Sylvicanthon candezei (Harold, 1869)

Cupello, Mario & Vaz-De, Fernando Z., 2018, A monographic revision of the Neotropical dung beetle genus Sylvicanthon Halffter & Martínez, 1977 (Coleoptera: Scarabaeidae: Scarabaeinae: Deltochilini), including a reappraisal of the taxonomic history of ‘ Canthon sensu lato’, European Journal of Taxonomy 467, pp. 1-205 : 73-80

Sylvicanthon candezei (Harold, 1869)

Figs 6B , 10C , 11J , 14 D–E, 15C, 17C, 20, 24, 25–26A–B

Canthon candezei Harold, 1869a: 96 .

Canthon candezei – Harold 1869b: 990. — Gillet 1911: 28. — Schmidt 1920: 133; 1922: 64, 73. — Balthasar 1939: 188. — Blackwelder 1944: 198. — Halffter & Martínez 1977: 62–63. — Krajcik 2012: 63.

Glaphyrocanthon (Glaphyrocanthon) candezei – Pereira & Martínez 1956: 126, 129. — Martínez et al. 1964: 5, 8–9, 14, 20. — Vulcano & Pereira 1967: 561.

Glaphyrocanthon (Glaphyrocanthon) candèzei – Vulcano & Pereira 1964: 661.

Sylvicanthon candezei – Halffter & Martínez 1977: 63. — Feer 2000: 32 (error: refers to S. seag sp. nov.); 2008: 62 (error: refers to S. seag sp nov.). — Vaz-de-Mello 2000: 195. — Escobar 2000a: 210 (error: probably refers to S. genieri sp. nov. or S. mayri sp. nov.). — Medina et al. 2001: 137 (idem). — Feer & Pincebourde 2005: 30 (error: refers to S. seag sp nov.). — Scheffler 2005: 19. — Medina & Pulido 2009: 59 (error: probably refers to S. genieri sp. nov. or S. mayri sp. nov.). — Carvajal et al. 2011: 117, 316 (error: refers to S. genieri sp. nov.).

Etymology

Eponym after the Belgian entomologist Ernest Candèze (1827–1898). One of the specimens studied by Harold (1869a) belonged to the Candèze collection – nowadays largely housed at the ISNB ( Horn & Kahle 1935) – and this may have been the reason for the homage.

Material examined

Lectotype (here designated, Fig. 26 )

BRAZIL: ♂, Pará, the Tapajós River (some point at the 170 km above its mouth in the Amazon River or across the banks of the Cupari River ( Fig. 26B ); see more details below), type locality cited by Harold (1869): “ Tapajos ”. Labels: (“ Candezei / Harold. Hefte V.”, “Muséum Paris / 1952 / Coll. R. Oberthur”, “ Tapajos ”, “Ex-Mus ӕo / H. W. Bates / 1892”, “ LECTOTYPE ♂ / Canthon / candezei / Harold / des. F. Z. Vaz-de-Mello, 20 14 ”) ( MNHN).

Paralectotypes (4 ♂♂, 4 ♀♀)

BRAZIL: 1 ♂, (“ Tapajós ”, “Ex-Mus ӕo / H. W. Bates / 1892”, “Muséum Paris / 1952 / coll. R. Oberthür”, “ PARALECTOTYPE / ♂ / Canthon / candezei / Harold / des. F. Z. Vaz-de-Mello, 20 14 ”) ( MNHN); 1 ♂, (“ Tapajos ”, “ Candezei / Harold ”, “Ex-Mus ӕo / E. Harold”, “S ylvicanthon / candezei / Harold / G. H. y A. M. det . 76 ”, “PARALETOTYPE / Canthon candezei / Harold, 1869 ♂ / des. Cupello & Vaz-de-Mello, 2015”) ( MNHN); 1 ♂ (dissected) (“ Tapajos ”, “ Candezei / Harold ”, “Ex-Mus ӕo / E. Harold”, “S ylvicanthon / candezei / Harold / G. H. y A. M. det . 76 ”, “ PARALECTOTYPE / ♀ / Canthon / candezei / Harold / des. F. Z. Vaz-de-Mello, 2015”) ( MNHN); 1 ♀ (“ Tapajos ”, “Ex-Mus ӕo / H. W. Bates / 1892”, “ candezei / Harold / [ilegível] Har. [illegible]”, “Muséum Paris / 1952 / coll. R. Oberthür” “ PARALECTOTYPE / ♀ / Canthon / candezei / Harold / des. F. Z. Vaz-de-Mello, 2014”) ( MNHN) ( Fig. 26 ); 1 ♀, (“ Tapajos ”, “Ex-Mus ӕo / H. W. Bates / 1892”, “ PARALECTOTYPE / ♀ / Canthon / candezei / Harold / des. F. Z. Vaz-de-Mello, 20 14 ”) ( MNHN); 1 ♀ (“ Tapajos ”, “Ex-Mus ӕo / H.W. Bates / 1892”, “Muséum Paris / 1952 / coll. R. Oberthür” “ PARALECTOTYPE / ♀ / Canthon / candezei / Harold / des. F.Z. Vaz-de-Mello, 2014”) ( MNHN); 1 ♀, (“ Tapajos ” “ Candezei / Harold ”, “Type”, “Collection / E. CANDÈZE”, “ PARALECTOTYPE / ♀ / Canthon / candezei / Harold / des. F.Z. Vazde-Mello, 2014”) ( ISNB); 1 ♂ (“ Tapajos ”, “ PARALECTOTYPE / ♂ / Canthon / candezei / Harold / des. F. Z. Vaz-de-Mello, 2014”) ( CEMT, ex MNHN).

Additional material (103 ♂♂, 54 ♀♀)

BRAZIL: Mato Grosso: 1 ♀, Araputanga, Fazenda Araputanga, Bacia Jauru, 15º21′48″ S, 58º26′03″ W, Dec. 2002, M. Santos Filho leg. ( CEMT); 1 ♂, Araputanga, Fazenda Bandeirantes, 15º22′18″ S, 58º26′23″ W, 297 m, 21–23 Jan. 2013, pitfall with human faeces, R. J. Silva leg. ( CEMT); 1 ♂, Araputanga, Fazenda Bandeirantes, 15º22′14″ S, 58º26′02″ W, 338 m, 20–22 Jan. 2013, pitfall with human and swine dung, R.J. Silva leg. ( CEMT); 1 ♂, Curvelândia, Fazenda Calça Vermelha, Bacia Cabaçal, 15º33′02″ S, 58º00′57″ W, Feb. 2004, M. Santos-Filho leg. ( CEMT); 1 ♀, Diamantino, Fazenda São João, 14º14′10″ S, 56º08′11″ W, 11 Jan. 2001, flight interception trap, Génier and Vazde-Mello leg. ( CMNC); 1 ♂, Diamantino, Fazenda São João, 14º14′10″ S, 56º08′11″ W, 400 m, 13 Jan. 2001, flight interception trap, Génier and Vaz-de-Mello leg. ( CMNC); 1 ♀, Diamantino, Fazenda São João, 14º23′49″ S, 56º09′30″ W, 480 m, 13 Jan. 2001, trap with faeces, Génier and Vaz-de-Mello leg. ( CMNC); 1 ♂, 1 ♀, Indiavaí, Fazenda Nova Canaã, Bacia Jauru, 15º16′31″ S, 58º40′06″ W, Jul. 2004, M. Santos-Filho leg. ( CEMT); 1 ♂, 1 ♀, Nobres, Jan. 2001, A. Bello leg. ( AMBC); 1 ♀, Nova Mutum, 13º48′07″ S, 56º05′22″ W, 23 Jan. 2011, human faeces, M.F. Souza leg. ( CEMT); 2 ♂♂, Nova Mutum, 13º48′07″ S, 56º05′22″ W, 25 Jan. 2011, human faeces, M.F. Souza leg. ( CEMT); 8 ♂♂, 4 ♀♀, Nova Mutum, 13º48′07″ S, 56º05′22″ W, 18 Apr. 2011, human faeces, M.F. Souza leg. ( CEMT); 1 ♂, Nova Mutum, 13º48′07″ S, 56º05′22″ W, 2 May 2011, human faeces, M.F. Souza leg. ( CEMT); 1 ♀, Nova Mutum, Trivelato, 15 Dec. 1995, W.O. Silva-Filho leg. ( CEMT); 1 ♂, 2 ♀♀, Querência, Fazenda São Luiz, 12º39.68′ S, 52º22.14′ W, 8 Jul. 2008, flight interception trap, R. Andrade leg. ( CEMT); 2 ♀♀, Querência, Fazenda São Luiz, 12º39.81′ S, 52º22.74′ W, 17 Jul. 2008, pitfall, R. Andrade leg. ( CEMT); 2 ♀♀, Querência, Fazenda São Luiz, 12º39.94′ S, 52º21.85′ W, 14– Jul. 2008, pitfall, R. Andrade leg. ( CEMT); 1 ♂♂, 5 ♀♀, Querência, Fazenda São Luiz, 12º39.64′ S, 52º22.74′ W, 17 Jul. 2008, pitfall, R. Andrade leg. ( CEMT); 2 ♂♂, Querência, Fazenda São Luiz, 12º40.48′S, 52º21.86′W, 13 Feb. 2009, pitfall, R. Andrade leg. ( CEMT); 1 ♂, 3 ♀♀, Tangará da Serra, 16 Jul. 2008, pitfall with human faeces, R.J. Silva leg. ( CEMT); 2 ♂♂, Tangará da Serra, Fazenda “Ap. Da Serra”, 14º19′15″ S, 57º43′51″ W, 640 m, 20–22 Apr. 2017, pitfall with human faeces, R.J. Silva leg. ( CEMT); 8 ♂♂, 1 ♀, Tangará da Serra, Fazenda Bahia, 14º37′19″ S, 57º25′07″ W, 419 m, 12–14 Jan. 2011, pitfall with human and swine dung, R.J. Silva leg. ( CEMT); 1 ♂, 1 ♀, Tangará da Serra, Fazenda Bahia, 14º37′13″ S, 57º24′50″ W, 428 m, 26–28 Jan. 2012, pitfall with cow dung, R.J. Silva leg. ( CEMT); 1 ♂, Tangará da Serra, Fazenda Curitiba, 14º21′47″ S, 57º28′17″ W, 345 m, 2–9 Apr. 2012, flight interception trap, R.J. Silva leg. ( CEMT); 11 ♂♂ (1 dissected), 1 ♀, Tangará da Serra, Fazenda Filé do Boi, 14º38′07″ S, 57º24′41″ W, 439 m, 25–27 Jan. 2011, pitfall with human and swine dung, R.J. Silva leg. ( CEMT); 1 ♂, Tangará da Serra, Fazenda Fontosa, 14º35′36″ S, 57º50′37″ W, 296 m, 6 Feb. 2012, pitfall with human and swine dung, R.J. Silva leg. ( CEMT); 2 ♂♂, Tangará da Serra, Fazenda Netolândia, 14º39′56″ S, 57º54′08″ W, 304 m, 20–22 Mar. 2012, pitfall trap baited with human faeces and pig dung, R.J. Silva leg. ( CEMT); 5 ♂♂, 1 ♀, Tangará da Serra, Fazenda Netolândia, 14º39′54″ S, 57º55′08″ W, 329 m, 12–14 Mar. 2012, pitfall with human and swine dung, R.J. Silva leg. ( CEMT); 11 ♂♂, 8 ♀♀, Tangará da Serra, Fazenda Netolândia, 14º39′56″ S, 57º54′08″ W, 304 m, 20–22 Mar. 2012, pitfall with human and swine dung, R.J. Silva leg. ( CEMT); 6 ♂♂, 4 ♀♀, Tangará da Serra, Fazenda Netolândia, 14º41′05″ S, 57º54′08″ W, 263 m, 25–27 Mar. 2012, pitfall with human and swine dung, R.J. Silva leg. ( CEMT); 1 ♀, Tangará da Serra, Fazenda Paraíso, 14º41′46″ S, 57º24′40″ W, 503 m, 13–15 Jan. 2011, pitfall with human and swine dung, R.J. Silva leg. ( CEMT); 2 ♂♂ (1 dissected), 1 ♀, Tangará da Serra, Fazenda Paraíso, 14º41′45″ S, 57º24′38″ W, 520 m, 15–22 Jan. 2011, flight interception trap, R.J. Silva leg. ( CEMT); 4 ♂♂, Tangará da Serra, Fazenda Rosa Branca, 14º33′59″ S, 57º52′33″ W, 312 m, 18–25 Feb. 2011, flight interception trap, R.J. Silva leg. ( CEMT); 3 ♂♂, Tangará da Serra, Fazenda Rosa Branca, 14º33′57″ S, 57º52′34″ W, 321 m, 20–27 Feb. 2011, flight interception trap, R.J. Silva leg. ( CEMT); 2 ♂♂, Tangará da Serra, Fazenda Rosa Branca, 14º34′00″ S, 57º52′24″ W, 468 m, 18–20 Feb. 2011, pitfall with human and swine dung, R.J. Silva leg. ( CEMT); 7 ♂♂, 4 ♀♀, Tangará da Serra, Fazenda Sudamata, 14º37′18″ S, 57º58′01″ W, 354 m, 10–17 Mar. 2012, flight interception trap, R.J. Silva leg. ( CEMT); 3 ♂♂, 1 ♀, Tangará da Serra, Fazenda Sudamata, 14º37′18″ S, 57º58′01″ W, 354 m, 21–23 Feb. 2012, pitfall with human and swine dung, R.J. Silva leg. ( CEMT); 1 ♂, Tangará da Serra, Sítio Mauá, 14º39′39″ S, 57º24′20″ W, 479 m, 19–26 Jan. 2011, R.J. Silva leg. ( CEMT). – Pará: 1 ♀, Belterra, 54º32′36″ W, 02º36′50″ S, 15 Jul. 2016, human and pig dung, F. França leg. ( CEMT); 1 ♂ ([dissected), Óbidos, 4 Mar. 1938, Zellibor-Hauff leg. ( MNRJ); 1 ♂ (dissected), Óbidos, Nov. 1955, C.A.C. Seabra leg. ( CMNC); 1 ♂, 1 ♀, Pau-d’Arco, Fazenda Marajoara, 07º50′ S, 50º16′ W, 13 Oct. 1998, P.Y. Scheffler leg. ( CEMT); 1 ♂ (dissected), Redenção, Pinkaiti-Aik, 07º44′ S, 52º02′ W, Jun. 1999, P.Y. Scheffler leg. ( CEMT); 1 ♂, Redenção, Pinkaiti-Aik, 07º46′ S, 51º58′ W, Oct. 1999, P.Y. Scheffler leg. ( CEMT); 1 ♀, Redenção, Pinkaiti-Aik, 07º50′ S, 50º16′ W, Nov. 1999, P.Y. Scheffler leg. ( CEMT); 3 ♀♀, Tapajós River [1852] , H.W. Bates leg. ( BMNH); 2 ♂♂, 2 ♀♀, Santarém, May 2010, V.H. Oliveira leg. ( CEMT).

Ambiguous data: 1 ♂, “ Amazon ” ( ISNB – Coll. J. Thomson).

Redescription

COLOURATION. Surface of the entire body with very dark shades. Head predominantly purple with greenish reflections, especially at centre. Pronotum with green shine at centre and purplish on the sides. Elytra usually dark green with purplish striae; in some specimens, elytra dark blue. Metaventrite very dark, with coppery or purple reflections. Meso- and metafemora dark brown or reddish-brown. Pygidium dark green.

HEAD. Tegument shiny, with strong alveolar microsculpture and covered by dense micropunctation, which is almost imperceptible or even absent only at the external edge of head ( Fig. 6B ). Clypeus with two apical teeth obtuse and contiguous at base; with a single transverse row of very short setae covering the base of both teeth. Genae with a weak tooth immediately behind clypeal-genal juncture. Posterior edge of head unmargined between eyes.

THORAX. Tegument of pronotum with diffuse shine, dark and with dense micropunctation at centre, denser and more marked than on head; towards the sides, micropunctation less dense and less marked, being completely absent or only very weakly impressed on the lateral margin; tegument with strong microsculpture throughout pronotal surface (southern populations) or with weak or almost absent microsculpture (northern populations). Posterior edge of head with a fine transverse line at centre (usually extending a little beyond the second elytral stria). Hypomeral cavity with tegument entirely glabrous or with only some few long or short yellowish setae at posterior region near to hypomeral carina, but always glabrous at centre; external margin simple, without any trace of tubercle. Metaventrite entirely glabrous; anterior region with tegument with distinct rivose microsculpture; centre and posterior region with dense alveolar microsculpture obliterating micropunctation.

LEGS. Ventral surface of all femora and tibiae dark ( Fig. 25B ). Profemora with tegument with strong alveolar microsculpture at their anterior half and with strong rivose microsculpture at posterior half. Protibiae narrow and straight on internal margin; at their apical seventh, with two tiny, acute external teeth – the apical one slightly larger than the second ( Fig. 11J ). Mesofemora margined anteriorly only at their basal three-fourths; unmargined portion of anterior edge with a row of short setae; tegument with strong rivose microsculpture. Metafemora margined only anteriorly; apical third of anterior edge covered by a row of setae; tegument with strong rivose microsculpture obliterating micropunctation, which is almost imperceptible; without coarse elongate punctation at base. Metatarsomeres II and V subequal in length and longer than the others; metatarsomere IV shorter than the others.

ELYTRA. With only seven narrow visible striae: first four to five striae strongly impressed, very finely carinulate and widened at base; fifth, sixth and seventh striae progressively more effaced and interrupted; all striae lack carenule before reaching apex of elytra, where they are marked only by microsculpture or are completely indistinct; humeral carina absent. Tegument of interstriae either with alveolar microsculpture and very dense micropunctation (southern populations) or with very weak or even absent alveolar microsculpture (northern populations).

ABDOMEN. Ventrite VI smooth at centre and with weaker rivose microsculpture on the sides ( Fig. 14 D– E). Pairs of foveae absent in both sexes. Pygidium with tegument with strong alveolar microsculpture and dense micropunctation; average length equal in both sexes.

AEDEAGUS. Parameres short, less than half as long as phallobase, and slightly asymmetrical: external face of left paramere flat or concave and external face of right paramere convex (difference more easily seen

in ventral view). In lateral view, parameres without ventral keel and with a profound notch at middle ( Fig. 17C ).

SEXUAL DIMORPHISM. Males: Protibial spur wide and bifid, with external projection spiniform, straight, and not very long, and internal projection bent and wide ( Fig. 15B ). Ventrite VI strongly narrowed at middle by emargination of its posterior edge and covered anteriorly by medial projection of ventrite V ( Fig. 14D ). Females: Protibial spur spiniform, simple. Ventrite VI wide at middle and covered anteriorly by medial projection of ventrite V, which gives a narrowed appearance to it (but never as in male) ( Fig. 14E ).

Measurements

Males (N = 13). TL: AV: 6.7 ± 0.56; MX: 7.6; MN: 5.8. EW: AV: 4.8 ± 0.38; MX: 5.3; MN: 4.1. PL: AV: 2.0 ± 0.22; MX: 2.5; MN: 1.8. PW: AV: 4.1 ± 0.29; MX: 4.5; MN: 3.6. PgL: AV: 1.4 ± 0.19; MX: 1.9; MN: 1.1. PgW: AV: 2.0 ± 0.17; MX: 2.3; MN: 1.7.

Females (N = 14). TL: AV: 6.7 ± 0.48; MX: 7.4; MN: 6. EW: AV: 4.8 ± 0.25; MX: 5.3; MN: 4.4. PL: AV: 2.26 ± 0.16; MX: 2.5; MN: 1.9. PW: ME: 4.2 ± 0.2; MX: 4.6; MN: 3.9. PgL: ME: 1.4 ± 0.09; MX: 1.2; MN: 1.5. PgW: AV: 2.1 ± 0.11; MX: 2.3; MN: 1.9.

Geographical distribution

Humid tropical forests from the mouth of the Tapajós River down to the semideciduous forests of southern and southeastern Amazonia in Brazil.

Ecoregions

Uatuma-Trombetas Moist Forests, Madeira-Tapajós Moist Forests, Tapajós-Xingu Moist Forests, Mato Grosso Tropical Dry Forest, Chiquitiano Dry Forest.

Collecting sites ( Fig. 24 )

BRAZIL. Pará: Belterra, Óbidos, Pau-d’Arco, Redenção, Santarém. Mato Grosso: Araputanga, Curvelândia, Diamantino, Indiavaí, Nobres, Nova Mutum, Querência, Tangará da Serra.

Intraspecific variation and taxonomic discussion

Since the beginning of the 20 th century, specimens of Sylvicanthon collected throughout the Amazon region have been misidentified as S. candezei , both in collections and in publications. Nevertheless, even individuals from places as distant from that biome as Rio de Janeiro also suffered from that mistake, as a specimen of S. foveiventris colleted in Itatiaia in 1959 and now deposited in the MNRJ has shown. Two other species that were largely confused with S. candezei are S. seag sp. nov. and S. genieri sp. nov. The great confusion in the recognition of the true identity of S. candezei naturally brought to publication a large amount of incorrect information on the geographical distribution of this species. Schmidt (1922), for example, cited “ Amazonas, Surinam, Cayenne, Ecuador ” as being the distribution of S. candezei , and that was followed by Martínez et al. (1964), who considered this species was distributed in most parts of the Amazonia, including Venezuela, Suriname, French Guiana, Brazil, and Ecuador.

In the present work, we observed that S. candezei has actually a much more restricted distribution than previously thought, being limited to eastern Amazonia, from the mouth of the Tapajós River , in Pará, south to the state of Mato Grosso ( Fig. 24 ). Interestingly enough, in Mato Grosso and southern Pará, specimens of S. candezei were found only at sites in the outer limits of the Amazon forest, i.e., places close to the transition between that biome and the Cerrado and Pantanal; no specimens were collected farther inland in this part of the Amazon region despite extensive collections made in places such as Cotriguaçu, Alta Floresta, Carlinda, and Nova Bandeirante, in Mato Grosso, and Novo Progresso, in Pará. This information tells us that S. candezei has possibly its southern distribution more closely related to the peripherical areas of the Amazon Rainforest, whereas northern populations are present farther inland in that biome.

Apart from this supposed difference in habitat preference, southern and northern populations of S. candezei also exhibit clear morphological distinctions. Specimens collected in Óbidos, Santarém, and in the region of the lower Tapajós River , in Pará, have pronotal and elytral tegument with a more smooth microsculpture, which is sometimes almost absent, so giving a brighter appearance to the entire body and a greater contrast to the elytral striae. Conversely, southern population in southern Pará (Pau d’Arco and Redenção) and in Mato Grosso have a strong microsculpture, with a very dark pronotum and elytra and very subtle elytral striae. Concerning colouration, such a variable feature among other Sylvicanthon , in S. candezei it is much more uniform, with the majority of the specimens of all populations being dark green with a weak metallic sheen.

Comments

The type series of S. candezei was collected by the much-celebrated English naturalist and explorer Henry Walter Bates (1825–1892) ( Fig. 26C ) during his long and famous collecting period in the Amazon Basin between 1848 and 1859. As the type locality of S. candezei, Harold (1869a) cited “ Tapajos ” and this information is indeed present on the labels of all the syntypes. Nevertheless, the Tapajós River is one of the largest tributaries of the Amazon, covering almost eight hundred kilometres from its formation in the convergence of the Teles Pires and Juruena Rivers to its mouth situated near the city of Santarém, in Pará. Therefore, where exactly in this vast area would have Bates collected the oldest known specimens of S. candezei ?

In his journey’s narrative, Bates (1863) dedicated a full chapter to his exploration of the Tapajós River. After spending almost six months in Santarém, a city on the banks of the Amazon, Bates decided to explore the lower region of the Tapajós . With that goal in mind, he rented a suitable vessel, hired a small crew and set off on the 8 th of June 1852. For the next four months, he navigated approximately 170 km upstream in the Tapajós and made collections in four main localities: Alter do Chão and Aveiro, on the right banks of the Tapajós , and Santa Cruz, on the left banks, besides having gone up for a few days the much narrower Cupari River, a tributary on the right banks of the Tapajós ( Fig. 26B ). At the end of this trip, Bates returned to Santarém, from where he then left to go up the Amazon River towards Ega (nowadays known as Tefé). By reading his accounts, it is not possible to find any mention to dung beetle collections, although it is clear that Bates collected a large number of insects in that region. Therefore, although we do not have the exact locality where Bates caught the type series of S. candezei , we do know at least that it was in the lower Tapajós area, very likely either in Alter do Chão, Aveiros, Santa Cruz or at some point along the Cupari River, between June and October 1852.

Apart from the nine specimens listed in the type series above, three other females collected by Bates in the Tapajós were found in the BMNH collection. Two of them bear circular blue labels handwritten “ Tapayo ”, on one side, and “ 54 / 18 ”, on the other. According to Max Barclay (personal communication to MC, 2015), curator of Coleoptera at the BMNH, this code refers to a lot consisting of 237 beetles collected by Bates in “Amazon (Alta de Chia) 100 miles from Santaren on the Japayos”. This lot was bought in 1854 by the BMNH at the J.C. Stevens Auction Rooms, London, an auction house specialized in natural history and antiquities and then property of J.C. Stevens, brother of Samuel Stevens, English naturalist who arranged Bates’ travel to the Amazon region and from whom he received material there collected ( Anonymous 1899; Carrington 1899). ‘Alta de Chia’ certainly refers to Alter do Chão, whereas ‘Japayos’ is a corruption of Tapajós . The third female in the BMNH, differently, has a circular blue label handwritten “ Braz / Tapajos ”, on the one face, and “ 53 / 27 ”, on the other, code referring to, according to Barclay, 232 beetles collected by Bates in “ Brazil, Banks of the Rio Japayos” and also auctioned by Stevens in 1854.

As none of those three females bears any label with Edgar von Harold’s handwriting, we do not consider them syntypes of S. candezei . Nonetheless, two of them have rectangular pinkish identification labels with other names: one was identified as “ castanipes Reiche ”, nomen in litteris that we do not know whether it was cited in the literature before, while the other specimen is labelled “S errimanus mihi ” (mihi, from the Latin, meaning “belonging to me”; Schenk & McMasters 1956). According to Harold (1868a, 1869b), “ serrimanus ” was a nomen in litteris used by the French entomologist Auguste Chevrolat (1799–1884) to refer to the species named by Harold as C. chevrolati Harold, 1868 , nowadays a subspecies of Canthon indigaceus LeConte, 1886 , from North and Central America . Therefore, in principle, Chevrolat should have been the author of that identification label; indeed, although the majority of his Scarabaeinae collection is currently housed in the SMTD (via C. Felsche collection), other Coleoptera groups of his collection, such as Cerambycidae , were deposited at the BMNH ( Horn & Kahle 1935). Even so, as C. indigaceus chevrolati is completely distinct from S. candezei in both morphology and distribution, it would be curious that such an experienced entomologist as Chevrolat could have confused them. Alternative explanations are that Harold perhaps confused the actual application of the name “ serrimanus ” by Chevrolat and that this latter entomologist used that name to refer to the current S. candezei , or that, in reality, the label’s author was not Chevrolat but, instead, a third unidentified entomologist who used the name ‘ serrimanus ’ independently from Chevrolat to refer to S. candezei . Although these nomina in litteris are unavailable and, therefore, cannot affect the validity of the name Canthon candezei , tracing their histories is worthy because it allows us to better understand the dynamics of discovering and naming taxa in the 19 th century, as well as communication between entomologists during the period.

Natural history

No information about the biology of S. candezei has been published until now. Label data state the species occurs in altitudes between 263 and 520 m and is attracted to pitfall traps baited with human faeces and cow and pig dung, and that it can also be collected using flight interception traps. Apparently, adults of S. candezei are active throughout the year, having been collected between January and May, and in July, October and December. Nonetheless, it is worth noting that it was during the first four months of the year that 115 out of the 157 examined specimens were caught (January: 35 specimens; February: 20; March: 47; April: 13), indicating that, although active throughout the year, S. candezei should be more abundant during the rainiest and hottest months than during the dry season in the middle of the year.

Figures

Fig. 6. Head, dorsal view. A. Sylvicanthon enkerlini (Martínez et al., 1964) comb. nov. B. S. candezei (Harold, 1869). C. S. genieri sp. nov. D. S. proseni (Martínez, 1949) stat. et comb. nov. E. S. seag sp. nov. F. S. securus (Schmidt, 1920) comb. nov. G. S. furvus (Schmidt, 1920).

Fig. 10. Lateral view. A. Canthon (Francmonrosia) sp. B. Sylvicanthon enkerlini (Martínez et al., 1964) comb. nov. C. S. candezei (Harold, 1869).

Fig. 11. Morphological diversity in the protibiae of Sylvicanthon Halffter & Martínez, 1977. A. S. enkerlini (Martínez et al., 1964) comb. nov. B. S. aequinoctialis (Harold, 1869) comb. nov. C. S. securus (Schmidt, 1920) comb. nov. D. S. obscurus (Schmidt, 1920). E. S. furvus (Schmidt, 1920). F. S. bridarollii (Martínez, 1949) (southern populations). G. S. bridarollii (northern populations). H. S. attenboroughi sp. nov. I. S. edmondsi sp. nov. J. S. candezei (Harold, 1869). Note the variation related to the presence and degree of development of an expansion on the internal edge of protibiae and to the number, size and position of the external teeth.

Fig. 14. A. Ventral view of the anterior region of the pterothorax of S. proseni (Martínez, 1949) stat. et comb. nov. Abbreviations: MsEs = mesoventrite; SEs = meso-metaventrite suture; MsC = mesocoxa; RAMtEs = anterior region of metaventrite. B–E. Sexual dimorphism in the abdominal ventrites of Sylvicanthon Halffter & Martínez, 1977. B. S. foveiventris (Schmidt, 1920), ♀. Note the three lateral foveae between ventrites I–II, II–III, and III–IV and the broad ventrite VI at the middle, without emargination on its posterior edge and without its anterior edge covered by a medial flange of ventrite V. C. S. proseni stat. et comb. nov., ♀, ventrite VI broad. D. S. candezei (Harold, 1869), ♂. Note how the ventrite VI is narrow at the middle due to the emargination on its porterior edge and to the medial flage of the posterior edge of the ventrite V. E. S. candezei (Harold, 1869), ♀. Despite its posterior edge not being emarginate, its anterior edge is covered by a well-developed medial flange of ventrite V, which gives a narrowed aspect to the centre of ventrite VI.

Fig. 26. Sylvicanthon candezei (Harold, 1869) type material. A. a. Lectotype. b. Labels of lectotype. c. Labels of paralectotype 3. B. The four localities where S. candezei type series may have been collected by Bates in the lower Tapajós region in 1852 (see discussion in the text). C. The English naturalist Henry Walter Bates (1825–1892), the collector of the S. candezei type series (modified from Bidau 2014).

Fig. 25. Sylvicanthon candezei (Harold, 1869). A. Dorsal view. B.Ventral view.

Fig. 17. Aedeagus morphological diversity in Sylvicanthon. A. S. enkerlini (Martínez et al., 1964) comb. nov. B. S. foveiventris (Schmidt, 1920). C. S. candezei (Harold, 1869) (arrows points to the ventral notch). D. S. genieri sp. nov. (idem). E. S. aequinoctialis (Harold, 1868). F. S. proseni (Martínez, 1949) stat. et comb. nov.

Fig. 15. Morphological variation and sexual dimorphism in the protibial spur of Sylvicanthon. A. S. enkerlini (Martínez et al., 1964) comb. nov., ♂. B. S. foveiventris (Schmidt, 1920), ♂. C. S. candezei (Harold, 1869), ♂. D. S. genieri sp. nov., ♂. E. S. aequinoctialis (Harold, 1868), ♂. F. S. aequinoctialis comb. nov., ♀. G. S. proseni (Martínez, 1949) stat. et comb. nov., ♂. H. S. proseni stat. et comb. nov., ♀. I. S. bridarollii (Martínez, 1949), ♂. J. S. seag sp. nov., ♂. K. S. edmondsi sp. nov.,♂. L. S. attenboroughi sp. nov., ♂. M. S. obscurus (Schmidt, 1920), ♂. N. S. furvus (Schmidt, 1920), ♂. O. S. securus (Schmidt, 1920) comb. nov., ♂.

Fig. 24. Distribution of the three species of the candezei subgroup.