Phoneutria boliviensis (F. O. Pickard-Cambridge, 1897)
Hazzi, Nicolas A. & Hormiga, Gustavo, 2021, Morphological and molecular evidence support the taxonomic separation of the medically important Neotropical spiders Phoneutria depilata (Strand, 1909) and P. boliviensis (F. O. Pickard-Cambridge, 1897) (Araneae, Ctenidae), ZooKeys 1022, pp. 13-50 : 13
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|Phoneutria boliviensis (F. O. Pickard-Cambridge, 1897)|
Ctenus boliviensis : F. O. Pickard-Cambridge, 1897: 80, pl. 3, (female holotype from Madre de Dios, Bolivia, fig. 3a-c (male), The Natural History Museum, London not found; see Schiapelli and Gerschman de Pikelin 1973: 36, and Simó and Brescovit 2001: 74.
Ctenus nigriventroides Strand, 1907: 426 (female holotype from Bolivia, Museum für Natur und Umwelt der Hansestadt, Lübeck presumed lost; see Eickstedt 1979: 111, and Simó and Brescovit 2001: 74).
Ctenus valdehirsutulus Strand, 1910: 318 (syntypes: female from Sara, W. Bolivia, 60 m, 14 March 1907, J. Steinbach leg., in ZMB 30615 View Materials ; female from Sara, Dpto. Sta. Cruz de la Sierra, Bolivia, 500 m, Steinbach, in ZMB 30616 View Materials , see Simó and Brescovit 2001: 74) .
Ctenus nigriventoides : Petrunkevitch, 1911: 475 (only citation of Strand 1907), 735.
Ctenus chilesicus Strand, 1915: 128 (female holotype from Chile, 1902, O. Hohenemser leg., in SMF-4557) .
Phoneutria boliviensis : Schmidt, 1954: 414; 1956: 28; Bücherl 1968: 188; 1969a: 49; Schiapelli and Gerschman de Pikelin 1973: 31, 33-38 (redescription male and female).
Phoneutria nigriventroides : Bonnet, 1958: 3621 (in part, only material from Bolivia); Eickstedt 1979: 111.
Justification of the neotype designation.
We have designated a neotype for P. boliviensis in accordance with Article 75 of the International Code of Zoological Nomenclature ( ICZN 1999 ). The type material of Ctenus boliviensis was considered lost after examination of the spider material at the Natural History Museum , London ( Simó and Brescovit 2001). The epigynum of the syntype female was reported to be damaged by Schiapelli and Gerschman (1973). In absence of type material, we consider necessary to designate a neotype to clarify the taxonomic status of P. boliviensis . Although the original type locality of P. boliviensis is the Madre de Dios area of Bolivia (F.O. Pickard-Cambridge 1897), the locality of the neotype (in Peru) belongs to the same region. The region takes its name from the Madre de Dios river, which is part of the Amazon river watershed. The Madre de Dios basin spreads across Bolivia and Peru. This area is called Inambari and is considered as a single biogeographic area because of its unique composition of species ( Da Silva et al. 2005). In addition, the neotype locality is very close to the Bolivian border (30 km in linear distance) .
The syntypes of Ctenus valderhirsutulus were revised by Simó and Brescovit (2001) and this species was deemed to be a junior synonym of Phoneutria bolivienesis . The syntype localities of Ctenus valderhirsutulus (the Sara Province of Bolivia, in the Santa Cruz Department) are within the distribution area of Phoneutria bolivienesis which corroborates the synonymy proposed by Simó and Brescovit (2001). The type of Ctenus chilesicus comes from an undisclosed locality in Chile and was deemed to be conspecific with Phoneutria bolivienesis by Simó and Brescovit (2001). The records of Phoneutria from Chile are of introductions of P. fera ( Zapfe 1963; Canals et al. 2004). Although we have no reason to question the synonymy of Ctenus chilesicus with Phoneutria bolivienesis , which was based on the examination of type specimens, future work should revise the type of Ctenus chilesicus . We suspect that the only specimen of Ctenus chilesicus is an introduction of an already described species (as suggested by Simó & Brescovit) or the result of labeling error.
Other material examined.
Colombia: Caqueta: two males, Universidad de la Amazonia (1.4998°N, 75.6632°W, 240 m) Florencia, 30.VII.2019, N. Hazzi, L. Martínez, and E. Across-Valencia (MUSENUV) GoogleMaps ; Amazonas: Comunidad Monifue Amena (4.1128°N, 69.9311°W, 70 m) 03.X.2005 ( MPUJ) GoogleMaps . Peru: Loreto: two males and two females, Reserva Nacional Allpahuayo Mishana, Biological Station “José Alvarez Alonso" (3.9663°S, 73.4368°W, 120 m) Iquitos, 02.IX.2019, N. Hazzi, E. Vargas and G. Gagliardi ( MCZ IZ 162185 View Materials ) GoogleMaps ; one female and one male, Universidad Nacional de la Amazonia Peruana (3.8466°S, 73.3671°W, 110 m), Puerto Almendras, Iquitos, 01.IX.2019, N. Hazzi and E. Vargas ( MCZ IZ 162186 View Materials ?) GoogleMaps ; one female, San Rafael (3.5617°S, 73.1191°W, 90 m), 04.IX.2019, N. Hazzi and E. Vargas ( MCZ IZ 162187 View Materials ?) GoogleMaps ; Inahuaya, Cerros Orullana (7.1158°S, 75.2709°W, 150 m), 9.VII.1988, R, Fernandez and P. Hocking ( MUSM-ENT 511187 ) GoogleMaps ; Ucayali: four females and four males, Panguana Biological Research Station (9.6137°S, 74.9352°W, 220 m), 15.IV.2019, N. Hazzi ( MCZ IZ 162188 View Materials ) GoogleMaps ; Madre de Dios: one female, same data as neotype ( MUSM-ENT 054122 ) ; one female Zona Reservada de Manu (11.96°S, 71.30°W, 250 m), 01.X.1987, D. Silva & J. Coddington ( USNM) GoogleMaps ; three females and one male, Zona Reservada Tambopata (12.83°S, 69.283°W, 290 m) ( MUSM-ENT 507653 , 507657 , 507658 and 507659) GoogleMaps ; Zona Reservada Pakitza (11.96°S, 71.30°W), 26. V.1987, ( MUSM-ENT 509196 ) GoogleMaps , one male, Explorers Inn (12.8455°S, 69.2942°W), 19. VI.2009 ( MUSM-ENT 500807 ) GoogleMaps ; Santuario Nacional Pampas del Heath (12.042°S, 71.7248°W), 27. VI.1987, V. Morales ( MUSM-ENT 509147 ) GoogleMaps ; Huánuco: one female and one male, Dantas la Molina (9.633°S, 75°W, 270 m), SW Puerto Inca, 18. V.1987 ( MUSM-ENT 507582 , 511349 ) GoogleMaps ; San Martin: one female, Juanji (7.1669°S, 76.7395°W, 350 m), 16.VIII.1998 ( MUSM-ENT 511348 ) GoogleMaps ; Pasco: one male, Santa Maria, Rio Palcazu (9.9369°S, 75.2471°W), 8.III.1998, P. Hocking ( MUSM-ENT 511043 ) GoogleMaps ; Amazonas: one male, Condorcanqui (4.59841°S, 77.8599°W), 18.VII.1994, M. Ortega ( MUSM-ENT 509062 ) GoogleMaps .
Males of P. boliviensis resemble those of P. depilata by the truncated apex of the RTA (Fig. 9C, D View Figure 9 ), but differ by the smaller tegulum (Figs 5B View Figure 5 , 9A View Figure 9 ), round median apophysis enlarged at the base (Figs 5B View Figure 5 , 9A View Figure 9 ), locking lobes located posteriorly (Figs 5B View Figure 5 , 9A View Figure 9 ), in contrast with the narrow base of the median apophysis and pronounced lateral locking lobes in P. depilata ; and embolus without internal bulge (Figs 5B View Figure 5 , 9A View Figure 9 ). Females of P. boliviensis also resemble those of P. depilata by the general configuration of the epigynum but differ by the wider area of the EMF (Figs 6A View Figure 6 , 10A View Figure 10 ), copulatory ducts strongly sclerotized (Figs 6B View Figure 6 , 10B View Figure 10 ), and reduced spermatheca heads (Figs 6B View Figure 6 , 10B View Figure 10 ), in contrast with the less sclerotized copulatory ducts and larger spermatheca heads of P. depilata . In addition, both females and males can be distinguished from P. depilata and the remaining Amazonian species ( P. perty and P. fera ) by the two lateral conspicuous white-yellow bands in the anterior area of the carapace which are also absent in all other congeneric species (Fig. 4A View Figure 4 ).
Male (from Madre de Dios, Puerto Maldonado, Finca Las Piedras, Peru; MUSM-ENT 54118). Coloration (Figs 1D View Figure 1 , 4A, B View Figure 4 ): Carapace brown with a longitudinal black line, transversal black stripes and two lateral conspicuous white-yellow bands in the anterior area. Ocular area with dark black-blue setae and back oblique band from PLE to anterior dorsal shield of prosoma edge. Chelicerae brown. Sternum, endites and labium yellowish-brown. Dorsal abdomen yellow-brown, with a longitudinal black line reaching to the median region; ventrally dark brown with four series of pale brown dots. Total length 20.93. Carapace 10.91 long and 13.18 wide, eye diameters: AME 0.41, ALE 0.23, PME 0.72, PLE 0.46. Clypeal height 0.26, sternum 4.57 long, 4.00 wide; labium 1.31 long, 0.84 wide. Leg measurements: I: femur 12.20, patella 4.20, tibia 13.52, metatarsus 17.98, tarsus 5.00, total 52.90; II: 17.60, 7.49, 18.81, 13.65, 3.94, total 61.49; III, 14.09, 6.62, 12.42, 8.18, 2.43, total 43.74; IV 11.79, 4.42, 10.8, 12.58, 3.42, total 43.01. Leg spination: I tibia v2-2-2-2-2, d1-1-1, p0-1-0, r1-1-0, metatarsus v2-2-2, p1-0-0 r1-0-0, II tibia v-2-2-2-2-2, d1-1-1, p1-1-0, r1-1-0, metatarsus v2-2-2, p1-0-0 r1-0-0, III v2-2-2, d1-1-1, p1-0-1-0, r1-0-1-0, metatarsus v2-2-2-2, p1-1-2, r1-1-2, IV tibia v2-2-2, d1-1-1, p1-0-1-0, r1-0-1-0, metatarsus v2-2-2-2, d0-1-0, p1-1-2, r1-1-2. Palp. RTA small and truncated at the apex (Figs 5C View Figure 5 , 9C View Figure 9 ); embolus curve without internal bulge (Figs 5B View Figure 5 , 9A View Figure 9 ); cup-shaped median apophysis constrained at the base (Figs 5B View Figure 5 , 9A View Figure 9 ); conductor membranous, hyaline and C-shaped (Figs 5B View Figure 5 , 9A View Figure 9 ); tegulum with probasal rounded projection (Figs 5B View Figure 5 , 9A View Figure 9 ).
Female (from Madre de Dios, Puerto Maldonado, Finca Las Piedras, Peru; MUSM-ENT 054122). Coloration (Figs 1C View Figure 1 , 4A, B View Figure 4 ): Carapace brown with a longitudinal black line and two lateral conspicuous white-yellow bands in the anterior area. Ocular area with dark brown setae and back oblique band from PLE to anterior dorsal shield of prosoma edge. Chelicerae brown with red setae. Sternum, endites and labium yellowish-brown. Dorsal abdomen yellow-brown, with a yellow dot; ventrally dark brown with four series of pale brown dots. Total length 20.19. Carapace 9.70 long and 7.57 wide, eye diameter: AME 0.45, ALE 0.29, PME 0.46, PLE 0.53. Clypeal height 0.44, sternum long 3.94 and 3.55 wide, endites 3.89 long and 2.50 wide, labium 1.43 long and 1.25 wide. Leg measurements: I: femur 9.06, patella 3.98, tibia 9.93, metatarsus 8.01, tarsus 2.33, total 33.31; II, 8.45, 4.19, 8.67, 6.90, 2.27, total 30.48; III 6.92, 3.20, 5.97, 5.46, 1.59, total 23.14; IV 8.66, 3.51, 8.06, 9.00, 1.58, total 30.81. Leg spination: tibia I-II v2-2-2-2-2, metatarsus I-II v2-2-2-2-2; III tibia v2-2-2, d1-1-1, p1-0-1-0, r1-0-1-0; metatarsus v2-2-2-2, p1-1-2, r1-1-2; IV tibia v2-2-2, d1-1-1, p1-0-1-0, r1-0-1-0, metatarsus v2-2-2-2, d0-1-0, p1-1-2, r1-1-2. Epigynum (Figs 6A View Figure 6 , 9A View Figure 9 ): middle field convex with straight edges, anteriorly divergent and posteriorly convergent; lateral field with lateral apophysis. Vulva (Figs 6B View Figure 6 , 9B View Figure 9 ): copulatory ducts strongly sclerotized and reduced spermatheca heads, fertilization ducts small and posteriorly located.
Males (n = 6): Total length 9.70-10.60, carapace 4.86-5.90, femur I 5.90-6.72. Females (n = 5): Total length 12.22-15.22, carapace 6.33-6.97, femur I 5.20-5.86.
Phoneutria boliviensis is the smallest species of the genus and it inhabits in sympatry with P. fera and P. reidyi . Torres-Sánchez and Gasnier (2010) indicated that P. boliviensis seems to be restricted to periodically indudated forests because they have never been detected in "terra firme" forests. In Peru, this species was also very common in swamp forests (aguajales) dominated by the large, dioecious palm Mauritia flexuosa . However, we also found that Phoneutria boliviensis is not exclusive to inundated forests but also can be found in "terra firme" forests and even in the Amazonian foothills in Caqueta, Colombia. In these non-inundated ecosystems, P. boliviensis is found in secondary forests and forest edges. This species lives in the leaf litter and low vegetation. It is interesting to highlight that in the Amazon of Colombia, Ecuador and Peru, we always found P. boliviensis in sympatry with P. fera but never with P. reidyi .
Museum f�r Naturkunde Berlin (Zoological Collections)
Museum of Comparative Zoology
Instituto de Zoologia
Departamento de Geologia, Universidad de Chile
Smithsonian Institution, National Museum of Natural History
Royal British Columbia Museum - Herbarium
Mykotektet, National Veterinary Institute
Západoceské muzeum v Plzni
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