Marma nigritarsis ( Simon, 1900 )

Salgado, Alexandre & Ruiz, Gustavo R. S., 2020, Taxonomic revision of Marma Simon, 1902 (Araneae: Salticidae: Euophryini), Zootaxa 4899 (1), pp. 287-353 : 324-334

Marma nigritarsis ( Simon, 1900)

Figures 1A, D , 34– 43 , 57E , 58G

Amphidraus nigritarsis Simon, 1900: 60 (Holotype ♁: VENEZUELA: Fracatal, Caroní [08°14’37.8”N, 62°47’33.4”W], deposited in MNHN—collection of L. de Bertier—examined by photographs); Simon (1901: 419, fig. 477).

Pseudoamphidraus variegatus Caporiacco, 1947: 30 (Holotype ♁: GUYANA: Esequibo, Two Mouths, leg. Romiti, 14.VII.1936, deposited in MZLS 597—examined); synonymized by Ruiz, Brescovit & Lise (2007: 380); Caporiacco (1948: 700, fig. 125).

Marma trifidocarinata Caporiacco, 1947: 34 (Holotype ♀: GUYANA: Alto Demerara, leg. Beccari, XII.1931, deposited in MZLS 549—examined); synonymized by Ruiz, Brescovit & Lise (2007: 380); Caporiacco (1948: 731, fig. 165).

Marma nigritarsis: Galiano, 1962: 40 , pl. I, figs 6–9, pl. II, figs 4–5; Galiano 1963: 397, pl. XXVIII, figs 1–2; Galiano 1981: 12; Ruiz, Brescovit & Lise 2007: 380; Zhang & Maddison 2015: 57, figs 215–220.

Note. The species Agelista rosea Mello-Leit „o, 1941, Thysema dorae Mello-Leit „o, 1944 and Paralophostica centralis Soares & Camargo, 1948 were synonymized with Marma nigritarsis by Galiano (1962). Ocnotelus argentinus Mello-Leit „o, 1941 was also synonymized with M. nigritarsis by Galiano (1981). These synonymies are rejected herein.

We concluded, by comparisons among the types of the previously synonymized species ( Figs 35 A–I, 45A–F, 48A–D, 49) with the material provided by MPEG, MACN, UFMG and IBSP, that the synonymies of Pseudoamphidraus variegatus Caporiacco, 1947 and Marma trifidocarinata Caporiacco, 1947 with M. nigritaris should be maintained. In addition, we concluded that M. rosea (Mello-Leit„o, 1941) and M. argentina (Mello-Leit„o, 1941) are morphologically different from M. nigritarsis , and therefore, these species are revalidated and redescribed.

We provide new diagnostic illustrations based on freshly collected material for the male ( Figs 37 A–E, G) and based on the holotype of Marma trifidocarinata for the female ( Figs 37 H–I).

Revised diagnosis. Among all Marma species, M. nigritarsis shares more similarities with M. pipa sp. nov. The males of both species have an embolic disc with similar width, with straight prolateral border and curved retrolateral border ( Figs 25C , 27A , 36C , 37A , 57C, 57E ). In the females, the copulatory ducts do not touch each other and encircle the primary spermathecae ( Figs 26 C–D, 27F–G, 35H–I, 37H–I, 58E, G). The males of M. nigritarsis differ from those of M. pipa sp. nov. by having a longer embolus and a longer process on the embolic disc (PED; Figs 57C, E ), and the females differ by having a homogenous diameter along the primary spermathecae (broad proximally and narrow distally in M. pipa sp. nov.; Figs 58E, G ).

Comparisons among the species revalidated in this study: Males of M. nigritarsis differ from those of M. rosea by having the embolic disc with a smaller process and its retrolateral edge with less pronounced curvature ( Figs 36C , 37A, C , 48C , 50A, C , 57E, H ). Also, males of M. nigritarsis have the prolateral border straight in the embolic disc ( Figs 36C , 37A , 57E ), whereas it has a more pronounced curvature in M. rosea ( Figs 48C , 50A , 57H ) and M. argentina ( Figs 44C , 46A , 57G ). Besides, the process of the embolic disc (PED), embolus and cymbium are smaller (proportionally when compared to the bulb) in M. nigritarsis than in M. argentina ( Figs 57E, G ). Females of M. nigritarsis differ by having the copulatory ducts far from each other ( Figs 35 H–I, 37H–I, 58G), whereas they are close (touching each other) in M. rosea ( Figs 49C, F , 50 F–G, 58J) and M. argentina ( Figs 45C, F , 46 C–D, 58I). In addition, the primary spermathecae of M. nigritarsis are approximately straight, whereas they are accentuated arched in M. argentina ( Figs 58G, I ).

Description. Male (MPEG 34357). Total length: 3.62. Carapace 2.00 long, 1.34 wide, 1.01 high. Ocular quadrangle 0.96 long. Anterior eye row 1.33 wide, posterior 1.11 wide. Legs 1432. Length of legs: I 4.06 (1.34 + 1.51 + 1.21); II 3.69 (1.19 + 1.33 + 1.17); III 4.01 (1.31 + 1.34 + 1.36); IV 4.02 (1.26 + 1.32 + 1.44).

Leg macrosetae: Femur I d1-1-1, p1di, r0; II d1-1-1-1 (or d1-1-1), p1di, r0-1-1 (or r1di); III–IV d1-1-1, p1di, r1di. Patella I–II 0, III–IV p0, r1. Tibia I p0-1-0, r0, v1r-2-2; II p0-1-0, r0, v1r-2-1p (or v1r-2-2); III p1-1-0, r1-1-0, v2-0-2; IV p0-1-1-0, r1-1-1-0 (or r0-1-1-0), v1p-0-2. Metatarsus I p1di, r0, v2-2; II p1-1, r1di, v2-2; III p1-0-2, r1- 0-2, v1r-0-2; IV p1-1-2, r1-1-2, v1p-0-2.

Color in alcohol ( Figs 36 A–B): carapace with triangle of setae pointing backwards; abdomen ventrally with three longitudinal dark stripes; legs: femur I–II with dark distal ring, III–IV with dark distal ring and proximal region with dark proventral spot; tibia I–IV with proximal and distal dark ring (less colorful on leg I); tarsus I almost black, II with dark tip, III–IV with dark proximal ring.

Palp: RTA finger-shaped ( Figs 36 C–D); embolic disc thick, with straight prolateral edge and curved retrolateral edge ( Fig. 37A ); PED longer than exposed portion of embolic disc (about twice the length) and emerging from the prodistal part of embolic disc ( Figs 37A, G ); embolus tip ends slightly beyond tip of PED ( Figs 37A, E ).

Female (MPEG 34358). Total length: 3.44. Carapace 1.78 long, 1.27 wide, 0.88 high. Ocular quadrangle 0.99 long. Anterior eye row 1.24 wide, posterior 1.08 wide. Legs 4123. Length of legs: I 3.76 (0.90 + 1.05 + 1.81); II 2.67 (0.93 + 1.00 + 0.74); III 3.53 (1.26 + 1.14 + 1.13); IV 3.78 (1.21 + 1.21 + 1.36).

Leg macrosetae: Femur I d1-1-1, p1di, r0; II d1-1-1, p1di, r1di; III d1-1-1, p1d1, r0 (or r1di); IV d1-1-1, p0, r1di. Patella I–II 0, III–IV p0, r1. Tibia I p0-1-0, r0, v2-2-2; II p0-1-0, r0, v1r-2-2; III p1-1-0, r1-1-0, v1p-0-1p; IV p1-1-1-0 (or p0-1-1-0), r1-1-1-0, v1p-0-2. Metatarsus I p1di, r0, v-2-2; II p1-1, r1di (or r0), v2-2; III p1-0-2, r1-0-2, v2-0-2; IV p1-1-2, r1-1-2, v1p-0-2.

Color in alcohol ( Figs 36 E–F): carapace and abdomen as in male; legs: femur I–II with dark distal ring and proximal region with dark retrolateral spot, III–IV with dark distal ring and proximal region with dark proventral spot; tibia I–IV with dark proximal ring and distal region with prolateral and retrolateral dark spot; tarsus I–IV without dark marks.

Epigyne ( Figs 35 H–I, 37H–I, 41–43): copulatory openings very close to each other and placed slighty more anteriorly than end portion of primary spermathecae; copulatory ducts long, encircling primary spermathecae; proximal copulatory duct shorter than distal section (twice as short); primary spermathecae with homogenous diameter and anteriorly projected.

Other material examined. FRENCH GUIANA: Maripasoula : [03°38’44.1”N, 54°02’02.3”W], leg. Quintero, 18–19.XII.1972, 1♁ (MACN-Ar 39565) . BRAZIL: Amapá: Rio Matapi [00°19’38.2”N, 51°12’39.6”W], leg. M.E. Galiano, 18.VI.1966, 1♁ 1♀ (MACN-Ar 39523) . Pará: Maracan „, Praia da Princesa, Ilha de Algodoal, 00°34’54.7”S, 47°34’27.9”W,leg. G.R.S.Ruiz & R.Valente, 3–9.VIII.2017, 1♁( MPEG37137 ); Soure[00°43’13.3”S, 48°31’08.6”W], leg. S.B. Farias, 2016, 1♁ ( MPEG 37138 ); Ilha de Cotijuba, Ruínas do Presídio, 01°16’13.7”S, 48°33’30.1”W, leg. A. Salgado, 13.VII.2017, 1♀ ( MPEG 37139 ); Santa Izabel do Pará, BR-316, 01°16’31.5”S, 48°05’52.8”W, leg. G.R.S Ruiz, 2017, 1♀ ( MPEG 37140 ); Benevides, [01°21’41.8”S, 48°15’17.3”W], leg. A. Salgado, 11.IX.2017, 2♁ 2♀ ( MPEG 37141 ); Belém, Utinga, [01°25’27.1”S, 48°26’39.8”W], leg. M.E. Galiano, VIII.1970, 1♁ (MACN-Ar 39506); Belém, near to Bosque Rodrigues Alves, 01°25’58.5”S, 48°27’23.0”W, leg. A. Salgado, 17.VII.2017, 1♁ ( MPEG 37142 ), 2♁ 1♀ ( MPEG 37143 ); Belém, Instituto de Ciências Biológicas, Universidade Federal do Pará, 01°28’20.6”S, 48°27’28.9”W, leg. A Salgado, VI–VII.2017, 1♁ ( MPEG 34357 ), 1♀ ( MPEG 34358 ), 1♁ 1♀ ( MPEG 37147 ), 1♀ ( MPEG 37148 ), 7♁ 1♀ ( MPEG 37149 ). Paraíba: Jo„o Pessoa, Hotel Globo, 07°06’44.9”S, 34°53’20.2”W, leg. A. Salgado, 23.VII.2019, 1♁ ( MPEG 37151 ) , 1♀ ( MPEG 37152 ) , 1♁ ( MPEG 37153 ) , 1♀ ( MPEG 37154 ) , 1♁ ( MPEG 37155 ) , 1♁ ( MPEG 37156 ) . Pernambuco: Santa Cruz do Capibaribe, Serra do Pará , 07°52’29.2’’S, 36°24’10.06’’W, leg. F.M.G. Las-Casas, 19–21.X.2011, 2♁ ( MPEG 37157 , 37158 ) . Piauí: Parque Nacional da Serra das Confusıes [09°13’17.9”S, 43°29’24.0”W], leg. L.S. Carvalho et al., 13–15.XII.2010, 1♁ ( MPEG 37159 ) , 1♀ ( MPEG 37160 ) , 1♁ ( MPEG 37161 ) , 1♁ ( MPEG 37162 ) , 2♁ ( MPEG 37163 ) , 1♀ ( MPEG 37164 ) . Acre: Rio Branco [09°58’52.6”S, 67°49’46.8”W], leg. M.D.F. Santos & O.C.R. Azevedo, 21.XI.1995, 1♀ ( IBSP 7010 ) . Bahia: Salvador, Hotel Sol Bahia , 12°57’46.1”S, 38°24’16.9”W, leg. A. Salgado, 7.VII.2018, 1♀ ( MPEG 37165 ) ; Lafaiete Coutinho , [13°39’22.3”S, 40°12’46.3”W], leg. J. Romão, VII.2006 – VII.2007, 1♁ ( IBSP 140421 ) . Mato Grosso: Poconé, Pantanal , Fazenda Retiro Novo , 16°22’44’’S, 56°18’47’’W, leg. A.C.C. Castilho et al., 2003, 1♁ ( IBSP 87300 ) , 1♁ ( IBSP 87301 ) , 1♁ ( IBSP 87306 ) , 1♀ ( IBSP 87312 ) , 1♀ ( IBSP 87313 ) , 1♁ ( IBSP 87314 ) . Minas Gerais: Santana do Riacho, Cardeal Mota , Serra do Cipó , 19°20’15.84’’S, 43°38’18.72’’W, leg. P.H. Martins et al., 17.VII.2012, 1♁ ( UFMG 12581 ) ; Prudente de Morais, Fazenda Sapé [19°28’26.6”S, 44°09’35.0”W], leg. E.S.S. Álvares, 13.X.2001, 1♁ 1♀ ( IBSP 44239 ) ; Caeté, Tauá Resort , 19°46’08.6”S, 43°37’41.9”W, leg. A. Salgado, 3–8.XII.2017, 1♀ ( MPEG 37166 ) , 1♁ ( MPEG 37167 ) , 1♁ ( MPEG 37168 ) , 1♁ ( MPEG 37169 ) , 1♁ ( MPEG 37170 ) ; Belo Horizonte [19°53’11.5”S, 43°56’53.8”W], leg. A. Salgado, 2.XII.2017, 2♁ 3♀ ( MPEG 37176 ) ; Belo Horizonte, Bairro Buritis , 19°58’43.9’’S, 43°58’04.3W, leg. D.F. Santos, 24.II.2016, 1♁ ( UFMG 19529 ) ; Santa Barbara, Santuário do Caraça, RPPN, Pico do Sol , 20°04’08.17”S, 43°30’17.21’’W, leg. W.L.N Perillo, 15.IX.2010, 1♁ ( UFMG 6701 ) ; Três Corações, Terminal Rodoviário , 21°42’00.3”S, 45°15’34.3”W, leg. A. Salgado, 2.XII.2017, 1♁ ( MPEG 37171 ) ; São Thomé das Letras, Vale das Borboletas , 21°43’31.5”S, 45°00’24.7”W, leg. A. Salgado, 10–12.XII.2017, 2♀ ( MPEG 37172 ) , 1♀ ( MPEG 37173 ) ; São Thomé das Letras, Eubiose Waterfall , 21°43’32.3”S, 44°57’36.1”W, leg. A. Salgado, 10–12.XII.2017, 1♁ ( MPEG 37174 ) . Mato Grosso do Sul: Corumbá, Sub-regions Miranda e Abobral , Passo do Lontra [19°34’34.3”S 57°01’40.1”W], leg. J. Raizer et al., VI.1998 – XI.1999, 4♁ 1♀ ( IBSP 87141 ) , 2♀ ( IBSP 87885 ) , 2♁ ( IBSP 87886 ) , 1♁ ( IBSP 87887 ) , 2♁ ( IBSP 87888 ) , 4♁ ( IBSP 87889 ) , 5♁ ( IBSP 87890 ) , 1♁ ( IBSP 87891 ) , 2♁ ( IBSP 87892 ) , 2♁ 3♀ ( IBSP 87893 ) , 1♀ ( IBSP 87894 ) , 1♁ ( IBSP 87895 ) , 4♁ ( IBSP 87896 ) . São Paulo: Vargem Grande do Sul [21°50’08.1”S, 46°53’38.0”W], leg. G. P. Perroni, 26.XII.2007 – 5.I.2008, 2♁ 1♀ ( IBSP 99252 ) ; Tupã [21°56’11.3”S, 50°31’02.1”W], leg. G.R.S Ruiz, I.2007, 2♁ 1♀ ( IBSP 71099 ) ; São Carlos, Bosque Pinus , UFSCar [21°59’04.1”S, 47°52’48.7”W], leg. M.A. Rezarde, II.2007, 1♁ ( IBSP 78795 ) ; Primavera, Usina Hidrelétrica Sérgio Motta [22°30’25.6”S, 52°52’08.2”W], leg. D. Candiani & R.P. Indicatti, II.2001, 1♁ ( IBSP 53438 ) ; Botucatu, Fazenda do Lageado , UNESP [22°51’27.3”S, 48°25’58.5”W], leg. E.F. Santos & C.P.S. Santos, 8.VI.2003, 1♁ ( IBSP 39664 ) . ARGENTINA: Salta: Pocitos [22°03’15.3”S, 63°41’18.5”W], leg. Braber, 21.XI.1951, 1♀ (MACN-Ar 39497) . Jujuy: Parque Nacional Calilegua [23°40’15.4”S, 64°51’59.5”W], leg. Ramírez & Comagnucci 7.VIII.1997, 1♀ (MACN-Ar 39598) . Misiones: Parque Nacional Iguazú ( Área Cataratas ) [25°40’59.4”S, 54°27’16.6”W], leg. M. Ramírez & L. Lopardo, 11–16.XII.1999, 1♁ (MACN-Ar 39546) ; San Ignacio [27°15’23.5”S, 55°32’06.3”W], leg. Galiano, 1.X.1983, 1♀ (MACN-Ar 39537) ; San Javier [27°51’52.6”S, 55°08’15.4”W], leg. Biraben, XII.1948, 2♀ (MACN-Ar 39520, 39582) ; Santa María [27°56’02.1”S, 55°24’24.4”W], leg. Viana, XI.1962, 1♁ (MACN-Ar 39502) . Santiago del Estero: Dto. Moreno, Mercedes 10 Km E de Amamá [27°21’55.1”S, 62°28’20.4”W], leg. D. Vezzani, I–VIII.1996, 3♀ (MACN-Ar 39524), 1♀ (MACN-Ar 39525) .

Distribution. The most widespread species in the genus. Known from the Venezuelan state of Bolívar; the Esequibo and Alto Demerara regions of Guyana; the Maripasoula region of French Guyana; the Brazilian states of Acre, Amapá, Amazonas, Bahia, Pará, Paraíba, Pernambuco, Piauí, Mato Grosso, Mato Grosso do Sul, Minas Gerais and São Paulo; the Argentine provinces of Salta, Jujuy, Misiones and Santiago del Estero ( Fig. 59A ).

Natural history. The specimens were collected from tree trunks, beach sand and urban structures (camouflaged on dark walls or places covered with moss) ( Figs 1A, D ).

Figures

FIGURE 1. Marma Simon, 1902. A–E live specimens in different habitats [A M. nigritarsis from Belém/Pará/Brazil, on urban construction (Photo credit: César Favacho); B M. linae sp. nov. from São Geraldo do Araguaia/Pará/Brazil, on tree trunk; C M. rosea from São Geraldo do Araguaia/Pará/Brazil, on a wall of an urban construction; D M. nigritarsis from Belém/Pará/Brazil, on a substrate with moss; E M. wesolowskae sp. nov. from Buíque/Pernambuco/Brazil, on rocks].

FIGURE 25. Male holotype of Marma pipa sp. nov. (MPEG 34352). A–B habitus (A dorsal, B ventral); C–D left palp (C ventral; D retrolateral).

FIGURE 26. Female paratype of Marma pipa sp. nov (MPEG 34349). A–B habitus (A dorsal, B ventral); C–D cleared epigyne/ vulva (C ventral, D dorsal). Abbreviations: BG—Bennett’s gland; cd—copulatory duct; co—copulatory opening; ets—endothe- cal spikes; fd—fertilization duct; PS—Primary spermatheca; SS—secondary spermatheca.

FIGURE 27. Marma pipa sp. nov. A–E left male palp (A ventral; B retrolateral; C proventral; D dorsal; E cleared bulb, ventral); F–G epigyne/vulva (F ventral; G cleared, ventral). Abbreviations: BG—Bennett’s gland; cc—cymbial conductor; cd—copu- latory duct; co—copulatory opening; dh—distal hematodocha; e—embolus shaft; ed—embolic disc; fd—fertilization duct; PED—process on embolic disc; PS—primary spermatheca; RTA—retrolateral tibial apophysis; SS—secondary spermatheca.

FIGURE 34. Marma nigritarsis (Simon, 1900), live specimens from Belém/Pará/Brasil.A–C male (A dorsal, B lateral, C frontal); D–F female (D dorsal, E lateral, F frontal). Black arrow in A shows black tarsus in male front leg.

FIGURE 35. Marma nigritarsis (Simon, 1900). A–D male holotype, collection of E. Simon deposited in the MNHN (A habitus, dorsal; B ventral; C right palp reflected as left palp, proventral; D label data). E–F male holotype of Pseudoamphidraus variegatus Caporiacco, 1947, MZLS 597 (A habitus, dorsal; B ventral). G–I female holotype of Marma trifidocarinata Caporiacco, 1947, MZLS 549 (G habitus, dorsal; H cleared epigyne/vulva, ventral; I dorsal). Abbreviations: BG—Bennett’s gland; cd—copulatory duct; co—copulatory opening; fd—fertilization duct; PS—primary spermatheca; SS—secondary spermatheca.

FIGURE 36. Marma nigritarsis (Simon, 1900), specimens from Belém/Pará/Brasil. A–D male, MPEG 34357 (A habitus, dorsal; B ventral; C left palp, ventral; D retrolateral). E–F female, MPEG 34358 (E habitus, dorsal; F ventral).

FIGURE 37. Marma nigritarsis (Simon, 1900). A–E, G specimen from Belém/Pará/Brazil. F holotype illustrated by Galiano (1962), reflected.A–G left male palp (A ventral, B retrolateral, C proventral, D dorsal, E–F retroventral; G cleared bulb, ventral). H–I female holotype of M. trifidocarinata Caporiacco, 1947, epigyne/vulva (H ventral; I cleared, ventral).Abbreviations: BG— Bennett’s gland; cc—cymbial conductor; cd—copulatory duct; co—copulatory opening; dh—distal hematodocha; e—embolus shaft; ed—embolic disc; fd—fertilization duct; PED—process on embolic disc; PS—primary spermatheca; RTA—retrolateral tibial apophysis; SS—secondary spermatheca.

FIGURE 38. Male of Marma nigritarsis (Simon, 1900).A–B, D–E Left male palp, expanded (A retrolateral, partially expanded; B totally expanded; D prolateral, partially expanded; E totally expanded); C left bulb, cleared, dorsal. Abbreviations: bh—basal hematodocha; Cy—cymbium; dh—distal hematodocha; e—embolus shaft; ed—embolic disc; fnd—fundus; RTA—retrolateral tibial apophysis; sp—spermophore; st—subtegulum; T—Tegulum; Ti—tibia.

FIGURE 39. Pair of Marma nigritarsis (Simon, 1900), A–D courtship, ventral view (A male approximation; B the male insemi- nating the female; C after the insemination; D detail of partially expanded male palp). Abbreviations: bh—basal hematodocha; dh—distal hematodocha; e—embolus shaft; ed—embolic disc.

FIGURE 40. Male of Marma nigritarsis (Simon, 1900). A–D left palp (A apical region of cymbium, ventral; B tibia and proximal region of cymbium, ventral; C dissected embolic disc and embolus, dorsal; D detail of embolus shaft, dorsal). Abbreviations: cc—cymbial conductor; e—embolus shaft; ed—embolic disc; PED—process on embolic disc. Black arrow in B shows retroventral projection of cymbium; in C shows groove; white arrow in C shows projection of embolus base; white arrows in D show spikes of embolus shaft.

FIGURE 41. Female of Marma nigritarsis (Simon, 1900).A–D Epigyne/vulva (A ventral; B dorsal; C–D primary spermatheca, dissected, showing internal integumental structures). Abbreviations: BG—Bennett’s gland; co—copulatory opening; ets—en- dothecal spikes; fd—fertilization duct; PS—primary spermatheca; SS—secondary spermatheca.

FIGURE 42. Female of Marma nigritarsis (Simon, 1900). A–D Epigyne/vulva (A initial portion of copulatory duct and end of primary spermatheca, left side, dissected, posterior view; B detail of previous image, showing Bennett’s gland, endothecal spikes and pore leading to fertilization duct; C detail of spiky Bennett’s gland; D detail of dissected copulatory duct).Abbrevia- tions: BG—Bennett’s gland; cd—copulatory duct; ets—endothecal spike; fd—fertilization duct.

FIGURE 43. Female of Marma nigritarsis (Simon, 1900). A–D Epigyne/vulva (A initial portion of copulatory duct, left side, anterior view; B detail of previous image showing secondary spermatheca). C–D detail of glands on initial portion of copula- tory duct. Abbreviations: cd—copulatory duct; co—copulatory opening; SS—secondary spermatheca. White arrows in C show glands near copulatory opening.

FIGURE 44. Male of Marma argentina (Mello-Leit„o, 1941), specimen from Cura Brochero/Córdoba/Argentina (MACN-Ar 39650). A–B habitus (A dorsal, B ventral); C–D left palp (C ventral, D retrolateral).

FIGURE 45. Females of Marma argentina (Mello-Leit„o, 1941). A–C holotype of Ocnotelus argentinus, MLP 14980 (A habitus, dorsal; B ventral; C cleared epigyne, ventral); D–F holotype of Thysema dorae, MLP 16300 (D habitus, dorsal; E ventral; F cleared epigyne, ventral). G–I specimen from Cura Brochero/Córdoba/Argentina, MACN-Ar 39648 (G habitus, dorsal; H ven- tral; I cleared epigyne, left side, ventral). Abbreviations: BG—Bennett’s gland; cd—copulatory duct; co—copulatory opening; ets—endothecal spikes; fd—fertilization duct; PS—primary spermatheca; SS—secondary spermatheca.

FIGURE 46. Marma argentina (Mello-Leit„o, 1941). A–B left male palp, specimen from Cura Brochero/Córdoba/Argentina, MACN-Ar 39650 (A ventral, B retrolateral); C–D Epigyne/vulva, holotype of Thysema dorae, MLP 16300 (C ventral; D cleared, ventral). Abbreviations: BG—Bennett’s gland; cc—cymbial conductor; cd—copulatory duct; co—copulatory open- ing; e—embolus shaft; ed—embolic disc; fd—fertilization duct; PED—process on embolic disc; PS—primary spermatheca; RTA—retrolateral tibial apophysis; SS—secondary spermatheca.

FIGURE 48. Males of Marma rosea (Mello-Leit„o, 1941). A–D types of Paralophostica centralis [A habitus (Lectotype ♁, MZUSP E.740 C.1334), dorsal; B ventral; C left palp (paralectotype ♁, MZUSP E.740 C.1335), ventral; D retrolateral]; E–G specimen from São Geraldo do Araguaia/Pará/Brasil, MPEG 34360 (E habitus, dorsal; F ventral; G left palp, ventral). White arrows in F show black tarsi on legs I and II.

FIGURE 49. Females of Marma rosea (Mello-Leit„o, 1941). A–C paralectotype of Paralophostica centralis (MZUSP E.740 C.1335) (A dorsal; B ventral; C cleared epigyne, ventral); D–F holotype of Agelista rosea (MLP 14965) (D dorsal; E ventral; F cleared epigyne, ventral).Abbreviations: BG—Bennett’s gland; cd—copulatory duct; co—copulatory opening; fd—fertilization duct; PS—primary spermatheca; SS—secondary spermatheca.

FIGURE 50. Marma rosea (Mello-Leit„o, 1941). A–E male paralectotype of Paralophostica centralis, E.740 C.1335 (A left male palp, ventral; B retrolateral; C proventral; D dorsal; E cleared bulb, ventral). F–G epigyne/vulva, holotype of Marma rosea, MLP 14965 (F ventral, G cleared, ventral).Abbreviations: BG—Bennett’s gland; cc—cymbial conductor; cd—copulatory duct; co—copulatory opening; dh—distal hematodocha; e—embolus shaft; ed—embolic disc; fd—fertilization duct; PED—process on embolic disc; PS—primary spermatheca; RTA—retrolateral tibial apophysis; SS—secondary spermatheca.

FIGURE 57. Left male palp, ventral [A Marma baeri Simon, 1902; B M. linae sp. nov.; C M. pipa sp. nov.; D M. wesolowskae sp. nov.; E M. nigritarsis (Simon, 1900); F M. sinuosa sp. nov., G M. argentina (Mello-Leit„o, 1941), H M. rosea (Mello- Leit„o, 1941), I M. spelunca sp. nov.]. Colors represent homologous structures.

FIGURE 58. Epigyne/vulva, A–C, E–K ventral, D dorsal [A Marma baeri Simon, 1902; B M. linae sp. nov.; C M. abaira sp. nov.; D M. femella (Caporiacco, 1955); E M. pipa sp. nov.; F M. wesolowskae sp. nov.; G M. nigritarsis (Simon, 1900); H M. sinuosa sp. nov.; I M. argentina (Mello-Leitão, 1941); J M. rosea (Mello-Leitão, 1941); K M. spelunca sp. nov.]. Colors represent homologous structures; arrow in D shows hypothetical position of secondary spermatheca.

FIGURE 59. Maps of South America showing records of Marma species (A species previously described; B new species). Record from the state of Amazonas (Brazil) in Fig. 59A is taken from Zhang & Maddison (2015).