New species of the....Part II

[Vera Santos]

Continução

 

 

Table 2.

Likelihood-based tests for alternative topologies. SH and AU are probability values obtained from the Shimodaira-Hasegawa and the Approximately Unbiased tests (Shimodaira 2002). Asterisks denote significant values (P<0.05 for SH and P<0.01 for AU and ELW) that imply the topology is rejected.

Test Topology - Ln L ∆ - LnL ELW SH AU ML 11910.81 1 Pseudancistrus zawadzkii sister group to Pseudancistrus pectegenitor +Pseudancistrus sidereus a 11952.41 41.60 <0.001* 0.021* <0.001* 2 Pseudancistrus zawadzkii sister group to Guyanancistrus members a 11962.24 51.43 <0.001* 0.011* <0.001* 3 Pseudancistrus zawadzkii sister group to Lithoxancistrus members a 11966.25 55.44 <0.001* <0.001* <0.001* 4 Pseudancistrus zawadzkii sister group to Pseudancistrus genisetiger a 12033.30 122.49 <0.001* <0.001* <0.001*

a The alternative topology was defined as the ML tree forcing the desired relationship.

 

Table 3.

Taxa list, specimen and sequence data analyzed in the present study (n=44). Institutional acronyms followFricke and Eschmeyer (2013).

Species Catalog Number Field Number GenBank Nº F-RTN4 Ref. Corydoras oiapoquensis MHNG 2682.023 GF06-186 GU210997 Alexandrou et al. (2011) Hemipsilichthys gobio LBP 2368 15363 EU817547 Chiachio et al. (2008) Harttia guianensis MHNG 2643.016 GF00–351 FJ013232 Chiachio et al. (2008) Hypostomus sp. MHNG 2721.062 PE08-198 JN855790 Covain and Fisch-Muller (2012) Hypostomus boulengeri (Eigenmann & Kennedy, 1903) MHNG 2519.23 ASU7 EU817560 Chiachio et al. (2008) Hypostomus gymnorhynchus (Norman, 1926) MHNG 2621.098 SU01-160 JN855789 Covain and Fisch-Muller (2012) Ancistrus cirrhosis (Valenciennes, 1836) MHNG 2645.037 MUS 202 HM623638 Rodriguez et al. (2011) Dekeyseria picta (Kner, 1854) MHNG 2588.046 MUS 162 JN855755 Covain and Fisch-Muller (2012) Dekeyseria scaphirhyncha (Kner, 1854) AUM 43874 V5528 JN855756 Covain and Fisch-Muller (2012) Hemiancistrus medians (Kner, 1854) MHNG 2664.078 GF00-084 JF747011 Fisch-Muller et al. (2012) Guyanancistrus brevispinis MHNG 2725.099 GF00-103 JN855772 Covain and Fisch-Muller (2012) Guyanancistrus brevispinis MHNG 2621.073 SU01-121 JN855773 Covain and Fisch-Muller (2012) Guyanancistrus longispinis MHNG 2725.100 GF99-204 JN855757 Covain and Fisch-Muller (2012) Guyanancistrus niger MHNG 2722.089 GF99-185 JN855759 Covain and Fisch-Muller (2012) Guyanancistrus sp. MHNG 2679.099 MUS 300 JN855774 Covain and Fisch-Muller (2012) Hopliancistrus tricornis Isbrücker & Nijssen, 1989 MHNG 2588.051 MUS 146 JN855765 Covain and Fisch-Muller (2012) Lasiancistrus aff. caucanus MHNG 2586.043 MUS 118 JN855786 Covain and Fisch-Muller (2012) Lasiancistrus heteracanthus (Günther, 1869) MHNG 2613.037 CA 013 JN855787 Covain and Fisch-Muller (2012) Lasiancistrus planiceps (Meek & Hildebrand, 1913) STRI-01805 Stri 3526 JN855785 Covain and Fisch-Muller (2012) Lasiancistrus saetiger Armbruster 2005 MHNG 2602.016 BR98-148 JN855754 Covain and Fisch-Muller (2012) Lasiancistrus schomburgkii (Günther, 1869) MHNG 2651.009 PE08-719 JN855782 Covain and Fisch-Muller (2012) Lasiancistrus schomburgkii MHNG 2651.068 GY04-308 JN855783 Covain and Fisch-Muller (2012) Lasiancistrus schomburgkii MHNG 2710.055 PE08-277 JN855784 Covain and Fisch-Muller (2012) Lasiancistrus tentaculatus Armbruster, 2005 MhnG uncat. MUS 573 JN855788 Covain and Fisch-Muller (2012) Lithoxus lithoides Eigenmann, 1912 MHNG 2651.087 GY04-136 JN855777 Covain and Fisch-Muller (2012) Lasiancistrus pallidimaculatus Boeseman, 1982 MHNG 2621.066 SU01-096 JN855778 Covain and Fisch-Muller (2012) Lasiancistrus planquettei Boeseman, 1982 MHNG 2722.060 GF03-055 JN855779 Covain and Fisch-Muller (2012) Lithoxancistrus orinoco AUM 43725 V5246 JN855766 Covain and Fisch-Muller (2012) Lithoxancistrus orinoco AUM 42179 P4527 JN855767 Covain and Fisch-Muller (2012) Pseudancistrus barbatus MHNG 2653.059 GF00-074 JN855761 Covain and Fisch-Muller (2012) Pseudancistrus corantijniensis MHNG 2672.092 SU05-296 JN855781 Covain and Fisch-Muller (2012) Pseudancistrus depressus MHNG 2674.026 SU05-020 JN855780 Covain and Fisch-Muller (2012) Pseudancistrus genisetiger MHNG 2593.061 MUS 173 JN855764 Covain and Fisch-Muller (2012) Pseudancistrus nigrescens MHNG 2651.069 GY04-313 JN855770 Covain and Fisch-Muller (2012) Pseudancistrus nigrescens MHNG 2650.087 GY04-260 JN855771 Covain and Fisch-Muller (2012) Pseudancistrus pectegenitor AUM 42202 V5363 JN855769 Covain and Fisch-Muller (2012) Pseudancistrus pectegenitor ANSP 182801 V5433 JN855768 Covain and Fisch-Muller (2012) Pseudancistrus sidereus AUM 43443 P4871 JN855775 Covain and Fisch-Muller (2012) Pseudancistrus sidereus AUM 42180 P4537 JN855776 Covain and Fisch-Muller (2012) Pseudancistrus zawadzkii LBP 15045 61628 KJ028080 Present study Pseudancistrus sp. L17 MHNG 2586.046 MuS 132 JN855763 Covain and Fisch-Muller (2012) Pseudolithoxus cf. kelsorum MHNG 2679.043 MUS 260 JN855762 Covain and Fisch-Muller (2012) Pseudancistrus dumus (Armbruster & Provenzano, 2000) MHNG 2708.080 MUS 288 JN855760 Covain and Fisch-Muller (2012) Pseudancistrus tigris (Armbruster & Provenzano, 2000) AUM 42215 V5292 JN855758 Covain and Fisch-Muller (2012)

Pseudancistrus zawadzkii, Pseudancistrus corantijniensis, and Pseudancistrus nigrescens share the presence of whitish colored snout odontodes and a dark colored body covered with white spots. The new species can be easily distinguished from Pseudancistrus corantijniensis and Pseudancistrus nigrescens by having large hypertrophied odontodes on the posteriormost portion of the non-evertible check plates, and marginal odontodes that increase gradually in length from tip of snout to cheeks. Pseudancistrus

Biogeography and dispersal routes

Named species of the Pseudancistrus barbatus group are distributed in rivers draining to Guyana Shield into the Atlantic Ocean, and the new species described herein is from Tapajós river draining of Brazilian Shield into the Amazon. In our phylogeny, species from the eastern Guyana Shield (Pseudancistrusbarbatus and Pseudancistrus depressus) form a clade sister to a group composed of species from the western Guyana Shield (Pseudancistrus corantijniensis and Pseudancistrus nigrescens) and Amazon basin (Pseudancistrus zawadzkii and Pseudancistrus sp. L17) (Fig. 6). Therefore, based on this interpretation and our results of phylogenetic analysis, we suggested two hypotheses that could generate the distribution pattern of Pseudancistrus barbatus group extant-species. The first hypothesis is that the ancestral stock of the Pseudancistrus barbatus group was widely distributed through all Guyana Shield rivers and Amazon Brazilian Shield rivers, and the species Pseudancistrus zawadzkii and Pseudancistrussp. L17 are in the limit of the distribution for the group in Tapajós and Xingu rivers, respectively. Gaston (1998) and Hubbell (2001) suggested that when allopatric divergence is the dominant mode of speciation, many daughter species are expected to arise from geographically widespread ancestral species. This is a reasonable interpretation given that named species of the group are widespread in rivers draining Guyana Shield into the Atlantic Ocean; the new species Pseudancistrus zawadzkii are from Tapajós river drainage of Amazon Brazilian Shield; the possible new and undescribed species Pseudancistrus sp. L17 are from Xingu river which also belongs to drainages of Amazon Brazilian Shield and others possible new and undescribed species of Pseudancistrus barbatus group may be present in drainages of Guyana Shield into Amazon (Pseudancistrus sp. L220 from rio Paru; Pseudancistrus sp. L251 from rio Cuminá (rio Erepecuru);Pseudancistrus sp. L383 from rio Trombetas; Pseudancistrus sp. L440 from rio Jatapu (Seidel 2008)). However, phylogenetic and taxonomic studies are necessary to confirm that the latter undescribed species belong to Pseudancistrus barbatus group.

Figure 6.

Distribution and phylogenetic relationships of species of the Pseudancistrus barbatus group based on F-reticulon 4 gene. Based in our first hypothesis of extand-species distribution of this group the ancestral was widespread through all Guyana Shield rivers and Amazon Brazilian Shield rivers, the species Pseudancistrus zawadzkiiand Pseudancistrus sp. L17 are in the limited distribution of this group in Tapajós and Xingu rivers, drainages of Brazilian Shield into Amazon.

The second hypothesis suggests that the ancestral stock of Pseudancistrus barbatus group should have been distributed through Guyana Shield rivers and there existed several dispersal routes through Guyana and Amazon rivers, permitting that the ancestral lineages of Pseudancistrus sp. L17 and Pseudancistruszawadzkii reached the rivers of Amazon basin (see Fig. 7 for dispersal routes). Therefore, examples of connections and areas of movement among Guyana drainages and the north tributaries of Amazon basin was reported by several authors: (1) the Rupununi portal, an example of seasonal connection among Takutu and Rupununi rivers (Armbruster and Werneke 2005; Lujan and Armbruster 2011; De Souza et al. 2012); (2) the corridor among Sipalawini (Corantijn river basin) and the Paru do Oeste (Amazon basin), also connected only in the rainy season (Nijssen 1972; Lujan and Armbruster 2011); (3) the Cassiquiare Canal, a large and permanently navigable corridor between the upper Orinoco and the upper Rio Negro (Amazon) (Chernoff et al. 1991; Buckup 1993; Schaefer and Provenzano 1993; Lovejoy and Araújo 2000;Turner et al. 2004; Moyer et al. 2005; Willis et al. 2007; Winemiller et al. 2008; Winemiller and Willis 2011); (4) Proto-Berbice, a river system which had its headwaters in an ancient mountain range draining northward to Guyana system (Rupununi and Essequibo rivers) and suffered a major sedimentation, erosion and/or corrosion of the highlands and at the end of the Pliocene had its head waters captured by the Amazon system; (5) the Atlantic coastal corridors resulted in a coastal marine corridor with reduced salinity due to the westerly Amazon River discharge, coastal junctions during times of marine regressions and expanded coastal plains, and stream captures (Eigenmann 1912; Boeseman 1968; Cardoso and Montoya-Burgos 2009; Lujan and Armbruster 2011).

 

Figure 7.

Hypothesized dispersal routs between basins of the Guiana Shield and Amazon Shield of ancestror of thePseudancistrus barbatus group (based on Lujan and Armbruster 2011). Our second hypothesis of the Pseudancistrusbarbatus group extent-species distribution is based on the assumption of a widespread ancestral through all Guyana Shield rivers and dispersal events enable the ancestor of Pseudancistrus zawadzkii (red star) and Pseudancistrus sp. L17 (yellow star) to colonize the Amazon Brazilian Shield rivers in Tapajós and Xingu rivers.

Additionally, the mainstream of Amazon River can act as a permeable barrier for endemic taxa on the respective Guiana and Brazilian shields. Several genera known to tolerate more lowland conditions (e.g.Ancistrus Kner, 1854, Lasiancistrus, and Hypostomus Lacepéde, 1803) may be able to cross the Amazon basin, but such dispersal is unlikely among most species of Ancistrini (Lujan and Armbruster 2011). Also historically, epochs of cooler climate, as during glacial periods, could produce reduced precipitation, marine regressions, expansion of the coastal plain, and deepening of river channels. During such arid periods, rapids would have been more widespread, and deep-channel habitats that may currently work as barriers to fish dispersal would have been reduced (Schubert et al. 1986; Latrubesse and Franzinelli 2005;Lujan and Armbruster 2011). Drier climate will hardly change the Amazon river in a rapid, but can reduce its water flow allowing fish dispersal. Among Neotropical fishes Psectrogaster essequibensis Günther, 1864 (Characiformes: Curimatidae; see Vari (1987)), Parotocinclus aripuanensis Garavello, 1988, andPseudancistrus britskii Boeseman, 1974 (Loricariidae: Hypoptopomatinae) are species known to support dispersal via the northern Brazilian Shield.

Also, the dispersal routes around adjacent drainages of southern and northern Guyana Shield and northern parts of the Brazilian Shield could allow the dispersal of the ancestral form of Pseudancistruszawadzkii and Pseudancistrus sp. L17, as well as others ancestral species of the Pseudancistrus barbatusgroup and even species of Ancistrini (Lujan and Armbruster 2011). The movement of fish species around adjacent drainages could be explained by two hydrographic reconfiguration process: headwater capture events (geomorphological phenomenon) and marine regressions (sea level oscillation). Changes in the earth’s surface involving changes in the courses of rivers, as stream captures, portions of tributaries of a river in a watershed could be “captured” by adjacent basins resulting in isolated populations and at the

same time letting species to move, or disperse, between adjacent drainages (Almeida and Carneiro 1998;Bishop 1995; Wilkinson et al. 2006, 2010; Roxo et al. 2012). Montoya-Burgos (2003) hypothesized that dispersal (followed by allopatric population divergence) among Amazon and North-eastern coastal rivers probably occurred by temporary connections between adjacent rivers during periods of lower sea level about 6–5 Ma (see fig. 5 in Montoya-Burgos 2003). Cardoso and Montoya-Burgos (2009) suggested the same process to explain dispersal of Pseudancistrus brevispinis along coastal rivers of the Guyana. Therefore, temporary lowland connections and headwater capture events, together with the previously related hypothesis of colonization routes, likely explain the widespread distribution of the Pseudancistrusbarbatus group extant species on Guyana and Brazilian Shields, as well as how the ancestral lineages ofPseudancistrus zawadzkii and Pseudancistrus sp. L17 reached the drainages of the northern Brazilian Shield, in Tapajós and Xingu rivers.

Comparative material

Pseudancistrus barbatus (Valencienes, 1840): ANSP 177366, 2, 76.5−103.7 mm SL, Burro Burro river, Water Dog Falls, Essequibo river basin, Guyana. ANSP 189119, 3, 75.1−151.5 mm SL, Lawa river, Sipalawini, Suriname. Pseudancistrus brevispinis (Heitmans, Nijssen & Isbrücker, 1983): ANSP 189128, 3, 56.8−125.7 mm SL, Marowini river, Sipalawini, Suriname. Pseudancistrus nigrescens Eigenmann, 1912: ANSP 177379, 5, 96.4−133.5 mm SL, Burro Burro river, Water Dog Falls, Essequibo river basin, Guyana.Pseudancistrus orinoco (Isbrücker, Nijssen & Cala, 1988): ANSP 160600, 6, 68.0−78.5 mm SL, Orinoco river, Venezuela. Pseudancistrus pectegenitor Lujan, Armbruster & Sabaj, 2007: ANSP 190755, 1, 206, 2 mm SL, Ventuari river, Orinoco river basin, Venezuela. Pseudancistrus sidereus Armbruster, 2004b: ANSP 185321, 4, 148.6−154.1 mm SL, Casiquiari river, Venezuela. Pseudancistrus sp. L17: LBP 16551, 2, 75.3−101.0 mm SL; rio Xingu, Altamira, Pará State, Amazon river basin, Brazil. ANSP 193074, 3, 51.7−188.7 mm SL, Xingu river, Altamira, Pará State, Amazon river basin, Brazil. Pseudancistrus sp. ANSP 191153, 6, 49.2−75.7 mm SL, Ventuari river, Orinoco river basin, Venezuela.

 

Acknowledgements

We are grateful to M.H. Sabaj Pérez (Academy of Natural Sciences of Philadelphia) and J.S. Albert (University of Louisiana at Lafayette) for loans of specimens and curatorial assistance; to CEPTA’s team (Centro de Pesquisa Treinamento em Aquicultura - formerly Centro Nacional de Pesquisa e Conservação

de Peixes Continentais) for collecting specimens; B. Waltz and again to M.H. Sabaj Pérez for reading the manuscript and providing valuable suggestions. Fishes collected in accordance with Brazilian laws, under a permanent scientific collecting license issued to Dr. Claudio Oliveira by IcmBio/CEPTA. Research supported by Brazilian agencies FAPESP (Fundação de Amparo à Pesquisa do Estado de São Paulo, proc. 2010/01610-9 to FFR, proc. 2012/01622-2 to GSCS and proc. 2011/00269-4 to RB), MCT/CNPq (Conselho Nacional de Desenvolvimento Científico e Tecnológico) and CAPES (Coordenação de Aperfeiçoamento de Pessoal de Nível Superior). Comparative material from rio Xingu made available by iXingu Project funded by the U.S. National Science Foundation (DEB-1257813).

 

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http://zookeys.pensoft.net/articles.php?id=3788&display_type=element&element_type=7&element_id=0&element_name=Pseudancistrus

Editado Maio 1, 2016 por Vera Santos