Tetragnatha cladognatha Bertkau, 1880
Castanheira, Pedro de Souza, Baptista, Renner Luiz Cerqueira & Oliveira, Francisca Samia Martins, 2022, Five new species of the long-jawed orb-weaving spider genus Tetragnatha (Araneae, Tetragnathidae) in South America, with a key to the species from Argentina and Brazil, Evolutionary Systematics 6 (2), pp. 175-210 : 175
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|Tetragnatha cladognatha Bertkau, 1880|
Tetragnatha cladognatha Bertkau, 1880: 79, pl. 2, fig. 27 (♀)
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) GoogleMaps ; • 2 males, 4 females, 3 juveniles, Departamento Cainguás, Parque Provincial Salto Encantado, Arroyo Cuna-Piru , 27°07'S, 54°48'W, 12.i.2005, C Grismado, L Lopardo, L Piacentini, A Quaglino and G Rubio leg. (MACN-Ar 31757) GoogleMaps ; • 1 male, 1 female, 1 juvenile, same data (MACN-Ar 31763) GoogleMaps ; • 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) GoogleMaps ; • 1 female, same data, (MACN-Ar 24645) GoogleMaps ; • 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) GoogleMaps ; • 1 male, 2 females, 1 juvenile, Cataratas del Iguazu , 25°41'28.1"S, 54°26'43.6"W, xi.1954, BC Schiapelli leg. (MACN-Ar 39613); BRAZIL, Mato Grosso do Sul GoogleMaps • 1 male, Jaraguari, Furnas de Dionisio , 20°09'21.5"S, 54°43'34.4"W, 14.xi.2015, D Araújo leg. (IBSP 167038); Minas Gerais GoogleMaps • 3 males, 4 females, Alto Caparaó, Parque Nacional do Caparao , 20°05'S, 41°09'W, 01-07.v.2002, Equipe Biota leg. (IBSP 220017) GoogleMaps ; • 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) GoogleMaps ; • 2 females, same data, (MNRJ 1586); Paraná GoogleMaps • 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) GoogleMaps ; • 1 female, same locality, 15.xi.2015, AC Domahovski leg. (MCTP 39130) GoogleMaps ; • 1 male, same locality, 25°36'12.65"S, 49°11'33.58"W, x.2015, AC Domahovski leg. (MCTP 39055); Rio de Janeiro GoogleMaps • 1 male, 2 females, 2 juveniles, Cachoeiras de Macacu, Reserva Ecologica de Guapiacu (REGUA), river nearby accommodation, 22°27'12.4"S, 42°46'21.1"W, RLC Baptista leg. (UFRJ 1503) GoogleMaps ; • 1 female, same locality, trilha cinza, 29.viii.2019, AA Alves leg. (UFRJ 1620) GoogleMaps ; • 1 male, 3 females, Guapimirim , 22°35'25.1"S, 43°06'15.7"W (MNRJ 1572) GoogleMaps ; • 1 female, same locality, ii.1996 (MNRJ 1570) GoogleMaps ; • 3 males, 2 females, 1 juvenile, Itatiaia, Cachoeira Veu da Noiva , 1250 m, 22°25'35.6"S, 44°37'12.6"W, 06.i-03.ii.2016, M Monné leg. (MNRJ 4260) GoogleMaps ; • 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) GoogleMaps ; • 1 male, 1 female, Magé, Piabetá, 22°36'21.7"S, 43°10'36.8"W, xi.1986 (MNRJ 1574) GoogleMaps ; • 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) GoogleMaps ; • 1 female, same locality, Rio das Flores, 22°24'06.7"S, 42°29'19.4"W, BHL Sampaio leg. (UFRJ 0073) GoogleMaps ; • 1 female, Nova Iguaçu, Parque Municipal de Nova Iguacu (PMNI), 22°46'44.6"S, 43°27'31.8"W, 01.v.2004, BHL Sampaio leg. (MNRJ 07445) GoogleMaps ; • 1 male, 2 females, same locality, 28.v.2004, BHL Sampaio leg. (MNRJ 07449) GoogleMaps ; • 1 female, same locality, 10.vii.2004, BHL Sampaio leg. (MNRJ 07446) GoogleMaps ; • 2 females, same locality, 31.vii.2004, C Lima leg. (MNRJ 07444) GoogleMaps ; • 1 male, 1 female, same locality, 26.xi.2004, BHL Sampaio leg. (MNRJ 07447) GoogleMaps ; • 1 female, Petrópolis, 22°29'01.5"S 43°15'10.8"W, Mello-Leitão leg. (MNRJ 60003) GoogleMaps ; • 1 female, Pinheiral, Pinheiro , 22°30'24.3"S, 44°01'22.2"W, Mello-Leitão leg. (MNRJ 60009) GoogleMaps ; • 1 male, Resende, Vila da Fumaça, Estrada Falcao-Fumaca , 22°17'57.1"S, 44°13'07.9"W, 10.xi.2016, LBN Coelho leg. (UFRJ 1366) GoogleMaps ; • 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) GoogleMaps ; • 1 female, 1 juvenile, same data (UFRJ 1623) GoogleMaps ; • 1 female, same data (UFRJ 1624) GoogleMaps ; • 1 female same data but RLC Baptista and PdS Castanheira leg. (UFRJ 1625) GoogleMaps ; • 3 males, 1 female, 7 juveniles, same locality, 09.i.2014, RLC Baptista leg. (UFRJ 1626) GoogleMaps ; • 5 females, 4 juveniles, same data (UFRJ 1629) GoogleMaps ; • 1 female, same locality, 07.iv.2014 (UFRJ 1627) GoogleMaps ; • 1 male, 1 female, 1 juvenile, same locality, Camorim ( Veu da Noiva ); 04.x.2017, ALD Ferreira leg. (UFRJ 1483) GoogleMaps ; 1 male, same data (UFRJ 1523) GoogleMaps ; • 1 female, same data (UFRJ 1524) GoogleMaps ; • 1 female, same data (UFRJ 1525) GoogleMaps ; • 1 male, Sepetiba , 22°58'06.9"S, 43°42'46.8"W, iv.1994, EH Wienskoski leg. (MNRJ 1585) GoogleMaps ; • 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) GoogleMaps ; • 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 GoogleMaps : • 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) GoogleMaps ; • 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) GoogleMaps ; • 5 males, Rio Uruguai , 29°27'34.6"S, 56°43'54.2"W, 02.ix.2010, RC Francisco leg. (MCTP 43349); Santa Catarina GoogleMaps • 1 male, Nova Teutonia , 27°09'40.4"S, 52°25'31.8"W, 13-15.x.2006, ELC Silva et al. leg. (MCTP 28703); São Paulo GoogleMaps • 3 females, Botucatu , 22°56'15.3"S, 48°23'32.6"W, ii.2002, EH Wienskoski leg. (MZUSP 62396) GoogleMaps ; • 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) GoogleMaps ; • 1 male, 1 female, same locality, 10.iii.2002, EH Wienskoski leg. (MZUSP 62036) GoogleMaps ; • 1 male, 1 female, same locality, i.2003, EH Wienskoski leg. (MZUSP 62430) GoogleMaps ; • 1 male, Fazenda Paulina , 22°56'15.3"S, 48°23'32.6"W, i.2002 (MZUSP 62395) GoogleMaps ; • 1 male, 2 females, Cabreúva, Sitio do Sol , 23°18'54.4"S 47°05'48.4"W, 05.iv.2009, AB Ribeiro leg. (IBSP 145209) GoogleMaps ; • 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) GoogleMaps ; • 1 male, same locality, Corrego do Boqueirao , 19.xii.2010, Entomologia UFRJ leg. (UFRJ 0550) GoogleMaps ; • 1 male, 1 female, Rio Claro, Lidice, RPPN Fazenda Sambaiba , 22°50'55"S, 44°13'03"W, 23.iv.2012, C Bragagnolo et al. leg. (IBSP 213057) GoogleMaps ; • 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) GoogleMaps .
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 View Figure 1 , 3C View Figure 3 ). 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 View Figure 2 , 3A, B View Figure 3 ); Gl and L2 straight with much larger basis (Figs 2E, F View Figure 2 , 3B View Figure 3 ); upper teeth row with lesser teeth and lower row with more teeth (Figs 2D, F View Figure 2 , 3A, B View Figure 3 ); narrower pedipalp tibia (Figs 2H-J View Figure 2 , 3D View Figure 3 ); narrower, thinner and more pointed conductor tip (Figs 2I View Figure 2 , 3F View Figure 3 ) and epiandrous field almost straight, with narrower median division, with more spigots (26 vs. 18) (Fig. 3H View Figure 3 ; 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 View Figure 2 , 3A, B View Figure 3 ). 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 View Figure 2 , 3D-G View Figure 3 ).
Female (based on neotype UFRJ 1628): Carapace elongated, oval and reddish brown, slightly elevated anteriorly (Fig. 1A, B View Figure 1 ). Fovea reddish brown, slightly darker than carapace, with dark borders (Fig. 1A, B View Figure 1 ). Labium wider than long and dark brown (Fig. 1C View Figure 1 ). Sternum oval and light brown (Fig. 1C View Figure 1 ). Eyes with procurved parallel rows, and evenly separated, AME and PME separated by its length, ALE and PLE almost touching (Fig. 1A, B View Figure 1 ). Legs reddish brown, with few spines on femora (Fig. 1B, C View Figure 1 ). 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 View Figure 1 , 3C View Figure 3 ). AXu absent (Figs 1D, E View Figure 1 , 3C View Figure 3 ). Upper row with nine teeth distalward projected (Figs 1D, E View Figure 1 , 3C View Figure 3 ): 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 View Figure 1 ). Lower row with 20 teeth distalward projected (Fig. 1E, F View Figure 1 ): 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 View Figure 1 , 3C View Figure 3 ). Abdomen around 4.1 × longer than carapace, cylindrical and anteriorly enlarged, dorsally greyish and completely covered by guanine crystals (Fig. 1B View Figure 1 ). Venter colour as dorsum, with a median brown stripe from genital fold towards spinnerets (Fig. 1C View Figure 1 ). Genital fold elevated, 1.4 × wider than long, with parallel borders and ending in concave and wide excavated tip (Fig. 1H View Figure 1 ). 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 View Figure 1 , 3I View Figure 3 ).
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 View Figure 2 ). 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 View Figure 2 , 3A, B View Figure 3 ). ‘a’ elongated, thin, pointed, and distally projected, located on edge of paturon close to fang groove (Figs 2D, E, G View Figure 2 , 3A View Figure 3 ). AXu short with large basis (Figs 2D, E View Figure 2 , 3A View Figure 3 ), ‘t’ absent (Figs 2D, E View Figure 2 , 3A View Figure 3 ). Upper row with eight teeth distalward projected (Figs 2D, E View Figure 2 , 3A View Figure 3 ): 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 View Figure 2 , 3B View Figure 3 ). Lower row with 17 teeth and two additional ones, one besides U4 and another besides U6, all distalward projected (Figs 2E, F View Figure 2 , 3B View Figure 3 ): 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 View Figure 2 , 3A, B View Figure 3 ). Abdomen of similar colour as female, but much slimmer (Fig. 2B, C View Figure 2 ). Epiandrous field much wider than high and curved, with a narrow division, and bearing thirteen spigots on each side (Fig. 3H View Figure 3 ). Pedipalps with median-sized cymbium, around the same size as the rounded, narrow tibia (Figs 2H-J View Figure 2 , 3D View Figure 3 ); tegulum about 1.5 wider than long, spherical and inflated (Figs 2H View Figure 2 , 3D, E View Figure 3 ); 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 View Figure 2 , 3D-F View Figure 3 ); 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 View Figure 2 , 3D-F View Figure 3 ); 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 View Figure 2 , 3G View Figure 3 ).
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 View Figure 4 ), 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 View Figure 4 ). 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.
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 .
Life history and habitat preferences.
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 View Figure 5 ).
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