Tetragnatha bogotensis Keyserling, 1865,

Castanheira, Pedro de Souza, Baptista, Renner Luiz Cerqueira, Pizzetti, Daniela Dos Passos & Teixeira, Renato Augusto, 2019, Contributions to the taxonomy of the long-jawed orb-weaving spider genus Tetragnatha (Araneae, Tetragnathidae) in the Neotropical region, with comments on the morphology of the chelicerae, Zoosystematics and Evolution 95 (2), pp. 465-505: 467-469

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Tetragnatha bogotensis Keyserling, 1865
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Tetragnatha bogotensis Keyserling, 1865  Figs 1View Figure 1, 2View Figure 2, 3View Figure 3, 4View Figure 4, 20AView Figure 20, 21 A–GView Figure 21, 22AView Figure 22

Tetragnatha bogotensis  Keyserling 1865: 854, pl. 21, fig. 5 (♀).

Tetragnatha andina  Taczanowski 1878: 144, pl. 1, fig. 2 (♀); Levi 1981: 291 (removed from syn. with T. nitens  ) syn. nov.

Tetragnatha boydi  O. Pickard-Cambridge 1898: 389, pl. 31, fig. 4 (♀) syn. nov.

Tetragnatha peninsulana  Banks 1898: 246, pl. 15, fig. 12 (♂ ♀); Levi (1981): 291 (removed from syn. with T. nitens  ) syn nov.

Tetragnatha praedator  Tullgren 1910: 147, pl. 3, fig. 69 (♂)

Tetragnatha mandibulata bidentata  Gravely 1921: 442, fig. 3c, f (♂ ♀).

Tetragnatha eremita  Chamberlin 1924: 645, figs 89, 90 (♂); Levi 1981: 292 (removed from syn. with T. nitens  ) syn. nov.

Tetragnatha nitens  Lawrence 1927: 27, pl. 3, fig. 61, pl. 4, fig. 77 (♂ ♀ misidentified).

Tetragnatha bemalcuei  Mello-Leitão 1939: 67, figs 42-44 (♀) syn. nov.

Tetragnatha ramboi  Mello-Leitão 1943: 193, fig. 24, 24a, b (♂) syn. nov.

Tetragnatha haitiensis  Bryant 1945: 408, fig. 37 (♀); Levi 1981: 292 (removed from syn. with T. nitens  ) syn. nov.

Tetragnatha nitens kullmanni  Wiehle 1962: 379, figs 1-5, 6b, 9-11, 14, 15 (♂ ♀); Wunderlich 1992: 365 (removed from syn. with T. nitens  ) syn nov.

Tetragnatha infuscata  Benoit 1978: 667, fig. 2D, E (♂); Saaristo 2003: 23, figs 21A, B, 25 (syn. with T. boydi  rejected, see T. mandibulata  ).

Tetragnatha boydi praedator  Schmidt and Krause 1993: 6, fig. 5 (♂ ♀) syn. nov.

Type material.

Tetragnatha bogotensis  : COLOMBIA: ♀♀ syntypes, "Nova Granada" [surely Bogotá], 1859-1863? Alexander Lindig leg., not located, probably lost. Tetragnatha andina  : PERU: ♂♂, ♀♀ syntypes, "Amable Maria" [surely province of Tarma, region of Junín] (38 syntypes, Coll. K. Jelski, MIZ 225346-225446), examined (photos). Tetragnatha boydi  : YEMEN: ♀ syntype, Socotra, not located. Tetragnatha peninsulana  : MEXICO: ♂♂ ( CAS), ♀♀ ( MCZ 22587) syntypes: San José del Cabo, Baja California Sur, females examined (photos), males destroyed ( Levi 1981). Tetragnatha praedator  : KENYA: 4♂, syntypes ( NHRS JUST000000671-672), Kilimandjaro and Meru, examined (photos). Tetragnatha eremita  : MEXICO: ♂ holotype, Baja California, Puerto Escondido, Arroyo de Escondido (Coll. J. C. Chamberlin, 14.vi.1921, male pedipalp in MCZ 15283, labeled RVC 1111; whole specimen in CAS 1430), examined (photos of CAS 1430); ♀ paratype, same data as holotype ( MCZ 25228), examined (photos). Tetragnatha bemalcuei  : PARAGUAY: ♀ holotype, Asunción (Coll. C. Ternetz, 1895, NMB-ARAN 01092a), examined (photos). Tetragnatha ramboi  : BRAZIL: ♂, lectotype, Rio Grande do Sul (Coll. Father Balduíno Rambo, MNRJ 42467), examined. Tetragnatha haitiensis  : HAITI: ♀ holotype, “Hispaniola” [Ennery] (Coll. P. J. Darlington Jr., 07.vii.1934, MCZ 21516), examined (photos). Tetragnatha nitens kullmanni  : ITALY: ♂, holotype ( SMF 12741), 2♀, 1 immature, paratypes ( SMF 12742), Sardinia, Oliena (E. Kullmann leg.), not examined.

Extended diagnosis.

Males of T. bogotensis  are similar to T. nitens  (Audouin, 1826) and share chelicerae with an elongated ‘a’, and AXu and ‘t’ extremely elongated and distally bent. AXu and ‘t’ differ as follows: thick and triangular in T. bogotensis  ( Figs 1C, E, FView Figure 1, 3BView Figure 3), but thinner and ‘t’ sickle-like in T. nitens  ( Figs 14C, D, FView Figure 14, 16AView Figure 16). Gu longer, sharper, straight and larger than U2 in T. bogotensis  ( Figs 1C, E, FView Figure 1, 3BView Figure 3) while it is shorter than U2 and slightly curved downward in T. nitens  ( Figs 14C, DView Figure 14, 16AView Figure 16). Palps share medium-sized conductors without pleats ( Figs 1IView Figure 1, 3AView Figure 3, 14GView Figure 14, 15D, EView Figure 15). Tetragnatha bogotensis  palps differ by shorter tibias (ca 3 × longer than wide) ( Figs 1 I–KView Figure 1, 3AView Figure 3), conductors not twisted at their distal halves, with completely folded apexes enclosing the emboli tips ( Figs 1I, JView Figure 1; 3A, EView Figure 3; 20AView Figure 20) and longer paracymbia, overreaching the upper border of teguli, each bearing deeper notch, resulting in an elongated tip of the paracymbium proper, and thinner and relatively narrow translucent lobe ( Figs 1I, KView Figure 1; 3DView Figure 3). In T. nitens  , palps have longer and thinner tibias (almost 4 × longer than wide) ( Figs 14 G–IView Figure 14, 16EView Figure 16), conductors twisted in distal half with their apexes rounded and excavated, exposing the emboli tips ( Figs 14G, HView Figure 14, 16 D–FView Figure 16, 20EView Figure 20) and shorter paracymbia that do not reach the upper border of teguli, with expanded and very wide translucent lobes ( Figs 14I, JView Figure 14, 16FView Figure 16). The epiandrous field is straight, with 19 fusules, in T. bogotensis  ( Fig. 3FView Figure 3) but arched and smaller, with only 15 fusules, in T. nitens  ( Fig. 16CView Figure 16).

Females are similar to T. nitens  and also T. mandibulata  Walckenaer, 1841, sharing: bulky body, wide and pointed at the terminal end ( Figs 2A, BView Figure 2, 12 A–CView Figure 12, 15A, BView Figure 15), and elongate genital fold ( Figs 2GView Figure 2, 12HView Figure 12, 15JView Figure 15). Their chelicerae also bear an evident basal cusp (BC) in the fang ( Figs 2D, EView Figure 2, 12 D–GView Figure 12, 13C, DView Figure 13, 15D, F–IView Figure 15, 16BView Figure 16), Gu elongated and not contiguous to U2 ( Figs 2C, DView Figure 2, 12D, EView Figure 12, 13CView Figure 13, 15C, E, FView Figure 15, 16BView Figure 16); Gl large and pointed distalward, followed by smaller L2 and L3 ( Figs 2D, EView Figure 2, 3CView Figure 3, 12E, FView Figure 12, 13DView Figure 13, 15D, F, HView Figure 15, 16BView Figure 16). T. bogotensis  and T. mandibulata  differ from T. nitens  by distinct bulky AXl, which are elongated, pointed and with large bases, clearly visible and overreaching the claws even in upper view ( Figs 2 C–FView Figure 2, 3CView Figure 3, 12 D–GView Figure 12, 13C, DView Figure 13), while in T. nitens  it is short and not pointed ( Figs 15 F–HView Figure 15, 16BView Figure 16). T. bogotensis  and T. mandibulata  also differ by the conspicuous bulge (CB) in the area between both rows ( Figs 2C, EView Figure 2, 3CView Figure 3, 12D, FView Figure 12, 13C, DView Figure 13), which is absent in T. nitens  . Tetragnatha bogotensis  chelicerae can be distinguished from T. mandibulata  by the following characters: more robust basal cusp (BC), placed at the middle line of the lower side of the claw, compared to a smaller BC, displaced towards the outer face of the claw ( Figs 2D, EView Figure 2, 3 C–FView Figure 3, 12 D–GView Figure 12, 13C, DView Figure 13); Gu straight with a large basis, being separated from U2 by a wide and deep furrow, versus both teeth not quite spaced ( Figs 2C, DView Figure 2, 3CView Figure 3, 12D, EView Figure 12, 13CView Figure 13); AXl bulkier, with much larger basis, compared to a thinner and shorter tooth ( Figs 2 C–FView Figure 2, 3CView Figure 3, 12 D–GView Figure 12, 13C, DView Figure 13); Gl shorter, straight and pointed, with wider basis, and much smaller than AXl, versus a longer and slanted Gl, regularly tapered and just a bit smaller than AXl ( Figs 2D, EView Figure 2, 12E, FView Figure 12, 13DView Figure 13); and CB rounded and wide, extending from a bit above L2 to L3, and placed in the middle line between both rows of teeth, contrasting to a smaller and lower CB, extending from the basis of Gl to the middle of the gap between L2 and L3, and adjoined to the basis of L2 ( Figs 2C, EView Figure 2, 3CView Figure 3, 12D, FView Figure 12, 13C, DView Figure 13). In T. nitens  , BC is placed at a similar position to T. bogotensis  , but is larger ( Figs 15D, F–IView Figure 15, 16BView Figure 16) and Gu is connected to U2 by a thin and dark ridge, with a gap of similar size to T. bogotensis  ( Figs 15C, E, FView Figure 15, 16BView Figure 16). The genital fold ( Figs 2GView Figure 2, 12HView Figure 12, 15JView Figure 15) is similar in all three species but shorter in T. nitens  (genital fold length around 0.6 × the width), intermediate in T. bogotensis  (0.8 ×) and longer in T. mandibulata  (1.1 ×). The internal genitalia of T. bogotensis  is more similar to T. mandibulata  , with medium-sized spermathecae and a sclerotized and rounded fundus ( Figs 2H, IView Figure 2, 12IView Figure 12), in contrast to wider spermathecae without a well-defined fundus in T. nitens  ( Fig. 15K, LView Figure 15). The central membranous sac (CS) of T. bogotensis  is medium-sized, with almost the same size of the spermathecae, and placed at the same level of their bases ( Fig. 2H, IView Figure 2), while in T. mandibulata  it is massive, longer, and placed at the same level as the spermathecae ( Fig. 12IView Figure 12). In T. nitens  , the CS is about the same size as in T. bogotensis  , but with a stalk of variable size, sometimes longer than the spermathecae, placing the apical portion of the CS at the same level or anteriorly to the spermathecae ( Fig. 15K, LView Figure 15).

Synonymy and notes.

Keyserling (1865) described this species based on female specimens from "Nova Granada", which encompasses a large area ranging from Panama to Ecuador, and which were collected by Alexander Lindig, who gathered many animals and plants in Bogotá (Colombia) from 1859 to 1863 ( Meagher 2012). Therefore, the type specimens were most likely collected, at least partially, in this city, as indicated by the name T. bogotensis  . The drawing of one female by Keyserling (1865: pl. 21, fig. 5) allows the recognition of the species by showing the long and robust AXl placed near to a smaller and traverse Gl of the right chelicera. In the original description, Keyserling wrote that he had "many copies in my collection", without citation of any males ( Keyserling 1865: 855).

The first author of this paper visited three collections with type material by Keyserling: MIZ; NHM ( Beccaloni 2012), and ZMB ( Kretschmann 2006), and we also contacted curators of NHMW and USNM. Possible well-preserved type specimens were found in two vials at only the NHM. In the first vial ( Fig. 4AView Figure 4), there were three females labeled “type” and originated from “Taquara” (originally "Taquara do Mundo Novo", state of Rio Grande do Sul, Brazil). In the second vial ( Fig. 4BView Figure 4), there were two females and one male labeled “Bogotá”, but with no clear indication as type. All five females from both vials are T. bogotensis  but the male from Bogotá belongs to T. nitens  . We consider Keyserling’s syntypes to be lost, as there is no clear indication that the specimens above were used to describe the species: females from Taquara are not syntypes because Keyserling had not cited any specimens from that Brazilian locality, and females from Bogotá are also not syntypes as they were not labeled as type or "N. Granada" and are kept in the same vial as the additional male. Therefore, the confirmation of the identity of Keyserling’s species relies on the illustration in the original description and the fact that specimens collected in Bogotá and other cities from Colombia clearly display the key cheliceral teeth characters that allow the diagnosis of T. bogotensis  .

Tetragnatha boydi  has been redescribed and illustrated many times (see World Spider Catalog 2019). Pickard-Cambridge (1898) described this species based on a female from Socotra (Yemen), giving detailed illustrations of the lower and inner views of the chelicerae ( Pickard-Cambridge 1898: fig. 4a, b). Males were first described as T. praedator  Tullgren, 1910 from Kenya, a species considered junior synonym afterwards (see below). Unfortunately, the female type material was not located at NHM or in OUMNH, institutions where O. Pickard-Cambridge normally deposited his specimens, and remains lost. Lawrence (1927) (sub T. nitens  ) and Okuma (1983) were the first authors to correctly match both sexes in their papers. Lawrence (1927) gave very detailed illustrations of the male left palp and chelicera, clearly showing the diagnostic characters of this species. On the other hand, the females were not illustrated, but his description points to the "inferior margin with a large apical tooth" ( Lawrence 1927: 28), thereby rejecting the identification as T. nitens  . Okuma (1983) also correctly matched the couples, gave very detailed drawings of chelicerae and genitalia, synonymised T. mandibulata bidentata  and T. nitens kullmanni  with T. boydi  , and gave the first records for Brazil. Later, Okuma (1992) also provided new drawings of the species. After examining illustrations by Keyserling (1865: pl. 21, fig. 5), the NHM specimens of T. bogotensis  from the type locality ( Bogotá), and making comparisons with O. Pickard-Cambridge’s (1898) illustrations and the later illustrations by Tullgren (1910), Lawrence (1927), and Okuma (1983, 1992), we conclude that all specimens belong to the same common species and establish that T. boydi  is a junior synonym of T. bogotensis  .

Tetragnatha praedator  Tullgren (1910: fig. 69a, b) was described from four male specimens from Kilimandjaro and Meru (Kenya). Lessert (1915) also cited males of T. praedator  , and the species was later synonymised with T. boydi  by Roewer (1942). Finally, it was treated as a subspecies of T. boydi  by Schmidt and Krause (1993), who also described females from Comoros Island, forming the combination T. boydi praedator  Tullgren, 1910. After comparing Tullgren’s (1910) illustrations and photos of the syntypes we received ( Fig. 21CView Figure 21) with the specimens we identified as T. bogotensis  , we observed that they clearly match. Thus, T. boydi praedator  is synonymised here with T. bogotensis  . On the other hand, the females assigned to T. boydi praedator  by Schmidt and Krause (1993: fig. 5) probably belong to a different species judging by their illustrations, which are, however, too poor to allow a proper evaluation.

Tetragnatha bemalcuei  was described by Mello-Leitão (1939: 68, figs 42-44), who mentioned on the original description "a robust conical frontward apophysis" (AXl) on the lower row of the chelicerae. We examined detailed images of the holotype ( NMB) that show the characters of T. bogotensis  ( Figs 4DView Figure 4, 21EView Figure 21) and establish T. bemalcuei  as a junior synonym of T. bogotensis  Keyserling, 1865.

Mello-Leitão (1943) described T. ramboi  based on males and females from Rio Grande do Sul, south Brazil. He indicated the vial MNRJ 42467 as “tipo” (type in Portuguese) in the description, but did not label it as “typus”, in contrast to his common practice ( Fig. 4CView Figure 4). We examined the type series (one male, two females, and one immature specimen) and agree with Silva-Moreira et al. (2010) that the whole series of specimens should be treated as syntypes. In the original description, the male was cited first and its diagnostic chelicerae and palps were illustrated ( Mello-Leitão 1943: fig. 24a, b), whereas only the habitus of the female was illustrated ( Mello-Leitão 1943: fig. 24). Under the rule of the "First Reviser" ( ICZN 1999, article 24), we consider the sequence of descriptions and the presence of diagnostic illustrations in establishing the male as the lectotype of the species. This male clearly belongs to T. bogotensis  according to the chelicerae and palp diagnostic characters ( Figs 1C, E, JView Figure 1, 3A, B, EView Figure 3, 20AView Figure 20, 21FView Figure 21; Mello-Leitão 1943: fig. 24b) and must be newly synonymised with this species. Finally, we consider the female and juveniles of the type series of T. ramboi  as misidentified specimens of T. argentinensis  Mello-Leitão, 1931 which were erroneously attributed to T. ramboi  .

We also note that several species previously considered junior synonyms of T. nitens  should be newly synonymised with T. bogotensis  . For example, Levi (1981: 291, 292) established 13 junior synonyms of T. nitens  . Indeed, most of those species are correctly junior synonyms of it, but at least four should now be regarded as junior synonyms of T. bogotensis  (see T. nitens  below). It also seems that Levi (1981: figs 23-29) matched males of T. nitens  with at least some females of T. bogotensis  , as it is evident by the female illustrations he gave. These clearly depict the large AXl of T. bogotensis  ( Levi 1981: figs 23-25) and similar genitalia ( Levi 1981: figs 27-29), with a pattern very different from T. nitens  ( Fig. 15K, LView Figure 15, Zhu and Zhang 2011: fig. 125G). In the “Variation” and “Diagnosis” sections of his paper, Levi (1981: 292) pointed out "On Panamanian specimens the diagnostic tooth at the posterior base of the fang is as long as the chelicerae are wide, and is sometimes smaller than illustrated on the most northern specimens" and "The female chelicerae have a large posterior lateral tooth at the insertion of the fang". Unfortunately, he did not provide the collection site for the female specimen he illustrated, and although there is no formal record for T. bogotensis  or any of its synonyms from the United States, at least some of the females from the southern USA cited as T. nitens  may belong to that species instead.

Levi (1981: 291, 292) synonymised T. andina  Taczanowski, 1878 with T. nitens  based on multiple male and female syntypes from Amable Maria, Peru, however, Taczanowski’s (1878: fig. 2) poor illustration of the female chelicerae allows its recognition as T. bogotensis  , as verified in the photos we received from MIZ ( Fig. 21AView Figure 21). Thus, T. andina  is here removed from the synonymy of T. nitens  and newly synonymised with T. bogotensis  .

Banks (1898) described T. peninsulana  from two males and "several females" from San José del Cabo, Baja California Sur, Mexico. Photos of females provided by the MCZ ( Fig. 21BView Figure 21) include three specimens of T. bogotensis  and one of T. nitens  , which we consider a misidentification. Additionally, according to Levi (1981), the male syntypes were destroyed. Based on the elongated AXl ( Fig. 21BView Figure 21), this species must also be removed from the synonymy of T. nitens  and be newly synonymised with T. bogotensis  .

Tetragnatha eremita  Chamberlin, 1924 was based on a male holotype from Baja California, Mexico. Chamberlin (1924: figs 89, 90) provided a short description and two good illustrations that clearly show the characteristic shape and teeth formula of T. bogotensis  , besides citing a female paratype collected at the same time. The male holotype is represented by the right pedipalp in the MCZ ( MCZ 15283, RVC 1111), and by the whole specimen in CAS 1430. We were able to examine the holotype’s chelicerae through photos ( Fig. 21DView Figure 21), and thus confirmed its identity, removing T. eremita  from the synonymy with T. nitens  and newly synonymising it with T. bogotensis  .

Furthermore, Levi (1981: 292) followed Chickering (1957b: 2) in the synonymization of T. haitiensis  Bryant, 1945, which was based on a female from Haiti, with T. nitens  . Bryant (1945: fig. 37) illustrated the huge AXl tooth typical of T. bogotensis  , which we also observed in the photos we received from MCZ ( Fig. 21GView Figure 21). Therefore, T. haitiensis  Bryant, 1945 is also removed from the synonymy of T. nitens  and newly synonymised with T. bogotensis. 

Wunderlich (1992: 365) also mistook T. bogotensis  as T. nitens  , removing T. nitens kullmanni  from its proper synonymy with T. boydi  ( Okuma 1983: 70). We agree with Okuma (1983) that this species is "undoubtedly identical with T. boydi  ", as both males and females of T. nitens kullmanni  bear the same diagnostic characters of T. bogotensis  in comparison with T. nitens  : male chelicerae with Gu longer than U2 ( Figs 1EView Figure 1, 3BView Figure 3; Wiehle 1962: fig. 9) and female chelicerae with a long AXl ( Figs 2 D–FView Figure 2, 3CView Figure 3; Wiehle 1962: fig. 15). Therefore, the synonymy with T. nitens  is rejected and T. nitens kullmanni  is a junior synonym of T. bogotensis  .

Variation.

Males (n = 18): total length, 7.36-11.60; females (n = 17): total length, 7.52-11.76. The gap between Gu and U2 is variable in males of this species and can have almost the double of the length of the specimen illustrated.

Distribution.

This species is widespread in the Neotropics and Mexico, but potentially north into the southern United States; it also occurs in the Old World, with records from Africa, Yemen, India, Nepal, and China ( Fig. 22AView Figure 22).