Proceedings of the AcAdemy of nAturAl sciences of PhilAdelPhiA 162: 61-87 mArch 2013

Detection of shifts in coral reef fish assemblage structure over 50 years at reefs of New Providence Island, The Bahamas highlight the value of the Academy of Natural Sciences’ collections in a changing world

KAtriinA l. ilves

Department of Ichthyology, The Academy of Natural Sciences, 1900 Benjamin Franklin Parkway, Philadelphia, PA 19103-1195, USA.Phone/Fax: (1) 215 405 5080; E-mail: ilves@ansp.org

AndreA m. QuAttrini

Biology Department, Temple University, 1900 N 12th Street, Philadelphia, PA 19122, USA.

mArK W. WestneAt

Department of Zoology, Field Museum of Natural History, Chicago, IL 60605, USA.

ron i. eytAn

Department of Ecology and Evolutionary Biology, Yale University, 21 Sachem Street, #358D, New Haven, CT 06520, USA.

gordon W. chAPlin

Department of Ichthyology, The Academy of Natural Sciences, 1900 Benjamin Franklin Parkway, Philadelphia, PA 19103-1195, USA.

heidi hertler

Inter American University of Puerto Rico, Center for Environmental Education, Conservation and Research, PO Box 5100, San German PR 00683, USA.

John g. lundberg

Department of Ichthyology, The Academy of Natural Sciences, 1900 Benjamin Franklin Parkway, Philadelphia, PA 19103-1195, USA.

ABSTRACT.—Natural history collections contain historical records that present snapshots of organisms and communities, and are now more important than ever in the face of rapidly changing environments. The world’s largest collection of fishes from The Bahamas is housed at The Academy of Natural Sciences, Philadelphia. These vast Bahamas holdings primarily resulted from the collecting efforts of James Böhlke and Charles Chaplin in the 1950s-1970s. One of the many ways these historical records have been used since their collection is as baseline data for coral reef sites that were revisited decades later to answer questions about changes in coral reef fish community structure through time. Analyses of data collected from resurveys conducted in 2006 and 2010 provided evidence for a relative increase in the abundance of individuals belonging to different trophic groups and families; particularly, a relative increase in herbivores (mainly parrotfishes) and a relative decrease in planktivores. Furthermore, an interesting inverse trend was detected between two nocturnal groups of fishes, where the larger-bodied squirrelfishes increased while the smaller-bodied cardinalfishes decreased in relative abundance. These significant differences across trophic groups and families are consistent with the effects of a degraded reef habitat characterized by increased algal cover and fewer shelter spaces. Overall, these resurvey results suggest that neither the coral nor reef-fish communities reflect ‘baseline’ conditions. Continued monitoring of these assemblages using comparative data from multiple replicate samples across time periods will be needed to detect their further change or stabilization.

ISSN 0097-3157

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INtroduct Io N

Natural history collections represent a catalogue of biodiversity and are an invaluable resource for the study of evolution and ecology. In the bicentennial year of t he Academy of Natural Sciences, Philadelphia (ANSP), the current study documents one aspect of the importance of one of this institution’s collections – fishes of The Bahamas in ANSP Ichthyology.

t he ANSP Ichthyology collection houses the world’s largest collection of fishes from The Bahamas (Table 1). Most of the >60,000 specimens are reef fishes that were collected between 1955 and 1973 by Ichthyology c urator James Böhlke and ANSP associate Charles Chaplin, who were pioneers in the combined use of rotenone, a naturally-derived ichthyocide, and SCUBA in the earliest days of recreational diving (Böhlke 1997). Many specimens represented species that were new to science, as discovered by Böhlke who dedicated much of his career to understanding the taxonomy and systematics of c aribbean, and particularly Bahamian, reef fishes. These efforts resulted in Böhlke’s descriptions of 57 species with their type localities in The Bahamas, and a further 26 species for which the type locality is in the greater c aribbean (Eschmeyer 2012). In total, Böhlke described 143 new species from around the world, 115 from marine and 28 from freshwater environments (Eschmeyer 2012). Böhlke’s collections and work on Caribbean fishes were particularly significant with respect to descriptions of cryptic reef fishessuch as blennies and gobies, and important for studies of eel taxonomy led by ANSP ichthyologist Eugenie Böhlke, who was also an active participant in much of the Bahamas work (Smith 2002). Almost 15 years of study culminated in the 1968 publication of Böhlke and Chaplin’s landmark book “Fishes of the Bahamas and Adjacent Tropical Waters” that documented 507 species with identificationkeys, color plates, and detailed illustrations (Böhlke and Chaplin 1968). This book remains one of the best scientificguides to a regional fish fauna ever published

The original goals of Böhlke and Chaplin’s collecting and subsequent publications were to provide a scientificinventory of the Bahamian fish fauna and to further understand the taxonomy, morphology, distribution, and evolutionary relationships of these fishes, purposes for which the specimens remain in use. A perhaps unintended result of their work, however, was the provision of baseline snapshots of the reef fish assemblages at Bahamian islands. Most of these ‘historical’ collections were made prior to the escalation of major anthropogenic influences known to degrade reef environments, such as sedimentation, pollution, overfishing, development, and the effects of climate change (e.g., Gardner et al. 2005; Wilson et al.

2006; Hoegh-Guldberg et al. 2007; Weber et al. 2012). t hus, these data can be used as a close approximation to the natural state of regional reef fish assemblage structure.

t he practice of using baseline data and revisiting historically studied areas is not new. For instance, the Grinnell Resurvey Project at the Museum of Vertebrate Zoology at the University of California, Berkeley has undertaken ambitious resurveys in regions of California first studied almost 100 years ago. Insights from those 21st century resurveys include the detection of terrestrial bird and mammal range contractions or shifts over the past century that have been correlated with climate changes (e.g., Moritz et al. 2008; Tingley et al. 2009; Rubidge et al. 2012). There has also been extensive study of the effects of large-scale disturbances, such as extreme weather events, on coral and benthic invertebrate communities (e.g., Dahl and Lamberts 1977; Pearson 1981; Rogers et al. 1983; Baker et al. 2008). Furthermore, numerous resurveys of reef fish assemblages, often conducted in concert with coral surveys, have spanned multiple years and disturbances (e.g., Nagelkerken et al. 2005; Bellwood et al. 2006; Kellison et al. 2012). Despite the immense amount of ecological work focused on coral reefs and their associated fishes, the availability of reef fish data across multiple decades is exceedingly rare. To our knowledge, the only comparison of reef fish assemblage structure spanning a timeframe beyond 30 years is the study by Ilves et al. (2011).

Ilves et al. (2011) resurveyed four coral reef sites at New Providence Island, The Bahamas in 2006 to compare reef fish assemblage structure between historical (1955-1973) and recent (2006) collections, spanning a range of 33-51 years. Key findings included no significant differences in species diversity (H’, Shannon index) between time periods, significant increases in the relative proportion of individuals of herbivorous species, and in families such as parrotfishes and squirrelfishes, and a decreasing trend in the relative proportion of individuals of planktivorous species. Qualitative differences in coral health, including decreased live coral cover and an increase in algal cover were noted. c oral structure, however, was observed to be relatively intact. The finding of subtle changes in reef fish assemblages combined with the maintenance of coral structure led to the conclusion that the reef fish community was responding to a disturbance that occurred on the order of a decade.

For the years spanning 2006 and 2010 there was anecdotal evidence from locals and dive operators (e.g., S. Cove, personal communication, 2010) that reef community health at New Providence Island was improving, indicated by a perceived increase in live coral and decrease in algae. The present study reports on a further resurvey in 2010

detection of shifts in corAl reef fish AssemblAge structure At reefs of neW Providence islAnd 63

at the same reef sites around New Providence Island, t he Bahamas, sampled historically and/or in 2006 with two specific aims: (1) collect additional reef fish assemblage data at coral reef sites, and (2) assess whether (a) anecdotal evidence about the improved condition of coral reefs in this area is supported by a trend in reef fish assemblage structure towards a return to ‘historical’ conditions, or (b) patterns detected from the 2006 resurvey (Ilves et al. 2011) are reinforced and/or expanded by the addition of a new time period data point. Furthermore, this study aims to highlight the importance of the Bahamas holdings of The Academy of Natural Science’s Ichthyology collection in a comparison to seven other major US museums. Analyses of data from five sites resurveyed in 2010 yielded the same statistically significant differences reported by Ilves et al. (2011), and provided statistical support for qualitative trends apparent from the 2006 data. These results suggest that the reef fish assemblages have not recovered, and may have either reached a new assemblage state or are in continued flux.

METHo DS

Comparison of Bahamas holdings across major Ichthyology collections.—t he selection of institutions to which to compare ANSP Ichthyology Bahamas holdings was made based on the authors’ knowledge, in consulta-tion with ANSP Ichthyology collection manager M. Sabaj Pérez, of institutions likely to have significant collections from the region: Florida Museum of Natural History (UF), American Museum of Natural History (AMNH), Harvard Museum of c omparative Zoology (Mc Z), Smithsonian Institution National Museum of Natural History (USNM), Yale Peabody Museum (YPM), Field Museum of Natural History (FMNH), and California Academy of Sciences (c AS). c ollection data were either downloaded from the institution’s website or requested from the respective col-lection manager. Specimens collected from freshwater habitats were excluded, when such information was avail-able. As there are few freshwater habitats in The Bahamas, these exclusions were few in number and inadvertent in-

Table 1. Comparison of marine/brackish Bahamas holdings among Ichthyology collections at the Academy of Natural Sciences (ANSP), Florida Museum of Natural History (UF), American Museum of Natural History (AMNH), Harvard Museum of Comparative Zoology (MCZ), Smithsonian Institution National Museum of Natural History (USNM), Yale Peabody Museum (YPM), Field Museum of Natural History (FMNH), and California Academy of Sciences (CAS). * indicates known collections in 2010 and 2012 that have not yet been accessioned into the FMNH collection.

Museum Ichthyology CollectionCollection statistics ANSP UF AMNH MCZ USNM YPM FMNH CASLots 10,784 10,386 8,453 2,973 1,715 1,052 291 123Specimens 61,246 48,594 54,065 15,219 7,555 5,464 626 233Species (valid) 626 974 513 425 491 328 135 93Primary types 86 10 14 1 26 82 1 1

Specimens/time periodUnknown 674 97 59 283 288 12 36 41<1900 158 0 2 142 346 0 1 21900-1919 1 0 27 214 563 12 3 01920-1939 80 0 503 1,424 224 5,440 213 111940-1949 169 2,123 455 69 117 0 81 01950-1959 28,940 8,558 889 237 2,613 0 191 671960-1969 19,578 23,206 46,057 80 845 0 33 631970-1979 2,178 13,979 3,659 1 98 0 44 471980-1989 91 253 428 12,059 323 0 0 21990-1999 13 369 1 710 5 0 24 02000-2009 5,394 18 1,262 0 2,104 0 0 02010-09/2012 3,970 0 723 0 29 0 0* 0

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clusion of freshwater collections is unlikely to have af-fected the main findings. Species counts were conducted on refined databases where effort was made to account for misspellings and synonymy according to the c atalog of Fishes (Eschmeyer 2012). Counts of type specimens were limited to name-bearing specimens (primary types): holo-types, neotypes, syntypes, and lectotypes.

Sampling sites and methods.—Following Ilves et al. (2011), the sample sites were classified into three depth categories: shallow (< 6 m), mid (6 – 12 m), and deep (12 – 18 m), and designated by a two-letter site code followed by 1, 2, or 3 to represent the respective depth category. Three sites resurveyed in November 2010 corresponded to historical samples (NLI, d P1, Gc 3) and two additional sites (GC1, GC2) were first surveyed in July or November 2006 (Fig. 1). Site details are described in detail by Ilves et al. (2011). The original goal was to resample all four sites surveyed both historically and in 2006 (LC1, NL1, DP1, GC3) and one additional deep site from 2006; however, due to wind conditions and poor visibility it was not possible to survey Lc 1 or a deep site other than Gc 3.

t he sampling methodology closely followed that of the historical surveys by Böhlke and Chaplin and the 2006 collections described by Ilves et al. (2011): 5% powdered rotenone (TIFA International) was emulsified with Dawn® concentrated phosphate-free dish soap and seawater and, using SCUBA, was distributed at a specified patch reef. Following rotenone application, five or six divers col-lected all affected fishes using handnets for 60-90 min.

Fishes were sorted and given preliminary IDs daily in the field. As many specimens as possible were sub-sampled for tissues intended for molecular study prior to fixation of whole specimens in a 10% solution of seawater-buffered formalin; tissue samples were stored in 100% Eto H. All specimens were catalogued into the ANSP Ichthyology collection database and most whole specimens are physi-cally at ANSP, although a small subset was gifted to the Field Museum of Natural History. Tissue samples are cur-rently stored at -80 ºC in the ANSP Fish Tissue Collection.

Although attempts were made to replicate the Böhlke and Chaplin methodology, there were differences. Historically, Böhlke and Chaplin used both powdered and liquid rotenone emulsified with detergent, as confirmedby G. Chaplin who was a participant in the Historical and r ecent collections. t he switch from powdered to liquid rotenone occurred in the late 1950s, which in the currently analyzed dataset corresponds to collections from 1959 to 1973. Although there is anecdotal evidence that rotenone became less effective over time due to changes in regulations and formulations (J. t yler, personal communication, 2012), from observations in the fieldwe have no reason to believe such changes resulted in differential effects on different species. t he current analysis did not rely on quantitative comparisons, and as such we are confident that comparisons of relative abundances of individuals of species between time periods are not biased by different rotenone formulations.

A further potential concern about the comparability of sampling methodologies is the possibility that

Fig. 1. Map of sampling sites off the coast of New Providence Island, The Bahamas. Historical sampling sites are open crosshairs. Site abbreviations are as follows: Lyford Cay/Clifton Pt. (LC), Delaporte Point (DP), North/Long Cay (NL), and Green Cay (GC). Reprinted from Ilves et al. (2011) with permission from the Rosenstiel School of Marine and Atmospheric Science, University of Miami.

detection of shifts in corAl reef fish AssemblAge structure At reefs of neW Providence islAnd 65

collection practices differed between time periods. d uring historical expeditions, including some in which Böhlke was a participant, it was common to leave excess numbers of some species, particularly large and spinous taxa such as squirrelfishes, on the reef (J. Tyler, personal communication, 2012). Such sampling would certainly bias comparisons, even of relative abundances between time periods; however, G. Chaplin clearly remembers that for the Böhlke and Chaplin Bahamas expeditions the practice was to collect every specimen, or note when excess specimens were discarded. The sample data and fieldnotes also support these recollections; for example, many Historical samples included dozens of squirrelfishes. We are thus confident that the current sampling methodology is sufficiently congruent with the historical practice that comparisons of species’ relative abundances are valid.

Data analysis.—Each species was classified into one of five trophic groups following Paddack et al. (2009) and Ilves et al. (2011), with the exception that the separate groups ‘carnivore’ and ‘piscivore’ were combined as ‘carnivore’ in the current study. Ilves et al. (2011) excluded potentially biased or compromised sampling events from the historical and 2006 time periods based on information in field notes, and these same events were excluded from the present analyses. t he time periods used for comparison were classified as ‘Historical’ (1955-1973), ‘2006’, ‘2010’, and ‘Recent’ (combined 2006 and 2010 data). Species that are typically considered “non-reef” [Appendix 1, following Choat and Bellwood (1991) and Ilves et al. (2011)] were excluded from all analyses. Specimens identified only to family or genus due to immature life stage and/or condition (e.g., damage, poor preservation), were excluded from multivariate assemblage structure analyses, but were included in univariate analyses of family and trophic group relative abundance. Families that represented less than 1% relative abundance across sites were grouped as ‘o ther’. Due to uncertain identificationsof brotula species belonging to Ogilbia and Ogilbichthys, all such specimens were combined into the taxon ‘Ogilbia/Ogilbichthys sp.’. Similarly, specimens identified as the gobies Coryphopterus glaucofraenum and C. tortugae, and soapfishes of the newly described Rypticus carpenteri (Baldwin and Weigt 2012) and R. subbifrenatus were combined as C. glaucofraenum/tortugae and R. carpenteri/subbifrenatus, respectively. All of those composite taxa were included in all analyses. t he glassy sweeper, Pempheris schomburgkii, was excluded from the analyzed datasets due to an unprecedented catch from the d P1 site in 2010 (>800 specimens, many of which were left on the reef) that would have biased all analyses. Finally, because no mid depth sites were sampled historically, these sites

were excluded from most analyses. t he raw collection data across all years and sites included in this study appear in Appendix 1 and are available from K.L. Ilves in spreadsheet format upon request.

EstimateS (Colwell 2009) was used to assess the adequacy of the community sample and to compare species accumulation curves across time periods. Standard diversity metrics (number of specimens, number of species, H’) and relative abundances of trophic groups and families were calculated in PRIMER 6 (Clarke and Gorley 2006). To assess differences in means between samples for H’ and relative abundance of trophic group and family, t-tests were calculated using the ‘stats’ package in R (R Development Core Team 2008). This test of differences between means is robust to violations of the normality and equal variances assumptions, and is also suitable for testing for differences when only a single sample is available for one of the populations (Simpson et al. 2003), which is the case for the 2010 deep category. Because mid depth sites were only sampled during the Recent time period, comparisons of H’ were only conducted for mid vs. shallow and mid vs. deep sites for this period. Potential differences in assemblage structure between depths and time periods were assessed using the multivariate approach of Clarke (1993) and Clarke and Warwick (2001) implemented in PRIMER 6 (Clarke and Gorley 2006). Abundances were standardized and fourth-root transformed prior to the calculation of Bray-Curtis similarity coefficien s (Bray and Curtis 1957). Multi-dimensional scaling (Md S) and hierarchical cluster analyses were performed to visualize the degree of similarity among sites and depths. Analysis of similarity (ANo SIM; Clarke 1993) was conducted to test for differences between combinations of depth (shallow, mid, deep) and time periods (Historical, 2006, 2010, Recent). Where significant differences were detected, SIMPER (similarity percentage) analysis (Clarke 1993) was used to identify the species most responsible for these differences.

r e Su Lt S

Comparison of Bahamas holdings across Ichthyology collections.—The results of the comparisons of Bahamas holdings across eight major US Ichthyology collections are presented in t able 1. d ue to the inherent vagaries of all such databases and the flux of taxonomic revisions and new species descriptions, the numbers are best regarded as approximations to actual holdings. With 10,784 lots and 61,246 specimens, the ANSP Ichthyology collection has the world’s largest holdings of fishes from The Bahamas, and the highest number of primary types from this region, with 86. In terms of species numbers, ANSP is second, with 626, to UF, with 974 (Table 1). All

66 K.l. ilves et. Al.

collections compared, however, have significant holdings apart from raw numbers. ANSP undoubtedly has the most important collection of Bahamian reef fishes; however, this collection generally lacks specimens from a wide diversity of habitats, such as pelagic and deep-sea environments for which UF and MCZ are particularly strong.

Species accumulation curves and diversity metrics.—When data are combined across time periods, rarefaction curves plotting the number of species accumulated by the number of specimens collected by depth reached an asymptote at approximately 140 species, suggesting that these sites were well-sampled (Fig. 2A). When the data were further separated by time period the curves for the ‘Shallow, Recent’ and ‘Deep, Historical’ are approaching an asymptote of 120 and 140 species, respectively (Fig. 2B,C) The ‘Shallow, Historical and ‘Deep, Recent’

Fig. 2. Species accumulation curves for A) all samples segregated by depth zone ‘shallow’ (open diamonds) and ‘deep’ (closed diamonds), B) shallow depth zone segregated by ‘Historical’ (closed squares) and ‘r ecent’ (open squares) time period, and c ) deep depth zone segregated by ‘Historical’ (closed circles) and ‘Recent’ (open circles) time period. Values in parentheses for each curve’s legend represent the number of species and number of specimens collected, respectively.

Table 2. Number of specimens (N), number of species (S), and Shannon diversity index (H’) for each site, following exclusions. H’ values in bold are mean values across sample sites that were used in t-test comparisons (no significantdifferences were detected). Pempheris schomburgkii (glassy sweeper) was excluded from all analyses due to the 2010 DP1 site (*) where over 800 specimens were encountered. Sample site and depth codes as in Figure 1.Site (mm/dd/yyyy) N S H’

Historical, shallow 1229 102 3.48LC1 (05/08/1956) 161 44 3.35DP1 (04/11/1955) 130 44 3.50DP1 (07/27/1955) 293 55 3.30DP1 (09/05/1955) 366 59 3.60NL1 (08/5/1955) 279 65 3.63

2006, shallow 1794 104 3.33LC1 (11/12/2006) 517 64 3.455DP1 (11/12/2006) 443 65 3.57NL1 (07/14/2006) 462 60 3.04GC1 (11/16/2006) 372 50 3.24

2010, shallow 1173 83 3.28DP1 (11/17/2010)* 764 61 3.25NL (11/17/2010) 248 54 3.40GC1 (11/16/2010) 161 39 3.19

Recent (2006, 2010), shallow 2967 116 3.31

Historical, deep 3364 136 3.16GC3 (05/07/1956) 228 46 3.04GC3 (05/12/1957) 115 33 2.69GC3 (07/21/1957) 788 65 2.94GC3 (05/14/1959) 828 81 3.41GC3 (11/14/1959) 904 92 3.80GC3 (08/27/1969) 501 61 3.06

2006, deep 1558 102 3.47LC3 (11/10/2006) 118 35 3.16DP3 (11/14/2006) 251 52 3.53NL3 (07/13/2006) 185 48 3.43GC3 (11/15/2006) 1004 84 3.45

2010, deep 859 71 3.37GC3 (11/15/2010) 859 71 3.37

Recent (2006, 2010), deep 2417 117 3.45

Recent (2006, 2010), mid 2176 111 3.27LC2 (11/11/2006) 112 30 2.81DP2 (11/13/2006) 754 66 3.20NL2 (11/09/2006) 185 53 3.55GC2 (11/16/2006) 451 62 3.42GC2 (11/16/2010) 674 63 3.39

detection of shifts in corAl reef fish AssemblAge structure At reefs of neW Providence islAnd 67

t able 3. r esults of Analysis of Similarity (ANo SIM) tests between depths and time periods. Significant differences (P ≤ 0.05) highlighted in with *. Historical refers to samples from 1955-1973; Recent refers to combined 2006 and 2010 samples. Number of samples compared for each category in parentheses.

Comparison |R| P

Between depths Shallow (12) vs. Mid (5) 0.519 0.001* Shallow (12) vs. Deep (11) 0.829 0.001* Mid (5) vs. Deep (11) 0.288 0.04*

Between depths, within time periods Historical: Shallow (5) vs. Deep (6) 0.963 0.002* 2006: Shallow (4) vs. Deep (4) 0.802 0.03* 2010: Shallow (3) vs. d eep (N =1) 1 0.25 Recent: Shallow (7) vs. Deep (5) 0.895 0.001*

Shallow depth, between time periods Historical (5) vs. 2006 (4) 0.3 0.02* Historical (5) vs. 2010 (3) 0.333 0.05* 2006 (4) vs. 2010 (3) 0.241 0.83 Historical (5) vs. Recent (7) 0.486 0.001*

Deep depth, between time periods Historical (6) vs. 2006 (4) 0.413 0.01* Historical (6) vs. 2010 (1) 0.067 0.571 2006 (4) vs. 2010 (1) 0.25 0.40 Historical (6) vs. Recent (5) 0.304 0.01*

Mid depth, recent time periods Mid, 2006 (4) vs. Shallow, 2006 (4) 0.375 0.03* Mid, 2006 (4) vs. Deep, 2006 (4) 0.01 0.49 Mid, 2010 (1) vs. Shallow, 2010 (3) 0.556 0.25 Mid, Recent (5) vs. Shallow, Recent (7) 0.639 0.001* Mid, r ecent (5) vs. d eep, r ecent (5) 0.064 0.25

collections have not quite reached an asymptote with respect to the number of specimens collected (Fig. 2B,C).

Diversity metrics, ANOSIM, and SIMPER results.—

d iversity metrics for the sampled sites are listed in t able 2. No significant differences in mean H’ were detected for any comparison (P > 0.5 for all; data not shown). By contrast, significant differences in community composition were detected between shallow and deep sites for all comparisons, except when the 2010 collections, which had a single deep depth sample, were analyzed in isolation (t able 3). e xamination of the Md S and cluster plots (Appendix 2) shows that all sites, including collections from 2010, group with their respective depth categories. No significant differences in community composition were detected between the 2006 and 2010 collections at shallow or deep sites (t able 3). t his similarity between the two most recent time periods provides justification for combining these collections into a ‘r ecent’ time period for further comparisons with the ‘Historical’ data.

At the shallow depth range we found significantdifferences for all comparisons between the Historical and more recent collections, (vs. 2006, vs. 2010, vs. Recent; Table 3). At the deep depth significant differences were found between the Historical and 2006 and the Historical and Recent time periods, but not between the Historical and 2010 collections (Table 3). SIMPER analysis did not clearly discriminate species that were responsible for the detected differences between depths and between time periods within depths: 100-113 and 42-46 species accounted for 90% and 50%, respectively, of the average dissimilarities between groups (data not shown; available from K.L. Ilves).

Trophic group and family relative abundance.—d ue to the significant differences in assemblage structure by depth, the relative abundances by trophic group (Fig. 3) and family (Fig. 4) were segregated into shallow and deep sites for comparisons across time periods (Historical, 2006, 2010) and (Historical, Recent). Significant differences in herbivore relative abundance were detected at shallow sites between the Historical and 2010 time periods (Fig. 3A), at deep sites between the Historical and 2006 time periods (Fig. 3B), and at both shallow and deep depths between the Historical and Recent (2006 and 2010 combined) time periods (Fig. 3C), where the proportion of herbivores increased over time. Further, planktivore relative abundance was significantly lower at deep sites in the Recent compared to the Historical time period (Fig. 3C). Finally, significant differences in the relative proportion of carnivores were detected at shallow sites between 2006 and 2010, and deep sites between Historical

and 2006 and between Historical and Recent (Fig. 3). The associated statistics for trophic group relative abundance are listed in Appendix 3.

With respect to changes in family relative abundance, the proportion of chaenopsid blennies (c haenopsidae) was significantly less in 2010 than in 2006 at shallow and deep sites (Fig. 4A,B) and also less in 2010 than in the Historical time period at deep sites (Fig. 4B). Significant differences

68 K.l. ilves et. Al.

structure over time. r esults from a previous resurvey conducted in 2006 (Ilves et al. 2011) provided evidence of recent disturbance to the coral reef environment around New Providence Island, The Bahamas. In terms of species diversity and differences in the relative abundances of trophic groups and families, the results from the 2010 collections reinforce and expand the findings from Ilves et al. (2011). Thus, despite anecdotal evidence of improved coral reef conditions, there is no support for the hypothesis that the reef fish communities sampled at New Providence Island are in the process of returning to historical conditions.

Sampling and differences between depth zones.—r arefaction curves plotting number of species by number of specimens collected indicated fairly complete coverage of the fish assemblages across depths and time periods, although the Shallow, Historical and Deep, Recent samples may not account for all species. o verall, these results provided confidence that there was an adequate community sample for subsequent analyses of assemblage structure.

in squirrelfish (Holocentridae) relative abundance were detected at shallow and deep sites, with a trend of increased abundance from Historical to Recent (Fig. 4). A trend in increased relative abundance of parrotfishes (Scaridae) was also seen, with a significantly higher proportion of scarids at deep sites in 2006, 2010, and the combined Recent time period than in the Historical samples (Fig. 4B,C). Further significant differences were detected between triplefins(Tripterygiidae) in 2006 and 2010 at the deep depth (Fig. 4B), and between the ‘o ther’ category at shallow depths between Historical and 2006 and Historical and Recent. t he associated statistics for family relative abundance are listed in Appendix 4.

dIS cu SSIo N

The significant Bahamas holdings of The Academy of Natural Sciences’ Ichthyology collection (t able 1), primarily a result of Böhlke and Chaplin’s efforts over a 20-year period starting in the early 1950s, provide a unique opportunity to examine changes in reef fish assemblage

Fig. 3. Relative abundance (%) of trophic group between Historical, 2006, 2010, and Recent collections at shallow and deep sites. Bars represent standard error and * indicates significant result at P ≤ 0.05 (Appendix 3).

detection of shifts in corAl reef fish AssemblAge structure At reefs of neW Providence islAnd 69

and Johnson 1990a), blennies (Blenniidae, Chaenopsidae, Labrisomidae, Tripterygiidae; Greenfield and Johnson 1990b), and squirrelfishes (Holocentridae; Lieske and Myers 1996) matched our collection data, again providing a measure of confidence in the assemblage sample

Comparison of Historical and Recent assemblage changes with new data.—The most striking result of adding an additional data point was the confirmation and expansion of significant differences and trends detected by a comparison of Historical and 2006 data alone. Ilves et al. (2011) only found three significant differences between the mean relative abundance of trophic groups and families between these latter time periods: increased relative abundance of herbivores and of herbivorous parrotfishes(Scaridae) at deep sites in 2006, and an increased relative abundance of squirrelfishes (Holocentridae) at shallow depths in 2006. These same significant differences were found in data from 2010 (Figs. 3, 4). The trend towards an increase in the relative abundance of herbivores at

It is a well-documented phenomenon that coral reef fish species segregate by depth (e.g., Arreola-Robles and Elorduy-Garay 2002; Greenfield and Johnson 1990a,b; McGehee 1994; Srinivasan 2003), a result seen here in the Historical and Recent samples with significant differences in assemblage structure across all depth comparisons apart from the Shallow, 2010 vs. Deep, 2010. Similarly, when assemblage structure was compared within depth zones but between Historical and Recent time periods, significantdifferences were detected in all comparisons apart from Deep, Historical vs. Deep, 2010, although the latter category was represented by only a single site (Gc 3). When data from the 2010 collection were combined with those from 2006, assemblage structure was significantly different between the Historical and Recent time periods, indicating the lack of significant differences are best attributed to the small 2010 sample size. Although SIMPER results did not yield any strongly discriminating species, similar to the results of Ilves et al. (2011), we found that known depth distributions for cardinalfishes (Apogonidae; Greenfield

Fig. 4. Relative abundances (%) of family between Historical, 2006, 2010, and Recent collections at shallow and deep sites. Families with less than a mean of 1% relative abundance across sites were grouped into ‘o ther’. Bars represent standard error and * indicates significant result at P ≤ 0.05 (Appendix 4).

70 K.l. ilves et. Al.

shallow depths in 2006 was followed in 2010. In contrast to the 2006 data alone, significant differences in herbivore relative abundances were detected between Historical and 2010 and Historical and Recent data. Furthermore, the 2010 data indicate an increased relative abundance of squirrelfishes at both shallow and deep depths.

Additional trends noted in Ilves et al.’s previous study included a decrease in planktivores and a decrease in car-dinalfishes (Apogonidae) through time at both shallow and deep sites. The addition of data from 2010 added statistical support to most of these previously qualitative observa-tions. There was a continuing trend of decreased plank-tivore abundance through time (Fig. 3), although the dif-ference was only statistically significant for a comparison of Historical and Recent collections at deep depths. This decrease in planktivores is most likely a result of a continu-ous decrease in apogonids through time. d ue to the large variance associated with the collection of apogonids their decrease was not statistically significant (Fig. 4; Appendix 4). Finally, the significant differences observed in the car-nivore trophic group, and the chaenopsid, trypterygiid, and ‘o ther’ families (Figs. 3, 4) were unexpected. Relative increases in the collection of carnivorous bythitids, labri-somids, and serranids between time periods appear to be contributing to the detection of a significant increase in carnivore relative abundance through time. An explanation for this difference is not readily clear, but may be associ-ated with the trend in decreased planktivore representation (discussed below). Similarly surprising were the inverse differences seen in chaenopsid and triplefin blennies be-tween the 2010 and other collections; however, these may be best explained by sampling artifacts with only a single deep site sampled in 2010. Likewise, the differences at the shallow depths in the ‘o ther’ grouping of families with <1% representation across all sites was primarily driven by bythitids (brotulas), which had a higher relative abun-dance in recent than in historical samples, and gobiesocids (clingfishes), which showed the opposite pattern (data not shown). We hesitate to make any conclusions about actual changes in the relative abundance of these taxa given their rarity of collection at the shallow depth zone.

Ecological context of reef fish assemblage changes from Historical to Recent.—t he coral reefs at New Providence Island appear to be in a disturbed state (Ilves et al. 2011; Jaap et al. 2008). In such disturbed habitats increases in herbivores due to an increase in food availability are well-documented (e.g., Cheal et al., 2008; Sheppard et al. 2002; Wilson et al. 2009). In relatively undisturbed reef habitats the presence of herbivores is generally considered to be beneficial for live coral reefs, as grazing reduces algal cover that may otherwise hinder

settlement and/or growth of coral colonies (e.g., Mumby et al. 2006, 2007); however, in areas where much of the coral is dead, herbivore grazing may actually precipitate a decline in coral integrity, particularly through the feeding behavior of scarids that scrape algae off coral skeletons and contribute to bioerosion of the coral structure (e.g., Carreiro-Silva et al. 2005; Graham et al. 2006; Pratchett et al. 2009). Furthermore, studies of the relationship between corals and herbivorous fishes suggest that herbivore species richness, not simply individual abundance, is important in both controlling diverse algal populations and limiting damage to the coral (Burkepile and Hay 2008, 2010).

t he current study documents a clear trend of an increasing representation of herbivores, represented by scarids (parrotfishes) (Fig. 4). There was no apparent trend for acanthurids (surgeonfishes), the other dominant herbivorous reef fish group on these reefs. These changes may indicate the reef structure, which remains mostly intact, is at risk from both continued anthropogenic effects as well as the resident fishes (i.e., herbivorous scarids). Preliminary results from benthic habitat surveys conducted concurrently with the 2010 fish surveys indicate that live coral cover at the sampled sites has either remained the same or decreased since 2006 (H. Hertler, unpublished data). c ontinued monitoring of the integrity of New Providence Island reefs and the associated fishassemblages will provide an opportunity to assess whether the communities recover from disturbance, maintain a new assemblage structure, or undergo further degradation.

Another result of particular interest is the apparent inverse trend seen in holocentrid and apogonid relative abundance (Fig. 4). Both groups are primarily nocturnal feeders that use caves and crevices for shelter during the day (Chave 1978; Gladfelter and Johnson 1983; Greenfieldand Johnson 1990a). Luckhurst and Luckhurst (1978) showed that the much larger-bodied holocentrids have longer residency times than apogonids within particular coral shelters. t hus, even though apogonids are not habitat specialists (Greenfield and Johnson 1990a), a possible explanation for the observed trends is that the smaller apogonids are out-competed for shelter space in degraded coral habitats. To our knowledge this inverse relationship between holocentrid and apogonid relative abundance has not been documented elsewhere, and it would be of great interest whether this pattern is a general trend in disturbed coral reef environments.

In conclusion, the availability of historical reference points in the Ichthyology collection at the Academy of Natural Sciences provided baseline data from which to study reef fish assemblage structure through time at reefs at New Providence Island, The Bahamas. The results from resurveys in 2006 and 2010 indicated that since the his-

detection of shifts in corAl reef fish AssemblAge structure At reefs of neW Providence islAnd 71

torical collections over 30 years ago, reef fish assemblage structure has changed in a consistent manner at four coral reef sites, with an increase in the relative abundance of her-bivores, scarids, and holocentrids, and a decrease in the relative abundance of planktivores and apogonids. The ul-timate causes of coral degradation in this region are likely a complex combination of anthropogenic effects, includ-ing pollution, such as run-off from adjacent developments in Nassau and Paradise Island, and large-scale changes in ocean temperature and chemistry resulting from climate change. t he availability of replicate samples at multiple time periods are necessary and invaluable for continued monitoring of the trajectory of these reef fish assemblages.

ACKNo WLEDGEMENTS

We thank the government of The Bahamas for their continued support of our ongoing studies of Bahamian reef fish assemblages, particularly M. Braynen, Director of Marine Resources, and R. Albury of the Fisheries Department for granting scientific research permits. We are grateful to Bahama Divers for the boat charter and SCUBA support, and especially to Captain Roscoe for his expert piloting and knowledge of the waters. Many thanks to R. and M. Ilves for their field support efforts underwater and onshore, and to M. Sabaj Pérez for remote field support and assistance with specimen cataloguing. We also appreciate the contribution of Bahamas collection data from Ichthyology collection managers G. Watkins-Colwell (YPM), K. Hartel (MCZ), R. Arrindell (AMNH), and particularly R. Robins (UF). An early version of this manuscript was improved by comments from J. t yler. t he 2010 fieldwork was funded by a Wistar-Morris grant from the Academy of Natural Sciences, and the current study was made possible by the generous support of K. Ilves at ANSP by G. c haplin and S. t eale.

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74 K.l. ilves et. Al.

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LC2-

2006

LC3-

2006

DP1

-20

06D

P2-

2006

DP3

-20

06LC

1-19

56D

P1-

1955

cD

P1-

1955

bD

P1-

1955

aN

L1-

1955

GC

3-19

56G

C3-

1957

aG

C3-

1957

bG

C3-

1959

aG

C3-

1959

bG

C3-

1969

Aul

osto

mid

aeAu

lost

omus

m

acul

atus

c0

10

01

10

20

42

00

01

35

10

22

12

01

01

1

Ble

nniid

aeB

lenn

iidae

sp.

H0

00

00

00

00

00

00

00

00

00

00

00

00

00

1

Ble

nniid

aeEn

tom

acro

dus

nigr

ican

sH

00

00

00

00

00

00

00

10

00

00

00

00

00

00

Ble

nniid

aeH

yple

uroc

hilu

s sp

ring

eri

c0

00

15

00

12

00

00

00

00

00

00

00

00

00

0

Ble

nniid

aeO

phio

blen

nius

m

accl

urei

H3

00

1146

120

014

00

10

012

00

016

51

60

00

00

0

Bot

hida

eBo

thus

luna

tus*

I0

00

00

10

00

00

00

00

00

00

00

00

00

00

0

Bot

hida

eBo

thus

oce

llatu

s*I

01

00

30

00

00

00

00

00

00

00

00

03

02

00

Byt

hitid

aeB

ythi

tidae

sp.

c0

10

00

00

00

00

00

00

00

00

00

00

00

00

0

Byt

hitid

aeC

alam

opte

ryx

gosl

inei

c0

00

00

00

00

00

00

10

00

00

00

00

00

01

0

Byt

hitid

aeG

ram

mon

us

clau

dei

c0

08

00

00

00

00

10

00

00

00

00

00

00

04

0

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hitid

aeO

gilb

ia b

oehl

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c0

00

00

36

43

10

20

05

50

00

00

00

00

1111

7

Byt

hitid

aeO

gilb

ia sa

baji

c0

00

00

00

13

11

00

00

00

00

00

00

00

01

0

Byt

hitid

aeO

gilb

ia sp

.c

00

00

10

00

00

00

00

00

00

00

00

00

00

02

Byt

hitid

aeO

gilb

ia su

arez

aec

00

00

01

10

00

00

00

00

00

00

00

00

00

00

Byt

hitid

aeo

gilb

ia/

ogi

lbic

hthy

s sp.

c1

52

90

00

00

00

00

00

00

00

00

00

00

00

0

Byt

hitid

aeO

gilb

icht

hys k

akuk

ic

00

00

00

00

40

00

00

00

00

00

00

00

00

20

Byt

hitid

aeO

gilb

icht

hys

long

iman

usc

00

00

01

00

02

00

00

01

00

00

00

00

02

30

Byt

hitid

aeO

gilb

icht

hys s

p.c

00

00

130

00

00

00

00

00

00

00

00

00

00

10

Byt

hitid

aeSt

ygno

brot

ula

late

bric

ola

c0

00

00

00

00

00

00

00

00

00

00

01

00

01

0

cal

liony

mid

aeC

allio

nym

us b

aird

ic

00

00

00

10

00

00

00

00

01

00

01

00

00

00

car

angi

dae

Car

anx

rube

r*c

00

00

00

00

00

00

00

00

00

00

00

00

00

01

cha

enop

sida

eAc

anth

embl

emar

ia

aspe

raI

012

20

00

152

22

06

00

813

28

40

51

10

09

1413

cha

enop

sida

eAc

anth

embl

emar

ia

chap

lini

I0

00

00

30

01

00

00

00

00

00

00

80

00

00

0

cha

enop

sida

eAc

anth

embl

emar

ia

mar

iaI

00

01

01

00

10

00

00

01

00

00

01

00

00

00

cha

enop

sida

eAc

anth

embl

emar

ia

spin

osa

I0

02

00

00

00

02

00

00

00

50

00

05

00

1211

0

cha

enop

sida

eC

oral

lioze

tus

car d

onae

I0

00

00

30

00

00

00

00

00

00

00

00

00

00

0

76 K.l. ilves et. Al.

App

endi

x 1.

con

t. R

aw a

bund

ance

dat

a fo

r His

toric

al, 2

006,

and

201

2 co

llect

ions

at s

ites G

reen

Cay

(GC

), N

orth

/Lon

g C

ay (N

L), L

yfor

d C

ay/C

lifto

n Po

int (

LC),

and

Del

apor

te

Poin

t (D

P). T

roph

ic g

roup

(TrG

) abb

revi

atio

ns: C

arni

vore

(C),

Her

bivo

re (H

), In

verti

vore

(I),

om

nivo

re (o

), Pl

ankt

ivor

e (P

). Sp

ecie

s mar

ked

with

an

* w

ere

excl

uded

as ‘

non-

reef

’ and

+ in

dica

tes s

peci

es e

xclu

sion

due

to u

npre

cede

nted

cat

ch a

t DP1

-201

0.

Fam

ilySp

ecie

sTr

GG

C1-

2010

GC

2-20

10G

C3-

2010

NL1

-20

10D

P1-

2010

GC

1-20

06G

C2-

2006

GC

3-20

06N

L1-

2006

NL2

-20

06N

L3-

2006

LC1-

2006

LC2-

2006

LC3-

2006

DP1

-20

06D

P2-

2006

DP3

-20

06LC

1-19

56D

P1-

1955

cD

P1-

1955

bD

P1-

1955

aN

L1-

1955

GC

3-19

56G

C3-

1957

aG

C3-

1957

bG

C3-

1959

aG

C3-

1959

bG

C3-

1969

cha

enop

sida

eEm

blem

aria

pa

ndio

nis

P0

00

00

00

00

12

00

00

02

00

00

00

00

00

0

cha

enop

sida

eEm

blem

ario

psis

ba

ham

ensi

sI

00

00

00

00

00

00

00

00

01

00

00

00

00

20

cha

enop

sida

eEm

blem

ario

psis

oc

cide

ntal

isI

00

00

00

00

00

00

00

00

00

00

00

01

00

00

cha

enop

sida

eEm

blem

ario

psis

si

gnife

raI

00

00

00

00

00

00

00

00

00

00

00

00

14

70

cha

enop

sida

eEm

blem

ario

psis

sp.

I0

11

00

00

00

00

00

00

00

00

00

00

00

00

0

cha

enop

sida

eH

emie

mbl

emar

ia

sim

ulus

c0

00

00

01

20

03

00

00

13

00

00

00

02

01

0

cha

enop

sida

eLu

caya

blen

nius

zi

ngar

oc

04

20

00

37

00

00

03

00

00

00

00

20

33

02

cha

enop

sida

eSt

athm

onot

us

hem

phill

iic

00

00

00

01

00

00

00

00

00

00

00

00

00

00

cha

enop

sida

eSt

athm

onot

us st

ahli

c0

00

00

17

00

00

00

00

00

00

00

00

00

00

0

cha

enop

sida

eSt

athm

onot

us te

kla

c0

20

00

00

00

00

00

00

00

00

00

00

00

00

0

cha

etod

ontid

aeC

haet

odon

ca

pist

ratu

sI

00

00

00

00

00

00

00

10

00

10

00

00

02

00

chl

opsi

dae

Cat

esby

a ps

eudo

mur

aena

c0

04

00

00

00

00

00

00

00

10

00

00

00

00

0

chl

opsi

dae

Chi

lor h

inus

su

enso

nii

c0

00

00

01

00

00

00

00

05

00

00

00

00

00

0

chl

opsi

dae

Kau

pich

thys

hy

opr o

roid

esc

14

71

23

05

31

20

12

22

22

51

21

00

16

163

chl

opsi

dae

Kau

pich

thys

nu

chal

isc

00

20

00

01

00

00

00

00

00

00

00

00

00

20

cirr

hitid

aeAm

blyc

irr h

itus

pino

sI

00

00

00

00

01

44

10

00

01

12

04

00

00

10

clu

peid

aeJe

nkin

sia

sp.*

I0

01

00

00

793

00

00

00

00

00

00

00

029

00

0

con

grid

aeAr

ioso

ma

bale

aric

umc

00

00

00

01

00

00

00

00

00

00

00

00

02

05

con

grid

aeC

onge

r tri

pori

ceps

c0

01

00

00

20

00

00

00

00

00

00

00

00

40

0

con

grid

aeH

eter

ocon

ger

long

issi

mus

P0

00

00

00

00

00

00

00

00

00

00

01

00

20

1

cyn

oglo

ssid

aeSy

mph

urus

om

mas

pilu

s *I

00

00

00

00

00

00

00

00

00

00

00

00

00

01

dac

tylo

scop

idae

Dac

tylo

scop

us

trid

igita

tus

c0

30

05

00

00

00

00

00

00

03

03

00

00

01

0

dac

tylo

scop

idae

Gill

ellu

s gre

yae

c0

10

12

00

10

00

00

00

00

01

02

00

00

00

1

dac

tylo

scop

idae

Gill

ellu

s ura

nide

ac

00

00

00

00

00

02

00

23

00

00

00

00

00

20

detection of shifts in corAl reef fish AssemblAge structure At reefs of neW Providence islAnd 77

App

endi

x 1

cont

. Raw

abu

ndan

ce d

ata

for H

isto

rical

, 200

6, a

nd 2

012

colle

ctio

ns a

t site

s Gre

en C

ay (G

C),

Nor

th/L

ong

Cay

(NL)

, Lyf

ord

Cay

/Clif

ton

Poin

t (LC

), an

d D

elap

orte

Po

int (

DP)

. Tro

phic

gro

up (T

rG) a

bbre

viat

ions

: Car

nivo

re (C

), H

erbi

vore

(H),

Inve

rtivo

re (I

), o

mni

vore

(o),

Plan

ktiv

ore

(P).

Spec

ies m

arke

d w

ith a

n *

wer

e ex

clud

ed a

s ‘no

n-re

ef’ a

nd +

indi

cate

s spe

cies

exc

lusi

on d

ue to

unp

rece

dent

ed c

atch

at D

P1-2

010.

Fam

ilySp

ecie

sTr

GG

C1-

2010

GC

2-20

10G

C3-

2010

NL1

-20

10D

P1-

2010

GC

1-20

06G

C2-

2006

GC

3-20

06N

L1-

2006

NL2

-20

06N

L3-

2006

LC1-

2006

LC2-

2006

LC3-

2006

DP1

-20

06D

P2-

2006

DP3

-20

06LC

1-19

56D

P1-

1955

cD

P1-

1955

bD

P1-

1955

aN

L1-

1955

GC

3-19

56G

C3-

1957

aG

C3-

1957

bG

C3-

1959

aG

C3-

1959

bG

C3-

1969

dac

tylo

scop

idae

Leur

ochi

lus a

con

c0

00

00

02

00

00

00

00

00

02

01

00

00

00

0

dac

tylo

scop

idae

Plat

ygill

ellu

s ru

broc

inct

usc

13

18

130

22

20

01

00

10

00

41

10

00

02

10

dio

dont

idae

Dio

don

holo

cant

hus*

I1

00

00

00

00

00

10

01

00

00

00

00

00

00

0

Gob

ieso

cida

eAc

yrtu

s art

ius

I0

00

00

00

00

00

00

00

00

021

11

20

03

04

1

Gob

ieso

cida

eA

cyrtu

s ru

bigi

nosu

sI

02

20

00

00

00

00

00

00

00

00

00

00

00

00

Gob

iidae

Cor

ypho

pter

us

allo

ides

o0

00

00

00

80

00

00

00

00

00

00

00

01

16

0

Gob

iidae

Cor

ypho

pter

us

dicr

uso

144

76

00

91

11

04

20

53

30

00

02

01

25

60

Gob

iidae

Cor

ypho

pter

us

eido

lon

H0

00

00

01

20

01

00

60

01

00

00

00

00

30

0

Gob

iidae

Cor

ypho

pter

us

glau

cofr

aenu

m/

tort

ugae

o0

195

00

013

50

00

00

23

11

05

11

22

011

1915

5

Gob

iidae

Cor

ypho

pter

us

hyal

inus

o0

05

00

00

70

00

00

150

11

00

00

00

08

02

0

Gob

iidae

Cor

ypho

pter

us

pers

onat

usP

00

70

00

021

00

00

18

00

10

00

00

00

181

716

Gob

iidae

Cor

ypho

pter

us sp

.o

02

60

00

30

00

00

00

00

00

00

00

00

00

00

Gob

iidae

Cor

ypho

pter

us

thri

xo

00

00

00

00

00

02

00

02

12

00

00

00

00

00

Gob

iidae

Cte

nogo

bius

sa

epep

alle

nsI

00

00

00

00

00

00

00

00

00

00

00

00

30

10

Gob

iidae

Elac

atin

us

evel

ynae

I0

00

00

00

10

02

00

00

00

00

00

02

06

51

0

Gob

iidae

Elac

atin

us g

enie

I0

67

33

42

66

22

10

03

33

14

30

25

112

21

7

Gob

iidae

Gna

thol

epis

ca

uere

nsis

o0

10

00

00

20

76

130

12

21

20

00

20

14

32

7

Gob

iidae

Gob

iidae

sp.

o0

491

00

00

00

00

00

00

00

00

00

00

00

00

1

Gob

iidae

Lyth

rypn

us e

lass

ono

00

00

00

00

00

22

07

00

00

00

00

21

45

70

Gob

iidae

Lyth

rypn

us

hete

r och

rom

ao

00

00

00

838

00

00

00

04

00

00

00

51

1749

730

Gob

iidae

Lyth

rypn

us o

kapi

ao

00

00

00

00

00

00

00

00

00

00

00

00

00

10

Gob

iidae

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rypn

us sp

ilus

o0

00

00

00

00

01

00

00

00

00

00

00

01

1723

10

Gob

iidae

Prio

lepi

s hip

oliti

I0

016

10

10

70

10

00

02

211

01

50

33

020

613

7

Gob

iidae

Psilo

tris

ba

trac

hode

so

01

00

00

10

00

00

00

01

00

20

00

00

05

54

Gob

iidae

Psilo

tris

cel

sus

o0

00

00

00

00

00

00

00

00

00

00

00

02

10

0

78 K.l. ilves et. Al.

App

endi

x 1c

ont.

Raw

abu

ndan

ce d

ata

for H

isto

rical

, 200

6, a

nd 2

012

colle

ctio

ns a

t site

s Gre

en C

ay (G

C),

Nor

th/L

ong

Cay

(NL)

, Lyf

ord

Cay

/Clif

ton

Poin

t (LC

), an

d D

elap

orte

Po

int (

DP)

. Tro

phic

gro

up (T

rG) a

bbre

viat

ions

: Car

nivo

re (C

), H

erbi

vore

(H),

Inve

rtivo

re (I

), o

mni

vore

(o),

Plan

ktiv

ore

(P).

Spec

ies m

arke

d w

ith a

n *

wer

e ex

clud

ed a

s ‘no

n-re

ef’ a

nd +

indi

cate

s spe

cies

exc

lusi

on d

ue to

unp

rece

dent

ed c

atch

at D

P1-2

010.

Fam

ilySp

ecie

sTr

GG

C1-

2010

GC

2-20

10G

C3-

2010

NL1

-20

10D

P1-

2010

GC

1-20

06G

C2-

2006

GC

3-20

06N

L1-

2006

NL2

-20

06N

L3-

2006

LC1-

2006

LC2-

2006

LC3-

2006

DP1

-20

06D

P2-

2006

DP3

-20

06LC

1-19

56D

P1-

1955

cD

P1-

1955

bD

P1-

1955

aN

L1-

1955

GC

3-19

56G

C3-

1957

aG

C3-

1957

bG

C3-

1959

aG

C3-

1959

bG

C3-

1969

Gob

iidae

Psilo

tris s

p.o

00

20

00

00

00

00

00

00

00

00

00

00

00

00

Gob

iidae

Tigr

igob

ius d

ilepi

so

00

00

03

10

00

00

00

00

00

00

00

00

00

00

Gra

mm

atid

aeG

ram

ma

lore

toI

011

165

63

09

214

02

2021

1214

420

285

50

49

5225

6711

610

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536

1

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00

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Labr

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09

83

00

16

00

17

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70

Labr

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416

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00

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02

00

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00

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03

00

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00

Labr

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00

130

00

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00

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Labr

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detection of shifts in corAl reef fish AssemblAge structure At reefs of neW Providence islAnd 79

App

endi

x 1

cont

. Raw

abu

ndan

ce d

ata

for H

isto

rical

, 200

6, a

nd 2

012

colle

ctio

ns a

t site

s Gre

en C

ay (G

C),

Nor

th/L

ong

Cay

(NL)

, Lyf

ord

Cay

/Clif

ton

Poin

t (LC

), an

d D

elap

orte

Po

int (

DP)

. Tro

phic

gro

up (T

rG) a

bbre

viat

ions

: Car

nivo

re (C

), H

erbi

vore

(H),

Inve

rtivo

re (I

), o

mni

vore

(o),

Plan

ktiv

ore

(P).

Spec

ies m

arke

d w

ith a

n *

wer

e ex

clud

ed a

s ‘no

n-re

ef’ a

nd +

indi

cate

s spe

cies

exc

lusi

on d

ue to

unp

rece

dent

ed c

atch

at D

P1-2

010.

Fam

ilySp

ecie

sTr

GG

C1-

2010

GC

2-20

10G

C3-

2010

NL1

-20

10D

P1-

2010

GC

1-20

06G

C2-

2006

GC

3-20

06N

L1-

2006

NL2

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06N

L3-

2006

LC1-

2006

LC2-

2006

LC3-

2006

DP1

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06D

P2-

2006

DP3

-20

06LC

1-19

56D

P1-

1955

cD

P1-

1955

bD

P1-

1955

aN

L1-

1955

GC

3-19

56G

C3-

1957

aG

C3-

1957

bG

C3-

1959

aG

C3-

1959

bG

C3-

1969

Labr

idae

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asso

ma

bifa

scia

tum

I5

1235

2063

5425

4617

17

443

231

1914

1514

176

2412

15

1416

7

Labr

isom

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00

00

00

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00

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00

00

00

00

00

00

0

Labr

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00

00

00

00

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00

00

00

00

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00

00

00

00

Labr

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3510

2846

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330

84

60

019

20

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710

173

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12

1

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26

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60

180

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00

260

00

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85

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158

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50

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91

00

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00

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rolin

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00

00

32

00

00

00

00

20

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00

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Labr

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50

01

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01

20

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09

50

00

00

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280

1315

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00

02

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30

00

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00

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Labr

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Labr

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126

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00

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Labr

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00

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00

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Labr

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181

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20

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73

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01

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00

1

80 K.l. ilves et. Al.

App

endi

x 1

cont

. Raw

abu

ndan

ce d

ata

for H

isto

rical

, 200

6, a

nd 2

012

colle

ctio

ns a

t site

s Gre

en C

ay (G

C),

Nor

th/L

ong

Cay

(NL)

, Lyf

ord

Cay

/Clif

ton

Poin

t (LC

), an

d D

elap

orte

Po

int (

DP)

. Tro

phic

gro

up (T

rG) a

bbre

viat

ions

: Car

nivo

re (C

), H

erbi

vore

(H),

Inve

rtivo

re (I

), o

mni

vore

(o),

Plan

ktiv

ore

(P).

Spec

ies m

arke

d w

ith a

n *

wer

e ex

clud

ed a

s ‘no

n-re

ef’ a

nd +

indi

cate

s spe

cies

exc

lusi

on d

ue to

unp

rece

dent

ed c

atch

at D

P1-2

010.

Fam

ilySp

ecie

sTr

GG

C1-

2010

GC

2-20

10G

C3-

2010

NL1

-20

10D

P1-

2010

GC

1-20

06G

C2-

2006

GC

3-20

06N

L1-

2006

NL2

-20

06N

L3-

2006

LC1-

2006

LC2-

2006

LC3-

2006

DP1

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06D

P2-

2006

DP3

-20

06LC

1-19

56D

P1-

1955

cD

P1-

1955

bD

P1-

1955

aN

L1-

1955

GC

3-19

56G

C3-

1957

aG

C3-

1957

bG

C3-

1959

aG

C3-

1959

bG

C3-

1969

Mon

acan

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anth

us

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eri

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00

00

00

04

11

00

00

00

00

00

00

00

13

0

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acan

thid

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hisp

idus

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00

01

00

00

00

00

00

00

00

00

00

00

00

0

Mor

ingu

idae

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ingu

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war

dsi

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90

11

212

134

62

70

010

46

35

150

20

13

1217

3

Mul

lidae

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loid

icht

hys

mar

tinic

usI

01

00

00

01

00

00

00

00

00

00

00

00

10

00

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lidae

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00

00

00

00

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20

00

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00

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00

00

00

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00

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ac

21

223

163

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100

58

20

810

02

160

44

41

01

175

Mur

aeni

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nigr

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sc

10

00

77

00

00

00

00

10

00

27

13

00

00

00

Mur

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11

00

72

00

00

02

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10

10

59

11

00

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Mur

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00

03

20

10

02

01

00

60

00

00

00

00

00

00

Mur

aeni

dae

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pter

ygiu

s m

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ariu

sc

00

04

03

00

71

03

00

80

00

00

01

00

00

10

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halid

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00

00

00

00

00

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oph

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00

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00

00

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00

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00

00

00

00

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01

00

31

19

oph

icht

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00

00

00

00

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oph

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00

00

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00

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00

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10

detection of shifts in corAl reef fish AssemblAge structure At reefs of neW Providence islAnd 81

App

endi

x 1

cont

. Raw

abu

ndan

ce d

ata

for H

isto

rical

, 200

6, a

nd 2

012

colle

ctio

ns a

t site

s Gre

en C

ay (G

C),

Nor

th/L

ong

Cay

(NL)

, Lyf

ord

Cay

/Clif

ton

Poin

t (LC

), an

d D

elap

orte

Po

int (

DP)

. Tro

phic

gro

up (T

rG) a

bbre

viat

ions

: Car

nivo

re (C

), H

erbi

vore

(H),

Inve

rtivo

re (I

), o

mni

vore

(o),

Plan

ktiv

ore

(P).

Spec

ies m

arke

d w

ith a

n *

wer

e ex

clud

ed a

s ‘no

n-re

ef’ a

nd +

indi

cate

s spe

cies

exc

lusi

on d

ue to

unp

rece

dent

ed c

atch

at D

P1-2

010.

Fam

ilySp

ecie

sTr

GG

C1-

2010

GC

2-20

10G

C3-

2010

NL1

-20

10D

P1-

2010

GC

1-20

06G

C2-

2006

GC

3-20

06N

L1-

2006

NL2

-20

06N

L3-

2006

LC1-

2006

LC2-

2006

LC3-

2006

DP1

-20

06D

P2-

2006

DP3

-20

06LC

1-19

56D

P1-

1955

cD

P1-

1955

bD

P1-

1955

aN

L1-

1955

GC

3-19

56G

C3-

1957

aG

C3-

1957

bG

C3-

1959

aG

C3-

1959

bG

C3-

1969

oph

idiid

aeO

toph

idiu

m

dorm

itato

r*c

00

00

00

00

00

00

00

00

00

00

00

00

11

00

oph

idiid

aePe

troty

x

sang

uine

usc

00

141

50

00

10

00

00

00

01

50

50

00

20

140

opi

stog

nath

idae

Opi

stog

nath

us

auri

frons

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00

00

00

00

02

00

00

00

00

00

00

00

00

00

opi

stog

nath

idae

Opi

stog

nath

us

max

illos

us*

c0

00

04

10

00

02

00

02

00

00

20

00

00

00

0

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raci

idae

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neso

mus

tri

quet

er*

I0

00

01

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00

00

00

00

00

00

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00

0

Pem

pher

idae

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pher

is

scho

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i+P

131

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acan

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us

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olor

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8321

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eri

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00

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04

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00

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02

00

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idae

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us sp

.H

00

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00

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00

00

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00

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Scar

idae

Scar

us ta

enio

pter

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us v

etul

aH

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00

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00

00

10

04

00

02

01

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idae

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isom

a au

rofr e

natu

mH

06

00

114

46

01

47

02

14

24

00

17

00

00

10

82 K.l. ilves et. Al.

App

endi

x 1

cont

. Raw

abu

ndan

ce d

ata

for H

isto

rical

, 200

6, a

nd 2

012

colle

ctio

ns a

t site

s Gre

en C

ay (G

C),

Nor

th/L

ong

Cay

(NL)

, Lyf

ord

Cay

/Clif

ton

Poin

t (LC

), an

d D

elap

orte

Po

int (

DP)

. Tro

phic

gro

up (T

rG) a

bbre

viat

ions

: Car

nivo

re (C

), H

erbi

vore

(H),

Inve

rtivo

re (I

), o

mni

vore

(o),

Plan

ktiv

ore

(P).

Spec

ies m

arke

d w

ith a

n *

wer

e ex

clud

ed a

s ‘no

n-re

ef’ a

nd +

indi

cate

s spe

cies

exc

lusi

on d

ue to

unp

rece

dent

ed c

atch

at D

P1-2

010.

Fam

ilySp

ecie

sTr

GG

C1-

2010

GC

2-20

10G

C3-

2010

NL1

-20

10D

P1-

2010

GC

1-20

06G

C2-

2006

GC

3-20

06N

L1-

2006

NL2

-20

06N

L3-

2006

LC1-

2006

LC2-

2006

LC3-

2006

DP1

-20

06D

P2-

2006

DP3

-20

06LC

1-19

56D

P1-

1955

cD

P1-

1955

bD

P1-

1955

aN

L1-

1955

GC

3-19

56G

C3-

1957

aG

C3-

1957

bG

C3-

1959

aG

C3-

1959

bG

C3-

1969

Scar

idae

Spar

isom

a ra

dian

sH

06

00

00

00

03

10

00

00

00

00

04

00

01

10

Scar

idae

Spar

isom

a ru

brip

inne

H0

00

10

00

00

00

10

00

00

00

10

10

00

00

0

Scar

idae

Spar

isom

a sp

.H

00

00

00

20

00

01

00

00

00

00

00

00

10

01

Scar

idae

Spar

isom

a vi

ride

H8

1416

523

156

39

21

62

014

31

05

30

60

04

12

0

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enid

aeEq

uetu

s pun

ctat

usc

00

00

00

00

00

01

00

00

00

00

00

00

00

00

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enid

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s sp.

c0

00

00

00

00

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10

00

00

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00

00

00

00

0

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paen

idae

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ois v

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ns/

mile

sc

02

10

00

00

00

00

00

01

00

00

00

00

00

00

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paen

idae

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paen

a in

erm

isc

00

00

00

11

00

00

00

00

10

00

00

20

00

00

Scor

paen

idae

Scor

paen

a pl

umie

ric

00

00

00

00

00

03

10

00

00

00

00

00

00

00

Scor

paen

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paen

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ribb

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14

314

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41

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35

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00

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0

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anid

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36

11

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00

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00

00

00

10

01

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10

00

00

0

Serr

anid

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s ad

scen

sion

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00

00

00

00

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00

00

01

01

00

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00

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anid

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00

00

00

14

01

00

00

00

00

00

20

00

00

00

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anid

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s st

riat

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00

00

00

00

00

00

00

01

00

00

00

00

00

00

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anid

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lect

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00

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00

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00

00

00

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01

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00

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00

00

00

00

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00

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anid

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00

00

00

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anid

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olor

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00

00

00

10

00

00

00

00

00

00

00

09

00

0

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anid

aeLi

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pom

a ca

rmab

ic

00

00

00

00

00

10

00

00

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00

00

00

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anid

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pom

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00

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00

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00

0

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anid

aeM

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rca

tigri

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00

00

00

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ma

greg

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05

10

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11

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01

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24

40

10

11

023

2

detection of shifts in corAl reef fish AssemblAge structure At reefs of neW Providence islAnd 83

App

endi

x 1

cont

. Raw

abu

ndan

ce d

ata

for H

isto

rical

, 200

6, a

nd 2

012

colle

ctio

ns a

t site

s Gre

en C

ay (G

C),

Nor

th/L

ong

Cay

(NL)

, Lyf

ord

Cay

/Clif

ton

Poin

t (LC

), an

d D

elap

orte

Po

int (

DP)

. Tro

phic

gro

up (T

rG) a

bbre

viat

ions

: Car

nivo

re (C

), H

erbi

vore

(H),

Inve

rtivo

re (I

), o

mni

vore

(o),

Plan

ktiv

ore

(P).

Spec

ies m

arke

d w

ith a

n *

wer

e ex

clud

ed a

s ‘no

n-re

ef’ a

nd +

indi

cate

s spe

cies

exc

lusi

on d

ue to

unp

rece

dent

ed c

atch

at D

P1-2

010.

Fam

ilySp

ecie

sTr

GG

C1-

2010

GC

2-20

10G

C3-

2010

NL1

-20

10D

P1-

2010

GC

1-20

06G

C2-

2006

GC

3-20

06N

L1-

2006

NL2

-20

06N

L3-

2006

LC1-

2006

LC2-

2006

LC3-

2006

DP1

-20

06D

P2-

2006

DP3

-20

06LC

1-19

56D

P1-

1955

cD

P1-

1955

bD

P1-

1955

aN

L1-

1955

GC

3-19

56G

C3-

1957

aG

C3-

1957

bG

C3-

1959

aG

C3-

1959

bG

C3-

1969

Serr

anid

aery

ptic

us

carp

ente

ri/su

bbifr

enat

usc

70

190

61

06

11

20

02

02

01

31

12

10

20

11

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anid

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ptic

us

sapo

nace

usc

10

00

01

00

00

00

00

10

00

03

00

00

00

00

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anid

aery

ptic

us sp

.c

00

10

00

00

00

00

00

00

00

00

00

00

00

00

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anid

aeSe

rran

us b

aldw

ini

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00

00

00

00

30

00

00

00

00

00

00

00

00

0

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us

taba

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00

00

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anid

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us ti

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00

20

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01

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nath

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Cos

moc

ampu

s el

ucen

sI

00

10

10

00

00

00

00

00

00

00

00

00

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00

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nath

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Hal

icam

pus

crin

itus

I0

00

00

00

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00

00

00

00

00

00

00

00

1

Syno

dont

idae

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00

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00

00

00

00

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20

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10

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23

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35

12

tetra

odon

tidae

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thig

aste

r ro

stra

tao

316

70

23

1122

32

55

19

612

94

212

09

31

222

2114

tetra

odon

tidae

Sphe

roid

es sp

.*I

00

00

00

00

00

00

00

00

01

00

00

00

00

00

trip

tery

giid

aeEn

nean

ecte

s al

tivel

isI

243

112

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821

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234

1434

14

trip

tery

giid

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nean

ecte

s at

roru

sI

00

00

00

00

00

00

01

00

00

00

00

01

00

00

trip

tery

giid

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nean

ecte

s bo

ehlk

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30

01

317

10

50

00

00

60

00

20

00

00

00

00

trip

tery

giid

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nean

ecte

s jo

rdan

iI

11

01

60

00

60

00

00

10

00

10

016

10

00

00

trip

tery

giid

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nean

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s pe

ctor

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I0

00

021

00

01

00

30

014

00

036

36

00

00

00

0

trip

tery

giid

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511

00

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00

00

00

00

0

84 K.l. ilves et. Al.

Appendix 2. Multidimensional scaling and hierarchical cluster plots of Bray-Curtis similarities of standardized, fourth root transformed abundance data across the historical, 2006, and 2010 sites. Contour lines mark sites with 50% similarity of reef fish assemblage structure.

detection of shifts in corAl reef fish AssemblAge structure At reefs of neW Providence islAnd 85

App

endi

x 3.

Mea

n, st

anda

rd er

ror (

SE),

and

t-tes

ts o

f diff

eren

ces b

etw

een

troph

ic g

roup

rela

tive a

bund

ance

mea

ns b

etw

een

time p

erio

ds: ‘

His

toric

’ (19

56-1

969)

, ‘R

ecen

t’ (2

006,

20

10),

2006

, and

201

0, w

ith n

umbe

r of s

ites

(N) f

or e

ach

dept

h/tim

e pe

riod.

one

-sam

ple

t-tes

ts w

ere

cond

ucte

d fo

r com

paris

ons

invo

lvin

g on

ly 2

010

deep

site

s (N

=1).

Bol

d va

lues

with

* a

re si

gnifi

cant

dif

eren

ces b

etw

een

time

perio

ds (P

≤ 0

.05)

.

Trop

hic

grou

pM

ean

rela

tive

abun

danc

e (S

E)t-

test

(Hist

oric

al, R

ecen

t)t-

test

(Hist

oric

al, 2

006)

t- te

st(H

istor

ical

, 201

0)t-

test

(200

6, 2

010)

His

toric

alr

ecen

t20

0620

10|t|

P|t|

P|t|

P|t|

P

SHA

LLO

WN

= 5

N =

7N

= 4

N =

3

Her

bivo

res

11.0

(1.4

)18

.4 (2

.4)

16.2

(3.6

)21

.4 (2

.6)

2.39

0.04

*1.

470.

183.

860.

01*

1.10

0.32

Pla

nktiv

ores

8.7

(3.9

)2.

9 (1

.6)

4.5

(2.6

)0.

8 (0

.8)

1.52

0.16

0.84

0.43

1.48

0.19

1.18

0.29

om

nivo

res

6.2

(1.7

)4.

2 (0

.8)

4.6

(1.3

)3.

6 (1

.0)

1.20

0.26

0.72

0.50

1.12

0.31

0.58

0.59

Inv

ertiv

ores

50.0

(2.5

)51

.7 (2

.9)

56.2

(3.4

)45

.8 (2

.0)

0.43

0.68

1.50

0.18

1.15

0.30

2.40

0.06

Car

nivo

res

24.1

(6.1

)22

.7 (2

.5)

18.5

(2.6

)28

.3 (1

.6)

0.23

0.82

0.76

0.47

0.52

0.62

2.92

0.03

*

DEE

PN

= 6

N =

5N

= 4

N =

1

Her

bivo

res

1.8

(0.3

)6.

2 (1

.2)

6.9

(1.3

)3.

74.

030.

003*

4.77

0.00

1*2.

290.

071.

130.

34

Pla

nktiv

ores

20.4

(3.3

)10

.1 (2

.5)

10.8

(3.1

)7.

52.

390.

04*

2.00

0.08

1.46

0.21

0.48

0.67

om

nivo

res

13.3

(3.2

)15

.6 (3

.9)

15.0

(4.9

)18

.10.

460.

650.

300.

770.

280.

800.

460.

65

Inv

ertiv

ores

54.6

(4.8

)53

.3 (4

.9)

52.7

(6.3

)55

.90.

180.

860.

240.

820.

100.

920.

230.

84

Car

nivo

res

10.0

(1.3

)14

.7 (1

.3)

14.7

(1.6

)14

.72.

620.

03*

2.32

0.05

*1.

400.

220.

010.

99

86 K.l. ilves et. Al.

App

endi

x 4.

Mea

n, s

tand

ard

erro

r (S

E), a

nd t-

test

s of

diff

eren

ces

betw

een

fam

ily r

elat

ive

abun

danc

e m

eans

bet

wee

n tim

e pe

riods

: ‘H

isto

ric’ (

1956

-196

9), ‘

Rec

ent’

(200

6,

2010

), 20

06, a

nd 2

010,

with

num

ber o

f site

s (N

) for

each

dep

th/ti

me p

erio

d. o

ne-s

ampl

e t-te

sts w

ere c

ondu

cted

for c

ompa

rison

s inv

olvi

ng o

nly

2010

dee

p si

tes (

N=1

). Fa

mili

es

with

< 1

% m

ean

rela

tive

abun

danc

e ac

ross

all

site

s wer

e gr

oupe

d as

“o

ther

”. B

old

valu

es w

ith *

are

sign

ifica

nt d

ifer

ence

s bet

wee

n tim

e pe

riods

(P ≤

0.0

5).

Fam

ilyM

ean

rela

tive

abun

danc

e (S

E)t-t

est

(Hist

oric

al, R

ecen

t)t-t

est

(Hist

oric

al, 2

006)

t-tes

t(H

istor

ical

, 201

0)t-t

est

(200

6, 2

010)

His

toric

alr

ecen

t 20

0620

10|t|

P|t|

P|t|

P|t|

PSH

ALL

OW

N =

5N

= 7

N =

4N

= 3

Aca

nthu

ridae

1.7

(0.7

)2.

9 (0

.3)

2.8

(0.4

)3.

0 (0

.3)

1.88

0.09

1.42

0.19

1.32

0.23

0.22

0.83

Apo

goni

dae

16.1

(6.3

)11

.5 (3

.6)

14.0

(5.9

)8.

2 (3

.0)

0.68

0.51

0.24

0.81

0.91

0.40

0.77

0.47

Ble

nniid

ae1.

8 (0

.7)

3.3

(0.8

)2.

5 (0

.8)

4.5

(1.4

)1.

380.

200.

640.

541.

900.

111.

370.

23 C

haen

opsid

ae3.

3 (1

.4)

0.9

(0.3

)1.

5 (0

.3)

0.1

(0.1

)1.

930.

081.

100.

311.

680.

143.

830.

01*

Gob

iidae

2.7

(0.5

)2.

3 (0

.6)

2.8

(0.6

)1.

7 (1

.2)

0.49

0.63

0.10

0.92

0.96

0.37

0.94

0.39

Gra

mm

atid

ae2.

1 (0

.6)

1.1

(0.6

)1.

2 (0

.9)

0.9

(0.7

)1.

180.

270.

820.

441.

200.

280.

230.

83 H

oloc

entr

idae

6.3

(1.3

)17

.6 (1

.4)

17.7

(2.0

)17

.5 (2

.5)

5.54

<0.0

01*

4.95

0.00

2*4.

340.

005*

0.07

0.95

Lab

ridae

7.5

(1.0

)10

.3 (1

.8)

10.4

(2.2

)10

.2 (3

.7)

1.19

0.26

1.26

0.25

0.90

0.40

0.03

0.97

Lab

risom

idae

14.5

(3.1

)16

.6 (1

.8)

14.4

(2.4

)19

.6 (1

.9)

0.64

0.54

0.02

0.98

1.18

0.28

1.59

0.17

Mor

ingu

idae

1.7

(0.8

)0.

8 (0

.3)

1.2

(0.4

)0.

2 (0

.1)

1.25

0.24

0.49

0.64

1.47

0.19

2.39

0.06

Mur

aeni

dae

4.0

(0.9

)3.

8 (0

.4)

3.9

(0.5

)3.

6 (0

.6)

0.26

0.80

0.07

0.95

0.33

0.75

0.41

0.70

Pom

acen

trida

e8.

4 (1

.8)

11.2

(2.3

)10

.6 (4

.2)

11.9

(0.9

)0.

880.

400.

520.

621.

390.

210.

260.

81 S

carid

ae3.

1 (1

.3)

4.4

(0.9

)4.

9 (1

.4)

3.8

(0.9

)0.

920.

380.

960.

370.

400.

700.

590.

58 S

erra

nida

e3.

9 (0

.9)

2.2

(1.0

)1.

6 (0

.7)

3.1

(2.2

)1.

220.

251.

910.

100.

420.

690.

720.

51 S

ynod

ontid

ae2.

4 (1

.4)

0.3

(0.1

)0.

5 (0

.2)

0.1

(0.1

)1.

840.

101.

260.

251.

260.

251.

420.

21 t

etra

odon

tidae

1.9

(0.7

)0.

8 (0

.2)

0.9

(0.1

)0.

7 (0

.6)

1.64

0.13

1.22

0.26

1.14

0.30

0.37

0.73

trip

tery

giid

ae6.

9 (2

.6)

3.7

(0.6

)4.

1 (1

.0)

3.1

(0.8

)1.

420.

190.

930.

391.

080.

320.

680.

53 O

ther

11.7

(2.4

)6.

2 (0

.8)

5.0

(0.6

)7.

7 (1

.2)

2.50

0.03

*3.

910.

006*

1.19

0.28

2.23

0.08

DEE

PN

= 6

N =

5N

= 4

N =

1 A

cant

hurid

ae0.

9 (0

.2)

0.7

(0.1

)0.

7 (0

.2)

0.7

0.45

0.66

0.39

0.71

0.24

0.82

0.06

0.95

Apo

goni

dae

21.2

(7.2

)15

.0 (3

.2)

17.4

(2.8

)5.

70.

730.

480.

420.

690.

820.

451.

870.

16 B

lenn

iidae

0.03

(0.0

3)0.

05 (0

.05)

0.03

(0.0

2)0.

00.

320.

750.

180.

860.

380.

720.

450.

69 C

haen

opsid

ae2.

6 (0

.6)

2.2

(0.5

)2.

6 (0

.5)

0.8

0.46

0.66

0.02

0.98

3.09

0.03

*3.

280.

05*

Gob

iidae

11.6

(2.0

)13

.1 (5

.0)

14.8

(6.0

)6.

30.

300.

770.

600.

571.

020.

360.

630.

57 G

ram

mat

idae

18.5

(3.6

)14

.7 (2

.2)

13.7

(2.5

)18

.70.

840.

420.

960.

360.

030.

980.

880.

44

detection of shifts in corAl reef fish AssemblAge structure At reefs of neW Providence islAnd 87

App

endi

x 4

cont

. Mea

n, st

anda

rd e

rror

(SE)

, and

t-te

sts o

f diff

eren

ces b

etw

een

fam

ily re

lativ

e ab

unda

nce

mea

ns b

etw

een

time

perio

ds: ‘

His

toric

’ (19

56-1

969)

, ‘R

ecen

t’ (2

006,

20

10),

2006

, and

201

0, w

ith n

umbe

r of s

ites (

N) f

or ea

ch d

epth

/tim

e per

iod.

one

-sam

ple t

-test

s wer

e con

duct

ed fo

r com

paris

ons i

nvol

ving

onl

y 20

10 d

eep

site

s (N

=1).

Fam

ilies

w

ith <

1%

mea

n re

lativ

e ab

unda

nce

acro

ss a

ll si

tes w

ere

grou

ped

as “

oth

er”.

Bol

d va

lues

with

* a

re si

gnifi

cant

dif

eren

ces b

etw

een

time

perio

ds (P

≤ 0

.05)

.

Fam

ilyM

ean

rela

tive

abun

danc

e (S

E)t-t

est

(Hist

oric

al, R

ecen

t)t-t

est

(Hist

oric

al, 2

006)

t-tes

t(H

istor

ical

, 201

0)t-t

est

(200

6, 2

010)

His

toric

alr

ecen

t 20

0620

10|t|

P|t|

P|t|

P|t|

P H

oloc

entr

idae

4.7

(0.7

)10

.1 (2

.6)

11.2

(3.0

)5.

52.

210.

05*

2.62

0.03

*0.

440.

680.

870.

45 L

abrid

ae3.

5 (1

.2)

6.1

(1.1

)5.

7 (1

.3)

7.9

1.56

0.15

1.25

0.24

1.34

0.24

0.68

0.53

Lab

risom

idae

7.3

(2.1

)11

.0 (2

.2)

9.3

(1.7

)17

.91.

210.

260.

660.

531.

890.

122.

230.

11 M

orin

guid

ae0.

9 (0

.3)

0.9

(0.4

)1.

2 (0

.5)

0.0

0.15

0.89

0.60

0.56

1.16

0.30

1.11

0.35

Mur

aeni

dae

1.0

(0.3

)1.

3 (0

.6)

1.0

(0.6

)2.

50.

480.

640.

001.

001.

660.

161.

130.

34 P

omac

entri

dae

9.8

(3.5

)4.

4 (0

.8)

3.9

(0.7

)6.

61.

350.

211.

330.

221.

330.

220.

340.

75 S

cari

dae

0.4

(0.1

)2.

2 (0

.4)

2.1

(0.5

)2.

74.

91<0

.001

*4.

240.

003*

6.60

0.00

1*0.

570.

61 S

erra

nida

e2.

6 (0

.8)

4.5

(0.7

)4.

6 (0

.9)

4.0

1.67

0.13

1.58

0.15

0.66

0.54

0.27

0.80

Syn

odon

tidae

1.3

(0.9

)0.

6 (0

.3)

0.8

(0.3

)0.

00.

690.

510.

490.

640.

540.

611.

020.

38 t

etra

odon

tidae

1.7

(0.4

)3.

4 (1

.1)

4.0

(1.2

)0.

81.

450.

182.

070.

070.

790.

471.

160.

33 T

ript

eryg

iidae

5.7

(3.1

)4.

8 (2

.3)

2.6

(0.6

)13

.90.

220.

830.

810.

441.

010.

367.

900.

004*

oth

er6.

5 (1

.2)

5.0

(0.6

)4.

8 (0

.6)

6.0

1.06

0.32

1.10

0.30

0.17

0.87

0.83

0.47