Long-term comparison of the fish community in a Costa Rican rocky shore marine reserve.
One strategy to conserve tropical marine biodiversity, including rocky shore fish communities, is through the establishment of no-take marine reserves (NRC 2001, Sobel & Dahlgren 2004). Marine protected areas may provide spatial refuge from harvest for adult fish, improve reproduction or recruitment by protecting vulnerable spawning areas or nursery sites, or prevent habitat destruction resulting from human activities. Benefits of marine protected areas, including increased density, biomass, and average body size of target species and increased community diversity relative to unprotected or pre-protection sites, have been well documented in diverse contexts (Halpern & Warner 2002, Halpern 2003, Lubchenco et al. 2003). In addition, the effects of reserve designation appear to operate independent of reserve size, so that even relatively small marine reserves can contribute to marine conservation goals (Halpern 2003).
We studied fish community composition in a rocky shore marine reserve located at the mouth of the Gulf of Nicoya, Costa Rica in the Panamic province of the Tropical Eastern Pacific. Our objectives were: 1) to document fish community structure present in the subtidal rocky shore habitat at the site and to relate our findings to other studies of fish community structure in the region, and 2) to assess change in fish abundance and community composition at the site following an eleven-year period of marine protection.
MATERIALS AND METHODS
Study site: The study was carried out in the Punta Leona Biological Reserve on the central Pacific coast of Costa Rica (9[degrees]40'41" - 9[degrees]40'44" N and 84[degrees]39'30"-84[degrees]40'15" W). The reserve includes approximately 3.3km of coastline comprised primarily of Mesozoic volcanic and sedimentary rock (Castillo-Munoz 1983). Three small sandy beaches, each < 800m in length, are located among the rocky outcroppings in the reserve. In April 1994, the marine environment around the southernmost beach was declared the Playa Blanca Marine Reserve, and a zone of absolute protection was established in the waters extending from the shoreline to a depth of 6m at low tide along approximately 1.3km of coastline. Since then, commercial and recreational fishing has been prohibited and boat traffic within the reserve has been limited as much as possible by local lifeguards. In March of 2010, Playa Blanca was one of only two beaches nationally to earn five stars in the Costa Rican Tourism Institute's Blue Flag Ecological Program (Programa Bandera Azul Ecologica). The award recognizes the quality of marine resources at the site and the Punta Leona administration's commitment to protection and management of the beach and surrounding marine environment.
Fish surveys: We conducted underwater visual surveys of the fish community in shallow (1.0-3.5m deep) sub-tidal habitat along six strip transects selected to represent the diversity of habitats present in the Playa Blanca Marine Reserve. Transects were 50m long x 10m wide. Surveys were conducted during March and April, the last two months of the dry season, in 1995 and 2006. A total of 124 fish surveys were conducted (60 in 1995, 64 in 2006) with each of the six transects surveyed at least ten times in both years.
Visual survey methods were adapted from Williams (1982) and Ferreira et al. (2001). Using snorkeling equipment, a single surveyor recorded the presence and abundance of fishes observed while swimming in a zigzag pattern across the width of each strip transect. To maintain correct transect orientation and width, a 50m tape measure was placed along the centerline and borders were marked with painted rocks or flags at 5m intervals. In addition, the surveyor actively counted fin kicks and kept visual reference with the transect centerline throughout the survey. Surveyors actively searched for fish under rock outcroppings and within crevices or caves. Surveys averaged 20min in duration and were conducted only within two hours of low tide.
Some fishes were difficult to identify to the species level while conducting underwater visual surveys. For morphologically similar species whose identification is based on meristic characteristics (e.g., Diodon holocanthus/hystrix, Kyphosus elegans/analogus and Mugil cephalis/curema), we recorded individuals only to genus-level and conservatively counted species richness as one for each genus. Other species occurred together in large, mixed aggregations (e.g., Haemulon scudderii, maculicauda, steindachneri and flaviguttatum) where the presence of individual species could be verified, but accurately assigning each individual to a species was not feasible. Here we chose to lump species into "species groups" for analysis (Table 1) but included each positively identified species in our counts of species richness. Data on changes in abundance of species within species groups should be interpreted cautiously, as an increase or decrease in abundance of the group does not necessarily imply that all species within the group followed the same trend. However, we believe this approach is conservative and preferable to erroneously generating species-specific results based on inaccurate counts.
We classified each fish species/species group into trophic groups using the categories applied by Dominici-Arosemena & Wolff (2006). We also modified the mobility categories applied by Floeter et al. (2004) and classified each species/species group as "highly mobile" or "relatively sedentary."
Data analysis: Fish abundance, species richness, Shannon's index of community diversity (H = -[summation][P.sub.i] (ln[p.sub.i])) and Simpson's reciprocal index of diversity (D = 1/[summation][([n.sub.i]/N).sup.2]) were calculated for each survey (n = 124), then averaged for each transect (n = 6) within the two survey years. Paired-t tests were employed to compare the variable means among years using the six transect means as replicates.
We tested for significant change in community composition among years using analysis of similarity (ANOSIM). Community composition varied significantly among transects within years ([R.sub.1995] = 0.6854 and [R.sub.2006] = 0.8054, p-values < 0.0001); therefore, we tested for change in community composition among years separately for each of the six transects to remove site effects. Each survey was assigned to an a priori defined group (survey year, 1995 or 2006). ANOSIM is a permutation technique that tests the a priori defined groups against randomly generated groups. The R-statistic generated by ANOSIM is a measure of (dis) similarity of the defined groups, where a value of zero (0) indicates no difference among groups and a value of one (1) indicates that all samples within groups are more similar to one another than to samples in other groups. We conducted ANOSIM using the Bray-Curtis similarity index and 10 000 permutations; we report R statistics and uncorrected p-values for each transect.
We used a variety of techniques to assess which species/species groups were responsible for change in community composition over time. First, we used similarity percentages (SIMPER) analysis to identify which species/species groups made the greatest relative contribution to dissimilarity within transects among years. In addition, after pooling data for all transects, we classified each species/species group as dominant, common, uncommon, or rare in each year (and for the pooled total) by plotting their relative abundance versus frequency of occurrence in surveys. Dominant species had high relative abundance (>1.5%) and frequency of occurrence (detected in >60% of surveys). Common species relative abundance ranged from 0.20 to 11.12% and frequency of occurrence from 23 to 68%; uncommon species, 0.08-0.92% and 5-30%. Species were classified as uncommon or common based on the combination of values and their position on the graph. Rare species had low relative abundance (< 0.1%) and frequency of occurrence (< 12%). Finally, the abundance of each species/species group was compared among years using the transect means as replicates and paired-t tests or nonparametric Wilcoxon paired-sample tests as appropriate. Analyses were performed using SYSTAT 12.0 (SYSTAT 2010) and PAST (version 2.02) software (Hammer et al. 2001).
Fish community composition: We recorded a total of 31 404 sightings of 72 fish species representing 30 families in the Playa Blanca Marine Reserve (Table 1). Mean abundance of fishes observed per survey was 253.3 (SD=165.8) with a range from 2 to 954, while mean species richness per survey was 18.8 (6.7) and ranged from 2 to 31 species.
Using the data pooled for both years, we classified each species/species group as dominant, common, uncommon or rare according to its relative abundance and frequency of occurrence (Fig. 1a). Eleven dominant species/ species groups (Abudefduf troschelii, Bodianus diplotaenia, Chromis atrilobata, Chaetodon humeralis, Haemulon scudderii/maculicauda/ steindachneri/flaviguttatum, Halichoeres notospilus/nicholsi, Sargocentron suborbitalis, Scarus rubroviolaceus/perrico/ghobban/compressus, Stegastes flavilatum/acapulcoensis, Sufflamen verres and Thalassoma lucasanum) accounted for 74.1% of total fish abundance. At the family level, Pomacentridae was most dominant, accounting for 42.5% of all individuals, with Labridae (16.6%) and Haemulidae (14.8%) the next highest ranking families.
[FIGURE 1 OMITTED]
Twenty-three common species/species groups were characterized by moderate abundance and frequency of occurrence. Some common species (e.g., Caranx sexfasciatus/ caballus, Lutjanus guttatus and Kyphosus sp.) were encountered less frequently but in large aggregations, while others were encountered in a high percentage of all surveys but as solitary individuals (e.g., species in Diodontidae, Fistulariidae, Serranidae and Tetradontidae). Common species accounted for 23.4% of total fish abundance.
Twenty-eight species/species groups were classified as uncommon or rare. Fifteen uncommon species had relative abundances from 0.08% to 0.62% and frequency of occurrence [less than or equal to] 21%. Uncommon species accounted for 2.2% of total fish abundance. Thirteen rare species had both low relative abundance (< 0.1%) and frequency of occurrence (< 12%) and accounted for just 0.3% of total abundance. The rarest families were Dasyatidae, Scorpenidae, Zanclidae and Ostraciidae, each represented by a single species accounting for < 0.01% of total abundance.
In terms of trophic groups, nearly one-third of fish species were carnivores (25 species, 19.2% total fish abundance), yet herbivores (11 species) accounted for the greatest proportion of total fish abundance (23.8%). Overall, the distribution of species among trophic groups was quite equitable: omnivores (19.4%, 13 species), invertivores (18.8%, 11 species), planktivores (15.6%, 6 species) and piscivores (3.2%, 6 species).
Community change over time: We recorded 12 893 sightings of 61 species from 26 families in 1995 and 18 511 sightings of 69 species from 28 families in 2006. Fish abundance (paired t-test, df = 5, t = -2.575, p = 0.050) and species richness (paired t-test, df = 5, t = -2.571, p = 0.050) were significantly greater in 2006 than 1995. Shannon's index of diversity, which is sensitive to changes in rare species, was significantly greater in 2006 (paired t-test, df = 5, t = -2.624, p = 0.047), but there was no difference in Simpson's index of diversity, which is sensitive to changes in the most abundant species among years (paired t-test, df = 5, t = -1.873, p = 0.120).
We detected significant change in fish community composition (all p < 0.01) among years in each of the six transects (RT1=0.2434; [R.sub.T2] = 0.7153; [R.sub.T3] = 0.7078; [R.sub.T4] = 0.6967; [R.sub.T5] = 0.8635; [R.sub.T6] = 0.7292). The five species/ species groups that contributed most to community dissimilarity among years within each transect are listed in Table 2. These species/ species groups accounted for 47.7 to 84.4% of the dissimilarity among years within transects. Much of the variation in community composition among years resulted from variation in the abundance of a few dominant species. For example, Abudefduf troschelli was identified as an influential species in all six transects; Thalassoma lucasanum and Chromis atrilobata in five; and Stegastes flavilatus/acapulcoensis in four. These were among the most abundant and omnipresent species in the study, but their average abundance varied substantially among transects and years. Not all species identified as influential by SIMPER were dominant; some, including Lutjanus guttatus, Eucinostomus currani and Nicholsina denticulate, were uncommon in 1995 but were more frequent and abundant and classified as common in 2006.
We also assessed change in community composition by comparing individual species/ species group classifications among years. Thirteen of the 19 species whose classification changed among years were more frequent and abundant in 2006 (Table 1; Fig. 1b and 1c). Six species classified as rare in 1995 were uncommon (Prionurus laticlavius, Apogon dovii, Cirrhitus rivulatus, Gerres cinereus and Rypticus bicolor) or common (Hoplopagrus guntheri) in 2006. Six uncommon species (Fistularia commersonii, Eucinostomus currani, Nicholsina denticulate, Lutjanus guttatus, Abudefduf concolor and Arothron meleagris) were classified as common, and Diodon sp. was uncommon in 1995 but dominant in 2006. Six species/species groups were less frequent and abundant in 2006, including Gnathanodon specious (uncommon to rare); Pseudobalistes naufragium, Holocanthus passer and Pomacanthus zonipectus (common to uncommon); Haemulon scudderii/maculicauda, steindachneri/flaviguttatum and Scarus rubroviolaceus/perrico/ ghobban/compressus (dominant to common).
We also directly compared the relative abundance of the 48 species/species groups that were recorded in both years, independent of their frequency of occurrence. Of these, thirty-seven (77%) were more abundant in 2006 and only 11 (23%) were more abundant in 1995 (Table 1). The abundance of seven species/species groups varied significantly by year. Of these, six (Diodon sp., Epinephelus labriformis, Hoplopagrus guntheri, Kyphosus sp., Nicholsina denticulate and Thalassoma lucasanum) were more abundant in 2006 than 1995 (Table 3). Only Halichoeres notospilus/ nicholsi were more abundant in 1995 than 2006. At the family level, serranids (Wilcoxon paired sample test, df = 5, z = 2.201, p = 0.028) were significantly more abundant in 2006. We found no significant difference in the abundance of fishes among years when pooled by trophic groups or mobility classes.
Finally, the presence/absence of less abundant species also affected community composition over time. Fourteen of 62 species/species groups were recorded in one year but not the other (Table 1). Eleven of these were recorded in 2006 but not 1995, including Balistes polylepis, Calamus brachysomus, Canthigaster punctatissima, Dasyatis longa, Gymnothorax undulatus, Lutjanus inermis, Lutjanus novemfasciatus, Ostracion meleagris, Spheroides lobatus, Trachinotus rhodopus and Zanclus cornutus. All of these were rare except for Lutjanus inermis, which was classified as uncommon and occurred in moderate numbers in <10% of surveys in 2006, and Canthigaster punctatissima, which was significantly more abundant (Table 3), occurred in 80% of surveys, and was classified as dominant in 2006. Two uncommon species (Elops affinis and Peprilus medius) recorded in 1995 were absent in 2006.
Community composition: The Playa Blanca Marine Reserve supports a diverse fish community similar in species composition to other reported sites in the Panamic Province of the Tropical Eastern Pacific (Table 4). Dominici-Arosemana & Wolff (2006) documented 126 species from 44 families in Bahia Honda, Gulf of Chiriqui, Panama (7[degrees]50' N-81[degrees]35 W), the most diverse Tropical Eastern Pacific reef fish community reported to date. However, Dominici-Arosemena & Wolff (2006) sampled 48 transects over a larger geographic area and used SCUBA to sample to depths up to 15m, while we used snorkeling to sample six transects at a localized site at depths < 3.5m. As a consequence of this variation in sampling effort and methods, total diversity at Playa Blanca was predictably lower; however, species composition between the two sites was relatively similar. Nearly 90% of the families and species recorded in our study were also reported by Dominici-Arosemena & Wolff (2006) (Table 4). The four families recorded at Playa Blanca (Belonidae, Elopidae, Sparidae and Stromateidae) absent from Bahia Honda were each represented by a single rare species.
Seventeen families reported by DominiciArosemena & Wolff (2006) from Bahia Honda were absent from the Playa Blanca Marine Reserve. Twelve of the 17 families were represented by a single species and none by more than three species. One of the most significant differences in community composition between the two sites was the underrepresentation of small-bodied, cryptic benthic fishes in our study. We believe this is a result of our sampling methods; we used relatively wide transects and did not use SCUBA equipment. Future studies employing SCUBA to more intensively sample benthic fishes along narrower transects would likely detect chaenopsids, gobiids, labrisomids, opistognathids, ophichthids and/or tripterygiids that would increase estimates of species richness in the Playa Blanca Marine Reserve.
Additionally, species richness of some families common to both sites was higher in Bahia Honda. For example, the species richness of carnivorous and piscivorous serranids (Bahia Honda: 10 species, Playa Blanca: 3), carangids (BH: 11, PB: 4) and lutjanids (BH: 8, PB: 5) were greater in Bahia Honda compared to Playa Blanca. Many of the species absent from Playa Blanca were reported from just one or a few of the 12 sites in Dominici-Areosemena & Wolff's (2006) study, and many of those sites were at depths >3.5m. Labridae (BH: 10, PB: 4) and Acanthuridae (BH: 5, PB: 1) species richness was also lower in Playa Blanca.
Interestingly, while approximately 90% of families and species from Playa Blanca were also reported from Bahia Honda, Panama, the species composition of Playa Blanca was less similar to that of Culebra Bay, Gulf of Papagayo, Costa Rica (10[degrees]45' N-85[degrees]43 W), even though this site is closer geographically and had similar measures of total diversity (Table 4). Sixty-seven percent of families and 71% of species from Playa Blanca were reported from Culebra Bay (Dominici-Arosemana et al. 2005). Future studies using standardized survey methods at multiple sites may elucidate patterns of variation in species composition along this latitudinal gradient (Ferriera et al. 2004).
In comparison to sites in the Cortez Province of the Tropical Eastern Pacific, measures of species richness were comparable; however, community composition, especially at the species level, was less similar compared to other Panamic Province sites (Table 4). For example, while nearly 70% of the families recorded in our study were also recorded at sites in the Gulf of California, only about 57% of species were (Alvarez-Filip et al. 2006, Aburto-Oropeza & Balart 2001). Further, only 61% of species classified as "dominant" or "frequent and abundant" by Aburto-Oropeza & Balart (2001) at Los Islotes, Gulf of California were present in the Playa Blanca Marine Reserve.
While there was significant variation in species composition across Tropical Eastern Pacific sites, a common feature of reef fish communities in the region appears to be the dominance of pomacentrids, labrids, and haemulids. These families collectively accounted for 73.9% of total fish abundance in the Playa Blanca Marine Reserve and were among the most dominant families in Bahia Honda (Dominici-Arosemena & Wolff 2006), Culebra Bay (Dominici-Arosemena et al. 2005) and the Cabo Pulmo Reef (Alvarez-Filip et al. 2006). Pomacentrids, particularly Stegastes acapulcoensis and S. flavilatus, were ubiquitously observed throughout Playa Blanca Marine Reserve. They were especially prevalent where broken rock covered by dense brown algae (Padina and Sargassum) bordered sandy substrate. This combination of physical and biological substrate characteristics provided ideal habitat for territorial herbivores like Stegastes. Other abundant pomacentrids included Abudefduf troschelii and Chromis atrilobata, which appeared to partition food resources based on depth. Abudefduf troschelii, an omnivore, fed on plankton and invertebrates primarily in the shallowest parts of each transect, while C. atrilobata, a planktivore, occurred in greatest abundance in the upper half of the water column in the deeper, offshore sections of the transects.
The dominance of the labrids (16.6% total fish abundance) was primarily a function of Thalassoma lucasanum, a planktivorous feeder found with high frequency at consistently high abundance in all non-sand transects. The haemulids accounted for 14.8% of total fish abundance in our study. Sixty-two percent of total haemulid abundance was observed in a single transect where very large schools of juveniles congregated in large rock caves and overhangs. Given their high relative abundance, the haemulid's frequency of occurrence was relatively low (< 40% in 2006), indicating that these species likely move in and out of the Playa Blanca Marine Reserve frequently.
Change in community composition and marine reserve management: The Playa Blanca Marine Reserve appears to be fulfilling its conservation role. Most measures of fish abundance, species richness, and diversity were greater in 2006 (after 11 years of protection) compared to 1995 (1 year after reserve designation). Seventy-seven percent of species species/ species groups reported in both years were more abundant in 2006, and seven of eight species/species groups with significant differences in mean abundance among years were more abundant in 2006. Thirteen of 19 species/ species groups whose classification changed among years were more frequent and abundant in 2006. While we lack data from unprotected control sites over the same time period, we believe that the limitations on fish harvest and habitat degradation conferred by the site's protected status have promoted an increase in fish abundance and diversity. Extraction of ornamental fishes such as T. lucasanum for the aquarium trade was common at Playa Blanca in the early-to mid-1990's (C. Vaughan 2010, pers. observ.). Creation of the marine reserve in 1994 and subsequent enforcement of regulations effectively halted this activity and probably contributed to the increase in T. lucasanum and other species observed in 2006.
We had hypothesized that species/species groups at higher trophic levels and/or species classified as relatively sedentary might show a greater response to protection; however, we found no significant difference in the abundance of fishes among years when pooled by trophic groups or mobility classes. Instead, consistent with the results of Halpern (2003), the increase in fish abundance was distributed proportionally among trophic groups.
The Gulf of Nicoya is an important nursery area for many fishes (Campos et al. 1984), and our observations suggest that within the Gulf, small protected areas such as the Playa Blanca Marine Reserve can contribute to fish conservation in the region. For example, we observed a substantial increase in the abundance of juvenile lutjanids and serranids in the reserve in 2006 compared to 1995, and lutjanid species richness increased from three species in 1995 to five in 2006. Both lutjanids and serranids are relatively sedentary carnivores, and their increased abundance could be a result of reduced harvest and/or increased food availabil ity in the reserve after protection. In 2006, large adult L. novemfasciatus, a species absent in 1995, were observed among the rock outcrops and overhangs of the deeper transects, and small schools of L. inermis were frequently observed along vertical rock walls. Future studies and continued monitoring are needed to assess the effects of protection in the long-term; Micheli et al. (2004) found that the magnitude of positive response of piscivorous fishes to protection increased substantially after > 10yrs of protection. The potential role of the Playa Blanca Marine Reserve as a breeding and maturation zone for commercially exploited fishes and as a source of emigrants to adjacent fishing grounds also warrants further investigation.
Marine reserves may also promote habitat diversity by reducing human disturbance and habitat alteration. The transect with the greatest dissimilarity in fish composition among years (see Table 2, transect 3) was one characterized by the highest percentage of sandy substrate. Within this transect, seagrass coverage increased from < 5% in 1995 to approximately 20% in 2006. Fish response to the change in habitat characteristics was apparent; average fish abundance increased five-fold and average species richness doubled in 2006 compared to 1995. In addition, species composition between years was quite different. While planktivores and mobile invertebrate feeders like Mugil sp. and Eucinostomus currani were most abundant in 1995, in 2006 the transect was dominated by schools of juvenile carnivores (especially Haemulon maculicauda, Haemulon steindachneri, and Lutjanus guttatus) actively searching for prey within the seagrass.
In conclusion, we have documented significant increases in fish abundance, species richness, and diversity and significant change in community composition in the Playa Blanca Marine Reserve following eleven years of protection. Future studies incorporating unprotected controls and more frequent monitoring are needed to further elucidate the contribution of small marine reserves to regional conservation goals.
We are grateful to many individuals who provided expertise, information, and logistical support over the course of this study. Eugenio Gordienko and Miguel Fernandez allowed access to Punta Leona facilities and the Playa Blanca Marine Reserve. Memo Hernandez and Andrea Osorio provided logistical support on site. Fieldwork was supported by the Associated Colleges of the Midwest (Spring Semester Research Program in Costa Rica) and Sigma Xi. Institutional support was provided by the University of Northern Iowa for M. Myers and the Universidad Nacional for C. Vaughan. We thank Sarah Faust for her assistance with data curation and literature review and three anonymous reviewers for their insightful comments on this manuscript.
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Mark C. Myers (1), Jonathan Wagner (2) & Christopher Vaughan (3,4,5)
Department of Biology, University of Northern Iowa, 144 McCollum Science Hall, Cedar Falls, IA 50614 USA; email@example.com
College of Food, Agriculture, and Natural Sciences, University of Minnesota, St. Paul, MN 55108 USA; firstname.lastname@example.org
Department of Forest and Wildlife Ecology, University of Wisconsin-Madison, Madison, WI 53706 USA; email@example.com
International Institute for Wildlife Conservation and Management, Universidad Nacional, Costa Rica
Associated Colleges of the Midwest, San Pedro, Costa Rica; firstname.lastname@example.org
TABLE 1 Fish species/species groups recorded during surveys of the Punta Leona Marine Reserve during 1995 and 2006 ID (1) Family Species/ Tg (2) Mob (3) species group 1 Acanthuridae Prionurus h m laticlavius 2 Apogonidae Apogon dovii pl s 3 Balistidae Balistes o m polylepis 4 Pseudobalistes o m naufragium 5 Sufflamen verres i m 6 Belonidae Tylosurus p m crocodilus 7 Blennidae Ophoblennius h s steindachneri 8 Carangidae Caranx p m sexfasciatus or caballus 9 Gnathanodon p m specious 10 Trachinotus p m rhodopus 11 Chaetodontidae Chaetodon i s humeralis 12 Johnrandalia o s nigrirostris 13 Cirrhitidae Cirrhitus c s rivulatus 14 Dasyatidae Dasyatis longa i m 15 Diodontidae Diodon sp. i s 16 Elopidae Elops affinis p m 17 Fistulariidae Fistularia c m commersonii 18 Gerreidae Eucinostomus i m currani 19 Gerres cinereus i m 20 Haemulidae Anistotremus c s caesius 21 Anistotremus c s taeniatus 22 Haemulon c s scudderii, maculicauda, steindachneri or flaviguttatum 23 Haemulidae Haemulon c s sexfasciatum 24 Holocentridae Myripristis pl s leiognathus 25 Sargocentron pl s suborbitalis 26 Kyphosidae Kyphosus sp. o m 27 Labridae Bodianus i s diplotaenia 28 Halichoeres i s notospilus or nicholsi 29 Nicholsina h m denticulata 30 Scarus h m rubroviolaceus, perrico, ghobban, or compressus 31 Thalassoma pl m lucasanum 32 Lutjanidae Hoplopagrus c s guntheri 33 Lutjanus c s argentiventris 34 Lutjanus c s 35 Lutjanus inermis c s 36 Lutjanus c s novemfasciatus 37 Mugilidae Mugil sp. pl m 38 Mullidae Mulloidicthyes sp. c m 39 Muraenidae Gymnomuraena c s zebra 40 Gymnothorax c s castaneus 41 Gymnothorax c s undulatus 42 Muraena c s lentiginosa 43 Ostraciidae Ostracion i m meleagris 44 Pomacanthidae Holocanthus o s passer 45 Pomacanthus o s zonipectus 46 Pomacentridae Abudefduf o m concolor 47 Abudefduf o m troschelii 48 Chromis pl m atrilobata 49 Microspathodon h s bairdi or dorsalis 50 Stagastes h s flavilatus or acapulcoensis 51 Scorpaenidae Scorpaena c m plumieri mystes 52 Serranidae Cephalopholis c s panamensis 53 Epinephelus c s labriformis 54 Rypticus bicolor c s 55 Sparidae Calamus brachysomus c m 56 Stromateidae Peprilus medius c m 57 Tetradontidae Arothron hispidus o s 58 Arothron meleagris o s 59 Canthigaster o s punctatissima 60 Spheroides lobatus o s 61 Sphoeroides o s annulatus 62 Zanclidae Zanclus cornutus i s Total observed ID (1) 1995 2006 N Ab (4) Fr (5) Cl (6) N Ab (4) 1 10 0.08 3 r 62 0.33 2 11 0.09 3 r 32 0.17 3 nr 6 0.03 4 61 0.47 45 c 16 0.09 5 208 1.61 85 d 305 1.65 6 9 0.07 7 r 15 0.08 7 19 0.15 13 u 28 0.15 8 625 4.85 23 c 280 1.51 9 23 0.18 12 u 4 0.02 10 nr 6 0.03 11 375 2.91 78 d 587 3.17 12 23 0.18 15 u 29 0.16 13 8 0.06 12 r 25 0.14 14 nr 1 0.01 15 27 0.21 27 u 243 1.31 16 43 0.33 5 u nr 17 54 0.42 22 u 134 0.72 18 67 0.52 17 u 223 1.20 19 4 0.03 7 r 32 0.17 20 255 1.98 60 c 320 1.73 21 156 1.21 33 c 117 0.63 22 1068 8.28 63 d 2059 11.12 23 246 1.91 45 c 426 2.30 24 125 0.97 33 c 181 0.98 25 645 5.00 82 d 811 4.38 26 40 0.31 33 c 854 4.61 27 364 2.82 80 d 503 2.72 28 408 3.16 82 d 217 1.17 29 22 0.17 25 u 369 1.99 30 312 2.42 73 d 317 1.71 31 571 4.43 78 d 2141 11.57 32 8 0.06 12 r 55 0.30 33 56 0.43 50 c 44 0.24 34 49 0.38 17 u 534 2.88 35 nr 50 0.27 36 nr 7 0.04 37 118 0.92 10 u 78 0.42 38 15 0.12 7 u 23 0.12 39 1 0.01 2 r 6 0.03 40 1 0.01 2 r 2 0.01 41 nr 2 0.01 42 12 0.09 20 u 15 0.08 43 nr 1 0.01 44 42 0.33 42 c 35 0.19 45 29 0.22 32 c 25 0.14 46 70 0.54 22 u 130 0.70 47 1749 13.57 95 d 2163 11.68 48 1831 14.20 62 d 1073 5.80 49 350 2.71 55 c 439 2.37 50 2613 20.27 83 d 2935 15.86 51 1 0.01 2 r 1 0.01 52 26 0.20 35 c 45 0.24 53 58 0.45 47 c 131 0.71 54 3 0.02 5 r 22 0.12 55 nr 10 0.05 56 22 0.17 8 u nr 57 45 0.35 42 c 56 0.30 58 14 0.11 18 u 55 0.30 59 nr 220 1.19 60 nr 6 0.03 61 1 0.01 2 r 3 0.02 62 nr 2 0.01 18511 ID (1) 2006 Fr (5) C (16) 1 6 u 2 22 u 3 8 r 4 21 u 5 84 d 6 2 r 7 19 u 8 19 c 9 5 r 10 3 r 11 78 d 12 22 u 13 21 u 14 2 r 15 97 d 16 17 38 c 18 29 c 19 11 u 20 35 c 21 41 c 22 37 c 23 49 c 24 30 c 25 83 d 26 60 c 27 83 d 28 75 d 29 56 c 30 67 c 31 83 d 32 57 c 33 37 c 34 29 c 35 10 u 36 10 r 37 11 u 38 14 u 39 10 r 40 3 r 41 3 r 42 22 u 43 2 r 44 25 u 45 29 u 46 21 c 47 76 d 48 68 d 49 51 c 50 84 d 51 2 r 52 43 c 53 68 c 54 19 u 55 8 r 56 57 35 c 58 41 c 59 79 d 60 8 r 61 3 r 62 3 r (1.) ID = identification number used to label species/species groups in Fig.1. (2.) Tg = trophic group; c = carnivore, h = herbivore, i = invertivore, o = omnivore, p = piscivore, pl = planktivore (3.) Mob = mobility; m = highly mobile, s = relatively sedentary (4.) Ab = relative abundance expressed as % of total (5.) Fr = frequency of occurrence; % of surveys in which species/species group was recorded (6.) Cl = classification; d = dominant, c = common, u = uncommon, r = rare. TABLE 2 Fish species/species groups most responsible for dissimilarity in community composition among years in each transect Tr (1) Dis (2) Species/species group [Ab.sub.1995] (3) 1 42.75 Abudefduf troschelii 77.2 Thalassoma lucasanum 9.2 Stegastes flavilatus 57.6 or acapulcoensis Microspathodon bairdi 4.7 or dorsalis Chromis atrilobata 9.7 2 48.34 Chromis atrilobata 98.3 Haemulon scudderii, 60.9 maculicauda, steindachneri, or flaviguttatum Kyphosus sp. 1 Abudefduf troschelii 39.8 Thalassoma lucasanum 11.1 3 86.87 Haemulon scudderii, 0 maculicauda, steindachneri, or flaviguttatum Lutjanus guttatus 4.3 Eucinostomus currani 5.8 Mugil sp. 7 Abudefduf troschelii 4.3 4 57.1 Stegastes flavilatus 31.4 or acapulcoensis Thalassoma lucasanum 9.4 Chromis atrilobata 23.7 Abudefduf troschelii 25.5 Sargocentron 4.4 suborbitalis 5 56.1 Thalassoma lucasanum 10.3 Stegastes flavilatus 30.3 or acapulcoensis Nicholsina denticulata 0.5 Abudefduf troschelii 6.7 Chromis atrilobata 3.2 6 43.72 Haemulon scudderii, 39.3 maculicauda, steindachneri, or flaviguttatum Chromis atrilobata 48.2 Stegastes flavilatus 73.9 or acapulcoensis Thalassoma lucasanum 17.1 Abudefduf troschelii 21.4 Tr (1) [Ab.sub.2006] (3) Co% (4) Cu% (5) 1 78.6 12.76 29.86 30.6 4.849 41.2 69.7 3.845 50.2 16 2.473 55.98 4.73 1.997 60.65 2 26.1 7.356 15.22 132 6.284 28.22 69.1 6.278 41.2 74.9 4.04 49.56 52.4 3.936 57.7 3 66.3 35.57 40.95 39.3 20.27 64.29 18 10.43 76.29 0 4.102 81.01 0 2.954 84.41 4 64.5 8.397 14.55 33.8 6.004 24.96 10.1 4.675 33.06 20 4.331 40.56 21.1 4.091 47.65 5 49.5 13.05 23.26 52.1 7.464 36.57 15.2 4.934 45.36 11.9 3.824 52.18 9.64 3.235 57.95 6 0 6.399 14.64 54.3 6.015 28.39 47.8 4.116 37.81 36.3 3.592 46.02 19.8 3.104 53.12 (1.) Tr = transect (2.) Dis = average dissimilarity (3.) Ab = mean abundance (4.) Co% = % of Dis accounted for by species/species group (5.) Cu% = cumulative % contribution of listed species/species groups to Dis. TABLE 3 Species/species groups with significant differences in abundance among years Family Species/Species [Ab.sub.1995] (1) [Ab.sub.2006] group Diodontidae Diodon sp. 1.017 3.786 Kyphosidae Kyphosus sp. 0.750 3.572 Labridae Halichoeres 6.800 3.489 notospilus or nicholsi Nicholsina 0.100 1.950 denticulata Thalassoma 9.517 33.489 lucasanum Lutjanidae Hoplopagrus 0.133 0.852 guntheri Serranidae Epinephelus 0.967 2.069 labriformis Tetradontidae Canthigaster 0.000 4.014 punctatissima Family Test p statistic (2) Diodontidae t=-4.778 0.028 Kyphosidae z=2.023 0.043 Labridae t=4.105 0.009 z=2.023 0.043 t=-4.141 0.009 Lutjanidae t=-5.711 0.002 Serranidae t=-2.549 0.050 Tetradontidae t=-3.907 0.011 (1.) Ab=abundance. If t-statistic is presented, means are reported; if z-statistic, medians. (2.) Where the differences between paired abundance values were normally distributed, we employed paired-t tests and report t-statistics. Where differences were non-normal, nonparametric Wilcoxon paired-sample tests were applied and z-statistics are reported. TABLE 4 Comparison of number offish species and families reported from various Tropical Eastern Pacific sites Authors Site Species Families Current study Playa Blanca Marine 72 30 Reserve, Gulf of Nicoya, Costa Rica Dominici-Arosemena Bahia Honda, Gulf 126 44 & Wolff (2006) of Chiriqui, Panama Dominici-Arosemena Culebra Bay, Gulf 75 28 et al. (2005) of Papagayo, Costa Rica Alvarez-Filip et Cabo Pulmo Reef, 62 23 al. (2006) Gulf of California, Mexico Arbuto-Oropeza & Los Islotes, Gulf 74 28 Balart (2001) of California, Mexico Authors Shared Shared species (1) families (2) Current study Dominici-Arosemena 64 (89%) 27 (90%) & Wolff (2006) Dominici-Arosemena 51 (71%) 20 (67%) et al. (2005) Alvarez-Filip et 41 (57%) 20 (67%) al. (2006) Arbuto-Oropeza & 41 (57%) 21 (70%) Balart (2001) (1.) Number and percentage of species recorded in the Playa Blanca Marine Reserve also recorded in the study indicated. (2.) Number and percentage of families recorded in the Playa Blanca Marine Reserve also recorded in the study indicated.
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|Author:||Myers, Mark C.; Wagner, Jonathan; Vaughan, Christopher|
|Publication:||Revista de Biologia Tropical|
|Date:||Mar 1, 2011|
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