Uroxys epipleuralis ( Boucomont, 1928 )

Nazaré-Silva, Everton E., Moura, Ana B. G. & Silva, Fernando A. B., 2023, A new species of Uroxys Westwood, 1842 (Coleoptera: Scarabaeidae: Scarabaeinae) from central Brazil, and taxonomical remarks on Uroxys epipleuralis (Boucomont, 1928), Zootaxa 5374 (1), pp. 106-118 : 112-115

Uroxys epipleuralis ( Boucomont, 1928)

urn:lsid:zoobank.org:act:2C89217E-6E99-4FAB-9F8C-7C30B41656C6

( Figs. 1A–B , 2A–B, 2E–F , 3A–B, 3E–F, 3H , 4A–C , 5 )

Choeridium epipleurale Boucomont, 1928: 193–194 (original description); Balthasar 1939: 66 (identification key, taxonomic remarks); Blackwelder 1944: 204 (checklist and distribution).

Uroxys epipleurale : Martínez & Martínez 1987: 28 (catalog, distribution, and taxonomic remarks); Vaz-de-Mello 2000: 195 (checklist).

Uroxys epipleuralis : Krajčík 2006: 166 (checklist); Krajčík 2012: 262 (checklist); Tarasov & Génier 2015: 21–24, 27, 56, 65 (used as a terminal in a phylogenetic analysis, figs. 4–7, fig. 9, fig. 31b, fig. 36f); Tarasov & Dimitrov 2016: 7, 12 (fig. 3, fig. 4c, used as a terminal in phylogenetic analysis); Vaz-de-Mello et al. 2017: 4 (checklist).

Uroxys thoracalis Balthasar, 1940: 3 (original description); Krajčík 2006: 166 (checklist); Bezděk & Hájek 2011: 360 (catalog); Krajčík 2012: 262 (checklist). New subjective synonymy

Uroxys dureti Martínez, 1947: 48–51 (original description, Fig. 6); Martínez 1959: 66 (catalog and distribution); Martínez & Martínez 1987: 28 (catalog, distribution, and synonymy with U. epipleuralis ); Krajčík 2006: 166 (checklist, cited as valid species); Krajčík 2012: 262 (checklist, cited as valid species).

Diagnosis. Among the species of the genus, U. epipleuralis is similar to U. punctatus in having a clypeus with four strong acute teeth, elytra lacking humeral carina, elytra with strong longitudinal external carina near the seventh stria, and body strongly convex and shorter than 1.3 times its maximum width. However, specimens of U. epipleuralis can be distinguished by the surface of the head lacking ocellate punctures ( Figs. 2A–B ) (head with ocellate punctures over the frontoclypeal surface in U. punctatus , Figs. 2C–D ). In males, the surface of the pronotum bears ocellate punctures irregularly distributed ( Fig. 2E ), or dense ocellate punctures only laterally ( Fig. 2F ) (in U. punctatus , pronotum with dense ocellate punctures evenly distributed across the surface, Fig. 2G ). Additionally, differences were observed in the male genitalia, particularly in the medial endophallite. The cable of inferior portion of the medial endophallite of U. epipleuralis is thicker, and about 1.2 times shorter than that of U. punctatus ( Figs. 3F–G ) (inferior portion of the medial endophallite is slender and longer in U. punctatus , Fig. 3G ).

Type material. Choeridium epipleurale Boucomont : lectotype ♁ (here designated; MNHN): “REPUBLIQUE ARGNT | Territ des Missiones” // “ COLLON LE MOULT| Naturaliste, Paris” // “COLLECTION | WAGNER” // “MUSÉUM PARIS | 1938 | COLL A. BOUCOMONT” // “Boucomont det.192 8 | Choeridium | epipleurale Bouc. ” [Boucomont’s handwriting] // “ LECTOTYPE ” // “ Ch. Epipleurale | Bouc. ” [unknown handwriting] // “ LECTOTYPE ♁ | Choeridium | epipleurale | Boucomont | des. F.Z. Vaz-de-Mello, 20 14 ” [Vaz-de-Mello’s handwriting]. Paralectotypes ♁ ( MNHN): “ San Ignacio | Missions ” [unknown handwriting] // “COLLECTION | WAGNER” // “MUSÉUM PARIS | 1938 | COLL. A BOUCOMONT” // “ PARALECTOTYPE | Choeridium ♁ | epipleurale | Boucomont |des. F.Z.Vaz-de-Mello, 20 14 ” [Vaz-de-Mello’s handwriting]. ♀ ( MNHN): “REPUBLIQUE ARGNT ” // “COLLECTION | WAGNER” // “Typus” // “MUSÉUM PARIS | 1938 | COLL A. BOUCOMONT” // “ PARALECTOTYPE / Choeridium ♀ | epipleurale | Boucomont | des. F.Z. Vaz-de-Mello, 20 14 ” [Vaz-de-Mello’s handwriting]. ♀ ( MNHN): “ ARGENTINE | S. Ignacio” // “P. Chabanaud | 1922” // “MUSÉUM PARIS | 1938 | COLL. A BOUCOMONT” // “ PARALECTOTYPE | Choeridium ♀ | epipleurale | Boucomont | des. F.Z. Vaz-de-Mello, 20 14 ” [Vaz-de-Mello’s handwriting].

Uroxys thoracalis Balthasar : holotype, unsexed ( NMPC): “Typus” // “ Holotype ” [Vaz-de-Mello’s handwriting] // “Mus. Nat. Pragae | 65715 | Inv.” [unknown handwriting] // “ thoracalis | n sp. ” [Balthasar’s handwriting] // “ U. thoracalis | n. sp | DR. V. BALTHASAR. DET” [Balthasar’s handwriting] // “H. Jacob leg | Eing. Nr. 55.1933.” // “ Paraguay | Prov. Alto Parana | Hohenau”.

Uroxys dureti Martínez : holotype ♀ ( MACN): “ FICHADO ” [unknown handwriting] // “MACN-Em | 1045” // “ Uroxys | dureti ♀ | sp. n. | A. Martínez-Det 19 46 ” [Martínez’s handwriting] // “ HOLOTYPUS ” // “ El Naranjo / R. de la P. | SALTA / ARGENTINA I 44 | Duret-- Martínez ” .

Non-type material examined. 12♁ 111♀. BRAZIL: MATO GROSSO: Araputanga, Bandeirantes farm [semideciduous forest], [15°22’14’’S; 58°26’2’’W], 338 m, 20–22.I.2013, pitfall with human-pig dung, R.J. Silva – 1♀ ( CEMT); Indiavaí, Alto Jaurú farm [semideciduous forest], [15°26’19’’S, 58°34’51’’W], 308 m, 12.I.2013, R.J. Silva – 1♀ ( CEMT); Tangará da Serra, Fontosa farm [semideciduous forest], [14°35’52’’S, 57°50’13’’W], 325 m, 6.II.2012, pitfall with cow dung, R.J. Silva – 1♀ ( CEMT); Tangará da Serra, Fontosa farm [semideciduous forest], [14°35’53’’S, 57°50’2’’W], 312 m, 31.I.2012, pitfall with human-pig dung, R.J. Silva – 1 ♁, 2♀ ( CEMT); Tangará da Serra, Fontosa farm [semideciduous forest], [14°35’54’’S, 57°50’0’’W], 313 m, 6.II.2012, pitfall with cow dung, R.J. Silva – 1♀ ( CEMT); Tangará da Serra, Fontosa farm [semideciduous forest], [14°35’56’’S, 57°50’42’’W], 312 m, 1.II.2012, pitfall with cow dung, R.J. Silva – 2 ♁, 4♀ ( CEMT); Tangará da Serra, Fontosa farm [semideciduous forest], [14°35’58’’S, 57°50’42’’W], 312 m, 1.II.2012, pitfall with cow dung, R.J. Silva – 4♀ ( CEMT); Tangará da Serra, Fontosa farm [semideciduous forest], [14°36’1’’S, 57°50’17’’W], 310 m, 31.I.2012, pitfall with human dung, R.J. Silva – 1 ♁, 1♀ ( CEMT); Tangará da Serra, Netolândia farm [semideciduous forest], [14°41’5’’S, 57°54’8’’W], 263 m, 25–27.III.2012, pitfall with human-pig dung, R.J. Silva – 1♀ ( CEMT); Tangará da Serra, Paraíso farm [semideciduous forest], [14°41’37’’S, 57°24’31’’W], 492 m, 15–17.I.2011, pitfall with human-pig dung, R.J. Silva – 1♀ ( CEMT); Tangará da Serra, Paraíso farm [semideciduous forest], [14°41’40’’S, 57°24’41’’W], 498 m, 13–15.I.2011, pitfall with human-pig dung, R.J. Silva – 8♀ ( CEMT); Tangará da Serra, Paraíso farm [semideciduous forest], [14°41’42’’S, 57°24’38’’W], 498 m, 13–15.I.2011, pitfall with human-pig dung, R.J. Silva – 26♀ 3♁ ( CEMT); Tangará da Serra, Paraíso farm [semideciduous forest], [14°41’44’’S, 57°24’40’’W], 500 m, 13–15.I.2011, pitfall with human-pig dung, R.J. Silva – 1 ♁, 21♀ ( CEMT); Tangará da Serra, Paraíso farm [semideciduous forest], [14°41’45’’S, 57°24’41’’W], 503 m, 13–15.I.2011, pitfall with human-pig dung, R.J. Silva – 2♀ ( CEMT); Tangará da Serra, Paraíso farm [semideciduous forest], [14°41’46’’S, 57°24’40’’W], 503 m, 13– 15.I.2011, pitfall with human-pig dung, R.J. Silva – 2 ♁, 14♀ ( CEMT). MATO GROSSO DO SUL: Aquidauana , [no coordinates], 15.IV.2000, S.R. Rodrigues – 2♀ ( CEMT); Bataguassu, Caraguata River , [no coordinates], 23.III.1953, Martínez ‒ 2♀ ( CEMT); Bataguassu, Caraguata River , [no coordinates], 25.III.1959, F. Plaumann ‒ 1♀ ( CEMT); Bonito, Bonito farm, [21°6’S, 56°38’W], XI.2009, human dung, F.O. Roque – 6♀ ( CEMT); Bonito, Pitangueiras , [20°52’14’’S, 56°35’19’’W], XI.2009, human dung, F.O. Roque – 2♀ ( CEMT); Bonito, Remanso farm, [20°47’S, 56°43’W], XI.2009, human dung, F.O. Roque – 2♀ ( CEMT); Selvíria, Universidade do Estado de São Paulo farm, [no coordinates], 24.IX.1992], C.A. Flechtmann – 2♀ ( CEMT); Três lagoas, Barra do Moeda farm, [no coordinates], 10.XI.2010, P.F. Rosas – 2♀ ( CEMT). GOIÁS: Mineiros, Parque Nacional das Emas , [17°54’15’’S, 52°59’21’’W], 826 m, 15.III.2011, human dung, M.F. Souza – 1♀ ( CEMT). DISTRITO FEDERAL: Planaltina, Estação Ecológica Águas Emendadas , [15°32’31’’S, 47°36’49’’W], 16–28.II.2010, pitfall, M.R. Frizzas – 1♀ ( CEMT). SÃO PAULO: Riolândia, Vovó Júlia farm, [19°59’46’’S, 49°38’23’’W], 31.XII.2014, pitfall, L.G. Nunes – 2♀ ( CEMT). ARGENTINA: MISIONES, Parana River , [no coordinates] – 1 ♁, 3♀ ( CEMT – ex. Wagner and Le Moult collections) .

Distribution. Known from Brazil (Mato Grosso, Mato Grosso do Sul, Goiás, Distrito Federal, and S„o Paulo) and Argentina (Misiones) ( Fig. 5 ). Areas of endemism [South America]: Brazilian subregion: South Brazilian dominion: Rondônia province. Chacoan subregion: Chacoan dominion: Cerrado and Chacoan provinces. Parana dominion: Parana forest province.

Wide geographical disjunction. One of the remarkable aspects of the distribution of this species is the disjunction between a record in Salta and other in San Ignacio (both in Argentina), separated by the xerophytic forests of the Dry Chaco. According to Morrone (2014), Salta is in the Chacoan province , while the record in San Ignacio is in a transitional area between the Atlantic Forest and the Dry Chaco ( Fig. 5 ). The known records of Uroxys epipleuralis with geographic coordinates indicate that this species occurs in forested locations with semideciduous vegetation. Semideciduous seasonal forests are formations less humid than those where dense rainforests develop. In general, they occupy transitional environments between the coastal humid zone and the semi-arid and arid zones. Therefore, based on its current records in less humid forest areas, it is possible that this species is also found in xerophytic forests of the Dry Chaco. The gap visible in the map is due to the lack of collections. We hope that new specimens will come to light and get recognized by other researchers.

Notes about the lectotype of Choeridium epipleurale Boucomont, 1928 . The original description ( Boucomont 1928) presents no designation of a holotype or explicit mention as to whether it was based on a single specimen. According to Recommendation 73F ( International Commission on Zoological Nomenclature 1999), we assume that the type series was based on more than one individual. Fernando Vaz-de-Mello photographed four syntypes of Choeridium epipleurale during his visit to the MNHN between 2013 and 2014 [see Vaz-de-Mello & Cupello (2018) for more details about his visits to European museums]. These specimens were recognized as part of the type series since the labels match the collecting data provided by Boucomont in his original description, namely “République Argentine, Missiones: S. Ignacio (Chabanaud); Rio Parana.” Additionally, all specimens examined are from Boucomont’s collection as stated on their labels, and the lectotype designated here has the species identification label dated 1928, the year of the publication of the original description, and is in Boucomont’s handwriting. Finally, the specimens match Boucomont’s description.

During his visit to the Institut royal des Sciences naturelles de Belgique (ISNB), Brussels, Belgium, Fernando Vaz-de-Mello found four specimens from Rio Parana (one specimem with labels from Wagner and Le Moult collections; and three with only label from Le Moult collection). These specimens were donated to CEMT and are housed now in this collection. Although these specimens are from the Parana River and from collections in which Boucomont received specimens when describing C. epipleurale (see label data of lectotype and paralectotypes), we are not sure that they are in fact syntypes due to the following reasons: 1 - According to Vaz-de-Mello (personal communication), Le Moult and Wagner were insect merchants and they sent specimens to different collections. Furthermore, part of the specimens from their collections were incorporated into other collections after their deaths; 2 - All specimens found at the MNHN and which were recognized by us as syntypes of C. epipleurale have labels from Boucomont’s collection, which does not occur in these specimens housed at CEMT (via ISNB collection); 3 - None of these four specimens bears any label with Boucomont’s handwriting. We believe that part of the material collected in the Parana River and from the Le Moult and Wagner collections was made available to Boucomount, being incorporated into his collection and used by him when describing C. epipleurale . Another part of the material from the same collection events may have been destined for other collections. This is our hypothesis based on the evidence we have available. Therefore, we have chosen at this time to list these specimens in the non-type material section. However, if other evidence emerges over time, the status of these specimens may change.

Remarks. We examined the type specimens of C. epipleurale , U. thoracalis , and U. dureti from photographs and compared them with the non-type specimens examined here. The lectotype of C. epipleurale and the holotypes of U. thoracalis and U. dureti have the head surface lacking ocellate punctures, and the pronotal surface with ocellate punctures irregularly distributed ( Figs. 4A–C ). Despite some subtle differences in the position and distribution of these punctures between the type specimens, these morphological differences co-occur in individuals from the same locality across the distribution range of the examined material. Additionally, the analysis of the male genitalia confirms the hypothesis that this external variation represents intrapopulational polymorphism since we found no significant differences in the shape of the tegmen or the endophallites. In the original descriptions of U. thoracalis and U. dureti, Balthasar (1940) and Martínez (1947) did not mention the morphological similarity these species names share with U. epipleuralis . This is probably related to the period in which the species were described since U. epipleuralis was still placed in the genus Choeridium . Balthasar (1940) mentioned that U. thoracalis would be similar to U. sulcicollis Harold, 1880 U. micros Bates, 1887 and U. simplex Waterhouse, 1891 but the author only mentioned the differences between these species. He stated that, as opposed to U. thoracalis , the other species have a smooth pronotum and a clypeus not quadridentate. Martínez (1947) mentioned that U. dureti is unmistakable compared to other Uroxys since it does not have cephalic process, the clypeus has four teeth, and its general body shape is similar to Trichillum Harold, 1868 specimens. Furthermore, we also note that when establishing the synonymy of U. dureti with U. epipleuralis, Martínez & Martínez (1987) did not expose the morphological hypothesis on which the synonymy was based. Therefore, we establish our hypothesis, maintaining the previous synonymy proposed by Martínez & Martínez (1987) and establishing here U. thoracalis Balthasar, 1940 as a new junior subjective synonym of U. epipleuralis ( Boucomont, 1928) .

Morphological variation. Among the U. epipleuralis specimens analyzed here, polymorphism in the distance between the medial teeth of the clypeus and in the intensity of the pronotal punctation have been observed in populations from Brazil (Goiás: Mineiros; Mato Grosso do Sul: Bonito) and Argentina (Misiones). These populations show specimens with two patterns for the median clypeal teeth, specifically: teeth separated by a V-shaped or a Ushaped space ( Figs. 2B, 2A , respectively).Additionally, some individuals present the lateral surface of the pronotum entirely covered with dense ocellate punctures ( Fig. 2F ), while others have ocellate punctures irregularly distributed in the same region ( Fig. 2E ). The other populations present the median clypeal teeth separated by a V-shaped space and lateral surface of the pronotum entirely covered with dense ocellate punctures. No strict correlation was found between patterns of clypeal teeth and pronotal punctation.

Figures

FIGURE 1. Dorsal habitus of Uroxys species. A–B, Uroxys epipleuralis (Boucomont, 1928), males; C–D, U. punctatus new species, holotype and paratype, respectively (C, male; D, female).

FIGURE 2. Head and pronotum of Uroxys species. A–B, Uroxys epipleuralis (Boucomont, 1928), head, males; C–D, U. punctatus new species head (C male; D female); E–F, U. epipleuralis (Boucomont, 1928), pronotum, males; G–H, U. punctatus new species pronotum (G, male; H, female).

FIGURE 3. The male genitalia of two species of Uroxys epipleuralis (Boucomont, 1928) and U. punctatus new species. A–B, Uroxys epipleuralis tegmen (A, lateral view; B, dorsal view); C–D, U. punctatus tegmen (C, lateral view; D, dorsal view); E, U. epipleuralis superior and inferior portions of medial endophallite, ventral view; F, U. epipleuralis inferior portion of medial endophallite; G, U. punctatus inferior portion of medial endophallite; H, U. epipleuralis superior right peripheral (SRP) endophallites, complex of axial (A), subaxial (SA), and frontolateral peripheral (FLP) endophallites.

FIGURE 4. Type specimens studied for this work. A, Choeridium epipleurale Boucomont, 1928 (currently, Uroxys epipleuralis), lectotype; B, U. thoracalis Balthasar, 1940, holotype; C, U. dureti Martínez, 1947, holotype. A and B courtesy of Fernando Z. Vaz-de-Mello; C courtesy of Mario Cupello.

FIGURE 5. Known distribution of Uroxys epipleuralis (Boucomont, 1928) and U. punctatus new species.