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Aereas de endemismo del Oceano Austral: un re-analisis basado en datos adicionales de hidroides bentonicos.

Southern Ocean areas of endemism: a reanalysis using benthic hydroids (Cnidaria, Hydrozoa)

Marine polar regions are often considered to have low biodiversity, a pattern generally thought to be associated with extreme abiotic factors (Clarke & Crame, 1992). However, several studies on biodiversity, biogeography and paleontology of polar regions (e.g., Beu et al., 1997; Clarke & Johnston, 2003; Adey et al., 2008) revealed greater than expected biodiversity, particularly for the Southern Ocean (SO; viz., Clarke & Johnston, 2003).

The SO is a unique oceanographic system in which the Antarctic Circumpolar Current (ACC) dominates (Barker & Thomas, 2004) and moves around the southern seas with no continental barriers. The ACC originated with the opening of the Drake Passage (ca. 30 Ma), thereby causing biogeographically and thermal isolation of the SO (Lawver & Gahagan, 2003). This, in turn, contributed to the isolation and development of endemic marine fauna (Clarke & Crame, 1989; Beu et al., 1997; Barker & Thomas, 2004; Clarke et al., 2004).

On the other hand, there are historical similarities of the Antarctic fauna to those from northern regions (Canete et al., 1999; Yasuhara et al., 2007; Kaiser et al., 2011), mainly in the Antarctic Peninsula and the subantarctic region of South America (Clarke & Johnston, 2003; Clarke et al., 2005). The connection between both continents is through the Scotia Arc, and since it will have influenced dispersal of their marine fauna, we may question exactly how isolated was the SO (Clarke et al., 2005).

Thus, several hypotheses have been proposed to explain the origin of the SO fauna: (H1) evolution in situ, (H2) derivation from adjacent deep-water basins, (H3) dispersal from South America through the Scotia Arc, and (H4) dispersal from Antarctica through the Scotia Arc (cf., Knox & Lowry, 1977). These notmutually-independent hypotheses are partially supported by fauna and geography and have been contrasted with abiotic factors of the SO (viz., Beu et al., 1997; Canete et al., 1999; Yasuhara et al., 2007; Kaiser et al., 2011).

Theoretical and practical frameworks concerning areas of endemism (cf. Harold & Mooi, 1994; Morrone, 1994; Szumik et al., 2002) are complicated in marine biogeography. The tridimensional nature of the marine realm, the dynamics of currents and oceanic fronts, the difficulties to establish thresholds in ecophysiological continuums and the amazingly diverse strategies of dispersal, all make for a unique definition of areas, or "volumes," of endemism (see Miranda & Marques, 2011). Clearly, this will also be an issue in the SO, and in which few studies examine the origin and evolution of all marine organisms, not just endemics (e.g., Clayton, 1994; Brandt, 1999; Clarke et al., 2004).

The hydroids--benthic hydrozoans of the orders Anthoathecata and Leptothecata (cf. Marques & Collins, 2004; Collins et al., 2006)--provide an example with many endemics in the SO (Pena Cantero, 2012). For example, a Parsimony Analysis of Endemicity (PAE) for the endemic SO genus Oswaldella (a single study using strict endemicity analysis) suggested four areas of endemism: (1) Magellanic Zone, (2) Antarctic Peninsula Zone, (3) Western High Antarctica Zone and (4) Eastern High Antarctica Zone (Marques & Pena Cantero, 2010). In another PAE for the SO, we used additional geographic data of endemic benthic hydroids to test previous hypotheses and to better understand the biogeography of the SO. We used a matrix of 5[degrees] latitude by 5[degrees] longitude and geographic records of 61 species of the genera Antarctoscyphus, Mixoscyphus, Oswaldella and Staurotheca (Table 1). PAE was carried out following Marques & Pena Cantero (2010), but using semistrict consensus trees. Eight areas of endemism were found for the SO, concentrated in the Magellan region, the Antarctic Peninsula, the subantarctic islands, the Ross Sea, the Weddell Sea and Wilkes Land (Figs. 1, 2). Areas I, II and V (Figs. 1, 2) are similar to the previously mentioned Magellanic and Antarctic Peninsula zones (Marques & Pena Cantero, 2010). These areas began with the ACC as a system of deep eastward currents connecting the Magellan region and Scotia Arc to the Weddell Sea, Queen Maud Land and Wilkes Land (Beu et al., 1997; Lawver & Gahagan, 2003; Marques & Pena Cantero, 2010). These currents caused dispersal towards Queen Maud Land (Marques & Pena Cantero, 2010), thereby supporting the third hypothesis of a South American origin for the SO fauna (Knox & Lowry, 1977).

Areas of endemism I, III, V and VI (Figs. 1, 2) coincide with the Scotia Arc of Marques & Pena Cantero (2010), and may be a transitional region for dispersal events of species distributed both in the Antarctic Peninsula and in the Magellan region (Pena Cantero et al., 1997; Pena Cantero & Vervoort, 2003, 2004--except the monotypic genus Mixoscyphus, which is exclusively in Antarctica (cf. Pena Cantero & Vervoort, 2005). These areas support the previously mentioned third and fourth hypotheses (Knox & Lowry, 1977). Nonetheless, this does not refute the hypothesis that vicariance influenced the isolation of the Magellanic (e.g., area V, Figs. 1, 2) from the Antarctic Peninsula (areas I and II, Figs. 1, 2). Thus, evolution in situ (hypothesis H1) may have also occurred with a fauna derived from the adjacent deepwater basin (hypothesis H2; cf. Knox & Lowry, 1977). Other areas of endemism (I, IV, VI to VIII; Figs. 1, 2) coincide with the Western High Antarctica Zone and Eastern High Antarctica Zone (Marques & Pena Cantero, 2010), and may be due to variations in depth, present oceanic currents and paleocurrents of the SO (Marques & Pena Cantero, 2010).

These results are coherent in part with ecological areas based on earlier informal biogeographic analyses (Hedgpeth, 1969; Briggs, 1974; Spalding et al., 2007). But, these results agree completely with previously hypothesized areas of endemism (Marques & Pena Cantero, 2010, cf. their Fig. 2), but now with more detail and defined subregions of those areas. These subregions suggest specific microhabitats for the benthic hydroid fauna of the SO that may be derived from dispersal or vicariant events.

If dispersal, then this suggests the formation of microhabitats, as a consequence of different strategies of larvae transportation, such as rafting of incrusting biota (e.g., on algae, wood) and oceanographic mechanisms (e.g., vortices and oceanic fronts). Both of these mechanisms are important for transportation of subantarctic/Antarctic plankton and benthos (including larvae of benthic or epipelagic organisms) along the southern polar region. If vicariance, historical and ecological barriers may have involved continental drift and climatic changes over time. Nevertheless, vicariance does not imply the absence of dispersal in the formation of the SO benthic hydroid fauna.

Considering the evolutionary history of the SO, an important question to be answered is how important were the intensity and periodicity of changes in sea level and ice (both in extent and quantity) in causing the depth and occupation of habitats along the Antarctic continental shelf (Clarke & Crame, 1989; Clarke et al., 2004). These phenomena influence marine areas of endemism because they contribute to the formation of new habitats and the availability of ecological niches, which in turn may alter the geographic distribution of the species. The SO biota has a complex evolutionary history associated with dispersal, vicariance and subsequent processes of oceanic restructuring. The use of different data sets and multiple evolutionary hypotheses will increase the explanatory power for understanding the peculiar processes leading to endemism and biogeographic patterns in the SO realm.

DOI: 103856/vol41-issue5-fulltext-20

ACKNOWLEDGEMENTS

This study was supported by CAPES (Proc. 9194/118; PROCAD and PROTAX), CNPq (Proc. 557333/ 2005-9; 490348/2006-8; 304720/2009-7, 562143/ 2010-6, 563106/2010-7, 564945/2010-2; 477156/ 2011-8), and FAPESP (Proc. 2004/09961-4; 2010/ 06927-0; 2010/52324-6; 2011/50242-5). This study is a contribution of NP-BioMar, USP. This study was also developed thanks to a research project (Ref. CTM2009-11128ANT) funded by the Ministerio de Ciencia e Innovacion of Spain and the Fondo Europeo de Desarrollo Regional (FEDER). James J. Roper revised the English in its entirety.

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Received: 28 February 2013; Accepted: 10 September 2013

Thais P. Miranda (1), Alvaro L. Pena Cantero (2) & Antonio C. Marques (1)

(1) Departamento de Zoologia, Instituto de Biociencias, Universidade de Sao Paulo Rua do Matao Trav. 14, 101, 05508-090, Sao Paulo, Brazil

(2) Instituto Cavanilles de Biodiversidad y Biologia Evolutiva, Departamento de Zoologia Universidad de Valencia, Apdo. Correos 22085, 46071 Valencia, Spain

Corresponding author: Thais P. Miranda (thaispmir@ib.usp.br)

Table 1. List of the 61 species of benthic hydroids used in PAE and
quadrants in which they are present.

Species                        Quadrant

Antarctoscyphus admirabilis    70-75[degrees]S, 5-10[degrees]W

Antarctoscyphus asymmetricus   55-60[degrees]S, 25-30[degrees]W;
                               50-55[degrees]S, 35-40[degrees]W;
                               50-55[degrees]S, 40-45[degrees]W;
                               60-65[degrees]S, 45-50[degrees]W;
Antarctoscyphus elongatus      60-65[degrees]S, 50- 55[degrees]W;
                               60-65[degrees]S, 55-60[degrees]W;
                               60-65[degrees]S, 60-65[degrees]W;
                               65-70[degrees]S, 90-95[degrees]W
                               65-70[degrees]S, 5-10[degrees]W;
                               70-75[degrees]S, 5-10[degrees]W;
                               70-75[degrees]S, 10-15[degrees]W;
                               70-75[degrees]S, 20-25[degrees]W;
                               55-60[degrees]S, 25- 30[degrees]W;
                               70-75[degrees]S, 25-30[degrees]W;
                               75-80[degrees]S, 25-30[degrees]W;
                               50-55[degrees]S, 35-40[degrees]W;
                               55-60[degrees]S, 35-40[degrees]W; 50-
                               55[degrees]S, 40-45[degrees]W;
                               75-80[degrees]S, 45-50[degrees]W;
                               60-65[degrees]S, 50-55[degrees]W;
                               75-80[degrees]S, 50-55[degrees]W;
                               60-65[degrees]S, 55-

                               60[degrees]W; 60-65[degrees]S,
                               60-65[degrees]W; 65-70[degrees]S,
                               60-65[degrees]W; 70-75[degrees]S,
                               60-65[degrees]W; 65-70[degrees]S,
                               65-70[degrees]W; 70- 75[degrees]S,
                               95-100[degrees]W; 75-80[degrees]S,
                               165-170[degrees]W; 50-55[degrees]S,
                               0-5[degrees]E; 45-50[degrees]S,
                               65-70[degrees]E; 65-70[degrees]S, 110-
                               115[degrees]E; 65-70[degrees]S,
                               135-140[degrees]E;65-70[degrees]S,
                               140-145[degrees]E; 75-80[degrees]S,
                               165-170[degrees]E;
                               70-75[degrees]S, 170-175[degrees]E

Antarctoscyphus encarnae       60-65[degrees]S, 55-60[degrees]W

Antarctoscyphus fragilis       70-75[degrees]S, 25-30[degrees]W
                               70-75[degrees]S, 5-10[degrees]W;
                               70-75[degrees]S, 10-15[degrees]W;
                               70-75[degrees]S, 20-25[degrees]W;
                               55-60[degrees]S, 25-30[degrees]W;
                               50-55[degrees]S, 35-

Antarctoscyphus grandis        40[degrees]W; 60-65[degrees]S,
                               50-55[degrees]W; 70-75[degrees]S,
                               50-55[degrees]W; 75-80[degrees]S,
                               50-55[degrees]W; 60-65[degrees]S,
                               55-60[degrees]W; 60- 65[degrees]S,
                               60-65[degrees]W; 65-70[degrees]S,
                               90-95[degrees]W; 75-80[degrees]S,
                               175-180[degrees]W; 65-70[degrees]S,
                               135-140[degrees]E; 65-70[degrees]S, 140-
                               145[degrees]E; 70-75[degrees]S,
                               170-175[degrees]E

Antarctoscyphus gruzovi        60-65[degrees]S, 50-55[degrees]W;
                               60-65[degrees]S, 55-60[degrees]W

Antarctoscyphus mawsoni        55-60[degrees]S, 25-30[degrees]W;
                               60-65[degrees]S, 50-55[degrees]W;
                               60-65[degrees]S, 55-60[degrees]W;
                               60-65[degrees]S, 60-65[degrees]W;
                               65-70[degrees]S, 65- 70[degrees]W;
                               70-75[degrees]S, 175-180[degrees]W;
                               65-70[degrees]S, 140-145[degrees]E;
                               70-75[degrees]S, 170-175[degrees]E
                               65-70[degrees]S, 5-10[degrees]W;
                               70-75[degrees]S, 10-15[degrees]W;
                               70-75[degrees]S, 20-25[degrees]W;
                               55-60[degrees]S, 25-30[degrees]W;
                               75-80[degrees]S, 25- 30[degrees]W;
                               50-55[degrees]S, 35-40[degrees]W;
                               60-65[degrees]S, 40-45[degrees]W;
                               75-80[degrees]S, 45-50[degrees]W;
                               60-65[degrees]S, 50-55[degrees]W; 75-
                               80[degrees]S, 50-55[degrees]W;
                               60-65[degrees]S, 55-60[degrees]W;
                               60-65[degrees]S, 60-65[degrees]W;
                               65-70[degrees]S, 60-65[degrees]W;
                               70-75[degrees]S, 60-

Antarctoscyphus spiralis       65[degrees]W; 65-70[degrees]S,
                               85-90[degrees]W; 65-70[degrees]S,
                               90-95[degrees]W; 70-75[degrees]S,
                               95-100[degrees]W; 75-80[degrees]S,
                               170-175[degrees]W; 70- 75[degrees]S,
                               175-180[degrees]W; 75-80[degrees]S,
                               175-180[degrees]W; 50-55[degrees]S,
                               0-5[degrees]E; 65-70[degrees]S,
                               135-140[degrees]E; 65-70[degrees]S, 140-
                               145[degrees]E; 65-70[degrees]S,
                               160-165[degrees]E; 70-75[degrees]S,
                               165-170[degrees]E; 75-80[degrees]S,
                               165-170[degrees]E; 70-75[degrees]S,
                               170-175[degrees]E; 70-75[degrees]S,
                               175-180[degrees]E

Mixoscyphus antarcticus        60-65[degrees]S, 60-65[degrees]W

Oswaldella antarctica          50-55[degrees]S, 55-60[degrees]W;
                               60-65[degrees]S, 55-60[degrees]W;
                               65-70[degrees]S, 90-95[degrees]W;
                               65-70[degrees]S, 135-140[degrees]E;
                               65-70[degrees]S, 140- 145[degrees]E

Oswaldella bifurca             75-80[degrees]S, 55-60[degrees]W;
                               70-75[degrees]S, 60-65[degrees]W;
                               75-80[degrees]S, 160-165[degrees]W;
                               70-75[degrees]S, 175-180[degrees]W;
                               75-80[degrees]S, 175-180[degrees]W;
                               65-70[degrees]S, 160-165[degrees]E;
                               70-75[degrees]S, 170-175[degrees]E

Oswaldella billardi            75-80[degrees]S, 30-35[degrees]W;
                               65-70[degrees]S, 90-95[degrees]E;
                               65-70[degrees]S, 110-115[degrees]E;
                               65-70[degrees]S, 135-140[degrees]E;
                               65-70[degrees]S, 140-145[degrees]E

Oswaldella blanconae           60-65[degrees]S, 60-65[degrees]W;
                               70-75[degrees]S, 170-175[degrees]E

Oswaldella crassa              60-65[degrees]S, 55-60[degrees]W

Oswaldella curiosa             60-65[degrees]S, 55-60[degrees]W

Oswaldella delicata            60-65[degrees]S, 50-55[degrees]W;
                               70-75[degrees]S, 50-55[degrees]W;
                               75-80[degrees]S, 50-55[degrees]W;
                               60-65[degrees]S, 55-60[degrees]W;
                               75-80[degrees]S, 55- 60[degrees]W;
                               60-65[degrees]S, 60-65[degrees]W;
                               75-80[degrees]S, 165-170[degrees]W

Oswaldella elongata            50-55[degrees]S, 30-35[degrees]W;
                               50-55[degrees]S, 35-40[degrees]W;
                               50-55[degrees]S, 50-55[degrees]W

Oswaldella encarnae            75-80[degrees]S, 45-50[degrees]W;
                               75-80[degrees]S, 50-55[degrees]W;
                               75-80[degrees]S, 55-60[degrees]W
                               50-55[degrees]S, 0-5[degrees]W;
                               70-75[degrees]S, 5-10[degrees]W;
                               70-75[degrees]S, 10-15[degrees]W;
                               70-75[degrees]S, 15-20[degrees]W;
                               70-75[degrees]S, 20-25[degrees]W;

Oswaldella erratum             55-60[degrees]S, 25-30[degrees]W;
                               70-75[degrees]S, 25-30[degrees]W;
                               75-80[degrees]S, 25-30[degrees]W;
                               75-80[degrees]S, 30-35[degrees]W;
                               60-65[degrees]S, 50-55[degrees]W;
                               70-75[degrees]S, 50-55[degrees]W;
                               60-65[degrees]S, 55-60[degrees]W;
                               60-65[degrees]S, 60-65[degrees]W;
                               65-70[degrees]S, 65-70[degrees]W; 50-
                               55[degrees]S, 0-5[degrees]E;65-
                               70[degrees]S, 10-15[degrees]E

Oswaldella frigida             60-65[degrees]S, 55-60[degrees]W;
                               60-65[degrees]S, 60-65[degrees]W

Oswaldella garciacarrascosai   75-80[degrees]S, 50-55[degrees]W

Oswaldella gracilis            75-80[degrees]S, 55-60[degrees]W

Oswaldella grandis             75-80[degrees]S, 45-50[degrees]W; 75-
                               80[degrees]S, 50-55[degrees]W; 60-
                               65[degrees]S, 55-60[degrees]W; 60-
                               65[degrees]S, 60-65[degrees]W

Oswaldella herwigi             50-55[degrees]S, 55-60[degrees]W;
                               50-55[degrees]S, 70-75[degrees]W

Oswaldella incognita           55-60[degrees]S, 40-45[degrees]W;
                               60-65[degrees]S, 55-60[degrees]W;
                               60-65[degrees]S, 60-65[degrees]W;
                               65-70[degrees]S, 160-165[degrees]W

Oswaldella laertesi            70-75[degrees]S, 170-175[degrees]E

Oswaldella medeae              60-65[degrees]S, 50-55[degrees]W; 70-
                               75[degrees]S, 175-180[degrees]W; 70-
                               75[degrees]S, 175-180[degrees]E

Oswaldella monomammillata      60-65[degrees]S, 50-55[degrees]W

Oswaldella niobae              60-65[degrees]S, 55-60[degrees]W

Oswaldella obscura             70-75[degrees]S, 5-10[degrees]W;
                               70-75[degrees]S, 20-25[degrees]W;
                               75-80[degrees]S, 45-50[degrees]W

Oswaldella rigida              70-75[degrees]S, 5-10[degrees]W;
                               70-75[degrees]S, 10-15[degrees]W;
                               70-75[degrees]S, 15-20[degrees]W;
                               70-75[degrees]S, 20-25[degrees]W;
                               75-80[degrees]S, 25- 30[degrees]W

Oswaldella shetlandica         55-60[degrees]S, 40-45[degrees]W;
                               60-65[degrees]S, 50-55[degrees]W;
                               60-65[degrees]S, 55-60[degrees]W;
                               50-55[degrees]S, 60-65[degrees]W;
                               55-60[degrees]S, 60- 65[degrees]W;
                               60-65[degrees]S, 60-65[degrees]W;
                               65-70[degrees]S, 60-65[degrees]W
                               70-75[degrees]S, 5-10[degrees]W;
                               70-75[degrees]S, 10-15[degrees]W;
                               60-65[degrees]S, 50-55[degrees]W;
                               60-65[degrees]S, 55-60[degrees]W;
                               60-65[degrees]S, 60-

Oswaldella stepanjantsae       65[degrees]W; 75-80[degrees]S,
                               170-175[degrees]W; 65-70[degrees]S,
                               110-115[degrees]E; 65-70[degrees]S,
                               160-165[degrees]E; 75-80[degrees]S,
                               165-170[degrees]E; 70-75[degrees]S,
                               170-175[degrees]E; 70-75[degrees]S,
                               175-180[degrees]E

Oswaldella terranovae          70-75[degrees]S, 170-175[degrees]E

Oswaldella tottoni             70-75[degrees]S, 10-15[degrees]W;
                               75-80[degrees]S, 160-165[degrees]E

Oswaldella vervoorti           55-60[degrees]S, 40-45[degrees]W;
                               60-65[degrees]S, 55-60[degrees]W;
                               60-65[degrees]S, 60-65[degrees]W;
                               50-55[degrees]S, 0-5[degrees]E

Staurotheca abyssalis          55-60[degrees]S, 55-60[degrees]W

Staurotheca affnis             50-55[degrees]S, 35-40[degrees]W

Staurotheca amphorophora       50-55[degrees]S, 35-40[degrees]W;
                               50-55[degrees]S, 40-45[degrees]W
                               70-75[degrees]S, 0-5[degrees]W;
                               70-75[degrees]S, 10-15[degrees]W;
                               70-75[degrees]S, 15-20[degrees]W;
                               75-80[degrees]S, 25-30[degrees]W;
                               50-55[degrees]S, 35- 40[degrees]W;
                               75-80[degrees]S, 45-50[degrees]W;
                               75-80[degrees]S, 50-55[degrees]W;
                               60-65[degrees]S, 55-60[degrees]W;
                               60-65[degrees]S, 60-65[degrees]W; 65-

Staurotheca antarctica         70[degrees]S, 60-65[degrees]W;
                               65-70[degrees]S, 65-70[degrees]W;
                               50-55[degrees]S, 70-75[degrees]W;
                               65-70[degrees]S, 70-75[degrees]W;
                               70-75[degrees]S, 80- 85[degrees]W;
                               75-80[degrees]S, 160-165[degrees]W;
                               75-80[degrees]S, 170-175[degrees]W;
                               70-75[degrees]S, 175-180[degrees]W;
                               75-80[degrees]S, 175- 180[degrees]W;
                               65-70[degrees]S, 5-10[degrees]E;
                               65-70[degrees]S, 10-15[degrees]E;
                               65-70[degrees]S, 110-115[degrees]E;
                               65-70[degrees]S, 135-140[degrees]E; 65-
                               70[degrees]S, 140-145[degrees]E;
                               75-80[degrees]S, 165-170[degrees]E;
                               70-75[degrees]S, 170-175[degrees]E

Staurotheca australis          65-70[degrees]S, 5-10[degrees]E;
                               65-70[degrees]S, 10-15[degrees]E
                               55-60[degrees]S, 25-30[degrees]W;
                               60-65[degrees]S, 50-55[degrees]W;
                               55-60[degrees]S, 55-60[degrees]W;
                               60-65[degrees]S, 55-60[degrees]W;
                               60-65[degrees]S, 60-

Staurotheca compressa          65[degrees]W; 65-70[degrees]S,
                               140-145[degrees]W; 65-70[degrees]S,
                               160-165[degrees]W; 65-70[degrees]S,
                               140-145[degrees]E; 65-70[degrees]S,
                               160-165[degrees]E; 70-75[degrees]S
                               165-170[degrees]E

Staurotheca cornuta            60-65[degrees]S, 45-50[degrees]W;
                               60-65[degrees]S, 55-60[degrees]W
                               60-65[degrees]S, 50-55[degrees]W;
                               60-65[degrees]S, 55-60[degrees]W;
                               60-65[degrees]S, 60-65[degrees]W;
                               65-70[degrees]S, 160-165[degrees]W;
                               70-75[degrees]S,

Staurotheca densa              170-175[degrees]E; 70-75[degrees]S,
                               175-180[degrees]E 70-75[degrees]S,
                               5-10[degrees]W; 55-60[degrees]S,
                               25-30[degrees]W; 70-75[degrees]S,
                               25-30[degrees]W; 50-55[degrees]S,
                               30-35[degrees]W; 55-60[degrees]S, 30-
                               35[degrees]W; 50-55[degrees]S,
                               35-40[degrees]W; 50-55[degrees]S,
                               40-45[degrees]W; 60-65[degrees]S,
                               45-50[degrees]W; 60-65[degrees]S,
                               55-60[degrees]W; 60-

Staurotheca dichotoma          65[degrees]S, 60-65[degrees]W;
                               70-75[degrees]S, 80-85[degrees]W;
                               65-70[degrees]S, 90-95[degrees]W;
                               75-80[degrees]S, 170-175[degrees]W;
                               70-75[degrees]S, 175- 180[degrees]W;
                               50-55[degrees]S, 0-5[degrees]E;
                               65-70[degrees]S, 5-10[degrees]E;
                               65-70[degrees]S, 10-15[degrees]E;
                               45-50[degrees]S, 35-40[degrees]E;
                               65-70[degrees]S, 160- 165[degrees]E;
                               75-80[degrees]S, 160-165[degrees]E;
                               75-80[degrees]S, 165-170[degrees]E;
                               70-75[degrees]S, 170-175[degrees]E

Staurotheca echinocarpa        45-50[degrees]S, 65-70[degrees]E;
                               45-50[degrees]S, 70-75[degrees]E
                               70-75[degrees]S, 5-10[degrees]W;
                               70-75[degrees]S, 10-15[degrees]W;
                               70-75[degrees]S, 20-15[degrees]W;
                               55-60[degrees]S, 25-30[degrees]W;
                               50-55[degrees]S, 30-

Staurotheca frigida            35[degrees]W; 55-60[degrees]S,
                               30-35[degrees]W; 50-55[degrees]S,
                               35-40[degrees]W; 55-60[degrees]S,
                               35-40[degrees]W; 50-55[degrees]S,
                               40-45[degrees]W; 60- 65[degrees]S,
                               40-45[degrees]W; 60-65[degrees]S,
                               45-50[degrees]W; 60-65[degrees]S,
                               60-65[degrees]W; 65-70[degrees]S,
                               65-70[degrees]W; 75-80[degrees]S, 160-
                               165[degrees]W; 65-70[degrees]S,
                               5-10[degrees]E; 65-70[degrees]S,
                               110-115[degrees]E; 70-75[degrees]S,
                               170-175[degrees]E 70-75[degrees]S,
                               0-5[degrees]W; 70-75[degrees]S,
                               10-15[degrees]W; 70-75[degrees]S,
                               20-25[degrees]W; 55-60[degrees]S,
                               25-30[degrees]W; 50-55[degrees]S, 35-
                               40[degrees]W; 60-65[degrees]S,
                               40-45[degrees]W; 75-80[degrees]S,
                               50-55[degrees]W; 60-65[degrees]S,
                               55-60[degrees]W; 60-65[degrees]S,
                               60-65[degrees]W; 65-

Staurotheca glomulosa          70[degrees]S, 60-65[degrees]W;
                               65-70[degrees]S, 70-75[degrees]W;
                               65-70[degrees]S, 160-165[degrees]W;
                               75-80[degrees]S, 160-165[degrees]W;
                               75-80[degrees]S, 175- 180[degrees]W;
                               65-70[degrees]S, 0-5[degrees]E;
                               65-70[degrees]S, 5-10[degrees]E;
                               65-70[degrees]S, 10-15[degrees]E;
                               65-70[degrees]S, 110-115[degrees]E;
                               65-70[degrees]S, 160-165[degrees]E;
                               70-75[degrees]S, 170-175[degrees]E;
                               70-75[degrees]S, 175-180[degrees]E

Staurotheca jaderholmi         50-55[degrees]S, 50-55[degrees]W;
                               50-55[degrees]S, 55-60[degrees]W;
                               50-55[degrees]S, 60-65[degrees]W;
                               50-55[degrees]S, 65-70[degrees]W;
                               50-55[degrees]S, 70- 75[degrees]W;
                               50-55[degrees]S, 75-80[degrees]W;
                               60-65[degrees]S, 90-95[degrees]W;
                               65-70[degrees]S, 135-140[degrees]E;
                               65-70[degrees]S, 140-145[degrees]E

Staurotheca juncea             70-75[degrees]S, 95-100[degrees]W

Staurotheca multifurcata       55-60[degrees]S, 25-30[degrees]W;
                               50-55[degrees]S, 30-35[degrees]W;
                               55-60[degrees]S, 30-35[degrees]W;
                               50-55[degrees]S, 35-40[degrees]W;
                               55-60[degrees]S, 35- 40W;
                               50-55[degrees]S, 40-45[degrees]W
                               70-75[degrees]S, 0-5[degrees]W;
                               70-75[degrees]S, 5-10[degrees]W;
                               70-75[degrees]S, 10-15[degrees]W;
                               60-65[degrees]S, 55-60[degrees]W;
                               75-80[degrees]S, 60-65[degrees]W;

Staurotheca nonscripta         75-80[degrees]S, 165-170[degrees]W;
                               75-80[degrees]S, 170-175[degrees]W;
                               65-70[degrees]S, 0-5[degrees]E;
                               65-70[degrees]S, 5-10[degrees]E;
                               65-70[degrees]S, 10- 15[degrees]E;
                               75-80[degrees]S, 160-165[degrees]E;
                               70-75[degrees]S, 165-170[degrees]E;
                               75-80[degrees]S, 165-170[degrees]E;
                               70-75[degrees]S, 170-175[degrees]E
                               70-75[degrees]S, 5-10[degrees]W;
                               70-75[degrees]S, 10-15[degrees]W;
                               70-75[degrees]S, 20-25[degrees]W;
                               55-60[degrees]S, 25-30[degrees]W;
                               75-80[degrees]S, 25-

Staurotheca pachyclada         30[degrees]W; 60-65[degrees]S,
                               50-55[degrees]W; 60-65[degrees]S,
                               55-60[degrees]W; 60-65[degrees]S,
                               60-65[degrees]W; 65-70[degrees]S,
                               160-165[degrees]W; 75- 80[degrees]S,
                               160-165[degrees]W; 75-80[degrees]S,
                               175-180[degrees]W; 70-75[degrees]S,
                               170-175[degrees]E

Staurotheca plana              70-75[degrees]S, 5-10[degrees]W;
                               70-75[degrees]S, 10-15[degrees]W

Staurotheca polarsteni         70-75[degrees]S, 10-15[degrees]W;
                               70-75[degrees]S, 15-20[degrees]W;
                               70-75[degrees]S, 20-25[degrees]W;
                               75-80[degrees]S, 25-30[degrees]W;
                               70-75[degrees]S, 30- 35[degrees]W;
                               60-65[degrees]S, 60-65[degrees]W;
                               65-70[degrees]S, 0-5[degrees]E;
                               65-70[degrees]S, 5-10[degrees]E;
                               65-70[degrees]S, 10-15[degrees]E

Staurotheca profunda           50-55S 55-60W

Staurotheca stolonifera        70-75[degrees]S, 5-10[degrees]W;
                               70-75[degrees]S, 80-85[degrees]W

Staurotheca undosiparietina    50-55[degrees]S, 30-35[degrees]W;
                               50-55[degrees]S, 35-40[degrees]W;
                               50-55[degrees]S, 40-45[degrees]W;
                               60-65[degrees]S, 50-55[degrees]W;
                               60-65[degrees]S, 55- 60[degrees]W

Staurotheca vanhoeffeni        70-75[degrees]S, 0-5[degrees]W;
                               70-75[degrees]S, 5-10[degrees]W;
                               70-75[degrees]S, 10-15[degrees]W;
                               60-65[degrees]S, 50-55[degrees]W;
                               60-65[degrees]S, 55-60[degrees]W;
                               60-65[degrees]S, 60-65[degrees]W;
                               65-70[degrees]S, 5-10[degrees]E;
                               50-55[degrees]S, 70-75[degrees]E;
                               70-75[degrees]S, 170-175[degrees]E

Staurotheca vervoorti          50-55[degrees]S, 30-35[degrees]W;
                               50-55[degrees]S, 55-60[degrees]W
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Article Details
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Title Annotation:articulo en ingles
Author:Miranda, Thais P.; Pena Cantero, Alvaro L.; Marques, Antonio C.
Publication:Latin American Journal of Aquatic Research
Date:Nov 1, 2013
Words:4656
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