Dasyus collaris LePeletier & Audinet-Serville, 1828

Fuhrmann, Juares, 2019, Taxonomy of Neotropical genera Compsodactylus and Dasyus and notes on claw movement in Macrodactylini (Coleoptera: Scarabaeidae: Melolonthinae), Zootaxa 4679 (1), pp. 139-163 : 147-148

Dasyus collaris LePeletier & Audinet-Serville, 1828

( Figs. 9 , 24–25 , 27–67 , 71–75 , 78, 80–81, 83–90 , 99 )

Dasyus collaris LePeletier & Audinet-Serville, 1828: 370 ; Laporte 1840: 143; Duponchel 1843: 616; Blanchard 1850: 96; Drapiez 1853: 383; Lacordaire 1855: 250; Harold 1869: 1139 (checklist); Dalla Torre 1913: 317 (checklist); Blackwelder 1944: 227 (checklist); Moser, 1918: 95; Evans 2003: 257 (checklist); Evans & Smith 2009: 226 (checklist); Katovich 2008: 38, figs. 77–81; Fuhrmann & Vaz-de-Mello 2017: 6, 9.

Dasyus fulvicollis Blanchard, 1850: 96 ; Burmeister 1855: 72; Duponchel 1843: 616; new synonymy.

Dasyus collaris fulvicollis; Harold 1869: 1139 (checklist).

Dasyus nigellus Blanchard, 1850: 96 ; Burmeister 1855: 72; Duponchel 1843: 616; Blackwelder 1944: 227 (checklist, synonym); synonymy confirmed.

Dasyus collaris nigellus ; Harold 1869: 1139 (checklist); Dalla Torre 1913: 317 (checklist); Moser 1918: 95; Evans 2003: 265 (checklist); Evans & Smith 2009: 226 (checklist).

Dasyus collaris luridipennis Moser, 1918: 95 ; Blackwelder 1944: 227 (checklist); Evans 2003: 257 (checklist, synonym); synonymy confirmed.

Dicrania hirsuta Frey, 1970a: 285 , fig. 7 (aedeagus); Frey, 1972: 69, fig. 1 (aedeagus); Evans 2003: 264 (checklist); Evans & Smith 2009: 227 (checklist); new synonymy.

Description. Body length: 7.2–11.6 mm, greatest width: 3.8–5.3 mm; entirely black ( Fig. 89 ), or black with pronotum red ( Fig. 87 ), or black with elytra yellow ( Fig. 84 ), or black with pronotum and elytra yellow ( Fig. 88 ). Head with epistomal suture distinct or not; eye small in dorsal view and interocular distance 6–7 times wider than eye width (measures taken at medial area of eyes). Clava longer than funicle (clava/funicle = 1.3–1.5), clava of males (length/width = 2.7–3) longer than clava of females (length/width = 2.5–2.6), funicle of females (length/width = 6.2–6.4) longer than funicle of males (length/width = 5–5.3). Prothoracic leg of males with tarsomere I bearing a small inner tooth ( Fig. 41d ) and I–IV with inner setal comb ( Fig. 41 ). Scutellum as wide as or wider than long (width/length = 1–1.6). Elytra of females (length/width = 2.6–2.7) relatively longer than elytra of males (length/ width = 2.2–2.3), disc densely setose. Metathoracic legs with tibia bearing dense thin setae, some setae slightly thicker than others. Abdomen with propygidium partially exposed in males by the resting elytra or almost hidden in female; pygidium of females with a shallow longitudinal sulcus; ventrite VI of males with a medial concavity surrounded anterolaterally by carina ( Fig. 25 ). Aedeagus, parameres as figured ( Figs. 65–67 ); endophallus ( Figs. 71–75 ) with distal piece bearing a dorsal lobe, asperites of distal raspula with some aciculate projection ( Fig. 75 details). Female terminalia with bar-like paraprocts ( Fig. 78 ) and genitalia with gland of spermatheca rounded ( Fig. 81 ).

Remarks. Dasyus collaris has a wide color variation, and four specific group names were proposed for some of these morphological variations ( Figs. 84, 87–89 ) (attributed name between brackets): black specimens with red pronotum ( D. collaris “typical form”), black specimens with yellow pronotum and elytra ( D. fulvipennis sensu Burmeister, 1855 ), black specimens with yellow elytra ( D. fulvipennis and D. collaris luridipennis ), and entirely black specimens ( D. nigellus ). Despite the color variation, no other morphological variation is found and all “color forms” are sympatric. The present study proposes D. fulvicollis and Dicrania hirsuta as new synonyms of Dasyus collaris and confirm the synonymy of D. nigellus and D. collaris luridipennis with D. collaris .

Distribution. Southeast Brazilian coast, in the extreme south of Bahia, in Espírito Santo and Rio de Janeiro ( Fig. 99 ). One specimen labeled Buenos Aires, but it is probably a mislabeled specimen (omitted in map: Fig. 99 ).

Type series. Dasyus collaris , D. fulvipennis , and D. nigellus: Three species were described based on an undetermined number of specimens and have their type series presumably deposited in MNHN, one of them is the series of the type species D. collaris . The names D. fulvipennis and D. nigellus were first used in Dejean’s second catalogue ( Dejean 1833: 164), but the names were only described and disponible by Blanchard (1850). The type series of these three species were not found in MNHN, Katovich (2008) studied a possible syntype of D. collaris in MNHN, a more careful study of the collection material is needed to locate these type series.

Dasyus collaris luridipennis: This subspecies was described based on an undetermined number of specimens. A male lectotype ( Figs. 84–85 ) at ZMHU is hereby designed; lectotype without external damage; labels ( Fig. 86 ): [Brasilia] [ Dasyus collaris | v. luridiennis | type Mos.] [LECTOTYPE ♂ | Dasyus collaris | luridipennis Moser, 1918 | des. J. Fuhrmann, 2014]. A female paralectotype at ZMHU with similar labels to the lectotype was also observed.

Dicrania hirsuta: Frey (1970a) described the species based on 11 specimens and noted that the holotype was housed at MZSP and the paratypes in his personal collection (currently on NHMB), but the holotype and two paratypes were found at IBSP, five paratypes at NHMB, and a paratype at ZMHU. Male holotype ( Fig. 89 ) with right mesotarsus lost; labels ( Fig. 90 ): [Rio de Janeiro | Corcovado | GB, x.1965 | Alvarenga col.] [Type] [ Dicrania | hirsuta | det. G. Frey, 1970 | n.sp.]. The label data “GB” is an acronym of Guanabara State, that existed from 1960 to 1975 and comprised only the city of Rio de Janeiro when it was the national capital.

Material examined. (62 specimens). BRAZIL: 2 ♂, 1 ♀ ( BMNH) , 1 ♂, 1 ♀ ( MNHN) , 1 ♂ ( D. collaris luridipennis lectotype) ( ZMHU) , 2 ♀ ( ZNSM) , 1 ♀ ( D. collaris luridipennis paralectotype) ( ZMHU) , 5 ♂, 2 ♀ ( ZMHU) ; BAHIA, Mucuri, P.C. Elias (collector), ii.1971 , 1 ♂ ( MZSP) ; ESPÍRITO SANTO, 1 ♀ ( ZMHU) , Santa Leopoldina, 1 ♀ ( ZMHU) ; RIO DE JANEIRO, Campos dos Goytacazes , “Morro Itaoca”, M.E. Resende (collector), in Vellozia sp., 2.x.2001 , 1 ♂, 2 ♀ ( CEMT) , Nova Friburgo, 1 ♂ ( ZNSM) , Brenske (collector), 1 ♀ ( ZMHU) , Sero- pédia, “Universidade Federal Rural do Rio de Janeiro” (“ Escola de Agronomia ”), Martínez (collector), xii.1952 , 1 ♂ ( CMNC) , Rio de Janeiro , 2 ♂, 1 ♀ ( BMNH) , “Corcovado”, Alvarenga (collector), x.1965, 1 ♂ ( Dicrania hirsuta holotype) ( IBSP) , 2 ♀ ( D. hirsuta paratypes) ( IBSP) , 2 ♂, 3 ♀ ( D. hirsuta paratypes) ( NHMB), # 1 m ( D. hirsuta paratype) ( ZMHU) , Alvarenga & Seabra (collectors), ix.1959, 1 ♀ ( DZUP) , x.1959, 1 ♀ ( DZUP) , ix.1961, 1 ♀ ( DZUP) , xii.1961, 1 ♂ ( DZUP) , ix.1969, 2 ♂, 1 ♀ ( DZUP) , Moure, Alvarenga & Seabra (Collectors), 15.ix.1961, 1 ♀ ( DZUP) , 18.ix.2961, 2 ♀ ( DZUP) , Maucer (collector), xi.1853, 1 ♀ ( MNHN) , G. Perina (collector), 1 ♀ ( NHMB) , “Parque Estadual da Pedra Branca – Pau da Fome” (“ Represa do Rio Grande ”), F.M. Oliverira (collector), 10.ix.1966 , 1 ♂ ( DZUP) , 15.x.1967, 1 ♀ ( DZUP) ; ARGENTINA (?): BUENOS AIRES, Buenos Aires , 1 ♀ ( ZMHU) ; NO DATA, 1 ♂ ( BMNH) , 2 ♂, 4 ♀ ( MNHN) , 1 ♀ ( MZSP) , 1 ♂, 1 ♀ ( ZMHU) .

Figures

FIGURES 9–20. Male appendages. 9, Dasyus collaris metacoxa; 10, Compsodactylus argentinus metacoxa; 11, C. argentinus apex of metatibia; 12, C. argentinus metafemur and tibia; 13, C. argentinus tibia, posterior view; 14, C. parvulus apex of metatibia; 15, C. parvulus metafemur and tibia; 16, C. parvulus tibia, posterior view; 17–18, C. argentinus aedeagus (lateral and parameres in dorsal view); 19–20, C. parvulus aedeagus (lateral and parameres in dorsal view). cxk, coxa-katepisternum articulation; cxp, coxopleural articulation; cxt, coxa-trochanter articulation; mbd, marginal bead; mip, metacoxal inner process (limit in dotted line). Scale = 1 mm.

FIGURES 24–26. Dasyus male. 24, Dasyus collaris habitus, lateral (without some appendages); 25, D. collaris abdomen, lateroventral; 26, D. variabilis abdomen, lateroventral. Scale = 1 mm.

FIGURES 27–29. Dasyus collaris. 27, epipharynx (a, phobae; b, minute setae; details with relative magnification of 3x); 28, hypopharynx; 29, labium (left palp and some setae omitted). Scale = 0.5 mm.

FIGURES 30–34. Dasyus collaris mandibles. 30–33, Left mandible (dorsal, ventral, outer, molar); 34, molar of right mandible. Scale = 0.2 mm.

FIGURES 35–37. Dasyus collaris left maxilla (dorsal, inner, ventral; seta details with relative magnification of 3x). Scale = 0.2 mm.

FIGURES 38–45. Dasyus collaris. 38, prothorax, anterior (right side with cervix and coxa omitted); 39, prothorax, posterior (right without intersegmental membrane and coxa omitted); 40, prothorax, ventral (right side with hypomeron bridge detached); 41, protibia-tarsus of males (with protracted pretarsus), details: a, tarsomere V with retracted pretarsus, b–c, tarsomere V, inner (with protracted/retracted pretarsus, respectively), d, tarsomere I detail; 42, 43, mesotibia-tarsus (female, male); 44, metatibiatarsus of male (with protracted pretarsus) and detail of tarsomere V with retracted pretarsus; 45, apex of metatibia. am1, angulate margin between hypomeron and posterior fold of pronotum; am2, angulate margin between dorsal and ventral areas of posterior fold of pronotum. Scale = 1 mm.

FIGURES 46–51. Dasyus collaris pterothorax. 46, anterior (right side with mesoalinotum omitted); 47, posterior; 48, metanotum, internoventral (right side with mesopostnotum complex); 49, internodorsal (right episternum-epimeron and associated region omitted); 50, lateral; 51, internolateral. Acronyms suffix 2 and 3 indicating mesothorax and metathorax elements, respectively (except to fragma: ph1–3); apn, apodeme of postnotum; awp, anterior notal wing process; ba2–3, basalare (* apodeme); dm3, dorsal arm of epimeron; em2–3, epimeron; es2–3, episternum; fu2–3, furca; lp2–3, lateral arm of postnotum; ls2–3, lateral arm of scutellum; msb, mesendosternite bridge; mwp, medial notal wing process; pe2–3, prealare; ph1–3, first, second and third phragma; pla, pleural arm; po2, postalare; pwp, posterior notal wing process; sc2–3, scutum; su3, subalare (* apodeme); st2–3, scutellum; vt2–3, meso- and metaventrite; wp2–3, pleural wing process. Scale = 1 mm.

FIGURES 52–56. Dasyus collaris pterothorax and abdomen. 52, pterothorax, dorsal (right side of mesoalinotum omitted); 53, mesoalinotum, internoventral; 54, episternum-epimeron complex and associated region, internal (apodeme limits of basalare and subalare in black line); 55–56, female abdomen (dorsal, ventral). Acronyms suffix 2 and 3 to thorax indicating mesothorax and metathorax elements, respectively (except to fragma: ph1–2); apn, postnotal apodeme; awp, anterior notal wing process; ba2–3, basalare; dm3, dorsal arm of metepimeron; em2–3, epimeron; es2–3, episternum; lp2–3, lateral arm of postnotum; ls2– 3, lateral arm of scutellum; mts, metathoracic spiracle; pcx, pleurocoxal articulation; pe2–3, prealare; ph1–2, first and second phragma; pla, pleural arm; po2, postalare; pyg, pygidium; sc2, mesoscutum; sn3–8, abdominal sternite 3–8; su3, subalare; st2, scutellum; tg1–8, abdominal tergite 1–8; va1–6, abdominal ventrite 1–6; wp2–3, pleural wing process. Scale = 1 mm.

FIGURES 57–63. Dasyus collaris, wing. 57, right posterior wing; 58–60, first axillary (dorsal, ventral, head anterior detail); 61–62, second axillary (dorsal, ventral); 63, third axillary, dorsal. Veins (anterior to posterior): Pc, postcosta; C, costa; Sc, ScA, subcosta, Sc anterior; R, RA, RP, radius, R anterior, R posterior; MP, medial posterior; Cu, CuA, cubital, Cu anterior; A, AA, AP, anal, A anterior, A posterior. Folds (dotted lines; anterior to posterior): dl1, dl2, longitudinal fold 1 and 2; mf, medial fold; tf, transversal fold; af, anal fold; jf, jugal fold. Scale = 0.5 mm.

FIGURES 64–70. Dasyus male terminalia. 64, Dasyus collaris genital ring; 65, D. collaris tegmen, lateral; 66, D. collaris parameres, distal; 67, D. collaris parameres, laterodistal; 68, D. variabilis tegmen, lateral; 69, D. variabilis parameres, distal; 70, D. variabilis parameres, laterodistal. phb, phallobasis (* apodeme); scm, second connective membrane (connecting genital ring to phallobasis); spg, spiculum gastrale (tergite IX element, * apodeme or cranial piece); sn9, abdominal sternite IX; tcm, third connective membrane (connecting abdominal segment VIII to genital ring). Scale = 1 mm.

FIGURES 71–77. Dasyus endophallus. 71, Dasyus collaris right side (distal piece introverted); 72, D. collaris dorsal detail of temones; 73, D. collaris ventral detail of V-sclerite; 74–75, D. collaris right and left side; 76–77, D. variabilis right (distal piece introverted) and left side. fcm, first connective membrane (connecting tegmen to endophallus); ldd, dorsal lobes of distal piece; lpd, dorsal lobe of proximal piece; lpm, ventromedial lobe of proximal piece; lpp, ventroproximal lobe of proximal piece; lpv, ventrodistal lobe of proximal piece; M(O), musculus phallobaso-endophalloapodemalis; M(S), musculus endophalloapodemalis; ttm, temones tendon. Asperites details with relative magnification of about 10x. Scale = 0.5 mm.

FIGURES 78–83. Dasyus female terminalia. 78, Dasyus collaris terminalia with detail of asperites of proctiger area; 79, D. variabilis terminalia; 80, D. collaris pygidium, internal (showing paraproct-pygidium articulation); 81, D. collaris genitalia (right side without external elements; left side with detail of gonocoxite setae); 82, D. variabilis spermatheca complex; 83, D. collaris longitudinal section, lateral (accessory gland in dotted line). acg, accessory gland; bcx, bursa copulatrix; gcx, gonocoxite; ovd, oviduct; ppt, paraproct; pyg, pygidium; ret, rectum; spt, spermatheca; stg, gland of spermatheca; tg8–9, abdominal tergite VIII and IX. Details with relative magnification of about 10x. Scale = 0.5 mm.

FIGURES 84–90. Dasyus collaris variation. 84–86, Dasyus collaris luridipennis male lectotype (dorsal, lateral, labels); 87, typical color (Rio de Janeiro, “Corcovado”); 88, female with same color attributed to D. fulvipennis sensu Burmeister, 1855; 89–90, Dicrania hirsuta male holotype (dorsal, labels). Scale = 1 mm.

FIGURES 98–99. South America distributional records. 98, Compsodactylus; 99, Dasyus. Scale = 500 km.