Epicadinus villosus Mello-Leitão, 1929

Prado, André Wanderley Do, Baptista, Renner Luiz Cerqueira & Machado, Miguel, 2018, Taxonomic review of Epicadinus Simon, 1895 (Araneae: Thomisidae), Zootaxa 4459 (2), pp. 201-234 : 223-229

Epicadinus villosus Mello-Leitão, 1929

( Figs 11–13 , 15 )

Epicadinus villosus Mello-Leitão, 1929: 103 .

Epicadinus helenae Piza, 1936: 50 , fig. 1. Soares 1943: 12, fig. 10. New synonymy.

Epicadinus marmoratus Mello-Leitão, 1947: 274 . New synonymy.

Type-material: Holotype of Epicadinus helenae : subadult female, BRASIL, São Paulo: Piracicaba, 1936 , M. Diniz (IBSP 138.217 [ex. MZLQ 117], Figs 13D–F ), examined. Holotype of Epicadinus marmoratus : juvenile, BRASIL, Paraná: Curitiba, Barigui, x.1944, R.B. Lange (MNHCI 0 2496 [Coll. Rudolf B. Lange], Figs 13G–I ), examined. Holotype of Epicadinus villosus : female, BRASIL, Bahia: “ S. Antônio da Barra ” (= Santo Antônio de Condeúba, MNHN AR-15078 [Coll. Simon 11.511], Figs 13A–C ), examined. Paratypes of Epicadinus villosus : female, PARAGUAY (MNHN [Coll. Simon 9752]); female, PARAGUAY (MNHN [Coll. Simon 19194]).

Additional material examined. ARGENTINA: Misiones: 1 male ca. 27°25'33"S, 55°56'44"W, February 1950 ( MACN 19053 ) ; idem, 1 female, October 1953, Schiapelli, De Carlo, Viana, Galiano (MACN 3802); idem, 1 male, October 1953 (MACN 3803); 2 females, General Manoel Belgrano , ca. 26°04'26"S, 53°59'35"W, 1954, Schiapelli & De Carlo ( MACN 19054 ) ; 1 female, Montecarlo , ca. 26°33'41"S, 54°46'08"W, January 1966, M. E. Galiano ( MACN 19064 ) ; idem, 1 female (MACN 19066); 2 females, Parque Nacional Iguazú , ca. 25°41'08"S, 54°27'39"W, August 1985, M. Ramirez ( MACN 19058 ) ; idem, 1 female, 8–15 February 1995, M. Ramirez (MACN 19060); 3 females, Parque Nacional Iguazú, Ruta 101, ca. 25°39'12"S, 54°19'36"W, January 1966, Galiano (MACN 19065); 1 male, Parque Nacional Iguazú, Area Cataratas, ca. 25°41' 05"S, 54°26'46"W, 13–15 September 1992, Goloboff (MACN 19052); idem, 1 male, November 1989, M. Ramirez & L. Lopardo (MACN 19056); idem, 1 male, 22–30 August 1986, M. Ramirez & L. Lopardo (MACN 19057); idem, 1 female, 11–16 December 1999, M. Ramirez & L. Lopardo (MACN 19062); 1 male, Parque Nacional Iguazú, RN 101, E. seccional Yacuy , ca. 25°36'27"S, 54°18'52"W, 14–16 December 1999, M. Ramirez & L. Lopardo ( MACN 19063 ) ; 1 female, San Pedro, Tobuna , ca. 26°28'03"S, 53°53'53"W, February 1952, W. Partridge ( MACN 04064 ) . BRASIL: Minas Gerais: 1 female, Belo Horizonte, Estação ecológica da UFMG 19°52'28"S, 43°58'22"W, January 2001, E.S.S. Álvares ( UFMG 129 ) ; idem, 1 male, 1 female, 1 juvenile, 14 November 2009, G.H.F. Azevedo et al. (UFMG 3389); idem, 1 male, 1 female, x.2000, no data (UFMG 5959); 1 female, Catas Altas, Serra do Caraça , ca. 20°08'33"S, 43°29'07"W, 23–26 November 1960, Araújo & Martim ( MZUSP 13626 ) ; 1 male, Sete Lagoas , Serra Santa Helena, 19° 26' 43"S, 44° 16' 20"W, 26 November 2001, E.S.S. Álvares ( IBSP 40263 ) . Pará: 1 male, 1 female, Santarém , ca. 2°27'11"S, 54°41'47"W ( MNHN Coll. Simon 16064) . Paraná: 1 female, ca. 25°25'35"S, 49°15'59"W, 15 April 2005, J. Ricetti ( PUCRS 37233 ) ; 1 male, 1 female, Candói, Usina Hidrelétrica de Segredo ca. 25°47'19"S, 52°06'25"W, April 1996, R. Bertani ( IBSP 11665 ) ; 1 female, Curitiba , ca. 25°25'35"S, 49°15'59"W, 1938, Pe. F. S. Pereira ( MZUSP 00004 ) ; 1 male, 1 female, Curitiba, Barigui ca. 25°31'22"S, 49°20'52"W, 20 April 1962, Lange ( MHNCI) ; 1 male, Curitiba, Bom Retiro, ca. 25°24'39"S, 49°16'32"W, 10 April 1987, A. D. Brescovit (FZBRS 16739); 1 female, Curitiba, Parque da Cidade , ca. 25°30'12"S, 49°21'07"W, 1 December 1990, A. D. Brescovit ( FZBRS 20627 ) ; 4 females, Fênix , Parque Estadual Vila Rica do Espírito Santo, ca. 23°55'00"S, 51°57'18"W, 22 November 1987, A. D. Brescovit ( IBSP 14184 ) ; 1 female, 2 juvenile, Foz do Iguaçu, Parque Nacional do Iguaçu , ca. 25°41'08"S, 54°27'39"W, 29 March 1993, A B. Bonaldo ( PUCRS 4310 ) ; 7 females, Foz do Iguaçu, Refúgio Biológico de Bela Vista , ca. 25°26'55"S, 54°33'22"W, 17 March 1994, A. B. Bonaldo ( FZBRS 20939 ) ; 1 female, Morretes, Serra da Graciosa ca. 25°25'19"S, 48°52'05"W, 9–20 January 1995, Eq. Lab. Aracnologia ( PUCRS 7026 ) ; idem, 1 male, 2 females, 2 juveniles (PUCRS 7209); 1 male, 5 females, A. A. Lise (PUCRS 7347); 1 male, 1 female, São José dos Pinhais , 25°36'18"S, 49°11'37"W, 9 December 2015, A. C. Domahovski ( PUCRS 39104 ) ; idem, 1 male, October 2015, A. C. Domahovski (PUCRS 39069); 2 females, 1 juvenile, Três Barras do Paraná, ca. 25°25'14"S, 53°11'04"W, 26 February 1993, A. B. Bonaldo ( PUCRS 3035 ) ; 1 female, Três Barras do Paraná, Rio Guarani, Foz do Córrego Três Barras , ca. 25°26'28"S, 53°07'23"W, 20-26 February 1993, A. B. Bonaldo ( FZBRS 23042 ) . Rio de Janeiro: 1 male, 1 female, Itatiaia, Parque Nacional de Itatiaia , ca. 22°22'12"S, 44°37'46"W, 28 October 1967 ( MACN 19055 ) ; 1 female, Mendes, Centro Marista São José das Paineiras , 22°30'35"S, 43°45'16"W, 21 November 2014, A. W. Prado ( UFRJ 1330 ) . Rio Grande do Sul: 1 male, Augusto Pestana , ca. 28°30'51"S, 53°59'40"W, 12 December 2008, L. V. Silva ( PUCRS 27091 ) , idem,1 female, 6 September 2009, L. V. Silva & L. B. Medeiros (PUCRS 30592); idem, 1 female, 25 October 2009, L. V. Silva & L. B. Medeiros (PUCRS 30639); 1 female, Bom Jesus, Fazenda Aver, ca. 28°40'23"S, 50°26'24"W, 24 March 1989, A. B. Bonaldo (FBZRS 18426), idem, 1 female, 1 April 1988 (FZBRS 17395); 1 male, 1 female, Cambará do Sul, Itaimbézinho , ca. 29°09'40"S, 50°04'39"W, 28 May 1983, A.A. Lise ( FZBRS 11760 ) ; 2 males, Canela , ca. 29°21'22"S, 50°49'10"W, 3 June 1966, A. A. Lise ( FZBRS 0238 ) ; idem, 1 female, 20 February 1972 (FZBRS 1026); idem, 1 female, 31 December 1973 (FZBRS 2056); idem, 1 female, 2 December 1973 (FZBRS 2207); idem, 2 males, 1 female, 5 juveniles, 26 December 1974 (FZBRS 2497); idem, 2 males, 20 February 1973 (FZBRS 8062); idem, 1 male, 3 females, 1 juvenile, 19 March 1976 (FZBRS 8063); idem, 1 male, 1 female, 4 juveniles, 26 December 1975 (FZBRS 8064); idem, 1 male, 8 November 1975 (FZBRS 8067); idem, 1 female, 20 March 1976 (FZBRS 8069); idem, 1 male, 2 March 1976 (FZBRS 8071); idem, 1 male, 2 March 1976 (FZBRS 8074); idem, 1 male, 1 female, 20 March 1976 (FZBRS 8078); idem, 1 female, 2 March 1977 (FZBRS 8129); 1 female, Usina Hidrelétrica da Canastra , ca. 29°23'39"S, 50°44'45"W, 20 November 1997, M. A. L. Marques ( FZBRS 28844 ) ; 1 female, Caxias do Sul, Faz. Souza , ca. 29°09'07"S, 50°57'21"W, 11–12 November 1995, Eq. Lab. Aracnologia ( PUCRS 7320 ) ; 7 males, 5 females, Derrubadas, Parque Estadual do Turvo , ca. 27°12'26"S, 53°56'07"W, 28–31 October 2003, R. Ott ( FZBRS 37824 ) ; idem, 1 male, 1 female, 27–31 October 2003 (FZBRS 38863); idem, 4 females, 4–08 May 2004 (FZBRS 38867), idem, 1 male, 1 female, 19–22 October 2004 (FZBRS 38875); idem, 7 males, 4 females, 19–22 October 2004 (FZBRS 38881); idem, 6 females, 25–30 April 2005 (FZBRS 39201); 2 males, Estrela Velha, Barragem Itaúba, ca. 29°15'40"S, 53°14'26"W, 7 March 2001, R. Ott (FZBRS 33698); 1 female, Farroupilha , ca. 2913'33"S, 51°20'18"W, 29 September 1978, A. A. Lise ( FZBRS 8271 ) ; idem, 2 juveniles, H. Bischoff (PUCRS 8272); 1 female, Guaíba , ca. 30°06'24"S, 51°19'27"W, 3 June 1994, A. A. Lise ( PUCRS 4817 ) ; 1 female, 1 juvenile, Guaíba, Fazenda São Maximiano, ca. 30°10'53"S, 51°22'56"W, 15 March 1996, A. A. Lise (PUCRS 8286); 2 males, 1 female, Iraí , ca. 27°11'32"S, 53°14'54"W, 21 November 1975, A. A. Lise ( FBZRS 8065 ) ; idem, 11 males, 7 females, 10 juveniles (FZBRS 8061); idem, 1 female, 18 November 1975 (FZBRS 8908); 1 female, Itaara , ca. 29°35'40"S, 53°45'34"W, 16 January 2007, A. A. Lise ( PUCRS 21344 ) , idem, 1 female, 13 February 2007 (PUCRS 21344), idem, 1 female, 12 May 2007 (PUCRS 21346); 1 male, 1 female, Itaúba, Arroio do Tigre , ca. 29°16'25"S, 53°9'01"W, 22 April 1978, A. A. Lise ( FZBRS 8037 ) , 2 males, 22 April 1978 (FZBRS 8038); 1 female, Machadinho, Linha do Tigre , ca. 27°35'17"S, 51°36'25"W, 9–10 May 2001, R. Ott & L. Moura ( FZBRS 33874 ) ; 1 female, Marcelino Ramos , ca. 27°28'31"S, 51°55'30"W, February 1989, A. Braul ( PUCRS 19807 ) ; 1 juvenile, Montenegro, ca. 29°41'03"S, 51°28'20"W, 3 November 1977, M. E. L. de Souza ( FZBRS 7093 ) ; idem, 1 male, M. H. Galileo (FZBRS 8070); 1 male, 3 females, 5 juveniles, Nonoai, Parque Estadual de Nonoai , ca. 27°25'47"S, 53°02'25"W, 14 January 1985, A. A. Lise ( FZBRS 13076 ) ; 1 female, Novo Cabrais, Parque Witeck, ca. 29°46'59"S, 52°58'18"W, 3 January 2008, R. G. Buss (PUCRS 20302); idem, 1 female, 19.x.2007 (PUCRS 28327); 2 males, 2 females, 1 juvenile, Porto Alegre, Ponta Grossa, ca. 30°10'48"S, 51°11'11"W, 7 May 1976, A. A. Lise (FZBRS 8073); 1 male, Porto Alegre, Rio Apuaê , BR 126, ca. 27°50'47"S, 51°49'59"W, September 1988, Equipe Proj. Ita-Machadinho ( PUCRS 0537 ) ; 1 female, Salto do Jacuí, Horto CEEE, ca . 29°04'49"S, 53°12'56"W, 19 October 1998, A. B. Bonaldo (FZBRS 30762); 1 female, Santa Maria, Perau Velho, ca. 29°39'14"S, 53°46'45"W, 19 January 1995, C. B. Kotzian & L. Indrusiak (PUCRS 10467); idem, 1 female, 19 November 1998 (PUCRS 38376); 1 male, São Borja, Reserva Biológica São Donato, ca. 29°01'06"S, 56°09'43"W, 18 January 2012, M. Machado (PUCRS 36898); 1 male, São Francisco de Paula , ca. 29°26'47"S, 50°34'48"W, 23–25 November 1998, A. Silva ( FZBRS 30819 ) ; 1 male, São Francisco de Paula, Barragem dos Bugres , ca. 29°20'41"S, 50°42'07"W, 1–4 February 1999, A. Silva ( FZBRS 30854 ) ; 1 female, São Francisco de Paula, Fazenda 3 Cachoeiras , ca. 29°27'29"S, 50°33'38"W, 5 November 1998, A. Silva ( FZBRS 30863 ) ; 1 female, 3 February 1999, A. B. Bonaldo (FZBRS 30961); 1 female, 2 juveniles, São Francisco de Paula, Usina Hidrelétrica Passo do Inferno , ca. 29°16'09"S, 50°45'W, 26 September 2000, M. A. L. Marques ( FZBRS 33241 ) ; idem, 4 females, 1 juvenile, 19 November 1997, E. H. Buckup (FZBRS 28815); 2 males, 2 juveniles, Tenente Portela , ca. 27°22'07"S, 53°45'35"W, 29 November 1978, H. Bischoff ( PUCRS 8394 ) ; 1 female, Tenente Portela, Parque Estadual do Turvo , ca. 27°12'26"S, 53°56'07"W, 15 January 1985, A. A. Lise ( FZBRS 13074 ) ; idem, 1 female, 1 juvenile, 4–6 February 1980 (FZBRS 8983); 7 males, 20 females, 7 juveniles, Tenente Portela, Parque Estadual do Turvo , Salto do Yucumã, ca. 27°08'37"S, 53°52'57"W, 16 January 1985, A. A. Lise ( FZBRS 13056 ) ; idem, 7 females, 7 juveniles, 17 January 1985, A. A. Lise (FZBRS 13065); 1 female, Triunfo , ca. 29°56'12"S, 51°42'44"W, 28 November 1989, E. H. Buckup ( FZBRS 19130 ) ; idem, 1 female, 25 January 1990, A. D. Brescovit (FZBRS 19343); idem, 1 female, 13 January 1994, L. A. Moura (FZBRS 24772); idem, 2 females, 16–17 March 1998, L. Moura (FZBRS 29239); idem, 1 male, 1 female, 19 May 1977, A. A. Lise (FZBRS 5391); 1 male, 1 female, Triunfo, Parque Copesul de Proteção Ambiental , ca. 29°55'46"S, 51°42'27"W, 28 November 2002, R. Araujo ( FZBRS 34998 ) ; idem, 1 female, 5 February 2003, R. Ott (FZBRS 35152); idem, 1 female, 29 July 2003, R. Ott & A. Barcellos (FZBRS 36093); idem, 1 female, 15 March 2007, M. A. L. Marques (FZBRS 43256); 2 females, Viamão , ca. 30°04'37"S, 51°01'40"W, 1 May 1976, A. A. Lise ( FZBRS 8066 ) ; idem, 1 male, 7 July 1995 (PUCRS 6669); 2 females, 1 juvenile, Viamão, Águas Belas, ca. 30°02'38"S, 50°59'59"W, 17 January 1977, A. A. Lise (FZBRS 4978); 1 female, 29 December 1976, A. A. Lise (FZBRS 5861); 1 female, Viamão, Estação Fitotécnica , ca. 30°02'15"S, 51°01'20"W, 6 May 1994, A. A. Lise ( PUCRS 4688 ) ; idem, 1 male, 12 August 1994 (PUCRS 5238); 1 male, Viamão, Parque Estadual Itapuã, ca. 30°20'42"S, 51°01'31"W, April 2002, L. Ernesto ( PUCRS 15331 ) . Roraima: 1 female, Ilha de Maracá, ca. 3°25'00"N, 61°40'00"W, 21–30 November 1987, J. A. Rafael ( INPA) . Santa Catarina: 2 males, 3 females, Rancho Queimado , ca. 27°40'17"S, 49°00'53"W, 13–15 January 1995, A. B. Bonaldo ( FZBRS 26470 ) ; idem, 2 males, 3 females, 13–18 November 1995 (FZBRS 27046), idem, 2 females, 8–12 October 1994 (PUCRS 5949), idem, 2 males 3 females, 4 juveniles, 13–15 January 1995 (PUCRS 6137). São Paulo: 1 female, Paranapiacaba , ca. 23°46'32"S, 46°17'49"W, 9 October 1956, W. Bockerman ( PUCRS 19738 ) ; 1 female, Rosana, ca. 22°34'47"S, 53°03'33"W, December 1986, Equipe IBSP ( IBSP 4484 ) ; 1 female, Salesópolis, Estação Biológica Boracéia, Trilha Laboratório , ca. 23°39'21"S, 45°53'29"W, 1 April 2001, S.T.P. Amarante et al. ( MZUSP 70593 ) ; 1 female, Santo André, Reserva Biológica do Alto da Serra de Paranapiacaba , ca. 23°47'10"S, 46°20'40"W, 14–16 December 2003, C.A. Rheims & R.P. Indicatti ( IBSP 52069 ) , idem, 1 male, ca. 23°48'24"S, 47°07'30"W, 28 October 2007, Equipe Butantan (IBSP 88546); 2 males, 1 female, São Paulo, Parque Estadual do Jaraguá , ca. 23°27'34"S, 46°45'19"W, 27–28 October 2005 ( IBSP 71290 ) ; 2 females, São Paulo, Horto Florestal , ca. 23°27'22"S, 46°37'48"W, 27 March 1948, O.P. Forattini ( MZUSP 13624 ) . URUGUAY: Cerro Largo: 1 female, Paso Centurión, 32°08'01"S, 53°43'57" W, 13 April 2017, Bruno da Silva (FCE Ar 6184) ; 1 female, 15 April 2017, S. Teijón et al. (FCE Ar 6352); idem, 1 female (FCE Ar 6382), idem, 1 female, 1 juvenile (FCE Ar 6399); idem, 1 male, 6 November 2017 (FCE Ar 8444) ; 1 male, Sierra de Ríos , 32°12'55"S, 53°45'40"W, 14 April 2017, S. Teijón et. al. (FCE Ar 6483).

Diagnosis. E. villosus closely resembles E. spinipes , both belonging to the spinipes -group. Females of E. villosus are distinguished from E. spinipes by the presence of a wide, median, brown longitudinal stripe along the carapace, bifurcating in the posterior portion, the dorsum of the opisthosoma dark-brown to black, without white lateral stripes, and dark-brown sternum with central yellowish area ( Figs 11A, B ). The paired macrosetal tubercles of the tibia and metatarsus of E. villosus are shorter than in E. spinipes , with a length less than half the width of the article. The epigynum has a rectangular median septum, with an anterior base almost as wide as the posterior one ( Fig. 11F ), and spiral copulatory ducts with approximately seven coils, the last one not covering the spermatheca ( Fig. 11E ). Males of E. villosus are distinguished from E. spinipes by the reddish-brown sternum bordered by dark brown spots ( Fig. 12C ), by the RTA just slightly longer than wide ( Fig. 12F ), and by the embolus forming about six turns around the tegulum ( Figs 12E, F ).

Notes. According to Mello-Leitão (1929), E. villosus was based on females from "Santo Antônio da Barra" (now Santo Antônio de Condeúba), Bahia, and Paraguay, and the type was deposited under #11511 from Coll. Simon. Since vial #11511 contains only one female labelled as “ type ” and collected in Bahia ( Figs 13A–C ), it is considered as the holotype of the species. The two females from Paraguay were only identified as E. villosus by Mello-Leitão but were not labeled as type material and were deposited under different numbers (Coll. Simon #19194 and #9752). Taking into account the article 72.4.1 of the ICZN (1999), these females are part of the typeseries and are considered as paratypes of E. villosus . The holotype of Epicadinus helenae ( Fig. 13D–F ) is not an adult female, as stated in the original description. It is a subadult female, with only a primordium of epigynum and not displaying fully-developed genitalic structures. However, this specimen presents the same color pattern and disposition of macrosetae and relative position of the conical projections of opisthosoma as adult females of E. villosus . In addition, there are several records of E. villosus for the interior of the state of São Paulo. The only other records for Epicadinus species for this state are a single specimen of E. spinipes , from northern coast of São Paulo, and seven specimens of E. trispinosus , from the border with the state of Mato Grosso do Sul. The female identified as a "homeotype" of E. helenae by Soares (1943), from the city of Curitiba, is also cospecific with E. villosus . The last synonym is Epicadinus marmoratus , described by Mello-Leitão (1947) based on a "male" holotype from Barigui, Curitiba. However, the holotype is a juvenile female ( Fig. 13G–I ), which also shows the typical color pattern of E. villosus and a wide and long posterior conical projection. Besides, the only species of the genus recorded from the Brazilian state of Paraná is E. villosus .

Description. Female (FZBRS 39201, Fig. 11A–C ): Total length: 4.87. Carapace: length 2.02, width 2.17, height 1.20. Clypeus: height 0.21. Chelicera: length 0.56, width 0.35. Sternum: length 0.84, width 0.95. Endites very setaceous: length 0.45, width, 0.26. Labium: length 0.25, width 0.37. Eye diameters and interdistances: AME 0.09, ALE 0.10, PME 0.07, PLE 0.06, AME–AME 0.013, AME–ALE 0.09, ALE–ALE 0.31, PME–PME 0.16, PME–PLE 0.10, PLE–PLE 0.56. MOQ trapezoidal. Legs 1234. Leg I: femur 1.91, patella 0.99, tibia 1.63, metatarsus 1.35, tarsus 0.71, total 6.59; II: 1.50-0.97-1.52-1.21-0.72-5.92; III: 1.00-0.55-0.66-0.48-0.31-3.00; IV: 1.30-0.55-0.70-0.62-0.40-3.57. Paired macrosetal tubercles in tibiae and metatarsi shorter than half width of the article. Opisthosoma: length 2.85, width 3.37, height 2.10, pentagonal; anterior margin of the dorsum concave in dorsal view and projecting over the posterior slope of the carapace, with projected anterior angles. Paired projection with blunt apex and placed transversally in the median portion. Posterior conical projection with blunt apex, slanted, almost parallel to the venter of opisthosoma, a little longer than the paired ones, and reaching 2/3 of the height of opisthosoma. In lateral view, anterior portion of the dorsum nearly flat and not clearly aligned to the posterior conical projection, and posterior portion slightly convex.

Epigynum ( Fig. 11D–F ) mushroom-shaped, slightly wider than long. Atrium reniform, approximately twice as wide as long, located anteriorly, formed by a deep depression, with the deepest areas forming two lateral foveas, whose margins are covered by a thin translucent chitinous keel. Anterior portion of the atrium rises gradually reaching the level of the surface of the opisthosoma. Median septum almost as long as wide, with an anterior base slightly wider than posterior base and projecting itself laterally, covering the copulatory openings and merging into the interior of the atrium. Its posterior base fuses to an area near the epigastric furrow, placed far from atrium. Median portion of the septum very high, covering the copulatory openings and the middle portion of the posterior margins of the atrium, both located in large and deep semicircular depressions. Margins of the epigynal plate present long cilium-shaped bristles, which partially cover the atrium. Vulva ( Fig. 11E ) with very long spiral copulatory ducts and a pair of posterior rounded spermathecae. Copulatory ducts initially formed by a simple, curved duct, followed by a spiral portion directed anteriorly, with approximately six coils lighter in hue, with a more dilated apical portion (secondary spermatheca). From the inner margin of this structure originates an inner darker spiral portion, formed by approximately seven coils, which runs posteriorly in opposite direction to first spiral, with a thicker ending portion which does not cover the spermathecae, and forming an almost straight distal duct connected to the posterior inner face of the rounded and well-developed spermathecae.

Coloration. Carapace predominantly dark brown with wide dark brown longitudinal median stripe ( Fig. 11A ), bifurcated only in its posterior third, and dark brown wide marginal stripes. Remaining areas yellowish around and at the splitting of posterior part of central stripe. Sternum mostly dark brown with a central light yellowish area. Labium, endites and chelicerae dark brown. Legs I and II predominantly dark brown, except for light yellow tarsi. Legs III and IV light yellow, with irregular dark brown spots and rings. Opisthosoma predominantly dark brown with venter brownish yellow and with light brown sigilas.

Male (MACN 19052, Fig. 12A–C ): Total length: 3.39. Carapace: length 1.59, width 1.46, height 0.88. Clypeus: height 0.17. Chelicera: length 0.43, width 0.23. Sternum very setaceous: length 0.65, width 0.63. Endites very setaceous: length 0.28, width, 0.26. Labium: length 0.13, width 0.26. Eye diameters and interdistances: AME 0.07, ALE 0.06, PME 0.05, PLE 0.04, AME–AME 0.13, AME–ALE 0.08, ALE–ALE 0.29, PME–PME 0.15, PME–PLE 0.11, PLE–PLE 0.46. MOQ trapezoidal. Legs 1243. Leg I: femur 1.28, patella 0.58, tibia 0.96, metatarsus 0.81, tarsus 0.48, total 4.11; II: 1.23-0.66-0.88-0.73-0.46-2.73; III: 0.63-0.38-0.57-0.39-0.33-2.30; IV: 0.75-0.34-0.62-0.43-0.33-2.47. Paired macrosetal tubercles in tibiae and metatarsi shorter than half the width of the article. Opisthosoma: length 1.80, width 2.20, height 1.30, pentagonal, covered by abundant macrosetae of variable size. Anterior margin slightly concave in dorsal view and anterior angles a little projected over the posterior slope of the carapace. Paired projections thin, with acute apex directed posterolaterally, and arranged transversally in median portion of opisthosoma. Posterior conical projection very slanted posteriorly, not reaching half the height of the opisthosoma itself, and similar in size to paired projections. In lateral view, anterior portion of dorsum nearly flat and almost aligned to posterior conical projection, and posterior portion slightly convex.

Palpus ( Fig. 12D–F ). Tegulum with transversal furrow at 1 o’clock position. Embolus basis gradually separating from tegulum, initially forming a wide stripe, slightly tapering during its median portion. Embolus describes about six turns around lateral margin of tegulum, from which at least four are visible in non-distended bulb in retrolateral view, and ends in a filiform apex, hidden by margin of tegulum. RTA fused to DTA, both connected to the tibia by an elongated and narrow base, forming a kind of pedestal. RTA short, slightly longer than wide; RTA also presents a large median concavity, marginal folds and apex describing a large rounded curve. Dorsal fold slightly projected over the median concavity, joining directly to base of DTA. Ventral fold slightly more projected on the median concavity than the dorsal one. DTA short, hook-shaped, with very thick base, excavated at the inner surface of the hook, and apex forming a blunt tip.

Coloration. Carapace dark brown, with two longitudinal median light brown stripes. Sternum reddish brown, radially bordered by dark brown spots. Endites, labium and chelicerae dark brown. Legs I and II with femur, patella and tibia predominantly dark brown, with irregular reddish-brown spots, metatarsus and tarsus reddish brown. Legs III and IV lighter, reddish brown with irregular dark brown patches and light brown patella. Opisthosoma dorsally dirty yellow with irregular black spots and a black transversal stripe between the paired conical projections. Two pairs of rounded reddish brown sigilla in the center of the dorsum of the opisthosoma. Projections dark brown with reddish brown hues at the sides. Opisthosoma ventrally and laterally almost black, with reddish-brown sigils.

Variation. Females may present opisthosoma predominantly light yellow with a single trapezoidal dark brown area on dorsum and some lighter irregular spots ( Fig. 11G ).

Distribution. Most records of E. villosus come from a wide range of the Brazilian Atlantic Forest, from southern-central Bahia to the north, to Rio Grande do Sul (and also north Uruguay) to the south, reaching northern Argentina and Paraguay to the west ( Fig. 15 ). Most localities of E. villosus are located in inland areas, especially in the northern part of their distribution, but there are coastal records from the states of São Paulo to Rio Grande do Sul, both in Brazil. The two isolated records of E. villosus from the Amazon are considered doubtful, pending additional material. The record from Santarém, Pará, is represented by a male and a female specimens collected at the 19th century, belonging to the Simon Collection (MNHN), and may represent a misplaced locality, as extensive collections by MPEG in that area produced only specimens of E. biocellatus . The other record is for Ilha de Maracá, Roraima, consisting of a single female, deposited in INPA. As the material of Ilha de Maracá was sorted and identified at Rio Grande do Sul, an area with abundant records of E. villosus , there is also the possibility of label exchange and mixing of localities.

Figures

FIGURE 11. Epicadinus villosus Mello-Leitão, 1929, female (FZBRS 39201). A–C habitus (A dorsal, B ventral, C lateral); D, F epigynum ventral; E vulva dorsal; G female with colour variation (FZBRS 39201). Abbreviations, A, atrium; CD, copulatory duct; MS, median septum; TK, translucent keel. Scale bars: A–C, G, 1 mm; D–F, 0.1 mm.

FIGURE 12. Epicadinus villosus Mello-Leitão, 1929, male (MACN 19052). A–C habitus (A dorsal, B lateral, C ventral); D–F left male palpus (D–E ventral, F retrolateral). Abbreviations, DTA, distal tibial apophysis; E, embolus; RTA, retrolateral tibial apophysis; TF, tegular furrow. Scale bars: A–C, 1 mm; D–F, 0.1 mm.

FIGURE 13. A–C Epicadinus villosus Mello-Leitão, 1929, female holotype (MNHN AR-15078 [Coll. Simon 11.511]) [total length: 7 mm] (A dorsal habitus, B ventral habitus, C original labels). D–F Epicadinus helenae subadult female holotype (IBSP 138.217 [ex. MZLQ 117]) (D dorsal habitus, E ventral habitus, F type labels). G–I Epicadinus marmoratus young holotype (MNHCI 02496 [Coll. Rudolf B. Lange]) (G dorsal habitus, H ventral habitus, I type label). Scale bars: D,E,G,H, 1 mm. Photo credits: A–C, Christine Rollard and MNHN Laboratory of Arachnology crew.

FIGURE 15. Geographic distribution records of Epicadinus, highlighting the type-locality of each of the four species (star symbols).