The presence of this structure seemingly provide significant advantage as D. denticollis is the most frequently collected species in the genus, suggesting that the species is represented by a larger population and it is also widely distributed.

Nomenclature and taxonomy. Dendropaemon denticollis lividus Blut, 1939 = Dendropaemon denticollis Felsche, 1909 , new synonymy. The type of D. denticollis lividus Blut, 1939 , from Bolivia, differs from the nominal species by the more deeply punctate pygidium and the dominant bluish tinge instead of greenish. After closer examination, other specimens from Brazil also show various degree of impression in pygidial punctures. We consider this character as being within the specific variation of the species and could not find any pattern in the distribution that could support maintaining this taxon as a separate entity.

Figures

FIGURES 37 – 41. Dorsal habitus. 37. D. latistriatus, male holotype; 38. D. nitidicollis, male lectotype; 39. D. quadratus, male neotype; 40. D. similis, male holotype; 41. D. denticollis, female holotype.

FIGURES 126 – 133. Metatarsus, ventral view. 126. D. ganglbaueri, holotype; 127. D. vazdemelloi, holotype; 128. D. refulgens, holotype; 129. D. fractipes; 130. D. fasces; 131. D. nitidicollis; 132. D. quadratus; 133. D. denticollis.