Dendropaemon (Sulcopaemon) quadratus ( Laporte, 1832 )

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) : 76-79

39. Dendropaemon (Sulcopaemon) quadratus ( Laporte, 1832)

( Figs. 39 , 104–105, 132, 153, 160)

Enicotarsus Quadratus Laporte 1832 , Ann. Soc. Ent. Fr. 1: 403 (original description) Enicotarsus quadratus: Castelnau 1840 , Hist. Nat . Ins. 2: 83 (diagnosis, distribution) Dendropemon quadratus: Harold 1869 , Cat. Col. IV: 1020 (mentioned as synonym) Dendropemon smaragdinus Waterhouse 1891 , Ann. Mag. Nat . Hist. 6 8: 56 (original description) new synonymy Dendropemon quadratus: Felsche 1909 , Deut. Ent. Zeit. 1909: 757 (comment)

Dendropemon quadratus: Gillet 1911 , Col. Cat. 38: 88 (mentioned as synonym)

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

Dendropaemon planus Olsoufieff 1924 , Insecta 13: 123 (original description) new synonymy Dendropaemon quadratus: Olsoufieff 1924 , Insecta 13: 124 (monograph)

Dendropaemon quadratus: Pessôa & Lane 1936 , Rev. Biol. Hygiene 7: 89 (identification key, comment) Dendropaemon planus: Blut 1939 , Arch. Naturg. (N.F.) 8: 293 (mentioned as synonym)

Dendropaemon smaragdinus: Blut 1939 , Arch. Naturg. (N.F.) 8: 293 (monograph)

Dendropaemon quadratus: Blut 1939 , Arch. Naturg. (N.F.) 8: 294 (monograph)

Dendropaemon smaragdinus Chevrolati Blut 1939 , Arch. Naturg. (N.F.) 8: 295 (original description) new synonymy Dendropemon planum: Blackwelder 1944 , U.S. Nat . Mus. Bull. 185: 211 (checklist)

Dendropemon quadratum: Blackwelder 1944 , U.S. Nat . Mus. Bull. 185: 211 (mentioned as synonym) Dendropemon smaragdinum: Blackwelder 1944 , U.S. Nat . Mus. Bull. 185: 211 (checklist) Dendropemon smaragdinum chevrolati: Blackwelder 1944 , U.S. Nat . Mus. Bull. 185: 211 (checklist) Dendropaemon quadratum: Edmonds 1972 , Univ. Kansas Sc. Bull. 49: 851 (comment taxonomy) Dendropaemon (D.) smaragdinus: Edmonds 1972 , Univ. Kansas Sc. Bull. 49: 851 (comment taxonomy) Dendropaemon planus: Arnaud 1982 , Rev. Fr. Ent. (N.S.) 4: 117 (lectotype designation)

Dendropaemon smaragdinum chevrolati : Vaz-de-Mello 2000, Hac. Proy. CYTED: 192 (faunistic) Dendropaemon smaragdinum smaragdinum : Vaz-de-Mello 2000, Hac. Proy. CYTED: 192 (faunistic) Dendropaemon planus: Vitolo 2000 , Rev. Acad. Colomb. Cienc. 24: 599 (identification key) Dendropaemon (D.) quadratus: Arnaud 2002 , Col. Monde 28: 15 (mention)

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

Dendropaemon (D.) smaragdinus : Vaz-de-Mello & Génier 2009, Col. Bull. 63: 365 (biology)

Type locality. Jatahy, Etat de Goyaz.

Diagnosis. The small size (less than 10.0 mm), with distinct metallic markings combined with the emarginate anterior edge of clypeus on external side of each clypeal tooth will place this species in the quadratus complex. It differs from all other species in the quadratus complex by the two-(instead of three-) segmented meso-and metatarsi.

Description. Male neotype ( Fig. 39 ). Body. Body small, length 8.5 mm, maximum width 4.5 mm; body subrectangular in dorsal view; dorsum largely flat. Color. Dorsal surface dark brown to black, glossy, with green metallic sheen; head black 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 black; pygidium with green metallic sheen; legs dark reddish brown to black. Head. Clypeus broadly arcuate, anterior portion slightly upturned; clypeal teeth 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, bordered posteriorly by a more or less regular row of punctures, clypeal surface with transverse blunt rugulae; clypeogenal suture well-defined, bluntly carinate internally; genal surface with fine blunt transverse rugulae laterally, minutely punctate internally, transversely tumescent; clypeofrontal carina low, more than 6 times wider than high, slightly arcuate in dorsal view, simply carinate, clypeofrontal carina apical edge slightly trilobate in frontal view; eyes small in dorsal view, interocular ratio 4.9. Pronotum. Pronotum transverse in dorsal view, pronotal width/length ratio 1.4; disc of pronotum minutely punctate throughout, with a fine longitudinal sulcus on posterior two-third; pronotal anterior margin wider and flat lateral to eyes; anterior portion with a medially sinuous sharp carina transversely tuberculate medially; anterior angles surface with fine tubercles laterally and few short longitudinal rugulae internally, slightly but distinctly sulcate along posterior edge of anterior margin; lateral fossae rounded and bordered anteriorly by a sharp carina; lateral portions unmodified; pronotal basal fossae very small and more or less rounded; posterior margin well-defined, lacking crenulation and setae. Elytra. Elytra approximately as long as wide in dorsal view, elytral combined width/length ratio 1.0; elytral base lacking distinct margin, simply convex; elytral striae 1–4 very wide basally and tapering toward apex, distinctly more impressed basally, elytral striae 5 similar to 4 on disc, strial punctures ill-defined throughout, stria 1 well-defined apically, going straight to elytral apical margin; interstriae slightly convex, minutely punctate throughout, surface glossy.

Thoracic sterna. Proepisternal carina reduced, present along coxal insertion only; metasternal median lobe flat, ventral ridge absent. Legs. Profemur posterior surface slightly but distinctly convex and glabrous internally, posterointernal margin rather thick, evenly developed, internal edge rather narrow, with a contiguous row of setae along anterointernal edge and few scattered long setae on anterior half, remaining surface smooth. Protibia with four teeth on lateral edge; internal basal angle lobate; anterior surface lacking aligned row of setae internally, with few aligned and isolated setiferous punctures only, surface coarsely microsculptured between punctures; posterior surface with irregular rugose punctures externally to median carina, surface finely and irregularly microsculptured between punctures, with a single interrupted setal row along lateral teeth. Mesofemur angularly produced 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, 2-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 median half. Metatibia robust, short, slightly widening toward apex in anterior view, anterior surface with distinct row of setae, surface glossy externally with distinct transverse microsculpture internally, metatibial posterior surface concave between internal and lateral edge, with transverse microsculpture. Metatarsus 2-segmented ( Fig. 132 ), first segment moderately elongate, approximately two times as long as wide at apex, with anterointernal carina well defined and almost reaching apical edge. Abdominal sternites. Sternites 3–6 longitudinally slightly convex; sternites 4–6 with 1–3 unaligned rows of setae laterally, glabrous medially; sternite 7 slightly longitudinally concave medially, shorter than segment 6 along midline; pygidium minutely punctate on disc. Male genitalia (Figs. 104–105). Parameres produced into a lobe laterally; surface smooth, glossy apically.

Measurements (9 males, 8 females). Length: male 7.5–9.5 (8.3±0.6), female 6.5–10.0 (8.6±1.2) mm.

Primary type data.

Enicotarsus quadratus Laporte ( Fig. 153 ). Male neotype (MNHN) present designation: [Trinidade/ (Goyaz)/ Ch.Pujol]; [WORLD/ SCARAB./ DATABASE/ WSD00016803]; [NEOTYPE / Enicotarsus / quadratus / Laporte,1832 / dés. Génier & Arnaud, 2014] red card; [ Dendropaemon / quadratus / Laporte/ dét. F. Génier, 2009] handwritten.

Dendrop[a]emon smaragdinus Waterhouse. Holotype male (BMNH): [Type] disc with red border; [67 45]; [TYPE] red card; [ Dendropemon / smaragdinus ,/ (Type) Waterh.] handwritten; [WORLD/ SCARAB./ DATABASE/ WSD00016804]; [HOLOTYPE / Dendrop[a]emon/ smaragdinus / Waterhouse, 1891] red card.

Dendropaemon planus Olsoufieff. Lectotype female (MNHN): [Jatahy/ (GOYAZ)] green card; [MUSÉUM PARIS/ 1936/ COLL. A. BOUCOMONT] green card; [Typus] red card; [Dendrop. planus n.sp. / det. G. OLSOUFIEFF] partly handwritten; [WORLD/ SCARAB./ DATABASE/ WSD00016805]; [ Dendropaemon / planus Ols. / LECTOTYPE / P. ARNAUD DET 1982] partly handwritten, red border.

Dendropaemon smaragdinus chevrolati Blut. Holotype female (MTD): [ Enicotarsus / quadratus / de Lap Bresil / type] handwritten; [ Brasilien] green card; [Coll. C. Felsche/ Kauf 20, 1918] green card; [♀]; [ Dendropaemon / s. Chevrolati ♀/ Brasilien.— Type.] handwritten, black border; [ viridis / Perty/ Paraguay] purple border; [WORLD/ SCARAB./ DATABASE/ WSD0016807]; [HOLOTYPE / Dendropaemon / smaragdinus chevrolati / Blut, 1939] red card; [ Dendropaemon ♀/ quadratus / ( Laporte, 1832)/ dét. Génier & Arnaud,2009].

Material examined. [NO DATA]: -., coll. [anonymous]— 1 male (holotype of D. smaragdinus Waterhouse ) ( BMNH); ARGENTINA: MISIONES, Dos de Mayo, elev. 500 m (27°1'39''S, 54°40'3''W), vii.1977, coll. R. Foerster— 1 female ( CPFA); Loreto, (27°18'22''S, 55°32'10''W), i.1956, coll. F.H. Walz— 1 male ( BDGC); same locality, [no date], coll. A. Ogloblin— 1 female ( CEMT); same locality, i.1955, coll. F.H. Walz— 1 female ( CMNC); same locality, xi.1958, coll. Martínez— 1 female, 2 males ( CMNC); same locality, i.1954, coll. F.H. Walz— 1 male ( CMNC); BRAZIL: [unspecified locality], [no date], coll. [anonymous]— 1 male ( CPFA); GOIÁS, Jataí, (17°53'S, 51°43'W), [no date], coll. [anonymous]— 1 female (lectotype of D. planus Olsoufieff ) ( MNHN); Rio Verde, (17°47'50''S, 50°54'0''W), [no date], coll. [anonymous]— 1 female, 1 male ( MTD); Trindade, (16°38'52''S, 49°29'53''W), [no date], coll. Ch. Pujol— 1 male (neotype) ( MNHN); same locality, [no date], coll. [anonymous]— 1 female ( ZMHB); same locality, [no date], coll. Ch. Pujol— 1 female (paralectotype of D. planus Olsoufieff ) ( MNHN); [unspecified locality], [no date], coll. [anonymous]— 1 female (holotype of D. smaragdinus chevrolati Blut ) ( MTD); MATO GROSSO DO SUL, UNESP Farm [=Fazenda Experimental da Universidade Estadual Paulista, câmpus de Ilha Solteira], Selvíria, (20°20'2.1''S, 51°24'26.3''W), 18.i.2008, coll. F. Oikawa— 1 female (MEFEIS); [NO DATA]: -., coll. [anonymous]— 1 female, 1 male ( MNHN, MNRJ); PARAGUAY: ITAPÚA, Puerto Cantera, (27°13'44''S, 55°36'9''W), xii.1956, coll. F.H. Schade— 1 female ( CMNC); same locality, xi.1956, coll. Walz— 1 male ( CMNC).

Natural history. A specimen was collected in a pitfall trap baited with Sus scrofa dung.

Remarks. Females differ in having the anterior pronotal carina straighter and set very close to the anterior margin, the carina is only finely tuberculate medially. The abdominal sternite 7 is also nearly flat medially.

Variation, as usual, occurs in the extent of the metallic marking on head and pronotum. The greenish tinge is rather uniform in the sample studied. The extent and density of the fine tubercles and rugulae on the pronotal anterior angles vary, in some specimens the sculpturing is finer and denser and set on a slightly larger area.

Nomenclature and taxonomy. The type of D. quadratus is presume destroyed (see comment under D. ater on Laporte’s collection). Historically, the status of this species has remain unsettled. In the first review of the group, Castelnau (1840) simply copied the description he made in 1832 omitting the Latin diagnosis. Harold (1869), in his catalogue, considered D. quadratus as a synonym of D. viridis . Felsche (1909) briefly commented Harold’s synonymy and suggest that D. quadratus cannot be synonymous with D. viridis as the meso- and metatarsi are three segmented and are only two segments in D. viridis . Felsche stated that the third segment is present but barely visible based on a specimen studied from Chevrolat’s collection and labeled as “ Enicotarsus quadratus de Cast.- Type ”. Olsoufieff (1924) discuss again the situation and concluded that D. quadratus is indeed a valid species, but do not attempt to settle the issue. Pessôa and Lane (1936) essentially repeat Olsoufieff’s observations and questionably leave D. quadratus as a synonym of D. viridis . Finally, Blut (1939) concluded that the status of D. quadratus remain uncertain and is the first author to use the measurements given by Laporte in the original description to suggest that D. quadratus cannot be a synonym of D. viridis which is distinctly larger. Blut also rightly point out that the specimen from Chevrolat’s collection possess only two tarsal segments and not three as stated by previous authors. Furthermore, Blut suggest that D. quadratus body size would be similar to those of D. refulgen s. However, in the end, Blut conclude that the name D. quadratus is to be forgotten as he considered the name to be in litteris.

The purpose of this work is to address all taxonomic problems pertaining to the genus and therefore we designate a neotype for Laporte’s species D. quadratus of which the original description meet requirements for being valid. In order to remain as conservative as possible we designate a male specimen from the Museum national d’Histoire naturelle that comply with the original description and has precise locality data.

This specimen matches approximately the size given by Laporte and the very rudimentary description. It also matches the historical specimen from Chevrolat’s collection. This specimen differs from the original description by the “[pronotum] très-fortement ponctué avec des espaces lisses”, which could correspond to certain Coprophanaeoides species. We however, interpret the heavy punctation as being the sculpturing of the lateral pronotal declivities.

Following the designation of a neotype for D. quadratus the following nomenclatural acts can now be implemented (the second epithet being valid):

1) D. smaragdinus Waterhouse, 1891 = D. quadratus Laporte, 1832 , new synonymy.

The holotype of D. smaragdinus is a very small individual of D. quadratus , aside being slightly teneral and having the median lobe of the metasternum slightly more convex no differences can be found.

2) D. planus Olsoufieff, 1924 = D. quadratus Laporte, 1832 , new synonymy.

The lectotype of D. planus has been studied and except for being a female and having the pronotal green metallic marking less extensive no other differences could be found.

3) D. smaragdinus chevrolati Blut, 1939 = D. quadratus Laporte, 1832 , new synonymy.

The holotype female of D. smaragdinus chevrolati has been compared to the holotype of D. quadratus and only differs in the extent of the pronotal metallic marking which are absent on most of the disc. The pronotal and cephalic microsculpture are slightly more pronounced. The fine tubercles and rugulae of the pronotal lateral declivities are denser and cover slightly more surface. The pronotal punctures are fine instead of minute laterally posterior to lateral fossae. Some specimens in the material studied show intermediate forms and this variation is here considered intraspecific.

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.

FIGURES 134 – 154. Primary type labels. 134. D. hirticollis; 135. D. pauliani; 136. D. renatii; 137. D. ater; 138. D. piceus; 139: D. telephus; 140. D. viridis; 141. D. viridipennis. 142. D. convexus; 143. D. fredericki; 144. D. angustipennis; 145. D. bahianus; 146. D. amyntas (= D. attalus); 147. D. ganglbaueri; 148. D. refulgens; 149. D. fractipes; 150. D. fasces; 151. D. haroldi; 152. D. nitidicollis; 153. D. quadratus; 154. D. similis; 155. D. denticollis.