Guérin-Méneville, 1829 : 6 Eripus heterogaster (Guérin-Méneville) Walckenaer 1833 : 438 White 1841 : 476 C. L. Koch 1845 : 46 Keyserling 1880 : 164 Göldi 1886 : 411 O. Pickard-Cambridge, 1877 : 557 Keyserling, 1880 : 164 E. quinquegibbosus , O. Pickard-Cambridge Epicadus heterogaster (Guérin-Méneville) Simon 1895 : 1052 F. O. Pickard-Cambridge 1900 : 160 Vellard 1924 : 9 Mello-Leitão 1929 : 94 Roewer 1954 : 755 Brignoli 1983 : 604 Piza, 1933 : 50 Taxonomic revision of the crab spider genus Epicadus Simon, 1895 (Arachnida: Araneae: Thomisidae) with notes on related genera of Stephanopinae Simon, 1895 Silva-Moreira, Thiago Da Machado, Miguel Zootaxa 2016 4147 3 281 310 6FRDT Guerin-Meneville, 1829 Guerin-Meneville 1829 [151,741,1367,1393] Arachnida Thomisidae Epicadus Animalia Araneae 4 285 Arthropoda species heterogaster     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: 94pl 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. Heterogasteris an adjective.   Type material: Holotypeof  Thomisus heterogaster(designated by Simon 1895): female, BRAZIL, Rio de Janeiro, [-22.9; -43.233333], M. Latreille leg. (BMNH,image examined).   FIGURE 2.  Epicadus heterogaster(Guérin-Menéville, 1829). Female (IBSP 06247). A–C habitus (A dorsal, B lateral, C ventral); D cephalic region, frontal view; E femur I, dorsal view. Scale bars: 2 mm (A–C); 1 mm (D–E).  Holotypeof  E. quinquegibbosus(designated by Pickard Cambridge 1877): male, BRAZIL, Minas Gerais [- 19.9167; -43.9333], H. Rogers leg. 1871 (BMNH, not located).  Holotypeof  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 typesof  E. heterogasterand  E. quinquegibbosuscould not be directly examined by us. However, J. Beccaloni ( BMNH) kindly photographed the typespecimen of  E. heterogasterand 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 typeof  E. quinquegibbosuswas 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 typecollection of ESALQwas acquired by IBSP, but some of the ESALQ typeswere too degraded or were never found (A. Brescovit, pers. comm.), including the typeof  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 typelocalities without discrete differences, we feel very confident in establishing the synonymy (in agreement with Mello-Leitão´s view).   FIGURE 3.  Epicadus heterogaster(Guérin-Menéville, 1829). Female (IBSP 08491). A–D epigynum/vulva (A ventral–arrows point the atrium, B dorsal–arrows points tto FD, C sagittal, D frontal—arrow points to FD (left one broken)); E metatarsus I, retrolateral view. Arrows point to bifid setae. Scale bars: 0.5 mm (A, E); 0.2 mm (B–D).   FIGURE 4.  Epicadus heterogaster(Guérin-Menéville, 1829). Male (MZSP 07721). A cephalic region, frontal view; B, D–E habitus (B lateral, D dorsal, E ventral); C opisthosoma, caudal view. Scale bars: 0.25 mm (A), 0.5 mm (C); 1 mm (B, D, E).  Other material examined. COSTA RICA:   Cartago: 1 female, Turialba, [9.9, -83.6833], O. Brenes-Gamezleg. (InBIO).    Heredia: 1 female, San Ramón de Sarapiquí, [9.4167, -83.7333], M. Monteroleg.,  6 November 2006(InBIO).    Puntarenas: 1 female, Estac. Agujas, [8.32, -83.25], A. Azofeileg.,  13 November 1999(InBIO). PANAMA: Canal Zone:  1 female, Barro Colorado Island, [9.16361, -79.8377], A.M. Chickeringleg.,  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], Zeterleg.,  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. Paulsleg.,  5 February 1979( MCN08961).   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. Schnebleleg.,  25 February 1975( MCZ).    Meta: 1 female, Lomalinda( Puerto Lleras), [3.3038, -73.3615], E.T. Carrolland B. Rothleg., 1989, ( CAS).   ECUADOR:  Orellana:  1 male, Francisco de Orellana( Naporiver) [-0.4486, -76.9939], Boidonleg.,  January 1972(  MACN17288).    Napo:  1 female, Cabañas Aliñahui, 25kmeast of Puerto Napo, [-1.05, - 77.7833],  450 malt., E.S. Rossleg,  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], Willnerleg.,  May 1976, (  MACN17286).   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. Weyrauchleg.,  December 1946(  AMNH41).  Madre de Dios:  1 subadultfemale, 35 kmEast of Puerto Maldonadoon rio Madre de Dios, Albergue Lodge, [- 12.6, -69.1833], G.C. Hunterleg.,  22 June 1983( CAS).   BRAZIL:  Pará: 1 female, Fazenda Jaboti, [-2.1332, - 47.5603], T. Pimentelleg.,  17 February 1972(  MPEG003387); 1 female, Belém (Maracangalha), [-1.45, -48.4833], R.F. Silva leg., 12 April 1984(MPEG 003386).    Rondônia: 1 femaleand 2 juveniles, Ouro Preto do Oeste, [- 10.6666, -62.3], J. Beckerleg.,  22 October 1986(  MNRJ11462).    Roraima: 1 male, Alto Alegre( Ilhade Maracá), [2.0438, -50.4504], A.B. Bonaldoleg.,  14 February 1992(  MCTP1772).     Bahia: 1 female, Lençóis, [-12.5667, - 41.3833],  30 April 1999(  IBSP26842).    Goiás:  1 subadultfemale, Niquelândia(formerly São José do Tocantins), [- 14.4738,-48.46], F. Laneleg.,  22 June 1942(  MZSP171); 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(  IBSP3787); 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 Tomédas Letras, [-21.7167, -44.9833], F. Frieiro-Costaleg.,  3 October 1992(  MNRJ11500); 1 female, V.R. Souza leg. (MNRJ 1659).    Espírito Santo: 1 female, Apiacá( Fazenda Santa Maria), [-21.1536, -41.5675], R.L.C. Baptistaleg.,  22 July 1985(  MNRJ4239); 1 female, Conceição da Barra, [-18.3582, -39.8455], Ruschi leg. (MLP 17896).    Mato Grosso do Sul: 1 subadultfemale, Selvíria, Fazenda Canaã, [-20.3669, -51.4188], F. Laneleg.,  April 1967(  MZSP13576).    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. Barbosaleg.,  7 August 1998(  MNRJ02699);  1 female, Rio de Janeiro, [-22.9, -43.2333] (  MLP909); 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. Giuponnileg.,  30 October 1995(  MNRJ11550).  São Paulo:  1 female, São Paulo( Campus IB USP), [-23.5653, -46.7324], S.A. Hogeleg.,  October 1997(  IBSP14546); 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 (  IBSP20206); 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(  IBSP56827); 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 maleand 1 female, Ilhado Cardoso, [-25.0167, -47.95], E.S.A. Marquesleg.,  5 April 1988(  IBSP6247); 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(  IBSP00359); 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. Imaguireleg.,  17 August 1947(  MZSP7850.;    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( MLP17899).  BOLIVIA:   Santa Cruz: 1 female, Buena Vista, [-17.45, -63.6667], A. Prosenleg.,  October 1949( MLP17898). PARAGUAY:   Distrito Capital: 2 males,  Assunción, [-25.2666, -57.6667] ( MCZ).   Concepción: 1 female, Agüerito, [-22.8991, -56.6673], Willnerleg.,  November 1966(  MACN17287).   ARGENTINA:  Missiones: 1 subadultfemale, Piñalitos, [-25.9822, - 53.9000], M.E. Galianoleg.,  January 1976(  MACN17289).   Diagnosis.Both males and females of  E. heterogasterhave a distinctive white medial spot and a well pronounced MS on the dorsal shield of the prosoma ( Figs. 2A, D; 4D). The female opisthosoma projections are vertically oriented ( Figs. 2B). Left and right CD run in parallel, clustered. CV positioned sideways, curving outward ( Figs. 3B; 15A). Male RTA flattened, with only its dorsal margin slightly folded; DTA is projected upwards, in parallel with the RTA ( Figs. 5A, 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;  COSTARICA: 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. heterogasterhas 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 heterogasterexhibits a high degree of polychromy in vivo( Figs. 1A–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. heterogasteris usually found on vegetation with flowers, where the spiders hunt as sit and wait predators ( Figure 16A–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. heterogasterexhibit 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. heterogastermove very slowly on leaves, in a fashion similar to a flower blown by the wind. All these data suggest that  E. heterogasteris 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. heterogasterplays an important role on foraging. Little is known about their mating behavior. Observations based on photos show that adult males of  E. heterogasterare found seated on the back of females ( Fig. 16B), 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. 16D). 1290989952 [714,816,1072,1097] Turialba & O. Brenes-Gamez Cartago 7 288 1 1 Cartago 1290989947 2006-11-06 San Ramon de Sarapiqui & M. Montero Heredia 7 288 1 1 Heredia 1290989941 [261,1432,1143,1168] 1999-11-13 Estac. Agujas & A. Azofei Puntarenas 7 288 1 1 Puntarenas 1290989943 1934-06-16 MCZ A. M. Chickering Iceland Colorado 7 288 1 1 Colorado 1290989936 1940-01 MCZ El Cermeno & Zeter Venezuela Bolivar 7 288 1 1 Bolivar 1290989955 1979-02-05 MCN La Esmeralda & S. Pauls Venezuela Amazonas 7 288 1 1 Amazonas 1290989939 [779,914,1357,1380] Venezuela Portuguesa 7 288 1 Portuguesa 1290989933 Santa Maria Colombia Boyaca 7 288 1 1 Boyaca 1290989942 1975-02-25 MCZ Parque Nacional Natural Chingaza & P. A. Schneble Colombia Cundinamarca 7 288 1 1 Cundinamarca 1290989946 CAS Lomalinda & Puerto Lleras & E. T. Carroll & B. Roth Colombia Meta 7 288 1 1 Meta 1290989949 [361,621,1498,1523] Ecuador Orellana 7 288 1 Orellana 1290989940 1972-01 MACN Boidon Ecuador Orellana 7 288 MACN 17288 1 1 Orellana 1290989951 [581,643,1534,1557] Ecuador Napo 7 288 1 Napo 1290989948 1994-02 CAS Cabanas Alinahui & E. S. Ross Ecuador 450 Napo 7 288 1 1 Napo 1290989973 1976-05 MACN Pichilingue & Willner Ecuador Los Rios 7 288 MACN 17286 1 1 Los Rios 1290989944 [472,673,1640,1665] Peru Huanuco 7 288 1 Huanuco 1290989945 1946-12 AMNH W. Weyrauch Peru Madre de Dios 7 288 AMNH 41 1 1 Madre de Dios 1290989979 1983-06-22 CAS Albergue Lodge & G. C. Hunter Peru Puerto Maldonado 7 288 1 1 Madre de Dios 1290989972 1972-02-17 MPEG Fazenda Jaboti & T. Pimentel Brazil Para 7 288 MPEG 003387 1 1 Para 1290989968 1986-10-22 MNRJ Ouro Preto do Oeste & J. Becker Brazil Rondonia 7 288 MNRJ 11462 3 1 2 Rondonia 1290989963 1992-02-14 MCTP Alto Alegre & A. B. Bonaldo Iceland Roraima 7 288 MCTP 1772 1 1 Roraima 1290989975 1999-04-30 IBSP Lencois Iceland Bahia 7 288 IBSP 26842 1 1 Bahia 1290989981 [596,663,1959,1984] Iceland Goias 7 288 1 Goias 1290989970 1942-06-22 MZSP Niquelandia & F. Lane Iceland Tocantins 7 288 MZSP 171 1 1 Tocantins 1290989950 [515,1411,151,176] 1984-03 IBSP Arcos Iceland Minas Gerais 8 289 IBSP 3787 1 1 Minas Gerais 1290989954 1992-10-03 MNRJ F. Frieiro-Costa Sao Tome and Principe Sao Tome 8 289 MNRJ 11500 1 1 1290989969 1985-07-22 MNRJ Apiaca & Fazenda Santa Maria & R. L. C. Baptista Sao Tome and Principe Espirito Santo 8 289 MNRJ 4239 1 1 Espirito Santo 1290989957 1967-04 MZSP Selviria & Fazenda Canaa & F. Lane Sao Tome and Principe Mato Grosso do Sul 8 289 MZSP 13576 1 1 Mato Grosso do Sul 1290989971 [1145,1315,471,496] Sao Tome and Principe Rio de Janeiro 8 289 1 Rio de Janeiro 1290989960 1998-08-07 MNRJ PN Tijuca & Estrada da Vista Chinesa & L. F. Ferreira & A. F. Barbosa Sao Tome and Principe Rio de Janeiro 8 289 MNRJ 02699 1 1 Rio de Janeiro 1290989961 [664,1280,541,566] MLP Sao Tome and Principe Rio de Janeiro 8 289 MLP 909 1 1 Rio de Janeiro 1290989934 1995-10-30 MNRJ Mangaratiba & A. P. L. Giuponni Sao Tome and Principe Sao Paulo 8 289 MNRJ 11550 1 1 Sao Paulo 1290989962 [236,1432,861,886] 1997-10 USP, IBSP Campus IB & S. A. Hoge Sao Tome and Principe Sao Paulo 8 289 IBSP 14546 1 1 Sao Paulo 1290989937 IBSP Salesopolis & Parque Nascentes do Rio Tiete & R. Martins Sao Tome and Principe Rio Tiete 8 289 IBSP 20206 1 1 Sao Paulo 1290989976 2005-06-02 SUCEN, IBSP Saint Vincent and the Grenadines Sao Vicente 8 289 IBSP 56827 1 1 1290989938 1988-04-05 IBSP E. S. A. Marques Iceland Ilha 8 289 IBSP 6247 2 1 1 1290989993 1935-10 IBSP Iceland Sao Paulo 8 289 IBSP 00359 1 1 Sao Paulo 1290989967 1947-08-17 MZSP Ponta Grossa & Parque Estadual Vila Velha & K. Imaguire Iceland Parana 8 289 MZSP 7850 1 1 Parana 1290989964 [736,1436,1996,2021] Blumenau Iceland Santa Catarina 8 289 1 1 Santa Catarina 1290989974 [532,1170,187,212] MLP Bolivia Rio Grande do Sul 9 290 1 1 Rio Grande do Sul 1290989956 1949-10 MLP Buena Vista & A. Prosen Bolivia Santa Cruz 9 290 1 1 Santa Cruz 1290989935 [1222,1412,222,247] Bolivia Distrito Capital 9 290 2 2 Distrito Capital 1290989966 [233,696,258,283] MCZ Bolivia Assuncion 9 290 1 Assuncion 1290989953 1966-11 MACN Aguerito & Willner Bolivia Concepcion 9 290 MACN 17287 1 1 Concepcion 1290989986 1976-01 MACN Pinalitos & M. E. Galiano Argentina Missiones 9 290 MACN 17289 1 1