The deepwater faunas of oceanic islands and seamounts of the Eastern Tropical Pacific are poorly known. From 11-22 September 2009 we conducted an exploration of the deepwater areas of the Isla del Coco Marine Conservation Area, Costa Rica and a nearby seamount using a manned submersible. The goal of the exploration was to characterize the habitats and biota, and conduct quantitative surveys of the deepwater portions of Isla del Coco National Park and Las Gemelas Seamount, located about 50km southwest of Isla del Coco. We completed a total of 22 submersible dives, spanning more than 80hr underwater, and collected a total of 36hr of video. We surveyed habitats from 50-402m and observed more than 45 species of fishes, some of which have not yet been described and are likely new to science. The diversity of fish species in deep water at Isla del Coco National Park was lower than the diversity of fishes in shallow water, and eight species groups accounted for more than 95% of the total fish biomass. The combined density of all fish species was higher at Las Gemelas Seamount (253 fishes/100m2) than at Isla del Coco National Park (138 fishes/100m2). The combined density of fishes in habitats comprised primarily of bedrock or large boulders outcrops was more than three times as high at Las Gemelas Seamount as it was at Isla del Coco National Park. This discrepancy was caused by the extremely high concentration of Anthiinae fishes in rocky habitats at Las Gemelas Seamount. Densities of fishes in the other habitats were similar between the two sites. Similarly, when estimates of fish density were plotted by slope categories the density was much greater on steep slopes, which were usually comprised of rock habitats. Also, the density of fishes was greatest on high rugosity habitats. Results of these submersible surveys indicate that seamounts in the tropical eastern Pacific Ocean may be an important source of biodiversity and that more quantitative surveys are needed to characterize the fauna of the region.
Key words: seamounts, deepwater habitats and fishes, submersible observations, biodiversity, census, fish assemblages.
Resumen
Las faunas de aguas profundas de islas oceánicas del Pacífico Tropical Oriental se conocen poco y de los montes submarinos nada. Del 11 al 22 de septiembre de 2009 llevamos a cabo una exploración de zonas profundos del Área de Conservación Marina Isla del Coco utilizando un submarino tripulado. El objetivo del estudio fue caracterizar los hábitats y las comunidades, y cuantificar las poblaciones de peces de profundidad en la Isla del Coco y los montes submarinos Las Gemelas, situados a 50km al suroeste de la Isla del Coco. Realizamos 22 inmersiones con el submarino, con más de 80 horas de observación submarina, y filmamos 30 horas de video. Investigamos hábitats entre 50-402m de profundidad y observamos más de 45 especies de peces, algunas de las cuales son especies nuevas para la ciencia. La diversidad de peces profundos en la Isla del Coco fue menor que en aguas someras, y ocho grupos de especies representaron más del 95% de la biomasa total de peces. La densidad combinada de peces fue 253 peces/100m2 en Las Gemelas y 138 peces/100m2 en la Isla del Coco.
Palabras clave: Isla del Coco, peces de profundidad, observaciones desde submergible, biodiversidad.
Isla del Coco, also ]]>
In recognition of the large ]]>
et al. Revista de Biología ]]>
In 2007, a conservation gap analysis was conducted, and Las Gemelas Seamount was identified as a possible location for inclusion into the system of marine reserves in ]]>
Methods
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From 11-22 September 2009, scientists from the National Geographic Society, University of Costa Rica, Moss Landing Marine Laboratories, Monterey Bay Aquarium Research Institute, Ocean Research & Conservation Association, and the University of California conducted an exploration of the deepwater areas around Isla del Coco National Park (~5°33’N, 87°02’W) and Las Gemelas Seamount (~4°59’N, 87°38’W), located about 50km southwest of Isla del Coco National Park. ]]>
DeepSee submersible to explore the water column and seafloor habitats to a depth of 400m (Cortés & Blum 2008).
At Las Gemelas Seamount, we surveyed a general area that was suggested to us by commercial fishermen. When we arrived in the general vicinity of the seamount, we conducted bathymetric surveys of the ]]>
DeepSee submersible to survey two of the shallow peaks. At Isla del Coco National Park, most submersible dives occurred at dive locations along the drop-off at the northern edge of the island, in areas normally visited by the DeepSee submersible during its regular trips with commercial passengers. At all dive sites, observers usually spent 30min to one hour exploring the area. After getting a sense of the ]]>
Submersible transects were patterned after strip transect surveys that have been commonly used to evaluate fishes in temperate environments (Stein et al. 1992, Starr
et al. 1996, Yoklavich et al. 2002). During these quantitative transects, observers looked forward and downward through the submersible dome, and recorded species composition, species-habitat associations, length distributions of fishes, and relative abundances and depth distributions of fishes. We identified and counted every fish observed in a swath that was 1m wide, for a set time period (usually 10min). Lasers that were mounted 33cm apart, on either side of the camera housing, shined parallel beams of light and allowed us to ]]>
Habitat data: Habitats were defined by a combination of substrate type and slope and rugosity of the seafloor, as described in Greene et ]]>
(1999) and Tissot et al. (2007). We used seven primary substrate codes: boulder (B), cobble (C), gravel (G), pebble (P), bedrock or rock outcrop (R), sand (S), and a code for a vertical pinnacle (T). We defined bottom type as a two-letter code representing the approximate percent cover of the two most prevalent substrata in a particular habitat patch. The first character of the code represents the substratum that accounted for at least 50% of the habitat, and the second represents the second most prevalent habitat, accounting for at least 20% of the patch (e.g., the code RB represents a ]]>
et al. 1989, Stein et al. 1992, Starr et al. 1996, Greene et al. 1999, Yoklavich et al. 2002, Tissot et al. 2007, Starr & Yoklavich 2008)
Analyses: We used only data from submersible dives that occurred at similar depths and covered similar habitats at each site to compare fish assemblages at Las Gemelas Seamount with those around Isla del Coco National Park. The dives and associated transects at Las Gemelas Seamount covered primarily rocky habitats at depths greater than 150m. Thus, for comparison purposes, we used only the nine submersible dives at Isla del Coco National Park that contained transects covering similar depths and habitats to contrast with the four submersible surveys at Las Gemelas Seamount. The comparisons ]]>
To evaluate species composition, we calculated species richness (the number of species) and diversity (Shannon-Wiener Index, Zar 1999) at each site. We then calculated ]]>
Results
Submersible dives and transects: We completed a total of 22 submersible dives. During the cruise, rough weather limited our opportunity to launch the submersible at Las Gemelas Seamount, and we were only able to conduct surveys there on only two days. We were able to launch the submersible on 12 days at Isla del Coco National Park. We conducted a total of four dives at the Las Gemelas Seamount and 18 dives around Isla del Coco National Park. Maximum depths of dives ranged from 50-402m, and dive duration averaged 3.7hr. Total duration of visual observations during dive explorations was more than 80hr. ]]>
Quantitative data were available from 16 submersible dives (Table 1). Four of these dives occurred at Las Gemelas Seamount, and 12 dives occurred around Isla del Coco National Park. A total of 38 quantitative transects were completed in this study. Transect lengths varied from 23-169m. The total area surveyed ]]>
2. At Isla del Coco National Park, we conducted 25 quantitative transects on 12 dives, and surveyed 1999m2. At Las Gemelas Seamount, we completed 13 video transects on four dives, and surveyed 1004m2.
We collected 36hr of video ]]>
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Habitat data: We encountered a total of 18 combinations of the seven substratum codes. At Isla del Coco National Park, 58% of the habitats surveyed were comprised primarily of rock and 34% were primarily sand, whereas at Las Gemelas, 73% of the transects covered rocky habitats and 25% covered sandy habitats (Tables 2, 3). Transects at Las Gemelas Seamount occurred over somewhat more rugose habitats. At Isla del Coco ]]>
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Species composition: A total of 4,544 fishes were observed from 46 taxa on the quantitative transects that occurred at depths below 50m (Table 4). All the species observed on transects above 50m have been described in Garrison (2005), and are not reported here because only a few submersible transects were conducted in shallow water. In deeper waters, observers encountered unfamiliar species, and we were unable to resolve species ]]>
Table 5). We know that some of the fishes we saw are either new species or are species that have not been reported for this region. For example, in addition to Pontinus clemensi and ]]>
Scorpaenodes rubrivinctus (recently described by Poss et al. 2010), we saw at least six other species of Scorpionfish (Scorpaenidae). Based on discussions with taxonomists, some of the unidentified species are likely to be Pontinus strigatus (identified from the Galapagos Islands by Grove & Lavenberg (1997)),
Scorpaena afuerae (identified from Ecuador by Jiménez & Béarez (2004)), Pontinus furcirhinus (described by Poss (1995)), and two of the species we observed had characteristics similar to Trachyscorpia osheri and Idiastion hageyi, new species described from the Galapagos Islands by McCosker (2008). It was not possible, however, to identify to the ]]>
Anthias noeli (Anderson & Baldwin 2000, Béarez & Jiménez Prado 2003) and some were clearly Pronotogrammus eos or P. multifasciatus. However, we saw at least two other types of fishes that were either different Anthiinae species or different morphological versions of the Anthiinae species we observed. These four or more species of Anthiids were usually in mixed aggregations on boulder or rocky habitats with medium-sized crevices.
Due primarily to the larger number of transects occurring in Isla del Coco National Park, species richness was greater there than at Las Gemelas Seamount. We encountered 28 taxa on quantitative transects at Isla del Coco National Park and 16 taxa at Las Gemelas. Because we encountered several fishes that have not yet been reported in the scientific literature, we ]]>
Table 5). Flagfins were relatively common at Isla del Coco National Park, but this taxon was absent from the Las Gemelas dives. The diversity index (H’) calculated for Isla del Coco National Park was 1.79 and species evenness (J) was 0.54. At Las Gemelas, the Shannon diversity index equaled 0.66 and species evenness was 0.24. These values include only species on quantitative transects and assumes that the several different ]]>
Fish density and biomass: The combined density of all fish species was higher at Las Gemelas Seamount (253 fishes/100m2) than at Isla del Coco National Park (138 fishes/100m2). One reason for this ]]>
2) of Anthiinae fishes (Serranids: Basslets) that we observed on submersible dives at Las Gemelas Seamount. These fish were distributed throughout all transects, as evidenced by the 100% occurrence on all dives. Threadfin Bass (Pronotogrammus multifasciatus) and other Anthiinae fishes were commonly seen at Isla del Coco National Park (58.0% and 41.7% of the ]]>
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Despite these differences, on a per-dive basis, there was no difference in biomass between the two sites when all species were lumped. The average biomass among dives at Las Gemelas (348.3 g/m2) was not significantly different than the average biomass among dives at Cocos Island (320.5 g/m2) in a two-sample t-test (p>0.05). There were differences, however, for species groups present at both locations. The ]]>
Anthias spp. complex was significantly greater at Las Gemelas than at Cocos Island in a two- sample T-test (p<0.05). The biomasses of the rest of the species groups (groupers, scorpionfishes, wrasses, codlings, threadfin basses) were not significantly different between Las Gemelas and Cocos Island (p>0.05). These same patterns occurred in biomass; the standing stock of Basslets was 21.6kg/100m2 at Las Gemelas Seamount and the Threadfin Bass comprised the largest component of biomass at Isla ]]>
2). We divided the biomass (kg/100m2) of each taxonomic group observed at Las Gemelas Seamount by the biomass of each species at Isla del Coco National Park to develop a biomass ratio for the two sites (Table 5). The biomass ratio was greater at Las Gemelas than Isla del Coco National Park in four out of the seven categories of taxa that were present at both locations.
Size frequency of fishes: We were able to estimate the lengths of 2,040 fish (Table 6). In addition to being more abundant, Basslets were significantly larger (t-test, p<0.001) at Las Gemelas Seamount than at Isla del Coco National Park. Conversely, Threadfin Bass, were more abundant and significantly larger (t-test, p<0.001) at Isla del ]]>
Although not statistically significant (K-S test, p=0.313), a plot of the percentage of total biomass by size class at each site indicates that larger fishes ]]>
Fig. 1). At Isla del Coco National Park, fishes greater than 50 cm long provide 28% of the total biomass, whereas at Las Gemelas, fishes longer than 50 cm comprise only 16% of the biomass. The difference is due to the lower numbers of medium-sized groupers and much larger numbers of small fishes at Las Gemelas Seamount. We observed the largest groupers on transects at the seamount, but relatively more groupers at Isla del Coco National Park. Also, the difference is caused by the densities of ]]>
Threadfin Bass and other Anthiinae fishes play the same role in the ecosystem (as predators of small fishes and prey of larger fishes such as groupers), and were stratified by depth, thus providing prey to larger fishes at a wide variety of ]]>
Fig. 2), and were most often observed in large aggregations around large rock boulders, usually at depths of about 180-200 m. Other species of Basslets, however, occupied generally deeper depth zones, and were most frequently observed in or near the bottom, often lodged in cracks and crevices of rock habitats. The mean depth of other Basslets (231m) was significantly different (t-test, p<0.001) than the mean depth of Threadfin Bass (181m). ]]>
Species-Habitat associations: When density of fishes is viewed by habitat type, the combined density of fishes in habitats comprised primarily of bedrock or large boulders outcrops (RR and RB habitat categories) was significantly different (Chi Square, p< 0.001) between the two areas; density was more than ]]>
Fig. 3). This discrepancy was caused by the extremely high concentration of other Anthiinae fishes in rocky habitats at Las Gemelas Seamount. Densities of fishes in the other habitats were similar between the two sites. Similarly, when estimates of fish density were plotted by slope categories (Fig. 4) the density was significantly different between the two areas (Chi Square, ]]>
Fig. 5). This is to be expected as the Anthiinae are adapted to avoid predation by taking refuge in holes and crevices throughout their range.
Occurrence of fishing gear: Lost fishing gear was observed on all submersible dives at Las Gemelas Seamount and on 50% of the dives at Isla del Coco National Park. On six submersible dives, discarded fishing line was noted on 33 occasions, during nine of the 38 quantitative transects. Aside from the observations of fishing line during quantitative transects on those dives, the presence of fishing line was noted by observers when the submersible was off transect on eight additional dives. Presence of fishing line was greatest at Las Gemelas Seamount; 30 of the 33 observations of fishing line occurred at Las Gemelas.
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Discussion
Seamounts are important features in the world’s oceans, but until recently have been poorly studied (Clark et al. 2010). An increasing amount of research has been devoted to studying the biological communities and patterns of benthic ]]>
et al. (2000) who suggested that southwest Pacific seamounts contained greater endemism than that of deep-sea vent communities.
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Although Samadi et al. (2006) indicated that their genetic studies in a relatively small area in the western Pacific might refute the concept of increased endemism on seamounts, they did agree with the hypothesis that seamounts are diversity hotspots, possessing benthic assemblages with particularly high species richness. Worm et al. (2003) provided a rationale for the increased relative abundance of organisms found on seamounts by suggesting that higher trophic level predators are found in higher ]]>
et al. (2000) also described high densities of fishes associated with seamounts and discussed concerns about the vulnerability of seamount communities to human impacts, especially with the development of large-scale bottom trawl fisheries in the deep sea. The high densities of fishes and invertebrates in the area around Isla del Coco National Park and Las Gemelas seamount signifies the need for increased protection and study of these areas, because of the ]]>
The diversity of fish species in deep water at Isla del Coco National Park was lower than the diversity of fishes in shallow water in the National Park, and eight species groups accounted for more than 95% of the biomass of fishes. Two taxa, the Threadfin ]]>
Pronotogrammus multifasciatus) and other Anthiinae species, comprised 44% of the biomass at Isla del Coco National Park. Habitats surveyed at Isla del Coco National Park included vertical rock walls and steep slopes comprised of volcanic rock outcrops and sand. Habitats surrounding Isla del Coco National Park were often highly fragmented and contained many cracks and crevices for small fishes to hide. The edge of the shelf, at about 180-220m deep contained the highest density of fishes; we often saw ]]>
]]>
The number of taxa observed in waters deeper than 50m at Isla del Coco National Park and at Las Gemelas Seamount during our study was similar to that reported by Pearcy et al. (1989) and Tissot et al. (2007) for limited number of dives in deep rocky banks in the temperate waters of Oregon, USA, but far less than the 110 taxa reported by Starr and Yoklavich (2008) for extensive submersible surveys off California, ]]>
et al. 2004), we expect that many of the deepwater taxa reported to occur in the Galapagos Islands by Grove and Lavenberg (1997) would occur near Isla del Coco National Park. Although we expect similarities in genera, we expect different species as fishes from the ]]>
Although species richness was higher at Isla del Coco National Park (due to the increased area surveyed there), the density of fishes, both in terms of numbers and biomass per unit area, was greater at Las Gemelas Seamount. Habitats we surveyed at Las Gemelas contained rich and dense communities of ]]>
2 are similar to those observed in eastern-Pacific ]]>
2 (Yoklavich et al. 2002), about 70 fish/100m2 (Tissot et al. 2007), and 65-185 fishes/100m2 (Starr & Yoklavich 2008). In the small sample size we have, we speculate that the deeper waters ]]>
In terms of fish diversity and abundance, Las Gemelas Seamount probably have a similar diversity of fishes as the deep water off Isla del Coco National Park, and overall contain a higher biomass of fishes. Importantly, however, Las Gemelas ]]>
Given the diverse ]]>
The observed differences between the two survey locations may be due to the greater depth range and variety of habitats surveyed at Isla del Coco National Park, the larger number of transects conducted at Isla del Coco National Park, or it may be an effect of the island biogeography typical of tropical islands. Until more surveys are conducted to enable an analysis of species-area ]]>
et al. Revista de Biología Tropical, this issue); this very rich invertebrate community composition indicates that habitats at Las Gemelas may be able to harbor a greater diversity and biomass of fishes than at Isla del Coco National Park.
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Given that the island groups in the Tropical Eastern Pacific (i.e., Galapagos, Malpelo, Isla del Coco) are known for endemic species (Grove and Lavenberg 1997, Garrison 2005), we expect that the seamount communities near the Tropical Eastern Pacific island groups would also be reflective of ecologically isolated communities. This highlights the need for increased study of the biological communities at Las Gemelas Seamount – to determine the role of the seamount in maintenance of marine biodiversity.
Acknowledgments
We thank Odalisca Breedy, Jorge Cortés, Shmulik Blum, Sylvia Earle, Avi Klapfer, Bruce Robison, Edith Widder, and ]]>
Argo (Undersea Hunter Group) for safe submersible operations and help with observations of fishes and macroinvertebrates. We thank John McCosker, Bob Lea, Douglas Long, Matt Craig, Rachel Arnold, and Gregor Cailliet for help with identifying species from photographs. Cheryl Barnes provided some statistical analysis and helped finalize the figures and tables. Comments from three anonymous reviewers greatly improved the manuscript. Funding was provided by the National Geographic Society, the Walton Family Foundation, the Waitt Foundation, California Sea ]]>
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*Correspondencia a: Richard M. Starr. University of California Sea Grant Program and Moss Landing Marine Laboratories, 8272 Moss Landing Road, Moss Landing, California 95039, USA; starr@mlml.calstate.edu Kristen Green. Alaska Department of Fish and Game, 304 Lake Street, Room 103, Sitka, Alaska 99835; kristen.green@alaska.gov Enric Sala. National Geographic Society, 20036 Washington, DC, USA; esala@ngs.org. Centre d’Estudis Avançats de Blanes, CSIC, 17300 Blanes, Spain
1. University of California Sea Grant Program and Moss Landing Marine Laboratories, 8272 Moss Landing Road, Moss Landing, California 95039, USA; starr@mlml.calstate.edu 2. Alaska Department of Fish and Game, 304 Lake Street, Room 103, Sitka, Alaska 99835; kristen.green@alaska.gov 3. National Geographic Society, 20036 Washington, DC, USA; esala@ngs.org 4. Centre d’Estudis Avançats de Blanes, CSIC, 17300 Blanes, Spain
Received 05-III-2012. Corrected 15-VI-2012. Accepted 24-IX-2012.