Tmesiphantes mirim sp. nov. (Araneae: Theraphosidae) from the Atlantic Forest of Bahia, Brazil, biogeographical notes and identification keys for species of the genus

Abstract

A new species of Tmesiphantes Simon, 1892 is described and illustrated, based on eight male specimens collected at the Una Biological Reserve, southern state of Bahia, Brazil. It is distinguished by the morphology of male palpal bulb and tibial apophysis. The new species is very small and is the smallest theraphosid described to date (body length 5.5 mm). It is distinguished from congeners by the size, which vary from 12 mm (T. riopretano) to 23.8 mm (T. nubilus) in other species of the genus, aspect of palpal bulb, sternal posterior sigillae close to sternal margin and by the aspect of tibial apophysis which lacks the prolateral branch. Tmesiphantes presently comprises nine species. Sixth have been described for the southern region of Bahia, a well known area of endemism in the Atlantic Forest. Identification keys for Tmesiphantes males and females are presented.

Mygalomorphae; Neotropics; tarantula; taxonomy; Theraphosinae; Una Biological Reserve

Tmesiphantes comprises small to medium sized tarantula spiders, with labium and maxillae with few cuspules, rounded sternum, urticating setae type III and IV, femur III incrassate, male palpal bulb with curved embolus bearing two keels (prolateral superior and inferior), and spermathecal receptacles with subapical constriction.

Tmesiphantes was originally described by Simon (1892) for T. nubilus, based on a male and a female from the state of Bahia. Later, Mello-Leitão (1923, 1926, 1943) described three species from Brazil: in 1923, T. montanus from Itatiaia, state of Rio de Janeiro; in 1926, T. physopus from Campina Grande, state of Paraíba; and in 1943, T. minensis from Ouro Preto, state of Minas Gerais. Caporiacco (1955) described T. chickeringi, from Venezuela. Gerschmann de Pikelin & Schiapelli (1958) revised the genus and described two species from Argentina, T. serratus and T. elegans. They provided an identification key that included all species but T. chickeringi, which was known only from an a immature female. Later Gerschmann de Pikelin & Schiapelli (1972), transferred T. montanus and T. elegans to Homoeomma and considered H. uruguayense Mello-Leitão, 1946 as senior synonym of T. serratus. Raven (1985) considered Tmesiphantes as junior synonym of Dryptopelma Simon, 1889 (currently under the synonym of Cyclosternum Ausserer, 1871 by Pérez-Miles et al. 1996: 46) and confirmed the taxonomic changes made by Gerschmann de Pikelin & Schiapelli (1972). Pérez-Miles et al. (1996) revalidated Tmesiphantes, emphasizing the morphology of the male palpal bulb and femur III incrassate as diagnostic features of the genus. Schaefer (1996) described T. spinopalpus from Paraguay, which was later transferred to Cyclosternum Ausserer, 1871 by Rudloff (2000).

As a result of Rudloff's (2000) contribution Tmesiphantes comprised four species: three from Brazil (T. nubilus, T. physopus and T. minensis) and one from Venezuela (T. chickeringi). Two years later, Yamamoto et al. (2007) revised Tmesiphantes and described three species in it (T. amadoi, T. bethaniae and T. caymmii). They also proposed the following taxonomic changes: they placed T. chickeringi as species inquirenda; and transferred T. physopus and T. minensis to Plesiopelma because they have a retrolateral basal nodule on male metatarsus I. Guadanucci & Silva (2012) described two species from Minas Gerais, Brazil (T. perp and T. riopretano). Bertani et al. (2013) described T. hypogeus, the first troglomorphic tarantula from Brazil, also from Bahia. Recently, Gonzalez-Filho et al. (2014) described T. aridai, from the state of Pará, Brazil, comprising the first record of the genus for the Amazon Forest. Currently, the genus includes eight species from Brazil: T. nubilus, T. amadoi, T.bethaniae, T. caymmii, T. riopretano, T. perp, T. hypogeus and T. aridai. The examination of extra material from southern Bahia, deposited at Butantan Institute, revealed two small male spiders that fit the generic definition of Tmesiphantes. This species is described here as Tmesiphantes mirimsp. nov.

MATERIAL AND METHODS

The material examined belongs to the arachnid collection of Butantan Institute (IBSP, São Paulo, Brazil, A.D. Brescovit). All measurements are in millimeters, taken with a millimetric ocular lens. The length of leg segments was measured between the joints in dorsal view. The lengths and widths of the carapace, eye tubercle, labium and sternum are the maximum values obtained. The total body length does not include chelicerae and spinnerets. The male palpal bulb was removed from the cymbium and photographed in dorsal, prolateral and retrolateral views. The tibial apophysis was photographed in ventral and prolateral views. Abbreviations: (AME) anterior median eyes, (ALE) anterior lateral eyes, (PME) posterior median eyes, (PLE) posterior lateral eyes. Spines: (d) dorsal, (v) ventral, (p) prolateral, (r) retrolateral. The number and disposition of spines follows the terminology of Petrunkevitch (1925); urticating hair types terminology follows Cooke et al. (1972). Pictures and measurements were taken with a stereomicroscope (Leica M205C with DFC295 digital camera and Leica Application Suite v.4.2).

The distributional dataset was based on three distinct sources: the literature, scientific collections (listed below) and field collecting. All records were assembled in Microsoft Excel (xls files). The xls files were converted into shapefile format (shp) and the map was built using DIVA-GIS v.7.5, available at http://www.diva-gis.org.

Distributional records from the following scientific collections were used: (IBSP) Instituto Butantan, São Paulo, Brazil; (MZSP) Museu de Zoologia da Universidade de São Paulo, Brazil; (MPEG) Museu Paraense Emilio Goeldi, Belém, Brazil; (MNRJ) Museu Nacional do Rio de Janeiro, Rio de Janeiro, Brazil; (CAD) Coleção Aracnológica Diamantina, Diamantina, Brazil; (UFMG) Universidade Federal de Minas Gerais, Belo Horizonte, Brazil; (UFPB) Universidade Federal da Paraíba, João Pessoa, Brazil.

The following material was examined: Tmesiphantes: male holotype of T. amadoi from Jussarí, Bahia, Brazil, March 2001 (IBSP 12930). Paratypes of T.amadoi: one male with the same data as the holotype (IBSP 11878); four males with the same data as the holotype (IBSP 11859). Male paratypes of T. caymmii from Poções, Bahia, Brazil, 3 June 2002, A. Zanotti leg. (MZSP 28194). Male holotype of T. bethaniae from Una, Bahia, Brazil, December 1999, K. Kato leg. (MZSP 24226). Paratypes of T. bethaniae: two males from Uruçuca, Bahia, Brazil, March 2002, J.P.L. Guadanucci leg. (MZSP 24125; MZSP 24129). Four males of T. nubilus from Uruçuca, Bahia, Brazil, March 2002, J.P.L. Guadanucci leg. (MZSP 24241). Male holotype of T. riopretano from Parque Estadual do Rio Preto, São Gonçalo do Rio Preto, Minas Gerais, Brazil, 20-25 October 2010, G. Monteiro, F. Sá, W.F. Silva, J.P.L. Guadanucci leg. (CAD 422). Paratypes: one female of T. riopretano from the same locality as holotype, 12-19 January 2010, J.P.L. Guadanucci, W.F. Silva, D. Moura leg. (CAD 016). Male holotype of T. perp from Parque Estadual do Rio Preto, São Gonçalo do Rio Preto, Minas Gerais, Brazil, 19 January 2010, J.P.L. Guadanucci, W.F. Silva, D. Moura, R.F. Fonseca, D. Weinmann leg. (CAD 024). Holotype female of T. hypogeus from Andaraí, Igatu, Gruta da Parede Vermelha, Bahia, Brazil, 29 October 2010, M.E. Bichuette and J.E. Gallão leg. (MNRJ 04357). Paratypes: one male and two females of T. aridai from Floresta Nacional do Tapajós, Santarém, Pará, Brazil, X. 2011, A.D. Brescovit et al. leg. (IBSP 165020, IBSP 149977, IBSP 165996). Melloleitaoina: holotype male of Melolleitaoina crassifemur from Salta, Argentina, IIX.1947, N. Orfila leg. (MACN-Ar 2285). Holotype male of Melolleitaoina mutquina from Mutquín, Catamarca, Argentina, 2.II.1981, E. Maury leg. (MACN-Ar 7737). Holotype male of Melolleitaoina uru from Salta, Argentina, undated collection, Goloboff, Coyle & Bennet leg. (MACN-Ar 26042). Paratypes: one female of Melolleitaoina uru from Punila, Araguaray, Salta, Argentina, IV.1948, Biraben leg. (MACN-Ar 6542). Holotype male of Melolleitaoina yupanqui from P. Nacional Calilegua, Jujuy, Argentina, 06-11.XII.2008, C. Grismado, M. Izquierdo, F. Labarque, G. Rubio, M. Burger, P. Michalik, P. Carrera, A. Ojanguren & C. Mattoni leg. (MACN-Ar 26041). Paratypes: one female of Melolleitaoina yupanqui from P. Nacional Calilegua, Jujuy, Argentina, 06-11.XII.2008, C. Grismado, M. Izquierdo, F. Labarque, G. Rubio, M. Burger, P. Michalik, P. Carrera, A. Ojanguren & C. Mattoni leg. (MACN-Ar 26044).

TAXONOMY

Tmesiphantes mirimsp. nov.Figs. 1-7, Table 1

Diagnosis. It differs from the other species by the aspect of palpal bulb (Figs. 2-4), with the embolus inserted perpendicularly to the tegulum, in a way that the inferior prolateral keel is perpendicular to the basal-apical axis in the retrolateral view (Fig. 4).

The apical end of embolus is as wide as the mid portion, resembling representatives of T. bethaniae and T. aridai. It differs from those species by the sternal posterior sigillae close to sternal margin and by the produced prolateral inferior keel. In addition, by the aspect of tibial apophysis (Figs. 6 and 7), which lacks the prolateral branch, which is present and conspicuous in other species of the genus.

Females are unknown.

Description. Male holotype. Coloration (in ethanol 70%): carapace beige (Fig. 1), abdomen and legs light beige (Fig. 1). Total length 5.5. Cheliceral basal article with 11 teeth each on the promargin. Carapace: length 2.7, width 2.0. Thoracic furrow straight. Clipeus narrow. Eye tubercle: length 0.3, width 0.5. Eyes sizes: AME 0.1, ALE 0.3, PME 0.1, PLE 0.1. Labium: length 0.2, width 0.3, with three cuspules (Fig. 5). Sternum rounded (Fig. 5) length 1.3, width 1.2. Posterior sternal sigillae elongated and next to sternal margin. Maxillae with 34-23 (left-right) cuspules (Fig. 5). Spines: palp articles and leg tarsi without spines. Legs: I - metatarsus v0, p0-1-0, r0. II - tibia v0, p0, r0-1-0; metatarsus v0, p0, r0-1-1. III - metatarsus v0-0-1, p0-1-1-0, r0-1-1-0. IV - tibia: v1-0-0-1, p1-1-1-1, r0-1-0-1; metatarsus v0-0-1-0, p0-0-0-1, r0-0-1-1. Palpal bulb piriform, with embolus slightly curved, two parallel keels (prolateral superior and inferior), inferior more developed than superior (Figs. 2-4). Tibial apophysis (Figs. 6 and 7) composed of a single branch (namely retrolateral branch in the other species) bearing a short spine on the apex; a short spine is present, next to the apophysis branch. Urticating hairs type III on the center of abdominal dorsum. Tarsal scopula I-IV entire. Scopula on metatarsi absent. Femur III incrassate. Single tooth on each of the paired claws. PMS absent, apical article of PLS digitiform.

Type material.

Distribution. Known only from the type locality.

Etymology. The specific epithet is a word in Tupi, a Brazilian indigenous language, meaning very small.

Table 1.
Tmesiphantes mirimsp. nov. (holotype), legs and palp articles lengths (mm).

Figures 1-7.
Tmesiphantes mirimsp. nov. male holotype (IBSP 9954): (1) dorsal habitus; (2-4) male palpal bulb: (2) prolateral view; (3) retrolateral view; (4) dorsal view; (5) cephalotorax, vental view; (6-7) tibial apophysis; (6) lateral view; (7) ventral view. Scale bars: 1, 5-7 = 1 mm, 2-4 = 0.5 mm.

Key to males of Tmesiphantes species

1. 1. Palp bulb with a strong curvature in the apical portion of the embolus (Yamamoto et al. 2007: figs. 3-5; Guadanucci & Silva 2012: figs. 11-13) .................... 2

2. 1'. Palp bulb without such curvature in the apical portion of the embolus .................... 3

3. 2. Presence of a membrane in the apical portion of embolus, which is thicker than the mid portion (Guadanucci & Silva 2012: figs. 11-13) metatarsus I slightly curved .................... Tmesiphantes perp Guadanucci & Silva, 2012

4. 2'. Absence of such a membrane, the apical portion of embolus with the same thickness (Yamamoto et al. 2007: figs. 3-5) of the mid portion; metatarsus I straight .................... Tmesiphantes nubilus Simon, 1892

5. 3. Apical portion of embolus wider than the mid portion .................... 4

6. 3'. Apical portion of embolus of the same width or thinner than mid portion .................... 5

7. 4. Presence of serrated keel at apical end of embolus (Yamamoto et al. 2007: figs. 9 and 11) .................... Tmesiphantes amadoi Yamamoto et al., 2007

8. 4'. Presence of a circular patch of short bristles on the mid portion of abdomen (Yamamoto et al. 2007: fig. 20) .................... Tmesiphantes caymmii Yamamoto et al., 2007

9. 5. Apical portion of embolus thinner than the mid portion (Guadanucci & Silva 2012: figs. 3-5) .................... Tmesiphantes riopretano Guadanucci & Silva, 2012

10. 5'. Apical portion of embolus of the same width as mid portion .................... 6

11. 6. Posterior sternal sigillae near sternal margin; very small spiders (not exceed 6 mm of body length) .................... Tmesiphantes mirimsp. nov.

12. 6'. Posterior sternal sigillae far from sternal margin, medium-sized spiders (more than 22 mm of body length) .................... 7

13. 7. Retrolateral branch of tibial apophysis very small, not longer than half the length of spine next to it (Yamamoto et al. 2007: figs. 24) .................... Tmesiphantes bethaniae Yamamoto et al., 2007

14. 7'. Retrolateral branch of tibial apophysis longer than spine next to it (Gonzalez-Filho et al. 2014: fig. 5) .................... Tmesiphantes aridai Gonzalez-Filho et al., 2014

Males of T. hypogeus are unknown.

Key to females of Tmesiphantes species

1. 1. Spermatheca with receptacles fused at base (Yamamoto et al. 2007: fig. 8) .................... Tmesiphantes nubilus Simon, 1982

2. 1'. Spermatheca with free receptacles .................... 2

3. 2. Strong constriction at subapical region of spermatheca receptacles .................... 3

4. 2'. Without constriction at subapical region of spermatheca receptacles .................... 4

5. 3. Subapical region of the spermatheca receptacles thinner than the base (Yamamoto et al. 2007: fig. 19) .................... Tmesiphantes caymmii Yamamoto et al., 2007

6. 3'. Subapical region of spermatheca receptacles wider than the base (Guadanucci & Silva 2012: fig. 8) .................... Tmesiphantes riopretano Guadanucci & Silva 2012

7. 4. Anterior median eyes greatly reduced, transparent ocular tubercle (Bertani et al. 2013: fig. 2C); two patches of urticating hair type III at dorsolateral areas on the abdomen (Bertani et al. 2013: figs. 1and 2A) .................... Tmesiphantes hypogeus Bertani et al., 2013

8. 4'. Anterior median eyes of regular size (González-Filho et al. 2014: fig. 8); single urticating hair patch on abdomen .................... Tmesiphantes aridai González-Filho et al., 2014

9. Females of T. amadoi, T. bethaniae, T. perp, and T. mirimsp. nov. are unknown.

DISCUSSION

Species of Tmesiphantes are present in the Amazon Forest (T. aridai), in rocky fields of Cerrado (T. perp and T. riopretano), in the Atlantic Forest (T. bethaniae, T. amadoi, T. caymmii, T. nubilus, and T. mirimsp. nov.), and in Caatinga (T. hypogeus and T. nubilus) (Fig. 8). The species described here is the sixth from southern Bahia and the fifth from the Ombrophilous Dense Forest that covers the southern part of the state. Moreover, T. bethaniae, T. amadoi, T. nubilus, and T. mirimsp. nov. are sympatric at the municipality of Una (Una Biological Reserve) (Fig. 8).

Figure 8.
Distribution map of Tmesiphantes. (CS) Cerrado/Savanna, (Am) Amazonian Forest, (Caa) Caatinga, (HAF) Humid Atlantic Forest, (IAF) Interior Atlantic Forest.

Southern Bahia is a well-known area of endemism in the Atlantic Forest (e.g., Prance 1982, Silva & Casteli 2005, DaSilva & Pinto-da-Rocha 2011). There, the forest has remained relatively stable from the late Quaternary to the present, acting as a refuge for forest species (Carnaval & Moritz 2008). The sympatric occurrence of the four species of Tmesiphantes at Una and the endemism of three of them in this area (Fig. 8) support the historic biogeographical hypothesis.

The sympatry of T. bethaniae, T. amadoi, T. nubilus, and T. mirim is remarkable since it is not a common pattern of geographical distribution among Theraphosidae spiders in the Neotropics. Closely-related spiders in this family are more commonly allopatric or parapatric (e.g., Catanduba Yamamoto, Lucas & Brescovit, 2012, Catumiri Guadanucci, 2004, Cyriocosmus Simon, 1903, Ephebopus Simon, 1892, Eupalaestrus Pocock, 1901, Homoeomma Ausserer, 1871, Plesiopelma Pocock, 1901, Proshapalopus Mello-Leitão, 1923, Typhochlaena C.L. Koch, 1850, Pachistopelma Pocock, 1901, Iridopelma Pocock, 1901, Nhandu Lucas, 1983). There are also cases of partial sympatry (e.g., Magula Simon, 1892, Maraca Pérez-Miles, 2006, Vitalius Lucas, Silva & Bertani, 1993, Acanthoscurria Ausserer, 1871). T. mirimsp. nov. is a tiny theraphosid spider (5.5 mm of body length - the smallest theraphosid described to date) that shares its habitat with congeners (ranging from 19 to 23 of body length). The great difference in the size of this species may be the result of sympatric speciation. It is possible that small spiders avoided competition with with larger ones, since they were able to explore smaller microhabitats and food items, which gave them an advantage and eventually led to speciation. This hypothesis, however, needs to be tested with ecological and behavioral approaches.

The intra and inter relationships of Tmesiphantes are far from being resolved. As already pointed out (Pérez-Miles et al. 1996, Yamamoto et al. 2007, Guadanucci & Silva 2012, Perafán & Pérez-Miles 2014), representatives of Tmesiphantes share the femur III incrassate with spiders of the genus Melloleitaoina Gerschman & Schiapelli, 1960. According to Perafán & Pérez-Miles (2014), the distinction between the two genera relies on the aspect of the sternal sigillae, which are more rounded in Melloleitaoina, and the leg spines, which are more numerous in Tmesiphantes. After description of T. mirim, along with the examination of all species of both genera, it has become obvious to us that all previous diagnoses fail to separate these genera due to interspecific morphological variation. It is very likely that these two genera are synonyms. In this case, Tmesiphantes would be the senior synonym, and for this reason we have chosen to place our new species in it. However, a comprehensive revision and phylogenetic analysis including representatives of all species of both genera, as well as close related genera (Grammostola Simon, 1892, Homoeomma Ausserer, 1871, Euathlus Ausserer, 1875, Cyriocosmus Simon, 1903, Maraca Pérez-Miles, 2006, Plesiopelma Pocock, 1901), is needed to ascertain the generic status of Melloleitaoina and Tmesiphantes.

ACKNOWLEDGEMENTS

We thank the curator of Butantan, Antonio Brescovit, for providing the material on loan for analysis and description of the holotype. Celso Feitosa Martins of the Departmento de Sistemática e Ecologia (UFPB) for granting us access to the Entomology Laboratory for photographic recording of the holotype. Capes/UFPB for graduate fellowship (W.F.) and CNPq for grant (M.B.D., 477383/2013-0).

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• Errata

  Page 153, second column, line 9:
  Where read: Type material.
  Should read: Type material. Holotype male from Una, Bahia, Brazil (IBSP 109954), from X.1999 to IX.2000, M.T. Dias leg. Paratype one male (IBSP 109958), two males (IBSP 111463), one male (IBSP 111482), two males (IBSP 111463) and one male (IBSP 111482).

Publication Dates

• Publication in this collection
  Mar-Apr 2015

History

• Received
  26 Dec 2014
• Reviewed
  23 Mar 2015
• Accepted
  25 Mar 2015