508

The southeast coast of mainland

Florida is within a biogeographic

transition zone of high marine

biodiversity (Briggs, 1974; Gilmore,

1995). This region is also undergoing some of the most rapid human

population growth of any area of the

United States (Culliton et al., 1990).

Due to the economic and recreational value of beaches, substantial marine dredging projects (up to

1.5 × 105 m3 of fill/project) are

commony used to widen beaches

that are subject to erosion in the

area (ACOE, 1996). Nearshore

hardbottom habitats are the primary natural reef structures of this

region at depths of 0–4 m and are

often buried or indirectly affected by

these projects. To date, no quantitative studies of the fish fauna of

these habitats or the effects of beach

dredge-and-fill projects on nearshore fishes are available (NRC,

1995).

Nearshore hardbottom habitats

of this area are derived from accretionary ridges of coquina mollusks,

sand, and shell marl which lithified

parallel to ancient shorelines during Pleistocene interglacial periods

(Duane and Meisburger, 1969;

Hoffmeister, 1974). The habitat

complexity of these limestone structures has been expanded by colonies

Nearshore hardbottom fishes of

southeast Florida and effects of

habitat burial caused by dredging

Kenyon C. Lindeman

Division of Marine Biology and Fisheries,

Rosenstiel School of Marine and Atmospheric Science,

University of Miami

4600 Rickenbacker Cswy., Miami, Florida 33149

E-mail address (for K. C. Lindeman): klindeman@rsmas.miami.edu

David B. Snyder

Continental Shelf Associates, Inc.

759 Parkway St.

Jupiter, Florida 33477

Manuscript accepted 28 August 1998.

Fish. Bull. 97:508–525 (1999).

Abstract.–Fish assemblages of nearshore hardbottom habitats of southeast

Florida were quantified at three sites

from April 1994 to June 1996. Random

2 × 15 m transects were visually censused

within two replicate areas at each site.

The hardbottom at one site was buried

by a dredge project to widen a beach

one year into the study. A total of 394

transects were sampled. Eighty-six

taxa (77 identified to species) from 36

families were censused. Grunts (Haemulidae) were the most diverse family

(11 species), followed by the wrasses

(Labridae) and parrotfishes (Scaridae)

with seven and six species, respectively.

The most abundant species were sailors choice (Haemulon parra), silver porgy

(Diplodus argenteus), and cocoa damselfish (Stegastes variabilis) with mean

abundances (individuals/transect) of

4.5, 3.8, and 3.7, respectively. Early life

stages (newly settled, early juvenile,

and juvenile) represented over 80% of

the individuals at all sites. Newly

settled stages of over 20 species were

observed in association with hardbottom reef structure. Outside of lagoons, nearshore hardbottom areas are

the primary natural structures in shallow waters of mainland Florida’s east

coast and were estimated to have nursery value for 34 species of fishes. After

one year, burial of approximately five

ha of hardbottom habitat at one site

lowered the numbers of individuals and

species by over 30× and 10×, respectively. Due to their early ontogenetic

stage, many of these species may not

be adapted for high mobility in response to habitat burial. Dredging effects may be amplified by burial prior

to and during spring and summer periods of peak larval recruitment.

of tube-building polychaete worms

(Kirtley and Tanner, 1968) and

other invertebrate and macroalgal

species (Goldberg, 1973; Nelson,

1989; Nelson and Demetriades,

1992). In southeast Florida, most

nearshore hardbottom structures

are within 200 m of the shore. These

habitats are often centrally located

between mid shelf reefs to the east

and estuarine habitats within inlets

to the west. Therefore, they may

serve as settlement habitats for

immigrating larvae or as intermediate nursery habitats for juveniles

emigrating out of inlets (Vare, 1991;

Lindeman, 1997a). Nonetheless,

most administrative reviews have

concluded that the fish habitat

value of nearshore hardbottom and

the effects of dredge-based beach

restoration projects are minimal

(e.g. ACOE, 1996).

This study quantifies nearshore

hardbottom fish assemblages on the

southeast coast of mainland Florida

over a 27-month period. The effects

of dredge-fill placement were also

examined because the hardbottom

habitat at one site was buried on

account of a beach restoration

project 12 months into the study.

Three primary objectives were examined. First, spatial and temporal

attributes of fish assemblages at
Lindeman and Snyder: Nearshore hardbottom fishes of southeast Florida 509

Figure 1

Primary study sites for fish surveys of nearshore hardbottom habitats at Jupiter, Florida (26°56'N, 80°04'W).

three undisturbed hardbottom sites were characterized. Second, abundances of different life stages were

compared to assess the potential nursery value of

nearshore hardbottom habitat. Third, effects of

dredge burial on numbers of individuals and species

were compared between a site subjected to burial and

a control site.

Methods

Study areas

Fish abundances were quantitatively surveyed on

two nearshore hardbottom sites approximately 2 km

north (Coral Cove) and 2 km south (Carlin Park) of

Jupiter Inlet, Florida (26°56'N, 80°04'W) from April

1994 through June 1996 (Fig. 1). Sampling at both

sites extended approximately 100 m offshore to a

depth of 4 m. Nearshore hardbottom of similar depth

and structure at Ocean Ridge, immediately south of

the South Lake Worth Inlet (26°31'N, 80°02'W) was

also surveyed for comparative purposes during the

summer of 1995.

Weathered limestone outcroppings were common

between depths of 0 and 4 m at all sites. These structures have a variety of names (e.g. Anastasia formation outcroppings, coquina reefs, worm reefs) but are

referred to by their most common name, “nearshore

hardbottom,” in the present study. In some areas,

the hardbottom extended 1.75 m above the bottom

and was highly convoluted. Shoreward portions of

the hardbottom were exposed at low tide. Epibiota

consisted of a variety of invertebrates and algae. The
510 Fishery Bulletin 97(3), 1999

most widespread encrusting organism was the reefbuilding sabellariid worm Phragmatopoma lapidosa

(=P. caudata; Kirtley, 1994), often covering over 50%

of the hardbottom at all sites.

A beach restoration project occurred at Carlin Park

in March and April of 1995. More than 350,000 m3 of

beach-compatible sediments were excavated by a

cutter-head dredge from a site 0.8 km offshore and

hydraulically pumped along 1.8 km of shoreline.

Bulldozers extended the fill seaward to an estimated

width of 60 m. An estimated total of 4.9 to 5.7 ha

(12–14 acres) of nearshore hardbottom was buried.1

Visual surveys of fishes were conducted for 12 months

before burial and 15 months after burial at both

Carlin Park (the impact site) and Coral Cove (the

control site). Little or no hardbottom was observed

in fish surveys at Carlin Park after the project.

Survey protocol

During each site visit, three to five transects within

two adjacent areas (=6–10 total transects/site) were

censused. These 2 × 15 m transects were randomly

located at depths ranging from one to four m.

Transects were deployed along random compass

headings at random distances between successive

transects. Random number tables were used prior

to site visits to determine the compass headings and

distances between transects (based on numbers of

fin kicks). All fishes observed within one m of each

side of the transect line were identified and tallied

by a snorkeler. The survey zone extended from the

bottom to the surface and 2 m in front of the observer.

Ledges and sand-rock interfaces were examined for

fishes. Rocks were not overturned. The proportion of

hardbottom to sand was estimated within each

transect. An estimated 35% of the area within all

transects was sand. Surveys were conducted between

0900 and 1700 and avoided twilight periods. To compare fish abundances at hardbottom and sand areas, identical transect methods were used at nearshore sand plains greater than 50 m from any hardbottom structures.

Monthly visual censusing occurred from April 1994

to June 1996 as permitted by nearshore visibility and

sea state. Discharges of turbid water from Jupiter

Inlet and wave resuspension of fine sediments sometimes resulted in turbidity levels that precluded sampling. Samples were obtained for all months except

October through January when waves and turbidity

were typically prohibitive.

1 Davis, P. 1998. Palm Beach County Department of Environmental Resources Management, 3323 Belvedere Rd., Bldg. 502,

W. Palm Beach, FL 33406. Personal commun.

In addition to total abundances, early life stages

were also enumerated. Fork length was used for size

estimation. Following Lindeman (1986; 1997a), life

stages of grunts (Haemulon and Anisotremus) were

recorded as follows: newly settled (<2 cm), early juvenile (2–5 cm), juvenile (5–15 cm), and adult (>15

cm). For other families, the same newly settled size

range (<2 cm) was used. The early juvenile designation was used only for grunts because of the distinct

morphological features of the 2–5 cm size range

(Lindeman, 1986). Juvenile and adult stages were

based on size and pigment patterns reported in the

literature (e.g. Robins and Ray, 1986; Humann, 1994).

Species identifications of the newly settled or juvenile stages for certain taxa were limited by very similar morphological features (e.g. scarids, kyphosids,

gerreids, haemulids, clupeids). Some early stage identifications were therefore recorded only at the genus

or family level. Collections of small schools of newly

settled grunts were made with hand nets at both

Jupiter sites in 1994 and 1995 to supplement field

identifications. All collections were deposited at the

Florida Museum of Natural History, University of

Florida.

Data analyses

To address the first objective of our study, two complementary multivariate methods were used to spatially

and temporally characterize the assemblages at the

three sites. The second objective was addressed by

univariate testing of the hypothesis that abundances

of different life stages would not differ significantly

within sites. The third objective was examined with

the hypothesis that numbers of individuals and species would not differ significantly between an impact

site where allmost all the hardbottom was buried and

a control site that was unaffected by the burial. In

univariate analyses, data were standardized as the

mean number of individuals per transect and as the

mean number of species per transect.

Samples were temporally unbalanced owing to the

inability to visually sample during portions of the

winter. Therefore, to examine the first objective,

multivariate ordination and classification (cluster

analysis) of a samples-by-taxa matrix for the entire,

unpooled data set were used. These analyses were

performed on a data set of 31 samples (16 from Coral

Cove, 12 from Carlin Park, and 3 from Ocean Ridge)

and 61 taxa. Each sample represented a site visit

where 6 to 10 transects were censused. Samples from

Carlin Park did not include postdredging site visits

because these samples contained few or no fishes.

The 61 taxa were those remaining from a total of 86

after eliminating taxa occurring only once across all
Lindeman and Snyder: Nearshore hardbottom fishes of southeast Florida 511

samples. Within each sample, counts for individual

taxa were averaged over all transects to provide values for the matrix. These values were log-transformed [log10(n+1)] to prevent abundant taxa from

dominating the ordination or classification results.

The transformed matrix was analyzed by correspondence analysis (CA), a method that employs a

two-way weighted averaging algorithm to produce

simultaneous ordination of sites and taxa (Gauch,

1982; Jongman et al., 1995). These analyses were

performed with the program CANOCO (ter Braak,

1988). From the same log-transformed data matrix,

normal (samples) and inverse (taxa) resemblance

matrices were generated by using the Bray-Curtis

dissimilarity index (Bray and Curtis, 1957). Normal

and inverse resemblance matrices were clustered

separately by the unweighted paired-group method

of averaging (UPGMA) (Sneath and Sokal, 1973). All

dissimilarity and cluster analyses were computed

with NTSYS-pc software (Rohlf, 1997).

To address the second project objective, numbers

of life stages per transect were compared within each

site. Data were analyzed by using a parametric oneway ANOVA when variances were homogeneous

(Bartlett’s test). A posteriori comparisons of differences among means employed Tukey’s HSD test.

Variances of numbers of life stages of grunts per

transect at the two Jupiter sites remained heterogeneous after log10(n+1) transformation and a KruskallWallis non parametric, single classification ANOVA

was used. Probability was calculated using the χ2

approximation. Two-sample t-tests for unequal variances were used to compare numbers of individuals

at hardbottom and natural sand sites. Only hardbottom samples from months when natural sand sites

were sampled (March and April 1995) were used for

these tests. In all statistical tests, differences were

considered significant at P<0.05.

The third objective, examining dredging effects at

the impact site (Carlin Park) and the control site

(Coral Cove), employed a BACIPS (before after control impact paired series) design (Stewart-Oaten et

al., 1986; Osenberg and Schmitt, 1996). This approach compares differences in variables between

sites over time before and after the impact. The differences in the paired series were examined by twosample t-tests by using the mean number of both

individuals and species as the variables (StewartOaten, 1996).

Results

A total of 352 transects was sampled at the two Jupiter sites: 204 at Carlin Park and 148 at Coral Cove.

At Carlin Park, 112 transects were sampled before

burial and 92 after. At Coral Cove, 58 transects were

sampled before the burial of the Carlin Park reef and

90 after. Eight transects were sampled over natural

sand habitats at Carlin Park and six at Coral Cove.

An additional 36 hardbottom and 6 sand transects

were sampled at Ocean Ridge.

Family and species abundances

Thirty-six families of fishes were censused among

the three hardbottom sites. The most speciose family was the grunts and margates (Haemulidae) with

11 species of Haemulon and Anisotremus. The

wrasses, parrotfishes, and damselfishes (Labridae,

Scaridae, and Pomacentridae) had seven, six, and

five species, respectively. Four species each of jacks

(Carangidae), snappers (Lutjanidae), and clinids

(Labrisomidae) were recorded. These seven families

contained 50% of the total species censused. Eightysix taxa (77 identified to species) and 10,491 individuals were censused at all sites (Appendix). At

Coral Cove, 64 species and 5093 individuals were

recorded. At Carlin Park, 53 species and 4438 individuals were recorded. At Ocean Ridge, 48 species

and 960 individuals were recorded. Mean numbers

of both species and individuals per transect were

similar among all sites (Fig. 2).

Figure 2

Mean number of individuals and species per transect (with

95% confidence intervals) for all nearshore hardbottom

sites. Only predredging data were used for Carlin Park

site. Sample sizes: Coral Cove: 148 transects; Carlin Park:

112; Ocean Ridge: 36.
512 Fishery Bulletin 97(3), 1999

Table 1

Mean number of individuals/transect and frequency of occurrence for the most abundant three families, genera, and species at all

nearshore hardbottom sites. Only predredging data were used for Carlin Park site. CC: Coral Cove (148 transects); CP: Carlin

Park (112 transects); OR: Ocean Ridge (36 transects); GM: grand mean.

Mean number/transect % frequency occurrence

CC CP OR GM CC CP OR GM

Family

Haemulidae 15.5 17.4 9.4 15.5 89 90 92 90

Pomacentridae 5.9 7.9 5.7 6.6 81 95 86 87

Sparidae 5.9 3.7 3.9 37 44 38

Labridae 3.2 3.0 65 64

Genus

Haemulon 9.8 15.3 6.2 11.4 75 80 42 75

Stegastes 3.4 6.1 4.3 72 89 73

Anisotremus 5.7 3.2 4.1 74 39 69

Diplodus 5.8 3.7 3.8 35 36 65

Species

Haemulon parra 4.4 5.0 3.4 4.5 62 64 33 59

Diplodus argenteus 5.8 3.7 3.8 35 36 36

Stegastes variabilis 5.4 3.7 86 71

Labrisomus nuchipinnis 3.1 2.7 73 69

Abudefduf saxatilis 3.1 2.3 17 31

Anisotremus surinamensis 3.5 2.2 36 43

The three most abundant species were the sailors

choice (Haemulon parra), silver porgy (Diplodus

argenteus), and cocoa damselfish (Stegastes variablis)

with means of 4.5, 3.8, and 3.7 individuals/transect

over all sites (Table 1). The most abundant species

at Coral Cove, sailors choice, black margate

(Anisotremus surinamensis), and hairy blenny

(Labrisomus nuchipinnis), represented 32% of all

individuals. Seven of the 15 most abundant species

at Coral Cove were grunts. At Carlin Park, silver

porgy, cocoa damselfish, and sailors choice represented 41% of all individuals. Eight of the 16 most

abundant species were grunts. At Ocean Ridge, the

most abundant species were silver porgy, sergeant

major (Abudefduf saxatilis), and sailors choice. Grunt

species ranked first in frequency of occurrence per

transect at Coral Cove and Ocean Ridge, and second

at Carlin Park (Table 1). Damselfish species ranked

first in frequency at Carlin Park and second at the other

sites. The most frequently occurring species overall

were cocoa damselfish, hairy blenny (Labrisomus

nuchipinnus), and sailors choice (Table 1).

Normal cluster analysis of samples from all sites

resolved three groups that broadly reflected temporal patterns (Fig. 3). No distinct spatial groupings

emerged in the normal analysis. Group 1 consisted

of 21 samples (eight from Carlin Park, ten from Coral

Cove, and three from Ocean Ridge) mostly taken in

spring and summer months. Group 2 consisted of 8

samples (four each from Carlin Park and Coral Cove)

taken in mid and late summer. Group 3 included the

only winter samples (February 1995 and 1996) taken

during the project.

Inverse cluster analysis revealed seven groups of

taxa (Fig. 4). Group A contained 26 common taxa

including the most frequently occurring and abundant species from visual surveys such as sailors

choice, cocoa damselfish, hairy blenny, and silver

porgy (Table 1). This group characterized the springsummer group of samples defined by normal group

1. The remaining six groups consisted of taxa that

were temporally variable in their abundance and

occurrence in the samples. Group B was characterized by species that occurred at lower abundances.

Groups F and G were represented by single taxa:

Apogon maculatus and Archosargus probatocephalus,

respectively. The latter species was important in defining normal group 3 (Fig. 3).

Ordination of samples projected on CA axes 1 and

2 produced a pattern that generally agreed with the

normal cluster analysis (Fig. 5A). The eigenvalue for

CA axis 1 was 0.218 and accounted for 16.9% of the
Lindeman and Snyder: Nearshore hardbottom fishes of southeast Florida 513

Figure 3

Clustering of visual census samples (sites and times)

by UPGMA analysis of a normal Bray-Curtis dissimilarity matrix. CC: Coral Cove; CP: Carlin Park;

OR: Ocean Ridge.

variation in the data set, whereas the eigenvalue for

CA axis 2 was 0.124 and accounted for 9.7% of the

variation in the data set. Samples from August and

September at Coral Cove and Carlin Park separated

from all other samples along CA axis 1. In general,

samples were not spread widely along CA axis 2;

however, two samples, May 1995 at Ocean Ridge and

February 1995 at Carlin Park, did separate from the

other sites.

The ordination of taxa on CA axes 1 and 2 showed

how the taxa were distributed in relation to the

hardbottom samples along these same axes (Fig. 5B).

The most common species (e.g. Table 1) clustered near

the origin of the ordination. Taxa with high scores

along CA axis 1 included infrequently occurring species such as Halichoeres poeyi, Haemulon aurolineatum, Mulloidichthys martinicus, and Caranx ruber.

Low scores on CA axis 1 were Echidna catenata,

Acanthurus chirurgus, Chaetodon ocellatus, and

Sciaenidae sp. Species with high scores on axis 2 were

Sparisoma aurofrenatum, Chaetodon ocellatus, and

Sparisoma viride. These were most abundant at Ocean

Ridge in May 1995 and were responsible for the separation of this sample from all others along CA axis 2.

In comparisons of hardbottom and natural sand,

20 transects over natural sand plains recorded only

four taxa. The clupeid, Harengula jaguana, was most

abundant (18 juveniles in two schools total). An unidentified Eucinostomus species, Gerres cinereus, and

Caranx bartholomaei were also recorded (four, one,

and one individuals, respectively). Hardbottom habitats typically had over thirty times the individuals

per transect as natural sand habitats. Two-sample

t-tests comparing hardbottom with sand habitats

rejected the hypothesis of no differences in mean

numbers of individuals per transect (P<0.005).

Life-stage abundances

At all sites, juveniles were the most abundant life

stage among the top ten species (Table 2). At the two

intensively sampled sites in Jupiter, the numbers of

juveniles of all species pooled were significantly

greater than any other life stage (ANOVA, P<0.01,

Tukey’s HSD). There were no significant differences

in abundances among newly settled, early juvenile,

and adult stages (Tukey’s HSD). At both Jupiter sites,

at least 80% of the individuals were early life stages

(pooled newly settled, early juvenile, and juvenile

stages)(Fig. 6). Abundances of newly settled and early

juvenile stages at Ocean Ridge were similar to the

Jupiter sites (Table 2), although numbers of juveniles and adults were lower. Newly settled stages of

over 20 species were recorded on nearshore hardbottom structures among the three sites.

Eight of the ten most abundant taxa by site were

represented primarily by early stages (Table 2). Cocoa damselfish and hairy blenny occurred most abundantly as adults. Adults of at least five of the top ten

species occurred as residents, not transients. These

included sergeant major, hairy blenny, cocoa damselfish, silver porgy, and black margate. Adults of a

variety of less common species occurred but were often less abundant than early life history stages.

Newly settled and juvenile stages often appeared to

display more site-fidelity with hardbottom structure

than did adults.

Six species of grunts (four Haemulon and two Anisotremus) and two species of damselfishes (Stegastes
514 Fishery Bulletin 97(3), 1999

Figure 4

Clustering of fish taxa co-occurrence at three nearshore hardbottom sites by UPGMA analysis of an

inverse Bray-Curtis dissimilarity matrix. Numeric codes used in correspondence analysis are next

to each name. NS: Newly Settled. Dashed lines delineate groups A–G.

and Abudefduf) ranked within the ten most abundant species from all three sites (Table 2). Relative

abundances of the life stages of all grunts censused

at the Jupiter sites are shown in Figure 7. Early juvenile stages of the most abundant species, sailors

choice, were significantly more abundant than any

other life stage at each of the three sites (KruskalWallis ANOVA, P<.001, and a posteriori pairwise

comparisons). Adult sailors choice were significantly

lower in abundance than juvenile stages (KruskalWallis ANOVA, P<0.001). Black margate and

porkfish, Anisotremus virginicus, ranked second and
Lindeman and Snyder: Nearshore hardbottom fishes of southeast Florida 515

third in overall abundance among grunts and were

represented by all life stages (Fig. 7). Tomtate, Haemulon aurolineatum, ranked fourth on the basis of large

but infrequent influxes of early stages. Outside of these

pulses, tomtate was not an abundant or frequently occurring species at any site during any life stage.

Some newly settled grunts could not be positively

identified during visual surveys and were pooled as

Haemulon sp. (newly settled larvae of Anisotremus

are distinctive, Lindeman, 1997a). This group contained epibenthic larvae of several species and

ranked tenth in abundance among all taxa (Table 2)

and fifth among haemulids (Fig. 7). The largest component of these unidentified schools was probably

sailors choice. This assumption is based on 1) the

greater relative abundances of sailors choice early

juveniles at all sites; 2) the close proximity of sailors

choice early juveniles to these newly settled Haemulon

sp.; and 3) collections of several newly settled Haemulon sp. schools most commonly contained sailors

choice upon microscopic examination.

Early stages of commercially valuable species occurred infrequently during the surveys, although

recreationally important species were common. The

most abundant commercial family at the nearshore

hardbottom sites was the Lutjanidae (snappers).

Four snapper species, totaling 58 individuals, were

recorded at all sites. Thirty-eight of these were lane

snapper, Lutjanus synagris. Thirty-three of these

were juveniles, the majority less than five cm. Five

newly settled individuals (<2 cm) were also recorded.

Ten yellowtail snapper, Ocyurus chrysurus, were recorded, nine small juveniles and one newly settled

individual. Unlike grunts, newly settled and small

juvenile snappers were not gregarious, occurring individually or in pairs near interfaces of hardbottom

structure and sand. Gray snapper, Lutjanus griseus,

and schoolmaster, Lutjanus apodus, occurred only

as older juveniles or adults and in low numbers (eight

and two individuals, respectively).

Comparisons of interannual and seasonal patterns

of life stage abundances were limited by temporally

unbalanced sampling. Wind and wave conditions

from September through February made collection

of nearshore visual data in fall and winter erratic or

impossible. Several hurricanes during July and August of 1995 produced wave and turbidity conditions

that interrupted summer sampling as well. Other

phenomena, including high winds and discharges of

turbid water from the Jupiter Inlet, also precluded

sampling for extended intervals. Nonetheless,

samples were obtained from Coral Cove (the control

site) for three consecutive years for the months of

April and June. Comparisons of both numbers of speTable 2

Mean number of individuals/transect by life stage for the ten most abundant taxa at each of three sites. NS: Newly Settled; EJ:

Early Juvenile (for haemulids only); J: Juveniles; A: Adults. Only predredging data were used for Carlin Park site. na = not

available.

Mean number individuals/transect

Coral Cove Carlin Park Ocean Ridge

148 transects 112 transects 36 transects

Species NS EJ J A NS EJ J A NS EJ J A

Haemulon parra 0.7 2.7 0.8 0.2 0.7 3.8 0.8 0.1 0.4 2.6 0.4 <0.1

Diplodus argenteus 0.3 na 1.7 0.3 1.6 na 4.1 0.1 0.7 na 2.8 0.2

Stegastes variabilis 0.2 na 1.4 1.3 0.4 na 1.4 3.5 0.4 na 0.9 0.6

Halichoeres bivittatus 0.2 na 2.3 0.5 0.3 na 2.1 0.5 0.3 na 1.1 <0.1

Labrisomus nuchipinnis 0 na 0.8 2.3 0 na 0.3 2.0 <0.1 na 0.5 1.8

Abudefduf saxatilis 0.4 na 2.0 0.1 0.3 na 1.1 0.4 1.3 na 1.7 0.1

Anisotremus surinamensis 0.4 1.9 1.1 0.1 0.1 0.3 0.3 0 0.5 0.6 0.4 0

Anisotremus virginicus 0.9 0.7 0.4 0.2 0.3 0.9 0.3 <0.1 0.9 0.2 0.4 0.3

Haemulon aurolineatum 0 0.7 1.0 0 0 0.1 2.5 0 0 0 0 0

Haemulon sp. 1.8 <0.1 0 0 2.1 <0.1 0 0 0.9 0 0 0

Haemulon flavolineatum <0.1 0.6 0.3 <0.1 0 0.9 1.0 <0.1 0 1.1 0.1 0

Haemulon chrysargyreum 0 <0.1 0.3 0 <0.1 0.3 1.7 00000

Total means of life

stages per site 4.9 6.6 12 5.0 6.3 6.3 15.5 6.6 5.0 5.4 7.4 3.0
516 Fishery Bulletin 97(3), 1999

Figure 5

(A) Ordination of sample scores on axes 1 and 2 from correspondence analysis of a samples by taxa matrix of census data from Coral Cove (CC), Carlin

Park (CP), and Ocean Ridge (OR). Groups 1–3 from Figure 3. (B) Ordination

of taxa scores from the same correspondence analysis. Numeric codes for taxa

are given in Figure 4.

cies and numbers of individuals per transect for April

among 1994–96 revealed no significant interannual

differences (ANOVA, P=0.34; ANOVA, P=0.21). Identical comparisons for June among the same threeyear period revealed no differences among mean

numbers of individuals (ANOVA, P=0.06), but significant differences among numbers of species

(ANOVA, P<0.05). Pairwise comparisons using

Tukey’s HSD showed that Coral Cove in June 1995

had significantly more species than in June 1996

(mean species numbers: 8.7 versus 5.3).

Seasonal occurrence of only newly settled stages

was examined at Coral Cove (Fig. 8). The timing and

abundance of species occurrences suggested seasonal

variations with peaks of newly settled stages in late

spring and early summer. Sizes and numbers of

individuals typically increased as summer progressed. Abundances of early stages for most species

appeared to be low in February and March prior to

peak spawning activity for many species (GarcíaCagide et al., 1994). However, difficulties in the collection of visual surveys constrained the examination of fall and winter patterns.

Ichthyofaunal characteristics

after habitat burial

Prior to habitat burial, fish assemblages

at the two Jupiter sites were similar in

species composition and relative abundance (Figs. 2, 3, and 5A). Pre- and

postburial numbers of individuals and

species at the control and impact sites

are plotted in Figure 9. The hypotheses

of no differences in total numbers of individuals and species before and after

dredge burial of hardbottom were both

rejected (P<0.001) in two sample t-tests

for equal variances following a BACIPS

design (Stewart-Oaten et al., 1986;

Osenberg and Schmitt, 1996).

No fishes or exposed hardbottom were

recorded in the first postdredging surveys at Carlin Park (13 April 1995). Several hardbottom outcroppings (1 m high

by 3 m wide), parallel to the shore, were

exposed at a depth of 1.5 to 2.5 m at the

site during the second postdredging surveys on 24 April. Two of ten transects

crossed narrow outcrops with dense

schools of newly settled stages of three

grunt species and one drum species

(Sciaenidae). Before burial of the reef

by extension of the beach width by approximately 60 m, such outcrops were

deeper and further offshore than the

original hardbottom sampling area.

Small outcrops were still present in May

and were occupied in two of ten transects by three species of grunts and one

damselfish (Fig. 9). Schools of newly

settled stages predominated. Such outcrops were not encountered during the

20 surveys in June, and no fishes were

present with the exception of several

round scad, Decapterus punctatus. Ten
Lindeman and Snyder: Nearshore hardbottom fishes of southeast Florida 517

Figure 6

Abundances of different life history stages at the Jupiter

hardbottom sites (with 95% confidence intervals). A: All

species pooled. B: Pooled grunt species only. Only predredging data were used for Carlin Park site. NS: Newly

Settled; EJ: Early Juvenile (for grunts only); J: Juveniles;

A: Adults; PE: Pooled Early Stages (=NS+EJ+J).

Figure 7

Comparative abundances of grunts among 12 taxa and 4

life history stages. Data pooled from all Coral Cove surveys and predredging Carlin Park surveys (260 transects

total). Species represented by abbreviated genus and species names.

surveys in September 1995 recorded no exposed outcrops or fishes. During the following winter, erosion

occurred and the width of the new beach was reduced.

Some outcrops were re-exposed by the loss of dredgefill. However, wind and waves prohibited visual sampling during this period. Surveys in February, April,

and May of 1996 (22 transects total) recorded no species (Fig. 9).

Discussion

Fish assemblages of nearshore hardbottom

The diversity of fishes utilizing nearshore hardbottom habitats of mainland Florida has not been

quantified. Qualitative studies by ichthyologists experienced with the substantial taxonomic problems

within these diverse, largely juvenile assemblages

are also lacking. Three studies have included sections on nearshore hardbottom fishes as part of larger

project goals. Gilmore (1977) listed 105 species in

association with “surf zone reefs” at depths less than

two m. Two additional species were added in later

papers (Gilmore et al., 1983; Gilmore, 1992). Using

visual surveys, Vare (1991) recorded 118 species from

nearshore hardbottom sites in Palm Beach County.

Futch and Dwinell (1977) included a list of 34 species obtained from several ichthyocide collections on

“nearshore reefs.” Including species from these prior

studies, 192 species have now been recorded in association with nearshore hardbottom habitats of mainland southeast Florida (Table 3.3 in Lindeman,

1997a). Numbers of labrisomid, blenniid, gobiid, and

apogonid species may be underestimated owing to

their small size or cryptic behaviors. Other hardbottom habitats of the southeast United States occur in areas with substantially different physi-
518 Fishery Bulletin 97(3), 1999

Figure 8

Mean number of newly settled individuals per transect for all

species pooled at Coral Cove, 1994-1996 (n =148 transects).

ographic regimes (Sedberry and Van Dolah, 1984;

Chiappone and Sullivan, 1994) and may show differing patterns of fish diversity.

Spatial and temporal attributes of fish assemblages at the three sites in the present study were

examined by using ordination and cluster analysis. Visual census samples collected from March

through July were similar in species composition

and relative abundance among sites (Fig. 3). This

finding is in agreement with the similar plots of

individual and species abundances among sites

(Fig. 2). The relative homogeneity of these samples

was further reflected in the co-occurrence of many

taxa including haemulids (Haemulon parra, H.

flavolineatum, H. chrysargyreum, Anisotremus

virginicus, A. surinamensis), pomacentrids (Stegastes variabilis, Abudefduf saxatilis), labrisomids

(Labrisomus nuchipinnis), sparids (Diplodus

holbrooki), labrids (Halichoeres bivittatus, Thalassoma bifasciatum) and scarids (Sparisoma rubripinne) (Fig. 4). With the exception of L. nuchipinnis and S. variabilis, most taxa occurred as

early life stages.

Samples from late summer (August and September) were distinct from the spring and early summer in both cluster analysis and ordination (group

Florida can serve as nursery areas for many coastal

fish species. Over 80% of the individuals at all sites

were early life stages. Eight of the top ten species

were consistently represented by early stages. Use

of hardbottom habitats was recorded for newly settled

stages of more than 20 species. In addition, other

natural habitats with substantial vertical relief were

absent from the shallow physiographic regimes

where nearshore hardbottom occurred.

Although suggestive of nursery value, these lines

of evidence need to be viewed in the appropriate context. High abundances of early life stages compared

with adults do not guarantee that a habitat is a valuable nursery. High mortality rates in many reef fish

populations (Sale, 1980; Shulman and Ogden, 1987;

Richards and Lindeman, 1987; Jones, 1991) suggest

that early stages will typically be more abundant

than adults. If spatial distributions of all life stages

are homogeneous, all habitats will have more early

stages than adults. However, the abundances of early

stages on nearshore reefs probably reflect more than

just larger numbers of homogeneously distributed

recruits. Newly settled stages of eight of twelve species of grunts and eight of nine species of snappers

of the southeast mainland Florida shelf have been

recorded primarily in depths less than ten m (Lindeman et al., 1998). Adults of most species are typically uncommon or absent in shallow habitats. There

is considerable evidence for cross-shelf habitat seg2, Figs. 3 and 5). The only two samples taken in winter (February) differed from all other samples in the

analyses (group 3, Figs. 3 and 5). These patterns

suggest that some seasonality in assemblage structure existed. This may reflect late spring and summer peaks in larval settlement in contrast to reduced

winter settlement and, possibly, influxes of older juveniles from inshore lagoonal habitats. Substantial

numbers of many species still settled in late summer but were possibly subject to higher predation

from older individuals that settled earlier in the year.

Various physical disturbances (e.g. winter cold fronts,

summer hurricanes) and biological phenomena

(variation in larval recruitment) affect the composition of fish assemblages of nearshore hardbottom.

The turbidity generated by physical disturbances

constrains the visual surveys needed to assess their

immediate effects.

Nursery habitats and nearshore hardbottom

With increasing human modifications of coastal areas, detailed knowledge of habitat usage is a key component of informed fishery and coastal land management. Identification of essential habitats includes the

evaluation of spatial distributions of structural habitats across the shelf and habitat requirements of key

taxa. Several lines of evidence suggest that nearshore

hardbottom habitats along the mainland coast of east
Lindeman and Snyder: Nearshore hardbottom fishes of southeast Florida 519

Figure 9

Mean numbers of individuals and species at control and impact sites

in Jupiter, FL. Arrows indicate timing of dredge burial of hardbottom

reef.

regation among life stages of many grunt

and snapper species from other regions as

well; early demersal stages appear to most

commonly use shallow habitats (Starck,

1970; Dennis, 1992). Similar ontogenetic

differences in distribution and abundance

exist for many other taxa that utilize

nearshore hardbottom habitats.

Determining if the availability of habitat

structure limits survival of early stages is

important in assessing nursery value. Absences of habitat structure can result in

increased predation or lowered growth

(Hixon, 1991). In southeast mainland

Florida, many natural nearshore marine

habitats outside of coastal lagoons and between 25°30'N and 26°20'N (Dade and

Broward Counties) are sand plains lacking

hardbottom and substantial three-dimensional structure (ACOE, 1996). Although

large stretches of nearshore hardbottom

exist between 26°20'N and 27°50'N (Palm

Beach, Martin, St. Lucie, and Indian River

Counties) these habitats are often separated

by kilometers of sand plains. There are no

other natural habitats in the same nearshore areas that can support equivalent

abundances of early life stages. These conditions could promote a demographic bottleneck that limits local adult populations

owing to limited habitat availability for

early stages.

Despite their shallow depth, nearshore

hardbottom reefs are positioned within current and tide regimes that can support considerable larval abundances. The occurrence of presettlement larvae in these areas is reflected by the abundances of newly settled

stages in the present study and larvae in nearshore

zones of Gulf of Mexico barrier islands (Ruple, 1984;

Ross et al., 1987). Newly settled individuals were not

recorded during any surveys of pure sand habitats

in the present study. However, the presence of

nearshore hardbottom promoted substantial colonization of shallow outcrops by larvae of many species, including haemulids, lutjanids, sparids, labrids,

gerreids, sciaenids, and scarids. Ecotones with high

vertical relief (e.g. hardbottom-sand interfaces near

ledges) sometimes had large aggregations of newly

settled stages of these taxa. However, microhabitatscale distributions of fishes on nearshore hardbottom

remain unquantified.

Use of nearshore hardbottom reefs as nurseries

may be bidirectional across the shelf. Both inshore

and offshore migrations during differing ontogenetic

stages can be facilitated by habitats positioned centrally on the shelf. Nearshore hardbottom may serve

a primary nursery role for incoming early life stages

that would undergo increased predation mortality

without shelter. Nearshore hardbottom may also

serve as secondary nursery habitat for juveniles that

emigrate out of inlets towards offshore reefs. This

pattern is seen in gray snapper and bluestriped grunt

which often settle inside inlets and primarily use

nearshore hardbottom as older juveniles. In addition,

some species use these structures as resident nurseries, settling, growing-out, and maturing sexually

as permanent residents (e. g. pomacentrids, labrisomids). A secondary nursery role may also result

from increased growth because of higher food availabilities in structure-rich environments. The intermediate cross-shelf positioning and other attributes

reviewed above suggest nearshore hardbottom rep-
520 Fishery Bulletin 97(3), 1999

2 Hackney, C. T., M. H. Posey, and S. W. Ross. 1996. Summary

and recommendations. In C. T. Hackney, M. H. Posey, S. W.

Ross and A. R. Norris (eds.), A review and synthesis of data on

surf zone fishes and invertebrates in the south Atlantic Bight

and the potential impacts from beach renourishment, p. 108–

111. Rep. to U. S. Army Corps of Engineers, Wilmington District, Wilmington, NC.

resents essential fish habitat for many species following NOAA (1996). Bidirectional use of nursery

habitats positioned between inshore grassbeds and

offshore reefs requires further study.

From abundance patterns of early life stages and

the absence of any nearby natural habitats with high

vertical relief, nearshore hardbottom of southeast

mainland Florida was estimated to have nursery

value for 34 species (Appendix). Empirical correlation of variation in early survival with adult population size is an important but rarely achieved component of nursery area evaluation. Combining experimental studies of habitat requirements with broad

field surveys can aid in connecting organism-scale

attributes with population-scale patterns (Serafy et

al., 1997). Early demersal stages of several of the

most representative taxa of nearshore hardbottom

(e.g. grunt and damselfish species) can be collected and

manipulated in the field and laboratory with relative

ease (Lindeman, 1986; 1997a). These taxa may serve

as useful models for nursery habitat studies that experimentally assess habitat requirements.

Effects of dredge-and-fill activities on ichthyofauna

Burial of the nearshore hardbottom habitat at Carlin

Park with dredged sand significantly lowered the

abundances of both species and individuals (Fig. 9).

Before burial, 54 species were recorded, with mean

abundances of 38 individuals and 7.2 species per

transect (n=112 transects). After burial, eight species were recorded with mean abundances of less than

one individual and species per transect (n=92

transects). No quantitative studies on the effects of

nearshore hardbottom burial on fishes are available

in the peer-reviewed literature for comparison.

The final supplemental environmental impact

statement (EIS) for the Carlin project (Palm Beach

Co. Dep. Environ. Resources Management, 1994)

summarized several agency and contractor surveys

between 1985 and 1990 at Carlin Park. Ten to fortyeight fish species were recorded from qualitative

surveys of the hardbottom. Statements regarding the

habitat value of nearshore reefs and dredging effects

in the Carlin Park EIS emphasized the variable nature of reef exposure and forecast that fish impacts

would be minimal and temporary. Primary impacts

predicted for fishes were 1) short-term displacement

during construction; and 2) temporary loss of food

sources. The EIS also emphasized that impacts would

be reduced by several features of the project design

and nearshore environment. These features included

the following: 1) the fishery value of impacted species was low; 2) some amount of hardbottom would

remain or would be constructed for mitigation if

needed; and 3) construction of the project would take

place when fish populations were at their lowest. No

mention of direct or indirect mortality upon fishes

was made.

The biological assumptions within this EIS are

similar to those found in related documents (e.g.

ACOE, 1996). For the following reasons, it is suggested that some of these assumptions may be tenuous. The majority of individuals displaced by

hardbottom burial in southeast Florida are early

stages of economically and ecologically valuable species (Appendix; Figure 9). Early demersal life stages

are particularly vulnerable to predators (e.g.

Shulman and Ogden, 1987). Displacement was permanent for most individuals because almost all prior

habitat was eliminated for at least 15 months (the

postburial duration of the present study). Because

of behavioral and morphological constraints on flight

responses, high mortalities are probably unavoidable

for many cryptic species, newly settled life stages, or

other site-associated taxa subjected to direct habitat burial (Table 4.10 in Lindeman, 1997a). Whether

a fish population is seasonally low at the time a

project begins is insignificant if dredging will bury

the habitat immediately before the peak period of

larval settlement,2 as in the Carlin Park project. In

addition, loss of reef-associated food sources was

probably substantial over this period.

No substantial habitat structure was present

within at least 0.8 km of the Carlin Park reef during

its burial. The closest natural structure was eastward at depths of at least 10 m. These deeper

midshelf habitats may be utilized by relatively few

grunt and snapper species during the newly settled

and early juvenile stages. To the south, no substantial hardbottom was present for at least 4 km. To the

north, the jetties of the Jupiter Inlet were approximately 2 km away. However, fishes in a northerly

flight response had to negotiate a water column with

zero visibility because dredge fill was dumped northto-south. Any early stages of fish reaching the jetties would probably encounter high predation from

older piscivores utilizing the large cavities among the

armor-stone boulders of the artificially deepened jetty

area (Lindeman, 1997a).

A postburial mitigation project using shallow artificial reefs of limestone boulders was proposed in the
Lindeman and Snyder: Nearshore hardbottom fishes of southeast Florida 521

Carlin Park EIS. In the summer of 1998, three years

after the burial, construction of approximately 1.6

ha of mitigation reefs began. If constructed before

burial and at similar depths, mitigation reefs may

have provided a refuge for a sizeable fraction of the

thousands of displaced fishes during the burial of the

hardbottom reef, as well as thousands of subsequent

new recruits. Even with prompt construction of artificial reefs, many factors can limit the net production of biomass (Grossman et al., 1997). Some buried outcroppings were uncovered because of erosion

of the project fill. However, structural support for

two years of larval recruitment, shelter from postsettlement predation, and food for growth, were probably eliminated at the hardbottom burial site.

Nearshore hardbottom areas, such as Carlin Park,

can be exposed to extended periods of wave energy

and turbidity, particularly during winter months.

However, conditions in winter do not dilute the potential significance of artificial burial during the

spring and summer months. These are the periods

of peak usage of hardbottom habitats by newly settled

and juvenile stages of fishes. In the absence of dredging, nearshore areas typically show high reef exposures and reductions in physiological stressors during the spring-summer recruitment window. Elimination of this recruitment window by habitat burial

for one or more years, regardless of winter dynamics, may substantially degrade the value of the primary natural nursery habitats along the windward

shorelines of Florida’s east coast. The above reasons

suggest a risk-averse approach to hardbottom burial,

as previously suggested for invertebrate fauna

(Nelson, 1989).

The cumulative effects on fishes of repeated burial

of nearshore habitats and other byproducts of these

projects remain unknown. Cascading disturbances

with ecosystem-scale effects can be hypothesized for

a number of cumulative anthropogenic modifications

in south Florida (e.g. Butler et al., 1995; Ault et al.,

1998). Habitats affected by dredging or filling can

show effects over temporal and spatial scales that

are rarely considered (Vestal and Rieser, 1995;

Lindeman, 1997b). For example, chronically elevated

turbidities could lead to declines in primary production for frequently dredged areas of the southeast

Florida shelf. Conclusive statements on the cumulative effects of large-scale dredging upon fishes will

ultimately depend on the correlation of variations in

early survival with adult population sizes, a rarely

achieved task, even when effects may be substantial

(Osenberg and Schmitt, 1996). However, the current

absence of basic information on both short- and longterm scales can also be treated as an opportunity.

Large dredge projects affecting midshelf and nearshore habitats will continue along the southeast

Florida shelf at one- or two-year intervals. Basic

questions on dredge-and-fill effects upon habitat use,

predation, and growth, await study within a diverse

assemblage of nearshore fishes.

Acknowledgments

J. Ault, J. Bohnsack, G. Dennis, G. Gilmore, P. Glynn,

M. Harwell, and H. Wanless provided substantial

review comments. Several anonymous reviewers

were also very helpful. Conversations with the late

David Kirtley on sabellariid reefs were consistently

valuable. The assistance of these agency personnel

is acknowledged: P. Davis, D. Ferrill, J. Iliff, A. Mager,

and C. Sultzman. Assistance was also provided by J.

Gonzalez, B. Hartig, R. Hudson, C. Leyendecker, M.

Perry, R. Pugliese, M. Ridler, P. Sale, E. Schoppaul,

J. Serafy, A. Stone, K. Snyder, G. Waugh, and D.

Wilder. Funding was provided by the Elizabeth

Ordway Dunn Foundation, the American Littoral

Society, Coastal Research and Education, Inc., the

South Atlantic Fishery Management Council, and the

Dorr Foundation.

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Ocean Office, Silver Springs, MD, 139 p.
524 Fishery Bulletin 97(3), 1999

Appendix

Total abundances of all species visually surveyed at three nearshore hardbottom sites, southeast Florida. Only predredging data

were used for Carlin Park site. * = hypothesized to use nearshore hardbottom as a nursery habitat (see discussion).

Coral Carlin Ocean

Rank Common name Species Cove Park Ridge Total

1 Sailors choice Haemulon parra* 649 555 122 1326

2 Silver porgy Diplodus argenteus* 344 647 132 1123

3 Cocoa damselfish Stegastes variabilis* 420 600 66 1086

4 Slippery dick Halichoeres bivittatus* 439 327 50 816

5 Hairy blenny Labrisomus nuchipinnis* 463 262 81 806

6 Sergeant major Abudefduf saxatilis* 367 199 112 678

7 Black margate Anisotremus surinamensis* 513 68 55 636

8 Porkfish Anisotremus virginicus* 331 174 61 566

9 Tomtate Haemulon aurolineatum* 245 295 8 548

10 Grunt sp. Haemulon sp.* 266 233 34 533

11 French grunt Haemulon flavolineatum* 134 210 43 387

12 Smallmouth grunt Haemulon chrysargyreum* 60 222 10 292

13 White grunt Haemulon plumieri* 70 150 1 221

14 Glassy sweeper Pempheris schomburgki* 153 21 32 206

15 Dusky damselfish Stegastes fuscus* 75 83 9 167

16 High-hat Equetus acuminatus* 54 59 13 126

17 Ocean surgeon Acanthurus bahianus* 51 12 17 80

18 Doctorfish Acanthurus chirurgus* 63 2 7 72

19 Redfin parrotfish Sparisoma rubripinne* 52 14 2 68

20 Mojarra sp. Eucinostomus sp. 37 20 2 59

21 Spanish grunt Haemulon macrostomum* 14 35 1 50

21 Yellow jack Caranx bartholomaei 9 41 0 50

23 Yellow goatfish Mulloidichthys martinicus 34 8 0 42

24 Lane snapper Lutjanus synagris* 23 12 3 38

25 Bluehead wrasse Thalassoma bifasciatum 22 7 7 36

25 Croaker sp. Sciaenid sp. 22 14 0 36

27 Redtail parrotfish Sparisoma chrysopterum* 16 14 3 33

28 Damselfish sp. Stegastes sp. 9 5 18 32

29 Parrotfish sp. Sparisoma sp. 14 14 0 28

30 Reef croaker Odontoscion dentex* 13 3 8 24

30 Bar jack Caranx ruber 2 20 2 24

32 Chub sp. Kyphosus sp. 10 4 9 23

33 Bridled goby Coryphopterus glaucofraenum 2 19 1 22

34 Clown wrasse Halichoeres maculipinna* 8 5 4 17

35 Anchovy sp. Engraulid sp. 15 0 0 15

36 Puddingwife Halichoeres radiatus* 4 6 4 14

36 Orangespotted filefish Cantherhines pullus 2 11 1 14

38 French angelfish Pomacanthus paru* 5 5 3 13

39 Seaweed blenny Parablennius marmoreus 2 5 5 12

40 Caesar grunt Haemulon carbonarium* 3 7 1 11

41 Yellowtail snapper Ocyurus chrysurus* 3 5 2 10

41 Striped croaker Bairdiella sancteluciae* 10 0 0 10

43 Stoplight parrotfish Sparisoma viride 4 149

continued
Lindeman and Snyder: Nearshore hardbottom fishes of southeast Florida 525

43 Redband parrotfish Sparisoma aurofrenatum 1 089

44 Gray snapper Lutjanus griseus 6 028

44 Porgy sp. Sparid sp. 0 8 0 8

44 Bluestriped grunt Haemulon sciurus 2 428

44 Spanish sardine Sardinella aurita 8 008

49 Molly miller Scartella cristata* 2 507

49 Blackear wrasse Halichoeres poeyi 6 107

51 Sheepshead Archosargus probatocephalus 1 506

51 Blue tang Acanthurus coeruleus* 3 306

51 Spotted goatfish Pseudupeneus maculatus 3 036

54 Saddled blenny Malacoctenus triangulatus* 1 315

55 Barbfish Scorpaena plumieri 2 204

55 Queen parrotfish Scarus vetula 0 134

57 Flamefish Apogon maculatus 0 303

57 Yellowfin mojarra Gerres cinereus 3 003

57 Blue runner Caranx crysos 3 003

57 Spotfin butterflyfish Chaetodon ocellatus 0 123

61 Balloonfish Diodon hystrix 1 012

61 Chain moray Echidna catenata 2 002

61 Scrawled cowfish Lactophrys quadricornis 1 012

61 Schoolmaster Lutjanus apodus 2 002

61 Blenny sp. Bleniid sp. 2 0 0 2

61 Cottonwick Haemulon melanurum 2 002

61 Great barracuda Sphyraena barracuda 1 012

61 Scrawled filefish Aluterus scriptus 1 012

69 Bicolor damselfish Stegastes partitus 0 011

69 Orangespotted goby Nes longus 0 101

69 Spanish hogfish Bodianus rufus 0 101

69 Spotted snake eel Myrichthys acuminatus 0 101

69 Gray angelfish Pomacanthus arcuatus* 1 001

69 Sharpnose puffer Canthigaster rostrata 1 001

69 Greater soapfish Rypticus saponaceus 1 001

69 Smooth trunkfish Lactophrys triqueter 1 001

69 Hogfish Lachnolaimus maximus 0 101

69 Puffcheck blenny Labrisomus bucciferus 1 001

69 Nurse shark Ginglymostoma cirratum 0 101

69 Squirrelfish Holocentrus rufus 0 101

69 Blue angelfish Holacanthus bermudensis* 0 011

69 Rosy blenny Malacoctenus macropus 0 101

69 Spotted moray Gymnothorax moringa 1 001

69 Goldentail moray Muraena miliaris 0 101

69 Atlantic spadefish Chaetodipterus faber 1 001

69 Sand drum Umbrina coroides 1 001

Total taxa 72 60 50 86

Total individuals 5093 4438 960 10491

Appendix (continued)

Coral Carlin Ocean

Rank Common name Species Cove Park Ridge Total