Epicadus heterogaster Guerin-Meneville 1829
Creators: name: Silva-Moreira, Thiago Da
name: Machado, Miguel
Publisher: name: Zenodo
Year: 2016
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Arachnida
Araneae
Thomisidae
Epicadus
Epicadus heterogaster

description: Epicadus heterogaster (Guérin-Méneville, 1829)
Figs 1 A–C; 2–6; 15A–C, 16A–D, 18
Thomisus heterogaster Guérin-Méneville, 1829: 6, pl. 1, fig. 4.
Eripus heterogaster (Guérin-Méneville). Walckenaer 1833: 438, 441; 1837: 541, pl. 12, fig. 6; White 1841: 476; C. L. Koch 1845: 46, figs 984–985; Keyserling 1880: 164, pl. 3, fig. 90; Göldi 1886: 411, pl. 11, figs. 1–7.
Eripus quinquegibbosus O. Pickard-Cambridge, 1877: 557, pl. 56, fig. 5.
Eripus quinquestriatus Keyserling, 1880: 164, pl. 3, fig. 91 [= E. quinquegibbosus, O. Pickard-Cambridge, lapsus calami].
Epicadus heterogaster (Guérin-Méneville). Simon 1895: 1052, fig 1085; F. O. Pickard-Cambridge 1900: 160, pl. 10, fig 34; Vellard 1924: 9, fig 4; Mello-Leitão 1929: 94 pl 31, fig 44; Roewer 1954: 755; Brignoli 1983: 604; World Spider Catalog 2015.
Epicadus heterogaster scholagriculae Piza, 1933: 50, fig 3. New synonymy.
Etymology. “ Heterogaster ” is the combination of Greek words heteros [ἕτερος,] “different” and gaster [γαστήρ] “abdomen” as an allusion to the unique ophistosoma shape of the species. Heterogaster is an adjective.
Type material: Holotype of Thomisus heterogaster (designated by Simon 1895): female, BRAZIL, Rio de Janeiro, [-22.9; -43.233333], M. Latreille leg. (BMNH,image examined).
Holotype of E. quinquegibbosus (designated by Pickard Cambridge 1877): male, BRAZIL, Minas Gerais [- 19.9167; -43.9333], H. Rogers leg. 1871 (BMNH, not located).
Holotype of E.h. scholagriculae (designated by Piza, 1933): female, BRAZIL, São Paulo, Piracicaba, [- 22.7167; -47.6333]], S. de T. Piza leg. (IBSP [formerly ESALQ-MZLQ A0120], not located, lost).
Note. Both the types of E. heterogaster and E. quinquegibbosus could not be directly examined by us. However, J. Beccaloni (BMNH) kindly photographed the type specimen of E. heterogaster and sent the images to us for study. Despite the poor conditions of the type, the photos were sufficient to locate similar specimens and to proceed with the morphological analysis. She also stated (in litt.) that the type of E. quinquegibbosus was not found among the BMNHcollections and that it is probably lost. Nevertheless we still believe that the synonym proposed by Göldi (1886) is accurate. When comparing the illustration of the original description with our specimens, we found no indications for specific differences. Moreover, several of the studied samples consisted of both males and females found and captured together, with adds even more strength to this hypothesis.
The type collection of ESALQ was acquired by IBSP, but some of the ESALQ types were too degraded or were never found (A. Brescovit, pers. comm.), including the type of E. h. scholagriculae. We located among the research papers and personal belongings of C. F. Mello Leitão (currently belonging to the Arachnology lab at MNRJ) the original photograph that illustrated the original description, which helped to locate similar specimens from the same region. We found the diagnostic characters on which the original description relied to be highly variable. The author itself (p 54) stated that he was not able to access more specimens in addition to the holotype, and he decided to nominate it a subspecies (instead of a nominal species) because Mello-Leitão (in litt.) suggested that the specimen was simply a variation of E. heterogaster. Given 1) the impossibility to examine the holotype, 2) the lack of strong diagnostic characters in the original descriptions, and 3) the abundant material from both the type localities without discrete differences, we feel very confident in establishing the synonymy (in agreement with Mello-Leitão´s view).
Other material examined. COSTA RICA: Cartago: 1 female, Turialba, [9.9, -83.6833], O. Brenes-Gamez leg. (InBIO). Heredia: 1 female, San Ramón de Sarapiquí, [9.4167, -83.7333], M. Montero leg., 6 November 2006 (InBIO). Puntarenas: 1 female, Estac. Agujas, [8.32, -83.25], A. Azofei leg., 13 November 1999 (InBIO). PANAMA: Canal Zone: 1 female, Barro Colorado Island, [9.16361, -79.8377], A.M. Chickering leg., 16 June 1934 (MCZ); 15 males and 1 female, Pipeline road, [9.3333, -82.25], E.S. Ross leg., 2 October 1976 (CAS). Arraiján: 1 male and 1 female, Cerro Galera, [8.93333; -79.6166], M. Robinson leg., 20 May 1976 (MCZ); 1 female, El Cermeño, [8.9344, -79.6167], Zeter leg., January 1940 (MCZ). VENEZUELA: Bolívar: 1 female, [8.1222; -63.5497], S. Pauls leg., 8 February 1980 (MCN 11422). Amazonas: 1 female, La Esmeralda, [3.1667; - 65.55], S. Pauls leg., 5 February 1979 (MCN 08961). Portuguesa: 1female; Guanare [9.0436, -69.7488], Cesar leg., 15 October 1975 (MACN 17285). COLOMBIA: Boyacá: 1 female, Santa Maria, [4.8617, -73.2642], 1994 (ICN-AR). Cundinamarca: 1 male, Parque Nacional Natural Chingaza, [4.5667, -73.75], P.A. Schneble leg., 25 February 1975 (MCZ). Meta: 1 female, Lomalinda (Puerto Lleras), [3.3038, -73.3615], E.T. Carroll and B. Roth leg., 1989, (CAS). ECUADOR: Orellana: 1 male, Francisco de Orellana (Napo river) [-0.4486, -76.9939], Boidon leg., January 1972 (MACN 17288). Napo: 1 female, Cabañas Aliñahui, 25km east of Puerto Napo, [-1.05, - 77.7833], 450 m alt., E.S. Ross leg, February 1994 (CAS); 1 female, E.S. Ross leg., December 1995 (CAS); 1 female, E.S. Ross leg., January 1991 (CAS). Los Ríos: 1 female, Pichilingüe, [-1.0789, -79.4922], Willner leg., May 1976, (MACN 17286). PERU: Huánuco: 1 subadult female, Tingo Maria, [-9.2952, -75.9975], E.I. Schilling & E.S. Ross leg., 10 November 1954 (CAS); 1 female, E.I. Schilling & E.S. Ross leg., 15 October 1954 (CAS); 1 male, E.I. Schilling & E.S. Ross leg., 5 May 1954 (CAS); 1 female, (W. Weyrauch leg., December 1946 (AMNH 41). Madre de Dios: 1 subadult female, 35 km East of Puerto Maldonado on rio Madre de Dios, Albergue Lodge, [- 12.6, -69.1833], G.C. Hunter leg., 22 June 1983 (CAS). BRAZIL: Pará: 1 female, Fazenda Jaboti, [-2.1332, - 47.5603], T. Pimentel leg., 17 February 1972 (MPEG 003387); 1 female, Belém (Maracangalha), [-1.45, -48.4833], R.F. Silva leg., 12 April 1984 (MPEG 003386). Rondônia: 1 female and 2 juveniles, Ouro Preto do Oeste, [- 10.6666, -62.3], J. Becker leg., 22 October 1986 (MNRJ 11462). Roraima: 1 male, Alto Alegre (Ilha de Maracá), [2.0438, -50.4504], A.B. Bonaldo leg., 14 February 1992 (MCTP 1772). Bahia: 1 female, Lençóis, [-12.5667, - 41.3833], 30 April 1999 (IBSP 26842). Goiás: 1 subadult female, Niquelândia (formerly São José do Tocantins), [- 14.4738,-48.46], F. Lane leg., 22 June 1942 (MZSP 171); 1 subadult female, Pirenópolis, [-15.85, -48.95], F. Lane leg., 20 June 1942 (MZSP 196). Minas Gerais: 1 female, Arcos, [-20.2833, -45.5333], March 1984 (IBSP 3787); 1 female, Uberlândia (Reserva de Caça e Pesca), [-18.9138, -48.2747], D. Cunha leg., October 1996 (IBSP 8303); 1 female, Barbacena, [-21.2258, -43.7902], J.M. Xavier leg., December 1964 (IBSP 01904); 1 subadult female, Carmo da Cachoeira, [-21.4667, -45.2167], J. Netto leg., 22 April 1948 (MZSP 13578); 1 female, Curvelo, [-18.75, -44.4167], A. Raimundo leg., September 1960 (IBSP 1597); 1 female and 150 juveniles, Paraopeba, [-19.3, - 44.4166], L.V.R. Souza leg. (MNRJ 13928); 1 female, São Tomé das Letras, [-21.7167, -44.9833], F. Frieiro-Costa leg., 3 October 1992 (MNRJ 11500); 1 female, V.R. Souza leg. (MNRJ 1659). Espírito Santo: 1 female, Apiacá (Fazenda Santa Maria), [-21.1536, -41.5675], R.L.C. Baptista leg., 22 July 1985 (MNRJ 4239); 1 female, Conceição da Barra, [-18.3582, -39.8455], Ruschi leg. (MLP 17896). Mato Grosso do Sul: 1 subadult female, Selvíria, Fazenda Canaã, [-20.3669, -51.4188], F. Lane leg., April 1967 (MZSP 13576). Rio de Janeiro: 1 female, Rio de Janeiro (Rio de Janeiro, PN Tijuca, Estrada da Vista Chinesa), [-22.95, -43.2833], L.F. Ferreira & A.F. Barbosa leg., 7 August 1998 (MNRJ 02699); 1 female, Rio de Janeiro, [-22.9, -43.2333] (MLP 909); 1 female, M. Zilkar leg. (MLP 910); 1 male and 1 female, C.F. Mello-Leitão leg. (MNRJ 00909); 1 female (MNRJ 02700); 1 female, C. Seigneur leg., 11 December 1984 (MNRJ 10382); 1 female, Itatiaia (Lagoa Azul), [-22.4961, -44.5633], E. Izeckson leg., 19 September 1956 (UBTU 02802); 1 female, Itatiaia, [-22.9, -43.2333] (MLP 00908); 1 female, Silva Jardim (Poço das Antas), [-22.5041, -42.2704], F.F.F. Ferraz leg., 24 Feburary 1995 (IBSP 27950); 1 female, Macaé (Restinga de Jurubatiba), [-22.5167, -42.2833], D.S.D. Araujo leg., 12 January 1985 (MNRJ 10397); 1 female, Mendes, [-22.5261, -43.7247], 1933, (IBSP 358); 1 subadult female, 18 December 1944 (MZSP 03546); 1 female, Resende (Serrinha do Alambari), [-22.3731, -44.5387], U. Caramaschi leg., 22 November 2006 (MNRJ 13927); 1 female, Mangaratiba, [-22.95, -44.0333], A.P.L. Giuponni leg., 30 October 1995 (MNRJ 11550). São Paulo: 1 female, São Paulo (Campus IB USP), [-23.5653, -46.7324], S.A. Hoge leg., October 1997 (IBSP 14546); 1 female, Iporanga (Fazenda Intervales), [-24.5833, -48.5833], A. Leme leg., 28 May 1996 (IBSP 8103); 1 female, Rodovia Raposo Tavares (Km 50), [-23.5333, -47.1333], I. Grant leg., 23 March 1979 (IBSP 2916); 1 female, Salesópolis (Parque Nascentes do Rio Tietê), [-23.6555, -45.76805], R. Martins, 1997 (IBSP 20206); 1 female, Jundiaí (Serra do Japi), [-22.2808, -47.5483], L.P.C. Morellato leg., 30 June 1990 (IBSP 6243); 1 male, E.N. Ramires leg., 12 April 1991 (IBSP 6248); 1 male and 1 female, E.N. Ramires leg., 14 May 1990 (IBSP 6251); 1 female, São Vicente (SUCEN), [-23.9630, -46.3919], 2 June 2005 (IBSP 56827); 1 female, Araçariguama, [- 23.4333, -47.0667], M.V. Gonçalves leg., 6 January 2003 (IBSP 37220); 1 female, Araçoiaba da Serra, [-23.5, - 47.6167], F.E. Sprenges leg., 10 February 2005 (IBSP 56977); 1 female, Bom Jesus dos Perdões, [-23.1333, - 46.4667], C.I. Kawoamoto leg., 1990 (IBSP 4451); 1 female, Corumbataí, [-22.22, -47.6258], S. Burian leg., 1935 (IBSP 357); 1 female, Cotia, [-23.6166, -46.9333], B. Peruzzo leg., 11 January 1996 (IBSP 8494); 1 female, D. Bibeneus leg., 21 January 2000 (IBSP 27563); 1 female, Embu das Artes, [-23.6488, -46.8522], 17 Ocotober 2005 (IBSP 58202); 1 female, Embú Guaçu, [-23.8167, -46.8], C. Casoy leg., 7 January 2002 (IBSP 35003); 1 male and 1 female, Ilha do Cardoso, [-25.0167, -47.95], E.S.A. Marques leg., 5 April 1988 (IBSP 6247); 1 female, Itapecerica da Serra, [-23.7166, -46.8333], A. Calmon leg., September 1959 (IBSP 1450); 1 female, L. Richtzenhein leg., 24 October 1995 (IBSP 14301); 1 female, (K. Ogawa leg., 2 November 1997 (IBSP 14474); 1 female, Itapetininga, [-23.6, -48.05], Z.R. Borges leg., October 1959 (IBSP 1464); 1 female, Jacupiranga, [- 24.6986, -48.0038], F. Mathiesen leg., 16 October 1985 (IBSP 42360); 1 female, Jundiaí, [-23.1833, -46.8667], P. Ganiparauskas leg., September 1974 (IBSP 3302); 1 female, Juquitiba, [-23.9272, -47.07], E.B. Daniel leg., 29 January 1996 (IBSP 6786); 1 female, C.A. Andalfatto leg., 1 September 2003 (IBSP 41990); 1 female, Mariporã, [- 23.3188, -46.5869], V.Q. Andrade leg., 9 February 1998 (IBSP 16238); 1 female, Osasco, [-23.5333, -46.7666], F.R. Ortiz leg., 24 October 2003 (IBSP 41040); 1 female, Salto de Itu, [-23.2, -47.2833] (IBSP 1466); 1 female, São Lourenço da Serra, [-23.8667, -46.95], C.E.M. Machado leg., 26 October 1998 (IBSP 20437); 1 female, São Paulo, [-23.8667, -46.95], October 1935 (IBSP 00359); 1 female, O. Franca leg., 18 October 1935 (IBSP 360); 1 female, 27 March 1952 (IBSP 694); 1 female, J.R. Pereira leg., 19 May 1952 (IBSP 708); 1 female, J. Carrari leg., March 1953 (IBSP 886); 1 female, T.I. Miller leg., 1950 (IBSP 1054); 1 female, B. Ditizius leg. (IBSP 1220); 1 female, M. Nogueira leg., July 1959 (IBSP 1430); 1 female, R. Arrivabene leg., October 1985 (IBSP 4356); 1 female, I. Ferreira leg., July 1989 (IBSP 4454); 1 female, B. Fritz leg., October 1985 (IBSP 4977); 1 female, M. Quesada leg., 8 May 1996 (IBSP 14069); 1 female, A.F. do Amaral leg., 6 November 1995 (IBSP 14300); 1 female, A.A. de Santis leg., 7 June 2001 (IBSP 28455); 1 female, São Roque, [-23.5333, -47.1333], A. Camargo leg., March 1988 (MNRJ 4238); 1 female, Vargem Grande Paulista, [-23.6027, -47.0258], J.M. de Oliveira leg., 23 January 2003 (IBSP 37209). Paraná: 1 female, Ponta Grossa (Parque Estadual Vila Velha), [-25.2166, -50.0333], K. Imaguire leg., 17 August 1947 (MZSP 7850.; Santa Catarina: 1 female, Blumenau, [-26.9333, -49.05], J.
Lange leg., March 1984 (IBSP 3792); 1 female, Santo Amaro da Imperatriz, [-27.6833, -48.7666], A.A. Lise leg., 30 October 1986, (MCN 16032). Rio Grande do Sul: 1 female (MLP 17899). BOLIVIA: Santa Cruz: 1 female, Buena Vista, [-17.45, -63.6667], A. Prosen leg., October 1949 (MLP 17898). PARAGUAY: Distrito Capital: 2 males, Assunción, [-25.2666, -57.6667] (MCZ). Concepción: 1 female, Agüerito, [-22.8991, -56.6673], Willner leg., November 1966 (MACN 17287). ARGENTINA: Missiones: 1 subadult female, Piñalitos, [-25.9822, - 53.9000], M.E. Galiano leg., January 1976 (MACN 17289).
Diagnosis. Both males and females of E. heterogaster have a distinctive white medial spot and a well pronounced MS on the dorsal shield of the prosoma (Figs. 2 A, D; 4D). The female opisthosoma projections are vertically oriented (Figs. 2 B). Left and right CD run in parallel, clustered. CV positioned sideways, curving outward (Figs. 3 B; 15A). Male RTA flattened, with only its dorsal margin slightly folded; DTA is projected upwards, in parallel with the RTA (Figs. 5 A, 5B, E). Tegulum discoid with tegular furrow at eight o’clock.
Description. Female (MNRJ 13927): Total length 15.33; prosoma width 6.20; opisthosoma width 8.73; prosoma height without MS 4.43; opisthosoma height without projection 6.47. Leg measurements: femur I 5.67; patella I 2.67; tibia I 3.20; metatarsus I 2.13; tarsus I 1.67; femur II 5.40; patella II 2.37; tibia II 3.33; metatarsus II 2.33; tarsus II 1.67; femur III 2.87; patella III 1.43; tibia III 2.0; metatarsus III 1.53; tarsus III 1.30; femur IV 3.67; patella IV 1.43; tibia IV 2.27; metatarsus IV 1.90; tarsus IV 1.27. Leg spination: tibia I 0-2-2; metatarsus I 2 -0-2- 2ap; tibia II 0-2-2; metatarsus II 2 -0-2-2ap. Epigynum and genital features as in Fig. 2
Male (IBSP 6251): Total length 3.7 prosoma width: 1.78; opisthosoma width 1.85; prosoma height without MS 1.09; opisthosoma height without projections 2.11. Leg measurements: femur I 1.51mm; patella I 0.62; tibia I 0.91; metatarsus I 0.64; tarsus I 0.55; femur II 1.62; patella II 0.58; tibia II 0.95; metatarsus II 0.71; tarsus II 0.48; femur III 0.88; patella III 0.34; tibia III 0.50; metatarsus III 0.53; tarsus III 0.42; femur IV 0.96; patella IV 0.35; tibia IV 0.59; metatarsus IV 0.45; tarsus IV 0.35. Leg spination: tibia I 0-2-2; metatarsus I 2 -0-2-2ap; tibia II 0-2-2; metatarsus II 2 -0-2-2ap. Male palp as in Fig. 5.
Distribution. ARGENTINA: Misiones; BOLIVIA: Santa Cruz; BRAZIL: Bahia, Espírito Santo, Minas Gerais, Mato Grosso do Sul, Pará, Paraná, Rio de Janeiro, Rio Grande do Sul, Rondônia, Roraima, Santa Catarina, São Paulo; COLOMBIA: Cundinamarca, Meta; Boyacá; COSTA RICA: Cartago, Heredia, Puntarenas; ECUADOR: Napo, Orellana; PANAMA: Bocas Del Toro, Panama; PARAGUAY: Distrito Capital, Concepción; PERU: Huánuco, Madre de Dios; VENEZUELA: Amazonas, Bolívar, Portuguesa (Fig. 18).
N.B. While a specimen of E. heterogaster has been recorded from French Guiana (a female from Charvein, collected by Benoist. [sic.] in December 1913) by Caporiacco (1954: 128) this locality is not included here. Although the record seems to be reliable, as it fits in the herein updated distribution range, there is no hint to a registered voucher (with collection number) in the aforementioned publication and the specimen itself could not be examined. Therefore we believe the best practice would be not to include this species distribution record.
Natural history. Epicadus heterogaster exhibits a high degree of polychromy in vivo (Figs. 1 A–C). While the most common colors are white and yellow, other colors such as pink, purple or a mix of those were already registered (as seen in Fig. 1). The color variations do not seem to follow a geographical pattern, and they might be related to camouflage at their hunting sites. While it is likely that those spiders are capable of changing colors, as other thomisids do like Misumena vatia (Clerck, 1757) (see Gabritschevsky 1927, Oxford & Gillespie 1998), there is yet no study corroborating this hypothesis.
Romero & Vasconcellos-Neto (2007) and Vasconcellos-Neto & Ramires (2007) made a series of ecological and behavioral observations on E. heterogaster. Like many other thomisids, E. heterogaster is usually found on vegetation with flowers, where the spiders hunt as sit and wait predators (Figure 16 A–C). However, their site selection varies with vegetation shifts. While flowers are available (cold and dry season) individuals were found sitting on flowers, whereas during the rest of the year, they prefer the leaves, positioning themselves where flowers usually occur. The observed prey consisted mainly of insects with flower visiting behavior: lepidopterans (Ithominae), dipterans (Syrphidae) and hymenopterans (Vespidae, Apidae). Further investigations by Romero & Vasconcellos-Neto (2007) have shown that females of E. heterogaster exhibit similar UV reflectance levels as the leaves they were found on, but they do display strong fluorescence under UV light (337.1 nm) and an emission peak at blue light (450–500 nm), which is a range perceived by many flower-visiting insects. They also observed that E. heterogaster move very slowly on leaves, in a fashion similar to a flower blown by the wind. All these data suggest that E. heterogaster is a cryptic predator that targets flower-visiting insects and, while hunting on leaves, it acts as both a lure to insect prey (by aggressively mimicking flowers) and conceals itself from predators. In a more recent study, Peixoto et al. (2012) investigated the dynamics of hunting site selection in E. heterogaster, testing for a) body color match b) floral model and c) olfactory (chemical) stimuli. Those experiments result in negative responses for all tested stimuli, failing to isolate a specific trend, however the authors discarded chemical or visual (i.e. human perceived color spectrum) cues as influential factors on this this process. Bhaskara et al. (2009) suggested that UV cues might play a role on selection of the hunting sites, which might reinforce the idea that the UV reflection seen in E. heterogaster plays an important role on foraging.
Little is known about their mating behavior. Observations based on photos show that adult males of E. heterogaster are found seated on the back of females (Fig. 16 B), suggesting mate guarding and/or opportunistic mating behavior, as reported for other spiders (Elgar & Bathgate 1996; Fahey & Elgar 1997; Prenter et al. 2003; Bel-Venner & Venner 2006), including thomisids (Dodson & Beck 1993; Holdsworth & Morse 2000). Females were also seen guarding eggs sacs (Fig. 16 D).
description: Published as part of Silva-Moreira, Thiago Da & Machado, Miguel, 2016, Taxonomic revision of the crab spider genus Epicadus Simon, 1895 (Arachnida: Araneae: Thomisidae) with notes on related genera of Stephanopinae Simon, 1895, pp. 281-310 in Zootaxa 4147 (3) on pages 285-292, DOI: 10.11646/zootaxa.4147.3.4, http://zenodo.org/record/263154