Tetragnatha extensa (Linnaeus, 1758)

Tetragnatha differs from Dolichognatha, Pachygnatha and Glenognatha by its elongate and tubular-shaped abdomen, normally covered by guanine crystals. It further differs from Dolichognatha in lacking an epigynum (having a genital fold instead) (Figs 1H, 15H), by having trichobothria on their femora (Fig. 1A–C, 2A–C, 6A–C), a modified cymbium with its tip elongate and thin and a free paracymbium (e.g., Figs 2I, J, 6I, J, 7I, J). From Glenognatha and Pachygnatha, it may be identified by the absence of a tapetum on the lateral eyes (see Levi 1981) and by the more horizontal and forward projecting chelicerae (e.g., Figs 10A, 18A, B). From Glenognatha, it differs by the normal position (not advanced) of the tracheal spiracle (Cabra-Garcia and Brescovit 2016). From Pachygnatha, it may be distinguished by the sternum not projecting between the coxae, the carapace not covered with short macrosetae immersed in pits and the absence of the sclerotized genital slit (Dimitrov and Hormiga 2009; Alvarez-Padilla and Hormiga 2011). Adult Tetragnatha build orb-webs, while adult Pachygnatha do not spin webs. Additionally, Tetragnatha differs from Cyrtognatha by the more elongated abdomen, the absence of the straight line of long and robust macrosetae with enlarged bases on the spinnerets, male pedipalps with paracymbium as a separate sclerite connected to the cymbium by a membrane on its base, and females with enlarged spermathecae (reduced in Cyrtognatha) (Figs 1B, I, 2A, B, H, J; Dimitrov and Hormiga 2009).

Carapace longer than wide, normally with an elevated anterior part, without projections or bands (e.g., Figs 1A, B, 2A, B, 6A, B). Eyes normally large, ringed in black, with touching ALE and PLE in some species (e.g., Fig. 6A, B). Fovea normally small and visible (e.g., Figs 1A, B, 2A, B). Sternum oval, longer than wide, sometimes with dusky edges, with or without a dark contour (e.g., Figs 1C, 2C, 6C, 7C). Legs varying from pale yellow to light brown, legs I and II normally slightly darker than III and IV, all very long, leg formula I–II–IV–III or I–IV–II–III, leg I always much longer than all others, twice as long as leg III and sometimes having multiple spines on femur (e.g., Figs 1A, B, 2A, B, 6A, B). Chelicerae elongated, with paturon bearing bulges or crests in some species (Figs 6D, F, 7D, F, 14D, F) and abundant teeth on both sides, with males bearing a dorsal apophysis to lock female’s fangs and additional or modified teeth, such as ‘sl’, ‘t’ and ‘T’ (Figs 2D–G, 6D–G, 7D–G, 10D–G, 14D–G); fangs may have basal (BC), outer (OC), inner (IC) or median cusps (MC) (e.g., IC and MC in Fig. 1D–F or BC in Castanheira et al. 2019, fig. 12D–F). Abdomen longer than carapace, normally covered by guanine crystals, without tubercules or spines, sometimes with a projection after the spinnerets varying from short (e.g., Fig. 10B, C) to extremely elongated (Castanheira and Baptista 2021b, figs 1B, C, 2 B, C, 4A, B). Male pedipalps with elongated cymbium, as long (e.g., T. elongata) or longer than the tibia (e.g., T. vermiformis) (Castanheira et al. 2019, figs 5H, 17H–J, 19C). Tibia very short (e.g., T. jaculator) (Castanheira and Baptista 2021a, figs 63–65, 78), short (e.g., T. cristata sp. nov., T. oncognatha sp. nov.) (Figs 7H–J, 8C, 14H–J), or elongated (e.g., T. elongata, T. nitens) (Castanheira et al. 2019, figs 5H, 14G–I, 16E). Tegulum oval, always wider than long, slightly (e.g., T. elongata, Castanheira et al. 2019, fig. 5H) or extremely slanted (e.g., T. jaculator, Castanheira and Baptista 2021a, figs 63–65, 78). Conductor always twisted, with pleats varying from three (e.g., T. elongata) (Castanheira et al. 2019, figs 5H, I, 7C, D) to zero (e.g., T. cristata sp. nov. (Figs 7H, 8C), and tip of conductor and embolus with or without tail-like projections (Castanheira et al. 2019, fig. 20A–F). Embolus tip may be completely covered by the pouch-like conductor (e.g., T. bogotensis) (Castanheira et al. 2019, figs 1I, J, 3A, E, 20A), partially exposed (e.g., T. argentinensis; T. cristata sp. nov.) (Figs 7H, I, 8C, D; Cargnelutti et al. 2022, figs 2H, I, 3E, F) or completely apart from the conductor (e.g., T. amazonica sp. nov.; T. tenuissima) (Fig. 6H, I; Castanheira and Baptista 2020, figs 12G, H, 14E, F). Paracymbium may be very elongated (e.g., T. bogotensis; T. nitens) (Castanheira et al. 2019, figs 1K, 3D, 14J, 16F) or reduced in size (e.g., T. amazonica sp. nov.; T. jaculator) (Fig. 6J; Castanheira and Baptista 2021a, figs 65, 80). Paracymbium notch (N) can be carved in two parts (e.g., T. bogotensis, T. nitens) (Castanheira et al. 2019, figs 1K, 3D, 14I, J, 16F), slightly dented (e.g., T. cristata sp. nov., T. oncognatha sp. nov.) (Figs 7J, 8E, 14J, 16G) or rounded and not divided (e.g., elongata, T. keyserlingi) (Castanheira et al. 2019, figs 5K, 7F, 8J, 10E); Paracymbium translucent lobe (L) can be very broad (e.g., T. nitens, T. chauliodus, T. renatoi) (Castanheira et al. 2019, figs 14I, J, 16F; Castanheira and Baptista 2020, figs 4I, 6F, 9K), reduced (e.g., T. keyserlingi) (Castanheira et al. 2019, figs 8J, 10E) or not visible (e.g., T. elongata, T. vermiformis) (Castanheira et al. 2019, figs 5K, 7F, 17J, 19E). Female genital fold varies from short and wider than long, with straight or curved posterior rim (e.g., T. megalocera, T. renatoi, T. tenuissima (Castanheira and Baptista 2020, figs 2H, 5H, 13G), to extremely elongated and longer than wide, with rounded posterior rim (T. cladognatha, T. bogotensis, T. keyserlingi, T. mandibulata, T. nitens, T. pradoi sp. nov.) (Figs 1H, 18G; Castanheira et al. 2019, figs 2G, 9H, 12H, 15J). Female internal genitalia usually with central membranous sac (CS), but it is sometimes absent (e.g., T. vermiformis, T. laboriosa) (Castanheira et al. 2019, fig. 18I; Castanheira and Baptista 2021a, fig. 47). CS may have its head varying from rounded or oval (e.g., T. oncognatha sp. nov., T. bogotensis, T. mandibulata, T. nitens) (Fig. 12I; Castanheira et al. 2019, figs 2H, I, 12I, 15K, L) to cylindrical or thin and slender (e.g., T. pradoi sp. nov., T. tenuissima) (Fig. 15H; Castanheira and Baptista 2020, fig. 13H), with variable stalk length, from short (e.g., T. oncognatha sp. nov., T. bogotensis) (Fig. 15I; Castanheira et al. 2019, fig. 2H, I) to very elongated (T. keyserlingi) (Castanheira et al. 2019, fig. 9I), or even having either long or short stalks depending on the specimen (T. nitens) (Castanheira et al. 2019, fig. 15K, L). Spermathecae varies in number, either two (one on each side) (e.g., T. cladognatha, T. elongata, T. mandibulata) (Figs 1I, 3I; Castanheira et al. 2019, figs 6H, 12I), but sometimes four (e.g., T. megalocera, T. guatemalensis, T. laboriosa) (Castanheira and Baptista 2020, fig. 2I; Castanheira and Baptista 2021a, figs 20, 47), and in shape, from perfectly globular (T. oncognatha sp. nov.) (Fig. 15I) to oval (e.g., T. bogotensis, T. mandibulata, T. nitens) (Castanheira et al. 2019, figs 2H, I, 12I, 15K, L) or even kidney-like (T. elongata, T. vermiformis) (Castanheira et al. 2019, figs 6H, 18I). Colour of specimens is variable, encompassing reddish, yellowish, brown and orange hues depending on time spent in alcohol (e.g., Figs 1A, B, 6A, 11A). Old specimens, such as the female T. paraguayensis, usually lose completely their original colour, presenting a light-yellow bleached tone (Castanheira and Baptista 2021b, fig. 4A).

Cosmopolitan.

Tetragnatha cladognatha: Brazil: female, holotype, Rio de Janeiro, not located (presumed lost). Brazil, Rio de Janeiro • Neotype female herein designated, Rio de Janeiro, Parque Estadual da Pedra Branca, Camorim (Sede), 22°58'12.0"S, 043°26'16.4"W, 160 m, 09.i.2014, RLC Baptista leg. (UFRJ 1628).

Argentina, Misiones • 1 female, Parque Provincial Uruzú, 1° Uruzú, 25°50′S, 54°08'W, 02.ii.1988, P. Goloboff and Szumik leg. (MACN-Ar 24553); • 2 males, 4 females, 3 juveniles, Departamento Cainguás, Parque Provincial Salto Encantado, Arroyo Cuña-Pirú, 27°07'S, 54°48'W, 12.i.2005, C Grismado, L Lopardo, L Piacentini, A Quaglino and G Rubio leg. (MACN-Ar 31757); • 1 male, 1 female, 1 juvenile, same data (MACN-Ar 31763); • 1 male, Saltos del Uruguay, 10 Km N Puerto Libertad, 25°55'08.0"S, 54°35'59.7"W, 23–25.ii.1997, M Ramírez leg. (MACN-Ar 24661); • 1 female, same data, (MACN-Ar 24645); • 1 male, 8 females, 6 juveniles, Santa Maria, 27°56'05.6"S, 55°24'54.9"W, x.1956, Viana leg. (MACN-Ar 24423); • 1 male, 2 females, 1 juvenile, Cataratas del Iguazú, 25°41'28.1"S, 54°26'43.6"W, xi.1954, BC Schiapelli leg. (MACN-Ar 39613); BRAZIL, Mato Grosso do Sul • 1 male, Jaraguari, Furnas de Dionísio, 20°09'21.5"S, 54°43'34.4"W, 14.xi.2015, D Araújo leg. (IBSP 167038); Minas Gerais • 3 males, 4 females, Alto Caparaó, Parque Nacional do Caparaó, 20°05'S, 41°09'W, 01–07.v.2002, Equipe Biota leg. (IBSP 220017); • 1 female, Rio Preto, 22°05'21.1"S, 43°50'11.1"W, 14–20.v.2002, RLC Baptista et al. leg. (MNRJ 1577); • 2 females, same data, (MNRJ 1586); Paraná • 2 females, São José dos Pinhais, 25°36'01.40"S, 049°11'24.66"W, 08.i.2002, A Chagas-Jr. leg. (MNRJ 03744); • 1 female, same locality, 15.xi.2015, AC Domahovski leg. (MCTP 39130); • 1 male, same locality, 25°36'12.65"S, 49°11'33.58"W, x.2015, AC Domahovski leg. (MCTP 39055); Rio de Janeiro • 1 male, 2 females, 2 juveniles, Cachoeiras de Macacu, Reserva Ecológica de Guapiaçu (REGUA), river nearby accommodation, 22°27'12.4"S, 42°46'21.1"W, RLC Baptista leg. (UFRJ 1503); • 1 female, same locality, trilha cinza, 29.viii.2019, AA Alves leg. (UFRJ 1620); • 1 male, 3 females, Guapimirim, 22°35'25.1"S, 43°06'15.7"W (MNRJ 1572); • 1 female, same locality, ii.1996 (MNRJ 1570); • 3 males, 2 females, 1 juvenile, Itatiaia, Cachoeira Véu da Noiva, 1250 m, 22°25'35.6"S, 44°37'12.6"W, 06.i–03.ii.2016, M Monné leg. (MNRJ 4260); • 1 male, Macaé, Sana, tributário 2 ordem do Rio Sana, 313 m, 22°19'39.6"S, 42°11'11.4"W, 16.ii.2009, Entomologia UFRJ leg. (UFRJ 0378); • 1 male, 1 female, Magé, Piabetá, 22°36'21.7"S, 43°10'36.8"W, xi.1986 (MNRJ 1574); • 1 male, Nova Friburgo: Macaé de cima, 956 m, 22°21'11.1"S, 42°24'40.5"W, 01.xii.2008, BHL Sampaio and APM Santos leg. (UFRJ 0071); • 1 female, same locality, Rio das Flores, 22°24'06.7"S, 42°29'19.4"W, BHL Sampaio leg. (UFRJ 0073); • 1 female, Nova Iguaçu, Parque Municipal de Nova Iguaçu (PMNI), 22°46'44.6"S, 43°27'31.8"W, 01.v.2004, BHL Sampaio leg. (MNRJ 07445); • 1 male, 2 females, same locality, 28.v.2004, BHL Sampaio leg. (MNRJ 07449); • 1 female, same locality, 10.vii.2004, BHL Sampaio leg. (MNRJ 07446); • 2 females, same locality, 31.vii.2004, C Lima leg. (MNRJ 07444); • 1 male, 1 female, same locality, 26.xi.2004, BHL Sampaio leg. (MNRJ 07447); • 1 female, Petrópolis, 22°29'01.5"S 43°15'10.8"W, Mello-Leitão leg. (MNRJ 60003); • 1 female, Pinheiral, Pinheiro, 22°30'24.3"S, 44°01'22.2"W, Mello-Leitão leg. (MNRJ 60009); • 1 male, Resende, Vila da Fumaça, Estrada Falcão-Fumaça, 22°17'57.1"S, 44°13'07.9"W, 10.xi.2016, LBN Coelho leg. (UFRJ 1366); • 1 male, 1 juvenile, Rio de Janeiro, Parque Estadual da Pedra Branca, Camorim (Sede), 22°58'12.0"S, 043°26'16.4"W, 160 m, 15.ix.2013, RLC Baptista leg. (UFRJ 1622); • 1 female, 1 juvenile, same data (UFRJ 1623); • 1 female, same data (UFRJ 1624); • 1 female same data but RLC Baptista and PdS Castanheira leg. (UFRJ 1625); • 3 males, 1 female, 7 juveniles, same locality, 09.i.2014, RLC Baptista leg. (UFRJ 1626); • 5 females, 4 juveniles, same data (UFRJ 1629); • 1 female, same locality, 07.iv.2014 (UFRJ 1627); • 1 male, 1 female, 1 juvenile, same locality, Camorim (Véu da Noiva); 04.x.2017, ALD Ferreira leg. (UFRJ 1483); 1 male, same data (UFRJ 1523); • 1 female, same data (UFRJ 1524); • 1 female, same data (UFRJ 1525); • 1 male, Sepetiba, 22°58'06.9"S, 43°42'46.8"W, iv.1994, EH Wienskoski leg. (MNRJ 1585); • 1 male, 4 females, 1 juvenile, Teresópolis, Serra do Subaio, 22°27'10.8"S, 42°56'48.8"W, 20–22.iv.1995, RLC Baptista and M Landim leg. (MNRJ 1568); • 3 females, 2 juveniles, Visconde de Mauá: Alto Penedo (Rio das Pedras), 22°23'03.1"S, 44°37'32.5"W, i.2006, EH Wienskoski leg. (MNRJ 05041); Rio Grande do Sul: • 1 female, Eldorado do Sul, 30°06'59.3"S, 51°40'28.0"W, 28.iii.1993, M Silveira leg. (MCTP 43347); • 1 female, Novos Cabrais, Parque Witeck, 29°46'59.1"S, 52°58'26.7"W, 18.ii.2008, RG Buss leg. (MCTP 28307); • 1 male, 2 females, São Francisco de Paula, Potreiro Velho, 29°23'56.1"S, 50°16'12.6"W, 16–17.iii.2001, AA Lise leg. (MCTP 14349); • 5 males, Rio Uruguai, 29°27'34.6"S, 56°43'54.2"W, 02.ix.2010, RC Francisco leg. (MCTP 43349); Santa Catarina • 1 male, Nova Teutônia, 27°09'40.4"S, 52°25'31.8"W, 13–15.x.2006, ELC Silva et al. leg. (MCTP 28703); São Paulo • 3 females, Botucatu, 22°56'15.3"S, 48°23'32.6"W, ii.2002, EH Wienskoski leg. (MZUSP 62396); • 1 male, 1 female, Cachoeira da Marta, 22°55'55.0"S, 48°24'19.7"W, 10.i.2002, EH Wienskoski leg. (MZUSP 62429); • 1 male, 1 female, same locality, 10.iii.2002, EH Wienskoski leg. (MZUSP 62036); • 1 male, 1 female, same locality, i.2003, EH Wienskoski leg. (MZUSP 62430); • 1 male, Fazenda Paulina, 22°56'15.3"S, 48°23'32.6"W, i.2002 (MZUSP 62395); • 1 male, 2 females, Cabreúva, Sítio do Sol, 23°18'54.4"S 47°05'48.4"W, 05.iv.2009, AB Ribeiro leg. (IBSP 145209); • 2 males, São José do Barreiro, Parque Nacional Serra da Bocaina, Fazenda do Bonito, 22°43'25.3"S 44°32'21.0"W, Vulcano leg. (MZUSP 14746); • 1 male, same locality, Córrego do Boqueirão, 19.xii.2010, Entomologia UFRJ leg. (UFRJ 0550); • 1 male, 1 female, Rio Claro, Lidice, RPPN Fazenda Sambaíba, 22°50'55"S, 44°13'03"W, 23.iv.2012, C Bragagnolo et al. leg. (IBSP 213057); • 11 males, 26 females, Mogi das Cruzes, Manoel Ferreira, Biritiba-Uçu, 23°38'20.6"S, 46°07'33.1"W, v.2001, EK Kashimata and R Martins leg. (IBSP 56320).

Males and females most resemble T. argentinensis (Cargnelutti et al. 2022). Females can be distinguished by the following differences in the chelicerae: fang almost sickle-shaped, with a semi-circular basal half and an abruptly slanted and straighter distal half basally to the inner cusp; Gu more sclerotized with larger basis; Gl bulkier and closer to fang basis; L2 larger with wider, bulged, rounded basis, median cusp (MC) more distally placed and presence of conspicuous inner cusp (Figs 1D–F, 3C). Males of T. cladognatha differ from T. argentinensis by slightly shorter and less pointed ‘T’; longer and bulkier Gu, Gl and L2 (Figs 2D–F, 3A, B); Gl and L2 straight with much larger basis (Figs 2E, F, 3B); upper teeth row with lesser teeth and lower row with more teeth (Figs 2D, F, 3A, B); narrower pedipalp tibia (Figs 2H–J, 3D); narrower, thinner and more pointed conductor tip (Figs 2I, 3F) and epiandrous field almost straight, with narrower median division, with more spigots (26 vs. 18) (Fig. 3H; Cargnelutti et al. 2022, fig. 3H). Additionally, males are also similar to T. keyserlingi with T. cladognatha differing from it due to Gu shorter, with larger basis, ‘T’ longer and more projected, Gl and L2 bulky and more elongated, with much larger bases (Figs 2D–F, 3A, B). Pedipalps of T. cladognatha are identified by the less protruding hook-like conductor and paracymbium with divided notch and wider translucent lobe (Figs 2H–J, 3D–G).

Figure 1. 

Tetragnatha cladognatha female. A. Live specimen, dorsal view (carapace and chelicerae); B–H. Neotype female (UFRJ 1628); B. Dorsal habitus; C. Ventral habitus; D–G. Left chelicera; D. Upper view; E. Inner view; F. Lower view; G. Outer view; H–I. Female genitalia; H. Genital fold, ventral view; I. Internal genitalia, dorsal view (UFRJ 1524). Scale bars: 2 mm (B, C); 1 mm (D–G); 0.5 mm (H); 0.1 mm (I). Photo of live specimen: Ederson Oliveira.

Figure 2. 

Tetragnatha cladognatha male. A. Live specimen, dorsal habitus (not collected); B. Dorsal habitus (UFRJ 1483); C. Ventral habitus (UFRJ 1483); D–G. Left chelicera (UFRJ 1483); D. Upper view; E. Inner view; F. Lower view; G. Outer view; H–J. Left pedipalp (UFRJ 1483); H. Mesal view; I. Dorsal view; J. Ventral view (paracymbium). Scale bars: 2 mm (A–C); 0.2 mm (D–J). Photo of live specimen: Ederson Oliveira.

Figure 3. 

Tetragnatha cladognatha SEM photos. A, B. Male left chelicera (UFRJ 1483); A. Upper view; B. Lower view; C. Female left chelicera upper view (UFRJ 1629); D–G. Left male pedipalp (UFRJ 1523); D. Mesal view; E. Bulb detail, distal-mesal view; F. Tip of conductor and embolus opening detail, dorsal view; G. Paracymbium detail, ventral view; H. Epiandrous field detail, ventral view (UFRJ 1523); I. Internal genitalia, dorsal view (UFRJ 1524). Scale bars: 0.5 mm (A–D); 0.2 mm (E); 0.02 mm (F); 0.1 mm (G, I); 0.05 mm (H).

Female (based on neotype UFRJ 1628): Carapace elongated, oval and reddish brown, slightly elevated anteriorly (Fig. 1A, B). Fovea reddish brown, slightly darker than carapace, with dark borders (Fig. 1A, B). Labium wider than long and dark brown (Fig. 1C). Sternum oval and light brown (Fig. 1C). Eyes with procurved parallel rows, and evenly separated, AME and PME separated by its length, ALE and PLE almost touching (Fig. 1A, B). Legs reddish brown, with few spines on femora (Fig. 1B, C). Chelicera paturon thick, around 4× longer than wide and about 1.6× longer than carapace, well curved outwards, around 50° from body median line (Figs 1B, D–G, 3C). AXu absent (Figs 1D, E, 3C). Upper row with nine teeth distalward projected (Figs 1D, E, 3C): Gu almost straight, pointed, almost as long as U3, bearing bulky and wide basis and apart from U2 by an extremely large gap; U2 small and pointed, almost as long as U4–U7; U3–U9 decreasing in size and pointed. AXl absent (Fig. 1E, F). Lower row with 20 teeth distalward projected (Fig. 1E, F): Gl bulky and very sclerotized and located on fang furrow; L2 elongated, pointed, with rounded bulging basis and apart from Gl by a small gap; L3–L12 decreasing in size. Cheliceral fang very elongated, thick, with pointed and large median cusp (MC) on its first third, facing upper row and becoming slanted and projected inward to its tip from small pointed inner cusp (IC) at around half of its length (Figs 1D, E, 3C). Abdomen around 4.1× longer than carapace, cylindrical and anteriorly enlarged, dorsally greyish and completely covered by guanine crystals (Fig. 1B). Venter colour as dorsum, with a median brown stripe from genital fold towards spinnerets (Fig. 1C). Genital fold elevated, 1.4× wider than long, with parallel borders and ending in concave and wide excavated tip (Fig. 1H). Internal genitalia formed by two oval spermathecae, more sclerotized on the lateral border, and connected to a wide uterus externus and an almost cylindrical central membranous sac (Figs 1I, 3I).

Measurements. Total length 12.5. Carapace 4.4 long, 2.3 wide. Abdomen 10.0 long, 2.9 wide. Left chelicera 2.9 long, 0.5 wide. Leg formula I–IV–II–III. Leg I: femur 13.3, patella 1.8, tibia 11.5, metatarsus 11.5 and tarsus 2.3. Leg II: patella + tibia 8.6. Leg III: patella + tibia 3.4. Leg IV: patella + tibia 7.3.

Male (based on UFRJ 1483): Carapace, fovea, eyes, legs, legs and sternum similar to female (Figs 2A–C). Chelicerae paturon with similar colour as female, around 2.2× longer than wide, about 1.15× longer than carapace, slightly curved outwards around 35° from body median line (Figs 2B, D–G, 3A, B). ‘a’ elongated, thin, pointed, and distally projected, located on edge of paturon close to fang groove (Figs 2D, E, G, 3A). AXu short with large basis (Figs 2D, E, 3A), ‘t’ absent (Figs 2D, E, 3A). Upper row with eight teeth distalward projected (Figs 2D, E, 3A): Gu with large basis, thick and pointed, located on fang groove; ‘sl’ absent; ‘T’ very very elongated, thin, and pointed, with wide basis, slightly projecting upward and ‘rsu’ with six straight pointed teeth decreasing in size, with large gap between ‘T’–U3 and U3–U4. AXl absent (Figs 2E, F, 3B). Lower row with 17 teeth and two additional ones, one besides U4 and another besides U6, all distalward projected (Figs 2E, F, 3B): Gl and L2 very similar, thick, bulky with large bases, apart by small gap, Gl slightly bulkier with larger basis; L3–L17 and two additional teeth with almost the same size, all very short, triangular and pointed. Cheliceral fang elongated, slightly wavy from midway and closing between teeth rows (Figs 2D–F, 3A, B). Abdomen of similar colour as female, but much slimmer (Fig. 2B, C). Epiandrous field much wider than high and curved, with a narrow division, and bearing thirteen spigots on each side (Fig. 3H). Pedipalps with median-sized cymbium, around the same size as the rounded, narrow tibia (Figs 2H–J, 3D); tegulum about 1.5 wider than long, spherical and inflated (Figs 2H, 3D, E); conductor larger mid-way, ribbon-like, and twisted, with thick edges, enfolding the embolus as a pouch, tapering towards its hook-like tip (Figs 2H, I, 3D–F); embolus thick, heavily sclerotized, S-shaped from mid-way, originating near cymbium at middle portion of bulb and opening from below conductor on a curved tip (Figs 2H, I, 3D–F); paracymbium 3.1× longer than wide, boomerang-shaped, slanted, and tapering towards its excavated notch at apex, with translucent lobe occupying little less than 50% of paracymbium length and reaching both its basis and apex, narrow and medially placed, and knob not projected and elbow-like (Figs 2J, 3G).

Measurements. Total length 6.40. Carapace 2.0 long, 1.2 wide. Abdomen 4.5 long, 1.0 wide. Left chelicera 2.6 long, 0.5 wide. Leg formula I–IV–II–III. Leg I: femur 7.4, patella 0.8, tibia 7.8, metatarsus 8.9 and tarsus 1.2. Leg II: patella + tibia 5.1. Leg III: patella + tibia 1.9. Leg IV: patella + tibia 4.5.

Females (n = 8): total length, 9.24 – 12.50; males (n = 6): total length, 6.4 – 9.9. Two different variations are noticeable in the median cusp of some specimens of T. cladognatha. The median cusp is absent in specimens collected in the highlands in Minas Gerais state (IBSP 220017) (Fig. 4A–D), while it is in a different position in the specimens from Botucatu (MZUSP 62036, MZUSP 62396), in comparison to the neotype herein described, more basally located and apart from the inner cusp by a straight slanted portion of the fang (Fig. 4E, F). These variations on the cheliceral median cusp may corroborate Levi’s observation (1981) on intraspecific chelicerae variations, but they may also represent small, isolated populations on the verge of speciation.

Figure 4. 

Tetragnatha cladognatha female, variation. A–D. Left chelicera without MC (IBSP 220017); A. Upper view; B. Inner view; C. Lower view; D. Outer view; E–F. Chelicerae fangs with longer area from basis to MC and shorter and more slanted area between MC and IC (MZUSP 62396); E. Both chelicerae ventral view; F. Left chelicera inner view. Scale bars: 1 mm.

The holotype female of T. cladognatha from Rio de Janeiro city is considered presumed lost as all Bertkau’s spider types are, because none of them have ever been found in European collections. Considering that the distribution of T. cladognatha and T. argentinensis overlap in Brazil, with specimens of both species in Rio de Janeiro state (the type-locality of T. cladognatha), we consider the designation of a neotype necessary to clarify the taxonomic status of this species and therefore avoid possible misidentifications with T. argentinensis. Even though the original illustrations depicting the median cusp (Bertkau 1880, figs 27, 27a) help to identify the species, the precise identification of T. cladognatha was only possible after the examination of a large number of specimens from its type-locality, where it is much more commonly collected than T. argentinensis.

Mature males and females of T. cladognatha were collected in all months except June, but with much fewer specimens in the winter (also July and August). There seems to occur some plasticity in the life cycle of this species, despite a large percentage of mature specimens collected during late summer/early spring. Original labels of the specimens and our personal observations in the field suggest an affinity of this species with water courses, as it was hand collected along rivers as informed by the original labels, “manually collected in Camorim river” or in insect traps “malaise traps” and “light traps”.

From Central (Mato Grosso do Sul state) and south-east (Espírito Santo state) Brazil to north-east Argentina (Misiones province) (Fig. 5).

Figure 5. 

Distribution of T. cladognatha.

Holotype male, Sipapo River and Orinoco River confluence (05°04'04.7"N, 67°47'37.4"W, Amazonas, Venezuela), 27.xii.2002, OM Villarreal leg. (MNRJ 1571).

The male of this new species is similar to T. tenuissima considering abdomen, chelicerae and pedipalp morphology. Both share a slender abdomen, laterally with five black patches, chelicerae with centrally placed ‘a’, large and bulky Gu apart from the extremely elongated ‘T’ (visible in lower view) by very large gap and pedipalps with filiform embolus not enfolded by the conductor and small triangular paracymbium (Figs 6A–I; Castanheira and Baptista 2020, figs 12A–I, 14A, B, E–G, 15A). However, T. amazonica sp. nov. can be easily differentiated from T. tenuissima by chelicerae with much thinner ‘a’, absent ‘sl’, ‘T’ with curved tip, presence of CRu and CRl, Gl straight and apart from L2 by much larger gap and pedipalps with embolus apart from conductor since its middle portion and longer paracymbium with notch longer and distalward projected (Fig. 6D–J).

Figure 6. 

Tetragnatha amazonica sp. nov. male (MNRJ 1571). A. Dorsal habitus; B. Lateral habitus; C. Ventral habitus; D–G. Left chelicera; D. Upper view; E. Inner view; F. Lower view; G. Outer view; H–J. Left pedipalp; H. Mesal view; I. Dorsal view; J. Ventral view (paracymbium). Scale bars: 2 mm (A–C); 0.5 mm (D–J).

Male (based on holotype MNRJ 1571): Carapace elongated, slightly elevated anteriorly and light brown (Fig. 6A, B). Labium longer than wide and reddish brown (Fig. 6C). Sternum oval and light brown, with no marks or contour (Fig. 6C). Eyes with ALE and PLE touching (Fig. 6A). Legs very elongated and yellow, legs I and II slightly darker (Fig. 6A–C). Chelicera paturon approximately 4.3× longer than wide and 1.15× smaller than carapace, heavily curved outwards, around 55° from body median line, very thick and with orange hue (Fig. 6A, D–G). ‘a’ tubular, with small dent on inner margin of its tip and located in midway position of paturon (Fig. 6D, E, G). AXu and ‘t’ absent (Fig. 16D, E). Upper row with eight teeth distalward projected (Fig. 6D, E): Gu very long and thick, with very broad basis and apart from ‘T’ by a conspicuous upper crest (CRu) on large gap; ‘sl’ absent; ‘T’ conspicuous and elongated, sclerotized, and pointed with a curved tip; ‘rsu’ composed of six pointed teeth, decreasing in size and apart by regular small gaps. AXl absent (Fig. 6E, F). Lower row with eight teeth (Fig. 6E, F): Gl triangular, very thick, sclerotized, straight and bearing rounded blunt tip, apart from L2 by large gap following extremely sclerotized dark lower crest (CRl), that goes slightly beyond L2; L2 also very thick, sclerotized, and straight (or slightly basalward projected) with large tip and apart from L3 by smaller gap than Gl–L2; ‘rsl’ composed of extremely reduced teeth, apart from one another by small gaps. Cheliceral fang thick, uniformly tapering to its tip, slightly wavy from midway and closing between teeth rows (Fig. 6D–F). Abdomen approximately 2.6× longer than carapace, slender, and pale beige, covered by sparse guanine crystals with five dusky dark patches on each size of dorsum (Fig. 6A–C). Pedipalps with extremely elongated cymbium, around 2.9× longer than wide and rounded tibia, bearing wide basis and no apical constriction (Fig. 6H–J); tegulum approximately 1.4× wider than long and inflated (Fig. 16H); conductor elongated, ribbon-like and distally wider, not enclosing embolus and ending in broad twisted tip (Fig. 6H, I); embolus filiform, basally thicker and enlarged, ventrally bending from mid-way and resting its slender tip on conductor (Fig. 6H, I); paracymbium short, only 2.3× longer than wide, with transparent lobe occupying around 1/3 of its width, notch rounded and undivided, and knob large and triangular pointed (Fig. 6J).

Measurements. Total length 9.4. Carapace 2.1 long, 1.1 wide. Abdomen 7.3 long, 1.1 wide. Left chelicera 2.2 long, 0.4 wide. Leg formula I–II–IV–III. Leg I: femur 8.0, patella 0.8, tibia 8.3, metatarsus 9.2 and tarsus 1.9. Leg II: patella + tibia 4.9. Leg III: patella + tibia 1.6. Leg IV: patella + tibia 4.9.

Female. Unknown.

The specific epithet “amazonica” means “from Amazon” in Latin, referring to the type-locality in Venezuela and the Amazon Forest biome.

Only known from type-locality in Amazonas, Venezuela (Fig. 19).

The single male holotype was collected in the summer (December). No information about habitat preferences were given on the original label.

Holotype male, Lebon Régis (26°37'45.0"S, 54°06'48.0"W, Santa Catarina, Brazil), 15.vii.2006, R Lignau leg. (MCTP 43332). Paratypes: Argentina – Misiones • 1 male, 4 juveniles, San Pedro, Parque Provincial Cruce Caballero, 26°28'S, 53°58'W, 13–16.i.2005, C Grismado et. al. leg. (MACN-AR 31756); Brazil, São Paulo • 1 male, Onda Verde, Fazenda São João, 20°36'50.0"S, 49°17'56.0"W, I.1949, F Lane leg. (MZUSP 11408).

Brazil – Rio Grande do Sul • 1 male, Cachoeira do Sul: Cordilheira, 30°13'S 52°50'W, 09.ix.1992, RG Buss leg. (MCTP 43334); • 2 males, 2 juveniles, same locality, 14.xii.1992, RG Buss leg. (MCTP 43333); • 1 male, same locality, Capanezinho, 30°18'S 52°59'W, 17.x.1992, RG Buss leg. (MCTP 3375); • 2 males, Caxias do Sul, Fazenda Souza, 29°10'04.0"S, 51°10'44.0"W, 11–12.x.1995, Eq. Lab Aracnologia leg. (MCTP 7313); • 1 male, Itaara, 29°36'36.0"S, 53°45'54.0"W, 05.i.2006, L Indrusiak leg. (MCTP 21569); • 1 male, same locality, 16.ii.2006, L Indrusiak leg. (MCTP 21570); • 4 males, 2 juveniles, same locality, 28.iii.2006, L Indrusiak leg. (MCTP 21571); • 1 male, Novos Cabrais, Parque Witeck, 29°44'06.0"S, 52°56'52.0"W, 11.ix.2008, RG Buss leg. (MCTP 28092); • 1 male, same locality, 01.xi.2008, RG Buss leg. (MCTP 28013); • 1 male, Pelotas, Capão do Leão, 31°46'19.0"S, 52°20'34.0"W, 27.xi.2000, ENL Rodrigues leg. (MCTP 11711); • 1 male, same data (MCTP 11727); • 1 male, same locality, 27.ii.2001, ENL Rodrigues leg. (MCTP 13180); • 4 males, 2 juveniles, Santa Maria, 29°41'02.0"S, 53°48'25.0"W, 15.x.1998, CB Kotzian and L Indrusiak leg. (MCTP 40632); São Paulo • 1 male, 3 juveniles, Itú, Fazenda Pau D’Alho, 23°15'50.0"S, 47°17'56.0"W, 17–18.ix.1960, P Biasi leg. (MZUSP 14747);

Tetragnatha cristata sp. nov. most resembles T. oncognatha sp. nov. and T. jaculator by its similar small cylindrical body and small chelicerae, a small ‘a’ and long distalward projected ‘T’ (Figs 7A–E, G, 8A, 14A–E, G, 16A; Castanheira and Baptista 2021a, figs 56, 57, 59, 60, 62, 74). Tetragnatha cristata sp. nov. is easily differentiated by chelicerae with indented ‘a’, no AXu or AXl, much smoother rounded cheliceral bulge, and higher Gl, with large sclerotized lower crest not reaching L2, embolus dorsally opening on a rift on the conductor tip and the paracymbium with wider lobe and smaller notch (Figs 7D–J, 8A–E).

Figure 7. 

Tetragnatha cristata sp. nov. male holotype (MCTP 43332). A. Dorsal habitus; B. Lateral habitus; C. Ventral habitus; D–G. Left chelicera; D. Upper view; E. Inner view; F. Lower view; G. Outer view; H–J. Left pedipalp; H. Mesal view; I. Dorsal view; J. Ventral view (paracymbium). Scale bars: 2 mm (A–C); 0.2 mm (D–J).

Figure 8. 

Tetragnatha cristata sp. nov. male SEM photos. A, B. Left chelicera (MCTP 21570); A. Upper view; B. Lower view; C–E. Left male pedipalp (MCTP 43334); C. Mesal view; D. Tip of conductor and embolus opening detail, dorsal view; E. Paracymbium detail, ventral view; G. Epiandrous field detail, ventral view (MCTP 43334). Scale bars: 0.2 mm (A, B); 0.1 mm (C.); 0.05 mm (D, E); 0.02 mm (F).

Male (holotype MCTP 43332): Carapace oval and light brown, with two slightly darker thin parallel lines from cephalic furrow, passing through the fovea, and reaching posterior rim of carapace (Fig. 7A). Labium subquadrate and dark brown (Fig. 7C). Sternum light brown with dusky strikes (Fig. 7C). Eyes with ALE and PLE touching each other (Fig. 7A). Legs very elongated, yellowish brown with anterior pairs darker (Figs 7A–C). Chelicerae paturon about 3× longer than wide and around 1.6× smaller than carapace, moderately curved outwards, around 35° from body median line, moderately thick and orange-brown, bearing a smooth conspicuous bulge (CB) between teeth rows (Figs 7A, D–G, 8A, B). ‘a’ very short, rounded and distalward projected, deeply dented on its inner margin from its middle up to tip (Figs 7D, E, 8A). AXu absent (Figs 7D, E, 8A). Upper row with six uneven teeth distalward projected (Figs 7D, E, 8A): Gu thick, pointed, slanted and displaced from fang furrow and row itself, apart from ‘T’ by large gap formed by a cheliceral bulge; ‘sl’ absent; ‘T’ straight and pointed, with narrow basis and ‘rsu’ with four almost straight pointed teeth, decreasing in size, with the last one much smaller than the others. AXl absent (Figs 7E, F, 8B). Lower row with two teeth of same size (Figs 7E, F, 8B): Gl not very elongated, thick, sclerotized and completely pointing distalward and slightly downward, being apart from L2 by a large, straight, and very sclerotized conspicuous lower crest (CRl); L2 thin, pointed and distalward projected, with sclerotized basis. Cheliceral fang as wide as its base, moderately thick, and closing between ‘T’ and U3 (Figs 7D–F, 8A, B). Abdomen around 2× longer than carapace, cylindrical and beige, dorsally covered by guanine crystals, and bearing lateral brown line (Fig. 7A–C). Epiandrous field wide and flat, with large distal part, and bearing six and seven spigots respectively, in two bands apart by a broad midway division (Fig. 8F). Pedipalps with elongated cymbium, distally bending outwards, around 2.2× longer than wide and rounded tibia (Figs 7H, I, 8C); tegulum around 1.6× longer than high, spherical and inflated (Figs 7H, I, 8C); conductor thin, twisted near mid-way, with thick edges wrapping around embolus as a thick pouch on its median portion, and projected as a keel near tip (Figs 7H, I, 8C, D); embolus thick, originating at the middle portion of the bulb, near the cymbium and dorsally opening in middle of the conductor wrap (Figs 7H, I, 8C, D); paracymbium very elongated, around 2.8× longer than wide, boomerang-shaped, and bearing a notch with curved tip, translucent lobe occupying around half of the paracymbium width, and knob not projected (Figs 7J, 8E).

Measurements. Total length 5.1. Carapace 1.5 long, 0.9 wide. Abdomen 3.7 long, 0.8 wide. Left chelicera 0.9 long, 0.3 wide. Leg formula I–II–IV–III. Leg I: femur 4.7, patella 0.6, tibia 5.1, metatarsus 5.3 and tarsus 1.4. Leg II: patella + tibia 2.9. Leg III: patella + tibia 1.2. Leg IV: patella + tibia 1.6.

Female. Unknown.

The specific Latin epithet “cristata” is an adjective meaning “with a crest or ridge”, and refers to the large crest between the two first teeth at the lower row of the male cheliceral furrow.

Males (n = 8): total length, 4.16 – 5.79. Males can have three additional lower teeth in the chelicerae, decreasing in size after L2 (e.g., Fig. 8B, MCTP 21570).

The distribution of this species ranges from São Paulo state to Rio Grande do Sul state, both in Brazil, passing through Misiones, Argentina (Fig. 9).

Figure 9. 

Distribution of T. cristata sp. nov.

Mature specimens of T. cristata sp. nov. were collected in the hottest months of the year: January, February, March, September, October, November and December, pointing to a possible early summer/late spring maturity. Only one specimen was collected in July, during winter. No information on habitat preferences was provided in the original labels.

Holotype male, Torres (29°20'06.0"S, 49°43'37.0"W, Rio Grande do Sul, Brazil), 26.iii.2006, ELC da-Silva leg. (MCTP 43335). Paratypes: Brazil – Pará • 1 male, Belém, Reserva Mocambo, 01°26'28.7"S, 48°24'46.2"W, 05.v.2008, BVB Rodrigues leg. (MPEG.ARA 031362); Paraná • 1 male, Cambará, 23°02'45.0"S, 50°04'26.0"W, iv.2011, AM Giroti leg. (IBSP 167755); Santa Catarina • 1 male, Lebon Régis, 26°55'44.0"S, 50°41'42.0"W, 15.vii.2006, R. Lignau leg. (MCTP 19580).

Males of Tetragnatha didorata sp. nov. resembles T. tenuissima and T. amazonica sp. nov. considering the elongated slender abdomen; T. bogotensis considering cheliceral morphology as both share long and straight ‘t’, and T. renatoi sp. nov. considering their pedipalp morphology with large subquadrate paracymbium (Fig. 6A–C, 10A–F, J, K, Castanheira et al. 2019, figs 1C, E, F, 3C, Castanheira and Baptista 2020, figs 4I, 6F, 12A, B). Tetragnatha didorata sp. nov. differs from T. tenuissima and T. amazonica sp. nov. by the presence of a projection after the spinnerets (Figs 10B, C), from T. bogotensis by ‘a’ not slanted and carved on its lower portion, absence of AXu, the more elongated and sclerotized ‘t’, Gu and L2 smaller, absent AXl and much smaller teeth on lower row (Figs 10D–G), and from T. renatoi sp. nov. by the narrower translucent lobe, larger notch and larger knob (Fig. 10J, K).

Figure 10. 

Tetragnatha didorata sp. nov. male (MCTP 43335). A. Dorsal habitus; B. Lateral habitus; C. Ventral habitus; D–G. Left chelicera; D. Upper view; E. Inner view; F. Lower view; G. Outer view; H–J. Left pedipalp; H. Mesal view; I. Dorsal view; J. Ventral view (paracymbium); K. Paracymbium detail, ventral view. Scale bars: 2 mm (A–C); 0.5 mm (D–J); 0.2 mm (K).

Male (holotype MCTP 43335): Carapace elongated, slightly elevated anteriorly and with orange hue (Fig. 10A, B). Labium longer than wide and yellowish brown (Fig. 10C). Sternum oval and yellowish brown, with no marks or contour (Fig. 10C). Eyes with ALE and PLE almost touching (Fig. 10A). Legs very elongated, with yellowish hue, with legs I and II slightly darker (Fig. 10A–C). Chelicera paturon very elongated, approximately 4.95× longer than wide and as long as carapace, slightly curved outwards, around 25° from body median line, moderately thick and yellowish brown (Fig. 10A, D–G). ‘a’ elongated, thin, straight distalward projected and constricted on inner margin of its tip, located close to fang groove on centre of paturon (Fig. 10D–G). AXu absent (Fig. 10D, E). ‘t’ bulky, sclerotized, pointed and straight, forming the letter “L” with the apophysis (Fig. 10D, E). Upper row with eight teeth (Fig. 10D, E): Gu small, sclerotized and slightly distalward projected, located on a dark ridge apart from fang groove by large gap; U2–U7 of similar size, pointed and straight, U2–U3 apart by a large gap and remaining teeth apart from one another by small gaps. AXl absent (Fig. 10E, F). Lower row with nine teeth distalward projected (Fig. 10E, F): Gl small, triangular, pointed and sclerotized, located on fang groove and apart from L2 by very large gap; L2–L4 pointed and decreasing in size, apart by large gaps; L5–L9 extremely reduced in size, L7–L9 reduced to denticles. Cheliceral fang not thick, uniformly tapering to its tip and closing between teeth rows (Fig. 10D–F). Abdomen slender, approximately 2.7× longer than carapace, with a pointed end projection posterior to the spinnerets, pale yellow and completely covered by guanine crystals, with no lateral bands (Fig. 10A–C). Pedipalps with elongated cymbium, only around 1.2× longer than long and rounded tibia, medially bending outwards, bearing wide basis and with no apical constriction (Fig. 10H, I); tegulum about 1.4× wider than long and inflated (Fig. 10H); conductor elongated, distally projected and completely enfolding the embolus from its middle portion as a pouch, ending in rounded and twisted tip (Fig. 10H, I); embolus thick, filiform, sclerotized and twisted from mid-way, opening from below the conductor (Fig. 10H, I); paracymbium very long, around 3.3× longer than wide, subquadrate, bearing transparent lobe that occupies around 50% of paracymbium width, notch carved and rounded, and knob enlarged and mushroom-like (Fig. 10J, K).

Measurements. Total length 9.0. Carapace 2.5 long, 1.4 wide. Abdomen 6.6 long, 1.0 wide. Left chelicera 2.4 long, 0.6 wide. Leg formula I–II–IV–III. Leg I: femur 7.1, patella 0.9, tibia 6.8, metatarsus 7.5 and tarsus 1.6. Leg II: patella + tibia 4.4. Leg III: patella + tibia 1.7. Leg IV: patella + tibia 4.9.

Female. Unknown.

The specific epithet “didorata” is the compound latinized form of the Greek words: “di” meaning “two” and “dory, dorata” meaning “spear”, referring to the spear-like shape of the thin and long apophysis and the elongated and pointed ‘t’ at the upper side of the male chelicerae.

Males (n = 3): total length, 7.0 – 9.0. Little variation in colour pattern.

This species is mainly found at Paraná, Santa Catarina and Rio Grande do Sul, in the South region, but it was also collected in Pará state, in the North region, all in Brazil (Fig. 19).

All specimens of T. didorata sp. nov. were collected during the autumn (late March, April and May) and the beginning of winter (July). No information on habitat preferences was given on the original labels of this species.

Argentina – • Tetragnatha americana: one female, Río Santa Cruz, Santa Cruz, Patagonia (MNHN 3140). Chile: 6 males, 2 females (T. americana: Simon det., MNHN 12628).

See the section on T. nitens for additional information. Some of the specimens Simon (1896, 1905) identified as T. americana belong actually to T. laboriosa. We were able to analyse several specimens from Chile (MNHN 12628) and just one of the three females from Santa Cruz, Argentina (MNHN 3140, fig. 11) cited by Simon (1905) and confirmed that they are all typical T. laboriosa.

Figure 11. 

Tetragnatha laboriosa under T. americana, female (MNHN 3140). A. Dorsal habitus; B. Original label citing Santa Cruz, Argentina; C. Ventral habitus; D. Left chelicera, inner view; E. Chelicerae, apical view.

Tetragnatha americana: Unspecified number of syntypes from Peñaflor, Santiago, Chile, Lataste Coll., MNHN?, MHNS?, presumed lost. Tetragnatha labialis: Unspecified number of adult female syntypes from Santiago, Chile, Nicolet Coll. (female MNHN 4209, herein designated as lectotype), examined.

Chile – • 1 male from Punta Arenas (T. americana: Simon det., MNHN 22312), examined.

Tetragnatha americana was a new name Simon (1896, p. CV–CVI) applied to a species he considered as the same one misidentified as T. extensa and redescribed by Nicolet (1849, p. 516). He only added the following expression in a list of spiders from Chile assembled by F. Lataste: “Tetragnatha americana E. Sim. (= extensa Nicolet)”. Those specimens were collected in Peñaflor, Santiago, Chile, and deposited at the “Museo Zoolojico de la Escuela de Medicina”, currently the Museo Nacional de Historia Natural, Chile (MHNS, see Díaz (2019) for additional information). Simon (1905, p. 10) applied again without any justification the name T. americana to the species he considered as T. extensa sensu Nicolet, citing three females from Santa Cruz, Argentina. He listed T. extensa sensu Nicolet under the title of Tetragnatha americana, and included the bibliographical reference to the book, again without mentioning the examination of Nicolet’s specimens or any morphological character. Simon (1905) added that the specimens he had from Santa Cruz “were not different from T. americana of Chile, where the species is very common”.

Despite Simon (1896, 1905) not clearly stating that he intended to add a new replacement name for T. extensa sensu Nicolet, no previous citation to T. americana was found after an analysis of Simon’s papers dealing with spiders from Chile. Following Bonnet (1959, sub “T. americana Simon, 1897”, p. 4317) and against Roewer (1942, p. 988) and the World Spider Catalog (2022), we consider 1896 and not 1905 as the year of the proposition of the new name T. americana. The citation by Simon (1896, p. CV) was apparently the first appearance of the name and it is available under the International Code of Zoological Nomenclature (ICZN 1999), following its article 12, which establishes that an unambiguous reference to a species’ description suffices to make available a name published before 1931.

The type-series of T. americana Simon, 1896 is therefore composed of the specimens from Peñaflor examined by Simon (1896) when proposing the new name and also the specimens from Valdivia province studied and illustrated by Nicolet (1849), but not by the specimens from Santa Cruz, Argentina (Fig. 11) and unspecified localities of Chile cited by Simon (1905). According to the article 72.4.1.1 and its example, all the specimens studied by the species’ author or referred unambiguously to in the original description constitute the type series. As stated by Levi (1967, 2001), Nicolet probably returned to Paris in 1846 and brought with him a large part of the types of Chilean species he described. Levi was able to find at MNHN types of many species of Theridiidae, but he considered most Araneidae types as lost. We also have not been able to find most Tetragnathidae types of species described by Nicolet in MNHN, excepting T. labialis. We requested information about Tetragnatha specimens from the MHNS but did not receive an answer and there is no evidence that any of Nicolet or Simon’s specimens are still available in this institution. Because T. labialis was found in MNHN, we consider that this was probably the same depository institution for the other species described by Nicolet (1849) and the specimens from Valdivia province described as T. extensa. However, except for T. labialis, no other original specimen from Nicolet (1849) was found at MNHN. Likewise, T. americana specimens identified by Simon from Peñaflor and Santiago were also not found in the general collection of the MNHN. Therefore, we consider the syntypes of T. americana presumed lost.

Nicolet’s description focused on the shape and size of structures such as carapace, eyes, abdomen, chelicerae and pedipalps, but did not include precise details on chelicerae and genital morphology to allow species identification. Fortunately, he also provided relatively good drawings (Nicolet 1849, plate 4, fig. 5), which clearly depict a female specimen of T. nitens, based for example on the sinuous cheliceral fang, Gu and U2 large and of similar size and well-spaced, and AXl large but not overreaching the fang basis. So, the description and illustrations by Nicolet (1849) are the only reliable information on the type series we recovered.

We were able to analyse all the specimens currently present in the MNHN collection and identified as T. americana by Simon. Some of these are specimens of T. laboriosa, represented by several specimens from Chile (MNHN 12628) and just one of the three females from Santa Cruz, Argentina (MNHN 3140, Fig. 11) cited by Simon (1905). On the other hand, there is a male from Punta Arenas, Chile, that is clearly T. nitens (MNHN 22312, Fig. 12). Although these specimens are just additional material examined, they were identified by Simon himself, thus helping to establish his concept of T. americana. Although some specimens Simon identified as T. americana are in fact T. laboriosa, the original drawings of a female syntype of “T. extensa” from Nicolet and secondarily the male from Punta Arenas (MNHN 22312, Fig. 12) identified by Simon allow us to indicate that Tetragnatha americana Simon, 1896 is Tetragnatha nitens (Audouin, 1826) syn. nov.

Figure 12. 

Tetragnatha nitens under T. americana, male. A. Left chelicera, inner view; B. Chelicerae, slanted lower view.

As mentioned above, the only species dealt with by Nicolet (1849) that we were able to locate at MNHN was T. labialis, described based on an unspecified number of female syntypes from Santiago, Chile. A female specimen (MNHN 4209), labelled as “Tetragnatha labialis Nicol. Gay Saint Iago” (Fig. 13) was examined. This female clearly belongs to the syntype series examined by Nicolet and is herein designated as lectotype. Following Castanheira et al. (2019), this specimen is clearly T. nitens, as shown for example by the following diagnostic cheliceral characters: Gl distalward followed by smaller L2 and L3, AXl reduced and point distalward, Gu elongated and not contiguous to U2 and robust basal cusp placed at the lower side of fang (Fig. 13A–C). Therefore, we propose Tetragnatha labialis Nicolet 1849 = Tetragnatha nitens (Audouin, 1826) syn. nov. Additionally, the male and female specimens from Nova Granada (probably Colombia) named T. labialis by Keyserling (1865) are misidentified. The original drawings of the chelicerae resemble T. argentinensis (Keyserling 1865, fig. 11–13) or perhaps some related, undescribed species from Colombia. Furthermore, the original locality (Nova Granada) is too far north of the currently known T. argentinensis distribution range (Cargnelutti et al. 2022). The Chilean specimens of T. labialis cited by Simon (1902, 1904) from Valdivia and Punta Arenas, respectively, were not found at MNHN. Only a male from T. americana from Punta Arenas was located as we explained above (Fig. 12).

Figure 13. 

Tetragnatha nitens under T. labialis, female lectotype (MNHN 4209). A. Left chelicera, upper view; B. Left chelicera, inner view; C. Left chelicera, lower view; D. Original label.

Holotype male, Rancho Queimado (27°40'22.0"S, 49°01'19.0"W, Santa Catarina, Brazil), 09–13.x.1995, AA Lise leg. (MCTP 7067). Paratypes: Brazil, Rio de Janeiro • 1 male, 1 juvenile, Itatiaia, Parque Nacional de Itatiaia, 22°29'29.0"S, 44°33'33.0"W, 20–22.xi.2008, DT Castro leg. (UFRJ 0423); Rio Grande do Sul • 1 male, Guaíba, Fazenda São Maximiliano, 30°06'50.0"S, 51°19'30.0"W, 14.vii.1995, AA Lise leg. (MCTP 6688); • 1 male, 1 female, São Francisco de Paula, 29°26'52.0"S, 50°35'02.0"W, 05–09.iii.1997, AA Lise leg. (MCTP 15965).

Brazil – Rio Grande do Sul • 1 female, Novos Cabrais, 29°44'06.0"S, 52°56'52.0"W, 27.i.2010, RG Buss leg. (MCTP 43654); • 1 male, São Francisco de Paula, Potreiro Velho, 29°24'47.9"S, 50°15'36.8"W, 12–14.xi.1998, AA Lise leg. (MCTP 12037); • 1 male, same locality, 04–06.i.1999, AA Lise leg. (MCTP 15792); • 2 males, 2 females, same locality, 14–17.xii.1996, AA Lise leg. (MCTP 43336); • 1 male, Santo Antônio da Patrulha, 29°49'04.0"S, 50°31'12.0"W, 27.viii.1994 (MCTP 4929); • 1 male, Viamão, 30°04'51.0"S, 51°01'22.0"W, 07.xi.1995, AA Lise et. al. leg. (MCTP 43337); • 1 male, Viamão, Itapuã, , 30°17'02.7"S, 51°01'23.3"W, 29.v.2004, BP Zambonato leg. (MCTP 33638); Santa Catarina • 1 male, Florianópolis: Morro das Aranhas, Costão do Santinho, 27°28'05.9"S, 48°22'49.8"W, 2007, F Albertoni leg. (IBSP 144176); São Paulo • 1 female, São Paulo, Parque Ilha dos Eucaliptos, Jardim Ângela, Reservatório de Guarapiranga, 23°44'00.0"S, 46°44'01.5"W, 07–13.x.2003, I Cizauskas and CRM Garcia leg. (IBSP 61361).

Males of T. oncognatha sp. nov. share with T. cristata sp. nov. and T. jaculator similar small-sized cylindrical bodies and small chelicerae, with ‘a’ small sized and ‘T’ long distalward projected (Figs 7A–E, G, 8A, 14A–E, G, 16A; Castanheira and Baptista 2021a, figs 56, 57, 59, 60, 62, 74). However, T. oncognatha sp. nov. is easily identified by chelicerae with small AXu, much more conspicuous and rounded cheliceral bulge, and deep lower crest beginning before Gl and ending after L2, embolus opening inside ventral portion of conductor and paracymbium with narrower lobe and longer notch (Figs 14D–F, H–J, 16A, B, E–G). Females are similar to T. guatemalensis in regard to chelicerae and genitalia, but T. oncognatha sp. nov. is distinguished by chelicerae with much smaller U2 and L2, much larger gaps between Gl and L2 and between L2 and L3, no outer cusp, more rounded genital fold and internal genitalia formed by much larger central sac and two much more rounded spermathecae (Figs 15D–I, 16C, D).

Figure 14. 

Tetragnatha oncognatha sp. nov. male holotype (MCTP 7067). A. Dorsal habitus; B. Lateral habitus; C. Ventral habitus; D–G. Left chelicera; D. Upper view; E. Inner view; F. Lower view; G. Outer view; H–J. Left pedipalp; H. Mesal view; I. Dorsal view; J. Ventral view (paracymbium). Scale bars: 2 mm (A–C); 0.2 mm (D–J).

Figure 15. 

Tetragnatha oncognatha sp. nov. female paratype (MCTP 15965). A. Dorsal habitus; B. Lateral habitus; C. Ventral habitus; D–G. Left chelicera; D. Upper view; E. Inner view; F. Lower view; G. Outer view; H–I. Female genitalia; H. Genital fold, ventral view; I. Internal genitalia, dorsal view. Scale bars: 2 mm (A–C); 0.2 mm (D–I).

Figure 16. 

Tetragnatha oncognatha sp. nov. SEM photos. A, B. Male left chelicera; A. Upper view (MCTP 15965); B. Lower view (MCTP 15792); C, D. Female left chelicera (MCTP 43654); C. Upper view; D. Lower view; E–G. Left male pedipalp; E. Mesal view (MCTP 4929); F. Tip of conductor and embolus opening detail, ventral view (MCTP 15792); G. Paracymbium detail, ventral view (MCTP 15792); H. Epiandrous field detail, ventral view (MCTP 15965). Scale bars: 0.2 mm (A–D); 0.1 mm (E); 0.05 mm (F, G); 0.02 mm (H).

Male (holotype, MCTP 7067): Carapace elongated and, tapering toward slightly elevated anterior part, with rounded borders and yellowish brown colour hue with thin dark line rebordering its edges (Fig. 14A, B). Labium subquadrate and dark brown (Fig. 14C). Sternum oval and light brown, with no marks or contour (Fig. 14C). ALE and PLE touching each other (Fig. 14A). Legs yellowish brown with four pairs of spines on femur (Fig. 14A–C). Chelicerae paturon approximately 3.3× longer than wide and 1.6× shorter than carapace, moderately curved outwards, around 30° from body median line, moderately thick and with orange-brown colour hue, bearing a large and rounded conspicuous bulge (CB) between teeth rows (Figs 14A, D–G, 16A, B).‘a’ short, thick, out- and distalward projected, continuously bent from its basis, except in its last third, which bears a more abrupt curvature; with a carved tip, and located in middle portion of paturon (Figs 14D, E, 16A). AXu very reduced, almost a nub, located on fang furrow (Figs 14D, E, 16A). Upper row with six uneven teeth distalward projected (Figs 14D, E, 16A): Gu small, thick, pointed, slanted and displaced from fang furrow and row itself, apart from ‘T’ by a large gap formed by CB; ‘sl’ absent; ‘T’ elongated, thick, and very pointed, with very large basis; ‘rsu’ with four almost straight pointed teeth decreasing in size. AXl absent (Figs 14E, F, 16B). Lower row with seven teeth distalward projected (Figs 14E, F, 16B): Gl elongated, thick, finger-like and located on edge of fang furrow, alongside L2 on soft lower crest (CRl); L2 and L3 pointed, L2 more sclerotized and apart from L3 by a conspicuous gap, partially covered by CRl and L3 apart from L4 by a gap of same size; L4–L7 very pointed, L4 with almost same size as L6, L5–L7 decreasing in size, with L7 very reduced. Cheliceral fang moderately thick, and closing between teeth rows (Figs 14D–F, 16A, B). Abdomen medium-sized, around 1.9× longer than carapace, cylindrical and beige, dorsally covered by guanine crystals, bearing a lateral dark line (Fig. 14A–C). Epiandrous field oval and flat, with wide distal part, bearing six and eight spigots in two bands apart by a large midway division (Fig. 16H). Pedipalps with elongated cymbium, around 2× longer than short, distally bending laterad, with rounded tibia (Figs 14H, I, 16E); tegulum around 1.3× wider than long, spherical and inflated (Figs 14H, I, 16E); conductor twisted, only medially thicker and sclerotized, with wide tip completely enfolding embolus as a wide and projected pouch, which projects beyond embolus tip as a cap (Figs 14H, I, 16E, F); embolus short, thick and sinuous, originating at the middle portion of the bulb, near the cymbium (Figs 14H, I, 16E, F); paracymbium around 3.4× longer than wide, triangular, thick and moderately downward slanted at its tip, with a finger-like notch, translucent lobe occupying around 40% of the paracymbium length, neither reaching its basis, nor its apex, knob large and not projected (Figs 14J, 16G).

Measurements. Total length 6.1. Carapace 2.0 long, 1.3 wide. Abdomen 4.2 long, 1.1 wide. Left chelicera 1.2 long, 0.4 wide. Leg formula I–II–IV–III. Leg I: femur 2.9, patella 0.8, tibia 4.9, metatarsus 4.1 and tarsus 0.7. Leg II: patella + tibia 3.8. Leg III: patella + tibia 1.6. Leg IV: patella + tibia 2.4.

Female (paratype MCTP 15965): Carapace colour, maxillae, fovea, eyes, labium, and legs similar to male, but sternum dusky, with light brown spots between coxae (Fig. 15A–C). Chelicera paturon around 2.6× longer than wide, 1.9× shorter than carapace and bending around 25° from body median line, with similar colour hue as male (Figs 15A, D–G, 16C, D). AXu absent (Figs 15D, E, 16C). Upper row with seven teeth distalward projected (Figs 15D, E, 16C): Gu large, thick, pointed, very sclerotized and triangular, located on fang groove and apart from U2 by medium-sized gap; U2 small, with size similar to U7, triangular and pointed, apart from U3 by medium-sized gap; U3–U7 decreasing in size and pointed. AXl absent (Figs 15E, F, 16D). Lower row with six teeth distalward projected (Figs 15E, F, 16D): Gl triangular, thick and very sclerotized, located on fang groove and apart from L2 by medium-sized gap; L2 small, triangular and pointed, apart from L3 by medium-sized gap; L3–L6 decreasing in size and pointed. Cheliceral fang thick, uniformly tapering, serrated and abruptly curving from midway to its tip (Figs 15D–F, 16C, D). Abdomen slightly longer than that of male, around 2.6× longer than carapace, and bearing wider lateral black line (Fig. 15A–C). Genital fold short, around 4.7× wider than long, with a straight posterior rim and with beige colour hue (Fig. 15H). Internal genitalia with two massive globular spermathecae connected to a rounded uterus externus, and a large and oval central membranous sac, with very short stalk (Fig. 15I).

Measurements. Total length 8.5. Carapace 2.5 long, 1.4 wide. Abdomen 6.0 long, 1.4 wide. Left chelicera 1.1 long, 0.5 wide. Leg formula I–II–IV–III. Leg I: femur 4.4, patella 0.8, tibia 5.1, metatarsus 5.4 and tarsus 1.3. Leg II: patella + tibia 3.5. Leg III: patella + tibia 1.6. Leg IV: patella + tibia 3.1.

The specific Greek epithet “oncognatha” refers to the gibbous, inflated chelicerae of both sexes, and is composed of the latinized form of the Greek “onkos” meaning “tumour, swell”, and “gnatha” meaning “mouthpart, chelicera”.

Males (n = 6): total length, 4.2 – 6.2. Male L3 may be absent, increasing the length of the gap (Fig. 14E, F vs. Fig. 16B). No conspicuous colour variations between specimens was detected.

The distribution of this species ranges from Itatiaia in South-centre Rio de Janeiro state, through Rancho Queimado in Centre-east Santa Catarina state, to Guaíba in South-centre and Viamão in Northeast Rio Grande do Sul state, all in Brazil (Fig. 17).

Figure 17. 

Distribution of T. oncognatha sp. nov.

Mature males and females of T. oncognatha sp. nov. were collected in January, March, May, July, August, and October to December, therefore the species does not seem to have a specific maturity period. No information on habitat preferences was provided on the original labels.

Holotype female, São Francisco de Paula, Potreiro Velho (29°24'47.9"S, 50°15'36.8"W, Rio Grande do Sul, Brazil), 05–08.xii.1996, AA Lise leg. (MCTP 14123). Paratypes: Argentina – Buenos Aires • 2 females, Sierra de la Ventana, Hotel Provincial, 38°08'17.5"S, 61°48'02.6"W, 18.ii.1973 (MACN-AR 24550).

Tetragnatha pradoi sp. nov. seems similar to T. nitens, as both species share very elongated and bulky bodies, with abdomen much wider anteriorly, and similar chelicerae (Fig. 18A, B, Castanheira et al. 2019, fig.15A, B). Differently from T. nitens, however, T. pradoi sp. nov. has a short tail behind the spinnerets and AXu of the chelicerae is much more elongated, Gu and U2 slightly closer and fangs with an outer cusp and an inner cusp (differing from the basal cusp of T. nitens) (Fig. 18A–E; Castanheira et al. 2019, figs 15A–I, 16B). Also, the genital fold of this new species is the longest within Tetragnatha, even longer than that in T. mandibulata, identifying this species from all other in the genus (Fig. 18G).

Figure 18. 

Tetragnatha pradoi sp. nov. female. A–G. Holotype female (MCTP 14123); A. Dorsal habitus; B. Lateral habitus; C–F. Left chelicera; C. Upper view; D. Inner view; E. Lower view; F. Outer view; G–H. Female genitalia; G. Genital fold, ventral view; H. Paratype female, internal genitalia, dorsal view (MACN-AR 2455). Scale bars: 2 mm (A, B); 0.5 mm (C–F); 1 mm (G); 0.2 mm (H).

Female (based on holotype MCTP 14123): Carapace elongated, flattened and orange-brown, with two dark parallel lines from eyes, passing through fovea towards posterior rim (Fig. 18A). Labium l longer than wide and light brown. Sternum oval and light brown. AME and PME almost touching, and ALE and PLE touching (Fig. 18A). Legs very long, orange-brown (Fig. 18A, B). Chelicera paturon around 3.9× longer than wide, 1.3× shorter than carapace and around 20° from body median line, with orange-brown colour hue, (Fig. 18A, C–F). AXu elongated, pointed and distalward projected, with very large basis (Fig. 18C, D). Upper row with eight teeth distalward projected (Fig. 18C, D): Gu long and pointed, similar to AXu. but with narrow basis, and located on fang furrow, apart from U2 by a moderate gap; U2 similar to Gu and apart from it and U3 by moderate gaps of the same length; U3–U8 pointed and decreasing in size. AXl absent (Fig. 18D, E). Lower row with eleven teeth distalward projected (Fig. 18D, E): Gl elongated, pointed and sclerotized, apart from L2 by a large sclerotized gap; L2 with a moderate size, pointed and slightly displaced from the row itself, apart from L3 by a small gap; L3–L11 pointed and decreasing in size, L10 and L11 extremely reduced. Cheliceral fang thick, anteriorly enlarged and uniformly tapering to its tip, bearing a large triangular outer cusp near its basis and a small inner cusp on its first third (Fig. 18C–F). Abdomen almost 3.2× longer than carapace, cylindrical and anteriorly bulging, with beige colour hue, covered by sparse guanine crystals, and having a small tail after the spinnerets (Fig. 18A, B). Genital fold extremely elongated, around 1.7× longer than wide, narrowed midway, with a rounded posterior rim, and light brown colour hue (Fig. 18G). Internal genitalia with two rounded sclerotized and longer than wide spermathecae, connected to a large uterus externus and a cylindrical central membranous sac (Fig. 18H).

Measurements. Total length 11.1. Carapace 2.6 long, 1.5 wide. Abdomen 8.7 long, 2.1 wide. Left chelicera 1.8 long, 0.6 wide.

Male. Unknown.

The specific epithet “pradoi” is a patronym honouring André Wanderley do Prado, work-colleague and friend of the authors, who gave a great deal of support throughout the development of all manuscripts on Tetragnatha.

Tibia, metatarsus and tarsus of the legs of the holotype were missing and consequently not measured.

Females (n = 3): total length, 8.2 – 11.1. Specimens do not conspicuously vary in colouration.

From Brazil (Rio Grande do Sul state) to Argentina (Buenos Aires province) (Fig. 19).

Figure 19. 

Distribution of T. amazonica sp. nov., T. didorata sp. nov. and T. pradoi sp. nov.

The only three females of T. pradoi sp. nov. we examined were collected in February and December. No information on habitat preferences was provided on the original labels.

Nomina dubia

The types of the following species are immature or presumed lost. Therefore, we are not able to correctly recognize the species. We cannot precisely identify them, nor clearly diagnose them in relation to other Tetragnatha. We considered a species as nomen dubium whenever its type-series was lost and original illustrations and descriptions were not very diagnostic, or when immatures could not be associated with adults.

Tetragnatha bishopi: Subadult female holotype from Demerara-Mahaica, Tibicuri-Cuyahà, Guyana, x.1931, Coll. Beccari (MZUF 531), examined.

This species was described by Caporiacco (1947) based on a female subadult specimen from Guyana, which was examined by the first author during a visit to MZUF. The holotype was in bad condition, with a broken abdomen, both small immature chelicerae detached from the body and had been dried and subsequently been placed into alcohol. Therefore, this species cannot be correctly identified and is considered a nomen dubium.

Tetragnatha linearis: Adult female syntypes from Chile, Nicolet? Coll., MNHN?, presumed lost.

Tetragnatha similis: Female and adult? male syntypes from “Central Provinces”, Chile, Nicolet? Coll., MNHN?, presumed lost.

Tetragnatha sternalis: Adult males and females syntypes from several localities in Chile, especially females from Llanquihue province, Nicolet? Coll., MNHN?, presumed lost.

In the same book in which Nicolet (1849) misidentified T. nitens specimens as T. extensa and described T. labialis, he also described three other Tetragnatha from Chile: T. linearis Nicolet, 1849, T. similis Nicolet, 1849 and T. sternalis Nicolet, 1849. From these, only T. similis was accompanied by drawings, illustrating the dorsal habitus and eyes of an apparently immature male specimen, but they are not useful to correctly identify the species (see Nicolet 1849; plate 4, fig. 6). Also, as stated above, Nicolet’s description were succinct and without precise details on chelicerae and genital morphology to allow species identifications.

Sixteen years later, Keyserling (1865) provided good illustrations for specimens from Nova Granada (current Colombia) he identified as T. linearis (see Keyserling 1865; plate 21, fig. 23) and T. similis (see Keyserling 1865; plate 20, figs 21–23). However, Keyserling (page 835) pointed out that the species described in works of Walckenaer, Nicolet, Hentz etc. cannot be determined with certainty, since the descriptions have not included enough characters to provide reliable clues for determining the species. He added that he preferred to apply the old names given by those authors rather than to propose possible superfluous new names to species that could be identical to the old ones. According to his drawings, T. similis sensu Keyserling looks very similar to T. laboriosa and T. linearis sensu Keyserling looks like T. guatemalensis. However, we could not confirm the identity of these specimens as we were not able to find the type-material for these species in MNHN, the same institution housing T. labialis. We consider Tetragnatha linearis, Tetragnatha similis, and Tetragnatha sternalis nomina dubia.